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SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM

Bulletin 140

BIRD PARASITES OF THE NEMATODE

SUBORDERS STRONGYLATA,

ASCARIDATA, AND

SPIRURATA

BY

ELOISE B. CRAM

Of the Bureau of Animal Industry, United States Department of Agriculture

lZ6i

o N

UNITED STATES

GOVERNMENT PRINTING OFFICE

WASHINGTON

1927

SMITHSONIAN INSTITUTION

UNITED STATES NATIONAL MUSEUM
Bulletin 140

BIRD PARASITES OF THE NEMATODE

SUBORDERS STRONGYLATA,

ASCARIDATA, AND

SPIRURATA

ELOISE B. CRAM

Of the Bureau of Animal Industry, United States Department of Agriculture

UNITED STATES

GOVERNMENT PRINTING OFFICE

WASHINGTON

1927

ADVERTISEMENT

The scientific publications of the National Museum include two
series, known, respectively, as Proceedings and Bulletin.

The Proceedings, begun in 1878, is intended primarily as a medium
for the publication of original papers, based on the collections of the
National Museum, that set forth newly acquired facts in biology,
anthropology, and geology, with descriptions of new forms and
revisions of limited groups. Copies of each paper, in pamphlet form,
are distributed as published to libraries and scientific organizations
and to specialists and others interested in the different subjects. The
dates at which these separate papers are published are recorded in
the table of contents of each of the volumes.

The Bulletin, the first of which was issued in 1875, consists of a
series of separate publications comprising monographs of large zoo-
logical groups and other general systematic treatises (occasionally
in several volumes), faunal works, reports of expeditions, catalogues
of type-specimens, special collections, and other material of similar
nature. The majority of the volumes are octavo in size, but a quarto
size has been adopted in a few instances in which large plates were
regarded as indispensable. In the Bulletin series appear volumes
under the heading Contributions from the United States National
Herbarium, in octavo form, published by the National Museum since
1902, which contain papers relating to the botanical collections of
the Museum.

The present work forms No. 140 of the Bulletin series.

Alexander Wetmore,
Assistant Secretary , Smithsonian Institution.

Washington, D. C, May 28, 1927.
ii

PREFACE

In the preparation of this paper the writer has been greatly bene-
fited by the advice and assistance of a number of eminent scientific
workers here in Washington. To Dr. Paul Bartsch I am conspicu-
ously indebted for cooperation and helpful advice in connection with
the preparation of this paper. The late Dr. B. H. Ransom, who was
chief of the Zoological Division of the Bureau of Animal Industry
of the Department of Agriculture for the greater part of the time
during which the paper was in preparation, made possible by his
constructive policy the carrying out of such a project by the present
writer. From Dr. Maurice C. Hall, his successor as chief of the
Zoological Division, I have received much valuable advice and, in
addition, I have used his study of the nematode parasites of rodents
as a model. To Dr. Albert Hassall I am indebted for aid in securing
the literature upon which much of this study is based. Dr. H. C.
Oberholser of the Biological Survey has revised the host names and
indicated the synomyns of these names which have been used in the
text; I am greatly indebted to him for this courtesy, involving as it
did in some cases a prolonged search for the identification of a re-
ported bird. To W. S. D. Haines is due much credit for copying the
illustrations. The National Geographic Society kindly aided me in
locating obscure places from which reports of collections had been
made.

E. B. C.

LIST OF FIGURES OF NEMATODES

1. Trichostrongylus tenuis. Male bursa. X150. After Railliet, 1893.

2. Trichostrongylus pergracilis. Bursa; lateral view. Original.

3. Trichostrongylus pergracilis^ Dorsal and externo-dorsal rays of bursa.

Original.

4. Trichostrongylus pergracilis. Terminal genitalia of female. After Ship-

ley, 1909.

5. Ornithostrongylus quadriradiatus^ Male bursa. After Stevenson. 1904.

6. Ornithostrongylus quadriradiatus. Telamon. After Stevenson, 1904.

7. Ornithostrongylus quadriradiatus. Tail of female. After Stevenson, 1904.

8. Ornithostrongylus faridi. Male bursa. After Travassos, 1921.

9. Ornithostrongylus douglasi. Male bursa. After Theiler and Robertson,

1915.

10. Ornithostrongylus douglasi. Tail of female. After Theiler and Robert-

son, 1915.

11. Ornithostrongylus papillatus. Male bursa. After Linstow, 1882.

12. Ornithostrongylus hastatus. Male bursa. After Linstow. 1905.

13. Amidostomum anseris. Anterior end. After Railliet, 1893.

14. Amidostomum anseris. Male bursa. After Railliet. 1898.

15. Amidostomum raillieti. Anterior end. After Skrjabin, 1916.

16. Amidostomum raillieti. Bursa of male. After Skrjabin, 1916.

17. Amidostomum henryi. Anterior end. After Skrjabin, 1916.

18. Amidostomum henryi. Male bursa. After Skrjabin. 1910.

19. Amidostomum henryi. Tail <>f female. After Skrjabin, 1916.

20. Amidostomum ohevreuxi. Anterior end. a. Dorsal view: b. lateral view.

After Seurat, 1918.

21. Amidostomum chevreuxi. Male bursa. After Seurat, 1918.

22. Amidostomum chevreuxi. outline of bursa; lateral view. After Seurat.

191S.

23. Amidostomum monodon. Anterior end. After Linstow, 1S82.

24. Epomidiostomum uncinatum. Anterior end. After Skrjabin. 1916.
'27,. Epomidiostomum uncinatum. Male bursa. After Skrjabin. 1916.

26. Epomidiostomum uncinatum. Female tail. After Skrjabin, 1916.

27. Epomidiostomum orispmum. Male bursa. After Molin, 1861.

28. Codiostomum struthionis. Head end. After Horst, 1885.

29. Codiostomum struthionis. Buccal capsule. After Horst. 1885.

30. GOdiostomum Struthionis. Male bursa, spread out. After Horst. 1885.

31. Codiostomum struthionis. Male bursa, lateral view. After Monnig, 1923.

32. Codiostomum struthionis. Female tail. After Monnig. 1923.

M8. Deletrocephalus dimidiatus. Head; a. lateral; i>, front view. After
Diesing. 1857.

34. Deletrocephalus dimidiatus. Male bursa. After Diesing. ls.~i7.

35. Deletrocephalus dimidiatus. Female tail. After Diesing, lsr>7.

VI BULLETIN 140, UNITED STATES NATIONAL MUSEUM

30. Syngamus trachea. Outline. After Chapin, 1925.

37. Syngamus trachea. Head, a, Teeth in S-toothed (normal) form; b, front

view of head ; c, teeth in 9-toothed form. After Chapin, 1925.

38. Syngamus trachea. Spicules. After Chapin, 1925.

39. Syngamus trachea. Male bursa. After Chapin, 1925.

40. Syngamus microspiculum. a, Male and female; b, head end; c, male

bursa. After Skrjabin, 1910.

41. Syngamus parvus. Outline of pair. After Chapin, 1925.

42. Syngamus parvus. Spicules. After Chapin, 1925.

43. Syngamus parvus. Dorsal portion of male bursa. After Chapin, 1925.

44. Syngamus gracilis. Male and anterior portion of female. After Chapin,

1925.

45. Syngamus gracilis. Male bursa. After Chapin, 1925.
40. Syngamus gracilis. Female tail. After Chapin, 1925.

47. Cyathostoma bronchialis. Head end. o, Side view; b, front view. From

Chapin. 1925, after Muehlig, 18S4.

48. Cyathostoma bronchialis. Male bursa. From Chapin, 1925, after Muehlig,

1884.

49. Cyathostoma tadornae. Male. From Chapin, 1925, after Chatin, 1874.

50. Cyathostoma tadornae. Female tail. From Chapin, 1925, after Chatin,

1874.

51. Cyathostoma lari. Female; entire worm. After Blanchard, 1849.

52. Cyathostoma lari. Female ; head end. After Blanchard, 1849.

53. Cyathostoma lari. Vulva. After Blanchard, 1849.

54. Cyathostoma- varicgatum. Male bursa. From Chapin, 1925, after Linstow,

1890.

55. Cyathostoma rariegatum. Head, a, Front view; b, side view. From

Chapin, 1925, after Linstow, 1890.

56. Cyathostoma boularti. Head ; side view. From Chapin, 1925, after Megnin,

1SS4.

57. Cyathostoma boularti. Male bursa. From Chapin, 1925. after Megnin,

1884.

58. Cyathostoma boularti. Head; front view. From Chapin, 1925, after

Megnin, 1884.

59. Cyathostoma americanum. Outline of female and male worms. After

Chapin, 1925 (a?=extent of uteri).

60. Cyathostoma americanum. Head, a, Side view, male; b, front view; c,

side view, female. After Chapin, 1925.

61. Cyathostoma americanum. Male, a, Bursa; b, gubernaculum. After

(liapin, 1925.

62. Cyathostoma americanum. a, Egg; b. vulva; c, tail (female). After

Chapin, 1925.

63. Cyathostoma coscorobae. Buccal capsules of female and male. After

Chapin, 1925.

64. Cyathostoma coscorobae. Bursa. After Chapin, 1925.

65. 66, and 67. Heterakis gullinae. 65, tail, lateral view; 60, tail, ventral

view ; 67, female tail. After Lane, 1917.
68. Heterakis alata. Male tail. After Schneider, 1866.
69 and 70. Eeterakis arquata. Male tail ; 69, after Schneider, 1866 ; 70, after

Travassos, 1913.

71. Heterakis bancrofti. Male tail. After Johnston, 1912.

72. Heterakis beramporia. Male tail. After Lane, 1914.

73. Heterakis bosia. a, Male tail and point of right spicule ; ft, left spicule

from dorsum ; c, right spicule from side. After Lane, 1914.

LIST OF FIGURES OF NEMATODES VII

74. Hetcrakis brasiliana. Male tail. After Travassos, 1918.
7.~>. Heterakis in-crispiculum. Male tail. After Gendre, 1911.
70. H< U rakis brcvispiculum. Spicule. After Gendre, 1911.

77. Hetcrakis brevispiculum. Region of sucker. After Gendre, 1911.

78. Heterakis gallinae. Region of sucker. After Gendre, 1911.

79. Hetcrakis caudata. Male tail. After Linstow, 1906.

SO. Heterakis chenonettae. Male tail. After Johnston, 1912.
81. Hetcrakis circumvallata. Male tail. After Linstow, 1900.

52. Heterakis dispar. Male tail. After Railliet, 1893.

53. Hetcrakis fariai. Male tail. After Travassos, 1913.
84. Hetcrakis hamulus. Male tail. After Linstow, 1900.

85 and sc. Hetcrakis interlahiatu. 85, anterior end; 86, male tail. After

Ortlepp, 1923.
S7. Hetcrakis isolonche. Male tail. After Linstow, 1906.

88. Heterakis neoplastica. Male tail. After Wassink, 1917.

89. Heterakis long c can data, a, Cuticle; b, male tail. After Linstow, 1879.

90. Hetcrakis macroura. Male tail. After Linstow, 18S3.

91. Hetcrakis papulosa. Male tail, ventral view. After Linstow, 1907.

92. Hetcrakis papulosa. Male tail, lateral view. After Stossich, 1892.

93. Hetcrakis parisi. Male tail. After Blanc, 1914.

94. Hetcrakis psophiac. Male tail. After Travassos, 1913.

95. Heterakis psophiac. Vulva. After Travassos. 1913.

96. Heterakis pusilla. Male tail. After Linstow, 1906.

97. Heterakis putaustralis. Male tail. After Lane, 1914.

98. Heterakis putaustralis. a, Left spicule from side; b, left spicule from

venter ; c, right spicule. After Lane, 1914.

99. Heterakis skrjabini. Male tail. After Skrjabin, 1916.

100. Heterakis tenuicauda. Male tail. After Linstow, 1883.

101. Heterakis tenuicauda. Male tail. After Seurat, 1918.

102. Hetcrakis valvata. Male tail. After Travassos. 1913.

103. Heterakis valvata. Male tail. After Schneider, 1866.

104. Heterakis bonasae. Male tail, a, Ventral view ; b, lateral view. Orig-

inal.

105. Heterakis bonasae. Vulva. Original.

106. Heterakis acuticauda. Male tail. After Cobbold, 1861.

107. Hetcrakis acuticauda. Female tail. After Cobbold, 1861.

108. Ascaridia hermaphrodita. Male tail. After Schneider, 186(5.

109. Ascaridia hermaphrodita. Male tail. After Skrjabin, 1917.

110. Ascaridia galli. Male tail. After Smit, 1922.

111. Ascaridia galli. Male tail. After Baylis and Daubney, 1922.

112. Ascaridia aegyptiaca. Male tail. After Linstow, 1902.

113. Ascaridia amblgmoria. Male tail. After Drasche, 1882.

114. Ascaridia australis. Male tail. After Linstow, 1898.

115. Ascaridia borcalis. Male tail. After Linstow, 1884.

116. Ascaridia brasiliensis. Male tail. After Magalhaes, 1S92.

117. Ascaridia catheturina. Male tail. After Johnston, 1912.

118. Ascaridia circularis. Male tail. After Linstow, 1903.

119. Ascaridia columbae. Male tail, a, Ventral view; b, side view. After

Johnston, 1918.

120. Ascaridia compar. Male tail. After Linstow, 1899.

121. Ascaridia cordata. Male tail. After Linstow, 1906.

322. Ascaridia cristata. a, Head; b, male tail. After Linstow, 1901.
123. Ascaridia cglindrica. Male tail. After Blome, 1909.

VIII BULLETIN 140, UNITED STATES NATIONAL MUSEUM

124. Ascaridia cglindrica. Spicule. After Blome. 1909.

125. Ascaridia dolichocerca. Male tail. After Stossich, 1902.

126. Ascaridia fasciata. Head. After Baylis, 1920.

127. Ascaridia fasciata. Cuticular rings of body. After Baylis, 1920.

128. Ascaridia fasciata. Male tail. After Baylis, 1920.

129. Ascaridia francolina. Male tail. After Linstow. 1899.

130. Ascaridia granulosa. Male tail. After Linstow, 1906.

131. Ascaridia lineata. Male tail. After Boulenger, 1923.

132. Ascaridia longccirrata. Male tail. After Linstow, 1879.

133. Ascaridia magalhacsi. Male tail. After Travassos, 1913.

134. Ascaridia magnipapilla. a, Head ; b, male tail. After Linstow, 1906.

135. Ascaridia numidac. Male tail. After Gendre, 1909.

136. Ascaridia orthocerca. Male tail, ventral view. After Stossich, 1902.

137. Ascaridia orthocerca. Male tail, lateral view. After Stossich, 1902.

138. Ascaridia ptcrophora. Male tail. After Travassos, 1918.

139. Ascaridia scrrata. Male tail. After Schneider, 1866.

140. Ascaridia strelnikowi. a, Head; b, male tail. After Skrjabin, 1916.

141. Ascaridia stroma. Male tail. After Linstow, 1899.

142. Ascaridia styphlocerca. Male tail. After Stossich, 1904.

143. Ascaridia trilabium. Male tail. After Linstow, 1904.

144. Ascaridia anseris. Male tail. After Schwartz, 1925.

145. Pscudaspidodcra pavonis. Head, lateral view. After Baylis and Daub-

ney, 1922.

146. Pscudaspidodcra pavonis. Male tail. Left, lateral view ; right, ventral

view. After Baylis and Daubuey, 1922.

147. Pseudaspidodcra pavonis. Vulva and vagina in lateral view, showing

cement plug in vulva. Arrow points in direction of head. After Baylis
and Daubney, 1922.

148. Subulura. diffcrcns. Anterior end. Original.

149. Subulura diffcrcns. a, Vulva; b, female tail; c, ovejector; d, egg. (Scale

the same for a and b.) Original.

150. Subulura diffcrcns. Male tail. X37.5. After Barreto, 1918.

151. Subulura acutissima. Head, a, Side view ; b, front view. After Drasche,

1882.

152. Subulura acutissima. Male tail. After Drasche, 1882.

153. Subulura allodapa. Anterior end. After Barreto, 1918.

154. Subulura allodapa. Head, front view. After Drasche, 1882.

155. Subulura allodapa. Male tail. After Drasche, 1882.

156. Subulura allodapa. a and 6, Ovejector. After Barreto, 191S.

157. Subulura bcntocruzi. Male tail, lateral view. After Barreto, 1918.

158. Subulura bentocruzi. Male tail, ventral view. After Barreto, 1918.

159. Subulura brumpti. a, Anterior end; b, male tail. After Lopez-Neyra,

1922.

160. Subulura brumpti. Male tail. After Gedoelst, 1916.

161. Subulura brumpti. Male tail. After Gendre, 1909.

162. Subulura carlosi. Male tail, a, Side view; b, ventral view. After Bar-

reto, 1918.

163. Subulura carlosi. Vulva and ovejector. After Barreto, 1918.

164. Subulura curvata. Male tail. After Linstow, 1883.

165. Subulura forcipata. Male tail. After Barreto, 1918.

166. Subulura. forcipata. Vulva and ovejector. After Barreto, 1918.

167. Subulura gallopcrdicis. Male tail. After Baylis and Daubney, 1922.

168. Subulura halli. o, Male tail; b, ovejector; c, buccal cavity. After Seurat,

1914.

LIST OF FIGURES OF NEMATODES IX

JGO. Subulura leprincei. a. Buccal cavity; b. ovejector. After Seurat, 1!)14.

170. Subulura leprincei. a, Male tail, ventral view; b, lateral view; c, spicule,

d, gubeinaculuni : e, egg. After Gendre, 1909.

171. Subulura hi hi. Anterior end. After Barreto, 1918.

172. Subulura lutzi. Male tail, ventral view. After Barreto, 1918.

173. Subulura lutzi. Male tail, lateral view. After Barreto, 1918.

174. Subulura lutzi. Vulva and ovejector. After Barreto, 1918.

175. Subulura noctuae. Male tail. After Seurat, 1914.

176. Subulura olympioi. Male tail, a. Ventral view; b, lateral view. After

Barreto, 1918.

177. Subulura olympioi. Ovejector. After Barreto, 1918.

178. Subulura papilloma. Male tail. From Barreto, 1918 after Drasche.

179. Subulura plotina. Male tail. After Baylis, 1919.
ISO. Subulura poculum. Male tail. After Linstow, 1909.

181. Subulura reclinata. Male tail. After Barreto, 1918.

182. Subulura recurvata. Male tail. After Linstow, 1901.
1S3. Subulura rima. Male tail. After Linstow, 1906.

184. Subulura rimula. Male tail. After Linstow, 1903.

185. Subulura seurati. a, Anterior end; b, male tail; e, ovejector. After

Seurat, 1914.

186. Subulura similis. Male tail. After Gendre, 1909.

187. Subulura strongi/lina. Male tail. After Travassos, 1913.

188. Subulura suctoria. Male tail. From Barreto, 1918 after Drasche, 1882.

189. Subulura suctoria. Head, front view. From Barreto, 1918 after Drasche,

1882.

190. Subulura suctoria. Ovejector. After Barreto, 1918.

191. Subulura travassosi. Ovejector. After Barreto, 1918.

192 and 193. Subulura travassosi. Male tail. After Barreto, 1918. (Figs. 191,
192, and 193 from Bucco chacuru. )

194. Subulura travassosi. From Bucco swainsoni. Male tail. After Barreto,

1918.

195. Subulura trogoni. Anterior end. After Barreto, 1918.

196. Subulura trogoni. Male tail. After Barreto, 1918.

197. Subulura acuticuuda. Male tail. After Linstow, 1901.

198. Subulura gracilis. Head. After Linstow, 1899.

199. Subulura gracilis. Male tail. After Linstow, 1899.

200. Porrocaecum angusticolle. Dorsal lip. After Baylis and Dauhney. 1922.

201. Porrocaecum ttngusticolle. Male tail. After Baylis and Daubney, 1922.

202. Porrocaecum depressum. Dorsal lip. After Baylis and Daubney, 1922.

203. Porrocaecum depressum. Male tail. After Linstow, 1875.

204. Porrocaecum cusicuutlatum. a, Dorsal lip; b, male tail; c, egg. After

Linstow, 1884.

205. Porrocaecum rcticulatum. Male tail. After Baylis and Daubney, 1922.

206. Porrocaecum n ■ticulatum. Dorsal lip. After Linstow, 1899.

207. Porrocaecum semilcrcfi. Anterior end (dissected). After Creplin, 1829.

208. I'orrociK cum serpent ul us. Male tail. After Monnig, 1923.

209. Porrocaecum serpentulus. Dead. After Linstow. 1899.

210. I'orrocaecum serpent ulus. Male tail. After Baylis and Daubney, 1922.

211. Porrocaecum spirale. Dorsal lip. After Linstow, 1875.

212. Porrocaecum spirale. Egg. After Linstow, 1875.

213. Porrocaecum spirale. Male tail. After Linstow, 1875.

214. Porrocaecum heteroura. Anterior end (dissected). After Creplin, 1829.
215«. Porrocaecum anguillac. Anterior end (dissected). After Linstow, 1899.

3012—27 2

X BULLETIN 140, UNITED STATES NATIONAL MUSEUM

215. b to e. Porrocaecum kirghisensis. b. Lip ; c, interlabium ; d, lateral and e,

ventral view of male tail. After Skrjabin, 1916.

216. Contracaecum spiculigerum. Head. After Linstow, 1909.

217. Contracaecum spiculigerum. a, Esophagus and appendices ; b, male tail.

After Schneider, 1866.

218. Contracaecum andersoni. a, Dorsal lip ; &, interlabium ; c, vulva ; d, male

tail. After Vevers, 1923.

219. Contracaecum engonium. a, Head ; b, male tail. After Baylis and Daub-

ney, 1922.

220. Contracaecum haliacti. a, Lateral view; b, front view of head; c, egg.

After Smith, Fox, and White, 1908.

221. Contracaecum haliaeti. a, Inner surface ; b, outer surface of lip. After

Smith, Fox, and White, 1908.

222. Contracaecum microcephalutn. a, Front view of head ; b, dorsal view of

head ; c, male tail ; d, free end of spicule ; e, cross section of spicule.
After Gedoelst, 1916.

223. Contracaecum micropapillatum. Anterior end (dissected). Original.

224. Contracaecum micropapillatum. a, Male tail ; b, lip and interlabia. After

Stossich, 1890.

225. Contracaecum multipapillatum. Digestive organs of male. After Skrjabin,

1916.

226. Contracaecum multipapillatum. a, Head; b, male tail. After Drasche,

18S2.

227. Contracaecum ovale. Head. After Linstow, 1907.

228. Contracaecum praestriatum. Anterior end. After Monnig, 1923.

229. Contracaecum praestriatum. Male tail. After Monnig, 1923.

230. Contracaecum punctatum. Male tail. After Gedoelst, 1916.

231. Contracaecum rodhaini. Head. After Gedoelst, 1916.

232. Contracaecum rosarium. Anterior end. After Connal, 1912.

233. Contracaecum rosarium. Male tail. After Baylis and Daubney, 1922.

234. Contracaecum scotti. a, Esophagus and appendices; b, head; c, lateral

view and d, ventral view of male tail. After Leiper and Atkinson, 1915.

235. Contracaecum tricuspe. a, Male tail ; b, dorsal view and c, front view of

head. After Gedoelst, 1916.

236. Contracaecum magnipapillatum. Male tail. After Chapin, 1925.

237. Spirura uncinipenis. a, Lateral view ; b, front view of head ; c, male

tail. After Drasche, 1884.

238. Spirura uncinipenis. a, Head ; b, egg ; c, free end of short spicule. After

Linstow, 1899.

239. Cyrnea eurycerca. a, Female tail, showing ovejector ; b, male tail ; c,

male (length 7.6 mm) ; d, lateral view and e, ventral view of head.
After Seurat, 1914.

240. Cyrnea excisa. a, Dorsal view ; b, lateral view of head end ; c, female ;

d, male tail. After Seurat, 1914.

241. Cyrnea seurati. Tail end of male and of female. After Lopez-Neyra, 1918.

242. Cyrnea colini. Head, a, Ventral ; b, front ; c, oblique lateral view.

Original.

243. Cyrnea colini. a, Male tail; b, female tail; c, ovejector. (Scale for b and

c the same.) Original.

244. Habronema colaptes. a, Dorsal ; b, lateral ; c, front view of head ; d,

spicules and gubernaculum ; e, male tail ; f , female tail. After Walton,
1923.

245. Habronema ficheuri. a, Lateral; b, dorsal view of head; c, male tail.

After Seurat, 1916.

LIST OF FIGURES OF NEMATODES XI

246. Habronema incerta. a. Male tail; b, eggs; c, female tail; d, transverse

section of anterior end; e, anterior end; f, female, showing position of
vulva; g, front view; h, dorso-ventral view of head. After Smith, Fox,
and White, 1908.

247. Habronema leptoptera. a, Anterior end, dorsal view; b, male tail. After

Seurat, 1914.

248. Habronema leptoptera. a, Head, lateral view ; b, receptacle seminale, join-

ing oviduct with uterus ; c, ovejector. After Seurat, 1914.

249. Habronema longistriata. a, Head, ventral view; b, head, front view; c,

male tail. After Drasche, 18S4.

250. Habronema mansioni. a, Anterior end, ventral view; 6, male tail; e, ove-

jector. After Seurat, 1914.

251. Habronema monoptera. a, Anterior end; b, lateral lip; c, median lip; d,

male tail ; e, left spicule ; f, right spicule ; a, female tail. After Gendre,
1922.

252. Habronema seurati. a. Transverse striation of body; b, cuticular struc-

ture of ventral surface of male tail ; c, male tail ; d, left spicule ; e, right
spicule. After Skrjabin, 1917.

253. Habronema spinoaa. a, Spines of cuticle; b, head end; c, median lip: <Z,

male tail ; e, left spicule ; f, right spicule ; g, gorgeret ; li, female tail.
After Gendre, 1922.

254. Habronema tulontoma. a, Anterior end; b, lateral lip; c, dorsal lip.

After Seurat, 1914.

255. Habronema tulostoma. a, Male tail; b, cloacal region, showing gorgeret;

e, ovejector. After Seurat, 1914.

256. Habronema unilateralis. a, Head, front view; b, male tail. After

Drasche, 1884.

257. Hadjelia Ihuillieri. a, Head end, lateral view; b, head, dorsal vie'v.

After Seurat, 1916.

258. Hadjelia Ihuillieri. Vulva and ovejector. After Seurat, 1910.

259. Hadjelia inermis. a, Male tail as usually seen (most posterior papillae

hidden) ; b, male tail straightened out; c, left spicule; d, right spicule;
e, female tail. After Gendre, 1922.

260. Hadjelia parva. a, Male tail ; b, female tail. xVfter Gendre, 1922.

261. Hadjelia truneata. a, Head; b, male tail. After Mueller, 1S97.

262. Hartertia obesa. a, Dorsal; b, lateral; e, front view of head; d, egg.

After Seurat, 1915.
26.3. Hartertia obesa. a, Ovejector; b, male tail. After Seurat, 1915.

264. Hartertia annulata. a. Lateral; b, front; e, dorsal view of head; d, egg;

e, female tail; /\ ovejector; g, enlarged view of cul-de-sac at union of
sphincter and trompe. Original.

265. Hartertia annulata. Male tail and spicules. Original.

266. Hartertia eonfusa. Male tail. After Linstow, 1900.

267. Hartertia fiallinaram. a, Lateral; b, front view of head; e, female tail.

Original.

268. Hartertia gallinarum. Ovejector and uteri. Original.

269. Hartertia i/alliiuirum. Male tail and spicules. Original.

270. Hartertia rotundata. a. Head, ventral view; b, male tail. After Seurat,

1914.

271. Hartertia rotundata. Ovejector. After Seurat, 1914.

272. Harteria zorillac. Third stage larva, a, Ventral view of head; b, lateral

view of head; C, ventral; d. lateral view of lail. After Seurat. 1919.

273. Viguiera euryoptera. a, Male tail; b, ovejector. After Seurat, 1913.

XII BULLETIN 140, UNITED STATES NATIONAL MUSEUM

274. Gongylonema ingluvicola. a, Head ; b, male tail. After Ransom, 1904.

275. Spiroptera penihamata. Male tail. After Drasche, 1884.

270. Spiroptera turdi. Head, a, Front; b, lateral view. After Drasche, 1884.

277. Desmidocerca aerophila. a, Head ; b, male : c, male tail ; d, female tail.

After Skrjabin, 1910.

278. Acuaria ptilopachydis. a, Anterior end; 6, female tail. After Gendre,

1920.

279. Acuaria qnadriloba. Head, front view. After Schneider, 1800.

280. Acuaria tarentolae. Anterior and posterior ends of third stage larva.

After Seurat, 1910.

281. Acuaria anthnris. Male tail, a, Cross section; b, ventral view. After

Schneider, 1800.

282. Acuaria anthnris. a, Male tail X about 50 ; b, female tail end. After

Linstow, 1873.

283. Acuaria anthnris. Ovejector. After Seurat, 1915.

284. Acuaria cordata. a, Head end ; b, lateral and c, ventral view of male tail.

After Mueller, 1897.

285. Acuaria depressa. Male tail. After Schneider, 1800.

280. Acuaria gracilis, a and b, Anterior end ; c, left and d, right spicule ; e,
male tail; f, female tail. After Gendre, 1912.

287. Acuaria ornata. a, Male tail; b, left spicule; c, head end; d, right

spicule ; e, cordons of the lateral and f, of the submedian fields. After
Gendre, 1912.

288. Acuaria papillifera. Head and tail ends of male. After Linstow, 1878.

289. Acuaria subitla. a and b, Head end ; c, male tail ; d, right and e, left

spicule. After Gendre, 1913.

290. Cheilospirura hamulosa. a, Cordon ; b, head, lateral view ; c, male tail.

After Drasche, 1883.

291. Cheilospirura grureli. a, Anterior end; b, head; c, cordon; d, female

tail. After Gendre, 1912. e, Right and f, left spicule; g, male tail.
After Gendre, 1913.

292. Cheilospimra rotundata. Male tail. After Linstow, 1907.

293. Cheilospirura spinosa. a, Anterior end ; b, part of cordon ; c, vulva ; d,

female tail ; c, spicule. Original.

294. Cheilospirura spinosa. Male tail. Original.

295. Chevreuxia revoluta. Anterior end. After Seurat, 1918.

296. Cosmocephalus diesingi. a, Anterior and middle parts of worm, showing

position of vulva; b, anterior end; c, female tail. After Molin, 1801.

297. Cosmocephalus aduncus. a, Head end; b, male tail. After Stossich,

1S92.

298. Cosmocephalus obvelatus. a, Dorso-ventral view and b, lateral view of an-

terior end ; c, female tail, ventral view. After Seurat, 1919.

299. Cosmocephalus obrelatus. Male tail. After Linstow, 1877.

300. Dispharynx nasuta. Head and tail of male. After Pujardin, 1845.

301. Dispharynx spiralis. Male. After Piana, 1S97.

302. Dispharynx spiralis. Female. After Piana, 1897.

303. Dispharynx spiralis, a, Head end. After Seurat, 1910. b, Ovejector.

After Seurat, 1919.

304. Dispharynx laplantei. Ovejector. After Seurat, 1919.

305. Dispharynx magnilabiata. Male tail. After Drasche, 1884.

300. Dispharynx noctuae. a, A r ulva ; b, anterior end, dorsal view; c, female;
d, anterior end, lateral view ; e, ovejector and vulva. After Seurat,
1913.

LIST OF FIGURES OF NEMATODES XIII

307. Disitharim.r rectovaginata. Head end and tail of male. After Drasche,

L884.

308. EcMnuria jugadornata. a. Cordon. After Soloviev, 1912. b. Head end;

c, male tail. From Skrjabin, 191G. after Soloviev. 1912.

309. EcMnuria uuciiiata. Head end; ventral view. After Seurat, 1919.

310. EcMnuria uuciiiata. a. Head end, dorsal view; It, cross-section and e,

ventral view of male tail. After Schneider, 1S66.

311. Ech in it rin- ardeae, a, Front view and b, lateral view of head; c, female

tail. After Smith, Fox. and White, 1908.

312. EcMnuria calcarata, a. Lateral and b, ventral view of head; c, female

tail. After Drasche, 1884.

313. EcMnuria contorta. a, Anterior end; b, male tail; c, female tail, showing

vulva and anus. After Molin, 1801.

314. EcMnuria decorata. a. Anterior end; b, cordon; c, female tail; d, vulva

and vestibule; e, vulva, ovejector, and uterus; f, male tail; g and h,
spicules. Original.

315. Echinuriu hargilae. a, Anterior end; b, right spicule. After Baylis and

Daubney, 1922.

316. EcMnuria horrida. Head end. After Schneider, 1866.

317. Ech in it rin horrida. Head end. After Stossich, 1S91.

318. EcMnuria leptoptili. a, Head end; ft, male tail; c, left spicule; d. right

spicule (the small drawing showing orifice at free end dilated on
pressure) ; e, female tail. After Gcndre, 1920.

319. EcMnuria longevaginaia. a, Head: b, male tail. After Drasche, 1S84.

320. EcMnuria phoenicopteri. Immature female, a and b, Head end; c, tail;

d, vulva and ovejector. After Seurat, 1916.

321. EcMnuria squamata. a. Anterior end; b, deta'l of cordon. After Linstow,

1883.

322. Rusguniella clonijata. Anterior end. a. Lateral view; b, ventral view.

After Seurat. 1919.

323. Rusguniella vanellL Anterior end. ventral view. After Linstow, 1884.

324. Sciadiocara unibellifera. Head, a, Dorso-ventral view; b, front view.

After Drasche, 1884.

325. Sciadiocara umbelUfera. (/, Male tail; b,. vulva; c, female tail. After

Skrjabin, 1916.

326. Seuratia sMpleyi. <i, Dorso-ventral and b, lateral view of anterior end; c,

male tail : d, female tail ; e, egg. After Stossich, 1900.

327. Seuratia sMpleyi. a and b. Proximal region of ovejector; c, distal region

of ovejector. After Senral, 1916.

328. Streptocara pectinifera. a. Head end; b, cervical papillae; c, male tail.

After Neumann, 1900.

329. Streptocara cirrohamata; a. Head; b, male tail. After Linstow. 1888.

330. Streptocara orassicauda. <i. Head; b, tail of female; c, vulva. After

Skrjabin,1916.

331. Streptocara crassicauda charadrii. a. Head; b, cervical papilla. After

Skrjabin, 1916.

332. Streptocara decora, a. Anterior end; I), male tail. After Dujardin, 1845.
:;:;::. Streptocara recta, a, Dorso-ventral ami h. lateral view of head; c, male

tail; d, spicules. After Mueller, 1S97.

334. Streptocara triaciiucha. Cervical papilla. After Wright, 1897.

335. Streptocara tridentata. a, Male tail. After Linstow. 1909. b, Cervical

papilla. After Linstow, 1N77.

336. Diagram of cordons (as found in SynMmantus) as they would appear

if spread out horizontally. After Schneider, 1866.

XIV BULLETIN 140, UNITED STATES NATIONAL MUSEUM

337. Synhimantus affinis. a, Female genitalia. After Seurat, 1915. b, Head

end. After Seurat, 1916.

338. Synhimantus brevicaudata. a, Head end; b, female tail. After Skrjabin,

1917.

339. Synhimantus dcnticulata. Head end. After Dujardin, 1845.

340. Synhimantus laticeps. Male tail. After Seurat, 1916.

341. Synhimantus laticeps. Head. After Schneider, 1S66.

342. Synhimantus laticeps. Female genitalia. After Seurat, 1920.

343. Synhimantus elliptica. Male tail. After Molin, 1861.

344. Synhimantus hamata. a, Head end. After Linstow, 1879. b, Male tail.

After Linstow, 1877.

345. Synhitnantus mvaginata. a and b, Head end ; c, male tail ; d, right and

e, left spicule; f, female tail. After Gendre, 1913.

346. Synhimantus recta. Male tail. After Drasche, 1884.

347. Synhimantus sagittata. a, Head end ; b, male tail. After Schneider,

1S66.

348. Synhimantus subrecta. a, Male tail ; b, right spicule ; c, profile view and d,

ventral view of left spicule ; e, female tail. After Gendre, 1921.

349. Schistorophus longicornis. a, Lateral view and b, front view of head ; c,

male tail. After Schneider, 1866.

350. Schistorophus acanthocephalicus. Head. After Drasche, 1884.

351. Schistorophus aulieatina. Head end. After Skrjabin, 1916.

352. Schistorophus bidens. Male tail. After Schneider, 1866.

353. Schistorophus bidens. a, Head ; b, male tail. After Lepri, 1898.

354. Histiocephalus laticaudatus. a, Front view and b, lateral view of head ; c,

male tail. After Drasche, 1884.

355. Histiocephalus laticaudatus. Male. After Skrjabin, 1916.

356. Histiocephalus tridens. a, Front view and b, profile view of head ; c, male

tail ; d, anterior end ; e, posterior end, lateral view ; f, left spicule, ven-
tral view ; g, right spicule, lateral view ; h, right spicule, dorsal view ;
i, showing notched appearance when spicule removed from body. After
Gendre, 1921.

357. Serticeps vulvoinflata. a, Head ; b, male tail. After Drasche, 1884.

358. Serticeps vulvoinflata (probably). Labelled Spiroptera mediospiralis but

apparently incorrectly so. See text for discussion. After Drasche,
1884.

359. Physaloptera acuticauda. a, Terminal part of female genitalia ; b, male

tail. After Ortlepp, 1922.

360. Physaloptera alata. a, Head ; b, lip ; c, male tail. After Linstow, 1877.

361. Physaloptera alata Rudolphi of Ortlepp, 1922. Possibly equivalent to

P. galinieri Seurat. See text for discussion. After Ortlepp, 1922.

362. Physaloptera megalostoma. After Creplin, 1829.

363. Physaloptera crosi. a, Head end; b, male tail; c, female tail. After

Seurat, 1914.

364. Physaloptera fusiformis. Male tail. After Linstow, 1902.

365. Physaloptera gaUnieri. a, Head, lateral view; 6, anterior half of body

of female, showing ovejector ; c, male tail. After Seurat, 1914.

366. Physaloptera gemina. Male tail. After Linstow, 1S99.

367. Physaloptera subalata. a, Lip; b, male tail. After Schneider, 1866.

368. Physaloptera truncata. a, Lip; b, male tail. After Schneider, 1S66.

369. Physaloptera brevicauda. Male tail. After Linstow, 1909.

370. Physaloptera malleus, a, Head ; b, male tail. After Linstow, 1883.

371. Physaloptera, species. Head end. After Parona, 1885.

LIST OF FIGURES OF NEMATODES XV

372. Physaloptera oiata. Male tail. After Llnstow, 1907.

373. Physaloptera bulbosa. Male tail. After Linstow, 1906.

374. Thelazia campanulata. a, Head end; b, male tail; c, female tail. After

Travassos, 1918.

375. Thelazia eholodkowskii. Female, a, Head end ; b, cuticular annulations ;

c, tail. After Skrjabin, 1922.

376. Thelazia dacelonis. a, Head end, showing vulva; 6, tail of female: c, tail

of male. After Breinl, 1913.

377. Thelazia lutzi. Female, a, Head end, showing vulva ; b, tail. After

Travassos, 1918.

378. Thelazia lutzi. Male tail. After Travassos, 1918.

379. Thelazia stefeura. Female, a, Head end, showing vulva; b, tail. After

Skrjabin, 1922.

380. Thelazia papulosa. Head ; front view. After Drasche, 1S84.
3S1. Thelazia anolabiata. Head. After Drasche, 18S4.

382. Thelazia (?) cirrura. Head end, showing vulva. Original.

383. Thelazia (?) digitata. Female, a, Head end, showing vulva; b, tail.

After Travassos, 1918.

384. Ceratospira vesiculosa. Male tail. After Schneider, 1866.

385. Ceratospira ophthalmica. a, head end ; b, male tail. After Linstow, 1898.

386. Oxyspirura eephaloptera. a, Male tail : b, head, front view. After

Drasche, 1884.
3S7. Oxyspirura anacanthura. a, Right spicule ; b, male tail. After Drasche,

1884.
388. Oxyspirura anthocliaerae. a, Head end ; b, tail end of female. After

Johnston, 1912.
3S9. Oxyspirura mansoni. a, Dorsal view and b, front view of head : c, egg :

d, embryo escaping from the egg- (e.o.r., circumoral cuticular ring; es.,
esophagus; l.p., amphids or so-called "lateral papillae"; m, mouth; ph..
pharynx, s.lp.. sublateral papillae; s.m.p., submedian papillae.) After
Ransom, 1904.

390. Oxyspirura mansoni. Posterior end of female. (int., intestine; or.,

ovary ; ret., rectum ; vg., vagina ; vul., vulva ; x., cells surrounding rec-
tum.) After Ransom, 1904.

391. Oxyspirura mansoni. Posterior end of male. a, Lateral view. (dct. cj.,

ejaculatory duct; int., intestine; sp., spicules; v.s., seminal vesicle.)
b, ventral view. After Ransom, 1904.

392. Oxyspirura ophthalmica. a, Head ; b, male tail. After Linstow, 1903.

393. Oxyspirura parvovum. a, Head ; b, male tail. After Sweet, 1910.

394. Oxyspirura parvovum. Posterior end of female. After Sweet, 1910.

395. Oxyspirura siamensis. Male tail. After Linstow, 1903.

396. Oxyspirura sygmoidea. Male tail. After Skrjabin, 1916.

397. Oxyspirura tanasijtchnki. a, Head ; ft, right spicule ; e, male tail. After

Skrjabin, 1916.
39S. Oxyspirura brevipenis. Male tail. After Drasche, 1884.

399. Tetrameres paradoxa. a, Female, natural size; b, lateral view, and c,

view from above, of same, enlarged ; d, egg. Enlarged. After Diesing,
1835.

400. Tetrameres paradoxa. a, Head (whether of male or female not stated) ;

6, male tail. After Drasche, 1884.

401. Tetrameres americana. Posterior end of male. Original.

402. Tetrameres americana. a. Head of male; b, junction of testis and vas

deferens; c, head end of female; d, junction of uterus and ovary; e,
copulatory receptaculum of female; f, cross-section of female. Original.

XVI BULLETIN 140, UNITED STATES NATIONAL MUSEUM

403. Tetrameres coccinea. Female, a, Total; b, anterior extremity; c, ovejec-

tor ; d, seminal receptacle at junction of uterus and oviduct. After
Seurat, 1914.

404. Tetrameres cochlcariae. Ovejector. After Travassos, 1919.

405. Tetrameres confusa. a, Female : b, male tail. After Travassos, 1919.

406. Tetrameres dubia. Male. After Travassos, 1919.

407. Tetrameres fissispina. a and b, Head end of male, lateral view (a), dorsal

view (ft), showing the two bifid cephalic spines and the simple body
spines ; c, head end of female, dorsal view. After Seurat, 1920.

408. Tetrameres flssispina. Head and tail of male. After Travassos, 1919.

409. Tetrameres fissispina, a, Ovejector; b, copulatory receptaculum ; c, fe-

male genital system. After Seurat, 1914. d, Tail of larva, ventral
view. After Seurat, 1919.

410. Tetrameres gigas. Anterior and posterior ends of male. After Travassos,

1919.

411. Tetrameres gynaecophila. a, Head of female ; b, tail of male. After

Seurat, 1915.

412. Tetrameres micropenis, a, Tail of male; b, ovejector, showing striated

appearance of musculature. After Travassos, 1919.

413. Tetrameres nouveli. a, Female ; b, ovejector ; e, seminal receptacle and

oviduct ; d, egg with filaments ; e, head end of male ; f, tail end of male.
(500ft scale for a to c; 100/a scale for d to /.) After Seurat, 1914.

414. Tetrameres tetrica. Male. After Travassos, 1919.

415. Microtetramercs cruzi. a. Female ; b, male. After Travassos, 1914.

416. Microtetramercs eontorta. a and b, Males; c, youngest female, showing

tendency to coil ; d and e, mature females ; f, tail of female ; g, ova,
unilateral bib represented on one ; h, mature female, showing course,
of esophagus and intestine ; i, half grown female, coiled in one plane
only. After Weidman, 1913. ;', Wax reconstruction of female. After
Weidman, 1923.

417. Microtetramercs helix, a, Male tail ; b and e, females ; d, female tail.

Original.

418. Microtetramercs inermis. a, Male; b, ventral view and c, lateral view of

tail of same : d, head end of same. (Scale applies to b, c, and d.) After
Seurat, 1913.

419. Microtetramercs incrmte. Female, a, Adult ; b, adult examined perpen-

dicularly to determine manner of coiling ; c, immature ; d, anterior end.
After Seurat, 1913.

420. Microtetra meres inermis. a, Ovejector. After Seurat, 1914. b, Part of

same; c, ovary, oviduct, and initial region of uterus; d, vulva; e and f,
ha (cbing of eggs. After Seurat, 1913.

421. Microtetramercs minima. Male. After Travassos, 1914.

422. Microtetramercs pusilla. Male. After Travassos, 1919.

423. Microtetramercs spiralis. Head and tail of male. After Seurat, 1915.

424. Ancijracanthopsis bihamata. a and b, Head end; c and d, male tail;

e, spicule: f, female tail; g, vulva. After Mueller, 1897.

425. Gtiatliostoma accipitri. a, General appearance ; b, hooks of cephalic

region. After Skrjaliin. 1916.

426. Qnathostoma pelecani. a, General appearance; 6, glandular apparatus :

c, vulva. After Chatin, 1874.

427. EustrongiiUdes iul>ifc.r. a. Male tail; b, head end; c, female tail. After

Jaegerskiold, 1909.

LIST OF FIGURES OF NEMATODES XVII

428. Eustrongylides larvae. Species not determined. «, Front view of head;

b, anterior end; c, outline of male tail; (/, outline of female tail. After
Ciurea, 1924.

429. Eustrongylides africanus, a, Head end; b, female tail; c, exterior of egg;

d, pitting of shell ; e, interior of egg. After Jaegerskiold, 1909.

430. Eustrongylides excisus. a, Head ; b and c, tail of male. After Jaegers-

kiold, 1909.

431. Eustrongylides ignotus. a. Head; b, female tail; c and d, male tail; e

and f, egg. After Jaegerskiold, 1909.

432. Eustrongylides mergorum. a, Female; b, male; natural size; c. head.

After Linstow, 1S77.

433. Eustrongylides mergorum. a, Front view; b, lateral view of head; c, egg.

After Jaegerskiold, 1909.

434. Eustrongylides mergorum. a. Head end ; ?>, female tail ; c and d, inal< j

tail. After Jaegerskiold, 1909.

435. Eustrongylides papillosus. Male tail. After Jaegerskiold, 1909.

430. Eustrongylides perpapillatus. a. Front view: b, lateral view of head;

c, male tail ; d, female tail ; e and f, eggs. After Jaegerskiold, 1909.
437. Hystrichis tricolor. Head and tail ends of female. After Jaegerskiold,

1909.
43S. Hystrichis acanthocephalicus. a, Side view; &, front view of head; c and

d, male tail; e, female tail; f, egg. After Jaegerskiold, 1909.

439. Hyxtrichis coronatus. a. Head end; b, female, natural size; r, female

tail. After Molin, 1860.

440. Hystrichis cygni. Head end. After Molin, 1SG1.

441. Hystrichis neglectus. f/, Head end; b, male tail; c, female tr.il; d, egg.

After Jaegerskiold, 1909.

442. Hystrichis orispinus. Head end. From Jaegerskiold, 1909, after Molin.

443. Hystrichis varispvrvosus. a, Head end; b, spine of anterior cervical region.

After Jaegerskiold, 1909.

444. Hystrichis wedli (Linstow's material), a, Anterior portion of body; b,

one of the most anterior spines, lateral view: c, same, front view;
</, one of the most posterior spines; e, posterior end of body {x, probable
primordiuni of vagina). After Jaegerskiold, 1909.

BIRD PARASITES OF THE NEMATODE SUBORDERS
STRONGYLATA, ASCARIDATA, AND SPIRURATA

By Eloise B. Cram
Of the Bureau of Animal Industry, United States Department of Agriculture

INTRODUCTION

The present Avork attempts to assemble under one cover the de-
scriptions of the species of nematodes of the suborders Strongylata,
Ascaridata, and Spirurata, exclusive of the Filarioidea, found in
birds. No such compilation has until now been made for the nema-
todes of these hosts, although special groups have been worked up;
Ransom has made a study of the nematodes parasitic in the eyes of
birds, and Skrjabin has recently studied the Thelaziidae of birds
and possibly other groups, but his papers are written in Russian and
the publications are not available to us at the present time. Ap-
proximately 50 genera, containing about 500 species, have been dealt
with in the present paper. Until now the descriptions of very few
of these have been available in English; it is hoped that the trans-
lation and assembling of the descriptions, many of them from ob-
scure and little available publications, will be of use to workers in
this field.

The growing economic importance of birds furnishes an additional
reason for the desirability of a study of their parasites. Poultry
raising is of very widespread general interest in this country ; water-
birds, ostriches, and to some extent pheasants are of importance from
an economic point of view. Game birds are being imported and
released in the United States in increasing numbers. Some of the
nematode parasites of these various birds have been shown to have
a marked pathological effect on the host. Nodules and verrucous
growths are produced in chickens and pheasants, and spindle-celi
sarcoma of pheasants has been attributed to species of Ileterakis,
while nematodes of this same genus have been shown to act as vectors
for the organism causing blackhead in turkeys and chickens. Species
of Echinuria form large tumors in the gizzard of birds. It is seen
therefore that a knowledge of nematode parasites of birds has real
practical as well as scientific value, and it is hoped that a study such

l

2 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

as the present one will facilitate identification of nematodes and
stimulate interest in them.

The author has attempted to make a critical study of the species
in the assignment of them to genera. This is at times very difficult,
due to inadequate descriptions. For the same reason the keys which
have been given for all groups dealt with, from species up to orders,
in many cases fall far short of what could be desired. If authors in
describing new species of nematodes would hold to certain minimum
essentials in the details of the description, comparable data would be
available and the making of a key a comparatively easy task ; when,
however, one writer confines his attention to the head and another
to the tail of their respective worms it is often not only difficult but
impossible for a third person to recognize and distinguish the one
from the other.

Viewing the classification of nematodes as a filing system, and as
such capable of being enlarged or contracted to suit the convenience
of workers, the present writer has made several new groups that
seemed necessary, or at least desirable, for coordination. Classifica-
tion has evolved upward from species to genera, genera to families,
etc., and as the number of forms increases below, new groups are
needed above in order to indicate and maintain relative rank. The
present writer has accordingly made several new families and super-
families and has recognized the suborders as previously made by
various authors and the two orders as made by Ward.

Only seven neAV species have been described ; this part of the work
is considered quite subordinate to the assembling and systematic
arrangement of published material. Redescriptions have been given
of several species in which details were lacking and new names to
two previously described species.

The Filarioidea and Trichurata have not been dealt with here;
both these groups have a large number of species in birds, but they
are so poorly known and demand such a great amount of study that
it was deemed inadvisable to attempt to deal with them at the
present time.

The main emphasis has been placed upon the Spiruroidea. as that
is the group involving the largest number of species and the greatest
variety in respect to genera and families of the nematodes of birds.
It may be regarded as primarily and characteristically a group of
bird parasites, the spirurids in other host groups making a much
smaller group. This is due largely to the fact that the spirurids
are heteroxenous nematodes, which, unlike the great majority of
the Filarioidea. except in rare instances, have intermediate hosts
that are eaten. Small arthropods, such as entomostracans and in-
sects, are the usual intermediate hosts of spirurids in the cases where

NEMATODE PARASITES OF BIRDS 6

these hosts are known, and since birds feed to a great extent on these
intermediate hosts, the water birds eating entomostracans and the
insectivorous birds eating insects, it is not surprising that in birds
one finds a great variety of the adult spirurids. The origin of
spirurids in birds of prey calls for a different sort of explanation,
and a clue to this explanation is afforded in the literature on
spirurids. notably in the works of Seurat. Seurat finds that infective
third-stage spirurid larvae in arthropods, when eaten by hosts other
than the final or primary host, as by rodents, migrate into the tis-
sues of such a host and again encyst as third-stage larvae. This
has been regarded as a form of aberrant parasitism, but it appears
quite likely that this is a customary link in the life cycle of spirurids
in birds of prey, the rodent or other host of the encysted third-stage
larva serving as a passive vector for the worms.

In the host list at the end the bird names have been corrected
to conform to present-day usage by ornithologists, and the synonyms
indicated, whereas throughout the text the hosts are listed as previ-
ously reported.

Class NEMATODA

Class diagnosis. — Body limited by a cutinous (chitinous) cuticle
which may be either plain, striate, or ornamented with markings
which may be simple or elaborate. A simple, complete digestive
system in typical forms, consisting of a terminal mouth at the an-
terior end of the body, followed by an esophagus, and this in turn
by an intestine which terminates in an anus in the posterior por-
tion of the body. The intestine a tube, rarely with appendages,
made up of a single la} r er of cells which face the lumen of the intes-
tine on the one side and on the other form the external surface.
Body cavity of problematical nature, containing a body fluid which
has been the subject of considerable investigation but concerning
which opinion varies. Nervous system consisting of a nerve ring
.surrounding the esophagus and of nerve cords directly or indirectly
connecting with this ring. Muscular system consisting of muscles
lining the skin on the inside, excepting on the lateral fields, being
arranged longitudinally, the muscles themselves not lined by a lim-
iting membrane. Sexes usually separate, though in a number of
genera, especially in free-living forms, hermaphroditic, or what
Cobb terms syngonic, a condition which perhaps grades into
parthenogenesis in some forms. In syngonism the same gonad pro-
duces sperm cells and, later, eggs. Genital glands of both sexes con-
sisting of tubular structures lying in the body cavity. In the male
the genital ducts open to a cloaca. Males usually smaller than
females and usually equipped with cutinous (chitinous) copulatory
organs known as spicules. Usually one testis in parasitic forms,

4 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

but often two testes in free-living forms. Males frequently pro-
vided with a membranous structure at the posterior extremity of
the body known as a bursa, this structure reaching its highest de-
velopment in the strongyliform nematodes. Spermatozoa of vari-
able shape, being spherical, conical, elongated, or discoid, without
so-called tails but capable of ameboid motion. Vulva may be
located anywhere on or near (Arduenna) the ventral surface from
near the mouth to near the anus. Ovary and uterus forming a con-
tinuous structure. Ovary with double function of an ovary (s. str.)
and a vitellarium, there being no vitellarium distinct from the ovary.
Usually two ovaries and uteri, but there may be more than 2 (up to
10 or 11 (Turgida) or 15 (Physaloptera) or even more) or only one.
Worms oviparous (in which case the egg may or may not contain a
developed embryo when oviposited), ovoAiviparous, or viviparous.
The word " ovoviviparous " is commonly used in two senses, but
the correct use refers to an egg in which the embryo develops-
and from which it later escapes while still in the uterus; the con-
dition in which an egg containing a developed embryo is oviposited
is correctly referred to as oviparous, with a specification as to the
presence or absence of a developed embryo. Eggs simple, not com-
pound, usually ovoid or elliptical in outline.

KEY TO ORDERS OF NEMATODA

Forms with muscular esophagus of tripartite cross-section__ Myosyringata, p. 4-

Forms with tubular capillary esophagus, the tube embedded iu or otherwise in

relation to a single row of cells, the cells usually extending almost, but not

quite, to the head end of the body and the tube continuing anteriorly to the-

mouth Trichosyringata (not dealt with in this paper).

The validity of the two orders proposed some years ago by Ward
has been generally concurred in by subsequent writers, such as
Rauther, Magath, etc. So far as the present writer is concerned it
is perhaps sufficient to state that the distinction between the two
forms of esophagus noted above seems decidedly important.

Order MYOSYRINGATA Ward, 1917

Order diagnosis. — Nematoda (p. 3) : Nematodes with a muscular
esophagus of tripartite cross-section, as opposed to the capillary
tubular esophagus of the Trichosyringata (whipworms and allied
forms).

KEY TO St BORDERS OF MYOSYRINGATA

1. Heterogamic forms, the parasitic generation consisting of females with no-
males, and the free-living generation consisting of males and females.

Rhabdiasata (not dealt with in this paper) ^

Homogamic forms, the parasitic generation consisting of males and

females 2.

NEMATODE PARASITES OF BIRDS

2. Males usually with a well-developed membranous bursa supported by a sys-

tem of rays, usually 6 paired rays and a single unpaired median dorsal ray ;
or, sometimes, with paired dorsal rays ; rarely without bursa ; buccal

capsule present or absent ; eggs usually thin-shelled Strongylata, p. 5.

Males without a well-developed membranous bursa of the type noted 3.

3. Heteroxenous forms, the larval stages occurring in various intermediate

hosts ; mouth usually with 2 lips, or a larger number of lips or lobes
capable of resolution into 2 basic lips, or without lips ; oviparous, vivi-
parous, or ovoviviparous Spirurata, p. 162.

Monoxenous forms as a rule, exceptional forms (seal ascarids) with inter-
mediate hosts ; mouth usually with 3 lips, exceptionally without lips :
oviparous 4.

4. Meromyarian ; males with 1 spicule, at times reduced, imperfectly chitinized,

or absent Oxyurata (not dealt with in this paper).

Polymyarian ; males usually with 2 spicules Ascaridata, p. 48.

Several of the suborders listed above have already been proposed,
the Strongylata by Railliet and Henry in 1913 and the Ascaridata
and Spirurata in 1915, and the Trichurata by Skrjabin in 1916. It
is, therefore, hardly a radical action to recognize these groups or to
propose coordinate groups for the nematode groups of coordinate
rank and importance. This gradual elevation of groups, which
often begins with the splitting of composite species to assemble the
resultant species in a genus, is a characteristic feature of taxonomy.
It is correlated with the fact that as new species are added at the
bottom, newer and larger filing units must be added at the top to
accommodate the large number of lower groups and to maintain
them as far as possible in their order of relationships and relative
importance. In the nematodes the elevation of groups is crowding
up generic distinctions to subfamily rank, and thereby forcing up all
the groups above. Only the method of experiment, of trial and
error in taxonomy, will determine where the movement should stop,
but at present it seems advisable and necessary to continue along
this line.

Suborder Strongylata Railliet and Henry, 1913

Suborder diagnosis. — Bursate nematodes, the bursa a membranous
structure on the male tail and primarily a clasping organ for attach-
ment to the female. The basic pattern of the rays supporting the
bursa is fairly constant and consists of 2 ventral and 3 lateral rays
on each side and a dorsal group of 1 externo-dorsal ray on each side
and an unpaired median dorsal ray or of 1 externo-dorsal and 1 dor-
sal ray or branch of a dorsal ray on each side. The musculature is
either polymyarian or meromyarian. The body shape ranges from
thick and cylindrical to threadlike or hairlike. The anterior ex-
tremity may be straight or curved dorsally or ventrally. A buccal
capsule of extremely variable form may be present or it may be

6 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

absent; if absent the mouth is usually a simple structure with cephalic
papillae, but without the 2 distinct lateral lips of the spirurids or the
3 distinct and prominent lips of the ascarids and oxyurids. Males
with 2 spicules and females usually with 2 ovaries (the Heligmo-
somidae and Ollulaninae having but one ovary). Vulva situated
anywhere from anterior to posterior body region. Oviparous or
ovoviviparous. First-stage larvae rhabditiform except in meta-
strongyles, in which case filariform.

This suborder is here held to include all of the bursate nematodes.
The grouping of all such forms is warranted on the very distinctive
and important bursa. At the same time it is recognized that the
group includes groups of apparent wide divergence on such char-
acters as musculature and form of larval development.

KEY TO SUPERFAMILIES OF STRONGYLATA

1. Polymyarian ; usually in respiratory or circulatory tract; the bursa is re-

duced, in relation to the size of the worm, by comparison with typical
members of the other superfamilies, and the dorsal ray is often very much
wider than the other bursa rays ; in rare instances the bursa is lacking.

Metastrongyloidea, p. 7.
Meromyarian ; usually in digestive tract as adults, but sometimes in respira-
tory tract (Synffamus), circulatory system (Stronffylits), or tissues
(Stephatiurus) as adults or agamic forms, and often in such locations as
larvae or agamic individuals ; bursa typical, relatively large, and with the
dorsal ray approximately as wide as other members of the bursa rays. 2.

2. Buccal capsule present ; usually relatively thick forms ; usually in digestive

tract, sometimes in respiratory system or tissue as adults.

Strongyloidea, p. 29.

Buccal capsule absent or present as a much-reduced structure (Amidos-

toinum) ; usually relatively slender and sometimes hair-like forms; always

in digestive system Trichostrongyloidea, p. 7.

It appears necessary to restrict the limits of the superfamily
Strongyloidea to those of what has been the family Strongylidae
and to create two new superfamilies from the old Strongyloidea as
defined by Hall in assigning superfamily rank to this group. The
lines of demarcation which previously existed, separating all forms
with a distinct buccal capsule, as the Strongylidae, from all those
without a distinct buccal capsule, the Trichostrongylidae and Meta-
strongylidae, have been broken down by the formation of such new
families as the Syngamidae, Diaphanocephalidae, etc. These latter
have a buccal capsule and should again be coordinated with the
now restricted Strongylidae. In order to reestablish this grouping
on the basis of the presence or absence of a buccal capsule, the
present writer has restricted the superfamily Strongyloidea, here-
tofore including all the bursate nematodes, to those families which
possess a buccal capsule, and is creating the coordinate superfami-
lies Trichostrongyloidea and Metastrongyloidea for those without

NEMATODE PARASITES OF BIRDS 7

a buccal capsule, thereby maintaining these groups in their ancient
status as on a par with the restricted Strongyloidea. This regroup-
ing is necessary in order to keep a definite indication of relative
importance.

METASTRONGYLOIDEA, new superfamily

Superfamndly diagnosis. — Strongylata (p. 5) : Polymyarian.
Usually in respiratory or circulatory tract. Bursa smaller than
those of other Strongylata in comparison with size of worm, and
with the dorsal ray often much wider than other rays, sometimes
several times the size of other rays; in exceptional cases the bursa
is lacking. Peculiar accessory structures, sometimes paired, occur
in some species in place of the single unpaired gubernaculum of
other Strongylata, and there are distinctive cutinized (or chitinized,
in the usual terms) structures at the place where the telamon of
other Strongylata occurs.

Type family. — Metastrongylidae Leiper, 1908.

Family METASTRONGYLIDAE Leiper, 1908

Famihj diagnosis. — Metastrongyloidea (p. 7) : Characters of the
superfamily.

Type genus. — Metastrongylus Molin, 1801.

It has seemed advisable to create this new superfamily to balance
the two others in the Strongylata and to maintain in existence groups
of the same relative status, but as the members of this superfamily
have no known interest in connection with nematode parasites of
birds the group is not given further consideration here.

TRICHOSTRONGYLOIDEA, new superfamily

Superfamily diagnosis. — Strongylata (p. 5) : Meromyarian.
Buccal capsule absent or in some bird nematodes, present but rudi-
mentary. Relatively slender and sometimes hairlike forms. Bursa
of typical form for the suborder, not reduced in relative size.
Always in digestive system.

Type family. — Trichostrongylidae Leiper, 1908.

KKV T<S FAMILIES OF TRICHOSTRONGYLOIDBA

1. Body frequently spirally curled. Females with ] ovary.

Heligmosomidae, p. 8.
Body not spirally curled. Females with 2 ovaries 2.

2. A small buccal capsule present, or if rudimentary (absent?) the head has 4

papillae and epaulets Amidostomidae, p. 17.

Head simple; no buccal capsule and cephalic structures as above.

Trichostrongylidae, p. S.

8 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

HELIGMOSOMIDAE, new family

Family diagnosis. — Trichostrongyloidea (p. 7) : Body frequently
coiled in a spiral, but not so rolled in some genera. The presence of
1 ovary is characteristic of this group among the trichostrongyles.

Type genus. — Heligmosomum Eailliet and Henry, 1909.

Subfamily Heligmosominae Travassos, 1914c

Subfajiuily diagnosis. — Heligmosomidae (p. 8) : Characters of the
family.

Type genus. — Heligmosomum Railliet and Henry, 1909.

As this family and subfamily are not yet represented among
nematodes of birds they are not discussed further and the family
is erected here only in order to maintain uniformity of rank in
connection with other necessary changes.

Family TRICHOSTRONGYLIDAE Leiper, 1912

Family diagnosis. — Trichostrongyloidea (p. 7) : Buccal capsule
absent. Male with well-developed caudal bursa of typical form and
size, and with 2 equal spicules, sometimes accompanied by a gorgeret
or telamon. Eggs segmenting when deposited. Development direct
arid simple. Parasites of the digestive system.

Type genus. — Trichostrongylus Looss, 1905.

Subfamily Trichostrongylinae Leiper, 1908

Subfamily diagnosis. — Trichostrongylidae (p. 8) : Characters of
the family.

Type genus. — Trichostrongylus Looss, 1905.

It will undoubtedly be necessary in the near future to break up
the new superfamily Trichostrongyloidea into a number of families
aiid to create new subfamilies, but in view of the meager representa-
tion of trichostrongyles among birds no consideration has been
given this matter in this paper other than proposing the family
Heligmosomidae.

KEY TO GENERA OP TRICHOSTRONGILINAE

1. Spicules typically trifurcated for two-thirds of their length; prebursal
papillae present; gubernaculum (or, more properly, telamon) typically
with lateral processes tending to form a ring partially encircling the cloaca

as a spicule guide Ornithostrongylus, p. 11.

Spicules not trifurcated, but with twisted appearance due to ridges and with
proximal end thickened at one side ; prebursal papillae rudimentary or
absent; wedge-shaped gubernaculum present Trichostrongylus, p. 9.

NEMATODE PARASITES OF BIRDS 9

Genus TRICHOSTRONGYLUS Looss, 1905

Generic diagnosis. — Trichostrongylinae (p. 8) : Small and
slender worms. Head small, with 3 small lips and with nodular or
punctiform papillae. Esophagus long. Cervical papillae lacking.
Male with caudal bursa with large lateral lobes, without well de-
veloped median lobe, each lateral lobe with G supporting rays; the
dorsal ray short and cleft at the end. Spicules short, spoon-shaped
or spatula-shaped, with a twisted appearance due to the arrangement
of some ridges on them; the proximal end of each spicule thickened
by a disk-like process towards one side. Gubernaculum wedge-
shaped. Prebursal papillae rudimentary or absent. Female with
vulva in posterior half of body, slit-shaped or crescentic, surrounded
by somewhat protruding chitinous lips. Uteri divergent; ovejectors
well developed. Postanal portion of body relatively short, with a
pair of small caudal papillae (?) near the tip. Eggs of moderate
size, thin-shelled, segmenting when deposited. Parasitic in the
duodenum, more rarely the stomach, of herbivores, rodents, man and
birds, and in the ceca of birds.

Type species. — Trichostrongylus retortaeformis (Zeder, 1800)
Looss, 1905.

KEY TO SPECIES OF TRICHOSTRONGYLCS

1. Dorsal ray of bursa unbroken until near tip where it divides into 4 short
digitate processes; gubernaculum in profile the shape of a Turkish slipper;

vulva J,; of body length from tail end Trichostrongylus pergracilis, p. 9.

Dorsal ray split into 2 branches in the posterior third of its length, each
branch being forked at tip; gubernaculum elongate lemon-shaped; vulva
V7 of body length from tail end Trichostrongylus tenuis, p. 10.

TRICHOSTRONGYLUS PERGRACILIS (Cobbold, 1873) Raalliet and Henry, 1909

Synonym. — Strongylus per gracilis Cobbold, 1873.

Host. — Colinus virginianus and Lag opus scoticus.

Location. — Ceca.

Morphology. — Trichostrongylus (p. 9) : Body hair-like, attenu-
ated anteriorly. Mouth with 3 minute lobes. Transverse striation
distinct, especially in anterior portion of body where it gives the
worm a serrate outline as viewed under the microscope. Longitu-
dinal striations not seen.

Male G to 8 mm. long. Bursa (figs. 2 and 3) with 2 lateral lobes
and a small sharply pointed median lobe. The latero-ventral ray
ci lives anteriorly and the externo-lateral ray slightly posteriorly so
that the tips of these 2 rays are separated by a considerably greater
interval than that beween the tips of the 3 lateral rays. Externo-
dorsal rays (50^ in length) are longer than the dorsal ray (36/x in

10

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

length). The dorsal ray split at its tip into 4 digitate processe,-',
the 2 external ones being the most prominent. Spicules short,
strongly chitinized. with thickened edges, crossing each other when
extruded, according to Shipley, so that the right spicule projects to
the left and vice versa. Gubernaculum in profile is the shape of a
Turkish slipper.

Female 8 to 10 mm. long. Tail rather sharply pointed at tip.
Vulva (fig. 4) y 5 to y Q of body length from posterior end, the vulva
a transverse slit with crenelated edges. Eggs 72 to 75/t long by 46^
wide, in the morula stage as passed by host.

Life history. — According to Leiper, the embryo may develop and
the eggs hatch in 36 to 48 hours after the eggs leave the bird, or
under unfavorable conditions in summer hatching may require a
month. The young first-stage larva is 360^ long by 15/a wide, and
has an esophageal bulb and a cylindrical buccal capsule. In 4 or 5
days it may molt to form a second-stage larva 46G> long, which

aosrrm

Fics. 1-4. — 1, Trichostrongylus tenuis. Mali: bursa. X 150. After Rail-
liet, 1893. 2, Trichostrongylus pergracilis. Bursa ; lateral view.
Original. 3, Same. Dorsal and externo-dorsal rays of bursa. Original.
4. Same., Terminal genitalia of female. After Shipley, 1900

slowly loses its buccal capsule. Following the next molt the re-
sultant third-stage larva retains its old cuticle as a sheath. This
larva is very resistant and has the habit of ascending vegetation in
the presence of moisture. When swallowed by a grouse the larva
apparently undergoes two more molts and becomes an adult, ap-
parently reaching the stage of egg production in as little as 4 days.
Distribution. — Europe and North America (United States
(Georgia) ).

TRICHOSTRONGYLUS TENUIS (Mchlis, 1846) Railliet and Henry, 1909

Synonyms. — Strongylus tenuis Mehlis, 1846 (in Creplin. 1846) ;
Strongylus serratus Linstow, 1876.

Hosts. — Anas boschas, A. boschas dofnestlcus., Anser albifrons, A.
anser, A. cinereus, A. cinereus do?nesticus i A. domesticus, A. ferus,

NEMATODE PARASITES OF BIRDS 11

A. ferns domesticux, Gallus //alius, Otis tarda, Perdix cinerea, and
Phasianus colchieus.

Location. — Cecum and small intestine.

Morphology. — Trichostronyylu.s (p. 9). Body hair-like, strongly
attenuated anteriorly. Mouth surrounded by 3 small papillae.
Cuticle longitudinally striated.

Male 5 to 6.5 mm. long. Bursa (fig. 1) with 2 lateral lobes and a
small median lobe which is divided into 2 lobules by a median
depression on its ma rain. The ventro- ventral ray curves anteriorly
and is widely separated from the latero-ventral ray, which runs
parallel with the lateral rays to form a group of 4 comparatively
straight rays extending to the bursal margin; the externo-dorsal ray
is short, being slightly shorter than dorsal ray, and nearer to the
postero-lateral ray than to the dorsal ray; total length of dorsal ray
45/*.: it has a common stem which forms 2 branches in the posterior
third of its length, each branch being forked at the tip. The two
spicules are short (112/* long), twisted, and slightly dissimilar; there
is an elongate lemon-shaped gubernaculum, 63/x long.

Female 7.3 to 7.8 mm. long. The tail terminates in a sharp cone.
The vulva is near the posterior end, about I of the body length from
the tip of the tail. The eggs are elliptical, 65 to 75/u, long by 35 to
42ju wide, and segmenting when deposited.

Life history. — Development is doubtless direct, without inter-
mediate host. The eggs pass in the droppings, develop, and hatch,
the young worms reaching a stage where they will infect birds when
swallowed by them. Experiments on mice have suggested that
cutaneous infection may occur in the trichostrongyles.

Distribution. — Europe (Germany, France, Russia), Asia (Russian
Turkestan), and Xorth America (collected and determined by Foster
from A. cin>evem domesticu*. "Washington, D. C, in 1910, but not
reported until the present time).

Genus ORNITHOSTRONGYLUS Travassos, 1914b

Synonym. — Cephalostrongylus I rwin-Smith, 1920.

Generic diagnosis. — Trichostrongylinae (p. 8) : Slender worms,
attenuated anteriorly. Head swollen or inflated. Chitinous lining
of mouth aperture reduced. Esophagus of medium length. Male
with trilobed caudal bursa : posterior lobe sometimes much reduced;
dorsal ray typically bifurcated and each branch also bifurcated, and
(lie internal of these resultant secondary branches usually bifid at
the tip; ventral rays usually close together; externo-dorsal ray orig-
inating from a common trunk with the dorsal. Prebursal papillae
present. Spicules equal, typically trifurcated for two-thirds of their
length <1 ist ally. Telamon present in type species and one other; it is

12 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

elongate, with 2 lateral processes partially inclosing the cloacal lumen
and forming an incomplete ring through which the spicules pass.
Female with vulva in posterior half of the body. Eggs usually seg-
mented or embryonated in uterus. Parasitic in intestine or proven-
triculus of birds.

Type species. — Ornithostrongylus fariai Travassos, 19145.

The pattern of the bursal rays is so divergent in the species included
by Travassos in this genus as to raise a doubt as to whether all of
these species are congeneric.

KEY TO SPECIES OF ORNITHOSTRONGYLUS

1. Males less than 5 mm. and females less than 6 mm. long. Spicules 140 to

150/t long. Anus 80/i and vulva 800/t from tip of tail. In ostrich.

Ornithostrongylus douglasi, p. 14.
Males more than 6 and females more than 8 mm. long 2.

2. Males less than 7 mm. and females less than 9 mm. long. Spicules 140/*

long, with 2 spines at their anterior third. Eggs 80//, long. In Otis tarda.

Ornithostrongylus papillatus, p. 16.

Males more than 8 mm. and females more than 16 mm. long. Spicules 150 to

285/t long. Eggs not over 75/4 long 3

3. Spicules 220/1 long. Eggs 30/t by 21m- In Europe.

Ornithostrongylus hastatus, p. 17.

Spicules less than 200/x or more than 275/t long. Eggs not less than 70/x by

38/z. In Western Hemisphere and Australia 4

4. Male up to 12 mm. and female up to 24 mm. long. Anus 400/i from tip of

tail. Spicules 150 to 160//. long. From pigeon; United States and

Australia Ornithostrongylus quadriradiatus, p. 12.

Male not over 10 and female not over 20 mm. long. Anus 250 to 284/1 from
tip of tail. Spicules 285/a long. From Leptotila ruf axilla; Brazil.

Ornithostrongylus fariai, p. 13.

ORNITHOSTRONGYLUS QUADRIRADIATUS (Stevenson, 1904) Travassos. 1914

Synonyms. — Strongylus quadriradiatus Stevenson, 1904; Tricho-
strongylus quadriradiatus (Stevenson, 1904) Shipley, 1909; Cepha-
lostrongylus quadriradiatus (Stevenson, 1904) Irwin-Smith, 1920.

Hosts. — Goluiriba livia domestica.

Location. — Intestine.

Morphology. — Ornithostrongylus (p. 11) : Delicate, slender worms.
The cuticle about the head is inflated to form a vesicular enlarge-
ment. The mouth is simple, unarmed and without visible papillae.
Cuticle longitudinally striated. Worm red when freshty collected,
apparently from ingested blood in the intestine. Esophagus 450/^
long.

Male 9 to 12 mm. long. The bursa (fig. 5) is bilobed, with no dis-
tinct dorsal lobe. The ventro-ventral rays are close together and
parallel, their tips curving anteriorly; the medio-lateral ray is com-
paratively straight, the postero-lateral and dorso-lateral rays di-

NEMATODE PARASITES OF BIRDS

13

verging from it and the divergence emphasized at the tips, which
tend to recurve; the externo-dorsal ray is shorter than any of the
previous rays and is bent posteriorly near its middle ; the dorsal ray
is much shorter than the other rays, not extending half way to the
bursal margin; it bifurcates near its tip to form 2 short branches
which are deeply cleft, the internal branches of the cleft terminat-
ing in 2 short tips, and has a stumpy process on each side near its
base. The 2 spicules are 150 to 160^ long, somewhat curved and each
terminates in 3 pointed processes; the spicules are united by a mem-
brane to form a protrusible tube. The telamon (fig. 6) is 65 to 70/i
long, attached to the dorsal wall of the cloaca and has 2 longitudinal
processes extending backward and forward along the cloaca and 2
lateral processes forming a partial ring through which the spicules
protrude.

Figs. 5-7. — Ornithostrongylus quadriradiatus. 5. Male bursa. G,
Telamon. 7, Tail of Female. All after Stevenson, 1004

Female 18 to 24 mm. long. The vulva is 5 mm. from the end of
the tail. The short vagina is followed by 2 powerful muscular ove-
jectors. The tail (fig. 7) tapers to a narrow, blunt end, bearing a
short spine. The anus is 140^ from the end of the tail. Eggs 70 to
75//, long by 38 to 40/x wide, and apparently may be segmenting or
contain embryos when deposited. In normal saline eggs may hatch
in the uterus. ,

Larvae, first stage, 500/* long, with a blunt head, thick body and
slender, pointed tail.

Life history. — Unknown ; probably simple and direct.

Distribution. — United States and Australia.

ORNITHOSTRONGYLUS FARIAI Travassos, 1914b

Host. — Leptotila rufaxilla.
Location. — Intestine.

Morphology. — Ornithostrongylus (p. 11) : Body red, with fine
transverse and longitudinal striations. Head with a dilated cuticle

14 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

extending 78 to 85/a from the end of body. Mouth with a slight dis-
tinct chitinous armature. Excretory pore 26G> from head end.
Esophagus short, 350 to 450> long, slightly dilated posteriorly.

Male 9 to 10 mm. long by 110/x wide. Prebursal papillae quite dis-
tinct. Bursa (fig. 8) trilobed and large. Ventral rays close to-
gether and directed anteriorly; externo-lateral ray remote from
ventrals, but directed anteriorly; medio-lateral and postero-lateral
rays directed posteriorly; externo-dorsal ray more slender than
laterals but almost as long; dorsal ray orginates in a short common
trunk with externo-dorsals, bifurcates distally for two-thirds its
length, these primary branches in turn bifurcating, the resultant
external secondary branches being longer than the internal and
recurved in such a manner that their tips are anterior to those of
the internal secondary branches, which have bifid extremities; the
primary branches have also a short external branch near the proximal
extremity of each. Spicules equal, trifurcated distally, 285/x long.
Telamon elongated, with a transverse process on each side directed
ventrally in the lateral walls of the cloaca, the body of the telamon
64 to 71//. long by 21/* wide.

Female 17 to 20 mm. long by 150/* wide. Vulva in the posterior
half of the body ; vagina short ; ovejector slightly developed. Anus
250 to 284/x from tip to tail. Tail terminates in a very fine spine-
like point 21 to 22/* long. Eggs 71/x by 42/*, segmenting in uterus.

Life history. — Unknown; probably simple and direct.

Distribution. — South America (Brazil).

ORNITHOSTRONGYLUS DOUGLASI (Cobbold, 1882, emend. Gedoelst, 1911)
Travassos, 1920a

Synonyms. — Strongylus douglmsii Cobbold, 1882; Strongylus
douglasi (Cobbold, 1882, emend. Gedoelst, 1911) Gedoelst, 1911;
Trichostrongylus douglasi (Cobbold, 1882, emend. Gedoelst, 1911)
Theiler and Robertson, 1915.

Host. — Strut Mo camelus.

Location. — Proventriculus.

Monphology. — Ornithostrongylus (p. 11) : Body slender, tapering
slightly and suddenly in the cervical region. Worms yellowish red
by reflected light and rather colorless by transmitted light. Trans-
verse striations especially evident in mid-body. Head 20 to 27/i
in diameter. Mouth with 3 small lips, each having a small pro-
tuberance; the mouth aperture triradiate. Four submedian papillae
and 2 lateral structures, either lateral papillae or amphids, in shallow
grooves. Esophagus filariform, 480 to 500/* long. Excretory pore
about 300/i behind head end.

Male length averages 4.65 mm. ; width about 93 to 95/«,. Posterior
portion of body slightly twisted longitudinally and tail end slightly

NEMATODE PARASITES OF BIRDS

15

curved. Bursa (fig. 9) with a large lateral lobe and a small dorsal
lobe. The dorsal, lateral, and ventral ray systems originate in 3
stems, one to each system. Ventro-ventral ray slender and curved
ventrally, diverging from latero-ventral; the latter thicker and
closer to laterals. Postero-lateral ray thin and bent dorsally ; medio-
latcral and externo-lateral rays thicker and bent ventrally. Main
stem of dorsal ray branches, each main branch in turn bifurcating
near its base to form secondary branches, and the internal secondary
branches bifurcating near their tips; externo-dorsal ray short, ex-
tending only halfway to bursa margin. Prebursal papillae, slender,

Figs. 8-10. — 8. Ornithostrongylus fabiai. Male bursa. After Travassos, 1021. 9,
Ornithostrongylus douglasi. Male bursa. 10, Same. Tail of female. Nos. 9-10
after theiler and robertson, 1915

easily overlooked. Spicules 140 to 158/a long, dark brown, the proxi-
mal ends knob-like, the body longitudinally grooved and with a
convexly curved ridge on one side of the groove, and the distal end
terminating in a spine, a second spine originating in the posterior
third of the spicule. The telamon is lancet-shaped in outline, but
curved about the cloaca, with a short blunt spine directed backwards,
and is light brown.

Female length averages 5.G3 mm. ; width about 105 to 109/x. Body

terminates in a blunt point 80/x long, its tip slightly curved (fig. 10).

Vulva a transverse or oblique slit about 800/x from the tip of the tail.

A short vagina leads to the muscular ovejector, which is about

3(512—27 3

16 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

300/* long. Posterior uterus smaller than anterior. Eggs 59 to 74/*
by 36 to 44/*.

Life history. — Eggs segmenting in ovejectors; in morula stage in
fresh feces. Under favorable conditions eggs hatch in 59 to 98 hours.
First-stage larva 240 to 750/* long by 18 to 28/* wide; rhabditiform
and with a tapering tail continued in a thin cylindrical appendage
bearing a small knob at its tip. Molting under favorable conditions
occurs in 50 hours; the second-stage larva 600 to 900/* long, with a
tail longer and finer than in preceding and with rounded tip. Molt-
ing under favorable conditions occurs in 60 hours; the third-stage
larva is ensheathed at first and has a small spined knob on the end
of the tail; with the old cuticle the larva averages 745/* long and
without it 530/* long. Molting occurs in the host in 4 to 5 days ; tl
fourth-stage larva gradually developing to an adult under its cuticle,
with evident sex differences in the larvae. Molting to adults occurs
in about 3 weeks. Eggs are present in the ovejectors in 33 days.
This life history was ascertained by Theiler and Robertson.

Distribution. — South Africa.

ORNITHOSTRONGYLUS PAPILLATUS (Linstow, 1882) Travassos. 1920a

Synonym. — Strongylus papillatus Linstow, 1882.

Host. — Otis tarda.

Location. — Intestine.

Morphology. — Ornithostrongylus (p. 11) : Small and delicate
forms, transversely striated, especially at cephalic extremity. Cervi-
cal papillae present a short distance from the head end and very
small.

Male 6.7 mm. long by 72/*. wide, attenuating anteriorly. Esophagus
1/9 of body length or 740/* long. Spicules 140//, long, thick and with
2 spines in the anterior third. Telamon fusiform, 72/* long. Bursa
(fig. 11) with 2 large lateral lobes and a small median lobe. Lateral
rays and latero-ventral ray form a symmetrical group apart from
the ventro-ventral and the externo-dorsal ; the extremities of the rays
are curved. Two dorsal rays originate with their respective externo-
dorsal and converge to meet and fuse at their tips where they form
a bifid termination; previous to the fusion they each give off an
external branch. Prebursal papillae present. (If the structure of
this dorsal ray is correctty described and figured, it raises a doubt as
to whether this species should be included in Ornithostrongylus.)

Female 8.4 mm. long by 84/* wide. Esophagus 1/11 of body length
or 760/* long. Vulva in posterior body, dividing body in ratio of
41 : 7, or about 1.45 mm. from tail end. Eggs colorless, 80/* by 36/*.

Life history. — Unknown; probably simple and direct.

Distribution . — Europe.

NEMATODE PARASITES OF BIRDS

17

ORMTHOSTRONGYLUS HASTATUS (Linstow, 1905) Tr.ivassos, 1920a

Synonym. — Strongylus hastatus Linstow, 1905.

Hosts. — Lyrurus tetrix {Tetrao tetrix).

Location. — Intestine.

Morphology. — Ornithostrongylus (p. 11): Cuticle transversely
striate and with a lateral cephalic ala originating 230/* from the head
end. Head with 6 rounded papillae. Buccal aperture leads into a
vestibule with parallel walls and 78/* deep. Esophagus 5/32 of body
length or about 100 to 300^ long. Excretory pore at level of posterior
third of esophagus.

Male 8.7 mm. long by 190/* wide. Bursa (fig. 12) with 3 lobes, the
dorsal small but sharply defined. Ventral rays close together and
parallel ; the posterolateral set off from the medio-lateral and ex-
terno-lateral, which are close together and parallel ; the externo-

Figs. 11-12. — 11, Ornithostrongylus papu.latus. Male bursa. After Linstow,
1882. 12, Ornithostrongylus hastatus. Male bursa. After Linstow, 1905

dorsal close to and parallel to the externo-lateral most of its length,
but diverging distally; the dorsal with a long stem bifurcating near
its tip and with 2 small external branches near the bifurcation and
extending to incisions in the dorsal lobe. Spicules 220/* long, each
terminating in a lancetlike prolongation which is enlarged in its
median portion.

Female 21.5 mm. long by 530/* wide. Vulva near the posterior
extremity. Eggs thick-shelled, 30/* by 21/* in diameter.

Life history. — Unknown; probably simple and direct.

Distribution. — Europe.

Family AMIDOSTOMIDAE Baylis and Daubney, 1926

Family diagnosis. — Trichostrongyloidea (p. 7): Forms with a
reduced buccal capsule present; in forms with the capsule much re-
duced or, according to Skrjabin. absent, the head has 4 papillae or
papilliform lips and 2 epaulets.

Type genus. — Amidostomum Railliet and Henry, 1909.

18 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Subfamily Amidostominae Travassos, 1920b

Subfamily diagnosis. — Amidostomidae (p. 17) : Characters of the
family.

Type genus. — Amidostomum Railliet and Henry, 1909.

Although this family and its subfamily are a little difficult to
characterize, it appears desirable to have these groups to cover the
included genera, Amidostomum and E pomidiostomum. These genera
appear to be closely related in their general characteristics and in
their locations in bird hosts. The presence of a reduced buccal cap-
sule in a bursate nematode which has affinities in other respects with
the trichostrongyles suggests that Amidostomum is a transitional
form between the trichostrongyles and the group of strongyles with
a buccal capsule, a group here regarded as the Strongyloidea. Epo-
midiostomum has a buccal capsule, according to Seurat, but accord-
ing to Skrjabin, it does not have a buccal capsule; its affinities ap-
pear to be with Amidostomum in any case. The combination of a
buccal capsule and trichostrongyle affinities has caused a diversity
of opinion among helminthologists as to the group to which such
worms should be referred, Travassos referring them to the Strong-
ylidae and Skrjabin and Seurat to the Trichostrongylidae. While
the decision as to the closest affinities of transitional forms does not
appear to be very important, some decision must be made and the
present writer is inclined to concur with Skrjabin and Seurat in re-
ferring them to the trichostrongyles. In passing it may be noted
that the cuticular cephalic structures of amidostomes are very simi-
lar to those of the spirurids in birds, and that another spirurid, lack-
ing a strongyle bursa, as for instance, Rictularia, has a buccal
capsule.

KEY TO GENERA OF AMIDOSTOMINAE

1. Mouth followed by a globular depressed buccal capsule provided with pointed
teeth at its base. Head without 4 outwardly directed papillae and 2 pos-
teriorly directed epaulets. Spicules divided into 2 branches. Gorgeret

or telamon present Amidostomum, p. 19.

Mouth with a much reduced buccal capsule (Seurat) or without buccal cap-
sule (Skrjabin). Head with 4 outwardly directed papillae and 2 pos-
teriorly directed epaulets. Spicules divided into 3 branches. Gorgeret or
telamon absent Epomidiostomum, p. 26.

Since the above key was written, Boulenger (1926) has made a
new genus, Pseudamidostomum, which probably should be included
in this subfamily ; however, since the male is unknown, only a tem-
porary assignment can be made. This genus (see Addenda, p. 383)
differs from Amidostomum in having no teeth at the base of the

NEMATODE PARASITES OF BIRDS 19

buccal capsule and from E ' pomidiostomum in having no posteriorly
directed epaulets.

Genus AMIDOSTOMUM Raillet and Heary, 1909

Generic diagnosis. — Amidostominae (p. 18) : Worms of general
trichostrongyle conformation and affinities, but with a buccal cap-
sule provided with 1 or 3 pointed teeth at its base; the buccal cap-
sule is comparatively wide and thick-walled. The esophagus has 3
chitinous axial lamellae along its entire length or except for a
terminal unarmed region, the bulb. Tail of female elongated and
digitiform, with lateral caudal pores about the middle of its length.
Tail of male with bursa having 2 large lateral lobes; a median lobe
distinctly or indistinctly defined. Dorsal ray bifurcates to form
2 branches, each of which ends in a bifurcation with the tips approxi-
mately on the border of the dorsal lobe. Externo-dorsal ray origi-
nates in some cases (A. chevreuxi) at the base of the dorsal ray in
a common stem, but in other cases (A. raillieti) it originates in a
mass from which all the rays except the dorsal take origin. There
is a shortening of the externo-dorsal and the externo-lateral evi-
denced by a failure to closely approach the bursal margin in any
case and to be greatly reduced in extreme cases, which may be re-
garded as a feature of distinctive generic value. The medio-lateral
and postero-lateral rays are contiguous proximally, but divergent
distally. The ventro-ventral and latero-ventral rays are widely di-
vergent, the latter more closely associated with the shortened ex-
terno-lateral ray than with the ventro-ventral. A pair of volumi-
nous almost contiguous papillae occur on the posterior lip of the
cloaca. Prebursal papillae present. Spicules equal, each divided
into 2 separated prolongations posteriorly along most of their length.
A straight gorgeret or telamon present. Vulva a transverse slit,
posterior to the middle of the body. Ovejectors divergent and op-
posed; uteri opposed; ovaries parallel, very long, extending towards
the anterior end of body.

Type species. — Amidostomum anseris (Zeder, 1800) Railliet and
Henry, 1909.

At the time this genus was proposed by Railliet and Henry, they
designated A. anseris (Zeder) as type. Subsequently Seurat has
designated Strongylvs nodulosus Rudolphi as type, stating that Zeder
confused worms belonging in 2 genera {Amidostomum and Epomi-
diostomuni) in his Strongylus anseris. Under the zoological code,
Amidostomum is fixed by its type and must stand or fall with it; a
new type can not be designated for the reason given by Seurat. The
proper procedure appears to be to restrict S. anseris to one of
Zeder's species.

20 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

KEY TO SPECIES OF AMIDOSTOMUM

1. Female 16.5 mm. long ; buccal capsule apparently with only 1 large tooth ;

vulva not described as prominent ; male unknown ; imperfectly described

species from Oidemia nigra in Europe Amidostomum monodon, p. 2G.

Female not over 13 mm. long, or, if longer, then with a buccal capsule having
3 teeth, or vulva with prominent anterior lip (A. aciitum) 3.

2. Males 11.25 to 13.5 mm. long; females 13.5 to 15.75 mm. long; vulva % of

body length from end of tail ; imperfectly known.

Amidostomum acutum, p. 24.
Males less than 9 mm. long, or (A. anseris) 10 to 17.1 mm. long 3.

3. Spicules 120m long ; 1 tooth in buccal capsule ; externo-dorsal ray originates

in common with dorsal ray Amidostomum chevreuxi, p. 22.

Spicules over 150m long; 3 teeth in buccal capsule and externo-dorsal ray
originates in common with other rays, not with dorsal, where known
(unknown for A. fulicae) 4.

4. Males 10 to 17 mm. long ; females 12 to 23.7 mm. long ; spicules 200 to 300m

long; female tail 437m long; buccal capsule with 3 teeth and 6 paired

ridges Amidostomum anseris, p. 20.

Males less than mm. long; females up to 14.5 mm. long; spicules not over
204m long; paired ridges not reported for buccal capsule 5.

5. Spicules 175m long; telamon 70m long Amidostomum fulicae, p. 25.

Spicules 166m or 204m long; telamon 90 to 100m long 6.

-8. Spicules 166^ long; telamon 90/* long; buccal capsule 27.5m wide in male, with

1 large and 2 very small teeth Amidostomum henryi, p. 22.

Spicules 204/x long; telamon 100m long; buccal capsule 15m wide in male,
with 1 large and 2 small teeth Amidostomum raillieti, p. 24.

AMIDOSTOMUM ANSERIS (Zeder, 1800) Railliet and Henry, 1909

Synonyms. — Ascaris mucronata Froelich, 1791, not Schrank, 1780;
Strongylus anseris Zeder, 1800 in part; Strongylus nodulosus Ru-
dolphi, 1803; Strongylus nodularis Rudolphi, 1809; Trichostrongylus
nodularis (Rudolphi, 1809) Shipley, 1909; Amidostomum nodu-
losum (Rudolphi, 1803) Seurat, 1918.

Hosts. — Anas querquedula, Anser acuta, Anser albifrons, Anser
anser, A. anser domestica, A. cinereus, A. clangula, A. crecca, A.
fabalis, A. fuUgula, A. fusca, A. leucops, A. marila, A. mollissima,
A. nigra, A. penelope, A. segetum, Chloeplxaga poliocephala, Fulica
atra, FuUgula cristata, F. marila, Cfatlvnula chloropus, Nyroca clan-
gula, N. fuUgula, N. marila, and Soniateria dresseri.

Location. — In or under the mucosa of the gizzard, proventriculus,
the connection between these two, the esophagus, and, according to
Skrjabin, the duodenum.

Morphology. — Amidostomum (p. 19) : Slender reddish worms.
Cuticle transversely striated, with a longitudinal pseudo-striation
due to the musculature. Head end (fig. 13) slightly enlarged and
bearing 2 pairs of large submedian papillae. Mouth aperture oval.
The short wide buccal capsule has 3 pointed teeth at its base and
has 6 ridges grouped in pairs. Esophagus has 3 chitinous ridges
along its entire length.

NEMATODE PARASITES OF BIRDS

21

Male 10 to IT mm. Ion*: by 250 to 350/x wide. Bursa (fig. 14) with
2 large lateral lobes and a small median lobe. A pair of lame con-
tinuous papillae on the posterior lip of the cloacal aperture. Ventro-
ventral ray small, its tip ending at an incision in the bursal margin.
Latero-ventral ray longer, enlarged proximally, its tip ending at an
incision in the bursal margin. Externo-lateral ray short, the tip not
near the bursal margin. Medio-lateral ray long, extending almost
to bursal margin. Externo-dorsal ray long and thick, originating
near the base of the common stem from which all rays in the lateral
lobes arise, and not extending to near the bursal margin. Dorsal ray
short, bifurcating posteriorly and the bifurcations forked and ter-
minating in 2 tips; this ray alone has an independent origin. The

Figs. 13-16. — 13, Amidostomum ansekis. Anterior end. 14, Same. Male
. bursa. Xos. 13-14 after Railliet, 1893. 15, Amidostomum raillieti. An-
terior end. 16, Same. Bursa op Male. Nos. 15-16 after Skkjabin, 1916

maroon-colored spicules are 200/x long (Skrjabin says 280 to 300//.)
and slender and are cleft near their middle; the internal branch ends
in a spatulate tip ; the gubernaculum is slender and 95/x long.

Female 12 to 24 mm. long. The body is slender anteriorly, widens
at the vulva to 300 or 400^, and thins abruptly behind the anus; the
tail is long and straight and bluntly pointed. The vulva is a trans-
verse slit, 160jm long, Vr, of the body length from the tip of the tail,
and is sometimes covered with a projecting appendix. The eggs are
thin-shelled, 85/x, long by 50/x wide, or, according to Skrjabin, 110/i
long by 82/a wide, (or 100/x, by G6ju in American material) and con-
tain an embryo when deposited.

Life history. — Unknown; probably simple and direct.

Distribution. — Europe, Asia, Africa (Algeria) and North America
(United States).

22

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

AMIDOSTOMUM HENRYI Skrjabin, 1915

Host. — Vanellus cristatus.

Location. — Gizzard.

Morphology. — Amidostomum (p. 19) : Buccal capsule (fig. 17)
very small, 15 to 18.5/* wide, with 1 large tooth and 2 very small,
scarcely perceptible, teeth.

Male 8 mm. long by 130 to 150/* wide. Buccal capsule 15/*
wide. Dorsal ray (fig. 18) has an origin separate from all other rays,
including the externo-dorsal ray, all these having origin in a large
common stem; the dorsal ray is the only one supporting the small
but distinct dorsal lobe. The externo-dorsal ray and postero-lateral
ray are short; all other rays in the lateral lobes extend to the bursal
margin. Spicules 166/* long, irregular in shape and each cleft dis-

Figs. 17-19. — 17, Amidostomum henryi. Anterior end. 18, Same. Male
bursa. 19, Same. Tail of female. All after Skrjabin, 1916

tally along almost half its length, judging from Skrjabin's figure.
Telamon (gorgeret) slender, 90/* long.

Female 14.5 mm. long by 180 to 187/* wide. Tail (fig. 19) elon-
gate conical, and, in Skrjabin's figure, slightly curved. Esophagus
850/* long. Buccal capsule 18.5/* wide. Vulva 2.72 mm. from
tail end. Eggs oval, 92.5 to 103.6/* by 70 to 80/*, arranged as a
rule perpendicular to the long axis of the body.

Life history. — Unknown; probably simple and direct.

Distribution. — Asia (Russian Turkestan).

AMIDOSTOMUM CHEVREUXI Seurat, 1918d

Synonym. — Amidostomum skrjahini Boulenger, 1926 (see Ad-
denda, p. 383).

Host. — Himantopus himantopus and (A. skrjahini) Anser albi-
frons.

Location. — Gizzard, under corneus lining.

NEMATODE PARASITES OF BIRDS

23

Morphology. — Amidostomum (p. 19) : Body slender, faintly red
with transverse striations. No lateral alae; postcervical papillae
very small, subsymmetrical ; excretory pore ventral, between nerve
ring and postcervical papillae. Buccal cavity (lig. 20) 7 to 8fi
deep and 10/i. wide, with very thick walls, and with a large tri-
angular dorsal tooth having a wide base and a sharp tip which turns
dorsad. There are 4 small sessile cephalic papillae. No pharynx.
The cylindrical esophagus is armed with 3 axial triturating lamellae
and is in relation posteriorly with an unarmed bulb of the same
width, not differentiated externally, which is without masticatory ap-
paratus. Nerve ring at middle of esophagus.

•Male 7.25 to 8 mm. long by 80^ wide, and terminating in an un-
cinate tail concave ventrally. Esophagus and bulb 63C/x long.

Figs. 20-23. — 20, Amidostomum chevkicuxi. Anteiuor end. a, Dorsal view ; b,

LATERAL VIEW. 21, SAME. MALE BURSA. 22, SAME. OUTLINE OF BURSA; LATERAL

view. Nos. 20-22 after Seurat, 1918. 23, Amidostomum monodon. Anterior
end. Aftf.b Linstow, 1882

Bursa (figs. 21 and 22) with 2 large lateral lobes, 105/* long, with
their free borders folded towards the ventral surface, and with a dor-
sal lobe which is not distinctly delimited. The externo-dorsal ray
originates from the dorsal stem, but its relations otherwise are with
the lateral lobes and the rays other than the dorsal ; it is short, as is
the externo-lateral ray; all other rays extend to the bursa margin ex-
cept the latero-ventral which extends almost to the bursa margin.
A pair of large sessile papillae, contiguous or almost so, on the poste-
rior margin of the cloacal aperture. Prebursal papillae subsymmet-
rical and briefly pedunculated. Spicules 120^ long, each cleft dis-
tally for half its length into 2 unequal branches. Telamon (gor-
geret) straight, falciform, G0/x long.

Female 12.8 mm. long by 120/x wide, terminating in an elon-
gate digitiform tail 265/u. long. Nerve ring 3/7 of esophagus
length from head. Esophagus and bulb 840/x long. Vulva a trans-
verse slit, 90/x long, slightly salient, in the posterior fourth of the
3613—27 4

24 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

body and 2.835 mm. from anus. Ovejector branches divergent; vesti-
bule 510/z long and containing only 3 eggs; trompe or varnish
gland (Seurat's "trompe " or "glande vernissante " might be termed
in English the glandular ovejector, making the parts of the ovejector
or the vestibule, glandular ovejector and muscular ovejector) 40/x
long; sphincter 105/* long. Uteri opposed; ovaries parallel. Eggs
80/* by 45/*, thick-shelled, and in blastula stage as oviposted.

Life history. — Unknown; probably simple and direct.

Distribution. — Africa ( Bone ) .

AMIDOSTOMUM RAILLIETI Skrjabin, 1915

Host. — Fulica atra and Fulica, species.

Location. — Cecum and " stomach " wall.

Morphology. — Amidostomum (p. 19) : White, cylindrical worms,
slightly attenuated anteriorly. Buccal capsule (fig. 15) with 1 large
tooth and 2 very small teeth disposed laterally.

Male 5.5 to 7.9 mm. long by 200 to 255/* wide. Buccal capsule 27.5/x
wide. Bursa (fig. 16) with 2 large lateral lobes and a small but dis-
tinct dorsal lobe. Externo-dorsal ray originates in common with all
of the rays except the dorsal and is very short ; the postero-lateral is
also very short. All rays except the dorsal are confined to the lateral
lobes and all except the postero-lateral and the externo-dorsal extend
to the bursa margin. Dorsal ray bifurcating near distal end, each
branch again dividing unequally. Prebursal papillae present; geni-
tal cone with a pair of large papillae. Spicules 204/* long, irregular
in shape and each cleft distally along most of its length. Telamon
slender, 100/* long.

Female 6.8 to 9.3 mm. long. Anus 110 to 145/* from posterior end
of body, the tail thus being considerably shorter than that of the
type-species. Vulva 1.3 to 1.8 mm. from tail end. Combined length
of ovejectors (including sphincters) 270 to 290/*. Eggs 90 to 105/x
by 50 to 65/*.

Life history. — Unknown; probably simple and direct.

Distribution. — Asia (Russian Turkestan) and Egypt.

AMIDOSTOMUM ACUTUM (Lundahl, 1848) Scurat, 1918d

Synonym. — Strongylus acutus Lundahl, 1848.

Hosts. — Anas crecca, Oidemia fusca (Anas fusca, Fuligula fusca),
Oidemia nigra (Anas nigra, Fuligula nigra), Somateria mollissima
(Anas mollissima, Fuligula mollissima), Nyroca fidigula (Anas
fuligula) and Fuligula cristata.

Location. — Gizzard.

Morphology. — Amidostomum (p. 19) : Mouth aperture simple and
spherical. Body red, of rather uniform diameter except for an at-

NEMATODE PARASITES OF BIRDS 25

tenuation anteriorly. Esophagus 1.125 to 1.2375 mm. long, widening
posteriorly and joining a much narrower intestine.

Male 10 to 14 mm. long by 225/* wide. Lateral lobes of bursa
large. Ray structures uncertain. Lundahl says the anterior are
small and directed anteriorly and Linstow says the anterior rays are
doubled; one may assume from Lundahl that he refers to an anterior
ray on each side, the ventro-ventral, and this interpretation would
accord with the generic pattern, as this ray is set off from the suc-
ceeding rays in other species of the genus. Linstow's statement
would give the impression that the 2 ventral rays on each side w T ere
close together and parallel, a thing which would not accord with the
assignment of this species to the genus Amidostomw/m. Lundahl
states that the 4 middle rays extend to the margin of the bursa ; in
other species the distinct shortening of the externo-lateral ray, which
does not extend to the bursa margin, is rather distinctive; Linstow
states that the median rays have a common broad stem, and in most
species, except for A. chevreuxi, all rays except the dorsal ray have
such an origin where this point is covered or illustrated. Linstow
states that the posterior rays are doubled; Lundahl that the posterior
ray is straight and thick ; one may assume that there is a dorsal ray
associated with small dorsal lobe if the species belongs in this genus,
but the exact facts do not appear to be covered in the literature.
Spicules and telamon (gorgeret) not described.

Female 14 to 17 mm. long. Tail elongate and pointed, directed
posteriorly and then outwards (dorsally ?). Vulva y 5 of body length
from tail end, strongly defined and limited anteriorly by a prominent
lip.

Life history. — Unknown; probably simple and direct.

Distribution. — Europe ( Sweden ) .

AMIDOSTOMUM FULICAE (Rudolphi, 1819) Seurat, 191Se

Synonyms. — Sj)in>i>l( ra fulicae Rudolphi, 1819.

Host. — Fulica atr<i.

Morphology. — Amidostomum, (p. 19) : Characters of the genus.

Male 8.58 mm. long by 155/* wide. Tail 100/* long. Buccal cavity
16/* wide. Esophagus 884/* long. Spicules 175ft long. Telamon
(gorgeret) 70/* long.

Female 9 mm. long by 175/u, wide. Buccal cavity 14/* wide. Esoph-
agus 972/* long. Vulva 1.56 mm. from end of tail. Eggs 105/* by
52/*.

This description is taken from Seurat's table without critical con-
sideration of the works of Creplin, Stossicli, and Mueller, on which
Seurat bases it. Little information is available from Rudolphi or
Dujardin.

26 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

AMIDOSTOMUM MONODON (Linstow, 1882) Skrjabin, 1915

Synonyms. — Strongylus monodon Linstow, 1882 ; Sclerostomy
monodon (Linstow, 1882) Stossich, 1889.

Host. — Oldemia nigra.

Location. — Gizzard.

Morphology. — Amidostomum (p. 19) : Mouth cavity (fig. 23)
armed with a large triangular tooth. Wall of cavity without sup-
porting ribs.

Male unknown.

Female 16.5 mm. long by 14G> wide, the ratio of width to length
being 1 : 116. Esophagus 1/23, tail 1/55 of total body length, the
tail conical and rounded. Vulva at posterior fifth of body, dividing
body length in ratio of 17 : 4. Eggs 92/* long by 62/x wide.

Life history. — Unknown; probably simple and direct.

Distribution. — Europe.

Genus EPOMIDIOSTOMUM Skrjabin, 1916

Generic diagnosis. — Amidostominae (p. 18) : Meromyarian. Body
filiform, the anterior extremity attenuate. Cuticle thick and trans-
versely striated. Postcervical papillae present, but not prominent.
Head distinct, bearing on its dorsal and ventral surface a pair of
nodules (also referred to as lips or papillae) which are directed
posteriorly and are either uncinate or obtuse at their free extremity.
According to Seurat the head bears a pair of lateral papillae on each
side. Cephalic cuticle ornamented with a pair of epaulets or fes-
toons, which, according to Skrjabin, have zig-zag incisions in their
posterior portion. Buccal capsule lacking, according to Skrjabin, or
short and reduced, according to Seurat. Esophagus with 3 chitinous
axial lamellae. Male with uncinate tail, concave ventrally, with a
bursa which, according to Skrjabin, is delicate or, according to
Seurat, has thick lateral lobes marked with a strong reticulate stria-
tion and folded over each other ventrally. The ventro-ventral and
latero-ventral rays are in relation with each other as opposed to the
other bursal rays ; Seurat says they are parallel, but Skrjabin says they
are separated and figures them as slightly divergent. According to
Seurat, the postero-lateral and medio-lateral rays are slender, contig-
uous, and elongate, and the externo-lateral ray is short ; according to
Skrjabin, the externo-lateral ray is related to the 2 other lateral rays,
which are so united as to leave only a slight cleft between them ; Skrja-
bin's figure indicates that the lateral rays have a common stem which
divides to form the externo-lateral ray and the common stem of the
2 other laterals, and that this latter common stem divides a half to
two-thirds of its length from its base to form these 2 rays. The

NEMATODE PARASITES OF BIRDS 27

externo-dorsal ray is short and massive, originates directly at the
tip of the tail, and does not reach the bursal margin. The dorsal
ray, in the type-species, divides dichotomously distally to form 4
small tips and is the only ray in relation with the small dorsal
lobe of the bursa. There are 2 voluminous sessile papillae, contigu-
ous or almost so, on the posterior lip of the cloaca. A pair of briefly
pedunculated prebursal papillae present. Spicules equal and short,
each dividing distally, in the type-species, to form 1 dorsal and 2
latero-ventral branches. No telamon (or gorgeret) present. Female
with vulva posterior to middle of body. Branches of ovejector
divergent or parallel. Uteri opposed. Eggs segmenting when
deposited. Parasitic in corneus tunic of gizzard in palmipeds.

Type species. — Epomidiostomum. iineinatwm (Lundahl, 1848)
Seurat, 1918 (=E. anatinum Skrjabin, 1916).

KEY TO SPECIES OF EPOMIDIOSTOMUM

1. Males 6.03 to 7.13 mm. long; females 10 to 11.5 mm. long. Spicules 120 to

130^ long Epomidiostomum uncinatum, p. 27.

Males 8.8 mm., or more, in length ; females 13 mm., or more, in length.
Spicules 180 to 210/* long 2.

2. Spicules dividing distally into 2 branches.

Epomidiostomum querquetulae (Addenda, p. 3S4).
Spicules dividing distally into 3 branches 3.

3. Head, in addition to lateral festoons, has 8 processes (4 lateral papillae and

4 (2 dorsal, 2 ventral) posteriorly directed hooklike structures).

Epomidiostomum orispinum, p. 28.
Head, in addition to lateral festoons, has total of 12 sharply pointed processes
(4 anteriorly directed at mouth opening; external to these, 4 horizontally
directed; and posterior to these, 4 (2 dorsal, 2 ventral) anteriorly di-
rected) Epomidiostomum skrjabini (Addenda, p. 3S4).

EPOMIDIOSTOMUM UNCINATUM (Lundahl, 1848) Seurat, 1918e

Synonyms. — Strongylus uncinatus Lundahl, 1848; Epomidiosto-
mum anatinum Skrjabin, 1916.

Hosts. — Anas acuta, A. nigra, A. penelope, A. boschas domestica,
Anser albifrons, Fuligula nigra, Mareca penelope.

Location. — Under the cuticular layer of the gizzard.

Morphology. — Epomidiostomum (p. 26) : Slender worms, with
very narrow head ends. No buccal capsule; mouth with papillae
(lips?) directed outward. Cuticle of head end (fig. 24) with 2
epaulette-like ornaments, each terminating posteriorly in a free
border with 3 tooth-like structures.

Male 6.3 to 7.13 mm. long by 15G> wide, yellowish. Cuticle trans-
versely striated. Esophagus cylindrical, 800/t long. Bursa (fig. 25)
with 2 large lateral lobes and a small dorsal lobe. Ventral rays
divergent; externo-lateral ray divergent from the latero-ventral ray

28

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

and almost parallel to the main trunk which bifurcates to form the
medio-lateral and postero-lateral rays; the externo-dorsal rays arise
at the base of the dorsal ray and are short, with a swelling project-
ing near their union with the dorsal ray; the dorsal ray bifurcates,
each branch in turn apparently bifurcating at its tip. There are a
pair of large caudal papillae near the base of the dorsal ray. Two
equal spicules, brown, 120 to 130/* long, divide to form 3 terminations
posteriorly. No gubernaculum.

Female 10 to 11.5 mm. long by 250// wide. Tail (fig. 26) forms a
conical appendix 140 to 170//, long, with a button at its apex. The
vagina is a transverse cleft, 110/* long, situated 2.2 mm. from the tip
of the tail. Eggs elliptical, 74 to 80/* long by 48 to 50/* wide.

Life history. — Unknown.

Figs. 24-27. — 24, Epomidiostomum uncinatum. Anterior end. 25, Same. Male
bursa. 26, Same. Female tail. Nos. 24-26 after Skrjabin, 1916. 27, Epo-
midiostomum orispinum. male bursa. after molin, 1861

Distribution. — Europe (Sweden), Central Asia (Province of Syr-
Daria), and Africa (Algeria (Ain-Mokra) ).

EPOMIDIOSTOMUM ORISPINUM (Molin. 1861) Seurat, 1918

Synonyms. — Strongylus anseris Zeder, 1800, in part; Strongylus
orispinus Molin, 1861.

Hosts. — Anas albifrons, A. anser domestica, A. anser fera, A.
clangula, A. crecca, A. fuligula, A. fusca, A. leucops, A. mollissima,
A. nigra, A. penelope, A. segetum, Anser anser, Fulica atra.

Location. — Under the mucosa of the esophagus and proventriculus.

Morphology. — Epomidiostomum (p. 26) : Body robust, slender in
head region, transversely striated. Head distinct, with a pair of
large sessile lateral papillae and festoons on each side, and with 2
dorsal and 2 ventral hook-like nodules (lips or papillae), directed
posteriorly.

NEMATODE PARASITES OF BIRDS 29

Male 10.8 mm. long by 210/x, wide. Bursa (fig. 27) with a strongly
reticulate striation and with its free edge folded over the ventral
surface. Rays somewhat similar to those of Amidostoinum anseris
(p. 21), and similar papillae on the posterior edge of the cloacal
aperture. The spicules are 200,u. long, divided proximally into 3
branches, of which the dorsal is longitudinally striated.

Female 16.7 mm. long by 27;")^ wide. Body reddish in esophageal
region and white elsewhere, abruptly truncated on the ventral sur-
face at the level of the anus and terminating in a short, digitiform
tail. Vulva transverse, 140//, long, with salient lips, i/5 of the body
length from the tail end. Eggs ovoid, thick-shelled, 95/* long by
55//, wide, segmenting when deposited.

Life history. — Unknown; probably simple and direct.

Distribution. — Europe and Africa (Algeria).

Superfamily STRONGYLOIDEA Weinlancl, 1858

Superfamily diagnosis. — Strongylata (p. 5) : Meromyarian.
Buccal capsule present. Bursa usually of typical strongyle struc-
ture, with a dorsal ray or 2 dorsal rays of approximately the same
width as the 6 other rays on each side of the bursa and not excessively
wider than these. Male with 2 spicules and female with 2 ovaries.
Vulva usually posterior to middle of body; occasionally anterior
{Syngamvs). Oviparous, the eggs segmenting when oviposited.
Embryo and first-stage larva rhabditiform. Usually thick and
rather stiff worms. Usually in digestive tract, sometimes in respira-
tory tract {Syngamus) or tissues (Stephanurus) , as adults; in circu-
latory system or tissues as agamic individuals {Stiongylus).
Development, so far as known, direct and without intermediate host,
but with wandering of larvae through body (Strongylus, Ancylos-
toma, etc.) or at least into tissues adjoining digestive tract {(Eso-
phagostomum) in a number of known cases.

Type family. — Strongylidae Baird, 1853.

The name Strongyloidea was originally proposed as a family
name by Weinland and was first used as a superfamily name by
Hall (1013). The superfamily diagnosis given by Hall (1016) is
in this paper substantially that of the suborder Strongylata, and
the above diagnosis for the superfamily is substantially that of the
family Strongylidae as recognized of recent years and up to the
present time.

KEY TO THE FAMILIES OF STRONGYLOIDEA

1. Vulva in posterior half of the body. Parasites of digestive tract.

Strongylidae, p. 30.

Vulva in anterior half of the body or rarely median to slightly posterior.

Parasites of respiratory tract Syngamidae, p. 33.

30 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Family STRONGYLIDAE Baird, 1853

Family diagnosis. — Strongyloidea (p. 29) : Vulvo in posterior half
of the bodj 7 . Corona radiata usually present. Spicules well devel-
oped. Bursa well developed. Parasites of the digestive tract as
mature adults.

Type genus. — Strongylus Mueller, 1780.

Subfamily Strongylinae Railliet, 1893

Subfacility diagnosis. — Strongylidae (p. 30) : Forms usually with
a corona radiata. Head not bent dorsally. Intestine straight, not
greatly convoluted. Bursa with rays long, not reduced.

Type genus. — Strongylus Mueller, 1780.

Tribe DELETROCEPHALEAE Railliet and Henry, 1911, emend. Stiles

and Hassall, 1920

Synonym. — Deletrocephalae Railliet and Henry, 1911.

Tribe diagnosis. — Strongylinae (p. 30) : Ventro-ventral and latero-
ventral rays close together and parallel, originating from a common
stem. Lateral rays originate in a common stem and are typically
divergent. Externo-dorsal ray originates at the base of the dorsal
ray. Main dorsal ray bifurcates and each branch subdivides in such
a way as to have 3 major and minor terminations ultimately. The
vulva is close to the anus and the ovejectors and uteri are con-
vergent.

Type genus. — Deletrocephalm Diesing, 1851.

This tribe is almost too close to the Cylicostomeae to make its
tribal characters impressive. The externo-dorsal ray originates at
the very base of the dorsal ray, instead of with the lateral rays,
but the two conditions approach each other so closely as to leave
some doubt as to whether the distinction is of value. Other features
are so similar as to make it appear that the point of origin of the
externo-dorsal ray in the forms involved can hardly be regarded as
of more than generic value.

KEY TO GENERA OF DELETHOCEPHALEAE

1. Oorona radiata present. Buccal capsule without teeth or longitudinal sus-
taining ribs or rays Codiostomum, p. 30.

Corona radiata absent. Buccal capsule with longitudinal sustaining ribs or
rays and usually with teeth Deletrocephalus, p. 32.

Genus CODIOSTOMUM Railliet and Henry, 1911

Generic diagnosis. — Deletrocephaleae (p. 30) : Strongyles with in-
ner and outer leaf crown and well developed dorsal gutter. Bursa

NEMATODE PARASITES OF BIRDS

31

with characters of the tribe and the following specific characters:
Dorsal lobe very long and attached perpendicular to body ; the com-
mon ventral stem unites with the common lateral stem to form a
large stem mass; the externo-dorsal ray originates at the base of
the dorsal ray near its union with this stem mass; the postero-lat-
eral ray comes off first from the lateral stem and diverges farther
from the medio-lateral than does the externo-lateral ; the medio-lat-
eral is the longest of the 3 rays. The main branches of the dorsal
ray give off external branches which again divide to form 2 small
branches. Genital cone large. Spicules equal. Gubernaculum or
telamon present. Vulva and anus close together. Ovejectors con-
vergent.

Type species. — C odiostomum struthionis (Horst, 1885) Railliet and
Henry, 1911.

Figs. 28-30. — Codiostomum steuthionis. 28, Head end. 29, Buccal cadsule.
30, Male bursa., spread out. After Horst, 1885

CODIOSTOMUM STRUTHIONIS (Horst, 1885) Railliet and Henry, 1911

Synonym. — Sclerostomy, struthionis Horst, 1885.

Hosts. — Struthio molybdophanes, S. camelus, S. aus traits.

Location. — Ceca, large intestine, and stomach.

Specific diagnosis. — Codiostomum (p. 30) : White or yellow, rather
stiff worms. Buccal capsule (fig. 29) 28G> deep by 100> wide at
oral aperture, and with well developed dorsal gutter. Submedian
and lateral papillae present, Head (figs. 28 and 29) with inner and
outer leaf crown of many elements, more in outer than inner, judging
from Monnig's figure, the outer long and slender, the inner shorter.

Male 13 to 17 mm. long by 420 to 450^, wide. Esophagus length
1/12 of body length. Bursa (figs. 30 and 31) characters those of the
genus. Prebursal papillae present. Spicules equal, about 870/x
to 1 mm. long, alate, transversely striated, curved, and widened in
the proximal third of their length. Gubernaculuiu boat-shaped,

32

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

84,u long, according to Monnig; Horst figures what is evidently a
telamon partly encircling the cloaca near its aperture. Testis closely
wound about anterior portion of intestine.

Female 17 to 23 mm, long by 560 to 700/* wide. Esophagus length
14 of body length. Lateral cervical papillae 960/* from anterior end,
excretory pore 800/*, nerve ring 660/*. Vulva (fig. 32) 250/* anterior
to anus. Anus 572/* from tip of tail; tail ends in a mucronate tip
128/* long. Cuticula around vulva often inflated and may form
a considerable protrusion. Vagina 120/x long. Ovejectors parallel,
the cuticular portions 620/* long and the musculo-epithelial portions
1.5 mm. long. Eggs oval, 63/* by 35/x.

Life history. — Unknown; probably simple and direct.

D istribution. — Africa.

Figs. 31-32. — Codiostomum struthionis. 31, Male bursa, lateral view. 32.
Female tail. After Monnig, 1923

Genus DELETROCEPHALUS Diesing, 1851

Generic diagnosis. — Deletrocephaleae (p. 30) : No corona radiata
present, the mouth aperture bounded by 6 equal or subequal papillate
lips. Buccal capsule provided with meridial ribs or rays and with
teeth in some species. Bursa with characters of the tribe ; one branch
of the dorsal ray sometimes very slender. Spicules slender and equal.
Vulva and anus close together. Ovejectors parallel.

Type species. — Deletrocephalus dimidiatus Diesing, 1851.

DELETROCEPHALUS DIMIDIATUS Diesing, 1851

Synonym. — Sclerostomy dimidiatum (Diesing, 1851) Stossich
1899.

Host. — Rhea americana.

Location. — Ceca and large intestine, especially at union with small
intestine.

Morphology. — Deletrocephalus (p. 32) : Head (fig. 33) compressed
lnterally, the elliptical buccal aperture with its long axis dorso-ven-

NEMATODE PARASITES OF BIRDS

33

tral and closed by 6 striated membranous expansions or lips. Buccal
capsule well developed, with C meridial ribs in its wall and with a
dozen small teeth at its base, two of these, somewhat larger than the
others, at the base of the dorsal gutter.

Male 11 to 18 mm. long. Branches of dorsal ray of bursa (fig. 34)
tridigitate, one of the subordinate branches often very slender.
Spicules slender, 900/a to 1.025 mm. long.

Female 17 to 21 mm. long. Vulva (fig. 35) close to the anus and
often covered by a cuticnlar expansion. Eggs thin-shelled, 120 to
125//. by TO to 75/t, segmenting when deposited.

Life history. — Unknown; probably simple and direct.

Distribution. — South America (Brazil) and Europe (in museum
material).

Figs. 33-35. — DelhtrcxXephamjs dimidiates. 33, Head ; a, lateral; b, front
view. 34, Male BURSA. 35, Female tail. After Diesing, 1857

Family SYNGAMIDAE Leiper, 1912

Family diagnosis. — Strongyloidea (p. 29) : Worms permanently
joined in copula (Syngamws) or not permanently joined (Cyathos-
toma). Buccal capsule large, thick-walled and armed at the base
with 6 to 9 teeth of 2 distinct sizes arranged about a center. Bursa
membrane thick or of thickness usual in strongyles in other families;
rays often short and thick, sometimes slender. Spicules short (Synga-
mus) or long (Cyathostoma). Vulva usually in anterior half of
body, rarely median or slightly postme.dian. Eggs operculated after
deposition.

Type genus. — Syngamus von Siebold, 1836.

KEV TO GENERA OF SYNGAMIDAE

1. Worms permanently joined In copula. Buccal capsule usually with 8 or 9
teeth rarely 3 (error?). Bursa rays short and thick. Spicules small.

150^ or less. Vulva in anterior half of body Syngamus, p. 34.

Worms not permanently joined In copula. Buccal capsule with (5 or 7 teeth
rarely 2 (error'.'). Bursa rays slender. Spicules Long, over 40Qp. Vulva
anterior, median or slightly posterior Cyathostoma, p. 41.

34 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Genus SYNGAMUS von Siebold, 1836

Generic diagnosis. — Syngamidae (p. 33) : Sexes permanently
joined in copula. Buccal capsule in both sexes large, thick- walled,
usually provided with 8 or 9 teeth, rarely 3, arranged about the
center at the base, the teeth of 2 distinct sizes. Male bursal mem-
brane thick; bursal rays short and thick; spicules small to very
small, 150/* to 25/*. Vulva in anterior third of body; tip of female
tail blunt or acute. Eggs of moderate size, operculated after deposi-
tion.

Type species. — Syngamus trachea (Montagu, 1811) Chapin, 1925.

KEY TO SPECIES OF SYNGAMUS

1. Each branch of dorsal ray bifurcated ; spicules 115 to 150/x long.

Syngamus microspiculum, p. 38.

Each branch of dorsal ray trifurcated or simple ; spicules less than 100/*

long 2.

2. Each branch of dorsal ray trifurcate ; spicules 60*i long.

Syngamus trachea, p. 34.
Each branch of dorsal ray simple, not subdivided 3.

3. Spicules subequal, 50/* long. Parasites of Nucifraga.

Syngamus parvus, p. 39.

Spicules distinctly unequal, the right 79ii, the left 69/t* long. Parasites of

Corvus Syngamus gracilis, p. 39.

The following species are nomina nuda and are omitted from the
above key; Syngamus coelebs from Falco lag opus, S. mucronatus
from Picus canis and P. major, S. pugionatus from Corvus pica and
Sturnus vulgaris.

This key as well as the one to species of Cyathostoma (p. 41) are
modifications of keys from Chapin, 1925.

SYNGAMUS TRACHEA (Montagu, 1811) Chapin, 1925

Synonyms. — Fasciola trachea Montagu, 1811 ; Syngamus trachealis
Siebold, 1836 ; Syngamus primitives Molin, 1860 ; Strongylus primi-
tivus (Molin, 1860) Hutyra and Marek, 1910; S. trachealis (Siebold,
1836) Nathusius, 1837; Sclerostomy tracheale (Siebold, 1936) Dies-
ing, 1851. It is regretted that the well-established name, Syngamus
trachealis, should be dropped into synonomy, but as Montagu's name
has 25 years' priority, it must be regarded as the correct name, as
noted b/y Chapin.

Hosts. — Meleagris gallopavo and Gallus gallus. Also reported
from Anas boschas, Anser anser, Ciconia alba, C. nigra, Corvus
comix, C. corone, C. frugilegus, C. monedula, Cypselus apus, "jay,"
Lagopus scoticus, Otis tarda, Pavo cristatus, Pelecanus onocrotalus,
Perdix cinerea, Phasianus colchicus, Phasianus gallus, Phasianus
pictus, P. reevesi, Pica caudata, Pica pica, Picus canus, P. vh^idis,

NEMATODE PARASITES OF BIRDS

35

Pyrrocorax alpimts, Strix noctua, Sturnus vulgaris, Tetrao urog al-
ius, "thrush." Chapin notes that the specimens from birds other
than the Galliformes need further study; those from Corvus, spp.
may be S. gracilis.

Location. — In the trachea and bronchi as adults; in the lungs as
larvae; immature worms have been found in the peritracheal tissue
and air sacs (Megnin).

Morphology. — Syngamus (p. 34) : Cylindrical red worms, the
color more pronounced in the female. Head (fig. 37) enlarged and

Fig. 36. — Stngamds trachea. Outline. After Chapin, 1025

truncated. Mouth orbicular, with a hemispherical chitinous capsule,
at the base of which are usually 8, occasionally 9, sharp teeth; the
mouth is surrounded by a chitinous plate, the outer margin of which
is incised to form 6 festoons opposite each other, with 4 smaller fes-
toons between them in opposed pairs; a lateral papilla is between
each pair of small festoons, and 4 submedian papillae are present, 1
at each end of the large festoons. The male is permanently at-
tached in copula to the female, forming a Y (fig. 36).

FlG. P»7. — SYNGAM0S TRACHEA, HEAD. (I. TKETII IN 8-TOOTHED (NORMAL) FORM \
b, FRONT VIEW OF HEAD; c, TEETH IN 9-TOOTHEI> FORM. AFTER CHAPIN, 1925

Male 2 to mm. long by 20G> wide. The bursa (fig. 39) is ob-
liquely truncated, provided with rays somewhat similar to those of
C. bronchialis (p. 42), sometimes with strikingly asymmetrical dor-
sal rays. The male is permanently attached to the female about the
vulva. Two equal, slender, short spicules (fig. 38), 57 to G4/x long
according to Chapin, who states that the reports of 140/x spicules in
this species probably refer to another species.

Female 5 to 20 mm. long (longer in the turkey) by 350/* wide,
somewhat more slender anteriorly and irregularly swollen when

36 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

filled with eggs, the uterine coils extending almost to posterior end
of body (x in fig. 36). The conical tail end bears a pointed process.
The prominent vulva in immature worms is about one-fourth, in
gravid worms about one-sixth, of the body length from the anterior
end. The ellipsoidal eggs are operculated and 85 to 90/a long by 5(V
wide, or, according to Ortlepp, 78 to 100^ long by 43 to 46/*, wide.

Embryo 280/* long; anterior end blunt, tail elongate conical and
pointed. The embryo in the shell may be the first or second stage
larva. The description applied to the second stage larva.

Larva, third stage, in lungs, of 2 types. Male with obliquely trun-
cated tail and up to 1.16 mm. long. Female with pointed tail and
up to 1.42 mm. long. Simple buccal capsule 9/t deep.

Larva, fourth stage, separate or
copulating. Male up to 1.44 mm.
long. Female up to 1.98 mm.
long. Most of adult characters
present.

Life history. — As the fully formed
eggs develop, they pass out of the
~^r- vulva under the margin of the male

figs. 38-39.— syngamus trachea. 38. bursa and get to the lumen of the
spicules. 30, male bursa. After trachea and bronchi. These eggs,

Chapin, 1925 ... ,. , -, 5; t

which are segmenting when deposited,
up to the 16-cell stage, are coughed up and swallowed, passing out
in the droppings. In the external world they develop infective
larvae under favorable conditions, as in well aerated water, in
about a week, and hatch in about 2 weeks (some writers say 1
week) or longer if conditions are less favorable. Eggs hatch at
25° C, according to Ortlepp, but not at 20° C. The first molt may
take place in the shell or after hatching, and the larvae appear to
be infective as second-stage larvae. These larvae sometimes lose
their sheaths after hatching. The fact that birds swallow a large
amount of mineral matter, to furnish sand, gravel and small peb-
bles for the gizzard, ensures their ingesting the eggs containing lar-
vae or the larval worms present in soil, and it is not necessary to
assume that the larvae are ingested on vegetation or in food or
water, though this too may happen. When infective larvae are
swallowed, they make their way to the lungs where they have been
found by Ortlepp 24 hours after feeding eggs. Within a week they
are found there as young worms not yet paired. The third larval
stage is reached during the third day of parasitic life, and the fourth
stage on the fourth or fifth day. After pairing in the lungs while
still fourth-stage larvae, the young worms leave the smaller air
passages, gradually working their way to the larger air passages

NEMATODE PARASITES OF BIRDS 37

and attaining their full growth, both the male and female attaching
to the mucous membrane. Sexual maturity is reached 10 to 14 days
after the worms reach the trachea. Females attain a length of 15
nun. and begin depositing eggs as early as 2 weeks after infective
larvae are swallowed. Ransom has reported both mature worms
in the trachea and young worms in the lungs 2 weeks after feeding
infective material. Ortlepp finds eggs in the feces IT to 20 days
after infection.

A matter of great importance from an economic standpoint is
the fact that in chickens, as a rule, only young chicks can be in-
fested or do become infested with gapeworms. Post-mortem ex-
amination of hundreds of mature chickens have been almost always
negative, and attempts to infest such birds are likewise failures, or
at least to the extent that the worms seldom become mature and
usually die soon if they mature; commonly they either fail to de-
velop or undergo only partial development. (Ransom has reported
1 well-developed pair and 1 immature pair in a chicken at least 2
years old). The worms can only undergo complete development
and remain for any length of time in chickens during the first few
weeks of life, as a general rule. Ransom has noted that when ma-
ture chickens were successfully infected with gapeworms, the worms
were often buried in a mass of mucus, the males were dead in many
cases, and some of the females were apparently nonfertile and un-
natural in appearance.

On the other hand, gapeworms occur in turkeys, and turkeys be-
come infested at any time of life, so far as experiments and ob-
servations show. As a result, adult turkeys, which are but little
affected by the presence of gapeworms, serve as carriers of infec-
tion and young chickens on premises so infected become victims
of gapeworm disease. It has been noted in many cases that this,
association of turkeys and chickens has been a prominent feature
of gapeworm disease of chickens. Apparent!} 7 , the turkey is the
normal host of the gapeworm, the disease having been observed in
this bird, a native American animal, and in chickens at Baltimore in
1799. In 1800, 1807, and 1809, it was observed in chickens in Eng-
land, the turkeys and ducks associated with these chickens not show-
ing the disease. The turkey, being the normal host, evidently has
a normal immunity to the bad effects of the worm, although nor-
mally susceptible to infection at any age. On the other hand,
chickens, not the normal host, are immune to infection as adults, but
are susceptible to infection as chicks and at the same time devoid
of immunity to the bad effects.

Railliet has regarded the magpie {Pica pica) as a carrier of im-
portance in France, and it may be that a number of wild birds will

38

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

be found of importance in this connection. However, wild birds can
hardly play the part that domesticated birds can in carrying and
maintaining infection, and the turkey must be looked on as especially
dangerous in this connection.

Distribution. — More or less cosmopolitan (North America, South
America, Africa, Europe, Asia (Formosa), and Australia).

SYNGAMUS MICROSPICULUM Skrjabin, 1915a

Host. — Phalacrocorax carbo.

Location. — Trachea.

Morphology. — Syngamus (p. 34) : Base of buccal capsule (fig.
405 ) armed with 3 small triangular teeth.

Male 3.9 to 4 mm. long by 270 to 290//, wide, brown in color. Buccal
capsule 170/x deep by 220/* wide ; its basal teeth are 70/* long. Esoph-
agus 580,u long and is widened posteriorly. Spicules 150/* long, ac-
cording to Skrjabin (1915), or 115/* long, according to Skrjabin

Fig. 40.

-Syngamus microspiculum. a, Male and female; b, head end; c, male
bursa. After Skrjaben, 1910

(1916). Dorsal ray (fig. 40c) divides, apparently half its length
from the base, and each branch bifurcates distally. The ventral rays
are close together and parallel. The laterals are close together and
parallel. There is a distinct separation between the bases of the
ventral group, the lateral group, and the externo-dorsal ray, and a
wider separation between the base of the externo-dorsal and the
dorsal ray, according to Skrjabin's figure.

Female 11 mm. long by 420 to 500/*, attenuating to 250/* at the anus.
Tail pointed and curved, according to Skrjabin's figure. Buccal cap-
sule 250/* deep by 340/* wide. Esophagus 765/* long, widened pos-
teriorly. Vulva 4.45 mm. from head end. Eggs 75/* by 48/*, of char-
acteristic shape, flattened on one side.

Life history. — Unknown; probably similar to that of 8. trachea
(p. 36).

Distribution. — Asia (Russian Turkestan).

NEMATODE PARASITES OF BIRDS

39

SYNGAMUS PARVUS Chapin, 1925

Host. — Nucifraga caryocatactes.

Location. — Trachea.

Morphology. — Syngamus (p. 34) : Similar in general shape (fig.
41) to S. trachea, but much smaller and differing in secondary sexual
characters of male.

Male about 2.4 mm. long by 220/x, wide, cylindrical, with neck
region slightly constricted. Buccal capsule heavily chitinized, 178/x
in depth, 207^ in its greatest inside diameter; walls 29/*. thick. Buc-
cal teeth and circumoral papillae as in S. trachea. Esophagus about
326/x long; nerve ring and excretory pore not located. Bursa (fig.
43) 350/u. in diameter; ventral rays short, stout, and approximate;
lateral rays stout; medio-lateral ray very stout; externo-lateral ray
arising from the side of the medio-lateral; postero-lateral ray
slender in comparison with the other two. Externo-dorsal ray more

Pigs. 41-43. — Syngamus parvus. 41, Outline of pair. 42, Spic-
ules. 43, Dorsal portion of male bursa. After Chapin, 1925

slender and parallel than postero-lateral; dorsal trunk bifurcated
near tip, each bifurcation simple. Spicules (fig. 42) short, about
49/x, similar in shape to those of S. trachea.

Female about 7.8 mm. long by 65Gyt maximum width; width just
posterior to head 350^. Buccal capsule 300^ in depth, 440^ in aver-
age diameter. Buccal teeth as in S. trachea; nerve ring and ex-
cretory pore not located. Esophagus short and thick. Vulva in
gravid worm dividing total length into ratio as 1:5.2; uterine coils
extending posteriorly to 1.25 mm. from extremity (x in fig. 41) ;
anus subterminal; tip of tail very blunt. Eggs 74^ by 44^, not yet
segmenting in uterus.

Life history. — Unknown; probably similar to that of S. trachea
(p. 36).

Distribution. — Not given. (Host occurs in Europe.)

SYNGAMUS GRACILIS Chapin. 1925

Synonym. — Syngamus trachealis Weidman (part), Fox, 1923.
Host. — Corvus brachyrhynchos.
Location. — Trachea.

40

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Syngamus (p. 34) : Superficially (fig. 44) resem-
bling S. trachea but smaller and more slender.

Male 3 to 3.3 mm. long by 270/* wide. Buccal capsule 165/* deep
by 200/* wide near the anterior edge; wall 45/* thick. Buccal teeth
as in JS. trachea, the largest 45/* high. Esophagus nearly 600/* long,
clavate, its greatest diameter 135/*. Nerve ring, excretory pore, and
cervical papillae not seen. Bursa (fig. 45) narrow an\l deep. As
in JS. parvus, the dorsal ray branched, the branches simple ; however,
whereas in JS. parvus the bifurcation is only near the tip, in the
present species the bifurcation is basal, resulting in a total suppres-
sion of the common dorsal trunk, the branches being 150/* long.
Externo-dorsal ray slightly shorter, measuring 132/*. Lateral rays
mutually contiguous, parallel, and about the size of either branch

WOju

Figs. 44-46. — Syngamus gracilis. 44, Male and antekiok
portion of female. 45, male- bursa. 46, female tail.
After Chapin, 1925

of the dorsal ; ventral rays smaller and more slender. Spicules dis-
tinctly unequal; the right, 79/* long, is bent, the left, G9/* long, is
nearly straight.

Female 8 to 11 mm. long and proportionately stouter than the
male. Buccal capsule relatively enormous, its internal dimensions
525/* wide by 300/* deep ; wall 50/* thick. Teeth as in JS. trachea.
Esophagus 825/* long, attaining its greatest diameter, 225/t, near its
posterior end. Vulva 1.4 mm. from anterior end, or at about the
anterior eighth of body length. Vagina longer than in related
species; uteri parallel, posteriorly directed. Anus (fig. 46) about
300/* from posterior end, which is very blunt. Eggs 79 to 83/* by
40 to 46/*.

Life history. — Unknown; probably similar to that of S. trachea
(p. 36).

Distribution. — North America (Philadelphia Zoological Park).

Chapin has noted that in all probability it is this species that has
formerly been found in the American crow and reported as JS.

NEMATODE PARASITES OF BIRDS

41

trachea, and that the assumption that the crow may act as a reservoir
for the turkey and chicken gapeworm is probably not well founded.

Genus CYATHOSTOMA E. Blanchard, 1849

Generic diagnosis — Syngamidae (p. 33): Sexes not permanently
joined in copula. Buccal capsule in both sexes large, thick-walled,
provided with 6 or 7 teeth arranged about the center at the base, the
teeth of 2 distinct sizes. Male bursal membrane of usual thickness
for strongyles in general; bursal rays slender and sometimes
branched; spicules long (over 400/x) and filiform. Vulva anterior,

Figs. 47-48. — Cyathostoma bronchialis. 47, Head end. a, Side view ; b, front
view. 48, Male bursa. From Chapin, 1925, after Muehlig, 1884

median, or slightly posterior to equator of body; tip of female tail
acute. Eggs of moderate size, operculated after deposition.
Type species. — Cyathostoma lari E. Blanchard, 1849.

KEY TO SPECIES OF CYATHOSTOMA

1. Vulva at or near the anterior third of the body 2.

Vulva near the middle of tin- body 5-

2. Spicules 500m long; eggs 56/t long; in Tadorna tadoma.

Cyathostoma tadornae, p. 42.
Spicules 600/* long or longer, where known : eggs 80/t long or longer 3.

3. Eggs 80/* by 40/* or 92^ by 53/*; spicules 600/u long (Skrjabin) ; in ciconiform

birds Cyathostoma variegatum, p. 44.

Eggs SO to 90/x long by 55 to G0/z wide or wider ; spicules 600 to 650/* h>ng__ 4.

4. Eggs 90/* by 60//. ; spicules 600/* long; in nnseriforni birds.

Cyathostoma bronchialis, p. 42.
Eggs SO/* by 55/* j spicules G50/* long; in Casuarius galeatus.

Cyathostoma boularti, p. 44.

42 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

5. Male unknown ; female small, up to 13 mm. long, in orbital cavity of Larus

ridibundus Cyathostoma lari, p. 43.

Male known ; female up to 20 mm. long or longer 6.

6. Spicules 490// long; gubernaculum 67//. long; female up to 30 mm. long; in

thoracic air sac of Buteo borealis Cyathostoma americanum, p. 45.

Spicules 660 to 720//. long ; gubernaculum 92/t long ; female up to about 20 mm.
long; in trachea of Coscoroba coscoroba Cyathostoma coscorobae, p. 47.

CYATHOSTOMA BRONCHIALIS (Muchlig. 1884) Chapin, 1925

Synonym. — Syngamus bronchialis Muehlig, 1884.

Hosts. — Anas boschas domestica, Anser cinereus domesticus, and
Cascara cascara.

Location. — Larynx, trachea, and bronchi, as adults; also, appar-
ently accidently, in the abdominal air sacs.

Morphology. — Cyathostoma (p. 41) : Very similar to S. trachea
(p. 34), but larger and less firmly united in copula (head, fig. 47).

Male 8 to 12 mm. long by 200 to 600/* wide. Bursa (fig. 48) inte-
gral, not incised, with all the rays tending to be parallel on each side
of the dorsal ray. The 2 ventral rays are close together; the short
externo-lateral ray is alone and midway between the postero-lateral
and the dorsal rays ; the dorsal ray has a thick stem which bifurcates
to form 2 branches, each of which bears a knob near its base and
a small branch near its tip. There are 2 slender spicules, 600/a long,
slightly hooked at their distal extremities.

Female 20 to 30 mm. long (30 to 40 mm., according to Hayem)
by 1 to 1.5 mm. wide. The conical tail end bears a pointed process.
The somewhat prominent vulva is about one-third of the body length
from the head end. Eggs 80 to 90//, long by 60/* wide and provided,
according to Railliet, with a single polar operculum.

Life history. — Similar to that of Syngamus trachea (p. 36). Rail-
liet found that embryos developed in the egg in 2 weeks. On hatch-
ing the tail end of the worm emerged first. An attempt to infect 1
adult duck and 1 adult goose failed, but after feeding eggs to 3 geese
2 to 3 months old, 1 female worm developed in 1 goose.

Distribution. — Europe (France and Russia) and Asia (Turkestan
and Japan).

CYATHOSTOMA TADORNAE Chatin, 1874

Synonyms. — Sclerostoma tadornae (Chatin, 1874) Linstow, 1878;
Syngamus tadornae (Chatin, 1874) Railliet, 1898.

Host. — Tadorna tadorna.

Location. — Trachea.

Morphology. — Cyathostoma (p. 41): Buccal capsule with 2
pharyngeal teeth at its base (Chapin regards this as an error of
observation in a worm examined from the side only, and thinks there
are probably 6 or 7 teeth ) .

NEMATODE PARASITES OF BIRDS 43

Male 9.6 mm. long and of rather uniform thickness throughout
(fig. 49), brick-red in color. Cuticula transversely striated at 20/«
intervals. Esophagus length 640/i or 1/15 of body length. Bursa sup-
ported by 10 rays of which 4 are simple and G are bifurcated at their
apices. Spicules 50G> long; brown in color.

Female 23 mm. long, brighter red than male. Head 90G> wide.
Esophagus 1.8 mm. long. Vulva near anterior third of body. Body
ends in a conical process (fig. 50) set at an angle to the long axis of
the body. Eggs 56/* long (Chatin says 0.56 mm., but this is evidently
an error and Skrjabin has given the figures as 56//,).

Life history. — Unknown; probably similar to that of Syngamus
trachea (p. 36).

Distribution. — Europe.

Figs. 49-53. — Cyathostoma tadornae. 49, Male. 50, Female tail. From Chapin,
1925, after Chatin, 1874. 51, Cyathostoma laki. Female ; entire worm. 52,
Female ; head end. 53, Vulva. After Blanchard, 1849

CYATHOSTOMA LARI E. Blanchard, 1849

Synonyms. — Strongylus, species Siebold, 1837; Sclerosto-ma cya-
thostomum Diesing, 1851; Sclerostomy lari (E. Blanchard, 1849)
Molin, 1861a; Syngamus lari (E. Blanchard, 1849) Railliet, 1898.

Hosts. — Lams rudibundus, L. fuscus, and Larus, species.

Location. — Nasal and orbital cavities.

Morphology. — Cyathostoma (p. 41) : Red worms, attenuated an-
teriorly. Buccal capsule (fig. 52) marked in front by an annulation.
Esophagus muscular, gradually widening posteriorly. Intestine
sinuous.

Male 6 to 8 mm. long. Body ends abruptly at tail end. Bursa
said to have 6 rays, of which the middle are the most marked (no
satisfactory data on this feature). Spicules equal. Testis a single
very wide tube, sinuous anteriorly.

Female 6 to 13 mm. long by 500/i. to 1 mm. wide; body width
(fig. 51) approximately uniform except for anterior third, in which
it diminishes anteriorly to the head. Tail with slender conical tip.
Cuticula finely striated. Esophagus length y 5 of body length.
Ovaries join capacious uteri which unite to form a vagina 2 mm.

44

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

long which gradually diminishes in width to the vulva. Vulva (fig.
53) a large prominent opening with salient lips slightly posterior
to the middle of the body.

Life history. — Unknown; probably similar to that of Syngamus
trachea (p. 36).

Distribution. — Europe (Belgium and Sicily).

CYATHOSTOMA VARIEGATUM (Creplin, 1849) Chapin, 1925

Synonyms. — Strongylus trachealis Nathusius, 1837; Strongylus
variegatus Creplin, 1849; Syngamus variegatus (Creplin, 1849)
Railliet, 1898.

Hosts. — Ciconia nigra, G. alba ( ? ) , and Grus viridirostris.

Location. — Trachea.

Morphology. — Cyathostoma (p. 41) : Mouth aperture (fig. 55)
circular ; buccal capsule with 6 teeth ; 6 circumoral papillae. Buccal
capsule 260/* long by 140/* wide.

Figs. 54-55. — Cyathostoma variegatum. 54, Male bursa. 55, Head, a, Front
view ; b, side view. From Chapin, 1925, after Linstow, 1890

Male 7.8 to 9.5 mm. long by 350//, wide. Body slightly attenuated
anteriorly and posteriorly; terminating posteriorly in a truncate
tail. Esophagus 730/* long. Dorsal ray (fig. 54) bifurcated. Spic-
ules similar and equal, 690/* long and filiform.

Female 13.5 to 21.5 mm. long by 550/* wide. Esophagus 1.075
mm. long. Anus just anterior to tip of tail; tail acute. Vulva one-
third of body length from anterior end, or 7.6 mm. Eggs 80/* by
40/* (Chapin) or 92/* by 53/* (Skrjabin).

Life history. — Unknown; probably similar to that for Syngamus
trachea (p. 36).

Distribution. — Europe and Eastern Asia.

CYATHOSTOMA BOULARTI (Megnin, 1884) Chapin, 1925

Synonyms. — Sclerostoma boularti Megnin, 1884; Syngamus bou-
larti (Megnin, 1884) Railliet, 1898.
Host. — Casuarius galeatus.
Location.— Trachea.

NEMATODE PARASITES OF BIRDS

45

Morphology. — Cy at ho stoma (p. 41) : Body soft, colored bright red
by intervisceral fluid. Intestine spiral, much longer than body.
Head (figs. 56 and 58) narrower than neck. Buccal capsule with 6
teeth.

Male 7 mm. long by 450//, wide. Head 200//, wide. Buccal cap-
sule (or aperture?) 120//, wide. Bursa (fig. 57) with 2 lobes, each
supported by 5 rays. Spicules slender, 650/* long.

Female 18 to 20 mm. long by 850//, wide. Head 500//, wide. Buc-
cal capsule (or aperture?) 350//, wide. Body shaped like a fish-
hook, with a short conical tail. Anus just anterior to tip of tail.
Vulva prominent, 6 mm. from head end or about one-third of body
length from head end. Eggs 80//, by 55//,, operculated at small end.

Life history. — Unknown; probably similar to that of Syngamus
trachea (p. 36).

Figs. 5G-58. — Cyathostoma boularti. 56. Head ; side view. 57, Male bursa.
58, Head; front view. From Chapin, 1925, after Megnin, 1884

Distribution.

Paris).

-Australia and Europe (in zoological garden,

CYATHOSTOMA AMERICAN UM Chapin. 1925

Host. — Buteo borealis.

Location. — Posterior thoracic air sacs.

Morphology. — Cyathostoma (p. 41) : Cuticle smooth, without
transverse striations.

Male 12 nun. long: body (fig. 59) somewhat attenuated in ante-
rior fifth. Buccal capsule about as wide as deep, its internal trans-
verse diameter 185/x, the depth of its chitinous portion 170/*; wall of
capsule 25//, thick. Six or seven triangular buccal teeth, occupying
the entire floor of the capsule but not continued up the side of the
capsule in the form of ridges. The largest (lateral) teeth are about
57//, high. Head with circle of 6 papillae, situated 60/t behind the
anterior extremity, each papilla in form of rounded knob, 10//, in

46

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

diameter. Esophagus 730/* long, clavate, its diameter 74//, at the
anterior end, 130/x at the posterior end ; nerve ring at the middle of
the length, cervical papillae at about posterior seventh of esophagus,
excretory pore just anterior to beginning of intestine. Bursa (fig.
61) well developed, 750/x in transverse diameter when outspread.
Ventral rays similar and approximate; externo-lateral shorter than
other lateral rays and with a prominent ventral hump ; medio-lateral
and postero-lateral rays similar and approximate; externo-dorsal

Fig. 59.

-Cl'ATHOSTOMA AMERICANUM. OUTLINE OF FEMALE AND MALE WORMS.
CHAPIN, 1925 (a> = EXTENT OF UTERI)

After

ray more slender than, but equal in length to, the externo-lateral ray,
arising at the base of the dorsal ray. Dorsal ray divided near its
extremity into 2 simple branches. Spicules filiform, equal, from 470
to 490/x long, united at their tips and each bearing a finely striated
wing.

Female up to 30 mm. long; similar in form (fig. 59) to the male.
Buccal capsule (fig. 60) much broader than deep, its internal
transverse diameter 370/i, the depth of its chitinous portion 280/x;

Fig. 60. — Cyathostoma americanum. Head, a, Side view, male ; b,
front view ; c, side view, female. After Chapin, 1925

wall 20/t thick. Buccal teeth shorter and blunter than in male;
largest tooth 60/x high. Head papillae as in male, slightly more
anterior in location. Esophagus 960/i long, clavate, its diameter 130/x
at anterior end, 250/x at posterior end ; nerve ring at middle of length,
cervical papillae opposite the thickest portion of esophagus, excre-
tory pore near its posterior end. Vulva (fig. 62fr) just anterior to
middle of body, its lips prominent. Anus (fig. 62c) just anterior
to slender caudal appendage. Eggs (fig. 62a) 72/t by 42/x, thin
shelled, with a small operculum at one pole.

NEMATODE PARASITES OF BIRDS

47

Life history. — Unknown; probably similar to that of Syngamus
trachea (p. 36).
Distribution. — United States (Fairfax County, Virginia).

CYATHOSTOMA COSCOROBAE Chapin, 1925

Host. — Coscoroba coscoroba.

L (" [ation . — Trachea.

Morphology.— Cyathostoma (p. 41) :

Mule 5.5 nun. long (estimated) ; anterior portion attenuate. Buc-
cal capsule (fig. 63) with straight sides, 83/* wide by 100/* deep;
lateral walls 13/* thick. Six buccal teeth, the alternate teeth (about
33/* high) are about twice the size of the others. Head papillae ap-
parently as in ('. americanum. Esophagus about 700/* long, attaining
its' greatest diameter (150/t) at its posterior end. Nerve ring just

som 62

Bigs. 61-62. — Cyathostoma americanum. 0J, Male, a, Bursa; b,
gubebnaculum. 62, (i , egg ; b. vulva; c, tail (female). after
Chapin, 1925

anterior to middle of esophagus; position of cervical papillae and
excretory pore not determined. Bursa (rig. 64) well developed, 720/*
in transverse diameter when outspread. Ventral rays similar, slen-
der and approximate, about 105//, long; lateral rays parallel and con-
tinuous throughout their length: externo-lateral ray the shortest
(135/* long); medio-lateral and postero-later-aJ rays L50/t and 210/t
long, respectively; externo-dorsal ray 165/* long, arising near the
base of the main trunk of the dorsal ray, which is 150/* to its bifurca-
tion; branches of dorsal ray sinuous, each with 3 terminations, as in
('. bronchiaZis. Spicules filiform, from 660/* to 720/* long, united at
tips and each bearing a finely striated wing. Gubernaculum present.
92/* long.

Female about 20 mm. long. Buccal capsule (fig. 63) trapezoidal
in optical section, 210/* deep by 225/< wide at base, 285/t wide at apex:
3612- 27 5

48

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

walls 30/a thick. Six teeth, their height proportionately less than in
the male, the highest teeth being 60/*. Esophagus 900/* long, attain-
ing its greatest diameter (225/*) near its posterior end. Nerve ring
at anterior % of length of esophagus ; cervical papillae and excretory
pore not located. Vulva at about the middle of the body length;
vagina very short; uteri divergent. Tail acute; anus 225/* anterior
to tip. Eggs oval, 80/a long by 50/t wide, with a minute operculum
at the slightly smaller end.

Life history. — Unknown; probably similar to that of Syngamus
trachea (p. 36).

Distribution. — South America (and from captive bird in North
America, Philadelphia Zoological Park, Philadelphia, Pa., U. S. A.).

65.

Figs. 63-64. — Cyathostoma coscorobae. 63, Buccal capsules of female and male.
64, Bursa. After Chapin, 1925

Suborder Ascaridata Railliet and Henry, 1915

Suborder diagnosis. — Myosyringata (p. 4) : Polymyarian. Mouth
with 3 or 6 lips or without lips. When 3 lips are present, one is
median and dorsal, the others are submedian and are approximated
in the ventral line. Buccal capsule absent. Males with one or two
spicules. Females usually with two ovaries, occasionally more than
two (as in ascarids of snakes), oviparous. Development usually di-
rect and without intermediate host; it may be complicated by the
larvae journeying through the body, before maturity can be reached;
exceptionally (as in ascarids of seals) there is an intermediate host
(fish).

Superfamily ASCAROIDEA Railliet and Henry, 1915

Synonym. — Ascaridea Diesing, 1861, of Travassos, 1914.
Superfamily diagnosis. — Ascaridata (p. 48) : Characters of the
suborder.
Type-family. — Ascaridae Baird. 1853.

NEMATODE PARASITES OF BIEDS 49

KEY TO FAMILIES OP ASCAROIDEA

Mouth with 3 lips or without lips. Males with a preanal sucker which may
be limited by a ring, or formed by a simple longitudinal depression.

Heterakidae, p. 49.

Mouth with 3 prominent lips, or with 3 main lips and 3 intermediate lips. Male
without preanal sucker Ascaridae, p. 135.

Family HETERAKIDAE Railliet and Henry, 1914

Synonym. — Heteracidae Railliet and Henry, 1914.

Family diagnosis. — Ascaroidea (p. 48) : Polymyarian. Mouth pro-
vided with 3 lips or without lips and of variable shape. Esophagus
cylindrical or club-shaped, often followed by a distinct bulb. Males
with a preanal sucker which may be limited by a chitinous ring or a
delicate cuticular membrane, or formed by a simple longitudinal de-
pression. Two spicules, one or both of which may tend to atrophy
or show imperfect chitinization, and with gubernaculum present or
absent. Vulva usually near middle of body.

Type- genus. — Heterakis Dujardin, 1845.

The Heterakidae, including Heterakis, Ascaridia, and Suhulura,
are placed in the superfamily Ascaroidea in accordance with Railliet
(1916), Baylis (1923), etc. Travassos (1920) separated Heterakis
and Ascaridia, putting the former in the Oxyuroidea, the latter in
the Ascaroidea. The musculature, however, is identical in the two
genera and is polymyarian, whereas that of the Oxyuridae is mer-
omyarian; this similarity of the musculature of Heterakis and As-
caridia is accepted as the fundamental basis of classification by the
present author. Ascaridia has features very comparable with those
of Ascaris, the chief difference between the two being the sucker, the
presence of which in Ascaridia forms the transition between Ascaris
and Heterakis. The differences between Ascaridia and Heterakis
seem to merit only generic rank, not superfamily rank, as in Travas-
sos' classification. Seurat (1918) states that the genital tubes ©f
Ascaridia galli (=A. perspicillum) are identical with those of the
heterakids, showing the affinities of Ascaridia with Heterakis.

KEY TO SUBFAMILIES OF HETERAKIDAE

Mouth with 3 well-defined lips. Esophagus with 01' Without bulb; sucker of
male nearly circular and having a cutinous (chitinous) rim.

Heterakinae, p. 49.

Mouth with lips inconspicuous or wanting, followed by a vestibule; esophagus
with bulb; sucker of male spindle-shaped, without a cutinous (chitinous)
rim Subulurinae, p. 104.

Subfamily Heterakinae Railliet and Henry, 1912

Subfamily diagnosis. — Heterakidae (p. 49) : Mouth with 3 well-
defined lips; esophageal bulb present or absent. Preanal sucker

50 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

nearly circular and limited by a cutinous (chitinous) ring. Spicules
equal or unequal. Gubernaculum present or absent.
Type-genus. — Heterakis Dujardin, 1845.

1. Esophageal bulb lacking; caudal alae feebly developed Ascaridia, p. 77.

Esophageal bulb present ; caudal alae well developed 2.

2. Head with " cordons " consisting of tubular grooves- Pseudaspidodera. p. 102.
Head without " cordons " Heterakis. p. 50.

Genus HETERAKIS Dujardin, 1845

Sj/nonijms. — Heteracis Molin, 1858; Ganguleterakis Lane, 1911.

Generic diagnosis. — Heterahinae (p. 49) : Mouth with 3 lips.
Esophagus subcylindrical, progressively swollen toward its posterior
extremity and with a distinct bulb. Two lateral membranes present
or absent. Males with caudal alae well developed and sustained by
papillae of ray-like appearance. Spicules equal, subequal, or unequal,
without accessory piece. Preanal sucker with a distinct cutinous
(chitinous) ring. Females with vulva toward the middle of the
body, the uterine branches passing in opposite directions. Eggs with
thick shell.

Parasitic in intestines (especially the ceca) of birds, mammals, and
reptiles.

Type-species. — Heterakis gallinae (Gmelin, 1790) Freeborn, 1923
( =H. vesicularis ) .

KEY TO SPECIES OF HETERAKIS

1. Description incomplete; from Ciconict (Euxenura) rttagitari.

Heterakis valdenmcroiiata, p. 74.
From other hosts than above 2

2. Male with not more than 9 pairs of caudal papillae ?>.

Male with more than 9 pairs of caudal papillae 4.

3. Male with 6 pairs of caudal papillae; sucker 35/u anterior to elo'aeal aperture:

tail of female 1/6:8 of total length Heterakis bancrofti, p. 50.

Male with 9 pairs of caudal papillae; sucker 194,ot anterior to cloacal aper-
ture; tail of female 1/14 of total body length_ Heterakis chenonettae, p. 60.

4. Spicules unequal 5.

Spicules equal or subequal (not more than r>0,u difference in their lengths ) _ 13.

5. Longer spicule over 1 mm. in length 6

Longer spicule less than 1 mm. in length 12

6. Longer spicule over 2 mm. in length 7.

Longer spicule less than 2 mm. in length 10.

7. Short spicule 640/x long Heterakis parisi, p. 69.

Short spicule 700/t or longer 8.

S. Tail short in both sexes, the anus of female being 780^, the cloacal aperture

of male 170fi, from posterior end Heterakis valvata, p. 74.

Tail long compared with above, the anus of female being not less than 9SS,u.

the cloacal aperture of male not less than 416/i, from posterior end 9.

9. Preanal sucker 60 to 75/* in diameter; right spicule 2 to 2.17 mm. long;
female JO to in mm. long; eggd 03 to Tin I6og in 38 to !S,u wide.

Heterakis gallinae, p. 52.

NEMATODE PARASITES OF BIRDS 51

Preanal sucker 80 to 90/* in diameter; righl spicule 2.3 nun., long; female
T ! to 9.6 nun. long; eggs T."i to 78m long by 43 to 48/* wide.

Heterakis longecaudata, p. 66.

10. Preanal sucker 250/* in diameter: cluneal aperture I.;', mm. from posterior

extremity; female 9.5 mm. long; anus 1.5 mm. from posterior extremity.

Heterakis bosia, p. 57.

Preanal sucker not over 124/* in diameter: cloaca] aperture not over 488/t

from posterior extremity: female 7 to 8 mm. long; anus not more than

1 mm. from posterior extremity 11.

11. Preanal sucker 83/* internal diameter, 124/* external diameter; 5 pairs of

preanal papillae and 1 pair of very large adanal papillae: length of caudal
appendage of male less than distance of cloacal aperture from most

posterior papillae Heterakis bonasae, p. 75.

Preanal sucker only 36/* in diameter; 3 pairs of preanal and 1 pair of small
ventral adanal papillae: caudal appendage almost twice as long as dis-
tance from Cloacal aperture to most posterior papillae.

Heterakis fariai, p. 63.

12. Short spicule 150/* long; female 5.1 mm. long; its anus 540/* from posterior

end: vulva at about posterior third of body, dividing body length in ratio

of 12 : 5 Heterakis pusilla, p. 70.

Short spicule 260/* long; female 8.2 mm. long, its anus 830/* from posterior
end ; vulva slightly anterior to middle of body.

Heterakis putaustralis, p. 71-

13. Spicules over 1 mm. long 14.

Spicules less than 1 mm. long 17.

14. Male 14 to 22 mm. long: female 22 to 31 mm. long; eggs 52/* by 29/*;

according to figure, spicules over 2 mm. long Heterakis arquata, p. 55.

Male not over 9 mm. long; female not over 12 mm. long; eggs 70,<* or more
by 42/* or more: spicules not. over 1.8 mm. long 15.

15. Head with 3 lips and .''. small interlabia ; no papillae described in vulvar

region : spicules about 1.25 mm. long Heterakis interlabiata, p. 64.

Head with 3 lips but no interlabia: vulvar region with variable number
of papillae; spicules equal or subequal, 1.4 to 1.8 mm. long 16.

16. Lateral alae extend only along esophageal region of body ; male 7.5 to 9

mm. long, its tail 1/13 of total length; female 9 to 12 mm. long, its tail

1/7.3 of total length Heterakis isolonche, p. 64.

Lateral alae extend almost the whole body length; male 15 nun. long, its
tail 1/23 of total length; female 17 mm. long, its tail 1/14 of total length.

Heterakis neoplastica, p. 65.

17. Spicules over 600/* long 18.

Spicules not over 500« lout; 23.

18. Male 4 mm. long; female 4.2 nun. long Heterakis nattereri, p. 75.

Male 9 mm. or longer; female 14 mm. or longer 19.

19. Preanal sucker 290/* in diameter; the most anterior pair of preanal caudal

papillae very small and situated a considerable distance anterior to the

sucker; spicules at least 890/* long Heterakis tenuicauda, p. 7.'!.

Preanal sucker not over 230/* in diameter; the most anterior preanal
papillae not situated as above: spicules not over 860/t long 20.

20. Length of females 25 mm. or more, the tail short (anus not over 4<!0/* from

po&terior end) 21.

Females not over 15 mm. long, the tail comparatively long (anus 800/* or
farther from posterior end) 22.

52 BULLETIN 110, UNITED STATES NATIONAL MUSEUM

21. Male with cloacal aperture 270/t from posterior end ; preanal sucker 180/x in

diameter ; spicules 630 to 650/* long ; female with anus 460/t from posterior

end ; vulva at anterior third of body Heterakis alata, p. 55.

Male with cloacal aperture 420/x from posterior end ; preanal sucker 230/x in
diameter ; spicules 850/t long ; female with anus 220/t from posterior end ;
vulva at about anterior fifth of body Heterakis skrjabini, p. 71.

22. Tail of male 1/2S of total body length ; preanal sucker 90/t in diameter ; 10

pairs of caudal papillae Heterakis psophiae, p. 69.

Tail of male 1/55 of total body length ; preanal sucker very promi-
nent, 220/* iu diameter; 8 pairs of caudal papillae.

Heterakis papillosa, p. 67.

23. Female not over 7.6 mm. long 24.

Female 11.6 mm. to 34 mm. long 26.

24. Preanal sucker broken on each side, as well as posteriorly, by a papilla ;

spicules equal, 400/i long ; eggs 63/t long by 35/t wide.

Heterakis brevispiculum, p. 59.

Preanal sucker not broken on each side, but only posteriorly, by a papilla ;

spicules subequal, there being 50/i difference in their lengths ; the longer

not over 370/* in length 25.

25. Male 5.5 mm. long by 210/j. wide; preanal sucker 55/t in diameter; female

6.6 mm. long by 300/i wide, its tail <ks of total body length ; eggs 50/t long

by 30/t wide Heterakis beramporia, p. 56.

Male 7.5 mm. long by 410/* wide ; preanal sucker 71/t in diameter ; female
7.6 mm. long by 460/i wide, its tail V s of total body length ; eggs 57/6 long
by 42/* wide Heterakis hamulus, p. 63.

26. Spicules 110 and 130/t long respectively; tail of female 1/7.8 of total body

length Heterakis macroura, p. 67.

Spicules 260/* long or longer ; tail of female not as long as above except
possibly in H. dispar where length not given 27.

27. Female 34 mm. long ; vulva in anterior part of body ; male 19 to 20 mm.

long ; spicules not over 390/* long Heterakis brasiliana, p. 58.

Female not over 23 mm. long ; vulva posterior to middle of body ; male
not over 18 mm. long; spicules 400 to 500/i long 28.

28. Male 7.8 mm., female 11.6 mm. long ; eggs 70/* long.

Heterakis caudata, p. 59.

Male 11 mm. or longer ; female 14.8 mm. or longer ; eggs not more than

62/* long 29.

29. Male 11 to 18, female 16 to 23 mm. long; two lateral membranes present;

vulva surrounded by a cuticular thickening ; in other hosts than below.

Heterakis dispar, p. 62.
Male 13.1 mm. long, female 14.8 mm. long ; no lateral membranes and no
cuticular thickening of vulvar region described ; in Cygnus atratus.

Heterakis circumvallata, p. 61.

This key does not include Heterakis acuticaudata (p. 77), as the
evidence is at present inadequate for the allocation of that species
to this genus in its restricted sense. The key does not include the
four new species of Heterakis recently described by Chandler (see
Addenda, p. 385).

HETERAKIS GALLINAE (Gmelin, 1790) Freeborn, 1923

Synonyms. — Ascaris gallinae Gmelin, 1790; Ascaris vesieularis
Froelich, 1791, part; Heterakis vesieularis (Froelich, 1791) Dujar-

NEMATODE PARASITES OF BIRDS

53

din, 1845; Heterakis papulosa Railliet, 1885, misdet., not Ascaris
papillosa Bloch, 1782 (=Heterakis monticelliana Stossich, 1892)
from bustard.

Hosts. — Anas boschas domes tica, A. tadorna, Anser anser, A.
cinereus domesticus, Bonasa sylvestris, Geriomis satyra, Ghenopsis
atrata, Ghrysolophus pictus, Golinus virginianus, Gorvus cajanus,
Goturnix communis, G. dactylisonans, Gupidonia cupido, Gallus
gallus, Grossiptodon manscliuricum, Lagopus mutus, L. scoticus,
Meleagris gallopavo, Numida meleagns, Ortyx virginianus, Otis
tarda, O. tetrax, Pavo cristatus, Perdix cinerea, P. cotumix, P.
perdix, P. saxatilis, Phasianus colcliicus, P. gallus, P. nycthemerus,
P. pictus, P. veneratus, P. versicolor, Tadorna tadorna, Tetrao
bonasia, T. lagopus, T. urogallus.

The specimens from Otis tarda are probably Heterakis papillosa
(Bloch, 1782).

Figs. 6o-67.

-Heterakis galmnae. 65, Male tail, lateral view ; 66, ventral
view ; g7, female tail. after l.ane, 1917

Location. — In cecum, usually; in small intestine, colon and rectum,
rarely.

Morphology. — Heterakis (p. 50) : Small, rigid, white worms, the
head end bent dorsally from the region of the esophageal bulb.
Mouth with 3 small, equal lips without teeth and each with 2 papillae,
according to Lane. Two narrow lateral membranes extend almost
the entire length of the body. Esophagus with 6 longitudinal rows
of transversely placed chitinous rods and with a well-developed bulb.

Male 7 to 13 mm. long. The straight tail (figs. 65 and 66) termi-
nates in a subulate point and has 2 large lateral bursal wings. Cloa-
cal aperture 450/x from caudal extremity. There are 12 pairs of
papillae and a well-developed preanal sucker 60 to 75//. in diameter
with strongly chitinized w r alls (figs. 66 and 78) ; there is a small
semicircular incision in the posterior margin of the wall of the
sucker. Four pairs of papillae are between the cloacal aperture and
the end of the tail, 4 pairs of ray-like papillae and 2 pairs of sessile
papillae are in the vicinity of the cloacal aperture, and 2 pairs of
ray-like papillae are in the vicinity of the sucker. The spicules are

54 BULLETIN 140, "UNITED STATES NATIONAL MUSEUM

dissimilar, the right being 2 to 2.17 mm. long and the left 700/* to
1.1 mm. long.

Female 10 to 15 mm. long. The tail (fig. G7) is long, narrow, and
pointed, the anus 1 mm. or more ( ? ) from the tip. Vulva not salient,
situated slightly posterior of the middle of the body. From vulva
to bifurcation of the uterus is about 4.5 mm. Eggs thick-shelled,
ellipsoidal, 63 to 71/a long by 38 to 48/a wide, according to most
authors, or 68 to 75/x long by 36 to 38/x wide, according to Uribe, not
yet segmenting when deposited; Uribe notes that the shell is thick-
ened at one end of the egg, and that this thickening may enclose a
lenticular clear space.

Life history. — Eggs pass in the feces of the host and develop in
7 to 12 daj^s under favorable conditions of temperature (18 to 29° C.)
and moisture to the point where each contains an infective embiyo.
Eggs develop in water, physiologic saline solution, on salt solution
agar, in 1 : 1,000 corrosive sublimate or 1 per cent sulphuric acid.
When these eggs are swallowed b}' suitable birds, the embryos are
released from the shell and develop to adult worms. Within l 1 /^
hours after ingestion of the eggs, the larvae are found to have
emerged into the small intestine. At the end of 21 hours they are
present in the ceca, and sometimes in the colon, and are 250/a long by
18/i wide. In 10 days they are 790//. long by 45/x wide. The entire
development in the bird takes place in the digestive tract, the worms
being mature in 21 days. There is no wandering of the larvae to
the lungs as is possibly the case in Asca/ridia galli (p. 82), although
the Oklahoma Experiment Station reports that all chicks fed with
larvae died of pneumonia in 8 to 10 days. Galli- Valerio and also
Latulle and Marotel have reported the occurrence of the larvae in
tumors in the ceca, and Graybill has found the larvae in the mucosa.
Uribe finds the young worms in the cecal glands from the second to
the fifth day after infection, subsequent development taking place in
the lumen of the intestines, but about the ninth day worms may be
found with the anterior third of the body inserted in the cecal gland.
He found mature females in 56 days. Earthworms may ingest the
eggs and carry them to the intestine, and birds may become infested
by eating such earthworms; the earthworms may also pass these eggs
in their casts and thus infect otherwise uninfected ground.

Eggs from worms collected from chickens killed and chilled at
34° F., then frozen for 3 days at 0° to 10° F., then kept at 0° to
10° F. for 6 months, were found to develop embryos in 75 per cent
of the eggs examined by Riley and James. According to Graybill,
ova may survive desiccation for 16 to 18 days; eggs in soil contained
live embryos after 8 months.

Distribution. — Cosmopolitan. This species is very common in the
United States.

NEMATODE PARASITES OF BIRDS QQ

HETERAKIS ALATA Schneider, 1866

Hosts. — Orypturus, species and Tinamus, species.

Location. — Intestine.

Morphology. — Heterakis (p. 50): Mouth with 3 small lips. Lat-
eral membranes large. Travassos says esophagus with bulb more or
less developed, with a diameter of about 240 to 300/*; the slender part
measures about 1.2 mm. long.

Male 17 to 20 mm. long. Caudal alae (fig. 68) wide; preanal
sucker about ISO//, in diameter, with a papilla on its posterior border.
Schneider described and figured only 9 pairs of caudal papillae but
Travassos states there are 14; his figure seems to show 13 pairs.
Spicules slender, about 030 to 650/x, long. Cloacal aperture about
279// from posterior end.

Female 25 mm. long, according to Schneider; 35 to 37 mm. long,
according to Travassos. Vulva at anterior third of body. Anus
about 460,u from posterior end. Eggs 59// long by 37// wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribution. — South America (Brazil).

HETERAKIS ARQUATA Schneider. 1866

Synonym. — Ascaridia arquata (Schneider, 1866) Railliet and
Henry, 1914. The allocation by these authors of this species to
Ascairidia evidently was based on Schneider's inadequate descrip-
tion: Travassos has shown, in a more complete study, that it belongs
in the genus Heterakis.

Hosts. — Grypturus cupreus and Psophia viridis.

Location. — Intestine.

Morphology. — Heterakis (p. 50) : Mouth with 3 very small lips
with no teeth. Lateral membranes present. Travassos says esopha-
gus 1.35 to 1.44 mm. long, with a small bulb and a slight dilation in
the anterior parr.

Male 14 to 22 mm. long. Preanal sucker 90// in diameter, a papilla
on its posterior border. In addition (fig. 69) 12 pairs of caudal
papillae, •"'> of them preanal. Spicules equal and similar: Travassos
says they are about 260// long, but he must mean 2.6 mm., judging
from his figure (fig. 70), since he says the cloacal aperture is 270//
from the posterior end.

Female 22 to 31 nun. long. Vulva salient, situated a little an-
terior to the middle of the body. Anus about 1.12 mm. from pos-
terior end. Eggs 52/* long by 2!)// wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribution. — South America (Brazil).
3612—27 6

56

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

HETERAKIS BANCROFT! Johnston, 1912

Host. — Oatheturus lathami.

L ocation. — Ceca.

Morphology. — Heterakis (p. 50) : Mouth with 3 prominent lips of
equal size. Alimentary canal of usual Heterakis type.

Male 4.3 mm. long by 220/* wide. Tail (fig. 71) with very narrow
delicate appendage. Sucker 73/* in diameter, with chitinous ring, its
posterior margin about 35/* anterior to cloacal aperture. Caudal
alae lobulated. Six pairs of papillae, of which 2 are preanal and 4
postanal. Spicules equal, 860/* long, sharply pointed.

W 71.

Wnfrn

Figs. 68-71. — 68, Heterakis alata. Male tail. After Schneider, 1866. Heterakis
arquata. Male tail. 69, After Schneider, 1866. 70, After Travassos, 1913. 71,
Heterakis bancrofti. Male tail. After Johnston, 1912

Female 6.25 mm. long by 330/* wide. Tail rather short (anus 910/x
from end) and sharply pointed. Vulva at about middle of body.
Life history. — Probably similar to that of H. gallinae (p. 54).
Distribution. — Australia.

HETERAKIS BERAMPORIA Lane, 1914

Host. — Gallus gallus.

Location. — Ceca ; adults in lumen, larvae in nodules.

Morphology. — Heterakis (p. 50) : Very similar to H. gallinae, but
with short spicules. Lateral membranes begin about 800/t from head
and extend almost entire length of body.

Male 5.5 mm. long by 210/* wide. The circular sucker (fig. 72) is
55/* in diameter. The spicules 350 and 300/* long, the longer with a
tapering curved point, the shorter expanding in the terminal third
end with a prominent angle on the ventral aspect near the point.

NEMATODE PARASITES OF BIRDS 57

Twelve pairs of papillae; 2 pairs near the sucker. 6 in the region of
the cloacal aperture, of which the third lateral pair is comparatively
small; the 2 ventral pairs of this group lie close together; posterior
to this group is a pair of moderate sized papillae ; the lateral papillae
of the caudal group are relatively distant from one another and the
posterior pair is much the larger.

Female 6.6 mm. long by 300/x wide. The body has the usual dorsal
curve anteriorly and also curves ventrally somewhat abruptly at the
level of the vulva, which is in the middle of the body; a posterior
flap projects forward over the vulva. The anus is 660/x from the tip
of the tail. The eggs are 50/* long by 30/*, wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — Asia (Intlia (Bengal) and Philippines).

HETERAKIS BOSIA Lane. 1914

Hosts. — Ceriomis satyra and T rag o fan satyra.

Location. — Intestine, probably ceca.

Morfhology. — Heterakis (p. 50) : Cuticle with cross-striations.

Fig. 72. — Heterakis BEitAMromA. Male tail. After Lane, 1914

Lateral alae throughout practically whole length of body, ending
near the sucker of the male and 30/*, from posterior extremity of
female. Esophagus 1.5 mm. long.

Male 8.1 mm. long. Cloacal aperture 1.3 mm. from caudal ex-
tremity; long slender appendage (that is, part of tail posterior to
the papillae) is 700/t long. Sucker 250/*, in diameter, its posterior
rim 350ju, from the cloacal aperture. Twelve pairs of caudal
papillae (fig. 73a) of which 4 are preanal, 2 adanal, 6 postanal.
Spicules unequal, the right (fig. 73c) 1.6 mm. long, its proximal end
wide (75/x) like a spear head, its distal end a long sharp point; left
spicule (fig. 73£>) 900//, long, with a one-sided expansion at about a
quarter of its length from the point.

Female 9.5 mm. long by about 400/z wide. Anus 1.5 mm., caudal
papillae 800/*, from posterior extremity. Vulva slightly posterior
to middle of body; posterior to it there are occasionally 5 or 6
cuticular tubercles. Eggs 75/*. long by 35/*, wide.

58

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Probably similar to that of II. gallinae (p. 54).
Distribution. — Asia (India).

HETERAKIS BRASILIANA Linstow. 1899

Synonym. — Ascaridia brasiliana (Linstow, 1899) Travassos, 1913.
In a later study, Travassos (1918) reassigns this species to Heterakis.

Hosts. — Perdix, species and Rhynchotus rufesoens.

Location. — Intestine.

Morphology. — Heterakis (p. 50) : Mouth with 3 lips; at the base
of the lips there occurs laterally a papilla. Lateral alae of cephalic
extremity large. Esophagus 1/11.5 of total length. Posterior ex-
tremity of both sexes with digitiform prolongation.

Male 19.1 mm. to 20 mm. long by 520 to 590^ wide. Tail 1/57
of total length, according to Linstow; 450/* long, according to

Fig. 73. — Heterakis bosia. a, Mali: tail and point of kigiit spicule; ?». left

SPICULE FROM DORSUM : C, RIGHT SPICULE FROM SIDE. AFTER I.ANE, 1014

Travassos. Preanal sucker 200/x in diameter, with a papilla on pos-
terior rim. The earlier descriptions (Linstow, 1899, and Travassos,
1913) describe and figure 10 pairs of caudal papillae but those
figures omitted all detail of the cloacal region, whereas according
to Travassos, 1918, there are 12 pairs of papillae, his figure (fig. 74)
showing a pair just anterior and another just posterior to the cloacal
aperture. The 12 pairs are arranged as follows: 2 preanal, 6 adanal,
4 postanal. There is in addition an unpaired papilla on the right
side slightly posterior to the sucker. Spicules short and wide, about
260/x long, according to early descriptions; 390/x, long, according to
Travassos, 1918.

Female 34 mm. long by 790/t wide. Vulva in anterior part of
body. Tail 1/30 of total iength. Eggs 05/x long by 44^ wide (Tra-
vassos says 0.065 mm. long by 0.944 mm. wide but the latter figure
is evidently a typographical error for 0.044).

NEMATODE PARASITES OF BIRDS

59

Lifi history. — Probably similar to that of II. gallinae (p. 54).
Distribution. — South America (Brazil).

HETERAKIS BREVISPICULUM Cendre, 1911

Hosts. — Francolinus bicalcaratus, Gallus gallus, and Numida
meleagris.

I o < <i t io 1 1 . — Cec a .

Morphology. — Heterakis (p. 50) : These worms are very similar
to //. gallinae (p. 53), but the males have short, equal spicules and
the sucker rim (fig. 77) is broken by papillae on each side, as well as
by the papilla in the posterior rim.

Male 5.. '14 to 8.35 mm. long by 250 to 300/* wide. The spicules
(fig. 76) 400/* long and equal; sucker as described above. The

Fig. 74. — Heterakis brasiliana. Male tail. After Travassos, 1918

spicules are shaped like a small nail, with an enlarged head and with
its tip united to a dorsal prominence by an oblique line. Travassos
has described II. gallinae from Brazil as having equal spicules 270/*
long. Apparently he had one of the species similar to this one, with
equal spicules. Caudal pupillae (fig. 75).

Femah 6.38 to 10.6 mm. long by 260 to 360/* wide. The vulva is
in the middle of the body. The eggs are elliptical. 63/* long by 30/*
wide, and deposited before the formation of the embryo.

Life history. — Probably similar to that of //. gallinae (p. 54).

Distribution. — Africa (Dahomey and Belgian Congo) and South
America ( Brazil).

HETERAKIS CAUDATA Linstow, 1906

Hosts. — Anas sj>ohs<i and Lampronessa sponsa.
Location. — Ceca.

Morphology. ^-Heterakis (p. 50): Three semicircular lips.
Esophagus with bulb.

60

BULLETIN - U0, UNITED STATES NATIONAL MUSEUM

Male 7.8 mm. long by 290/x, wide. Esophagus 1/8.5, tail 1/42, of
total body length, the latter (fig. 79) with a styloid prolongation at
its posterior end. Preanal sucker 180/* in diameter. Caudal alae
broad. Two long pedunculated papillae on each side of sucker; in
addition 4 pairs just anterior to the cloacal aperture, and 4 pairs sit-
uated more posteriorly. Spicules equal, 440//. long.

Female 11.6 mm. long by 290//, wide. Tail long and pointed,
1/15 of total body length. Vulva posterior to middle, dividing body
in ratio of 19 :14. Eggs 70/x long by 44/x wide.

©

Figs. 75-78. — Heterakis rreyispiculum. 75, Male tail. 7G,
Spicule. 77, Region of sucker. 78, Heterakis gallinae.
Region of sucker. All after Gendre, 1911

Life history.— Probably similar to that of H. gallinae (p. 54).
Distribution. — Europe (Germany (Zoological Museum, Koenigs-
berg) ).

HETERAKIS CHENONETTAE Johnston, 1912

Host. — G henonetta jubata.

Location. — Ceca.

Morphology.— Heterakis (p. 50) : Anterior end tapering rather
rapidly; posterior extremity pointed. Lips equal, small.

Male 6.8 mm. long by 190/* wide. Caudal alae prominent, with 9
pairs of papillae (fig. 80), of which 4 are preanal (3 just anterior

NEMATODE PARASITES OF BIRDS

61

to cloacal aperture and 1 at side of sucker). Cloacal aperture promi-
nent, the sucker, which is also prominent, 194/a anterior to it. Johns-
ton's description of the spicules is confusing. He states : " The
spicules are 1.17 mm. in length and 0.013 mm. in breadth. The
longer male spicule is strongly curved, and has a fairly uniform
breadth (0.008 mm.), but tapers slightly toward the extremity.
The length is 0.48 mm. The shorter spicule has a length of 0.18
mm., and a breadth (in its mid-region) of 0.012 mm., but the ex-
tremity is widened and rounded." The spicule which Johnston
figures agrees with the first length given, being about 1.1 mm. long.

Female 7.7 mm. long by 260/* wide. Vulva just posterior to middle
of body (4.2 mm. from anterior end). Anus 530/* from posterior
end. Eggs in uterus embryonated.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution . — Australi a ( Sydney ) .

Figs. 70-82. — 79, Heterakis caudata. Male tail. After Linstow, 1906. 80,
Hetekakis chenonettak. Male tail. After Johnston, 1!>12. 81, Hetera-
kis OIRCUMVALLATA. MALE TAIL. AFTER LlXSTOW, 190G. 82, HETERAKIS

dispar. Male tail. After Railliet, 1893

HETERAKIS CIRCUMVALLATA Linstow, 1906

Host. — Cygnus atratus, black swan, a species from Australia; not
Cygnus olor. Gedoelst lists this worm from G. olor, but it is doubt-
ful if it has ever been correctly reported from this host.

Location. — Cecum.

Morphology. — Heterakis (p. 50) : Head with 3 hemispherical lips.
Esophagus short, with terminal bulb : intestine at origin wider than
esophagus.

62 BULLETIN 140, "UNITED STATES NATIONAL MUSEUM

Male 13.1 mm. long by 410/* wide. Tail (fig. 81) 1/57 of total
body length. The spicules are equal and 480//. long. The sucker is
190/* in diameter and has a heavily chitinized wall. There are 2
long-stalked papillae on each side of the sucker, 4 pairs of digitiform
preanal papillae (the last pair is adanal, according to Linstow's
figure), and 4 pairs of postanal papillae, of which the last 3 are
grouped close together.

Female 14.8 mm. long by 400//. wide. The pointed tail comprises
1/12 of the total length. The vulva is somewhat posterior to the equa-
tor of the body, about 4/7 of the body length from the head. Eggs
62/* long by 44//, wide.

Life history. — Probably similar to that of II. galllnae (p. 54).

Distribution. — Europe (Germany (Koenigsburg Zoological Mu-
seum ) ) .

HETERAKIS DISPAR (Schrank, 1790) Dujardin. 1845

Synonym. — Ascaris dispar Schrank, 1790.

Hosts. — Anas anser domestica, A. boschas, A. boschas domestica,
A. canadensis, A. leucopsis, A. ??ioschata, A. tadorna, Anser anser ;
A. cinereus, A. fabalis, A. segetum, Bernicla sandwichensis, Sumia
passerina, /St?*i% passerina, Tadorna tadorna.

Location. — Cecum.

Morphology. — Heterakis (p. 50) : White worms, thinned pos-
teriorly. Two lateral membranes, which are widest in the anterior
portion of the body and gradually narrowed posteriorly. Mouth
with 3 small lips.

Male 11 to 18 mm. long. Caudal alae (fig. 82) with a sharply de-
limited anterior margin. Ten pairs of caudal papillae, of which 2
pairs are near the sucker and in relation with an anterior long bursal
expansion, 4 pairs of ray-like papillae and 1 pair of sessile papillae
are near the cloacal aperture in relation with a second shorter bursal
expansion, and 3 pairs posterior of these in relation with 2 smaller
expansions anterior of the narrow terminal tail end, according to
Railliet's figure ; Schneider and Fiebiger show 4 pairs in the second
and third groups. The 2 spicules are equal, slender and compara-
tively short, approximately 500//, long in American material, and
apparently 400//. long according to Railliet's figure. The sucker has
chitinous walls, interrupted on the posterior rim.

Female 16 to 23 mm. long. Tail straight, long and slender. Vulva
a little posterior of the middle of the body, surrounded by a cuticulai
thickening. Eggs 59 to 62//, long by 39 to 41//, wide.

Life history. — Probably similar, in a general way, to that of H.
gallinae (p. 54). It is less common in geese that are being ranged
on pasture than in those kept up and fed.

Distribution. — United States (Nebraska), Europe (England,
France, Germany, Italy) and Asia (Russian Turkestan (Aulieata)).

NEMATODE PARASITES OF BIRDS 63

HETERAKIS FARIAI Travassos, 1913

Host. — Odontophoi us capueira.

Location . — Intestine.

Morphology. — Heterakis (p. 50) : Mouth with 3 equal lips. Esoph-
agus about DOCV long, provided with a bulb.

Male 6 mm. long. Caudal extremity with a long tapering pro-
longation (fig. 83). Caudal alae wide. Preanal sucker circular,
about 3(>pi in diameter, with a papilla on its posterior rim. Twelve
pairs of papillae, some of them asymmetrically arranged; 3 pairs
are preanal. Spicules long and unequal. Travassos describes them
as 810/x and 17<V long but his figure does not show any such great
difference in the size. Judging from the position of the cloacal
aperture which he says is 488^ from the posterior end, the shorter
spicule would measure about 900/x and the longer spicule over 1
mm. (roughly 1.3 mm.) in length.

Fig. 83. — Hetkuakis faisiai. Mali: tail. After Travassos, 1913

Female 7 mm. long. Vulva near middle of body. Anus about 1
mm. from the posterior end. Eggs 74/* long by 33/a wide.
Life history. — Probably similar to that of H. gallinae (p. 54).
Distribution. — South America (Brazil).

HETERAKIS HAMULUS Linstow, 1906

Hosts. — Pa/oo cristatus and P. spicifer.

Location. — Cecum.

Morphology. — Heterakis (p. 50) : Head with 3 slightly prominent
lips. Short esophagus terminating in a bulb. Tail pointed. Cuti-
cle finely striated transversely.

Male 7.5 mm. long by 410/x wide. Esophagus length 1/9 of body
length. Right spicule (fig. 84) slender, 370/* long, ending in a hook
distally; left spicule straight, 320/u, long, surrounded by a broad cor-
neous sheath from which only the point projects. Sucker 71/a in
diameter. Two stalked papillae on each side of the sucker. 5 papil-
lae on each side of the cloacal aperture, the posterior pair being more
ventrally located than the others, and 3 pairs of papillae posterior
to these. Tail length is 1/21 of total length.

64 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 7.6 mm. long by 460/* wide. Esophagus % of body length.
Tail length y 8 of body length. Vulva almost in equator of body.
Eggs 57/t long by 42[x wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribution. — Europe (Germany) .

HETERAKIS INTERLABIATA Ortlcpp, 1923

Host. — Rhizothera longirostris.

Location. — Ceca.

Morphology. — Heterakis (p. 50) : Mouth with 3 lips which are cut
in at their base; between each pair of lips a small interlabium.
Dorsal lip with 2 papillae, the other lips with one papilla each.
Lateral alae well developed, conspicuous, commencing a little pos-
terior to the head (fig. 85), extending into posterior half of body.

Male 7 mm. long by 30(V wide. Bursa continuous anteriorly across
ventral surface. Tail with long filiform appendage. Twelve pairs
of caudal papillae of which 3 are preanal, 5 adanal, and 4 postanal.
Spicules equal, 1.25 mm. long, or slightly subequal, the left flanged,
the right tapering to an acute tip (fig. 86).

Female 8 to 9.5 mm. long by 360 to 400[a wide. Tail long and
pointed. Vulva at about middle of body. Eggs oval, 66//, long by
37[x wide.

Life history. — Probabty similar to that of H. gallinae (p. 54).

Distribution. — Asia (Straits Settlement (Malacca)).

HETERAKIS ISOLONCHE Linstow, 1906

Synonym. — Probably Heterakis neoplastica Wassink, 1917 (p. 65).

Hosts. — Crossoptilon manchurianum, Lophophorus impeyanus,
Ithagenes cruentus, Phasianus colchicus, P. chrysomelas, Thaumalea
amherstiae, T. picta, Tragopan satyra.

Location. — Ceca.

Morphology. — Heterakis (p. 50) : Mouth with 3 small lips, each
with 2 papillae. Esophagus region of body with narrow lateral
membranes. Esophagus with bulb.

Male 7.5 to 9 mm. long by 380 to 480^ wide. Esophagus 1/6.8
total length. Tail 1/13 of total length, ending in a long slender point
(fig. 87). Caudal alae continuous anteriorly across ventral surface
in specimens examined by the writer, though this is not shown in the
figures. Preanal sucker 130 to 150/x in diameter, with chitinous ring.
Twelve pairs of caudal papillae, according to Linstow, 11 pairs
according to Lucet and Henry; 2 pairs are in conjunction with the
sucker, 5 or 6 pairs are adanal, and 4 pairs postanal. Spicules equal
(1.41 mm.), according to Linstow; according to Lucet and Henry
subequal, 1.4 to 1.75 mm. long. They have, wide alae.

NEMATODE PARASITES OF BIRDS

65

Female 9 to 12 mm. long by 425 to 470/*, wide. Esophagus 1/8.4,
tail 1/7.3 of total body length; the tail ends in a very long slender
point as in the oxyurids. Vulva a little anterior to middle of body,
dividing the body in a ratio of 22 : 25 ; posterior to it a row of 2 to 3
papillae, and at times a papilla anterior to it. The number and posi-
tion of these vulvar " papillae " are not constant and Baylis and
Daubney (1922) suggest they are due to the action of the sucker of
the male in attempting copulation. Eggs 70 to 75/x long by 42
to 46/* wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

/mm

2/Wfl7

Figs. 84-86. — 84, Heterakis hamulus. Male tail. After Linstow.
1906. 85-86, Heterakis interlabiata. 85. Anterior end; 86, male
tail. Aftejr Ortlepp, 1923

Distribution. — Europe (Germany (Zoological Museum Koenigs-
berg), France, England), Asia (India), and North America (United
States and Canada).

Wassink considers that the specimens described by Lucet and Henry
as II. isolonche are identical with his species, II. neoplastica (p. 65),
but the evidence which he presents, based mainly on the crossing of
the extruded spicules, is not conclusive.

HETERAKIS NEOPLASTICA Wassink, 1917

Hosts. — Phasianus colchicus, P. satscheunensis, Thaumalea obscurus,
and T. pictus.

I ocation. — Ceca.

Morphology. — Heterakis (p. 50) : Head with 3 hemispherical lips,
each lip bearing 2 short papillae. Esophagus cylindrical with pyri-
form bulb. Lateral alae narrow, extending almost the whole length
of the body (to origin of caudal alae in male).

66 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 15 mm. long by 550//, maximum width. Esophagus, including
bulb, 1.9 mm. long; esophagus 79/*, wide; bulb 370/* long by 210/* wide.
Preanal sucker 950/*. from tail end and measuring 140//, in diameter,
its posterior edge interrupted by a depression. Caudal alae (fig. 88)
originating just anterior to sucker and extending almost to tail end.
Cloacal aperture 640/*, from tail end. Twelve pairs of caudal
papillae, their position and relative size identical with those of
Heterakis isolonche: 2 pairs at the level of the sucker, 6 pairs adanal,
and 4 pairs postanal. Two spicules of almost equal length, 1.8 mm.
(the left spicule a few microns longer than the right spicule) ; the
width of the left spicule is 76/*, that of the right spicule 49/*. If
extruded, the spicules cross each other, a condition which is seen in
H. isolonche also, as figured by Lucet and Henry.

Female 17 mm. long by 600/*, maximum width. Total esophagus
2.1 mm. long. Anus 1.2 mm. from tail end. Vulva 8 mm. from
head end; 2 prevulvar and 2 to 3 postvulvar papillae. Vagina very
long. Eggs 65 to 67/* by 35 to 40/*.

Figs. 87-88. — 87, Heterakis isolonche. Male tail. After Linstow, 1006. 88,
Heterakis neoplastica. Male tail. After Wassink, 1917

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — Europe (Holland (Amsterdam)).

This species is very close to H. -isolonche, but because of certain
morphological differences (size of worms, length of tail of both male
and female), and because of an apparent difference in pathogenicity
of the two nematodes, the present writer prefers to regard them as
distinct species at the present time. Spindle-cell sarcoma of the
pheasants listed above is attributed by Wassink to H. neoplastica,
whereas the nodular growths caused by H. isolonche have not been
shown to be of that nature.

Since the above was written, Baylis (1925) has examined TVas-
sink's specimens and has come to the conclusion that this species is
identical with II. isolonche (p. 64).

HETERAKIS LONGECAUDATA Linstow, 1879

Hosts. — Franco/ in its gula/ris, Galloperdi.r x/xulicea, Lophophorus
impeyanus, Megacephalon -maleo, Tragopan satyra.
Location. — Ceca.

NEMATODE PARASITES OF BIRDS 67

Morphology. — Heterakis (p. 50) : Mouth with 3 lips, followed by
a short vestibule. Esophagus 1 8.5 of total length, with well de-
veloped bulb with valve teeth. Lateral alae well developed, extern i-
ing for greater part of length of body.

Male 7.9 to 9.1 mm. long by 240/* wide. Tail (fig. 89) very long
and sharply pointed, 1/19 of total length. Caudal alae wide. Preanal
sucker 80 to 90/* in diameter, 100 to 150/* from cloacal aperture.
Twelve pairs of caudal papillae, of which 8 are postanal and 4 pre-
anal. Left spicule (7'20/i. long) much shorter than right (2.3 mm.
long).

Female 7.9 to 9.6 mm. long by 360/* wide. According to Linstow,
vulva very slightly anterior to middle of body, dividing body length
in ratio of 73:75; according to Baylis and Daubney slightly posterior
to middle, 3.7 to 4.75 mm. from tail end. Tail y s of total length.
According to Baylis and Daubney, there is a conspicuous pair of
caudal papillae at about 680/* from posterior end. Eggs 75 to
78/* long by 43 to 48/* wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution* — Asia (India (Zoological Garden, Calcutta)).

HETERAKIS MACROURA Linstow. 1883

Host. — Megaloperdix n igelii.

Location. — Intestine.

Morphology. — Heterakis (p. 50): Mouth with 2 (?) semicircular
lips. Tail pointed. Esophagus 1/7 of total body length, somewhat
swollen posteriorly.

Male 9 mm. long by 600/* wide. Tail 1/7.5 of total length. Caudal
alae (fig. 90) wide; preanal sucker very large. Ten pairs of papillae,
of which 4 pairs are preanal, 1 pair adanal, 5 pairs postanal. Spic-
ules sharply pointed, 130/* and 110/* long.

Female 12.6 mm. long by 660/* wide. Tail 1/7.8 of total length.
Eggs 72/* long by 48/* wide. Position of vulva not given.

Life history. — Probably similar to II . gallinae (p. 54).

Distribution. — Asia (Turkestan) .

HETERAKIS TAPILLOSA (Bloch, 1782) Railliet. 1885

Synon-yms. — Ascaris papillosa Bloch, 1782; Heterakis montioel-
I'tttna Stossich, 1892; Heterakis stylosa Linstow, 1907.

This species has been greatly confused with Heterakis galUnae
{II. vesicularis) . Railliet and Henry have consequently used the
name Heterakis monticellia/na to differentiate it but that is not
allowable under the rules of nomenclature.

Hosts.— Otis tarda and 0. tetrax.

Location. — Colon and ceca.

68

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Heterakis (p. 50) : The original descriptions of
both Ascaris papulosa and Heterakis monticelliana are inadequate;
that of H. stylosa is more complete. Baylis has examined speci-
mens of both II. monticelliana and H. stylosa and finds them identi-
cal. The combined descriptions follow : Cuticle smooth, very thick ;
head with 3 anteriorly directed lips; esophagus thin, with bulb at
posterior end. Baylis says that, contrary to Stossich's description,
narrow lateral alae are present along almost the whole length of the
body.

Male 9 to 13 mm. long by 620/x wide. Esophagus % of total
length. Caudal alae (figs. 91 and 92) wide, elliptical. Preanal

Figs. 89-92. — 89, Heterakis longecaudata. a, Cuticle ; b, male tail.
After Linstow, 1879. 90, Heterakis macroura. Male tail. After
Linstow, 1883. 91, Heterakis papillosa, Male tail, ventral view.
After Linstow, 1907. 92, Heterakis papillosa. Male tail, lateral
view. After Stossich, 1892

sucker very prominent, 220/x, large. Baylis states that the lateral
view of tail as figured by Stossich shows admirably the highly char-
acteristic shape of the sucker. Tail 1/55 of total length, with styloid
appendage 610/* long. Stossich described 7 pairs of caudal papillae
but evidently overlooked a pair, as Baylis found 8 pairs as described
by Linstow, 1 pair near the sucker, 2 lateral and 1 ventral just
anterior to cloacal aperture, and 4 postanal. Spicules equal, 610/x
long.

Female 15 to 20 mm. long. Esophagus 1/9, tail 1/9 of total body
length; tail finely pointed. Vulva posterior to middle of body; ac-
cording to Stossich, it is at the posterior third ; according to Linstow,
it divides the body length in ratio of 28 :17. Posterior to vulva in
ventral line 3 or 4 mushroom-shaped cuticular bosses.

NEMATODE PARASITES OF BIEDS 69

Life history. — Probably similar to that of H. gallmae (p. 54).
Distribution, — Europe (Germany (Leipzig)) and Asia (Russian
Turkestan).

HETERAKIS PARISI Blanc, 1913

Host. — " Nandou " (=Rhea americana).

Location. — Cecum.

Morphology. — Heterakis (p. 50) : Body cylindrical, attenuated
at both ends. Lateral membranes present, extending from level of
buccal cavity to near the posterior extremity of body. Cuticle with
fine transverse striations. Mouth with 3 small lips, 18/*, high, each
with 2 external papillae. Total length of esophagus 1 mm.; bulb
215/* in diameter.

Fig. 93. — Heteeakis parisi. Male tail. After. Blanc, 1914

Male 7 to 9 mm. long by 350 to 400/x wide. Caudal extremity (tig.
93) with long delicate point. Preanal sucker with chitinous ring,
notched in the median line on its posterior edge. Two bursal alae
with 12 pairs of papillae of which 2 are preanal, 2 adanal, and 8
postanal. Spicules very unequal, the right about 2.2 mm. and the
left 640/1 long, provided with large membranous alae 45/x in trans-
verse diameter.

Female 10 mm. long by 340 to 360/t wide. Tail very slender.
Vulva slightly anterior to middle of body. Eggs 68//, long by 45/i
wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — Europe (France (Dijon)).

HETERAKIS PSOPHIAE Travassos, 1913

Synonym. — Ganguleteralds psophiae (Travassos, 1913) Lane, 1917.
Host. — P sophia viridis.
Location. — Intestine.

70

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Heterahis (p. 50) : Mouth with 3 small equal lips.
Esophagus 1.13 to 1.2G mm. long, with small bulb.

Male 10 mm. long. Cloacal aperture (fig. 94) 352^ from posterior
end. Preanal sucker circular, about 90/x in diameter, with a papilli-
form nodule in its posterior rim. Ten pairs of caudal papillae,
asymmetrically disposed, 3 of them preanal, 7 postanal. Spicules
short, robust, and equal, measuring about 630/a long.

Female 12 mm. long. Anus 800 to 900/x from posterior end. Vulva
(fig. 95) a little posterior to middle of body. Posterior to the vulva,
papilliform cuticular swellings varying in number from 1 to 6
(usually 4) ; at times they are greatly reduced in size. Eggs 52/x,
long by 39ju, wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — South America (Brazil).

Figs. 94-95. — Heterakis psophiae. 94, Male tail.

assos, 1913

95, Vulva. After Trav-

HETERAKIS PUSILLA Linstow, 1906

Hosts. — Gallus gallus and Gallus lafayettU. This species is listed
from the chicken by Gedoelst. Linstow described it from G. lafay-
ettii and there appears to be no definite record from the chicken.

Location, — Ceca.

Morphology. — Heterakis (p. 50) : Head with small rounded lips.

Male 5 mm. long by 190/a wide. Esophagus 1/5 of body length.
Tail (fig. 96) comprises 1/12 of body length and is finely pointed.
Left spicule 530/* long; right spicule 150^, long. Caudal sucker
round. Four pairs large postanal papillae, the interval in the case
of the 2 posterior pairs greater than the intervals for other adjacent
pairs.

Female 5.13 mm. long by 240/* wide. Esophagus length equal to
5/29 of body length. Tail length equal to 2/19 of bod}^ length ; long
and pointed. Vulva 12/17 of body length behind head end. Eggs
thick-shelled, 65/u. long by 3Lu wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — Northern Province (Mamadu). This locality could
not be definitely located but is thought to be in the Uganda Protec-
torate, British East Africa.

NEMATODE PAEASITES OF BIRDS

71

HETERAKIS PUTAUSTRALIS Lane, 1914

Host. — (rail us g alius.

Location. — Ceca.

Morphology. — Heterakis (p. 50) : Small worms, curved dorsally
at the cephalic end. Lateral membranes present, extending almost
the entire length of body.

Male 7.6 nun. long by 300/* wide. Circular sucker (fig. 97), 80/*
wide with papilla interruption on posterior border of chitinous rim.
Twelve pairs of caudal papillae: 2 pairs near the sucker, 6 pairs in
the region of the cloacal aperture, of which the first (most anterior)
of the lateral pairs and the third of these are much the smallest; the

Pigs. <i«;-os. — «.iti, Heterakis pusilla. Male tail. After Dinstow, 1906. 97. Hetera-
kis pi taistkalls. Male tail. 9S, a, Left SPICULE from side; b, left spicule from
• i:\ti;u ; c, RIOHT spicules. Figs. 97 and 98 after Lane, 1914

ventral papillae of this group are fairly far apart; posterior of these
is a pair of medium sized lateral papillae; in the caudal group, the 2
lateral pairs are set far apart, and the most anterior pair is very
small and asymmetrically placed. Right spicule (fig. 98c) 550/*
long and tapering to a blunt conical point. Left spicule (fig. 98 a
and b) 260/* long and provided with lateral alae which are not quite
symmetrical. Caudal alae do not meet as a cuticular thickening in
front of the sucker.

Female 8.2 mm. long by 350/* wide. Vulva just anterior of tiie
middle of the body. Anus 830ju from tip of tail. Eggs G5/a long by
37 n wide.

Life history. — Probably similar to that of 77. gallinae (p. 54).

Distribution.— Asia, (India (Bengal)).

HETERAKIS SKRJABINI Cram, 1327, new name

Synonym. — Heterakis drquata Schneider. L866, of Skrjabin, 1910.
Hoxt. — Tina in us, species.
Location. — Abdominal cavil v.

72

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Heterakis (p. 50) : Color gray-white. Cuticle with
fine transverse striations.

Male 23 to 25 mm. long by 700/* wide. Esophagus 1.125 mm. long,
its cylindrical part 238/* wide; bulb 272/* long by 300/* wide. Tail
(fig. 99) pointed, provided with caudal alae; cloacal aperture 420/i
from posterior end. Preanal sucker 230/* in diameter, its posterior
border 630/* from the caudal extremity. Thirteen pairs of caudal
papillae of which 2 are preanal, 7 adanal, 4 postanal. Spicules equal,
850/* long, whip-shaped, the anterior thicker end resembling the
handle of the whip and the exceedingly fine and delicate posterior
or free end the whip itself.

Female 25 mm. long by 1 mm. wide. Esophagus, 1.125 mm. long,
its cylindrical part 200/* wide; bulb 255/* long by 272/* wide. Tail
conical, anus 220/* from posterior end. Vulva in anterior part of
body, 4.42 mm. from anterior extremity. Eggs oval, 76.5 to 85/* long
by 47/* wide.

Life history. — Probably similar to that of H. gallinae (p. 54).

Distribution. — South America (Paraguay).

At the time that Skrjabin identified the specimens from Ti7iamus,
species of Paraguay as Heterakis arquata, he had not seen Travassos's
paper (1913) but based his identification oif Schneider's scant de-
scription. Travassos gave a detailed description of Heterakis ar-
quata, agreeing in all particulars with Schneider's description, and
the specimens described by Travassos were from the type locality and
as far as can be ascertained, from the type host, Crypturus cupreus.
Skrjabin's description differed from that of Schneider in the number
of caudal papillae, and from that of Travassos in numerous very
marked respects, as seen below. It has been thought advisable by
the present writer, therefore, to give a new name to the species de-
scribed by Skrjabin.

A comparison of H. arquata as described by Schneider and Tra-
vassos, and of H. skrjabini as described by Skrjabin is as follows:

H. arquata

H. skrjabini

Preanal sucker (diameter) _
Cloacal aperture, from end

of tail.
Spicules

Caudal papillae

Anus of female from tail

end.
Vulva

Eggs

90 M

270 M -

Very long (according to
figuie over 2 mm.)

12 pairs

1.12 mm

A little anterior to mid-
dle of body.
52m by 29m

230m.
420m.

Relatively short (850m).

13 pairs.
220m.

In anterior fifth of body.
76.5m to 85m by 47m-

NEMATODE PARASITES OF BIRDS
HETERAKIS TENUICAUDA Linstow, 1883

73

Hosts. — Caccdbis petrosa and Perdix graeca.

Location. — Intestine.

MorpJtology. — HeteraJcis (p. 50) : Head rounded, with 3 slightly
projecting lips. Esophagus 1/10 or 1/8 of total length. Lateral alae
present, extending to caudal region in both sexes.

Male 22 mm. long by 1 mm. wide, according to Linstow, or 17.8
mm. long by 552/* wide, according to Seurat. Tail length given as
1/61 and 1/42 of total length by the two authors respectively, with
a fine sharply pointed appendage 215/* long. Preanal sucker 290/*
in diameter, located 450/* anterior to the cloacal aperture in Seurat -'s

Figs. 99-101. — 99, Heterakis skrjabi.m. Male tail. After Skrjabin, 1916. 100.
heterakis tenuicauda. male tail. after llnstow, 1883. 101, male tail. after
Secrat, 1918

specimen; its cutinous (chitinous) rim 50/* wide. Twelve pairs of
caudal papillae, according to Linstow (fig. 100) ; Seurat describes
only 12 but his figure (fig. 101) shows 13, there being an additional
pair at the sides of the sucker. The most anterior pair very small.
at level of origin of caudal alae (thus considerably anterior to
sucker). Spicules very wide, 980 and 9G0/* long, according to Lin-
stow, or 890/* according to Seurat.

Female 14 mm. long by G00/* wide, according to Linstow (this
being surprising as it is much smaller than the length given by him
for the male) or 19.5 mm. long by GT2/* wide according to Seurat.
Tail awl-shaped, 1/13 to 1/11 of body length. Vulva 10.1 mm. from

74

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

the head end in Seurat's specimen. Vestibule 5.5 mm. long; un-
paired trompe 1.8 mm. long. Eggs 60 to 63/* long by 40 to 42/* wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribution. — Asia (Turkestan) and Africa (Algeria (Medea)).

HETERAKIS VALVATA Schneider, 18G6

Hosts. — Grypturus cupreus and C. nocticagus.

Location. — Ceca.

Morphology. — Ileterakis (p. 50) : Mouth with 3 small equal lips.
Lateral membranes present. Esophagus with bulb about 217/* in
diameter; slender part about 1.45 mm. long.

Male 10 mm. long. Caudal alac small. Preanal
sucker 90/* in diameter, with papilliform nodule on
its posterior edge. The anterior lip of the cloacal
aperture, which is 170//, from the posterior extremity,
projects as a triangular valve, from which the
species derives its name. Schneider described and
figured (fig. 103) only 11 pairs of papillae, but
according to Travassos (fig. 102) there are 13 pairs.
Spicules unequal, 2.6 mm. and 1 mm. long.

Figs. 102-103. — Heterakis valvata. 102, Male tail. After Travassos, 1913. 103,
Male tail. After Schneider, 1860

Female 12 to 15 mm. long. Vulva slightly posterior to middle of
body, with 2 papilliform projections just below it. Anus 780/* from
posterior end. Eggs 74/* long by 37/* wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribution, — South America (Brazil).

(?) HETERAKIS VALDEMUCRONATA (Molin, 1860) Stossieh, 1887

Synonym. — Ascaris valdemueronata Molin, 1860.

Hosts. — Ciconia maguari and Euxenura maguari.

Location. — Proventriculus.

Morphology. — ? Heterakis (p. 50) : Head with alae: mouth with 3
lips. Body attenuated anteriorly.

Male 7 mm. long. Posterior extremity bent, acutely conical, with
long mucron. Anterior to the cloacal aperture a depression with
saucer-shaped muscular sucker.

Female 12 mm. long by 200/* wide. Posterior extremity obtuse,
with mucron.

NEMATODE PARASITES OF BIRDS 75

Distribution. — South America (Brazil).

Railliet and Henry do not list this species in the Heterakidae and
Stossich and Travassos say it is a doubtful species; the evidence
seems adequate, however, for its place in the Heterakinae and the
present writer is leaving it in this genus.

HETERAKIS BONASAE, new species

Host. — Bonasa umbellus.

Location. — Ceca.

Morphology,. — Hete?*akis (p. 50) : Head with 3 lips. Buccal cav-
ity present. Esophagus with bulb at posterior end. Lateral alae
present, very narrow (16.6/*), extending from a point about 133/*
posterior to the cephalic end, to the anal region of the female and
to the region of the anterior end of caudal alae of male.

Male 7 nun. long by 365/* maximum width. Buccal cavity 41.5/*
deep: esophagus, including bulb, 1.0 mm. long; bulb 216/* in diameter.
Tail (fig. 104) ending in a slender appendage 158/* long; bursal alae
wide. Cloacal aperture 415/* from posterior end. Preanal sucker
prominent; its posterior rim 116/* anterior to cloacal aperture and
interrupted by a papilla in the median line; internal diameter of
sinker 83/*, external diameter 100 to 120/*. Thirteen pairs of caudal
papillae, of which 5 are preanal, 1 adanal. and 7 postanal. Spicules
unequal, the longer 1.4 to 1.6 mm., the shorter 1.1 to 1.3 mm. long;
both alate. 37/* wide.

Female 7 to 8 mm. long by 365 to 398/* wide. Buccal cavity 50/i
deep; esophagus, including bulb, 1.1 mm. long; diameter of bulb
249/*. Anus 830 to 860/* from posterior end. Tail slender, sharply
pointed. Vulva (fig. 105) directly at middle of body or slightly
posterior to middle (dividing body length in ratio of 10:9). the
vulvar lips being slightly salient, in some cases the under lip
markedly so. Eggs 75 to 83/* long by 42/* wide.

Life history. — Probably similar to that of II. gallinae (p. 54).

Distribitfio'n.— North America (United States (Pennsylvania)).

Type material.— Nbi 2637$, l.S.X.M. (Bureau of Ajiimal Industry
Ilclniinthological Collections ) .

I am indebted for this material from the ruffed grouse to Dr. E. L.
Brunett, Cornell [Jniversity.

HETERAKIS NATTERERI Travassos, 1923

Etost. ' '/>>■'■ hhuiicnhiicii'i.

Location. — Ceca.

Morphology. — Heterakis (p. 50): Head with X lips, vestibule 2
to 28/* deep: total length of esophagus 600/*; pyriform bulb L70/* in

diameter, with heavilv chitinized valves.

76

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 4 mm. long by 300/* wide. Caudal alae regularly developed.
Caudal papillae number 13 pairs, arranged as follows: 3 pairs of
preanal papillae (1 at level of anterior and 1 at level of posterior
border of sucker, pedunculate; 1, sessile, just anterior to cloacal aper-
ture) ; 2 pairs of adanal papillae (1 pair situated externally
(laterally?), with long slender pedunculations, the other internally
(ventrally ?) with short pedunculations) ; 8 pairs of postanal papillae
(3 large, lateral, just posterior to cloacal aperture, 2 ventrals corre-
sponding to the laterals in position and, near the extremity, 3 pairs

0-3/77/7?

Figs. 104-107. — Heterakis bonasae. 104, Male tail, a, Ventral view ; &,
lateral view. 105, vulva. heterakis acdticauda. 106, male tail.
107, Female tail. Nos. 106-107 after Cobbold, 1861

which vary in shape and position, some of them being absent at times,
or on the other hand, folds of the alae simulating additional papillae
above this posterior group). Preanal sucker 78/* in diameter, situ-
ated about 42/* from the cloacal aperture. Spicules subequal, about
1 mm. long, their proximal ends slightly dilated. Gubernaculum
absent. Cloacal aperture 160/* from caudal end, which is a slender
point about 71/* long.

Female 4.2 mm. long by 400/* wide. Vulva 2 mm. from caudal
extremity, thus situated more or less at middle of body. Ovejector

NEMATODE PARASITES OF BIRDS 77

short, directed anteriorly ; posterior branch of uterus extends to point
300/x from the anus, anterior branch to 300/a from the esophageal
bulb. Eggs 63/*, long by 35/* wide. Tail conical, elongated; anus
300/* from extremity.

Life history. — Probably similar to that of H. gallmac (p. 54).

Distribution. — South America (Brazil (Matto Grosno)).

HETERAKIS ACUTICAUDATA (Cobbold, 1861) Travassos. 1918b

Synonym. — Strongylus acuticaudatus Cobbold, 1861.

Host. — Chloephaga poliocephala.

Location. — Ceca.

Morphology. — Head obtuse, bare. Male 12 to 19 mm. long. Tail
sharply pointed. Caudal alae (fig. 106) subterminal, large, each ala
bearing 5 ray-like papillae. Female about 12 mm. long by 333/x wide.
Tail (fig. 107) straight, pointed.

Distribution. — Europe (London Zoological Garden).

Xo further stud^y of specimens of this nematode has been made
since the original and unrecognizable description by Cobbold.
Travassos has transferred the species to the genus Heterakis, evi-
dently basing this transfer on the character of the male tail, and more
especially the preanal sucker, as figured by Cobbold. However, that
Cobbold's drawings may be misleading is seen in the case of Stron-
gylus spiculatus, described and figured by him at the same time as
Strongylus acuticaudatus. Both figures show the male tail of
Heterakis sensu stricto; however, Cobbold at a later date (1879)
wrote that Strongylus spiculatus was identical with his earlier species
Ascaris strongylina, which in more recent studies has been placed in
the genus Subulura. The allocation of Strongylus acuticaudatus,
therefore, is inadvisable except to leave it in Heterakis sensu lato.

Genus ASCARIDIA Dujardin, 1845

Generic diagnosis. — Heterakinae (p. 49) : Mouth with 3 lips.
Esophagus club-shaped, without a bulb. Two lateral membranes
usually present. Male with caudal alae feebly developed; spicules
usually equal or subequal. Gubernaculum absent. Preanal sucker
slightly salient, rounded, with chitinous ring; papillae relatively
large. Female with vulva near middle of body; uteri divergent.
Eggs with thick shell, with a clear granulation inside the shell at
one of the poles.

Parasitic in intestine (especially the small intestine) of birds and
possibly of reptiles and fish.

Type-species. — Ascaridia hermaphrodita (Froelich, 1789) Railliet
and Henry, 1914.

78 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

KEY TO SPECIES OP ASCARIDIA

1. Inadequately described; female 85 mm., male 53 mm. long; from Gallus

gallus, Australia Ascaridia compressa, p. 8S.

Description more adequate than above 2.

2. Male with 13 pairs of caudal papillae, in 2 unusually closely-set rows;

spicules equal, 630/* long by 2Afi wide; from Macropygia nighrostris.

Ascaridia australis, p. 84.

Male usually with less than 13 pairs of papillae; if has 13 pairs, spicules

different from above ; from other hosts than above 3.

3. Spicules described as unequal 4.

Spicules equal or subequal 5.

4. Female 39 mm., male 40 mm. long ; 9 pairs of caudal papillae ; from Capri-

mulgus campestris Ascaridia amblymoria, p. 83.

Female 55 mm., male 43 mm. long ; caudal papillae number 10 pairs plus
an occasional unpaired papilla ; from Tetrao urogallus.

Ascaridia cylindrica, p. 90.

5. Male with a total of not more than 18 caudal papillae 6.

Male with a total of 20 or more caudal papillae 16.

6. No preanal caudal papillae but 8 pairs of postanals.

Ascaridia dolichocerca, p. 90.
Preanal caudal papillae present < 7.

7. Male 36 to 50 mm. long or longer ; female 62 to 100 mm. long 8.

Male not over 31 mm. long; female not over 60 mm. long 10.

S. Esophagus 1/14 of total body length; 4 pairs of preanal caudal papillae;

tail of female 1/65 of total body length Ascaridia compar, p. 88.

Esophagus 1/12.8 to 1/12.3 of total body length; 3 pairs of preanal caudal
papillae ; tail of female not described in one species, 1/133 of total body

length in other species 9.

9. Female 90 to 100 mm. long ; male 50 mm. or longer ; ventral surface of
male tail covered with conspicuous granulations.

Ascaridia styphlocerca, p. 100.
Female 62 mm. long; male 40 mm. long; male tail without granulations.

Ascaridia francolina, p. 92.

10. Male with 6 or 7 pairs of postanal papillae 11.

Male with not more than 5 pairs of postanal papillae 14.

11. Spicules 570/i long; male tail with granulations in region of sucker.

Ascaridia granulosa, p. 92.

Spicules 1 mm. or longer ; male tail without granulations in region of

sucker 12.

12. Body not described as having lateral alae : spicules 3.63 mm. long; tail

of male 1/77 of total length ; eggs 96/u. long by 57/a wide.

Ascaridia magnipapilla. p. 95.

Body with lateral alae; spicules not over 2.1 mm. long; tail of male 1 52

to 1/46 of total body length; egg sizes not given 13.

13. Only male known, 30 mm. long: papillae scattered over the whole body;

spicules subequal, 2.1 and 1.9 mm. long Ascaridia longecirrata, p. 94.

Both male and female known. Male 21 mm. long ; no papillae described
on body as above; spicules equal, 1.05 mm. long.

Ascaridia catheturina, p. 85.

14. Vulva posterior to middle of body, dividing body length in ratio of 15:11;

spicules 790,u long Ascaridia aegyptiaca, p. 83.

Vulva slightly anterior to middle of body ; spicules 1.7 mm. or longer 15.

NEMATODE PARASITES OF BIEDS 79

35. Female tail 1/35, male tail 1/70 of total body length; eggs 91/t long by
57/* wide; spicules subequal, 2.06 and 2.37 mm. long.

Ascaridia cordata, p. 88.
Female tail 1/67, male tail 1/48 of total body length : eggs 73/* long by

42/* wide ; spicules equal, 1.74 mm. long Ascaridia circularis, p. 86.

1G. From Qallus (jaUus; male only, 24 mm. long by 600/* wide; preanal sucker

110/t in diameter Ascaridia brasiliensis, p. 84.

From other hosts than above or, if from Galht.s <j<iUii,s, total length of male
30 mm. or more and sucker over 200/t in diameter 17.

17. Spicules 1.87 mm. long, their middle part enlarged to form unilateral

wings on the face of which are 10 to 12 small projecting teeth.

Ascaridia hermaphrodita, p. 80.
Spicules of different length or different shape, or both, from above 18.

18. Male 45 mm. long ; 7 pairs of postanal papillae ; caudal sucker 340/* in

diameter; spicules 3.6 mm. long Ascaridia borealis, p. 84.

Male not agreeing with above in all particulars 19.

19. Body with 26 to 30 pairs of cervical papillae 20.

Body without numerous cervical papillae, or noue described 21.

20. Tail of female 1/78 of total body length ; spicules 950/* long by 42/* wide.

Ascaridia cristata, p. 89.

Tail of female 1/30 of total body length, or longer; spicules 1.2 to 1.9

mm. long Ascaridia columbae, p. S6.

21. Male only known ; 9 pairs of postanal papillae ; spicules 820/* and 827/*

long Ascaridia anseris, p. 101.

Both male and female known ; male with less than 9 pairs of postanal
papillae ; spicules, if length given, different from above 22.

22. Tail of female 1/117 of total length ; eggs 172/* long by 146/* wide

Ascaridia stroma, p. 99.

Tail of female, if length given, longer than above; eggs considerably

smaller than above 23.

23. Vulva posterior to middle of body 24.

Vulva at middle or anterior to middle of body 26.

24. Male with 12 pairs of caudal papillae Ascaridia orthocerca, p. 97.

Male with 10 pairs of caudal papillae 25.

25. Spicules equal, 3 mm. long; eggs 9S/* by 53/* Ascaridia numidae, p. 96.

Spicules subequal, 1.95 and 1.97 mm. long; eggs SI/* long by 47/t wide

Ascaridia trilabium, p. 101.

26. Male and female both 41 mm. long. Vulva at anterior third of body. In

Penelope humeralis Ascaridia serrata, p. 98.

Total length usually greater than above; vulva usually more posterior than
above. In other hosts than above 27.

27. Preanal sucker not over 162/* in diameter 28.

Preanal sucker 200/* or more in diameter 29.

28. Male with tail 1/80 of body length and with 1.! pairs of caudal papillae;

sucker 162/* in diameter; spicules 1.66 mm. long

Ascaridia magalhaesi, p. 95.

Male with tail 1/55 of total Length and with 10 pairs of caudal papillae;

sucker 100/* in diameter; spicules 00!)/* long__Ascaridia pterophora, p. 07.

29. Female not over 52 mm. long; eggs 68/f long; preanal sucker of male 340/*

in diameter; spicules 3.4 mm. long Ascaridia strelnikowi, p, 99.

Female 60 nun. of longer; eggs 75/* or longer, except possibly in A. Un-
eata; preanal sucker not over 250/* in diameter; spicules shorten- than
3.4 mm. except in A. galli where they may possibly he 4 mm. long— 30.
3012—27 7

80 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

30. Body without lateral alae but with prominent white lateral lines with a

rope-like twist ; male with 10 pairs of caudal papillae ; preanal sucker

not over 217/t in diameter Ascaridia lineata, p. 93.

Body with lateral alae of usual type ; male with 10 to 13 pairs of caudal
papillae ; preanal sucker 220 to 250m in diameter 31.

31. Cuticle with marked annulations which may be 100fi wide ; lateral alae 3

to 4 mm. long by 300//, wide; spicules 1.9 mm. long.

Ascaridia fasciata, p- 91.
No annulations of cuticle described ; lateral alae very slender, extending
throughout whole body length ; spicules longer than above.

Ascaridia galli, p. 81.

ASCARIDIA HERMAPHRODITA (Froelich, 1789) Railliet and Henry, 1914

Synonyms. — Ascaris hermaphrodita Froelich, 1789; Fusaria trun>-
cata Zeder, 1803; Ascaris truncata (Zeder, 1803) Rudolphi, 1809;
Ascaridia truncata (Zeder, 1803) Dujardin, 1845; Heterakis trun-
cata (Zeder, 1803) Schneider, 1866.

Hosts. — Amazona ochrocephala, Chrysotis festiva, Conurus pavua,
C. solstitialis, Pionus (Psittacus) aestivus, P. aracanga, P. ararauna,
P. do?ninicensis, P. festivus, P. kucoc, P. leucotis, P. menstruus, P.
pulverulentus, P. pertinax, P. phoenieurus, P. purpurevs, P, species,
P. sulfureus, P. vinaceus.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Mouth with 3 strongly developed
lips of almost equal size. Esophagus gradually enlarging posteriorly
but without forming a bulb. Distinct but delicate lateral membranes
present, according to Froelich and Schneider, apparently throughout
the whole length.

Male 29 mm. long, according to Schneider; 19 mm. long by 750/u.
wide, according to Skrjabin. Cloacal aperture 323/* from posterior
end (in 19 mm. specimen). Caudal alae feebly developed. Preanal
sucker 220/*, in diameter, provided with a chitinous ring. Schneider
described 11 pairs of papillae but figured 13 (fig. 108) ; according to
Skrjabin (fig. 109), this latter number is correct, 6 pairs being pre-
anal and 7 pairs postanal, of unequal size and irregular distribution.
Spicules equal, 1.87 mm. long, of characteristic shape; starting as a
handle with marked transverse striations, the middle part of spicule
follows with smooth edges, but enlarged to form a unilateral wing
on the face of which are found 10 or 12 small projecting teeth; distal
part of spicule slender and with rounded end.

Female 35 mm. long. No further description.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Europe (Germany (Leipzig, P. menstruus, prob-
ably in captivity) and Russia), and South America (British Guiana
and Brazil).

NEMATODE PARASITES OF BIRDS

81

ASCARIDIA GALLI (Schrank, 1788) Freeborn, 1923

Synonyms. — Ascaris galli Schrank, 1788 ; Ascaris gallopavonis
Gmelin, 1790; Ascaris perspicillum Rudolphi, 1803; Ascaris gibbosa
Rudolphi, 1809; Fusaria inflexa Zeder, 1800; Ascaris inflexa (Zeder,
1800) Rudolphi, 1809; Ascaridia inflexa (Zeder, 1800) Dujardin,
1845; II eterakis inflexa (Zeder, 1800) Schneider, 1866; II. perspicil-
lum (Rudolphi, 1803) Railliet, 1893; Ascaridia perspicillum (Ru-
dolphi, 1803) Dujardin, 1845.

Hosts. — Anus acuta, A. boschas domestica, A. b. fera, A. moschata,
Cairina moschata, Gallus gallus, Ithagenes cruentus, Numida mele-
agris. Phas-ianus gallus, Tetrao bonasia, T. urogallus, and, rarely, cat.

Figs. 108-109.

-Ascaridia hekmaphrowta. 108, Male tail. After Schneider,
18G6. 109, Male tail. After Skrjabin, 1917

Location. — Small intestine ; in the large intestine, esophagus, crop,
gizzard, egg or oviduct, and body cavity as wandering parasites.

Morphology. — Ascaridia (p. 77) : Large, yellowish-white worms,
with 3 lips of which the dorsal is larger than the 2 submedian; 3
dentigerous ridges on each lip. Lateral alae slender, thruout whole
length of body. Esophagus without bulb.

Male 30 to 80 mm. long. Tail 1/50 of total body length, according
to Dujardin, its end obliquely truncated and with a narrow bursal
membrane on each side. There arc 10 pairs of caudal papillae (figs.
110 and 111), of which 3 pairs of pedunculated papillae are near the
sucker; this sucker (220/x. in diameter) has a cutinous (chitinous)
wall, which is interrupted posteriorly, according to Schneider ; Smit
figures it as uninterrupted and followed by a median sessile papilla.
Three pairs of pedunculated and 2 pairs of sessile papillae lie beside
and just behind the cloacal aperture, and 2 pairs lie in a group still

82 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

farther back in relation with the bursal expansion preceding the
portion which narrows to the point of the tail. Schneider refers to
9 pairs and figures 9 in his diagram and 12 pairs with 1 extra and
variable papilla in his plate figure. The spicules are subequal, alate,
and about 4 mm. long, according to some writers, or the short one
2 mm. long and the other 2.5 mm. long, according to Smit ; they ter-
minate in small buttonlike enlargements.

Female 60 to 120 mm. long. Tail 1/50 to 1/60 of body length,
its end straight, conical, and mucronated. Vulva in the anterior por-
tion of the body, dividing body length in ratio of 38:49. Eggs
ellipsoidal, 75 to 80/* long by 45 to 50/* wide, not segmenting, as a
rule, when oviposited.

Life history. — A female worm may contain over 1,200 fertilized
eggs, or, according to Danheim, approximately 1,500. With suffi-
cient moisture, these eggs will develop to a point where each egg
contains an infective embryo in 9 or 10 days, when incubated at a
temperature of 28° C. ; at less favorable temperatures this develop-
ment may require weeks. Hatching normally occurs after these in-
fective eggs are swallowed by suitable fowls, though an occasional
egg hatches without being swallowed. Within 28 hours after swal-
lowing infective eggs, according to Ackert, the fowl shows young
worms free in the small intestine. Ackert has reported finding the
larvae penetrating the intestinal mucosa to some extent. According
to the Oklahoma Experiment Station, the larvae migrate to the lungs
and here they reach a length of 2 to 3 mm. They then ascend the
windpipe and are swallowed, completing their development in the
intestine. In 30 days they are half-grown. The possibility that
all or part of the young worms may have a migratory phase similar
to that of Ascaris lumbricoid,es was suggested by the fact that Ackert
found 7 larvae in the lungs, 1 in the trachea, and 2 in the liver in
feeding experiments. Ransom and also Schwartz do not find evi-
dence of larval migration. Ova will survive 7 days exposure to sun-
light at 23° to 33° C. or 15 hours freezing at -11.6° to — 8°C. Scott
has pointed out that the earthworm (Helodrilus parvus) may be a
means of spreading the eggs.

Distribution. — Cosmopolitan (North and South America, Europe,
Africa, Asia, and Australia).

Schwartz (1925) has pointed out, after a study of specimens from
various parts of the United States, that it is not Ascaridia galli which
is found commonly in chickens in this country, but A. lineata.
Therefore, the life history experiments cited above and other reports
of this species in the United States, probably should be referred to
A. lineata.

NEMATOPK PAKASITKS OF BIRDS

83

ASCARIDIA AEGVPTIACA (Linstow, 1902) Kailliet and Henry, 1914

Synonym.— Heterakis aegyptiaca Linstow, 1002.

Host. — Ardea garzetta.

Location. — Not given.

Morphology. — Ascaridia (p. 77) : Cuticle with cross-striations;
mouth with 3 large semicircular lips.

Male 24 mm. long by 590/t wide. Esophagus 1/13.6, tail 1/60 of
length. Spicules 790/*, long. At each side of the preanal sucker
2 papillae; between the sucker and the cloacal aperture 1 pair;
5 pairs of postanal papillae, thus making a total of 8 pairs of caudal
papillae. (Fig. 112.)

Female 60 mm. long by 1.18 mm. wide. Esophagus 1/15, tail 1/36
of total length; tail conical. Vulva posterior to middle of body,
dividing body length in ratio of 15 : 11. Eggs 78jn long by 48/x wide.

3 mm

Figs. 110-113. — 110, Ascakidia galli. Male tail. After S.uit, 1022. Ill, Male tail.
Aftlr Baylis am* Daubney, 1022. 112, Ascaridia aegyptiaca. Male tail. Afteb
Linstow, 1002. 113, Ascakidia amblymoria. Male tail. After Drasche, 1S82

Life history. — Probably similar to that of A. galli (p. 82).
Distribution. — Africa ( Egypt ) .

ASCARIDIA AMBLYMORIA (Drasche, 1883) Railliet and Henry, 1914

Synonym. — Heterakis amblymoria Drasche, 1883a.
Host. -Caprimidgus campestris.

L ocation . — Tntest ine.

Morphology. — Ascaridia (p. 77) : Mouth with 3 semicircular lips
provided with Laminae or "tooth plates" (Zahnplatten). Dorsal
lip with 2 large papillae.

Male 40 mm. long by 1.25 mm. wide. Preanal sucker large, more or
less circular, and provided with a eutinous (chitinous) ring. Nine
pairs of papillae, 3 of them preanal, and in addition a small papilla
on posterior rim of sucker (fig. 113). Spicules unequal.

84 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

%

Female 39 mm. long by 1 mm. wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution.South. America (Brazil).

ASCARIDIA AUSTRALIS (Linstow, 1898) Railliet and Henry, 1914

Synonym. — Heterakis australis Linstow, 1898.

Host. — Macropygia nigrirostris.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : Cuticle with cross-striations.
Specimens imperfect so that total length could not be determined.
Both sexes with finger-like appendage.

Male with large preanal sucker with cutinous (chitinous) rim;
13 pairs of large closely set papillae (fig. 114). Spicules 630/* long
by 2.4/* wide, with rounded ends.

Eggs thick-shelled, 75/* long by 42/* wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Oceania (Bismarck- Archipelago (Ralum)).

ASCARIDIA BUREAUS (Linstow, 1884) Railliet and Henry, 1914

Synonym. — Heterakis borealis Linstow, 1884.

Hosts. — Lagopus mutus and Tetrao lapopus.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) ;

Male only, 45 mm. long by 1.3 mm. average width. Preanal
sucker large, 340/* in diameter, its posterior rim interrupted by a
papilla (fig. 115) ; from the sucker there radiate out to the edge of
the body wide lines, evidently muscular structures. Ten pairs of
papillae, 1 pair very large, projecting at an angle at a level just
anterior to the cloacal aperture; 2 other pairs preanal and 7 pairs
postanal, all quite small. Spicules 3.6 mm. long, the left slightly
curved, the right described and figured as bent hook-like.

Female unknown.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Asia (Siberia (Kamtschatka) ).

ASCARIDIA BRASILIENSIS (Magalhaes, 1892) Railliet and Henry, 1912

Synonyms. — Heterakis brasiliensis Magalhaes. 1892.

Hosts. — Gallus gallus.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : Yellowish worms, thinned to-
ward the ends, especially posteriorly. Mouth with 3 large unequal
lips, each with distinct submedian papillae. Esophagus 2 mm. long.

Male 24 mm. long by 600/* wide. Sucker circular, 90//, in internal
diameter, 110/* in external diameter, with chitinous rim interrupted

NEMATODE PARASITES OF BIRDS

85

by a papilla on the posterior rim. Caudal membranes narrow.
Two slightly unequal spicules. Ten pairs of papillae. (Fig. 116.)
(Magalhaes says there are 9, but Travassos notes that his detailed
description totals 10 and he figures 10.) One pair is anterior of the
sucker, 2 very close together behind the sucker, 1 near the cloacal
aperture, 2 just behind the cloacal aperture, 1 pair some distance
posterior of these, then 2 small pair close together, and finally a pair
near the extremity of the tail; between the last 2 is an asymmetrical
median papilla. (The foregoing does not agree with the figure
reproduced by Travassos.) Travassos states that the foregoing is

'/smrri

Figs. 114-117. — 114, Ascaridia australis. Male tail. After Linstow, 1898. 115.
ascaridia borealis. male tail. after llnstow, 1s84. 116, ascaridia brasiliknsis.
Male tail. After Magalhaes, 1892. 117, Ascaridia catheturina. Male tail.
After Johnston, 1912

probably a description of a young male of A. lineata (p. 93) and
this appears very probable; according to him the asymmetrical
papilla is a refringent artefact.

Female unknown.

Life history. — Probably similar to that of A. Galli (p. 82).

Distribution. — South America (Brazil).

ASCARIDIA CATHETURINA (Johnston, 1912) Raillict and Henry, 1914

Synonym. — Heterakis catheturinus Johnston, 1912.

Host. — Taleg alius (Catheturvs) latJiami.

Location. — Intestine.

Morphology. — Ascaridia (p. 77): Anterior end rounded, narrow;
posterior end pointed in both sexes. Lateral alae extend for about
2 mm. posterior to mouth. Three equal lips.

Male 21 mm. long. Caudal alae narrow. (Fig. 117.) Cloacal
aperture 400/x from posterior end. Preanal sucker 275/z anterior to
cloacal aperture. One small median unpaired and 2 prominent

86

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

paired preanal papillae. One small unpaired and G paired postanal
papillae, thus a total of 18 caudal papillae. Spicules slightly curved
and pointed, 1.05 mm. long by 22/x wide.

Female 28 mm. long. Vulva at about middle of body, slightly
salient. Anus 86G> from posterior end.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Burnett River (Australia?).

ASCARIDIA CIRCULARIS (Linstow, 1903) Raillict and Henry, 1914

Synonym. — Heterakis circularis Linstow, 1903.
Host. — Centropus sinensis.
Location. — Not given.

Morphology. — Ascaridia (p. 77) : Mouth with 3 large semicir-
cular lips, a dorsal with 2 papillae, the other 2 ventro-lateral.

5mm

Pigs. 118-120. — 118, Ascaridia circulahis. Male tail. After Linstow, 1903. 119
Ascaridia columbae. Male tail, a, Ventral view ; 8, side view. After Johnston
1918. 120, Ascaridia compar. Male tail. After Linstow, 1899

Male 31 mm. long by 790/a wide. Esophagus 1/17, tail 1/48 of
total length. Preanal sucker large. Eight pairs of large papillae
(fig. 118), of which 3 are preanal, 5 postanal. Spicules 1.74 mm.
long.

Female 52 mm. long by 1.07 mm. wide. Esophagus 1/14, tail 1/67
of total length. Vulva slightly anterior to middle, dividing body
in ratio of 12G : 129. Eggs 73/i, long by 42^ wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Asia ( Siam ) .

ASCARIDIA COLUMBAE (Gmelin, 1790) Travassos, 1913

Synonyms. — Ascaris columbae Gmelin, 1790; Ascaris maculosa
lludolphi, 1802; Heterakis maculosa Schneider, 1866; Heterakis
columbae (Gmelin, 1790) Railliet, 1885.

NEMATODE PARASITES OF BIRDS 87

Hosts. — Columba arquatrix, 0. domestica, G. d. laticaitda, C.
gutturosa, C. licia. ('. picui, C. risoria. C. speciosa, C. talpacoti,
Crocejms phoenicopt&nis, PMogoenas luzonica, Stictoenas arquatrix,
Turtur sylvaticits.

Chatin has reported this species from the pheasant and Sweet from
the chicken; Skrjabin thinks these writers probably had Ascaridia
galli {A. perxpicillum).

Location. — Small intestine, usually; in esophagus, proventriculus,
gizzard, liver, and body cavity, occasionally.

Morphology. — Ascaridia (p. 77): White translucent worms,
thinning towards the ends. Mouth with 3 subequal lips; dorsal lip
provided with 2 small papillae; a transverse fold of cuticle just
behind lateral lips. Anterior extremity provided with 2 semi-
elliptical cervical membranes. According to Ba} T lis and Daubney,
there are 26 to 30 pairs of cervical papillae, the first 2 or 3 pairs in
the cervical membranes, the others posterior to this. Nerve ring
f.OO/z posterior to head end.

Male 16 to 31 mm. long, according to some writers; Johnston says
it is up to 40 mm. long; Baylis and Daubney say 60 to 70 mm. long
by 1.1 mm. wide. This is a considerable variation, possibty corre-
lated with host variation. Tail end obliquely truncated and
mucronated at the tip. Circular sucker, with chitinous walls, 150 to
200/x. long by 150 to 160/a wide. There is a small ala on each side of
the tail. According to Travassos, and to Baylis and Daubney, there
are 14 pairs of caudal papillae, of which 5 pairs are postanal; an-
terior of the sucker, 2 pairs on each side. The number and arrange-
ment is somewhat variable. (Schneider says there are 10 pairs of
papillae and figures 11 on one side and 12 on the other; Johnston
(fig. 119) finds 13 or 14 pairs.) Spicules equal, 1.2 to 1.9 mm. long.

Female 20 to 37 mm. long by 1.3 to 1.6 mm. wide, according to
some writers; Johnston says it is up to 55 mm. long. Neumann says
up to 40 or even 70 nun. long; Castejori says usually 50 to 60 mm.
long in adult birds; Baylis and Daubney say 70 to 95 nun. long by
2.E mm. wide. Vulva near the middle of body. Tail straight,
conical, mucronated. Anus 1.2 mm. from end of tail: posterior to
onus a longitudinal depression. Eggs 68/* long, according to
Travassos; 80 to 90/x long, according to Railliet, by 1" to 50/t wide:
72/t long by 48/* wide, according to Irwin-Smith; 60 to 70/* long by
40/* wide, according to Johnston.

Life history.- -Unterberger (1868) found that embryos developed
in eggs to the infective stage in 17 days, and thai when such egos
were fed to pigeons the young worms developed to adults in :'» weeks.
The occurrence of larval worms in the liver, reported by Bedel,
suggests that the larvae may migrate in a manner similar to ascarids.

3612—27 8

88 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Distribution. — North America (United States), Europe (France,
Germany, Italy, Russia, and Spain), Africa (Dahomey), Asia
(India, Russian Turkestan), South America (Brazil), and Australia.
Probably a cosmopolitan species.

ASCARIDIA COMPAR (Schrank, 1790) Travassos, 1913

Synonyms. — Ascaris corn-par Schrank, 1790; Heterakis compar
(Schrank, 1790) Stossich, 1887; Ascaris lagopodis Froelich, 1802.

Hosts. — Caccabis chucar, C. saxatilis, Colinus virginianus, Coturnix
communis, G. dactylisonans, Gallus gallus, Numida meleagris, Ortyx
virginianus, Perdix cinerea, Tetrao lag opus, T. tetrix, T . urogallus.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : White worms, comparatively
thick but thinning posteriorly. Mouth with 3 rounded lips, each
bearing a central papilla. Esophagus 1/14 of total length.

Male 36 to 48 mm. long. Tail 1/66.3 of total length, straight and
obliquely truncated. Tail cordate. Sucker oval, 340/x, by 300/i.
Nine pairs of papillae (fig. 120) , of which 4 are preanal (2 being near
the sucker and 2 just anterior to cloacal aperture) and 5 are postanal.

Female 84 to 96 mm. long. Tail 1/65 of total body length. Eggs
91/* long by 57/* wide, according to Linstow ; 80/* by 60/t, according to
Mueller.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — North America (United States), Asia (Philippines,
India and Formosa), Europe (England, Germany, Italy), and
Australia. Reported from the quail in Florida by Leidy.

ASCARIDIA COMPRESSA (Schneider, 1866) Railliet and Henry, 1912

Synonym. — Heterakis compressa Schneider, 1866.

Host. — Gallus gallus.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : Large worms. Mouth with 3
lips, the lips dissimilar (presumably the dorsal lip larger than the
2 submedian) and each lip having a larger median lobe and 2 smaller
lateral lobes.

Male 53 mm. long.

Female 85 mm. long.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Australia.

ASCARIDIA CORDATA (Linstow, 1906) Railliet and Henry, 1914

Synonym. — Heterakis cordata Linstow, 1906c.
Host. — Callipepla sguamata.
Location. — Intestine.

NEMATODE PARASITES OF BIRDS

89

Morphology. — Ascaridia (p. 77) : Mouth with 3 semicircular lips,
the dorsal lip with 2, the other lips with 1 papilla.

Male 27 mm. long by 880/x wide. Esophagus 1/12, tail 1/70 of
total length. Caudal extremity heart-shaped (fig. 121). Preanal
sucker circular. Eight pairs of caudal papillae, 3 of which are pre-
anal and 5 postanal. Spicules subequal, the right 2.06 mm., the
left 2.37 mm. long.

Female 42 mm. long by 1.11 mm. wide. Esophagus 1/16, tail 1/35
of total length. Vulva slightly anterior to middle of body, dividing
the body length in ratio of 16 : 17. Eggs 91/x long by 57ju wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — North America (Mexico).

Figs. 121-122. — 121, Ascaridia cordata. Male tail. After Linstow, 1908. 122,
Ascaeujia cristata. a, Head; b, male tail. After Linstow, 1901

ASCARIDIA CRISTATA (Linstow, 1901) Railliet and Henry. 1914

Synonym. — Heterakis cristata Linstow, 1901.

Hosts. — Belearica pavoniria, B. regulosum, Grus antigone.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Mouth with 3 lips, the ventro-
laterals larger and bearing 2 blunt teeth; dorsal lip with 2 papillae.
Lateral alae strongly developed, extending about 1.22 mm. posterior
from the head. Esophagus 1/9 of body length. According to
Baylis and Daubney (1922) there are 27 pairs of cervical papillae,
similar to those in A. columbae, extending from a point about 900/i,
from the head posteriorly for a distance of 6 to 6.5 mm., on the dorsal
surface.

Male 35 to 38 mm. long by 1.1 to 1.34 mm. wide. Tail 1/72 of
total length; it does not become narrower until very near the pos-
terior extremity. Preanal sucker longer than wide. According

90 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

to Linstow (fig. 122), 9 pairs of caudal papillae, of which 2 preanal,
7 postanal; according to Bayiis and Daubney, 13 pairs, of which
6 are preanal, 7 postanal. Spicules equal, 950/a long by 42/z wide,
alate.

Female 57 mm. long by 1.76 mm. wide, according to Linstow;
18 to 40 mm. long by 1.1 to 1.2 mm. wide according to Bayiis and
Daubney. Tail 1/98 of body length. Vulva 20 mm. from anterior
end in the specimens 38 to 40 mm. long; vagina short, transverse.
Eggs 85 to 91/a long by 58 to 62/x wide, with thick shells.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Africa (Langenburg, Nyassa-See) and Asia
(India).

ASCARIDIA CYLINDRICA (Blome, 1909) Railliet and Henry, 1914

Synonym. — Heterakis cylindriea Blome, 1909.

Host. — Tetrao urogallus.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : Body cylindrical, the anterior
and posterior halves of body equally wide. Lateral membranes ab-
sent. Head set off from body. Lips of unequal size, each lip carry-
ing on its inner surface an anteriorly directed " tooth-plate " (Zahn-
platte).

Male 43 mm. long by 1.3 mm. wide. Caudal alae (fig. 123) well
developed. Ten pairs of papillae; in an occasional specimen there
may be an additional unpaired papilla on one side (anterior to
the most anterior pair). Preanal sucker with heavy chitinous ring.
Spicules unequal in length (fig. 124) and thickness, according to
Blome's description, but his figure shows them equal. Through-
out the anterior half of their length they are surrounded by a
sheath, the inner surface of which is neatly folded, giving in optical
section a dentate appearance.

F&male 55 mm. long by 1.5 mm. wide. Tail pointed.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Europe (Germany (Breslau)).

Skrjabin (1916ff) has made a study of this species from the
moor hen; the present writer, however, has been unable to obtain
his article.

ASCARIDIA DOLICHOCERCA (Stossich. 1902) Railliet and Henry, 1911

Synonym,. — Heteralcis dolichocerca Stossich, 1902.

Host. — Circus s pilot h o rax.

Location. — Not given.

Morphology. — Ascaridia (p. 77) : Head distinct from body, with
3 large lips. Lateral membranes wide. Length of specimens 25
to 30 mm.

NEMATODE PARASITES OF IHUDS

91

Male with preana] sucker circular, surrounded by a large chiti-
nous ring. No preanal papillae observed. Eight pairs of postanal
papillae (fig. 125), 3 of them ventral, near the cloaca! aperture,
the other 5 lateral and large. Spicules simple and straight.

Female with long tapering caudal extremity. Vulva anterior to
middle of body.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Oceania (New Guinea).

m

125

FlGS. 123-125. — 123, Ascabidia cyundbica. Male tail. 124, Sficule. After Blome,

1909. 125, ASCARIDIA DOLICHOCEUCA. MALE TAIL. AFTER grassier!, 1902
ASCARIDIA FASCIATA Baylis. 1920«

Host. — f "inago delalandii.

Location. — Not given.

Morphology. — Asearidia (p. 77) : Mouth (fig. 12(>) with 3 large
semicircular lips, the dorsal with 2 large papillae, the others with a
single median papilla. Cuticle of body (fig. 127) with thickened
transverse lings resembling annulations, their maximum width, in
middle of body, being 100/x. Lateral alae well developed. 3 to 4
mm. long by 800/x maximum width. Esophagus club-shaped, meas-
uring 1/20 of total body length.

Male up to 43 mm. long by 1 mm. wide. Tail (fig. 128) straight,
pointed, long (700/x). Cuticle raised in 2 longitudinal ridges at the
sides of the sucker; sucker 250/x in diameter. Thirteen pairs of
caudal papillae, of which 7 are preanal, 1 adanal, 5 postanal.
Spicules equal, 1.9 mm. long, alate, about 70/* wide.

92

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female up to 74 mm. long by 1.9 mm. wide. Tail 1.25 to 1.6 mm.
long. Vulva slightly anterior to middle of body. Vagina and un-
paired portion of uterus together measure about 2 mm. long. Eggs
88/x long by 45/u. wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — East Africa (Dar-es-Salaam).

ASCARIDIA FRANCOLINA (Linstow, 1899) Railliet and Henry, 1914

Synonym. — Heterakis francolina Linstow, 1899.

Host. — Francolinus bicalcaratus.

Location. — Not given.

Morphology. — Ascaridia (p. 77) : Esophagus 1/12.3 of total length.

0-3 mm

/Z6

IZ7.

Figs. 126-128. — Ascaridia fasciata. 126, Head. 127, Cuticulak rings of body. 128,
Male tail. After Baylis, 1920

Male 40 mm. long by 870/i. wide; tail 1/148 of total length. Eight
pairs of caudal papillae (fig. 129), of which 3 are preanal.

Female 62 mm. long by 1.3 mm. wide; tail 1/133 of total length,
rounded. Eggs 78/* long by 49/* wide, with very thick shells.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Africa (Togo and Bismarckburg) .

ASCARIDIA GRANULOSA (Linstow, 1906) Railliet and Henry, 1912

Synonym. — Heterakis granulosa Linstow, 1903.

Host. — G alius gallus.

Location. — Small intestine.

Morphology. — Ascaridia (p. 77) : Large, thick worms.

Male 27 mm. long by 590/t wide. Esophagus length equal to 1/3
of body length. Tail length equal to 1/49 of body length. Sucker
long and oval and surrounded by granulations. Three pairs of pre-
anal and 6 pairs of postanal papillae. (Fig. 130.) The most anterior
preanal papillae are transversely elliptical; of the postanal papillae

NEMATODE PAEASITES OF BIEDS 93

the fourth, seventh, and ninth pairs are spherical and marginal.
Spicules straight and rod-shaped, 570/* long.

Female 55 mm. long by 790/* wide. Esophagus length equal to
1/14 body length. Tail length equal to 1/40 of body length. Vulva
just anterior to equator of body, dividing it in the proportion of
16 : 17. Eggs thick-shelled, 78/*, long by 42/* wide.

Life history. — Probably similar to that of A. gdlli (p. 82).

Distribution. — Asia (Ceylon (Colombo)).

ASCARIDIA LINEATA (Schneider, 1866) Railliet and Henry, 1912

Synonyms. — Heterakis lineata Schneider, 1866; Ascaridia liamia
Lane, 1914. A. hamia is regarded as a synonym of A. lineata by
Boulenger. Existing figures of the male tail of these species indi-
cate that there is considerable variation within the limits of what
might be expected in one species, but the description below notes
the species to which certain points apply where nothing is known
to the writer as to the same points in the other species.

Hosts. — Anas boscJias domestica, Bonasa umbellus, Gallus gallus
and " goose."

Location. — Small intestine, and according to Boulenger, stomach.

Morphology. — Ascaridia (p. 77) : Large, thick yellowish worms,
with 2 very prominent white lateral lines. Lane describes A. hamia
as having no lateral membranes but as having instead a rope-like
twist down the middle of the lateral lines. Head separated from
body by a slight neck (A. hamia). Head with 3 large, subequal
lips, the lips having only 2 dentigerous ridges (the second one being
very small) and thus distinguishing it from A. galli, according to
Schneider. Lane says A. hamia has 3 large lips, each consisting of
a central mass and 2 lateral flaps, and each lip bearing 2 papillae,
those on the dorsal lip being larger than the others; on the median
face each lip bears a sharp horizontal ridge; close to the head are
5 or 6 cervical or nuchal papillae on each side.

Male 55 to 68 mm. long; A. hamia 70 mm. long by 900/u, wide.
Anal sucker 200 to 250//. in diameter, with! strong cutinous (chitinous)
wall with a papilliform interruption on its posterior rim. Tail
with narrow bursal membranes and 10 pairs of caudal papillae.
There are 3 pairs of preanal papillae located ventrally, one pair of
these anterior to the sucker, one pair opposite the sucker, and one
pair between the sucker and the cloacal aperture. The next pair
is lateral but seems to have a variable relationship ; it is figured for
A. hamia by Lane as preanal and for A. lineata by Travassos as
adanal, by Schneider as postanal, and by Boulenger (fig. 131) as
somewhat adanal and somewhat postanal. Close behind this pair
is a pair figured as ventral by Schneider and Boulenger and as

94

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

lateral by Travassos and Lane. This is closely followed by a ven-
tral pair and this in turn by a lateral pair. Towards the tail end,
in a secondary expansion of the caudal alae, are 2 pairs of lateral
papillae with a pair of ventral papillae occupying variable posi-
tions between them; it is this posterior group that seems most
characteristic of the species and distinguishes it in particular from
A. galli, in which the ventral pair of papillae is lacking. Cloacal
aperture 540/* from tip of tail. Spicules equal and narrow, with
slightly enlarged rounded points, 1.6 to 2.4 mm. long.

Female 60 to 95 mm. long. Vulva about at union of anterior and
middle thirds of body length (in middle of body (A. hamia), accord-
ing to Lane (A,, hamia) and to Schwartz). Tail 1.08 mm. long;

Figs. 129-132. — 129,Ascaridia fkancolina. Male tail. After Linstow, 1899. 130
ascauidia granulosa. male tail. after ll.nstow, 190g. 131, ascaridia lineata.
Male tail. After Boulenger, 1923. 132, Ascaridia longecirrata. Male tail.
After Linstow, 1879

according to Lane, that of A. hamia is 1.3 mm. long and there are
lateral papillae 500m from the tip. Eggs elliptical, 80m long by 50m
wide; in A. hamia Lane says they are thick-shelled and 65m long
by 40m wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — North America (United States), South America
(Brazil), Asia (Turkestan and India (A. hamia)), Africa (Belgian
Congo and Zanzibar) and, according to Skrjabin, Europe. As noted
previously (p. 82) it is this species and not A. galli which is com-
monly found in domestic birds in the United States.

ASCARIDIA LONGECIRRATA (Linstow, 1879) Travassos, 1913

Synonym. — Heterakis longecirrata Linstow, 1879.
Host. — Geopelia, species.
Location. — Intestine.

NEMATODE PARASITES OF BIRDS 95

Morphology. — Ascaridia (p. 77): Cuticle cross-striated with pe-
culiar markings (the figure which Linstow gives of the cuticle of
Heterakis longecaudata (p. 68) may be mislabeled and belong to this
species, as he does not describe any peculiar markings in the hetera-
kid). Mouth with 3 Lips bearing a small papilla. Linstow states
that in addition to these, papillae are found scattered in an irregular
manner over the whole body.

Male 30 mm. long by 190ju wide. Esophagus 1/11. tail ! 46 of total
body length. Nine pairs of caudal papillae (fig. 132), of which 3
are preanal and 6 postanal. Both spicules very long, the one 2.1
mm., the other 1.1) mm. long.

Female apparently unknown, as not described.

Life history. — -Probabh' similar to that of A. galli (p. 82).

Distribution. — Not given.

ASCARIDIA MAGALHAESI Travassos, 1913

Host. — Geotrygon iTiontana.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Cuticle with distinct transverse
striations. Head with 3 more or less equal lips. Cephalic extremity
with 2 lateral alae about 500^ wide and 3 mm. long; esophagus
about 3.5 mm. long.

Male 35 mm. long. Cloacal aperture about 434^ from posterior
border. Twelve pairs of caudal papillae (fig. 133) of which 5 are
preanal, 4 adanal and 3 postanal. Spicules equal, 1.6G mm. long.

Female 41 to 48 mm. long. Vulva slightly anterior to middle of
body. Anus 1.07 mm. from caudal extremity. Eggs 74/x long by
44/i. wide.

Life history. — Probably similar to that of A. galli (p. 82).

Locality. — South America (Brazil).

ASCARIDIA MAGNIPAPILLA (Linstow, 1906) Railliet and Henry, 1914

Synonym. — Heterakis magnipapilla Linstow, 190Ck/.

Host. — Tetrao tetrix (Lyrrurus tetrix).

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Cuticle with eross-st nations and
additional deep contractions. Mouth with 3 wide low lips (fig.
134a) which measure 180^ deep by 370/x wide. Body thick; head
and tail ends markedly narrowed. Esophagus 1/9 of total body
length.

Male 30 mm. long by 1.46 mm. wide. Tail 1/77 of total length.
Caudal alae (fig. 134/>) wide. Nine pairs of large pedunculated
papillae of which 2 are preanal, 7 postanal. Preanal sucker 280/*
wide by 310/* long. Spicules 3.03 mm. long, witli pointed ends.

96

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 42 mm. long by 1.58 mm. wide. Tail conical, 1/42 of total
length. Vulva somewhat anterior to middle of body, dividing the
body length in ratio of 3 : 4. Eggs oval, 96/x long by 57/x wide, with
thick shells.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Europe (Prussia (Friedland) ).

ASCARIDIA NUMIDAE (Leiper, 1908) Travassos, 1913

Synonyms. — Heterahis numidae Leiper, 1908; Heterakis calcarata
Gendre, 1909.

Host. — Numida meleagris, N. ptilorhyncha, N. papillosa trans-
vaalensis.

Location. — Small intestine and ceca.

Morphology. — Ascaridia (p. 77) : Very similar to A. columbae
(p. 86). Body white, without lateral membranes. Three strong,

/JJ.

Figs. 133-135. — 133, Ascaridia magalhaesi. Male tail. After Tkavassos, 1913. 134,
ascaridia magnipapilla. c, head ; 6, male tail. after llnstow, 1906. 135,
Ascaridia ndmidae. Male tail. After Gendre, 1909

equal lips, each having 3 lobes, of which the median is the larger,
and bearing a papilla where the median lobe joins each of the
smaller lateral lobes. Esophagus without bulb.

Male 19.4 to 35 mm. long by 720 to 880/* wide. Tail end (fig. 135)
terminates in a conical bent point resembling a spur. The cloacal
aperture is on a comparatively high prominence. There are 2 nar'
row caudal membranes. The sucker has strong cutinous (chitinous)
walls, interrupted by a papilla on the posterior rim. There are 10
pairs of caudal papillae; 1 pair is immediately posterior of the
sucker, 1 pair about half way between the sucker and the cloacal
aperture, 2 pairs of smaller papillae near the cloacal aperture, 3

NEMATODE PARASITES OF BIRDS 97

pairs of marginal papillae in relation with the bursal papillae fol-
low, the first pair of these opposite the last pair of the foregoing,
then 1 pair of papillae not in relation with the bursal membrane,
and finally 2 pairs in relation with the bursal membrane just anterior
of the terminal spur of the tail. The two spicules are equal, about
3 mm. long, slender, almost straight, and with slender alae.

Female 30.6 to 50 mm. long by 1 to 1.28 mm. wide. The tail is
straight, conical and pointed, and bears 2 very small, symmetrical
papillae about two-thirds of the distance from the anus to the tip of
the tail. The vulva is a slightly salient transverse slit, posterior of
the middle of the body. The eggs are 98/x long by 53ju, wide, and
contain an embryo when oviposited.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Africa (Dahomey and on White Nile and
Transvaal).

ASCARIDIA ORTHOCERCA (Stossich, 1902) Railliet and Henry, 1911

/Synonym. — Heterakis orthocerca, Stossich, 1902.

Host. — Rhea americana.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Body 30 to 40 mm. long by 1 to
2 mm. wide; cylindrical, attenuated anteriorly. Cuticle transversely
striated. Mouth with 3 almost equal lips, the dorsal semicircular
with undivided pulp and with 2 conspicuous papillae. Caudal ex-
tremity ending in a small cylindrical prolongation.

Male with caudal alae weakly developed (figs. 136 and 137). Pre-
anal sucker subelliptical, provided with cutinous (chitinous) ring
with a small papilla in its posterior rim. Twelve pairs of caudal
papillae, of which 5 are preanal, 6 are postanal and 1, which is
double, is adanal. Spicules very long, alate.

Female with prominent vulva situated at % of the body length.
Eggs elliptical, with thick smooth shell.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Europe (Italy (Cagliari)) and South America
(Brazil).

ASCARIDIA PTEROPHORA (Creplin, 1854) Railliet and Henry. 1914

Synonyms. — Ascaris pterophora Creplin, 1854; Ascaris lati-
caudaMol'm, 1860; Heterakis laticauda (Molin, 1860) Stossich, 1887.

Hosts. — Cariama cristata (Dicholophus marcgraf) and Micro-
dactylies cristatus.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Mouth with 3 large lips. Lat-
eral membranes wide, semilanceolate.

98

BULLETIN 140. UNITED STATES NATIONAL MUSEUM

Male 30 to 40 mm. long (77. laticauda) or 20 mm. long by 500/*
wide (.4. pterophora). Caudal extremity (fig. 138) broad; cloaca!
aperture 360// from posterior end in a 20 mm. specimen. Caudal
alae narrow. Preanal sucker large (100//, in diameter), with a pa-
pilliform nodule in the posterior rim; it is 520/x from the posterior
end of body. Ten pairs of caudal papillae, 4 of which are preanal
(3 pairs ventral, 1 pair lateral). Spicules equal, 600/* long by 21//,
wide.

Female 45 to 60 mm. long. Posterior extremity sharply pointed.
Anus 1 to 1.5 mm. from posterior end. Vulva prominent, situated
in the median part of the body.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution*. — South America (Brazil).

Figs. 136-139. — 136, Ascaridia orthocerca. Male tail, ventral view. 137, Male
tail, lateral view. After Stossich, 1902. 138, Ascaridia pterophora. Male tail.
After Travassos. 1918. 139, Ascaridia serrata. Male tail. After Schneider, 1866

ASCARIDIA SERRATA (Schneider, 1866) Railliet and Henry, 1914

Synonym. — Heterakis serrata Schneider, 1866.

Host. — Penelope humeralis.

Location. — Intestine.

Morphology. — Ascaridia (p. 77) : Mouth with 3 lips, the dorsal lip
wider than the others. An anterior and posterior denticulated plate
(Zahnplatten), the former with 8 teeth in a row.

Male 41 mm. long. Preanal sucker circular, with cutinous (chitin-
ous) ring interrupted posteriorly by a papilliform nodule. A small
finely pointed appendage on posterior end of body. Ten pairs of
large papillae (fig. 139), 3 preanal, 2 adanal and 5 postanal.

NEMATODE PARASITES OF BIRDS 99

Female 41 mm. Long. Vulva at about anterior third of body (30
mm. from tail end).

Life history.- — Probably similar to that of A. r/aJIi (p. 82).
Distribution. — South America (Brazil).

ASCARIDIA STRELNIKOWI Skrjabin. 1916

Host. — Tina/m us, species.

Location. — Intestine.

Morphology. — Ascaridia (p. TT) : Cuticle with fine transverse
striations. Lips (fig. 140#) short, wide, made up of 2 lateral promi-
nences and a rounded anterior lobe.

Male 40 to 43 mm. long by 1.1 mm. wide. Dorsal lip 13G/* long
by 250/* wide. Esophagus 2.38 mm. long by 220/* wide. Caudal ex-
tremity (fig. 140&) conical, pointed; cloacal aperture 600/* from
posterior end. Preanal sucker circular, 340/* in diameter, with chi-
tinous ring; its posterior border, which has a median papilla, is 870/*
from caudal extremity. Thirteen pairs of caudal papillae, of which
8 are preanal, 5 postanal, and in addition 2 pairs of small papillae
located at some distance from the others, anterior to the preanal
sucker. Spicules equal, 3.4 mm. long alate.

Female 45 to 52 mm. long by 1.8 mm. wide. Dorsal lip 187/* long
by 37/* wide. Esophagus 2.72 mm. long by 600/* wide. Caudal ex-
tremity rounded, with styloid appendage 40/* long. Anus 1.1 mm.
from posterior end. Vulva in middle of body (in female 50 mm.
long, the vulva 25.5 mm. from anterior end). Eggs 68/* long by 51/*
wide.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — South America (Paraguay).

ASCARIDIA STROMA (Linstow, 1899) Railliet and Henry, 1914

Synonym. — Heterakis stroma Linstow, 1899.

Hosts. — Grus {T etrapteryx) paradisea, G. antigone and G. com-
munis.

Location. — Xot given.

Morphology. — Ascaridia (p. 77) : Head with 3 lips, the dorsal with
2 papillae, the others 1 papilla. Wide lateral membranes at head end
of body.

Male 25 mm. long by 800/* wide; esophagus 1/16, tail 1/47 of total
length. Preanal sucker circular with wide rim. Ten pairs of papil-
lae (fig. 141), of which 3 are preanal, 1 adanal, and 6 postanal. The
adanal pair and 2 pairs directly posterior to it are lateral and have
very large bases.

Female 5G nun. long by 1.7 wide. Tail 1/117 of total length,
with digitiform prolongation. Eggs 172/* long by 14G/* wide.

100

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Probably similar to that A. galli (p. 82).
Europe (Germany (Museum Berlin)) and Asia

Life history.-
Distribution.
(India).

ASCARIDIA STYPHLOCERCA (Stossich, 1904) Railliet and Henry, 1914

Synonyms. — Heterakis styphlocerca Stossich, 1904.

Hosts. — This worm was described from a domestic bird ( u Un
volatile domestico") with no indication as to which bird is meant.
The present writer has identified it from Gallus gallus.

Location. — Intestine.

Morphology. — Ascaridia (p. 77). Worms 90 to 100 mm. long,
according to Stossich, somewhat attenuated toward the extremities,
and with a transversely striated cuticle.

Pigs. 140-142. — 140, Ascaridia strelnikowi. a, Head ; b, male tail. After Skrjabin,
1916. 141, Ascaridia stroma. Male tail. After Linstow, 1899. 142, Ascaridia
styphlocerca. male tail. after stossich, 1904

Male with caudal alae slightly developed and showing small round
granulations. (Fig. 142.) Ventral sucker large, almost circular, with
strong wall interrupted on the posterior margin by a small papilla.
Eight pairs of caudal papillae, large and fungiform ; of these 2 pairs
are in the region of the sucker, 1 pair preanal and postsuctorial,
and 5 pairs postanal. Stossich figures 1 unpaired papilla on the
right side between the papillae of the last 2 pairs on that side; he
refers to it as on the left side. He also refers to an asymmetrical
papilla in connection with the first preanal pair; his figure sug-
gests that there is a median, but apparently symmetrical papilla
between the members of the last postanal pair.

The single male specimen from South Africa, examined by the
present writer, was very like that figured by Stossich except that
the median papilla figured between the members of the most pos-
terior pair was absent and the second from posterior end, figured

NEMATODE PAEASITES OF BIRDS 101

as an asymmetrical papilla, was a symmetrical pair, making pairs
of postanal papillae, 5 of them lateral. The granulations, very
refractive, covered the ventral surface of the body from the caudal
extremity anteriorly to the level of the sucker (Stossich figures them
only in postanal region). Esophagus 3.9 mm. long; preanal sucker
232/* long by 183/* wide (external measurements). Spicules at least
2.5 mm. long (free ends apparently broken).

Female with a caudal extremity forming an elongated cone with
an obtuse apex.

Life history. — Unknown; presumably somewhat similar to that
of A. galli (p. 82).

Distribution. — Africa (Gambia and (new record) Potchefstroom,
South Africa).

ASCARIDIA TRILABIUM (Linstow, 1904) Railliet and Henry, 1914

Figs. 143-144. — 143, Ascaridia TRILABIUM. Male tail. After Linstow, 1904. 144,

ASCAIUDIA ANSEBIS. MALE TAIL. AFTER SCHWARTZ, 1925

Synonym. — Heterakis trilabium Linstow, 1904.

Host. — Centropus sinensis.

Location. — Intestine.

Morphology. — Ascaridia (p. 77): Cuticle annulate; head with 3
semicircular lips, the dorsal with 2 papillae, the others with 1
papilla. Esophagus 1/13 of total length.

Male 2G mm. long by 690/* wide. Tail 1/49 of body length. Ten
pairs of caudal papillae (fig. 143), of which 4 are preanal, 6 postanal.
Preanal sucker circular. Spicules subequal, 1.95 and 1.97 mm. long,
the free end rounded.

Female 39 mm. long by 1.1 mm. wide. Tail 1/58 of total length,
conical, attenuated, with small digitate prolongation. Vulva
posterior to middle of body, dividing body length in ratio of 8:5.
Eggs 81/* long by 47/* wide, with thick smooth shells.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Asia (Ceylon (Horana)).

102 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

ASCARIDIA ANSERIS Schwartz, 1925

Host. — A riser domesticus.

Location.— -Small intestine.

Morphology. — Ascaridia (p. 77).

Male 32 mm. long by GOO/* wide. Head, separated from body, 172/*
wide near base. Esophagus 1.75 mm. long by 285/* in maximum
width. Nerve ring about 350/* from head end. Preanal sucker
circular, 138/* by 130/*, its posterior margin 172/* from cloacal aper-
ture and at least 700/* from posterior end of body (tip of tail
broken off in specimen described so that complete measurement not
possible). Caudal papillae asymmetrical, there being 14 papillae on
one side and 13 papillae on the other side. Of the 14 papillae, 5 are
preanal (4 ventral and 1 lateral) and 9 postanal (4 ventral and 5
lateral). Of the 13 on the other side of the body, 4 are preanal and
ventral and 9 are postanal but of different arrangement (there being
5 ventral and 4 lateral) than the postanal papillae of the opposite
side. Spicules nearly equal, 820/* and 827/* long, respectively, their
distal ends rounded (fig. 144).

Female unknown.

Life history. — Probably similar to that of A. galli (p. 82).

Distribution. — Asia (Hanoi (Tonkin) Indo-China).

Genus PSEUDASPIDODERA Baylis and Daubney, 1922

Generic diagnosis. — Heterakinae (p. 49) : Mouth with 3 lips and
with " cordons " resembling those of Aspidodera, opening in pairs
at the interlabial spaces and consisting of tubular grooves running
below the surface of the cuticle, with a narrow external opening
along their length. Members of each pair of cordons diverge, ex-
tending posteriorly a short distance then curve forward and end on
outer surface of lip, not joining the corresponding member of the
next pair as in Aspidodera. Narrow lateral alae present. Esopha-
gus muscular throughout, ending in a well-developed bulb.

Male with caudal alae and long pedunculated papillae ; spicules
very dissimilar; gubernaculum absent.

Female with vulva in median region of body.

Parasitic in alimentary canal, presumably ceca, of birds.

Type-species. — Pseud asp id odera pavonis Baylis and Daubney,
1922.

A second species and also a variety of it have been described
recently by Chandler. (See Addenda, p. 387.)

NEMATODE PARASITES OF BIRDS

103

PSEUDASPIDODERA PAVONIS Baylis and Daubney, 1922

Hoxts. — Pavo cristatus and Pavo muticus.

Location. — Not given; presumably ceca.

Morphology. — Pseuddspidodera (p. 102) : Small worms. Head
(fig. 145) with cordons as described in generic diagnosis. Diameter
of head at posterior limit of cordons, about 100/*. Narrow lateral
alae extend from a little anterior to nerve ring almost to tail.
Esophagus divided into short anterior portion and a long posterior
portion ending in a pyriform bulb; there appears to be some kind
of valvular apparatus at the union of the 2 portions; bulb 250 to
260/* long by 170 to 190/x wide and containing the usual valves.
Nerve ring 400 to 460/x from head end. Excretory pore GOO to G50/*
from head end.

Male G mm. long by 250/* wide. Esophagus 1.4 to 1.48 mm. long.

0-5 mm.

oamm.

Figs. 1 15 147. — Pseudaspidodera pavonis. 145, Head, lateral view. 146, Male
tail. i. kit. latebal view ; right, ventral view. 147, vulva and vagina in
lateral view, showing cement plug in vulva. arrow points in direction op
head. After Baylis and Daubnbt, 1922

Tail (fig. 14G) 380 to 430/* long, less than half of it provided in
anterior portion with wide alar expansions containing some caudal
papillae; posterior part of tail simple, slender, and finely pointed.
Circular preanal sucker, 120 to 130/* in diameter, with well developed
chitinous wall, 150 to 170ft anterior to cloaca] aperture; the
greatest diameter (antero-posterior) of sucker aperture is 70/*.
Spicules unequal and dissimilar. Right spicule slender and simple.
780/t long; left provided with broad alae and a barbed tip. and 450/;
long. No accessory piece. Caudal papillae, 12 pairs: of these 3
pairs just anterior to filamentous portion of tail, the middle pair the

104 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

more ventral and larger; a fourth pair, solitary, projecting into
alae; adanal group of 4 more or less lateral pairs, with long stalks,
and 2 small, sessile, ventral pairs, one of them anterior to and the
other posterior to the cloacal aperture. Of the 4 lateral pairs of the
adanal group, the most posterior is the stoutest and projects later-
ally; the next is directed more ventrally; the next is lateral; the
most anterior projects ventrally. There are 2 very slender and long-
stalked papillae on either side of the sucker.

Female 7 mm. long by 300/a wide. Esophagus 1.5 to 1.6 mm. long.
Tail 1 to 1.02 mm. long, straight, and tapering to a slender point,
with a very minute pair of caudal papillae about midway. Vulva
(fig. 147) posterior to the middle of the body, about 3 mm. from the
tail end. Vagina extends forward, turns in a semicircle toward the
ventral body wall, then turns to the right and dorsally, doubling
back. Two opposed uteri. Ova relatively large, somewhat oblong,
with a thin shell, 70/* long by 40/i wide, usually slightly thickened
internally at one pole; as seen in utero one end of the shell is occa-
sionally drawn out almost to a point; eggs not segmenting when
deposited.

Life history. — Unknown; probably somewhat similar to that of
Heterakis gallinae (p. 54).

Distribution. — Asia (India).

Subfamily Subulubinae Travassos, 1914

Synonym. — Kathlaniinae Lane, 1914.

Sub family diagnosis. — Heterakidae (p. 49) : Mouth with lips
inconspicuous or lacking, rarely with 3 lips, followed by a vestibule.
Esophageal bulb present. Preanal sucker of male fusiform and not
limited by a cutinous (chitinous) ring. Spicules equal or unequal,
one or both occasionally lacking or imperfectly chitinized. Guber-
naculum usually present, rarely lacking. Position of vulva variable,
may be in median or posterior part of body.

Type-genus. — Subulura Molin, 1860.

Genus SUBULURA Molin, 1860

Syno-nyms. — Ascaris Linnaeus, 1758, part; Heterakis Dujardin,
1845, part; Oxyuris Rudolphi, 1803, part; AUodapa Diesing, 1861.

Generic diagnosis. — Subulurinae (p. 104) : Head rarety with 3 lips,
usually with 6 conspicuous papillae. Mouth sometimes round, more
often oval or hexagonal with the large axis dorso-ventral, followed
by a buccal cavity (vestibule) at the base of which are often 3 teeth
at the entrance to esophagus. Esophagus club-shaped, followed by

NEMATODE PARASITES OF BIRDS 105

a distinct bulb. Lateral membranes often present. Male with
caudal alae feebly developed or lacking. Spicules equal or unequal;
gubernaculum present. Preanal sucker fusiform, without cutinous
(chitinous) ring. Caudal papillae in 2 longitudinal rows, number-
ing as high as 11 pairs. Female with vulva usually in median region
of body. Eggs ellipsoidal, with thin shells, usually embryonated
when deposited.

Parasitic in proventriculus and intestine (usually ceca) of birds,
intestine (usually large intestine) of mammals and intestine of rep-
tiles.

Type-species. — Subulura acutissima Molin, 1860.

KEY TO SPECIES OF SUBULUBA

1. Only female known; tail short (320/0 : from Turnix, species.

Subulura, species Baylis and Daubney, p. 132.
Male known; tail of female longer than above (443/* or longer except pos-
sibly in 8. forcipata where length not given and in S. papulosa where
female unknown) ; from other hosts than above 2.

2. Only male known (10 to 12 mm. long with 11 pairs of caudal papillae and

unequal spicules) ; from Corvus cajanus Subulura papillosa, p. 122.

Both male and female known ; from other hosts than above 3.

3. Spicules unequal 4.

Spicules equal 12.

4. Longer spicule 840/* in length ; 9 pairs of caudal papillae ; female 9.3 mm.

long Subulura rima, p- 125.

Longer spicule over 1 mm. in length; 10 or 11 pairs of caudal papillae;
female 10 mm. or longer 5.

5. Lateral alae said to be absent ; 10 pairs of caudal papillae ; gubernaculum

llO/i long Subulura subulata, p. 128.

Lateral alae present (with possible exception of S. curvata, where not men-
tioned) ; 11 pairs of caudal papillae; gubernaculum, if described, 144/*
or longer 6.

6. Long spicule 2.54 mm. in length ; lateral alae about 930/1 long.

Subulura carlosi, p. 113.

Long spicule not over 1.52 mm. in length; lateral alae 1.27 mm. or longer

(possible exception S. curvata where not described) 7.

7. Vulva of female slightly posterior to middle of body, dividing body length

in ratio of 14: 13; tail of female 1.54 mm. long__ Subulura curvata, p. 114.
Vulva anterior to middle of body; tail of female not over 1.28 mm. long_ 8.

8. Tail of female 443/* long Subulura lutzi, p. US-
Tail of female 1 mm. or longer 9.

9. Lateral alae extend to middle of body in male. % of body length in

female, thus for a length of at least 3 mm. ; cloacal aperture of male

169/1 from posterior end Subulura allodapa, p. 108.

Lateral alae extend only to anterior part of intestine, a length of not over
1.6 mm.; cloacal aperture of male 211/* or farther from posterior end_ 10.
10. Shorter spicule 850/* long; eggs 55a long by 45/* wide.

Subulura seurati, p. 120.

Shorter spicule 1 mm. or longer; eggs 76/* long by 50/t wide or 83/t long by

49/* wide 11.

106 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

11. Preanai sucker of male 280m long, its posterior edge 883m from tail end ;

ovejector of female 899m long Subulura bentocruzi, p. 110.

Preanai sucker of male 143m long, its posterior edge 508m from tail end ;
ovejector of female 1.28 mm. long Subulura reclinata, p. 124.

12. Male with 8 or 9 pairs of caudal papillae 13.

Male with 10 or more pairs of caudal papillae 15.

13- Spicules 590m long ; tail of female 1.16 mm. long Subulura gracilis, p. 134.

Spicules 880m long; tail of female not over 800m long 14.

14. Male 10.5 mm., female 14.8 mm. long; male tail 1/38, female tail 1/18.7 of

total body length Subulura acuticauda, p. 133.

Male 6.78 mm., female 9.26 mm. long ; male tail 1/25.7, female tail 1/14 of
total body length Subulura recurvata, p. 124.

15. Male with 10 pairs of caudal papillae 16.

Male with 11 or more pairs of caudal papillae 23.

16. Description incomplete; male 7 mm. long, female 12 mm. long; vulva of

female said to be posterior to middle of body Subulura acutissima, p. 107.

Size of worm usually greater than above; vulva anterior to middle
of body 17.

17. Spicules not over 790m long 18.

Spicules 880m or longer 20.

18. Male 4.5 mm., female 8.5 mm. long; spicules 790m long; eggs 65m long by

49m wide Subulura poculum, p. 123.

Male 8.2 mm. or longer ; female 13.6 mm. or longer ; spicules not over

700m long; eggs not over 50m long by 40m wide 19.

39. Male 8.2 mm., female 13.6 mm. long; gubernaculum rectilinear; ovejector
short, its total length 1.28 mm., the trompe being 385m long.

Subulura forcipata, p. 115.
Male 14.5 mm., female 22 mm. long; gubernaculum triangular, with horns
anteriorly ; ovejector very long, the trompe up to 5 mm. long.

Subulura noctuae, p- 119.

20. Spicules not over 900m long 21.

Spicules over 1 mm. long 22.

21. Cloacal aperture of male 318m, anus of female 600m from end of tail ; eggs

49m long by 41m wide . Subulura rimula, p. 125.

Cloacal aperture of male 200m, anus of female 800m from end of tail; eggs
65 to 75m long by 52 to 55m wide Subulura plotina, p. 122.

22. Cloacal aperture of male 170m, posterior end of sucker 500m from tail end ;

gubernaculum 100m long; eggs 59m long by 50m wide.

Subulura differens, p. 111.

Cloacal aperture of male 250 to 315m. posterior end of sucker 590 to 815m

from end of tail; gubernaculum 175 to 210m long; eggs 75 to 80m long

by 65 to 70m wide Subulura brumpti, p. 112.

23. Lateral alae extend entire length of body; male with 13 pairs of caudal

papillae; female 21 to 35 mm. long, its tail only l s of total body length.

Subulura leprincei, p. 118.

Lateral alae extend not farther than first part of intestine (except possibly

in 8. halli where alae not described) ; male with 11 pairs of caudal

papillae; female not over 19 mm. long (except possibly in S. mctoria

where may reach 23 mm.), its tail being longer than above 24.

24. Posterior end of preanai sucker 860m, or farther, from caudal extremity. 25.
Posterior end of preanai sucker not over 50/* from caudal extremity 26.

NEMATODE PARASITES OF BIRDS 107

25. Cloaca] aperture of male 500ft from end; spicules 1.5 nun. long; tail of
female 1/8.5 of total length.; eggs 80** long by G'V* wide.

Subulura halli, p. 117.
Cloacal aperture of male 210/* from end; spicules 7t;i> to .soo,u long; tail
of female 1/11 to 1/12 of total length ; eggs 65/* long by 35/t wide.

Subulura galloperdicis, p. 116:

2d. Preanal sucker 385/* long; spicules 1.71 mm. long_ Subulura travassosi, p. 131.

Preanal .sucker not over 186/* long; spicules not over 1.5 mm. long 27.

27. Lateral alae extend to first part of intestine 28.

Lateral alae do not extend to posterior end of esophageal bulb 2'.).

28. Preanal sucker 84/* long; spicules 899/* long by 16/* wide; gubernaculum

127/* long; tail of female 97o> long Subulura olympioi, p. 121.

Preanal sucker 186/* long; spicules .V>s,u long by 38/* wide; gubernaculum
22S/* long; tail of female 1.2 mm. long Subulura trogoni, p. 131.

29. Lateral alae extend to median part of bulb; preanal sucker IG'-V* long:

ovejector of female 950/* long, the trompe equivalent to % of its length.

Subulura strongylina, p. 128.
Lateral alae extend to middle of esophagus ; preanal sucker 135/t long ; ove-
jector of female 779/* long, the trompe equivalent to less than % of
its length 30.

30. The 2 most posterior pairs of caudal papillae similar, being the same size

and shape whereas the third ventral pair from posterior end is shorter
and slenderer than the former 2 pairs and removed by quite a space

from the second pair Subulura suctoria, p. 129.

The 2 most posterior pairs of caudal papillae are dissimilar, the second
being shorter and more slender; the third ventral pair is at least as large
as the first pair and is in close juxtaposition with the second pair.

Subulura similis, p. 127-

The above key does not include Subulura nndtipapillata recently
described by Chandler (see Addenda, p. 388).

SUBULURA ACUTISSIMA Molin, 1860

Synonyms. — Physaloptera sayinata strigis brasiliensis ' ; P. stron-
gylina cuculi-seniculi 1 ; Ileterakis acutissima (Molin, 1860) Stossich,
1887.

Hosts. — Guculus se/uculus, Coccyzus melacoryph/m^ PisorMna
atricapilla, Strix atricapilla.

Location. — Proventriculus and intestine.

Morphology. — Subulura (p. 104) : Head (fig. 151) rounded, with
2 lateral and 4 submedian papillae. At the entrance of the eso-
phagus 3 arcuate teeth.

Male 7 nun. long by 300/* yvide. Caudal extremity (fig. L52)
subulate with pointed appendage. Preanal sucker large, elongate,
remote from posterior end. Caudal alae present. Ten pairs of
papillae, of which 5 are preanal, 5 postanal. Two equal spicules,
long and curved.

•■Catalogue of the Vienna Museum.

108

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 12 mm. long by 300/a wide. Tail long, straight, pointed.
Anus remote from end. Vulva in posterior ( ? ) part of body, an-
terior to the anus.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA ALLODAPA (Creplin, 1853) Railliet and Henry, 1913

/Synonyms. — Oxyuris allodapa Creplin, 1853; Heterakis suctoria
Molin, 1860, part; Allodapa typica Diesing, 1861; Heterakis forci-

o-smm

ozmm

Figs. 148-150. — Subulura differens. 148, Anterior end. Original. 149, a, Vdlva ;
b, female tail; c, ove.tector ; d, egg. (Scale the same for a and b.) Original.
150, Male tail. X37.5. After Barreto, 1918

paria Schneider, 1866, part; Heterakis allodapa (Creplin, 1853)
Seurat, 1914, part; Allodapa allodapa (Creplin, 1853) Seurat, 1914,
part.

Hosts. — Cariama cristata, C. huppe, Dicholophus margravi.

Location. — Ceca.

Morphology. — Subulura (p. 104): Body yellowish color; anterior
extremity (fig. 153) conical with truncate apex, usually bent dor-
sally. Cuticle with transverse striations. Lateral alae present, nar-

NEMATODE PARASITES OF BIRDS

109

row, finely striated transversely. Mouth hexagonal, with 6 small
papillae (fig. 154). Mouth cavity straight, with thick walls which
have a characteristic bend. Three rounded teeth at entrance to
esophagus. Esophagus with bulb.

Figs'. .151— 152. — Subuluba acutissima. 151, Head, a, Side view; 6, front view. 152,
Male tail. After Dkasche, 1882

Male 7 to 10 mm. long by 330/* wide. Lateral alae extend from
head to middle part of body. Esophagus 1.29 mm. long. Tail (fig.
155) curved ventrally; cloacal aperture 169/* from end. Preanal
sucker 169/* long, without rim, 719/* from posterior end of body.
Eleven pairs of papillae, of which 3 are preanal and ventral, 2

. /J6 6

Figs. 153-156. — Slbuluha allodapa. 153, Anterior end. After IUkrbto. 1918. 154,
Head, front view. After Dhasciie, 1882. 155, Male tail. After Drasciie, 1882.
156, a and b, Ovbjkctoe, After Babrkto, 1018

adanal lateral and 6 postanal lateral. Barreto describes the spicules
as 1.525 and 0.465 mm. long but his figure shows no such great dif-
ference in length and suggests that the second measurement should
be 1.465 mm. Gubernaculum 152it long.

Female 10 to 14 mm. long by 440/*. wide. Lateral alae extend only
along anterior third of body. Esophagus, exclusive of bulb, 1.34

110

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

mm. long. Tail straight, pointed ; anus 1.03 mm. from end. Vulva
only slightty salient, of irregular shape, a little anterior to middle
of body. Ovejector (fig. 156) anteriorly directed, about 762/*, long;
vestibule short, 313/*; sphincter, 211/*; trompe 338/* long.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA BENTOCRUZI Barreto, 1918

Hosts. — Trogon, species and T. variegatus.
Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle with transverse stria-
tions. Lateral alae extend slightly beyond esophageal bulb, their

/57.

Figs. 157-158. — Subulura bentoceuzi. 157, Male tail, lateral view. 158, Male tail,
vhntsal view. after ba1ireto, 1918

length about 1.97 mm. Mouth hexagonal, with 6 small equal papiL
lae. Buccal cavity small, 3 teeth at its base. Esophagus claviform.
bulb spherical, 190/* in diameter.

Male 7.7 to 16 mm. long by 436/* wide. Buccal cavity 38/* deep.
Tail (figs. 157 and 158) curved ventrally, ending in an appendage 93/*
long. Cloacal aperture 245/* from posterior end. Preanal sucker
without chitinous edge, 280/* long, its lower limit 883/* from caudal
extremity. Caudal alae rudimentary. Eleven pairs of papillae, 3
of which are preanal, 2 adanal, 6 postanal. Spicules unequal, the
larger 1.5 mm. long, the smaller 1.1 mm. {% the length of the for-
mer.) Gubernaculum 169/* long.

NEMATODE PARASITES OF BIRDS 111

Female 13 to 21 mm. loiig by 643/* wide. Tail with appendage
140/i. long. Anus 1.1G mm. from posterior end. Vulva salient, ante-
rior to middle of body. Ovejector relatively long (899/*) ; vestibule
334/* long; sphincter small; trompe 541/* long. Two uteri divergent.
Eggs elliptical. 83/t long by 49/* wide.

Life history-. — Unknown; probably similar to that of Ascaridia
galll (p. 82).

Distribution. — South America (Brazil).

SUBULURA DIFFERENS (Sonsino, 1890) Kailliet and Henry, 1912

Synonym. — Heterakis different Sonsino, 1890.

Hirsts. — Francolinus bkalcaratus, Gallvs (/alius, and Numida mele-
dyris. The present writer has identified this species from the red-
headed pheasant. Pternistes, species, the nematodes having been col-
lected by R. O. Wahl at Potchefstroom, Union of South Africa.

Location. — Small intestine, in posterior portion.

Morphology. — Subulura (p. 104) : Straight, yellowish-white worms.
Lateral membranes (fig. 148) well developed, extending from the head
end past the first fifth of the intestine. Mouth with very indistinct
lips. Mouth capsule with thick cutinous (chitinous) walls, enlarged
in its posterior portion where it has 3 very small teeth. The esoph-
agus thickens posteriorly and is followed by a distinct sub-spherical
intestinal bulb.

Male 7 to 8.6 mm. long by 282/i wide. Cloacal aperture 1GG to 190/x
from tail end. The ellipsoidal caudal sucker is without a cutinous
(chitinous) wall; its posterior margin is about 500/<, from the tail end.
Ten pairs of caudal papillae (fig. 150), as follows: 3 large ventral
preanal pairs, 2 large lateral adanal pairs, 5 postanal pairs, 4 of them
ventral and 1 lateral. Spicules equal, strongly chitinized, a little
over 1 mm. long, their proximal ends infundibuliform and the distal
ends pointed. The gubernaculum is slightly curved and 100/x long.

Female 11.3 to 19 mm. long by 335/* wide. Anus (fig. 1495) 631
to 770/* from tail end. Vulva (fig. 149a) situated in a slight cuticular
depression, slightly anterior to the middle of the body, dividing body
length in ratio of 19 : 25. Ovejector (fig. 149<?) very unusual. It is
anteriorly directed and its most striking character is a bulbous or
bladder-like swelling which can be seen through the body walls of the
toto mount, after it has been cleared. On dissection, there is found at
some distance anterior to this, a sphincter; between the sphincter and
the bladder-like formation is a sac-like passage which opens not into
the anterior end of the bulbous compartment, but laterally, at which
point there is a thickened circular hyaline area to support it. This
structure does not seem to have been previously described in this
r.f>12— 27 9

112 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

species. Barreto says the ovejector of Si diff evens is 677/* long, the
vestibule being very long (338/*), and united to the trompe by a small
sphincter. The figures of various species of Subulura show inter-
esting variation in this general type of structure: S. seurati (p. 126)
and S. leprinei (p. 118) show the side entrance into a slightly devel-
oped bulbous cavity, while S. allodapa (p. 108) shows it even more
highly developed, as does also S. forcipata (p. 115). Eggs almost
spherical, 59/* by 50/i, thin-shelled, containing an embryo when
oviposited (fig. liQd).

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — Europe (Italy), Africa (Dahomey, Guinea, Belgian
Congo, Union of South Africa and Algeria) and South America
(Brazil).

SUBULURA BRUMPTI (Lopez Neyra, 1922) Cram, 1926

Synonyms. — Allodapa suctoria of Seurat, 1914, Heterakis suctoria
of Gendre, 1909; Subulura suctoria of Gedoelst, 1916; Allodapa
brumpti Lopez Neyra, 1922.

Hosts. — Gallus gallus and Meleagris gallopavo.

Location. — Ceca.

Morphology. — Subulura (p. 104). Yellowish worms with curved
cephalic extremity, the concavity of the curve dorsal. Cephalic alae
(fig. 159a) finely striated transversely and extending the anterior
sixth of the body length. Buccal cavity divided into well-differen-
tiated zones, with 3 small teeth situated at the origin of the esoph-
agus. The length of the esophagus and its bulb is 1/7.25 of body
length in male and 1/9.5 of body length in female. Nerve ring about
*4 of length of esophagus proper, without bulb, from anterior end.
Intestine enlarged at union with esophageal bulb.

Male 6.9 to 10 mm. long by 340 to 420/* wide. Tail ends in a pro-
longation about y 3 of length from cloacal aperture to end of tail.
Caudal sucker 170 to 220/t long, 340 to 500/* anterior to cloacal aper-
ture and 590 to 815/* anterior to end of tail. Five pairs of preanal
(according to Lopez-Neyra (fig. 159&), Seurat, and the present
writer; according to Gendre (fig. 161) and Gedoelst (fig. 160) 6
pairs) and 5 pairs of postanal papillae; of the postanal, 2 pairs of
small papillae are near the median line toward the end of the tail,
anterior to this a third and larger pair, more lateral, and then 2
pairs of larger papillae nearer the median line; the caudal glands
open between the second and third pairs, the pore apertures resem-
bling minute papillae; of the preanal papillae, the first 2 pairs
posteriorly may be regarded as adanal, a third pair is just preanal,
a fourth pair is about V3 of the distance from the posterior border
of the sucker to the cloacal aperture, and a fifth pair is at the side

NEMATODE PAEASITES OF BIBDS

113

of the sucker. Gendre and Gedoelst indicate an additional pair, sit-
uated in the submedian lines directly on the anterior lip of the
cloaca. The specimens identified by the present writer from the
turkey show thickenings of the lip at those points, but no true
papillae. The spicules are large, distinct, alate, and equal, 1.32 to
1.45 mm. long, or 1.5 mm. long in Seurat's specimens; gubernaculum
triangular, 175 to 210/* long.

Female 9 to 13.7 mm. long by 470 to 560/* wide at level of vulva.
Tail straight and conical, 650/* to 1 mm. long, terminating in a point
100/* long. Vulva slightly salient, 4.3 to 5.4, or 6.3 mm. in Seurat's
specimens, from the head end, or slightly anterior to the middle of
the body. Ovejector directed anteriorly, 980/* to 1 mm. long, the
vestibule 600/* long, and the sphincter 380/* long. Eggs subspherical,

Figs. 159-161. — Subuluka brumpti. 159, a, Anterior end; b, male tail.
After Lopez-Neyra, 1922. 160, Male tail. After Gedoelst, 1916. 161,
Male tail. After Gendre, 1909

with smooth shell, 75 to 80/* long by 65 to 70/* wide, and containing
an embryo when deposited.

Life history. — Unknown; probably similar to that of Ascaridi/t
gatti (p. 82).

Distribution. — Europe (Spain), Africa (Belgian Congo, Daho-
mey, and Algeria), and North America (Porto Rico).

SUBULURA CARLOSI Barreto, 1918

Host. — Piaya cayanna.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle with fine transverse
striations; lateral alac 930u long by 51/* wide, extending to level of
posterior end of esophagus. Mouth with small lips armed with 6
papillae, 2 of which are 1 huge and the oilier 4 small. Month cavity
short: esophagus about 1.1 mm. long; bulb subspherical.

114

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male i) to 10 mm. long by 290/x wide. Caudal extremity (fig. 162)
curved ventrally; caudal alae much reduced; cloacal aperature 186//.
from posterior end. Preanal sucker without chitinous ring, 212/x
long, its posterior end 465/x from caudal extremity. Eleven pairs
of papillae, of which 3 are preanal, ventral, and very large, 2 adanal,
and 6 postanal. Spicules very unequal in length and width, the
larger 2.54 mm. long by 16/* wide, the smaller 960/t long by 25,«
wide. Gubernaculum 169/* long.

Female 10 to 15 mm. long by 350/* wide. Tail acute; anus 1.16
mm. from posterior end. Vulva slightly salient, anterior to middle
of body (at about % of body length from head). Ovejector (fig.

Figs. 162-163. — Subulura carlosi. 162, Male tail, a, Side view; b, ventral, view.
163, Vulva and ovejector. After Barreto, 1018

163) long (1.11 mm.) ; vestibule 254/x long, made up of 2 charac-
teristic parts, with chitinous lining; sphincter short (135/*). Two
divergent uteri. Eggs elliptical, 84/x long by 67/x wide, embryonated
when deposited.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA CURVATA (Linstow. 1883) Railliet and Henry, 1914

Synonym. — Heterahis curvata Linstow, 1883.
Hosts. — C'accabis chukar and Perdix graeca.
Location. — Intestine.

Morphology. — Subulura (p. 104) : Body attenuated anteriorly.
Head with 3 weakly developed round projections, each bearing an-

NEMATODE PARASITES OF BIRDS 115

teriorly a small papilla; Esophagus 1/9.7 of total length, ending
in bulb: tail pointed.

Male 14 mm. long by 540/x wide. Tail 1 58.5 of total length.
Preanal sucker weakly developed. Two pairs of preanal, 2 pairs of
adanal and 7 pairs of postanal papillae (fig. 104). Of the postanal
papillae, 4 are ventral, 3 lateral. Spicules 1.2 mm. and 90.0/*. long
respectively.

Female 12.3 mm. long by 540^ wide. Tail Vs °1 body length,
pointed. Vulva slightly posterior to middle of body, dividing body
length in ratio of 14 : 13. Eggs 39/* by 26/* wide.

Life history. — Unknown; probably similar to that of Asoaridia
galU (p. 82).

Distribution. — Asia (Turkestan (by Linstow) and Russian Turk-
estan (Aoulie-ata: by Skrjabin) ).

SUBULURA FOKCIPATA (Rudolphi, 1819) Railliet and Henry, 1914

Synonyms. — Ascaris forcipata Rudolphi. 1819. part; Ascaris ford-
paria Schneider, 1866, part; HeteraMs forciparia Schneider, 1866,
part.

Hosts. — Bucco, species, Capito collaris, C. m/zcror., ('. melanoleucus,
C. rufiventris, C. striolatvs, C. tamatia, Coccyzus ?Jielanocoryphus, G.
minor, Cuculus naevius, C. senicidus, C. tingasu, Oaprirmtlgus
bacaurau, C. nacandua, 0. n/ficollis, C. urutau. DicJtolophus cristatus,
/'iplopterus naevius, Ovira guira, Monasa leucop*. M . tranquilla, .V.
i'.trbrosa, M. torquata, Piaya cayana, Tetrao uru.

Railliet and Henry say that the reports from Bucco and the
Caprimulgides are probably misidentificatkms.

Location. — Intestine and ceca.

Morphology. — Svbulura (p. 104) : Cuticle with transverse striatums.
Lateral alae short but wide (710/*, long by 74/* wide), extending to
posterior limit of bulb. Mouth with 6 papillae in two laterally
placed series. Buccal cavity small, with 3 teeth 30//. long at its
base. Esophagus 1.03 mm. long, bulb spherical. 17<V in diameter.

Male 8.2 mm. long by 282}x vide. Tail (rig. 1'63) strongly recurved,
its alae rudimentary ; cloacal aperture 211 /* from end. Preanal sucker
without chitinous ring, elliptical, 177/* long. 592/* from caudal ex-
tremity. Ten pairs of papillae of which 3 are preanal, 2 adanal, and
5 postanal. Spicules equal, 677/* long, (iubernaculuni rectilinear,
118/* long.

Female 13.6 mm. long by 308/* wide. Tail pointed. Vulva slightly
salient, anterior to middle of body, at 5.65 mm. from anterior end.
Ovejector (fig. 100) short (1.28 mm.), vestibule pyriform, strongly
ehitinized, 514/* long; sphincter equal to vestibule in length; trompe
relatively short (385/*). Eggs elliptical, 48/* long by 38/* wide, with
very thin shell; embryonated when deposited.

116

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil) and Africa (Algeciras).

SUBULURA GALLOPERDICIS Baylis and Daubney, 1922

Host. — Galloperdix spadieea.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Head small, buccal cavity 60/x
deep by 23/t wide anteriorly, with 3 teeth at base. Esophagus 1.5
mm. long, exclusive of bulb; bulb 200/t in diameter. Lateral alae
narrow, extend for about 1 mm. from anterior end of body.

Male 9.5 to 10 mm. long by 300/i wide. Tail 210/* long, drawn
out to fine point. Sucker spindle-shape, 650/* anterior to cloacal

Figs. 1G4-1G0. — 164, Slbulira curvata. Male tail. After Linstow, 1883. 165,

SlMILURA FORCIFATA. MALE TAIL. 166, VULVA A.VD OVEJECTOR. AFTER BARRETO, 1918

aperture. Eleven pairs of papillae (fig. 167), of which 4 are pre-
anal, 2 adanal, 5 postanal. Spicules equal, 760 to 800/* long, alate,
tapering. Gubernaculum 180/* long with a spur at about 60/* from
its anterior end.

Female 11.5 to 12.5 mm. long by 400/* wide. Anus 1.1 mm. from
tail end. Vulva anterior to middle of body, dividing body length
in ratio of 3:4. Vagina short, transverse, with ovejectors running
anteriorly and posteriorly from it. Eggs 65/i long by 35/* wide, em-
bryonated at time of deposit.

Life hhtory.— Unknown ; probably similar to that of Ascaridia
gaUl, (p. 82).

Distribution. — Asia ( India) .

NEMATODE PARASITES OF BIRDS

117

SUBULURA HALLI Barreto, 1918

Synonym. — Subulura forcipata Seurat, 1914, in part.

Host. — Otis tetrax.

Location. — Ceca .

Morphology. — SubvZura (p. 104) : Pharyngeal teeth (fig. 168c)
cuneiform, their free ends pointed.

Male 10.5 to 12 mm. long by 540/x wide. Esophagus, including
bulb, 1/6.5 of total length. Tail (fig. 168a) slender, ending in a long
fine point. Cloacal aperture 500/x from posterior end. Caudal alae
narrow. Eleven pairs of pedunculated papillae, 5 of which are

5W

Pigs. 167-168. — 167, Subuluba gallopeudicis. Malk tail. After Baylis and Daubnut
1922. 168, Subulura halli. a, Male tail; b, ovejector ; c, buccal cavity. Afteb
Seurat, 1914

preanal. Sucker far anterior (700^) of cloacal aperture. Spicules
equal, very long (1.5 mm.) with an alate expansion toward the free
end. Gubernaculum 120/* long.

Female 13.4 mm. long by 600/x wide. Esophagus, including bulb,
1/7.5 of total length. Tail long and slender, 1/8.5 of the total length.
Vulva anterior to middle of body, at % of its length. Ovejector
(fig. 168 b) with remarkably short sphincter. Eggs 80/a long by
60.U wide.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — Africa (Algeria (Maison-Carree) ).

118 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SUBULURA LEPRINCEI (Gendre, 1909) Travassos, 1913

Synonyms. — HeteraMs leprincei Gendre, 1909&; Allodapa leprin-
cei (Gendre, 1909) Seurat, 1914.

ffosts. — Caprimulgus aegyptius saharae; C. fossil; Macrodipteryx
macrodipterus.

Location. — Ceca.

Morphology. — Subulura (p. 104) : Head without lips and not
marked off from body, but with 6 papillae, 4 of them suhmedian
and 2 lateral; buccal cavity with a plate of chitinous teeth (fig.
169«). Two lateral membranes extending entire length of body but
especially well developed in anterior region. Esophagus with 2
swellings.

Male 10.5 to 18 mm. long by 300 to 420/* wide. Esophagus (in-
cluding pharynx) 1/9.3, tail 1/48 of total length, the latter ending
in a slender point. Caudal alae well developed. Thirteen pairs of
caudal papillae (fig. 170« and b), 7 of which are preanal and 6
postanal. Spicules (fig. 170<?) equal, 1 mm. to 1.4 mm. long. Gu-
bernaculum (fig. 170c?) triangular, 130/* long.

Female 21 to 35 mm. long by 400 to 600//, wide. Esophagus (in-
cluding pharynx) 1/15, tail 1/25 of total length, the latter slender and
pointed. Vulva inconspicuous, anterior to middle of body, at %
of the body length. Ovejector (fig. 169&) very long, similar to
that of S. allodapa. Eggs (fig. 170e) 60/* long by 51/* wide; in the
early stages the shell is thick and made up of polygonal plates but
later becomes thinner and smooth.

Life history. — Unknown ; probably similar to that of Ascarulia
galli (p. 82).

Distribution. — Africa (Dahomey) .

SUBULURA LUTZI Barreto, 1918

Host. — Strix, species.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Body white, filiform, cuticle
transversely striated. Lateral alae relatively short, about 1.27 mm.
long, reaching to the level of posterior end of esophagus. Mouth
elliptical, with 6 papillae, the lateral smaller than the median.
Buccal cavity (fig. 171) about 55/* deep. Three small equal teeth
at entrance to esophagus. Esophagus 1.24 to 1.6 mm. long; bulb
spherical, 250/t in diameter.

Male 14 mm. long by 370/* wide. Caudal extremity strongly
curved ventrally (fig. 173), with a sharply pointed appendage 279/*
long. Cloacal aperture 169/* from posterior end. Preanal sucker
elliptical, about 213/* long, without chitinous border, 798/* from

NEMATODE PARASITES OF BIRDS

119

caudal extremity. Caudal alae poorly developed. Eleven pairs of
papillae (fig. 172) of which 3 are preanal, 2 adanal, and 6 postanal.
Spicules unequal, the longer 1.27 mm. long, the other 3/4 as long.
Guhernaculum 144/* long.

Female 14 to 22.5 mm. long. by 460/* wide. Tail with slender
appendage similar to that of male, 110/* long. Anus 443/* from pos-
terior end. Vulva (fig. 174) markedly salient, of irregular shape,
anterior to middle of body, more or less at the point of union of

» 3 /

iSmmm \

100 /J

r/o.

ff&

/70.

Figs. 160-170. — Slbiltka lep.binCet. 100, a, Buccal cavity ; b, ov ejector.
After Seurat, 1014. 170, a, Male tail, ventral view; b, lateral view; c,
spicule; d, guberxacclum ; c, egg. After Gendbe, 1000

anterior third and posterior two-thirds of body. Ovejector long
(about 1.156 mm.); vestibule short, 275/*; sphincter small; trompe
elongate, 899/*. Two uteri. Eggs 83/* long by 55/* wide, elliptical,
embryonated at time of deposit.

Life hist 'or'/ ■. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA NOCTUAE (Seurat, 1914) Barreto. 1918

Synonyrii. — Allodapa noctuae Seurat, 1914.
Host. — Carine noctua glaux.
L ocation. — Intestine.
3612—27 10

120

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Subulura (p. 104) : Body large but slender. Lateral
alae present, narrow, finely striated transversely, extending from
head end to level of esophageal bulb. Mouth with 6 papillae ; buccal
cavity clearly divided into 2 parts, in the second or more posterior
are found 3 small rounded teeth.

Figs. 171-174. — Sdbuluba lutzi. 171, Anterior end. 172, Male tail,
ventral view. 173, male tail, lateral view. 174, volva and ove-
jector. After Barreto, 1918

Male 14.5 mm. long by 250/a wide. Esophagus, including bulb,
1/10 of total length. Tail short, ending in a point 85/* long. Cloacal
aperture 300/* from posterior end. Preanal sucker in form of a
longitudinal slit. Caudal alae present. Ten pairs of papillae (fig.
175), of which 3 are preanal, 2 adanal, 5 postanal. The caudal

Fig. 175. — Sdbuluba xoctuae. Male tail. After Seiirat, 1914

glands open between the second and third pairs of papillae (num-
bering from the posterior end). Spicules equal, 700/t long, filiform.
Gubernaculum triangular, elongated, prolonged anteriorly by 2 small
horns.

Female slender, 22 mm. long by 380/x wide. Esophagus short,
1/14.5 of total length. Tail short (630 M ). Vulva a little anterior to
middle of body, at 3/7 of body length. Ovejector posteriorly directed ;

NEMATODE PARASITES OF BIRDS

121

vestibule short; trompe remarkably long (up to 5 mm.). Uteri par-
allel. Eggs 50V l° n £ by 40/* wide, embryonated.

Life history. — Unknown; probably similar to that of Ascaridin,
galli (p. 82).

Distribution. — Africa (Algeria (Bou-Saada) ).

SUBULURA OLYMPIOI Barreto. 1918

Hosts. — Crypturus parvirostris, Nothura maculosa, Rhynchotus
mf esc ens.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle with transverse stria-
tions. Lateral alae extend to point a little posterior to the bulb.

FiGS. 176-177. — Subulura oi.ymi'ioi. 176, Male tail, a, Ventral view; l>,

LATERAL VIEW. 177, OvEJECTOR. AFTER BARRETO, 1918

Mouth with 6 papillae in 2 lateral series. Buccal cavity narrow.
Three teeth at entrance to esophagus; bulb spherical.

Male 5 to 8.4 mm. long by 411/* wide. Buccal cavity 41/* deep, 29/*
wide. Esophagus 899/* long. Caudal alae (fig. 176) poorly devel-
oped; cloacal aperture 228/* from posterior end. Preanal sucker
elliptical, without chitinous ring, 84/* long by 33/* wide, its posterior
end 4G5/* from caudal extremity. Eleven pairs of papillae, of which
3 are preanal, 2 adanal, and G postanal. Spicules equal, strongly
chitinized, 899/t long by 16/* wide. Gubernaculum 127/* long.

Female 7.7 to 15.6 mm. long by 462/* wide. Buccal cavity 67/* deep
by 33/i wide. Esophagus 1.13 mm. long. Anus 976/* from posterior
end. Vulva small, not salient, situated a little anterior to middle of
body. Ovejector (fig. 177) short (693/*) ; vestibule 211/* long; sphinc-
ter 127/t long; trompe short (338/t). Two uteri, with many circum-
volutions. Eggs elliptical, 67/* long by 50/* wide.

122 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA PAPILLOSA (Molin, 1860) Railliet and Henry. 1912

Synonyms. — Spiroptera corvi-cajani 2 , in Molin, 1860 ; Ascaris pa-
pulosa Molin, 1860d, not Bloch, 1782.

Host. — Corvus cajanus.

Location. — Intestine.

Morphology.— Subulura (p. 104) : Mouth with 3 lips bearing single
papillae, central, spherical. Body transversely crenated. Lateral
membranes narrow.

Male 10 to 12 mm. long by 100 to 300^ wide. Tail (fig. 178) slen-
der. Preanal sucker elliptical, without a chitinous ring, with strongly
developed radiating muscles. Caudal alae weakly developed. Spic-
ules unequal. At least 11 pairs of caudal papillae, of which 6 are
preanal and 5 postanal (posterior end lacking in specimen described
so that possibly there are more postanals). Preanal sucker situated
between the second and third pair of preanal papillae (that is, 2 pairs
of preanals are anterior to sucker).

Female unknown.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA PLOTINA Baylls, 1919a

Host. — Flotus rufvs.

L.ocation. — Not given.

Morphology. — Subulura (p. 104) : Body slender, tapering at either
end. Lateral alae present, lanceolate, extending from cephalic ex-
tremity to about the beginning of esophageal bulb. Mouth without
lips but with 6(?) very small papillae. Mouth cavity small, with 3
small teeth at entrance to esophagus. Esophagus with bulb.

Male 8.2 mm. long by 340/t wide. Esophagus 1.25 mm. long. Tail
(fig. 179) 200/a long; caudal alae absent. Preanal sucker elongated,
Avithout chitinous border, 400ft anterior to anus. Ten pairs of
papillae of which 3 are preanal, 1 adanal, and 6 postanal. Spicules
equal, 900^ long. Gubernaculum 150/a long.

Female 14.2 mm. long by 460/* wide. Esophagus 1.5 mm. long.
Tail 800yu, long. Vulva in middle third of body, 5.8 mm. from an-
terior end (thus anterior to middle). Eggs 65 to 75/x long by 52.5
to 55ju wide. Uterus extends posterior to anus into cavity of tail.

2 Catalogue of the Vienna Museum.

NEMATODE PARASITES OF B1EDS

i23

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution . — A i'rica ( Uganda ) .

SUBULURA POCULUM (Linslow. 1909) Kailliet airid Hcr.ry, 1914

Synonym* — Heterakis poeuZum Linstow, 1909.

Host. — Francolirms adspe rsus.

Location. — Intestine.

Morphology . — Subvlwra (p. 104) : Cuticula with line crOss-stria-
tions. Lateral membranes present, wide anteriorly. Mouth cavity
small; mouth with C> papillae. Esophagus with bulb.

; . ©

173.

Figs. 178-180. — 178, Si bdu k\ papillosa. Male tail. From Barreto, 1918. after
drasche. 179, subulitra plotina. male tail. after ilaylis, 1919. 180, subulura
pocdlbm. Male tail. Afteb Linstow, 1909

Male 4.5 mm. long by 280j* wide. Esophagus 1/5.3, tail 1/29 of
total length. Preanal sucker of long oval shape with radiating
muscles. Ten pairs of caudal papillae (fig. 180) of which 2 are pre-
anal. 2 adanal, and G postanal. Spicules 790/x long, sharply pointed.

Female 8.5 mm. long by 430^ wide. Esophagus 1/8.5, tail 1/9.7
of total length. Vulva anterior to middle of body, dividing body
length in ratio of 20 : 33. Eggs 65/x long by 39/* wide, embryonated.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — Africa (German South West Africa).

124 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SUBULURA RECLINATA (Rudolphi, 1819) Barreto, 1918

Synonym,. — Ascaris reclinata Rudolphi, 1819.

Hosts. — Crotophaga ani and 0. major and Piaya cajanea.

Morphology. — Subulura (p. 104) : Cuticle with transverse stria-
tions. Lateral alae 1.6 mm. long, extending to level of anterior part
of intestine. Mouth with 3 indistinct lips, armed, with 6 papillae,
arranged in 2 lateral series of 3 papillae each, the median a little
smaller than the laterals. Buccal cavity small, divided into 2 parts.
Esophagus about 1.21 mm. long; bulb spherical.

Male 11 mm. long by 359/* wide. Tail strongly curved ventrally
(fig. 181); caudal alae atrophied; cloacal aperture 211//, from pos-
terior end. Preanal sucker elliptical, 143/* long, its posterior end
508/* from caudal extremity. Eleven pairs of papillae, of which 3
are preanal, 2 adanal, 6 postanal. Spicules unequal, both in length
and width, the larger 1.52 mm. long by 19/* wide, the smaller 1.10
mm. long by 25/* wide. Gubernaculum 160/* long.

Female 14.3 to 20.5 mm. long by 514/* wide. Tail ending in chi-
tinous appendage 140/* long; anus 1.28 mm. from posterior end.
Vulva salient, a little anterior to middle of body. Ovejector long
(1.28 mm.); vestibule relatively short; sphincter small; trompe 4
times the length of vestibule, very muscular. Two uteri. Eggs
elliptical, 76/* long by 50/* wide, embryonated when deposited.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA RECURVATA (Linstow, 1901) Travassos, 1913

Synonym. — Heterakis recurvata Linstow, 1901.

Host. — Eurystomus afer.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle smooth ; head end
rounded, with G papillae set cross-wise. Tail pointed. Esophagus
with bulb.

Male 6.78 mm. long by 230/* wide. Esophagus 1/7, tail 1/25.7 of
total length. Preanal sucker long and narrow, with muscles radiat-
ing from it; Linstow describes 8 pairs of papillae, of which 2 are
preanal and 6 postanal; his figure (fig. 182) shows an additional
adanal pair, large and situated laterally. Spicules 880/* long.

Female 9.26 mm. long by 430/* wide. Esophagus 1/7.9, tail 1/14 of
total length. Vulva somewhat anterior to middle of body, dividing
body length in ratio of 11 : 15. Eggs 49/* long by 36/* wide.

Life history. — Unknown ; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — Africa (Langenburg, Nyassa See).

NEMATODE PARASITES OF BIRDS

125

SUBULURA KIMA (Linstow, 1906) Travassos, 1913

Synonym. — Ileterakis rima Linstow, 1906.

Host. — Otis houbara.

Location. — Ceca.

Morphology. — Subulura (p. 104) : Cuticle coarsely cross-striated.
Mouth with 6 papillae. Mouth cavity deep; 6 teeth at opening of
esophagus. Esophagus ends in small bulb. Cuticle at the head end
of body markedly widened (lateral membranes?); this gradually
disappears posteriorly.

Male 8.2 mm. long by 400/x wide. Esophagus 1/5.3, tail 1/21 of
total length. Gubernaculum present, three sided. Preanal sucker
very narrow and elongate, slit-like. Nine pairs of caudal papillae

Figs. 181-184. — 181. Subulura reclinata. Male tail. After Barreto, 1918.
182, Subulura recurvata. Male tail. After Linstow, 1901. 183, Subulura
rima. Male tail. After Linstow, 1906. 184, Subulura rimula. Male tail.
After Linstow, 1903

(fig. 183), of which 4 are preanal (the first pair near the sucker, thus
far removed from the other 3 which are near the cloacal aperture)
and 5 postanal. The right spicule 840/i long, the left 700/u, long.

Female 9.3 mm. long by 430^ wide. Esophagus 1/5, tail 1/6.7 of
total length. Vulva slightly anterior to middle of body, dividing
it in ratio of 51 : 53. Eggs immature in specimen described.

Life history. — Unknown; probably similar to that of Ascaridia

galli (p. 82).

Distribution. — Europe (Germany (Zoological Museum Koenigs-

berg)).

Baylis suggests that Subulura rima is a synonym of S. suctoria

but as the descriptions of the two species are not comparable on

certain points and as the number of caudal papillae and the spicule

lengths differ, the present writer prefers to keep S. rima as a distinct

species.

126 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SUBULURA RIMULA (Linstow, 1903a) Travassos, 1913

Synonym. — Heterakis rimula Linstow, 1903.

Host. — Cent/opus sinensis.

Morphology. — Subulura (p. 104) : Cuticle with cross-striations.
Head without lips or papillae. Tail conical, pointed. Esophagus
with bulb.

Male 8.6 mm. long by 340/* wide. Esophagus 1/7, tail 1/27 of total
length. Preanal sucker elongate, slit-like, with radiating muscles.
Ten pairs of caudal papillae (fig. 184), of which 3 are preanal,
7 postanal. Spicules 880/x long.

Female 12 mm. long by 580^ wide. Esophagus 1/8, tail 1/20 of
body length. Vulva 1/3 of body length from anterior end. Eggs
49/a long by 41,u wide.

Life history. — Unknown; probably similar to that of Ascarklia
galli (p. 82).

Distribution . — Asia ( Si am ) .

SUBULURA SEURATI Barrcto, 1917

Synonyms. — Subidura allodapa Seurat, 1914, part; Allodapa allo-
dapa Seurat, 1914, part.

Hosts. — Caccabis rufa and C. petrosa.

Location. — Ceca.

Morphology. — Subulura (p. 104) : Blood-red, thick body, much
attenuated posteriorly; 2 lateral alae in cephalic and esophageal
region; lateral lines prominent. Mouth hexagonal, surrounded b}^ 6
papillae. Three small teeth (fig. 185a) at entrance to esophagus.
Esophagus with bulb.

Male 14.5 mm. long by 500ju wide. Cloacal aperture 430,u from
posterior end. Preanal sucker eliptical, elongated, without chitinous
ring. Caudal alae narrow; 11 pairs of papillae (fig. 1855), 5 of
which are preanal. Spicules unequal, 1.35 mm. and 850ju, long re-
spectively, thus their relative lengths as 3 : 2. Gubernaculum triangu-
lar, 150/u, long.

Female 12.5 mm. long by 685/* long. Tail relatively short, 1.14
mm. long. Vulva in anterior part of body, at 1/3 the length from
anterior end. Ovejector (fig 185c) remarkably long (over 5 mm.) ;
vestibule pyriform; sphincter and trompe very long, the latter over
4 mm. The ovejector of Subulura leprincei is identical with that of
this species; in S. forripata and S. subulata it is very different.
Eggs 55/u. long by 45/x wide.

Life history. — Unknown ; probably similar to that of Ascaridia
gain (p. 82).

Distribution. — Africa (Bou-Saada, Algeria).

NEMATODE PARASITES OF BIRDS

1 f~ '

SUBULURA SIMILIS (Gendre, 1909) Travassos. 1913

Synonym. — Heterakis similis Gendre, 19095.

Hosts. — Centropus monachus, Coracias abyssinieus, Eurystomus
afer, Scops leucotis.

Itailliet and Henry suggest that the reports from Scops leucotis
(Strigi forme) and Centropus monachus (Coccigy forme) may be
confused. Barreto considers that Gendre was dealing with several
species from this wide variety of hosts and suggests that the material
from Eurystomus afer may be Subulura recurvata, described by Lin-
stow from that host in Africa.

Location. — Ceca.

Morphology. — Subulura (p. 104) : Quite similar to Subulura suc-
toria but, according to Gendre, differs in general body form, which

Fig. 185. — Scbulura seirati. a, Anterior end: b, male tail; c, ove.tector. After

Secrat, 1914

is much more slender in ,S Y . suctoria (this does not seem to the present
writer a marked difference, the width of the male in S. suctoria be-
ing given as 330 to 359/*, of the female as 400 to 600/i, whereas that of
S. similis as given by Gendre is 300 to 400^ in the male and 420 to
500/x in the female). Disposition of the second and third pairs of
postanal papillae (counting from posterior end of worm) (fig 186)
also different in the two species (see No. 30 of key, page 107).
Size of S. similis varies according to host:

in C. abyssinicus, male 9.5 mm. long by 400/x wide.

E. afer male 13.1 mm. long by 400/x wide.

female 18.8 mm. long by 500/x wide.

S. leucotis male 12.6 to 15 mm. by 300 to 3G0ti.

female 20.1 to 22.5 mm. by 420 to 500/x.

C. monachus male 9.4 to 13.1 mm.

female 11.1 to 18.4 mm.
Distribution, — Africa (Labe and Dahomey).

128 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

This species is probably identical with S. suctoria, the distinction
between them being very slight since the width of the nematode
may vary considerably due to fixation, etc. and the caudal papillae
have been shown to be quite variable in some species. Since the
hosts and locality are different, however, it is considered advisable
to leave the two species distinct at present.

SUBULURA STRONGYLINA (Rudolphi, 1819) Railliet and Henry, 1912

Synonym. — Ascai^is strongylina Rudolphi, 1819 ; Strongylus spicu-
lutus Cobbold, 1861; Heterakis spiculatus (Cobbold, 1861) Travassos,
1923.

Hosts. — Bucco capensis, B. melanoleucos, B. rufiventris, B. striola-
tus, B. swainsoni, B. tamatina, Gaprimulgus ruficollis, C. nacandua,
0. wutas, Chelidoptera tenehrosa, Crypturus noctivagus, G . species,
C . tatuapa, Cuculus melanorhynchus, C. tinguacu, Gallus gallus,
Malacoptila torquata, Microdactylus cristatus, Monasa leucops, M.
tranquilla, Nonnula 7iibecula, Odontophorus capueira, Perdix den-
tata, Podager nacunda, Tetrao turn, TinamAis, species, T. tataupa.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle finely striated trans-
versely. Lateral alae well developed, 1.37 mm. long by 65/i. wide,
extending from head to level of median part of bulb. Mouth irregu-
lar hexagonal, with 6 papillae arranged in two lateral series. Buccal
capsule 48/t deep; 3 rectangular teeth, 19/* high, at entrance to
esophagus. Esophagus 1.04 mm. long. Bulb spherical, 205/* in
diameter.

Male 4.36 to 12 mm. long by 308/x wide. Tail (fig. 187) conical,
with straight chitinous appendage 102/x long. Anus 186/i from end
of body. Preanal sucker fusiform, without chitinous ring, 169/*
long, its posterior end being 450/* from caudal extremity. Caudal
alae rudimentary. Eleven pairs of papillae of which 3 are preanal,
2 adanal, and 6 postanal. Spicules equal, 1.18 mm. long. Guber-
naculum 169/* long.

Female 5.6 to 18.7 mm. long by 411/* wide. Tail acute, ending in
a chitinous appendage 102/* long. Anus 1.05 mm. from posterior end.
Vulva very slightly salient, a little anterior to middle of body.
Ovejector 950/* long; vestibule sinuous, sphincter small; trompe
equivalent to % the length of ovejector. Two divergent uteri.
Eggs 84/* long by 67/* wide.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82.)

Distribution. — South America (Brazil).

SUBULURA SUBULATA (Rudolphi. 1819) Railliet and Henry. 1914

Synonyms. — Ascaris subulata Rudolphi, 1819; Heterakis subulata
Schneider, 1866.

]

NEMATODE PARASITES OF BIRDS

129

Hosts. — CapHmulgus aegyptius saharae and 0. rup'coWs. Other
reports, at least part of which are probably confused with other
species, are: Antrostomus vocifo'us, CapHmulgus candicans, C.
cortopan, C. diumis, C. europaeus, C. gwiatiensis, 0. mercurvus, 0.
natfereri, 0. scaphiuris, C. semitorquaius, C. trifurcus, Chordeiles
semitorquatus, Cuculus cay arms, C. melacoryphus, V. naevius, C.
tinguazu, Nyctihius aetherem, N. grandis.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Body equally slender through-
out; head not distinct, without lips or lateral alae. Valves small and
inconspicuous.

Male, according; to the original description, 16 to 18 nun. long;
according to Seurat, 7.5 mm. long. Tail slender, without caudal

Figs. 186-190. — 186, Subulura similis. Male tail. After Gendre, 1909.
187, Subulura strongylina. Male tail. After Teavassos, 1913. 188,
Subulura suctokia. Male tail. 189, Head, front view. Figs. 188-189
from Barreto, 1918 after Drasche, 1882. 190, Ovejector. After
Barreto, 1918

alae. Cloacal aperture 220/x from posterior extremity (in Seurat 's
specimens). Preanal sucker elliptical, elongate. Ten pairs of
papillae, the first (that is, most posterior) 3 pairs united by short
cuticular alae; 3 pairs of papillae are adanal; the most anterior pair
of papillae is at level of sucker. Spicules unequal, 1.2 and 1.8 mm.
long. Gubernaculum triangular, 110/x, long.

Female, according to the original description, 22 to 22.5 mm. long:
according to Seurat, 12 mm. long by 310pi wide. Esophagus, includ-
ing bulb, 1/8, tail 1/16 of total body length; tail conical, ending in a
fine point. Vulva conspicuous, slightly salient, situated anterior to
middle of body, 4.3 mm. from head end in Seurat's specimens. Ove-
jector with very short sphincter. Eggs 85ja long by 56/i wide.

130 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably similar to that of Ascaridia
(/alii (p. 82).

Distribution. — Europe (Austria (Vienna Museum), Spain, and
Corsica.)

SUBULURA SUCTORIA (Molin, 1860) Railliet and Henry, 1912

Synonyms. — Heterakis suctoria Molin, 1860; Allodapa suctoria,
(Molin. 1860) Seurat, 1914.

Hosts. — Athene noctua, Caprimulgus campestris, C. europaeus, C.
nigrescens, C. rtifus, C. species, C. vociferus, Coturnix delagorgnei,
Dicholophus margrafi, Francolinus oicalcaratus, Gallus gallus, Helio-
treptus anomalus, Hydropsalis climacocercus, Lurocalis semitor-
qiiatus, Meleagris gallopavo, Microdactylus cristatus, Numida melea-
gris, N. papillosa transvaalensis, Nyctibius aethereus, N. grandis, N.
jamaicensis, Nyctidromus albicollis, "Otis houbara' 1 (Houbara mac-
queeni or H. undvJata), Podager nacunda, Pternistes swainsoni,
Sephina francolinm, Stenopsis candicans.

L oca tion . — Cec a .

Morphology.— Subulura (p. 104) : Filiform worms, the anterior
portion curved with the curvature toward the dorsal face. Lateral
cephalic aiae small, terminating at the level of the middle of the
esophagus. Mouth with indefinite lips, provided with 2 lateral
groups of 3 papillae each. (Fig. 189.) Mouth cavity small, cylin-
drical, with thick chitinous walls, and with 3 triangular teeth 25^
long. Esophagus dilated slightly in the posterior portion and ending
in a piriform bulb.

Male 11.8 to 13.8 mm. long by 359/a wide. Posterior extremity in-
fundibular, straight, and terminating in a short appendix. Cloacal
aperture 211ju, from posterior end. Sucker ellipsoidal, without chi-
tinous wall, 135[x long; it is 296/x from the cloacal aperture. Caudal
alae slightly developed. There are 11 pairs of caudal papillae (fig.
188) arranged as follows: Three pairs of large, ventral preanal, of
which 1 pair is at the posterior end of the anterior third of the
sucker, 1 pair just behind the sucker, and 1 pair somewhat anterior
to the cloacal aperture ; 2 pairs of large adanal papillae ; and 6 pairs
of postanal papillae, of which 2 large ventral pairs are near the
cloacal aperture, followed by a large lateral pair and a small ventral
pair, and these by 2 moderately large ventral pairs near the end of
the tail. Spicules equal, curved, pointed posteriorly and infundibuli-
form anteriorly; Gedoelst writes that the end is curved in a hook;
they are 1.02 to 1.15 mm. long, or, according to Seurat, 1.5 mm. long.
Gubernaculum straight, 127 to 150/x long; Seurat says triangular and
175ju long.

Female 20 to 23 mm. long by 539pi wide at the level of the vulva.
Tail straight or slightly bent ventrally and terminating in a small

NEMATODE PARASITES OF BIRDS 131

appendix. Vulva small, not salient, situated near the middle of the
bod}\ Ovejector (fig. 190) relatively short (779/*), extending an-
teriorly, vestibule small (313/*) and heavily chitinized internally,
sphincter comparatively long (254/*) and usually containing 3 to 6
eggs. Two divergent uteri. Anus 1.156 mm. anterior to tip of tail.
Eggs elliptical, thin-shelled, and embryonated when deposited; 51y.
long by 45/* wide.

For the descriptions given for this worm by Gendre, by Seurat
and by Gedoelst see Sublura brumptl (p. 112).

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil, .French Guinea) and
Africa (Dahomey, Egypt, Algeria and Tunis).

SUBULURA TBAVASSOSI Barreto, 1918

Synonyms. — Ascaris forcipata Rudolphi, part; Heterakls forcl-
paria Schneider, 1866, part; Subulura strongylina Railliet and
Henry, 1912, part.

Hosts. — Bucco chacuru, B. collaris, B. macrorhynchus, B. rufiven-
tris, B. striolatus, B. swainsoni, B. tamatia, B. tectus, Chelidoptera
tenebrosa, Malacoptila torquata, Monacha morpheus, M. nigra,
Monusa nigrifrons, Nonnula rubecula.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle with fine transverse stria-
tions. Lateral alae very narrow (33/* wide), disappearing at the
level of the bulb. Buccal cavity 42/* deep.

Male 3 to 7. 7 mm. long by 230 to 360/* wide. Esophagus 1.03 mm.
long, bulb 231/* in diameter. Cloacal aperture 127|U from posterior
extremity. Preanal sucker elliptical, 385/x long, 508/a from caudal
extremity. Eleven pairs of papillae (figs. 192, 193, and 194), of
which 3 are preanal, 2 adanal, and 6 postanal. Spicules equal, 1.71
mm. long. Gubernaculum 211/x long.

Female 6 to 10 mm. long by 400 to 410/* wide. Esophagus 1.16 mm.
long; bulb 254//. in diameter. Vulva not salient, a little anterior to
middle of body. Ovejector (fig. 191) relatively short (771,u) ; vesti-
bule 406/i, sphincter 169/x, trompe 296/*. long. Two uteri, divergent.
Eggs elliptical, 77/* long by 58/* wide, embryonated when deposited.

Life history. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution.— -South America (Brazil).

SUBULURA TROGONI Barreto, 1918

Host. — Trogon viridis.
Location. — Intestine.

Morphology. — Subulura (p. 104) : Cuticle with transverse striations.
Lateral alae extend from head to initial part of intestine, a length

132

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

of 223/*; width 55/*. Mouth with 6 small equal papillae. Buccal
cavity (fig. 195) 45/* long by 38/* wide. Three subrectangular teeth
at entrance to esophagus. Esophageal bulb subspherical.

Male 7.4 mm. long by 572/* wide. Tail (fig. 196) slender, slightly
curved ventrally, with pointed appendage 93/* long. Cloacal aper-
ture 232/* from posterior end. Preanal sucker 186/t long, without
chitinous ring, 423/* from caudal extremity. Caudal alae rudimen-
tary. Eleven pairs of papillae, of which 3 are preanal, 2 adanal, 6
postanal. Spicules equal, 558/* long by 38/* wide. Gubernaculum
228/* long.

I9L

Figs. 191-194. — Subulura travassosi. 191, Ovejector. 192-193, Male tail. (Figs.
191, 192, and 193 from Bucco chacuru.) 194, From Bucco swainsoni. Male tail.
All after Barreto, 1918

Female 15.5 mm. long by 858/* wide. Caudal extremity acute, with
appendage 186/* long. Anus 1.2 mm. from posterior extremity.
Vulva small, only slightly salient, anterior to middle of body.
Ovejector with muscular layer poorly developed; vestibule retort-
shaped; trompe sinuous, difficult to differentiate. Two divergent
uteri. Eggs almost spherical, 68/* long by 60/* wide, embryonated
when deposited.

Life Mst&ry. — Unknown; probably similar to that of Ascaridia
galli (p. 82).

Distribution. — South America (Brazil).

SUBULURA, species Baylis and Daubney, 1922

Host. — Tumix, species.
Location. — Not given.

NEMATODE PARASITES OF BIRDS

133

Morphology. — Subulura (p. 104) : Lateral alae narrow, extending
from head to level of prebulbar esophageal swelling.

Male unknown.

Female 14 mm. long by 410/a wide. Head 80^ in diameter; buccal
cavity 35^ deep by 20y wide; esophagus 1.1 mm. long; bulb 150*i in
diameter. Tail acutely pointed; anus 320/x from posterior end.
Vulva 6.1 mm. from anterior end. Eggs 85/* long by 5G/* wide, em-
bryonated.

Figs. 195-196. — Scbulura trogoni. 195, Anterior end. 196, Male tail. After

Barreto, 1918

Life history. — unknown: probably similar to that of Ascaridia
galli (p. 82).

Distribution. — Asia (India).

SUBULURA (?) ACUTICAUDA (Linstow, 1901) Railliet and Henry, 1914

Synonym. — Oxysoma acuticauda Linstow, 1901.

Host. — Numida rikwae.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Head end rounded, with no
mouth cavity ; esophagus with spherical bulb.

Male 10.5 mm. long by 390/* wide. Esophagus 1/10.7, tail 1/38 of
total length; 8 pairs of caudal papillae (fig. 197) of which 3 are
preanal, 5 postanal. Preanal sucker elongate. Spicules equal, 880/*
long.

134

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 14.8 mm. long by 510 ^ wide. Esophagus 1/12.5, tail 1/18.7
of total length. Vulva anterior to middle of body, dividing body
length in ratio of 11 : 14. Eggs 47/x long by 34/a wide.

Life history. — Unknown; probably similar to that of Ascaridia
gulli (p. 82).

Distribution. — Africa (Usanga, Eukwa-See).

Figs. 197-199.' — Subulijra achticauda. 197, Male tail. After Linstow, 1901.
198, subulura gracilis. head. 199, male tail. after llnstow, 1899

SUBULURA (?) GRACILIS (Linstow. 1899) Railliet and Henry, 1914

Synonym. — Oxysoma gracile Linstow, 1899a.

Host. — Francolinus, species.

Location. — Intestine.

Morphology. — Subulura (p. 104) : Head (fig. 198) with mouth
cavity with strongly developed chitinous walls; on the anterior and
outer edge 2 small papillae. Esophagus ends in bulb with valve-
teeth ; tail long and pointed in both sexes.

Male 7.9 mm. long by 310/* wide. Esophagus 1/8, tail 1/40 of
total length. Spicules 590/x long, sword-shaped. Nine pairs of
caudal papillae (fig. 199) of which 3 are preanal, 6 postanal.

Female 9.48 mm. long by 390/x wide. Esophagus 1/9.3, tail 1/8 of
total length. Vulva slightly in front of middle of body, dividing the
body length in ratio of 11 : 13. Eggs 57/u. long by 44/a wide.

NEMATODE PARASITES OF BIRDS 135

Life history. — UnknoAvn: probably similar to that of Ascaridia
(j alii (p. 82).

Distribution. — Europe (Germany (Berlin Zoological Garden) ).

Family ASCARIDAE Baird, 1853

Family diagnosis. — Ascaroidea (p. 48) : Polymyarian. Mouth with
Si prominent lips supplied with papillae, the dorsal lip being median
and the 2 others submedian and approximated in the ventral line,
or with 3 main lips and 3 relatively prominent or inconspicuous inter-
mediate lips (interlabia). Male usually with two spicules. Caudal
extremity of female terminates conically and fairly abruptly.

Type-genus. — Ascaris Linnaeus, 1758.

Subfamily Anisakinae Railliet and Henry, 1912

Subfamily diagnosis. — Ascaridae (p. 135) : Cuticle with cross stria-
tions but without cuticular spines or other raised structures.
Esophagus may or may not be divided into anterior muscular por-
tion and posterior ventriculus. Anteriorly directed cecum often
present, springing from intestine and lying alongside of esophagus.
A posteriorly directed solid glandular esophageal appendix may also
be present. Interlabia present or absent. Dentigerous ridges on
lips present or absent.

Parasitic in alimentary canal of mammals, birds, reptiles, and
fishes, the hosts usually being aquatic or at least fish-eating. Inter-
mediate hosts, such as a fish, probably necessary in some if not all
cases.

Type-genus. — Anisakis Dujardin, 1845.

This diagnosis is the diagnosis of Railliet and Henry as emended
by Baylis, 1920, to include part of the Heterocheilinae of the former
authors.

KEY TO CEN'KRA OF ANISAKINAE

Intestinal cecum and esophageal appendix both present Contracaecum, p. 146.

Intestinal cecum present; esophageal appendix absent Porrocaecura, p. 135-

Genus PORROCAECUM Railliet and Henry, 1912

Sy7i&nym. — Terranova Leiper and Atkinson, 1914.

Generic diagnosis. — Anasakinae (p. 135) : Esophagus with anterior
muscular portion and posterior ventriculus of oblong shape, the
latter short in the genotype but in other species frequently long and
bent at an angle so as to open into the intestine laterally. Intesti-
nal cecum present. Esophageal appendix absent. Interlabia pres-
ent, usually small. Dentigerous ridges usually present.

Parasitic in intestine of birds, marine mammals, and fishes.

136 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

l J ype-species. — Porrocaecum, crassum (Deslongchamps, 1824) Rail-
liet and Henry, 1912.

KEY TO SPECIES OF PORROCAECUM

1. Larva only known, in eel, Anguilla vulgaris Porrocaecum anguillae, p. 146.

Adults known, In birds 2.

2. Lateral membranes present 3.

Lateral membranes absent or undescribed 5.

3. Cecum long (about 3 mm.) Porrocaecum serpentulus, p. 142.

Cecum short (apparently not longer than ventriculus) 4.

4. Lips with 1 papilla, situated medianly ; female tail 1/50 to 1/63 of total length ;

eggs 91/4 long by 85/i wide Porrocaecum semiteres, p. 141.

Lips with 2 large oval papillae ; female tail 1/46 of total length ; eggs 110/t
long by S5/i wide Porrocaecum ensicaudatum, p- 139.

5. Species unrecognizable ; possibly identical with F. ensicaudatum.

Porrocaecum heteroura, p. 144.
Species recognizable 6.

6. Conspicuous gubernaculum present ; male with 3 pairs of postanal papillae

(2 of them very small, on the caudal appendage, the other pair large but
simple, a little posterior to cloacal aperture).

Porrocaecum reticulatum, p. 141.
No gubernaculum described ; postanal papillae, where described, more nu-
merous than 3 pairs except in P. spirale, where the pair near the cloacal
aperture is made up of double papillae 7.

7. Vulva posterior to middle of body 8.

Vulva anterior to middle of body 9.

8. Lips described only as convex in shape ; male not over 30 mm., female not

over 53 mm. long ; from Anas, species and Numida meleagris.

Porrocaecum crassum, p. 136.
Lips six-sided, with digitif orm projections of pulp ; male up to 48 nam.
long ; female up to 64 mm. long ; from other hosts than above.

Porrocaecum spirale, p. 143.

9. Female 154 mm. long. Male said to have caudal alae anterior to the

cloacal aperture, supported by 5 pairs of papillae ; from Colymbus

auritus Porrocaecum praelongum, p. 140.

Female not over 112 mm. long ; male without caudal alae ; from other hosts

than above 10.

10. Dorsal lip with pulp divided into 2 bipartite lobes and between them a
rounded lobe ; pulp not notched toward base but follows outline of lip.

Porrocaecum depressum, p. 137.
Pulp divided into 2 rounded lobes from the inner surface of which project
2 broad flat plates ; toward base of lip pulp sharply notched with a promi-
nent cuticular band opposite the notch Porrocaecum angusticolle, p. 137.

This key does not include Ascaris kirghisensis (p. 145), which the
writer has placed tentatively in Porrocaecum but which may prove
to be a species of Contracaecum.

PORROCAECUM CRASSUM (Deslongchamps, 1824) Railliet and Henry, 1912

Synonym. — Ascaris crassa Deslongchamps, 1824.
Hosts. — Anas boschas, A.b. domestica, A moschuta, Numida mele-
agris.
Location. — Small intestine.

NEMATODE PARASITES OF BIRDS 137

Morphology. — Porrocaecum (p. 135) : Mouth with 3 convex lips.
Cuticle with marked transverse striations. Muscular esophagus
claviform, followed by a ventriculitis and accompanied by a narrow
cecum originating at the base of the intestine and running anteriorly

Male 12 to 30 mm. long by 500^. to 1.2 mm. wide. Tail slender,
conical, mucronate, without lateral alae; cloacal aperture 1/46.4 of
total length from posterior end. Two spicules, 52G> long, curved,
alate, with rounded points.

Female 43 to 53 mm. long by 2.2 mm. wide. Tail slender, pointed,
straight. Anus about 1/55 of total length from posterior end.
Vulva slightly posterior to middle of body, dividing body length in
ratio of 13:11. Eggs lOO/i. long, globular.

Life history. — Unknown.

Distribution. — Europe (France and Germany).

PORROCAECUM ANGUSTICOLLE (Molin. 1860) Baylis and Daubney, 1922

Synonym. — Ascaris angusticollis Molin, 1860.

Hosts. — Archibuteo (Buteo) lagopus, Buteo buteo (Buteo vul-
garis), Circus aeruginosus, Falco hali-aetus, Helotarsus albicilla, H.
ecaudatvs, Milvus govinda, Pandion haliaetos, Pernis species (prob-
ably P. apivorus) .

Location. — Intestine.

Morphology. — Porrocaecum (p. 135) : Dorsal lip (fig. 200) hexag-
onal, pulp with 2 lobes rounded anteriorly and joined by a saddle;
from the inner surfaces of the lobes 2 projecting plates, flattened
and expanded distally. Toward base of lip on each side a cuticular
band. A pair of papillae and dentigerous ridges present. Inter-
labia small, triangular. Esophagus, including ventriculus, 4.8 mm.
long; cecum 2.7 to 3 mm. long.

Male up to 55 mm. long by 1.1 mm. wide. Tail (fig. 201) conical,
390/* long. A distinct constriction halfway from cloacal aperture to
tip of tail. Five pairs of postanal papillae, 4 of them in posterior
half of tail, the other pair made up of double papillae, not far
posterior to cloacal aperture. Spicules equal, 950/x long, not alate.

Female 40 to 90 mm. long by 500/x to 1.5 mm. wide. Tail 700/x
long, blunt ; caudal papillae 200/x from end. Vulva in anterior half
of body, dividing body length in ratio of 3:5. Eggs 85 to 93/a
long by 58 to 74/x wide.

Life history. — Unknown.

Distribution. — Europe (Austria (Vienna Museum)), Africa
(Egypt), and Asia (India).

PORROCAECUM DEPRESSUM (Zeder, 1800) Baylis, 1920b

Synonyms. — Fusaria depressa Zeder, 1800; Ascaris depressa
(Zeder, 1800) Rudolphi, 1809.

138

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Accipiter bicolor, A. nisus, Aquila albicilla, A. chrysaetos,
A. fasciata, A. imperialis, A. naevia, A. pennata, Archibuteo lagopus,
A. vulgaris, Astur palumbarius, Bubo -maximus, B. virginmnus, Cir-
caetus gallicus, Gircestus pectoralis, Circus aeruginosus, C. cinera-
ceus, G. cyaneus, G. rufus, Falco aesalon, F. albicilla, F. apivorus, F.
ater, F. brachy dactyl us, F. brasiliensis, F. buteo, F. chrysaetos, F.
cyaneus. F. degener, F. gallicus, F. imperialism F. lithofalco, F.
lanarius, F. lagopus, F. milvus, F. naevius, F. nisus, F. palumbarius,
F. penrtaius, F. peregrinus, F. rufus, F. rutilans, F tinnunculus,
Gypaetus barbatus, Gyps fulvus, Haliaetus albicilla, Milvus ater,
Milvus milvus, M. regalis, Nisaetus fasciatus, Strix aluco, S. brachy-
otus, 8. bubo, S. dasypus, S. flammea, S. nyctea, S. otus, S. stridula,
S. tangmalmi, Vultur cinereus, V. fulvus, V. monachus. Larvae in
Sorex tetragonurus and Talpa europaea.

L ocatiori . — Intestine.

o/mm

Figs. 200-202. — 200, Porrocaecdm angusticolle. Dorsal lip. 201, Male tail.
202, porrocaecum depressum. dorsal lip. after baylis and daubney, 1922

Morphology. — Porrocaecum (p. 135) : Large lips (fig. 202) with
dentate edges and with pulp projecting in 2 bipartite processes,
and internal to them a large median lobe, rounded anteriorly, dis-
tinctly visible where it projects beyond the saddle joining the 2
main lobes; 2 large papillae on outer surface. Small interlabia
sharply pointed.

Male 24 to 100 mm. long. Six pairs of postanal papillae (fig.
203), the most anterior pair composed of double papillae near the
cloacal aperture and the other 5 pairs on a conical prolongation of
the posterior end. Seventeen pairs of preanal papillae.

Female 30 to 112 mm long. Vulva in anterior third of body,
according to Linstow; Schneider says 37 mm. from head end in a
97 mm. long specimen. Eggs with double shells, thickened at the
poles and with fine dots (Punktchen).

NEMATODE PARASITES OF BIRDS 139

Life history. — Unknown, although larval forms of this species
{Ascaris incisa Rudolphi) have been reported as found encapsulecl
in the peritoneum of the shrew and mole (see above, under Hosts).

Distribution. — Europe, Asia (Russian Turkestan and India),
South America (Brazil and British Guiana) and Africa (Trans-
vaal).

PORROCAECUM ENSICAUDATUM (Zeder, 1800) Baylia, 1920b

Synonyms. — Fusaria ensicaudata, Zeder, 1800; Ascaris ensicaudata
(Zeder, 1800) Rudolphi, 1809.

Hosts. — Acrocephalus amndinaceus, Aland a species. Anas boschas,
Charadrius dubius, C. hiaticula, C. moAnellus, C. pluvialis, Gallinula
chloropus, Hirnantopus melanopterus, Luscinia philomela, Merula
nigra, Mimus polyglottus, Motacilla alba, Oedicnemv.s crepitans,
" rooks," Pica caudata, Pluvialis apricarius, Salicaria turdoides,
Sguatarola helvetica. Stum us vulgaris, Sylvia turdoides. Tnrdus
iliacus, T. musicvs, T . mcrvla, T. pilaris, T . saxatilis, T. torquatus,
T. viscivorus, Vanellus cristatus, V. meJanog aster.

Location. — Intestine.

„ o o o o o«

Fig. 203. — Fobbocakcum bhpkessum. Male tail. Afteh Linstovv, 1875

Morphology. — Porrocaecum (see p. 135) : Large lips wider than
long, the pulp with 2 wing-like projections; each lip (fig. 204a)
with 2 large, oval, closely set papillae. Interlabia 3/5 the length
of large lips, with rounded ends. Lateral membranes present, 120/t*
wide. Intestinal cecum small, almost rudimentary.

Male 28 to 32 mm. long by 1.02 mm. wide. Esophagus 1/9.9, tail
1/8G of total body length. Body narrows suddenly posterior to
cloacal aperture. Six pairs of postanal papillae (fig. 2046), 5 of
which are near the caudal extremity (4 ventral, 1 lateral) and 1
pair, composed of double papillae, near cloacal aperture. Preanal
papillae in single row, numerous (13 to 19).

Female 50 to 58 mm. long by 1 to 1.8 mm. wide. Tail 1/46 of
total body length, conical. Vulva somewhat anterior to middle of
body, dividing body length in ratio of 4:5. Eggs (fig. 204c) 110/x
long by 85/i wide, the outer shell with delicate lattice-work markings.

Life history. — Unknown.

Distribution. — Europe.

140 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

PORROCAECUM PRAELONGUM (Dujardin, 1845) Baylis, 1920b

Synonym, — Ascaris praelonga Dujardin, 1845.

Host. — Colymhus auritus.

Location. — Intestine.

Morphology. — Porrocaecum (p. 135) : Body white, filiform, very
elongate. Head proportionately large, 400/t wide. Esophagus 4.3
mm. long by 400/* wide, followed by a ventriculus 400/* long and
accompanied by a thick cecum 3 mm. long by 600/* wide, joining
with the intestine. Cuticle with very conspicuous transverse stria-
tions.

Fig. 204. — Purrocaecum bnsicaddatum. c, Dorsal lip; b, male tail; c, egg.
After, Linstow., 1884

Male 90 mm. long by 900/* wide. Tail suddenly narrowed and end-
ing in a short conical point ; cloacal aperture, with a large tubercule,
330/* from posterior end. Membranous caudal alae present, anterior
to cloacal aperture, supported by 5 pairs of papillae. Spicules equal,
900/* long by 58/* wide.

Female 154 mm. long by 1.3 mm. wide. Tail straight, conical,
pointed ; anus 600/* from posterior end. Vulva just posterior to
anterior fourth of body, 44 mm. from the head. Ovejector (?)
sinuous, 45 mm. in length, divides into 2 uteri. Eggs globular,
measuring 110 to 112/*, with reticulated shell.

L ife Ji istory. — Unknown.

NEMATODE PAEASITES OF BIRDS 141

Distribution. — Europe (Austria (Vienna Museum)).

Several earlier authors have made this species a synonym of Con-
tracaecum spiculigerum, but the description of the 2 species shows
them to be different and the present species is seen to belong in
Porrocaecum as placed by Baylis.

PORROCAECUM RETICULATUM (Linstow, 1899) Baylis and Daubney, 1922

Synonym. — Ascaris reticulata Linstow, 1899.

Baylis and Daubney list Ascaris ardeae Smith, Fox, and White,
1908, from Ardea herodias, as a synonym of the species, but in view
of the fact that the latter authors compared their species with that
of Linstow and stated that it was different and as they described the
esophagus merely as expanding posteriorly in clavate fashion, with
a valve-like opening into the intestine, and the intestine as simple,
it appears that they alone can determine as to whether or not it is a
distinct species.

Hosts. — Ardea einerea, A. cocoi, A. maniUeitsis, N yctico-rax griseus,
and " an egret."

Location. — Intestine.

Morphology. — Porrocaecum (p. 135) : Interlabia and dentigerous
ridges absent according to Linstow, present according to Baylis and
Daubney. Dorsal lip (fig. 206) 350,* long by 280 M wide at the base;
pulp with 2 round converging projections anteriorly; 2 papillae a
little above middle of lip, above lateral pulp projections. Esophagus
1/15 of total length, with short oblong ventriculus; cecum well
developed, running forward beside the esophagus for a considerable
portion of the length of the latter.

Male (size not given). Tail (fig. 205) with finger-like appendage,
with 2 very small pairs of papillae; an additional postanal pair of
large papillae just anterior to the constriction. Preanal papillae
number 5 pairs. A gubernaculum present, according to Baylis and
Daubney.

Female 82 mm. long by 1.0 mm. wide. Eggs 110/a long by 91/x
wide, according to Linstow; those of the specimens of Baylis and
Daubney somewhat smaller. Shell reticulate.

Life history. — Unknown.

Distribution. — Africa (Porto Alegre), and Asia (India (Calcutta
Zoological Garden)).

PORROCAECUM SEMITERES (Zeder, 1800) Bayliss, 1920

Synonyms. — Fusaria semiteres Zeder, 1800; Ascaris semiteres
(Zeder, 1800) Rudolphi, 1802.

Hosts. — Corvus comix, Pluvialis apricarius, Tringa qanellus s
Vanellus cristatus, V. melanogaster.

Location. — Intestine.

»•

142 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Porrocaecum, (p. 135) : Body white. Head (fi
207) 230 to 300/x wide, with 3 large convex lips each carrying a pa-
pilla in the middle of the convexity. Esophagus 2.8 mm. long,
followed by a ventriculus from the side of which there arises a short
cecum joining with the intestine. Lateral membranes present
throughout whole body length, wide near the head and narrow the
remainder of the length. Cuticle with very pronounced cross-stria-
tions.

Male 18 to 53 mm. long. Tail slender, conical. Two spicules,
wide, slightly curved.

Female 40 to 63 mm. long by 900 to 1.14 mm. wide. Tail straight,
slender; anus 800//, from the posterior end. Vulva situated at 2/5
of the length of the body from the head end (22 mm. from cephalic

Z06.

207.

Figs. 205-208.— 205, Porrocaecum reticulatum. Male tail. After Bay-
lis and Daubney, 1922. 206, Dorsal lip. After Linstow, 1899. 207,
Porrocaecum semiteres. Anterior end (dissected). After Creplin,
1829. 208, Porrocaecum serpentulus. Male tail. After Monnig, 1923

extremity in the large specimens). Ovejector (?) slender, filiform,
running anteriorly for 3 mm., then turning and running posteriorly
for 4 mm. at which point it divides into 2 parallel branches. Eggs
elliptical, 91,u long by 84^. wide, with thick reticulated shell.

Rife history. — Unknown.

Distribution. — Europe.
Lewis (1926,10) has recently expressed doubt as to the authenti-
city of this species ; from a study of specimens he thinks it probably
the same as P. ensicaudatum.

PORROCAECUM SERPENTULUS (Rudolphi, 1809) Baylia. 1920b

Synonyms. — Ascaris serpentulus Kudolphi, 1809; Ascaris ardeae
Froelich, 1802.

Host*. — Anthropoides virgo, Ardea agami, A. cinerea, A. ccerulea,
A. grus, A. major, A. melanocephala, A. nycticorax, A. pileata, A.
purpurea, A. scapularis, A. violaeea, Grus australmiana, G. cinerea,

NEMATODE PARASITES OF BIRDS

143

G. communis, G. pavonina, Nycticorax nycticorax, Phoenicopterus
roseus.

Location. — Intestine.

Morphology. — Porrocaecum (p. 135) : Large lips (fig. 209) with
dentigerous ridges; length of lips 210/x, width at base 78/*, at level
of the 2 papillae 180/t. Interlabia large, ear-shaped, 2/3 the size of
the large lips. Esophagus 3.4 mm., ventriculus 400/x., intestinal
cecum 3 mm. long, in a 55 mm. long specimen. Lateral membranes
very wide in the anterior 3 mm. of their length; they extend the
entire length of the. body.

Various lengths have been given for this species; 13 to 39 mm.,
50 to 52 mm., and for the female 160 mm. long by 2 to 2.5 mm. wide.
A male collected bv Monnig measured 55 mm. long bv 1 mm. wide.

Pics. 209-210. — I'okrocaecim sbupentulus. 209, Head. After Linstow,
1899. 210, Male tail. After Baylis and Daubney, 1922

Male tail (figs. 208 and 210) with digitiform prolongation, on the
subdorsal and ventral surfaces of which are 2 pairs of papillae (a
total of 4 pairs) and a fifth pair is lateral; in addition 15 pairs of
preanal and 1 pair of double postanal papillae. Spicules 1.25 mm.
long in a moderately large specimen (Monnig describes them as
"apparently" 260/x long in his specimen), with wide alae.

Female (see above for size). Eggs 104/a long by 78/i wide, the
outer surface reticulate.

Life history. — Unknown.

Distribution. — Europe, South America (Brazil), North America
and Africa (Transvaal).

PORROCAECUM SPIRALE (Rudolphi, 1795) Baylis, 1920b

Synonyms. — Ascaris spiralis Rudolphi, 1795; Fusaria spiralis
(Rudolphi, 1795) Zeder, 1803.

3612—27 11

144

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Aegoltus otus, A. brachyotus, Bubo maxim/us, B. virgiud-
ahus, Falco pygargus, Flammea flammea, Nyctale tengmalmi, Otus
brachyotus, O. vulgaris, Strix aluco, S. bubo, & -flammea, S. nivea, S.
noctua, S. stridula, Syrnia nyctea, S. aluco, Ulula aluco.

Location. — Intestine.

Morphology. — Porrocaecum (p. 135) : Large lips (fig. 211) six-
skied, with dentate band anteriorly and the pulp with digitiform
projections; each lip with 2 papillae. Interlabia sharply pointed.

Male up to 48 mm. long by 1.5 mm. wide. Tail (fig. 213) with a
conical appendage with 2 pairs of papillae; slightly posterior to the
cloacal aperture 1 pair of double papillae. Six pairs of preanal
papillae, 4 of which form a group about halfway between the other,
most anterior, 2 pairs and the cloacal aperture. Spicules sickle-
shaped.

Figs. 211-213. — Porrocaecum spiralb. 211, Dorsal lip.
Male tail. After Linstow, 1875

212, Egg. 213,

Female up to 64 mm. long by 1.7 mm. wide. Vulva slightly pos-
terior to middle of body, dividing body length in ratio of 8 :7. Eggs
(fig. 212) elliptical, 102/* long by 60/* wide, with double shell, the
outer thickened at the ends and with small glittering prominences.

Life history.— Unknown.

Distribution. — Europe.

This species has been listed by several authors as a synonym of
Porrocaecum depressum but Linstow (1875) compared the two and
stated they were distinct species. The descriptions show the caudal
papillae of male and position of vulva of female to differ in the two
species.

PORROCAECUM HETEROURA (Creplin, 1829 emend. Mehlis, 1831) Baylis, 1920b

Synonyms. — Ascaris heteriira Creplin, 1829; Asearis heteroura
Mehlis, 1831.

Hosts. — Gharadrius morinellus, C. oedicnenius, C. pluvialis,
Himantopus melanopterus, Squatarola helvetica, Sturnus vulgaris,
Turdus musicus.

Location — Intestine.

Morphology.— Powocaecum (p. 135) : Body 14 to 30 mm. long,
thick, without lateral alae. Lips large (fig. 214) .

NEMATODE PARASITES OF BIRDS

145

Male with short, slender tail. Spicules of median length.

Female with short tail.

Life history. — Unknown.

Distribution. — Europe (Ireland, Germany, and Italy (Portofer-
rajo)).

This species is given as a synonym of P. ensieaudatum by numerous
authors but P. ensioauddtum has lateral membranes whereas they
are said to be absent in this species. It is therefore left as a distinct
species by the present writer, although it is unrecognizable from the
present description.

yirai

^ —

2/S

PORROCAECUM KIRGHISENS1S (Skrjabin. 1916) Cram, 1927

Synonym. — Ascaris Mrghixensis Skrjabin, 1916; Ascaris
sen-sis Skrjabin, 1916 (evidently misspelled).

Host. — Aquila imperialis.

Location. — Small intestine.

Morphology. — Porrocaecum
(p. 135) : Cuticle transversely
striated. Head with lips and
interlabia (fig. 215c). Lips
(fig. 2156) made up of 3 parts,
the central one with denticulate
e<]tre and with the pulp divided
into 2 equally large lobes. Two
small papillae situated far an-
terior on lips.

Male 75 to 99 mm. long by 1.3
to 2 mm. wide. Tail (fig. 215 d
and e) conical, pointed; cloacal
aperture 500/x from end. Spic-
ules equal, 1.18 mm. long. Five
pairs of postanal papillae, the
most anterior pair of these com-
posed of double papillae. Four-
teen pairs of preanal papillae.

Female (immature) 80 mm. long by 1.3 mm. wide.

This species, according to Skrjabin, is closely related to P. depres-
sum but differs distinctly in the shape of the lips and the disposition
of the papillae on the lips.

The allocation of this specie's to Porrocaecum is made tentatively
by the present writer' on the basis of Skrjabin's statement that it is
very close to P. depn fsitm. The main differential characters of the
two genera Porrocaecum and Contracae< -u//t. that is, the nature of
the esophageal appendages, not being known in connection with this
species, the allocation can not lie made with certainty but it is

Figs. 214— 215.— 214, Porrocaecum hete-
roura. Anterior end (dissected). After
Creplin, 182!). 215a, Porrocaecum an-
gdillie. Anterior end (dissected).
After Linstow, 1890. 215 b to e, Por-
HOCAECUM kiri;h!Sens;s. 6, Lip; c, in-

TEKl.AUII M ; (/. LATERAL AND C, VENTRAL
VIEW OF MALE TAIL. AFTER SKUJAUI...

1916

146 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

thought preferable to place it in Porrocaecum where attention will
be drawn to it than to leave it in the genus Ascaris, which position
is known to be incorrect.

PORROCAECUM ANGUILLAE (Linstow, 1899) Cram, 1927

/Synonym. — Ascaris anguillae.

Linstow described this as a larva from the eel, Anguilla vulgaris.
The larvae were rolled up and often encapsuled in the abdominal
organs. The adult may possibly be found in water birds; therefore
a brief description follows: Length of larva 38 mm., width 970/x.
Esophagus 1/9, tail 1/198 of total length, the tail rounded with a
small digitiform prolongation. The posterior 5/13 of the esophagus
(fig. 215a) is a ventriculus from which there is a cecum, anteriorly
directed, equal in length to the ventriculus, joining it to the intestine.

Genus CONTRACAECUM Railliet and Henry, 1912

Synonym. — Kathleena Leiper and Atkinson, 1914.

Generic diagnosis. — Anasakinae (p. 135) : Esophagus with reduced
posterior ventriculus, giving off laterally a posteriorly directed
appendix. An intestinal cecum present. Interlabia present, usually
very well-developed. Dentigerous ridges usually absent.

Parasitic in the proventriculus and intestine of fish-eating birds,
in intestine of mammals and of fishes.

Type-species. — Contracaecum spiculigcrum (Rudolphi, 1809)
Railliet and Henry, 1912.

Xi:V TU SI-KCIES OK CONTRACAECUM

1. Male alone described 2.

Both male and female, or female alone, described 3.

2. Male 13 mm. long; esophageal appendix 700/* long; spicules 1.8 mm. long.

Contracaecum engonium, p. 150.

Male 1G.5 to 32.5 mm. long; esophageal appendix 960/t to 1.2 mm. long;

spicules 4 mm. long Contracaecum punctatum, p. 157.

3. Vulva posterior to middle of body dividing body length in ratio of 2 : 1.

Contracaecum multipapillatum, p. 154.
Vulva anterior to middle of body 4.

4. Female alone known; eggs 80 to 90/* long by 60/* wide; from Haliaetus

species Contracaecum haliaeti, p. 150-

Both male and female known ; eggs different from above ; from other hosts
than above 5

5. Pulp of lips with 2 anteriorly directed bipartite processes; interlabia

strongly curved, the anterior ends split into 2 rounded processes; male
witb 3 pairs of double postanal papillae.

Contracaecum micropapillatum, p. 153.
Lips and interlabia different from above; male with more than 3 pairs

of caudal papillae, and not all pairs double 6.

G. Esophageal appendix not more than 1 mm. long 7.

Esophageal appendix more than 1 mm. long 11.

NEMATODE PARASITES OF BIRDS 147

7. Male 26 mm. long, female 30.8 mm. long; cuticle with regularly arranged

refractile dots ; vulva only 1.59 mm. from head end ; eggs 26/a long by 21/i

wide Contracaecum rosarium, p. 158.

Male not over 20 mm. long, female not over 25 mm. long; no dots described
in cuticle ; vulva more posterior than above ; eggs at least 55m long
by 45/* wide 8.

8. Esophageal appendix about 500/*, cecum 1.8 mm. long; spicules unequal,

2.7 and 3 mm. long ; eggs 100/t long by 60/t wide.

Contracaecum scotti, p. 159.

Esophageal appendix 720/t or longer, cecum 2.G mm. or longer ; spicules

equal and either shorter or longer than those above ; eggs not over 6'.*/x

long by 45/t wide 9.

9. Tail of female 130//. long ; vulva divides body length in ratio of 9.5 : 10.5 ; tail

of male S5/i long; spicules 1 mm. long Contracaecum andersoni, p. 149-

Tail of female 300/t or longer; vulva more anterior than in C. andersoni;
tail of male 175/i or longer; spicules 3.6 mm. or longer 10.

10. Dorsal lip with 2 single papillae; esophageal appendix 720/t to 1.04 mm.

long; male with 33 pairs of caudal papillae, 5 of which are postanal, the

most anterior of the 5 being double Contracaecum rodhaini, p. 157.

Dorsal lip with 2 double papillae ; esophageal appendix not over 730/t long ;
male with 24 pairs of caudal papillae, 5 of which are postanal, the third
papilla from the posterior end being double.

Contracaecum magnipapillatum, p. 161.

11. A gubernaculura present, 120/t long; spicules 1.9 mm. long; eggs 40/t long

by 32/t wide Contracaecum praestriatum, p. 156.

Gubernaculum absent or undescribed ; spicules longer than above; eggs
larger than above (except possibly in C. tricuspe where not de-
scribed) 12.

12. Male 13.S mm. long ; female 12 to 17.5 mm. long ; interlabia very large, end-

ing in 3 points ; spicules 4.6 mm. long with curved alae.

Contracaecum tricuspe, p. 160-

Male 18 mm. or longer: female 23 mm. or longer; interlabia different from

above; spicules of different length than above 13.

13. Esophageal appendix 3 to 4.5 mm. long ; interlabia low and rounded ; anus

of female 564/t, from caudal extremity Contracaecum ovale, p. 155.

Esophageal appendix not over 1.9 mm. long ; interlabia different from
above; anus of female not over 500/t from caudal end 14.

14. Vulva of female divides body length in ratio of 5 : 13 ; 7 pairs of postanal

and 38 to 56 pairs of preanal papillae ; spicules probably 7.2 mm. long.

Contracaecum spiculigerum, p. 147.
Vulva of female divides body length in ratio of 2 : 3 ; not more than 6 pairs
of postanal and 20 pairs of preanal papillae : spicules 2.3 to 2.8 mm.

long Contracaecum microcephalum, p. 152.

In connection with species of C ontracaecuni, Ascarh kirghisensis
(placed by the present writer in Porrocaecum) should be kept in
mind as possibly belonging in this genus (see discussion, p. 145).

CONTRACAECUM SPICULIGERUM (Rudolphi, 1809) Raillet and Henry. 1912

Synonym. — Ascaris spiculigerum, Rudolphi, 1809.
Hosts. — AIca torda, Anas clangvla. Carbo bras-iliens'is, C
cormorartus, C. cristatus, C dilophus, (J. graculus, C. pygvweus,

148 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Chroocephalus ridibundus, Colymbus articus, G. atrigularis, G.
nigricans, G. ru.fogularis, G. septentrionalis, Fulica leucopt.,
Haematopus ostralegus, Halieus brasiliensis, Larus argentatus, L,
canus, L. fuscus, L. marinus, L. ridibundus, L. tridactylus, Lestris
parasitica, L. pomarinus, Merganser castor, Mergus merganser,
Microcarbo pygmaeus, Pelecanus americanus, P. conspiciUatus, P.
.erythrorhynchos, P. fuscus, P. onocrotalus, P. pygmaeus, P. species,
P. trachyrhynchus, Phalacrocorax carbo, P. fuscicollis, P. graculus,
P. jawanicus, P. pelagicus, P. sulcirostris, P. urile, P. verrucosus,
Plotus anhinga, P. lavaiUanti, P. melanog aster, P. novae- hollan-
diae, Podiceps auritus, P. cristatus, P. dominicensis, P. minor, P.
vigricollis, T achy petes aguila, Uria grylle, U. troile, Utamania torda.
Location. — Intestine and, in Pelecanus, the gular pouch.

2/d.

Figs. 21G-217. — Contbacaecum spiculigekum. 216, Head. Afteb Linstow,

1000. 217, (I, ESOPHAGCS AND APPENDICES '■ b, MALE TAIL. AFTEK

Schneider, 1S6G

Morphology. — Gontracaecum (p. 146) : Large lips rounded, the
outer surface flat, the inner giving off' 2 rounded processes which
extend outward and forward, protruding beyond edge of lip.
Linstow describes and figures (fig. 216) dentigerous ridges. Inter-
labia hook-shaped and bent inward, just slightly shorter than the
lips and separated from them by a considerable space. Esophagus
1/4.3 or according to other authors 1/8 to 1/9 of total length.
Ventriculus indistinct; posteriorly directed appendix 1.2 to 1.86 mm.
long and up to 720> wide; anteriorly directed cecum very
voluminous, elongate cone-shaped (fig. 217<7).

Mate 32 to 45 mm. long by 800 to 900,* wide. Tail curled, end-
ing in conical point. Cioacal aperture 250/j, from posterior end
in small specimens. Spicules, according to Dujardin, 2 mm. long
by 330/x wide: according to Linstow, 7.2 mm. long; the latter is
more probably correct as the name undoubtedly refers to their being
a striking character. Seven pairs of postanal papillae (fig. 2176),
4 of them ventral. 3 lateral; preanal papillae vary from 38 to 56
pairs.

NEMATODE PARASITES OF BIRDS

149

Female 24 to 46 mm. long by 1 to 1.8 mm. wide. Tail conical, anus
about 400/x from caudal end in a specimen of median size. Vulva
in anterior third of body, dividing body length in ratio of 5 :13.
Ovejector ( % ) filiform and sinuous for a length of 8 to 10 mm., then
enlarging and dividing into the 2 uteri. Eggs spherical, 50 to 52/*
in diameter, according to Dujardin, 72/i in outer diameter and
42/a as diameter of the yolk, according to Linstow. The latter author
describes the shell as 4.9/x thick and covered with uniformly distri-
buted, shining elevations.

Life history. — Unknown.

Distribution. — Europe (Baltic Sea), South America (Brazil), Af-
rica (Nubia and Egypt), Asia (India), Australia and North Amer-
ica (Mexico and United States (Yellowstone Lake, Wyoming)).

FlG. 218. OoNTHACABrUM ANDKKSOXI. a, DOESAL LIP/ b. [nterlabium; c, vulva;

d, MALE TAIL. AFTBR VEVEKS, 1923

CONTRACAECUM ANDERSONI Vevers. 1923

Host. — Florida caerulea.

Location. — Small intestine.

Morphology. — Contracaecum (p. 146) : Cuticle transversely stri-
ated. Head 400//. wide by lOO/i long. Three fleshy lips and 3
large interlabia; the former (fig. 218#) each bear 2 papillae; the in-
terlabia (fig. 2186) are triangular, 150/* long. Esophagus 3.75 mm.
long; posteriorly directed appendix 750/* long by 150/* wide. An-
teriorly directed cecum 2.75 mm. long.

Male 15 to 17 mm. long by 350/t wide. Cloacal aperture 85/t
from posterior end. Five pairs of postanal papillae (fig. 218r/), the
most anterior pair being double; 5 pairs of adanal, and 20 to 25
pairs of preanal papillae. Spicules equal, 1 mm. long by 10/i wide.

Femiale 19 to 22 mm. long by 750/* wide. Anus 130/* from
caudal end which is conical and sharply tapering. Vulva (fig. 218c)
just anterior to middle of body, 9.5 mm. from head end in a 20
mm. specimen. Vagina short (150/«), muscular, with 3 pairs of

150 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

leaf-like valves. Eggs oval, 55/*, long by 45/x wide, with thick shells
of mosaic appearance.

Life history. — Unknown.

Distribution. — South America (British Guiana (Georgetown)).

CONTRA CAECUM ENGONIUM Baylis and Daubney, 1922

Host. — Ciconia. nigra.

Location. — Not given.

Morphology. — C ontracaecum (p. 146) : Head constricted from
body. Dorsal lip (fig. 219a) rounded anteriorly and with a pair
of double papillae situated near the anterior edge. Pulp follows
shape of lip but indented anteriorly; a pair of flattened processes
project anteriorly and laterally like 2 horns.

Male 13 mm. long by 570/a wide. Muscular esophagus 2.75 mm.
long, ventriculus 140/x long; posteriorly directed appendix 700/t long

0-//77/77

^ o/mm

Fig. 219. — Coxtracaecum engoxium. a, Head; b, male tail. After Basiis
and Daubney., 1922

by 150/m wide. Intestinal cecum broad, reaching to within 690//. of
head end. Cloacal aperture 125/a from caudal extremity. Ten
pairs of postanal papillae (fig. 219&), 4 of them being latero-ventral
and the other 6 lateral and pedunculated. No preanal papillae de-
scribed. Spicules equal, 1.8 mm. long by 22/x, wide.

Female unknown.

Life h is'tory. — Unknown .

Distribution. — Asia ( Indi a ) .

CONTRACAECUM HALIAETI Baylis and Daubney, 1923

Synonym. — Ascaris aquillae Smith, Fox, and White, 1908. not
Ascaris aquilae Gmelin, 1790.

Hosts. — Haliaetus Uitcocephnlus and H. leucogaster.

Location. — Intestine.

Morphology. — C ontracaecum (p. 146) : Cuticle transversely
striated. No lateral alae. Three thick lips (figs. 220 and 221),

NEMATODE PARASITES OF BIRDS

151

each with 2 papillae. Interlabia well-developed, the margins in-
folded. No dentigerous ridges. Dorsal lip 110> high by 150>
wide at base, 140/x wide at anterior edge, four-sided, anterior edge
deeply grooved. Cuticle projecting beyond pulp on each side of
the saddle (like a "finger-nail") with ear-like projections.

IQQh 200 300 y

Fig. 220. — Contracaecum haliaeti. a, Lateral view; b, front view of head:
c, egg. After Smith, Fox, and White, 1908

Esophagus long with small, inconspicuous, posterior bulb from
which there extends posteriorly an appendix. Intestinal cecum
large.

Male unknown.

Female 50 mm. long by 1.5 mm. wide. Head blunt, 250y wide at
base of lips. Tail acutely conical, anus subterminal. Vulva an-

"S-wje to J* 4* to

to to i^ to lo.

FIG. 221. — CONTRACAECCM HALIAETI. a, INNER SURFACE; ft, OUTER SURFACE OF

UP. After Smith, Fox, and White, 1008

terior to middle of body, small and inconspicuous. Vagina long,
slender. Eggs (fig. 220c) of varying shape, from 80 to 90ju. long
by 50 to 60/* wide; shell thin, covered with small tubercle-like pro-
jections.

Life history. — Unknown.

Distribution. — North America (United States (Philadelphia
Zoological Garden)) and Asia (India).

3(312— 27 12

152 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

CONTRACAECUM MICROCEPHALIA (Rudolphi, 1809) Baylis, 1920b

Synonyms. — Asca?is microcephala Rudolphi, 1809 ; Kathleena arc-
uata Gedoelst, 1916; C ontracaecum quadricuspe Walton, 1923.

Baylis and Daubney have examined the type specimens of Kath-
leen® arcuata and find that species identical with Rudolphi's. The
present writer considers C. quadricuspe identical also, the descrip-
tion given by Walton agreeing closely with that of Gedoelst in all
particulars except that the esophageal appendix of G. quadricuspe
is somewhat shorter and the vulva slightly more anterior.

Hosts. — Anas boschas, Anas domestica, Ardea cinerea, A. cornata,
A. herodias, A. minor, A. nycticorax, A. purpurea, A. species, A.
stettaris, Ardeolo grayi, A. rattoides, Botauras mugitans, B. stel-
laris, Butorides virescens virescens, Ciconia alba, C . nigra, Herodias

Fig. 222. — Contracaecdm microcethalom. a, Front view of head ; 6, dorsal
view of head ; c, male tail ; d, free end of spicule ; e, cross section of spicule.
After Gedoelst, 1916

egretta, H. tricolor, Nyctiardea grlsea, Nycticorax europaeus. N.
nyctieorase.

Location. — Proventriculus and intestine.

Morphology. — C ontracaecum (p. 146) : Color yellow-white. Cuticle
cross-striated; directly behind the head the furrows thus produced
are so deep that a shirred appearance results. Mouth with 3 lips and
3 interlabia, all with thick transparent cuticle. Lips equal, on their
internal surface a longitudinal furrow; ear-shaped projections
anteriorly. Dorsal lip (fig. 222&) with 2 large double papillae, the
ventro-lateral lips each with 1 large double papilla. Interlabia of
about the same height as the lips, with a free curved internal part
which is incised at its summit. No lateral alae. Cervical papillae
400 to 720/* from head end. According to Gedoelst's description
(K. arcuata) , esophagus 2.8 to 3.2 mm. long by 160 to 225/a wide, with
an appendix 1.1 to 1.25 mm. long by 145/x wide; cecum extends
anteriorly along esophagus for 3/4 the latter's length. According to

NKMAT0DE PARASITES OF BIRDS 153

Walton (('. quadrieuape) , muscular esophagus 3;2 to 3.6 mm. long,
ventriculus 175/x long, esophageal appendix 800 to 816,u long by 80//,
wide, intestinal cecum 2.8 to 2.9 mm. long.

Male 18 to 2G mm. long by 900/* to 1 mm. wide (A', arcuata) ; 15 to
45 mm. long (C. microcephalum). Tail (fig. 222c) curved ventrally;
no caudal alae. Cloacal aperture 280 to 240/* from posterior end.
About 26 pairs of caudal papillae arranged as follows : near the tail
end 4 pairs, 2 being ventral and 2 lateral; immediately posterior to the
cloacal aperture 2 pairs of large papillae; anterior to the cloacal
aperture at least 20 pairs in more or less regular rows (K. arcuata),
or 31 pairs (C. mieroeephalum). Spicules equal. 2.8 to 2.8 mm. long,
with curved alae.

Female 28 to 37 mm. long by 1.1 mm. or less wide (K. arcuata) ;
45 to 70 mm. long (C. mieroeephalum). Tail conical; anus about
440 to 500/x, from posterior end; 2 lateral papillae, 240/t from tail
end. Vulva anterior to middle of body, dividing body length in
ratio of 2 : 3 (in Walton's specimen, which was 87.9 nun. long, vulva
9.6 mm. from the head). Eggs of variable shape, globular or ellip-
tical, averaging 68/* long by 58.5/x wide (the range being 72/a by 48 to
64/i down to 64//, by 56 to 64/t) ; shell thin, with irregular surface.

L ife h istory. — Unkn own.

Distribution. — North America (United States), Europe (Italy
(Rimini, Albona, and Padua)), Asia (India and Russian Turkestan
(Lac Kul-Kainar) ), and Africa (Belgian Congo and Transvaal).

CONTRACAECUM MICROPAPILLATUM (Stossich, 1890) Baylis, 1920

Synonym. — Ascaris micropapUlata Stossich, 1890.

Host. — Pelecamis, species, P. crispus, P. erytkrorhynchos, and
Glaucionetta el an aula.

Location. — Intestine.

Morphology. — Contracaecum (p. 146). The present writer has
recently identified this parasite from P. erythrorhynchos and G.
clangvla^ but until this time the only reports appear to be those of
Stossich in 1890 and 1896. Stossich's description is as follows:
Cuticle with dense cross-striations, especially marked directly behind
the head where there is a collar-like fold. Lips (fig. 2246) large,
the anterior margin projecting laterally as 2 ear-like processes; pulp
with median cleft and with 2 anteriorly directed bipartite prolonga-
tions. Two papillae on lips. Interlabia large, strongly curved, the
anterior ends split into 2 rounded processes.

Male 20 to 2(5 mm. long. Three pairs of double postanal papillae
(fig. 224ff). Preanal papillae numerous, in •_' regular rows.

Female 35 to 40 mm. lonir.

154

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

The present writer is able to add the following description, based
on the American material: esophageal appendix short, its length
equal to 1/6 that of the esophagus (fig. 223) ; intestinal cecum large,
extending anteriorly for 2/3 to 3/4 the length of the esophagus. Male
15 to 20 mm. long by 950/* wide. In a specimen 18 mm. long, esopha-
gus 2.7 mm., esophageal appendix 440//,, intestinal cecum 2.1 mm.
long. Spicules 5.1 mm. long. Tail short, the cloacal aperture being
about 175/* from posterior end of body. Female 20 to 30 mm. long
by 1.2 mm. wide. In a specimen 23 mm. long, esophagus 3.1 mm.,
esophageal appendix 500/*, intestinal cecum 2 mm. long. Vulva 1/4
of body length from anterior end. Eggs 58/* by 54/*.

Fig. 223-224. — Contracabcum micropapii.latum. 223, Anterior end (dis-
sected). Original. 224. a, Male tail; b, lip and interlabia. After
Stossich, 1890

]Jfe history. — Unknown.

Distribution. — Europe (Jugoslavia (Semlin) and Herzegovina
(Narenta River)) and United States (Texas and New York).

The writer is indebted to Dr. E. W. Price, formerly of College Sta-
tion, Texas, for the specimens described above from P. enjthrorhyn-
chos.

CONTRACAECUM MULTIPAPILLATUM (Drasche, 1882) Baylia, 1920b

Synonyms. — Ascaris viultipapiUata Drasche, 1882 (1883«) Con-
tracaecvm ?rmltipapi?losa (Drasche, 1882) Skrjabin, 1916 (evidently
a misspelling).

Hosts. — Ardea, species and Tantalus loeulator.

Location. — Esophagus and proventriculus.

NEMATODE PARASITES OF BIRDS

155

Morphology. — Contraeaecum (p. 146) : Lips (fig. 220a) with ear-
like projections and with 2 papillae; dentigerous ridges absent. Ac-
cording to Drasche, intcrlabia about 2/3 the height of the lips, thick
and curved, their free ends being close to lips; Skrjabin states that
no interlabia are present. Cross-striations of cuticle give dentate
appearance to collar as seen at edge of body.

Male 13 to 18 mm. long by COO to 900/* wide. Esophagus 3.23 mm.
long, its appendix small, dactyliform (fig. 225). Intestinal cecum
7/8 the length of esophagus. Tail (fig. 2206) 150/* long in 18 mm.
specimen, with 10 pairs of large conical postanal papillae, with
sometimes an additional asymmetrical papilla on one side. Preanal
papillae in two rows of 31 to 50 papillae each, the number being
not always the same in both rows. Spicules 1.27 mm. long, alate.

Female 18 to 20.4 mm. long by 500/* to 1 mm. wide. Posteriorly
directed esophageal appendix 1/3 the length of the anteriorlv directed

226

Figs. 225-226. — Oontracakcum multipapillatcm. 225, Digestive organs ov male.
After Skrjabin, 191(5. 22G, «, Head; b, male tail. After Drasche, 1882

intestinal cecum, the latter being 3/4 the length of the esophagus.
Tail pointed, anus 370/* from end. Vulva in posterior part of body,
dividing body length in ratio of 2:1. Eggs oval, 68/* long by 60/*
wide.

Life history. — Unknown.

Distribution. — South America (Brazil and Paraguay).

CONTRACAECUM OVALE (Linstow, 1907) Baylie, 1920b

Synonym. — Ascaris oralis Linstow, 1907.

Host. — Podiceps cristatus.

Location. — Proventriculus and intestine.

Morpholoyy. — Contraeaecum (p. 146) : Cuticle with cross-striations.
Lips oval (fig. 227), wider than long, papillae projecting laterally.
Dentigerous ridges absent. Pulp in 2 anteriorly directed processes.

156

BULLETIN HO, UNITED STATES NATIONAL MUSEUM

Interlabia low, anteriorly rounded. Esophagus 1/6 of total length,
with appendix equal to 3/5 the length of the former; cecum extend-
ing anteriorly from the intestine for 5/6 the length of the esophagus.

Male 29 mm. long by 880/* wide. Tail 1/124 of total length. Ten
pairs of pre- and post-anal papillae. Spicules 2.8 mm. long.

Female 44 mm. long by 1.38 mm. wide. Tail 1/78 of total length.
Vulva anterior to middle of body. Eggs 68/* long by 57/* wide.

Life history. — Unknown.

Distribution. — Europe (Germany (Berlin Museum)).

CONTRACAECUM PRAESTRIATUM Monnig, 1923

Host.- — Podiceps capensis.
Location. — Not given.

Morphology. — Contracaecwm (p. 146) : Color
striations, 38/* wide, very marked in anterior 1/3

white. Cuticular
of bodv, then slid-

22$

229.

Figs. 227-229, — 227, Contracaeccm ovale. Head. After Lin stow, 1907. 223,
contracaecum ruaestriatum". anterior end. 229, male tail. after mottnig,
1923

denly becoming finer. Lips (fig. 228) thick and broad, of equal
size, the dorsal with 2 papillae, the others with 1, all the papillae
being double. Interlabia short, triangular with very broad base.
No dentigerous ridges. Esophagus 4.1 mm. long; posteriorly
directed appendix 2 mm. long; anteriorly directed intestinal cecum
2.3 mm. long, in a female 30 mm. long.

Male 20 mm. long by 650/* wide. Tail (fig. 229) 160/* long. Five
pairs of postanal papillae, 4 of which are grouped together just
below the middle of the tail and the fifth pair, of double papillae,
just posterior to cloacal aperture. At least 18 pairs of preanal
papillae extending about 3.6 mm. anterior to cloacal aperture.
Spicules equal, 1.9 mm. long, alate; a small gubernaculum present,
120/* long.

NEMATODE PARASITES OF BIRDS 157

Female 33 mm. long by 800> wide. Tail 360//. long, sharply
pointed. Vulva in anterior third of body, dividing body length in
ratio of 5 : 12. Eggs thin-shelled, very round, 40> long by 32/* wide.

Life h istory. — Unknown.

Distribution. — Africa (Transvaal) .

COXTRACAECUM PUNCTATUM (Gedoelst, 1916) Baylis, 1920b

Synonym. — Kathlcena punctata Gedoelst, 1916.

Host. — "Grand Heron a bee de Pelican," probably Pseudo-
t an talus ibis.

Location. — Pro ventri cuius.

Morphology. — Contracaecum (p. 146) : Color white. Cuticle with
cross-striations. Head small, 148/* in dor so -ventral diameter. Three
lips, the dorsal with 2 papillae, the ventrolaterals each with a single
papilla. Three interlabia present. Precervical papillae 470/i from
head end in small specimens. Esophagus 2.24 to 4.5 mm. long, with

230 — 23/.

Figs. 230-231. — 230, Contracaecum punctatum. Male tail. After Gedoelst,
1916. 231, Contracaecum rodhaini. Head. After Gedoelst., 1916

an appendix 960/* to 1.2 mm. long. Cecum extends from intestine
anteriorly for 1.6 to 3.6 mm.

Male 16.5 to 32.5 mm. long by 665/t to 1.15 mm. wide. Tail (fig.
230) curved, conical, measuring 1/70 of total body length. At least
23 pairs of caudal papillae arranged as follows: 4 pairs near tail
end. 2 of them ventral. 2 lateral; 2 pairs between the preceding
group and the cloacal aperture; at least 17 pairs of preanal papillae,
arranged in 2 regular rows. Spicules equal, 4 mm. long by 42.5 ( u
wide, with curved alae.

Female. — Unknown.

Life history. — Unknown.

Distribution. — Africa (Belgian Congo (Bukama. Katanga)).

CONTRACAECUM RODHAINI (Gedoelst, 1916) Baylis, 1920b

Synonym. — KathJeena rodhaini Gedoelst, 1916.
Host. — Plotus rufus.
Location. — Proventriculus.

Morphology. — Contracaecum (p. 146): Color yellow - white.
Cuticle with fine cross-striations. Head (fig. 231) 300/t wide by

158

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

150 to IGO/i. high. Three equal lips, the dorsal with 2 simple papillae,
the 2 latero-ventrals with a double papilla. Three large interlabia,
almost equal in height to the lips. No lateral alae. Cervical papillae
510/a from anterior end. Esophagus 3.3 to 4 mm. long, with post-
eriorly directed appendix 720ii to 1.04 mm. long; anteriorly directed
cecum 2.6 to 2.8 mm. long.

Male 18.5 to 20 mm. long by 830/u, to 1.12 mm. wide. Tail curved
ventrally, without caudal alae; cloacal aperture 175/* from posterior
end. At least 33 pairs of caudal papillae arranged as follows: a
group of 4 papillae (the description does not say 4 pairs but that is
evidently intended) near caudal end, a double papilla posterior and
lateral to the cloacal aperture and a series of at least 28 preanal
papillae. Spicules equal, 3.6 mm. long, alate.

232 ^v7

233

c-/mm

Figs. 232-233. — Contraoaecum rosarium. 232, Anterior end. After Con-
nal, 1912. 233, Male tail. After Baylis and Daobney, 1922

Female 22 to 25 mm. long by 940/* to 1.2 mm. wide. Tail conical,
with a papilla on each side, 120/x from posterior end. Anus 336/*
from end. Vulva not salient, at the anterior 3/10 of body length.
Eggs oval or globular, 69/* long by 54/* wide, with thick shells.

Life history. — Unknown.

Distribution. — Africa (Belgian Congo (Yumbi)).

CONTRACAECUM ROSARIUM (Connal, 1912) Baylis, 1920b

Synonym. — Ascaris rosarius Connal, 1912.

Hosts. — Nycticorax griseus and Nycticorax, species.

Location. — Proventriculus.

Morphology. — C ontracaecum (p. 146) : Cuticle cross-striated and
with numerous regularly arranged refractile dots, as of beads of
rosary, along whole length. Three large fleshy lips (fig. 232),
concave internally and laterally; 3 interlabia also quite large,
triangular. Distinct articular collar around base of lips. Esophagus
3.9 mm. long (in the male), its appendix 700/* long. Cecum of

NEMATODE PARASITES OF BIRDS

159

intestine runs forward for a length of 2.46 mm. Body abruptly
widened at level of anterior end of cecum.

Male 26 mm. long by 600/t wide. Cloacal aperture 220//, from
caudal extremity. According to Connal, 3 pairs of postanal papillae,
the first pair double and button-like, the second and third pairs
smaller, nipple-like; according to Baylis and Daubney (fig. 233), 9
pairs of postanal papillae, this being the particular by which this
species can be differentiated from C. microcephalum. Numerous
preanal papillae. Spicules equal, 3.28 mm. long.

1--M

Fig. L'34. — Contkacaecum scotti. «, Esophagus and appendices; b, head; c,

LATERAL VIEW AND d, VENTRAL VIEW OF MALE TAIL. AFTER LHIPER AND ATKIN-
SON, 1915

Female 30.8 mm. long by 700/i, wide. Vulva 1.59 mm. from head
end. Vagina about 170/a long, dividing into 2 parallel uteri. Eggs
oval, 26/a long by 21/x wide, with thick shells.

Life history. — Unknown.

Distribution. — Africa (Lagos) and Asia (India).

CONTRACAECUM SCOTTI (Leiper and Atkinson. 1914) Baylis, 1920

Synonym. — Kathlcena scotti Leiper and Atkinson, 1914.

Host. — Diom edea melanophrys.

Location . — Intestine.

Morphology. — Contracaecum (p. 146) : Three lips (fig. 2345) 135u
long by 130/x wide. Interlabia large, pentagonal, 80/* long by 50/*
wide near base. Esophagus (fig. 234a) 2.53 mm. long by 400/*. wide,
posterior part slightly differentiated into muscular bulb and with a
posteriorly directed appendix 550/x long by 220/x wide. Anteriorly

160 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

directed cecum 1.8 mm. long, extending to within 700> of head and
attaining a width of 400//,. Cuticle transversely striated.

Male 15.3 mm. long by 900//, wide. Cloacal aperture 340//, from
caudal extremity, the latter ending in a digitiform process. Spicules
(fig. 234c) unequal in size but of similar shape, their lengths 3 mm.
and 2.7 mm., respectively, their width 90/x. Four pairs of papillae
near tail end (fig. 234a 7 ), 4 pairs about halfway from end to cloacal
aperture; in addition, starting somewhat posterior to the cloacal
aperture and running far anterior, on each side of body, is a double
row of papillae, 20 or more double pairs.

Female 15 to 25 mm. long. Tail ending in digitiform process.
In a specimen 25 mm. long, anus 400//, from posterior end, vulva 4
mm. from head end. Eggs 100/x long by 60//. wide.

Life history. — Unknown.

Distribution. — Antarctic (in the southern Pacific (Campbell
Islands) ).

CONTRACAECUM TRICUSPE (Gedoelst, 1916) Baylis 1920b

Synonym. — Kathleena tricuspis Gedoelst, 1916.

Hosts. — Ardea, species and Plotus inelano g aster.

Location. — Proventriculus.

Morphology. — Contracaecum (p. 146) : Color yellow-white. Cuticle
with cross-striations, producing directly posterior to the head deep
furrows or a shirred appearance. Three lips (fig. 235 b and c) and
3 interlabia of elaborate structure, the lateral surfaces of the former
notched, with a point of the interlabia fitting into them. Interlabia
with large base and slender body, ending in 3 points, an internal and
2 laterals. Dorsal lip with 2 simple papillae, latero-ventral lips each
with 1 double papilla. No lateral alae. Cervical papillae 700//. from
head end. Esophagus 3.8 mm. to 3.9 mm. long by 100 to 160/t wide,
with an appendix 1.6 to 2 mm. long by 190 to 225/x wide. Cecum
originating from intestine extends to the anterior fourth of the
esophagus.

Male 13.8 mm. long by 785//, wide. Tail conical (fig. 235a), ending
in a pointed appendage, curved ventrally. Cloacal aperture 140//,
from posterior end. At least 56 pairs of caudal papillae arranged
as follows : 4 pairs near caudal end, 2 of them lateral, 2 ventral ; 2
pairs directly posterior to cloacal aperture; 1 pair of large adanal
papillae and a series of about 50 pairs of preanal papillae in 2 regu-
lar rows. Spicules equal, 4.6 mm. long, of similar construction to
those of C. microcepluihon.

Female 12.1 to 17.5 mm. long by 960// to 1.28 mm. wide. Tail
conical, 1/39 of total length, the anus being 350 to 440// and 2 papillae

NEMATODE PARASITES OF BIRDS

161

180>, from its end. Vulva at anterior 2/5 of body. Eggs not ma-
ture in specimen described.

Life history. — Unknown.

Distribution. — Africa (Belgian Congo, Leopoldville) and Asia
(India).

CONTRACAECUM MAGNIPAPILLATUM Chapin, 1925

Host. — Megalopterus hawaiiensis.

Location. — Not given.

Morplioloijtj. — Contt aeaecum (p. 146): Lips subequal, the width
of each at base greater than its height. Interlabia slightly less high,
subtriangular. Dorsal lip with two double papillae, each sub-
ventral lip with one double papilla. Amphids in interlabia adja-
cent to dorsal lip ; excretory pore near base of ventral interlabium.

Male 20 mm. long by 585/x wide near middle of body. Lips 75/x
high ; cervical constriction 75/* behind base of lips. Cervical papillae
645/i posterior to base of lips, oval, slightly prominent. Esophagus

Figs. 235-236. — 235, Cqntbacaecum tbicdspe. a, Male tail; b. dorsal view;

C, FRONT VIEW OF HEAD. AFTER UePOKLST. 1!)1C>. 236, CONTR ACABCC M MAG-
NIPAPILLATUM. Male tail. After Chapin, 1925

3.2 mm. long; esophageal appendix 3/14, intestinal cecum 3/4 as long
as esophagus, respectively. Cloacal aperture 200/a from posterior
extremity of body, which ends in an acutely conical tip 13^ long.
Caudal papillae (fig. 230) arranged as follows: A pair of very
large double papillae, situated 2G/x. posterior to the cloacal aperture,
on two bosses, each boss measuring 43/x by 23^. and separated from
the other by a distance equal to the shorter diameter of either.
Posterior to the double papillae there are 2 pairs of single papillae,
one pair being submedian and the other pair subjateral and slightly
more prominent. Proceeding anteriorly from each double papilla
is a single row of 21 single papillae, the most anterior being about
4 mm. in front of the cloacal aperture. Spicules equal and similar.
3.8 mm. long, the tips acute.

Female 23 mm. long by 750/i wide just posterior to vulva. Ante-
rior portion similar to that of male. Esophagus 3.37 mm. long;
ratios of length of esophagus to that of appendices virtually the

162 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

same as in male. Vulva on a slight prominence. 8 mm. from anterior
extremity, or at about the anterior third of the body. Vagina short
(120/a), straight, opening into a muscular sinuous oviduct about 2
mm. long. Oviduct bifurcates to form two posteriorly directed
uteri. (Anus 300/*. from posterior end of body, which is conical with
a terminal spike 10/i long. Eggs G8/a by 45/i, with thin shells.

Life history. — Unknown.

Distribution. — Hawaii, Lisianski Island.

Suborder Spirurata Railliet and Henry, 1915

Synonym. — Filariata Skrjabin, 1915, in part.

Suborder diagnosis. — Myosyringata (p. 4.) : Body usually long
and slender. Mouth with 2 lips or a larger number of lips or lobes
capable of resolution into 2 basic lips, or without lips and surrounded
by papillae or, occasionally, with other cephalic structures than lips
or papillae. Esophagus slender, without posterior bulb. Male with
1 or 2 spicules. Tail usually provided with papillae, usually curved
sjDirally, caudal alae present or absent or a closed muscular, bell-
shaped " bursal cup " present. Female larger than male, rarely sex-
ually dimorphic (Tetrameridae). Anus subterminal or occasionally
terminal. Vulva present or, less often, absent in gravid females, its
position variable (anterior of middle of body, near middle or some-
times near or at posterior extremity). Two, four, or more uteri,
rarely only one. Oviparous, viviparous, or ovoviviparous. Hete-
loxenous, the larval stages occurring in various intermediate hosts.

KEY TO SUrERFAMILIES OF SPIKIRATA

1. Male with closed muscular bursal cup at tail end; female with anus terminal.

Dioctophymoidea, p. 366.
Male without bursal cup of above description ; anus of female subterminal- 2.

2. Mouth without lips ; vulva near anterior extremity of body ; adults parasitic

subcutaneously, in blood or on serous surfaces (not discussed in this paper).

Filarioidea.

Mouth with lips, or, rarely without lips; position of vulva variable; parasitic

usually in digestive tract, occasionally in orbital region or respiratory

tract, or as larvae subcutaneously Spiruroidea, p- 162.

Superfamily SPIRUROIDEA Railliet and Henry, 1915

Swperfamily diagnosis. — Spirurata (p. 162) : Mouth with 2 lips or
a larger number of lips or lobes capable of resolution into 2 basic
lips ; or, more rarely, without lips. Male with caudal extremity com-
monly expanded and alate. Female with vulva usually in middle
portion of body, occasionally near anterior or posterior extremity.

Intermediate host usually not biting arthropods (exception, Hab-
ronema majus in Stomoxys).

Type-family. — Spiruridae Oerley, 1886.

NEMATODE PARASITES OF BIRDS 163

KEY TO FAMILIES OF SPIKUttOIDEA

1. Mouth with 2 large trilobed lips with a longitudinal ridge on their Inner

surface meeting the one on the opposite sides; adults in gastric tract of

carnivorous mammals ; larvae may occur in birds.,- Gnathostomidae, p. 863.

Month without lips or with lips different from above; adults (except in

Spirocerca) may occur in birds 'J.

2. In orbital region or respiratory tract 3.

In digestive tract 4.

3. In respiratory tract; distinctive larval characters maintained into adult life,

both male and female may have cluster of filiform papillae at caudal ex-
termity ; male without caudal alae or papillae of usual type.

Desmidocercidae, p. 208.
In orbital region. Adult male and female without larval characters as in
above. Male with or without caudal alae and with caudal papillae »f
usual type, the preanal papillae being especially numerous.

Thelaziidae, p. 311.

4. Marked sexual dimorphism, the female robust, more or less fusiform or

globular, the male small and slender, without caudal alae.

Tetrameridae, p. 333.
Sexes not dimorphic; male with caudal alae 5.

5. Anterior region of body with cuticular ornaments 6.

Anterior region lacking cuticular ornaments 7.

6. Head with 4 pinnate, posteriorly directed processes; mouth without lips or

with 2 small papilliform lips ; bird forms found in gizzard.

Ancyracanthidae, p. 301-
Cuticular ornaments of anterior region different from above; 2, 4, or G
triangular lips; in esophagus, proventriculus or gizzard of birds.

Acuariidae, p. 210.

7. Male with circumscribing caudal alae continuous anteriorly and with a group

of 4 or 5 pairs of long pedunculated papillae in cloacal region.

Physalopteridae, p. 295.
Caudal alae of male not continuous anteriorly and without above arrange-
ment of papillae Spiruridae, p. 163.

Family SPIRURIDAE Oerley, 1885

Family dmynosis. — Spiruroidea (p. 162) : Mouth with 2 or 4 ex-
panded lips, or without lips. Male with caudal alae and dissimilar
spicules. Female with vulva of variable position, but usually located
in the median portion of body.

Parasitic as adults in mammals and birds, oeuerally in the diges-
tive tract.

Type-geniis. — Sphwa E. Blanchard, 1849.

KEY TO SUBFAMILIES OF SI'IROUIDAE

1. Inadequately described species Subfamily uncertain, p. 205

Description adequate for subfamily determination 2.

2. Mouth without lips or disk; cervical region ornamented with cuticular

bosses; vulva near posterior extremity Gongyloneminae, p. 203.

Mouth with lips <>r with circular projecting disk; cervical region without
cuticular bosses; vulva usually remote from posterior extremity.

Spirurinae, p. 164.

164 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Subfamily Spirurinae Railliet, 1915

Suh family diagnosis.— Spiruridae (p. 163) : Female with 2 uteri
and with vulva usually in the middle portion of the body, rarely close
to anterior or posterior extremities. Pharynx without cuticular
rings or spirals.

Type-genus. — Sjrirura E. Blanchard. 1849.

KEY TO GENERA OF SPIRURINAE

1. Larvae encysted, probably accidentally, in birds ; adults in otber bosts.

Spirocerca, p. 202.
Adults in birds 2.

2. Cepbalic extremity covered by a circular projecting disk Viguiera, p- 201.

Cepbalic extremity possessing no such disk but with the usual lip structures
3.

3. Mouth with 4 lips 4.

Mouth with 2 lips 5.

4. No lateral alae on body ; sensory papillae postcervical ; postanal papillae

symmetrical Cyrnea, p. 166.

Lateral alae usually present ; sensory papillae precervical ; postanal papillae
asymmetrical Habronema, p. 172.

5. Mouth surrounded by Glutinous ring; lips small, integral Spirura, p. 164.

No chitinous ring around mouth ; lips comparatively large, trilobed. at

least internally 6.

6. Sensory papillae cervical (i. e., at level of nerve ring) ; vulva in anterior part

of body (anterior to end of esophagus) Hadjelia, p. 187.

Sensory papillae precervical ; vulva in median region of body.

Hartertia, p. 191.

Genus SPIRURA E. Blanchard, 1849

Genetic diagnosis. — Spirurinae (p. 164) : Two lips, sometimes bear-
ing papillae, sometimes teeth. Mouth straight and surrounded by
a cutinous (chitinous) ring. A buccal cavity and often a pharynx
present. Esophagus long and cylindrical. Body of moderate size,
attenuated toward the anterior extremity. Lateral alae sometimes
present. Male with unequal spicules and with caudal alae which,
however, are not continuous across the ventral surface; 4 pairs of
preanal papillae. Female with vulva anterior or posterior of middle
of body ; two uteri.

Parasitic in stomach and intestine of mammals and birds.

Type-species. — Sphmra talpae (Gmelin, 1790) E. Blanchard. 1849.

KEY TO SPECIES OF SPIRURA

Lateral alae extending whole length of body; a total of 4 teeth around the
mouth on inner surface of lips; male 30 mm. long__ Spirura zschokkei, p. 165.

No lateral alae described ; a total of 6 teeth around the mouth on inner surface
on lips ; male 15 to 20 mm. long Spirura uncinipenis, p. 165.

This key and the descriptions which follow do not include Spimra
talpae (Filaria stmmosa) which has been reported from Buteo vul-

NEMATODE PARASITES OF BIRDS 165

garis by Stossich. If the report is correct, it is probable that the
bird had eaten a mole or rat, the normal hosts of S. tctipae, and that
the parasite was accidental in the bird.

SPIRURA ZSCHOKKEI Railliet and Henry, 1911a

Synony?n. — Spiroptera alata Zschokke, 1889, not Rudolphi, 1819,
renamed.

Host. — Primary: Rhea americcma; secondary: Unknown: prob-
ably an insect.

Location. — Pro vent riculus.

Morphology. — Spirura- (p. 164) : Body cylindrical, cuticle trans-
versely striated. Two lateral alae extend the total length of body;
at the level of the posterior extremity of the esophagus they each
bear a projecting papilla. Head, somewhat distinct from the body,
with 2 strong lateral lips, hemispherical, slightly hollowed out like
a spoon, at the base of which there opens a narrow buccal slit. Each
lip supported by a cutinous (chitinous) armature consisting of a
thick triangular disk which is prolonged posteriorly by an attenu-
ated band ; on the internal edges there extends also a strongly chiti-
nized band. These internal edges, slightly arc-shaped, form anterior
to the mouth a sort of vestibule, slightly raised. The 4 chitinous
bands unite at the base to form a ring which surrounds the mouth and
carries 4 sharp teeth. In addition only the anterior half of the lips
is free; in the posterior half they are joined into a sort of simple
buccal cavity, the anterior edge of which also bears a chitinous
ring. Esophagus slightly enlarged, bulb-like, at posterior extremity.

Male 30 mm. long by 1 mm. wide. Posterior third of body rolled
in spiral. Two pairs of postanal papillae. Preanal papillae un-
described but presumably (see generic diagnosis) number 4 pairs.

Female unknown.

Life history. — Unknown; probably involving an insect as inter-
mediate host, as larvae of Spirura gastropkila have been found en-
c} r sted in cockroaches.

Distribution. — South America (Brazil) and Europe (Switzer-
land (Zoological Garden, Basel)).

SPIRURA UNCINIPENIS (Molin, 1860) Railliet and Henry. 1911a

Synonyms. — Spiroptera vnrinipenis Molin, 1800: Cheilospirura
uncinipenis (Molin, 18G0) Diesing, 1801.

Host. — Primary: Rhea americana; secondary: Unknown, probably
an insect.

Location. — Proventriculus.

Morphology. — Spirura (p.164) : Body cylindrical, attenuated at the
extremities. Cuticle transversely striated; in the anterior region of

166

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

the body the striations are very prominent and produce the appear-
ance of the body being crenated. Head (figs. 237# and b and 238a)
elaborate; 2 lateral lips, large and semilunar at the wide anterior
edge, and 2 small interlabial median lobes. Each of the lateral lips
has 3 teeth on its inner surface.

Male 15 to 20 mm. long by 500 to 550/x wide. Cloacal aperture
1/22 of total length from posterior end. Caudal region (fig. 237c),
recurved toward the ventral face, sometimes even twisted in a spiral ;
caudal alae wide, longitudinally striated. Six pairs of pedunculated
papillae, of which 4 are preanal, 2 postanal. Spicules unequal, the
one 2.6 to 3.3 mm. long, slender and pointed, the other only 310/x.
(Linstow) or 180/x long (Railliet and Henry), thick and ending
in a hook with rounded point (fig. 238c).

Figs. 237-238. — Spikura uncinipenis. 237, a, Lateral view ; b, front view of
head; e, male tail. After Drasche, 1884. 238, a, Head; 6, egg; c, free end

OF SHORT SPICULE. AFTER L.INSTOW, 1899

Female 18 to 26 mm. long by 700 to 800/x wide. Tail 1/100
total length. Vulva almost at middle of body, salient. Eggs (fig.
2386) 43 to 47/x long by 26/z wide, embryonated; shell thick, rein-
forced at each pole.

Life history. — Unknown; probably involving an insect as inter-
mediate host, as larvae of Sjnrura gastrophila have been found
encysted in cockroaches.

Distribution. — South America (Brazil) and Europe (France
(Melun)).

Genus CYRNEA Seurat, 1914c

Generic diagnosis. — Spirurinae (p. 164) : Mouth with 4 lips. No
lateral alae. Sensory papillae far posterior to nerve ring. Male
with 2 unequal spicules; a gorgeret present, Female with vulva
usually situated considerably posterior to middle of body. Ovejec-
tor with vestibule differentiated into an organ for storage of eggs.
Uteri parallel.

NEMATODE PARASITES OF BIRDS 167

Parasitic usually under corneous tunic of gizzard of birds.
Type-species. — Cymea euryecrca Seurat, 1914c.

KEY TO SPECIES OF CYRNEA

1. Vulva immediately In front of middle of body Cyrnea excisa, p. 167.

Vulva a considerable distance posterior to middle of body 2.

2. Vulva prominent Cyrnea semilunaris, p. 171.

Vulva not prominent 3.

3. Left spicule 2.0 mm., right spicule 365/t long; male with 10 pairs of caudal

papillae; tail of female 332/u. long Cyrnea colini, p. 168.

Left spicule not over 1.7 mm., right spicule 3S0m or longer ; male with 9
pairs of caudal papillae : tail of female not over 285/i long 4.

4. No papillae directly in front of cloacal aperture : caudal alae 410 to 550/* long,

their span 275 to 390m Cyrnea seurati, p. 172.

A pair of papillae directly in front of cloacal aperture; caudal alae about
290/n long, their span 250/z 5.

5. Left spicule 1.7 mm., right spicule 380m long; tail of female 285m long; vulva

720/i anterior to anus ; eggs 42m long by 18m wide_ Cyrnea eurycerca, p. 167.
Left spicule 1.3 mm., right spicule 490m long ; tail of female 140/t long ; vulva
1.14 mm. anterior to anus; eggs 52m long by 24/* wide.

Cyrnea parroti, p. 171.

Physaloptera bulbosa (p. 310), recently transferred to Cyrnea (see
Addenda, p. 390), may be differentiated from the above species in
that the vulva is in the posterior part of body and the spicules
measure 2.45 mm. and 945m long, respectively.

CYRNEA EURYCERCA Seurat. 1914

Hosts. — Primary: Caccabis petrosa, Coturnix coturnix, and "per-
dix rouge "; secondary: Unknown.

Location. — Under tunic of gizzard.

Morphology. — Cyrnea (p. 166) : Male 6.1 to 10.5 mm. long by 250m
wide (fig. 239c). Buccal cavity 55m l° n g> muscular esophagus 285m
long ; total length of esophagus 2.5 mm. Excretory pore and sensory
papillae 280m from anterior end. Caudal alae (fig. 239&) outspread,
short, the semilunar wings transversely striated, 290m l° n ^ a little
greater than the span from margin to margin, 250m- Nine pairs of
long pedunculated papillae, of which 3 are preanal; in addition a
pair of very small papillae directly anterior to the cloacal aperture.
Left spicule 1.68 mm., right spicule 380m long. Gorgeret 70m long.

Fern-ale 14.7 to 20.5 mm. long. Tail (fig. 239a) attenuated, rounded
at end, 285m l° n g- Vulva not salient, 720m anterior to anus. Vesti-
bule 1.05 mm. long. Eggs 42m l° n ^ by 1% wide, with thick shell.

Life history. — Unknown.

Distribution. — Europe (Corsica) and Africa (Algeria).

CYRNEA EXCISA (Molin, I860) Seural. 19Hc

Synonym*. — Spiroptera excisa Molin, 1860; Physaloptera striata
Linstow, 1883.

168

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Primary: Giconia alba, G. a. asiatica, G. cicoivia and G.
maguari. Linstow has also reported this species from the snake,
Tropidonotus hydras. Secondary : Unknown.

Location. — Esophagus, proventriculus, and gizzard.

Morphology. — Gyrnea (p. 166) : Blood-red in color. Sensory pap-
illae far posterior to nerve ring. The lateral lips (fig. 240a and b)
divided on their inner surface into 3 lobes ending in a tooth. Dorsal
and ventral lips have 2 large papillae at their base.

Male 16.5 mm. long by 600> wide. Tail (fig. 240d) straight. Six
pairs of caudal papillae, of which 4 are preanal ; in addition a large

Fig. 239. — Cyhnea euryckrca. a, Female tail, showing ovejector ; b, male tail ;

C, MALE (LENGTH 7.6 MM) ; (l. LATERAL VIEW; I', VENTRAL VIEW OF HEAD. AFTER
SHURAT, 1914

sessile unpaired papilla on anterior edge of cloacal aperture. Spic-
ules unequal, the left 2.4 mm., the right 620/x long. Gorgeret present.

Female 19 to 26 mm. long by 670/j, wide (fig. 240c). Tail short
(300^). Vulva not salient, little visible, immediately in front of
middle of body. Vestibule 1.6 mm. long, sphincter 800,u long. Eggs
42/x long by 2Lu, wide.

Life history. — Unknown.

Distribution. — South America (Brazil), Africa (Algeria), and
Asia (Turkestan).

CYRNEA COLINI, new species

Hosts. — Primary : Golinus virginianus ; secondary : Unknown.
Location. — In wall of proventriculus, at its junction with gizzard.
Morphology. — Gyrnea (p. 166) : Head with 4 lips. Dorsal and
ventral lips prominent, deeply divided into 2 parts in such a manner

NEMATODE PARASITES OF BIRDS

169

that in lateral view (fig. 242c) the head appears to bear 4 conspicu-
ous projecting papillae. Each of the four parts of these lips bears
on its outer edge a prominent thumblike extension. Lateral lips

300/n

JOO/U

Fig. 240. — Cyknea excisa. a, Dorsal view; h, lateral view of head end; c,

FEMALE ; d, MALE TAIL. AFTER SEURAT, 1914

(fig. 242£> and c) very large, each bearing 2 digitiform processes on
its inner surface and 2 lateral winglike expansions which project
into the median groove of the dorsal and ventral lips in such a

Fig. 241.— Cyknea skcrati.

Tail end of male and of female.
Neyra, 1918

After Lopez-

manner as to give the appearance in some views of being processes
from the latter lips. There are 2 obscure lateral papillae near the
base of the lateral lips, which Dr. G. Steiner, on examination, stated

170

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

were undoubtedly amphids. The cervical papillae could not be
located.

Male about 6 mm. long by 250> wide. Buccal cavity 58/* deep;
total length of esophagus 2 mm. Caudal alae (fig. 243a) outspread,
280/u long, their span 230/*; they bear coarse transverse striations.

Fig. 242. — Cyrnea colini. Head, a, Ventral ; b, fkont ; c, oblique

LATERAL VIEW. ORIGINAL

Cloacal aperture about 115/* from posterior end. Ten pairs of pedun-
culated papillae forming an unbroken series, the 4 most posterior
pairs slightly smaller than the others; the seventh pair from the
posterior end lies at the level of the cloacal aperture. Spicules very
unequal, the left 2 mm. long by 12/t wide, the right 365/* long by 24/t

Fig. 243.' — Cyrnea colini. a, Male tail; b, female tail; c, ovejector. (Sgalb

FOR 6 AND C THE SAME.) ORIGINAL

wide. Gorgeret delicate; the left side (68/* in length) is longer than
the right side.

Female 14 to 18 mm. long by 365/* wide. Buccal cavity 75/* deep ;
first part of esophagus about 280/* long; second part of esophagus
about 2.5 mm. long. Tail (fig. 2436) 33*2/* long. Vulva (fig. 2436)

NEMATODE PARASITES OF BIRDS 171

only slightly salient, 915/* anterior to aims. Vestibule, including
sphincter, 1.2 mm. long (fig. 243c). Eggs 40.5m by 22.5m.

Life history. — Unknown.

Distribution: — North America (United States (Georgia)).

Type material. — No. 26389 U.S.N.M. (Bureau of Animal Indus-
try Helminthological Collection).

The proventriculi of two quail, containing numerous specimens
of these nematodes, were sent to the Zoological Division of the
Bureau of Animal Industry by the Biological Survey from the head-
quarters of the Cooperative Quail Investigation in Georgia.

CYRNEA PARROTI Seurat, 1917

Host. — Primary: Caccabis petrosa; secondary: Unknown.

Location. — Under corneous tunic of gizzard.

Morphology. — Cyrnea (p. 166) : Body robust; cuticle thick, finely
striated transversely. No lateral alae. Cervical papillae asymmetri-
cal, at level of excretory pore. Intestinal papillae asymmetrical, in
a female 18.8 mm. long, the left papilla 3.5 mm., the right 7.75 mm.
from caudal extremity. Mouth with 2 lateral lips, rounded at free
edge, having 3 small teeth on internal surface, and a dorsal and ven-
tral lip deeply notched in the middle.

Male 11.7 mm. long by 265/* wide. Tail 160/* long. Spicules
unequal, the left 1.34 mm., the right 490/* long. Gorgeret 65u
long. Caudal alae and papillae as in Cyrnea eurycera (p. 167).

Female 18.8 mm. long by 360/* wide. Tail 140/* long. Vulva
1.14 mm. from caudal extremity, not salient. Ovejector like that of
some species of Habronerna of birds; vestibule very short (300/*) ;
unpaired trompe 660/*. Eggs 52/* long by 24/* wide.

IAfe history. — Unknown.

Distribution. — Africa (Algeria (MacMahon) ).

CYRNEA SEMILUNARIS (Molin, I860) Seurat, 1914f

Synonyms. — Sp-iroptera semilunaris Molin, 1860; Spiroptera lan-
ceolata Molin, 1860.

Hosts. — Primary : Crotophaga major, Trogon eollcwis, and T. mel-
anurus', secondary: Unknown.

Location. — Between the tunics of the gizzard.

Morphology. — Cyrnea (p. 166) : M ale 6 to 10 mm. long by 150,u
wide. Caudal alae not long but wide, semilunar; 9 fungiform pa-
pillae, of which 6 are very long. Spicules unequal.

Female 4 to 20 mm. long by 100 to 400/* wide. Caudal ex-
tremity straight, obtuse; anus remote from caudal apex. Vulva
prominent, in posterior part of body not far from anus.

Life history. — Unknown.

Distribution. — South America (Brazil).

172 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

CYRNEA SEURATI Lopez-Neyra, 1918, emended

Synonym. — Cyrnea seuratii Lopez-Neyra, 1918 ; Cyrnea scurati of
Lopez-Neyra, 1923, misspelt.

Host. — Primary : Caccabis rufa; secondary : Unknown.

Location. — Under corneous tunic of gizzard.

Morphology. — Cyrnea (p. 166) : Body robust, white, translucent,
the blood-red intestine showing through the body wall. Cuticle
finely striated transversely. Two large lateral lips, rounded at free
edge, teeth-like projections on inner surface. Dorsal and ventral lip
with free edge notched. No lateral alae.

Male 8 to 13.6 mm. long by 300 to 360 ft wide. Tail not rolled ;
caudal alae outspread, strongly striated transversely; their length
410 to 550/a, the span of the 2 alae 275 to 390/x,. Nine pairs of
long pedunculated papillae, of which 3 are preanal. Of the 6 post-
anal pairs, 4 pairs are small and grouped toward the caudal end.
No median papilla anterior to cloaca! aperture. Left spicule 1.25 to
1.39 mm., right spicule 450 to 500/* long. Gorgeret 65 to 70^ long.

Female 13 to 17 mm. long by 320 to 420 /x wide. Tail (fig. 241)
140 to ISOfi long. Vulva not salient, 700 to 950/x anterior to anus.
Ovejector made up of small pyriform vestibule 300 to 420/x. long and
a tubular sphincter. Eggs 45 to 48/u, long by 25 to 26^ wide.

Life history. — Unknown.

Distribution. — Europe ( Spain ) .

Genus HABRONEMA Diesing, 1861

GeneHc diagnosis. — Spirurinae (p. 164) : Mouth with 4 lips.
Body usually with 1 or 2 lateral alae. Sensory papillae precervical.
Male with unequal spicules; postanal papillae asymmetrically dis-
posed. A gorgeret present. Female with vulva in median region of
body though it may be anterior or posterior of middle. Ovejector
with vestibule not differentiated as an organ for the storage of ejrsrs.
Uteri divergent.

Parasitic in digestive tract, usually stomach, of birds and mammals.
Intermediate host, where known, arthropods (flies).

Type-species. — Habronema muscae (Carter, 1861) Diesing, 1861.

KEY TO SPECIES OF HABRONEMA

1. No lateral alae present Habronema ficheuri, p. 174.

One or more lateral alae present 2.

2. Only one lateral ala present 3.

Two lateral alae present although they may be poorly developed (//.

ieurati) _ ^

3. Long slender spicule 2y 2 times the length of the short broad one; vulva

prominent Habronema unilateralis, p. 186.

Long slender spicule over 3 times the length of the short broad one ; vulva
not prominent _ 4

NEMATODE PARASITES OF BIRDS 173

4. Lateral ala extends along anterior half of body ; left cervical papilla a little

anterior to the right ; short spicule 550 to OOO/a long ; gorgeret highly devel-
oped and of complex structure, being hollowed out and having a strong

dorsal spur Habronema tulostoma, p. 184.

Lateral ala does not extend posterior to termination of glandular esophagus :
right, cervical papilla anterior to the left ; short spicule not over 4G0m ;
gorgeret simple in structure or poorly developed and difficult to see 5.

5. Male 5.5 to 6.9, female 8 to 9.4 mm. long; cuticle with many fine needle-like

spines, 4.4-fi long; lateral ala 8 to 12/* wide; long spicule 1.2 to 1.4 mm.

long Habronema spinosa, p. 183.

Male 7.6 to 9.S, female 12.3 to 13.6 mm. long; no spines on cuticle; lateral
ala attains width of 18 to 22(i; long spicule 1.6 to 1.7 mm. long.

Habronema monoptera, p. 180.

6. Female 9.2 mm., male 5.5 mm. long. Vulva posterior to middle of body.

Longitudinal folds of caudal alae posterior to cloacal aperture have teeth-
like projections on free border Habronema seurati, p. 182.

Female 13 mm. or longer ; male 7 mm. or longer ; vulva anterior to middle
of body; no teeth-like projections on longitudinal folds of caudal alae 7.

7. Dorsal and ventral lips very small and apparently lacking a median pro-

jecting crest ; internal to the lips 2 rows of 3 papillae each, tipped by
denticles which project anteriorly beyond the lips.

Habronema colaptes, p. 173.

Dorsal and ventral lips well developed and having a median projecting crest ;

no rows of papillae as above S.

8. Lateral alae extending 1/2 of body length 9.

Lateral alae extending not more than 1/4 of body length 10.

9. Body rolled in spiral ; caudal alae symmetrical ; no unpaired papilla

immediately anterior to cloacal aperture Habronema longistriata, p. 178.

Body not rolled in spiral ; caudal alae slightly asymmetrical ; an unpaired
papilla present, immediately anterior to cloacal aperture.

Habronema mansioni, p. 179-

30. Lateral alae of unequal length, one about 1 mm. long, the other 50G> long,

their margins strongly crenulated ; anus of female about 200^ from

posterior extremity Habronema incerta, p. 175.

Lateral alae of equal length, extending 1/4 of body length ; margin not
crenulated ; anus of female about 360^ from posterior extremity.

Habronema leptoptera, p. 176.

HABRONEMA COLAPTES Walton, 1923

Host. — Primary: Colaptes auratus In frits; secondary: Unknown.

Location, — In the mucosa of the proventriculus.

Morphology. — Habronema (p. 172) : Yellowish-white worms of
medium thickness. Walton states that there are 2 lips but his figures
(fig. 244 <7, b and c) show, in addition, a dorsal and a ventral lip
present, though small, each divided into 2 lobes without a projecting
crest between them. Lips partially covered by a prepuce. Exter-
nally each of the 2 large lateral lips divided into 3 lobes, each lobe
with a small papilla; internally there are 2 rows of 3 papillae each.
tipped by denticles. Cervical papillae not observed. Lateral ahu-
ll arrow.

174

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 10 to 15 mm. long by 300/* wide. Caudal alae (Fig. 244e)
long and narrow, the ventral surface covered with longitudinal rows
of scale-like cuticular flaps. The alae are figured as slightly asym-
metrical. Eleven pairs of caudal papillae arranged as follows: 4
pairs of preanal, 2 pairs of postanal pedunculated, and 5 pairs of
small sessile ventral papillae at the caudal extremity. In addition
a single or occasionally paired sessile papilla on the left side of
the anterior edge of cloacal aperture. Spicules unequal (fig. 244^) , 2
mm. and 440/* long. Gorgeret present, 140/* long.

Female 15 to 30 mm. long by 400/* wide. Vulva not salient, an-
terior to middle of body. Ovejector short, claviform. Eggs 35 to

Fig. 244. — Habbonema colaptes. a, Dorsal; b, lateral; c, front view op head;

d. SPICULES AND GUBERNACULUM ; C, MALE TAIL ; f, FEMALE TAIL. AFTER WALTON,
1923

40/* long by 25 to 30/* wide, in the morula stage of development when
oviposited.

Life-history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution. — North America (United States (Monticello, Il-
linois)).

HABRONEMA FICHEURI Seurat, 1916d

Host. — Primary: Bubulcus lucidus; secondary; Unknown.

Location. — Gizzard.

Morphology. — Habronema (p. 172) :Body robust. No lateral alae.
Cuticle thick and heavily striated transversely. Two small precer-
vical symmetrical papillae and 2 intestinal papillae. Two large tri-
lobed lateral lips (fig. 245a), the median lobe with 3 teeth; dorsal
and ventral lips (fig. 2456) with keel-like projection.

Male 8.4 mm. long by 336/* wide. Tail (fig. 245c) 120/* long. Two
long caudal alae, transversely striated. Ventral region of body also
transversely striated for 450/* posterior to cloacal aperture. Four

NEMATODE PARASITES OF BIBDS

175

pairs of pedunculated preanal papillae; 1 large unpaired sessile
papilla on anterior margin of cloacal aperture; 2 pairs of large
postanal papillae and near the caudal extremity, a circular smooth
area carrying several very small papillae. Spicules unequal, the left
1.27 mm., the right 350/x long. Gorgeret 70^ long.

Female 12.8 mm. long by 385/x wide. Tail 170ft long. Vulva 6.6
mm. from anterior end of body, slightly salient. Vestibule 300/x
long; sphincter 240/x long; unpaired trompe 120/u. long, dividing into
2 branches 720> in length. Eggs 47/x long by 23/x wide, embryonated
at maturity.

Fig. 245. — Habronema fichecri. a, Lateral ; b, dorsal view of head ; o, male
tail. After Seurat, 1916

Life history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution. — Africa (Algeria ) .

HABRONEMA INCERTA (Smith. 1908) Gendrc. 1922

Synonym. — Spiroptera incerta Smith (A. J.), 1908.

Ilosts. — Primary : Bolborhynchus lineola, B. monachus, Chrysotis
auripalliata, C. guatemalae, C. leucocephalus, C. ochroptera, Conurtis
leucotis, G. pertinax, Eclectus roratus, Lophortyx californicus,
Palaeomis fasciatus, Platycercus barnardi, P. eximius, P. palladi-
ceps, Poeocephalus senegalus, Protogerys mrescens; secondary : Un-
known.

Location. — In mucosa of proventriculus.

Morphology.— Habronema (p. 172) : Cuticle finely striated trans-
versely. Two lateral alae (fig. 246 d and e) of unequal length, one
extending for about 1 mm. posterior from the head, the other 1/2
3612—27 13

176

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

that length; margin crenulated. Lateral lips (fig. 246 g and h)
large, thick, with 3 prominent teeth on anterior border, and on the
external surface midway is a horizontal row of 3 very small papillae.
Dorsal and ventral lips with sharp-edged projecting central prom-
inence and 2 lateral rounded unarmed latered lobes. Cuticle forms
collar around base of lips.

Male size differs in different birds, the average being 8 mm. long
by 31G> wide, the maximum 12 mm. long by 350/a wide. Caudal
alae slightly asymmetrical. Ventral surface of tail (fig. 246a)
covered with long rectangular plates arranged longitudinally. Ten
pairs of caudal papillae and 1 unpaired papilla on anterior margin
of cloacal aperture; 4 pairs are preanal and 6 pairs postanal. Clo-
acal aperture about 350/x from posterior end. Spicules unequal.

Fig. 246. — Habronema incerta. a, Male tail; b, eggs; c, female tail; d.

TRANSVERSE SECTION OP ANTERIOR END; e, ANTERIOR END; f, FEMALE, SHOWING
POSITION OF VULVA ; g, FRONT VIEW ; h, DORSO-VENTRAL VIEW OF HEAD. AFTER

Smith, Fox, and White, 190S

Female size varies in different birds, the average being 14 mm.
long by 45G> wide, the maximum 20 mm. long by 600/x wide. Anus
(fig. 246c) about 200^ from posterior end. Vulva (fig. 246/) difficult
to find, described as in the first third of length of body (figure
shows it behind the first third, but anterior to the middle of the
body). Eggs (fig. 2466) 34 to 42^ long by 16 to 20^ wide.

Life history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution,— North America (United States (Zoological Gardens,
Philadelphia, Pennsylvania)).

HABRONEMA LEPTOPTERA (Rudolphi. 1819) Seurat. 1914a

Synonyms. —Spiroptera leptoptera Rudolphi, 1819; Filaria lep-
toptera (Rudolphi, 1819) Schneider, 1866.

Hosts. — Primary: Accipiter nisus, Astur nisus, A. palumbarius,
Buteo vulgaris, Circus aeruginosus, C. cineraceus, C. cyaneus, C.

NEMATODE PARASITES OF BIRDS

177

rufus, Emberiza pecoris, Falco albicollis, F. ater, F. aurantius, F.
bidentatus, F. buteo, F. cachinans, F. cineraceus, F. cyaneus, F.
lananits, F. magnirostris, F. nisus, F. palumbarius, F. i-ufus, F. sub-
buteo, F. tinnunculus, F. tridentaius, F. xanthothorax, F. unicinctvis,
Harpagus bidentatus, Herpetotheres cachinans, Hypotriorchis sub-
buteo, MUvus regalis, Strix otus; secondary : Unknown.

Location. — Proventriculus.

Morphology. — Habronema (p. 172) : Body elongate, strongly stri-
ated. Two lateral alae on anterior 1/4 of body. Cervical papillae

Fig. 247.

-IIabuonkma leptoptera. a, Anterior end, dorsal view
After Seurat, 1914

b. male tail.

far anterior to nerve ring. Head (figs. 217a and 248a) distinctly
set off from body. Four lips, the 2 lateral very large, trilobed, with
3 large teeth; the dorsal and ventral lips carrying a prominent
median crest, and also ! papillae at the point of insertion of these
two lips. Muscular esophagus long and slender.

Male 7.1 mm. long by 720/x wide. Spicules unequal, the left long
and slender, 040/* in length ; the right 200//, long. Gorgeret 30//, long.
Two caudal alae (fig. 2475) each formed of 2 halves, the dorsal finely
striated transversely; the other, with longitudinal striations, is
folded over and built upon the preceding, partly hiding the ventral
surface of the body. Left ala thickened at edge, the papillae ter-

178

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

minating in it having a mushroom-like appearance. Four pairs of
preanal papillae, 2 pairs of asymmetrical postanal, and a group of
8 small sessile papillae immediately anterior to caudal extremity.

Female 14 mm. long by 325/x wide. Tail 360/i long. Vulva (fig.
248c) 8 mm. from posterior extremity. Eggs elliptical, 32/i, long by
20/u wide, embryonated at maturity.

Life history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution, — Europe and Africa (Algeria).

Fig. 248. — Habronema leptoptera. a, Head, lateral view ; b, receptacle

SEMINALE, JOINING OVIDUCT WITH UTERUS ; C, OVEJECTOR. AFTER SEURAT,

1914

HABRONEMA LONGISTRIATA (Molin, 1859) Gendre, 1922

Synonyms. — Spiroptera longistriata Molin, 1859, in part; Gheilo-
spirura longistriata (Molin 1859) Diesing, 1861, in part.

Hosts. — Primary : Pious campestris and P. jumana; secondary :
Unknown.

Location. — Proventriculus.

Morphology. — Habronema (p. 172) : Body rolled in spiral. Head
(fig. 249a and b) with 2 wedge-shaped lateral lips, each lip with
3 notches on its inner surface and 3 sharply pointed teeth on its
outer surface, and in addition 3 large papillae. Dorsal and ventral
lips each with a sharp tooth; 4 submedian papillae. Lateral alae
wide.

Male 6 to 9 mm. long by 200/* wide. Caudal alae (fig. 249c) wide,
symmetrical. No unpaired papilla anterior to cloacal aperture. In
other respects bursa apparently like that of H. inansioni (p. 179).

Female 10 to 16 mm. long by 400 to 500^ wide. Anus remote
from caudal apex. Vulva in posterior part of body.

Life history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution, — South America (Brazil).

NEMATODE PARASITES OF BIRDS
HABRONEMA MANSIONI Seurat, 1914

179

Synonyms. — Spiroptera longistriata Molin, 1850 in part.; Cheilo-
spirura longistriata (Molin, 1859) Diesing, 1861 in part.

Hosts. — Primary: Buteo vulgaris and Picus grammicus. Possibly
also in Milvus korschwm; secondary: Unknown.

Location. — Proventriculus.

Morphology. — Hah rone ma (p. 172) : Body robust, with wide trans-
verse striations; lateral alae on anterior half of body. Cervical
papillae far anterior to nerve ring. Mouth (fig. 250 a) with 2 large
lateral lips with 4 small teeth; dorsal and ventral lips prolonged in
middle by a sharp point.

Male 9 to 11 mm. long by 315/* wide. Tail (fig. 250 b) straight.
An unpaired papilla on left side of body 500/* anterior to origin of

Fig. 249.

-Habronema longistriata. a, Head, ventral view ; b, head, front
view ; c, male tail. After Drasche, 1884

caudal alae. Alae slightly asymmetrical, elongate (840/*). Each ala
composed of a dorsal part finely striated transversely, and a part
curved over the ventral face, with longitudinal striations. Four pairs
of pedunculated preanal papillae. 1 unpaired sessile papilla on the
left side of anterior edge of cloacal aperture, and 2 pairs of postanal
papillae of which the first pair is remarkably asymmetrical ; in addi-
tion at the caudal extremity a group of 8 very small papillae. Spi-
cules unequal, 680/* and 315/t long. Gorgeret present.

Female 13 to 17 mm. long by 400/* wide. Tail 240/* long. Vulva
(fig. 250 c) very small, situated slightly anterior to middle of body.
Ovejector like that of //. leptoptera (p. 17G). Eggs 28/* long by 19/*
wide, embryonated when oviposited.

Life history. — Unknown; probably involving insects as interme-
diate hosts.

Distribution. — South America (Brazil) and Europe (Corsica).

180

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Gendre's (1922) specimens from Milvus korschum conform to
Seurat's description of this species except that they are smaller in
size, the lateral alae are unequal, and one spicule length is differ-
ent, the left spicule measuring 870 to 930/i in length.

HABRONEMA MONOPTERA Gendre, 1922

Host. — Primary: ". Rapace nocturne"; secondary: Unknown.

Location. — Proventriculus.

Morphology. — Habronema (p. 172) : Body slender anteriorly,
thickened posteriorly. One lateral ala (fig. 251 a) , situated in cephalic
region on left side, originating 250 to 280/* from the anterior extrem-

3Q0JJ .

Fig. 250. — Habronema mansioni. a, Anterior end, vertical view :
tail ; c, ovejectou. After Seurat, 1914

h, male

ity and extending to the level of the termination of the glandular
esophagus, its greatest width 18 to 22/*. Cervical papillae in inverse
position from those of H. tulosto?na, the right one being anterior to
the left. Head (fig. 251 b and c) with 4 lips, the laterals trilobed,
rounded and flattened, with 4 long teeth on the inner surface of the
median lobe ; dorsal and ventral lip with central triangular tooth and
2 lateral semiglobular lobes, each bearing a large papilla.

Male 7.6 to 9.8 mm. long by 300 to 350/x wide. Lateral ala about
2.58 mm. long. Tail (fig. 251 d) 230 to 260/x long. Caudal alae long,
lanceolate, thick, each ala consisting of 2 folds. External or dorsal
surface of alae transversely striated;. internal surface covered with
longitudinal, parallel ribs and shields, which cover the ventral sur-

NEMATODE PARASITES OF BIRDS

181

face of the body for a distance of QlOfx anterior to cloacal aperture.
Eleven pairs of caudal papillae, of which 4 are preanal, 7 postanal,
and in addition a large unpaired papilla on the left side of the ante-
rior edge of cloacal aperture. Spicules (fig. 251 e and /) unequal, the

Fig. 251. — Habronbma monoptera. a, Anterior end; b, lateual up; c, mediax
lip; d, male tail; e, left spicule; f, right spicule; g, female tail. After
Gendre, 1922

one 1.56 to 1.68 mm. long, the other 380 to 460/* long. Gorgeret diffi-
cult to see.

Female 12.3 to 13.6 mm. long by 430 to 440^ wide. Lateral ala
about 2.98 mm. long. Tail (fig. 251 g) 180/* long. Vulva near middle
of body, sometimes a little anterior, sometimes a little posterior to
middle, not salient, difficult to find. Vestibule 810/x long. Eggs 38
to 42/x long by 17 to 19/x wide.

182

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably involving an insect as inter-
mediate host.

Distribution. — Africa (French Guinea (Labe)).

HABRONEMA SEURATI Skrjabin, 1917

Host. — Primary: Fateo cenchris; secondary: Unknown.

Location. — Gizzard.

Morphology. — Habronema (p. 172) : Body small, attenuated at two
extremities; cuticle transversely striated (fig. 252a), the striations
crenate. Lateral alae very feebly developed, originating a little pos-
terior to the level of the cervical papillae, which are 144/*. from the
anterior end. Two lateral lips, each with a tooth on the inner sur-
face; a dorsal and a ventral lip, each with a small acuminate tooth.
In general, lips like those of H. mansioni (p. 179).

Fig. 252. — Habronema seubati. a, Transverse striation of body ; b, cuticular

STRUCTURE OF VENTRAL SURFACE OF MALE TAIL ; C, MALE TAIL ; d, LEFT SPICULE ; €,
RIGHT SPICULE. AFTEB SKRJABIN, 1917

Male 5.5 mm. long by 300/*, wide. Caudal alae (fig. 252c) elongate,
longitudinally striated, the striations giving appearance of undula-
tions. Cuticle of ventral surface of caudal region anterior to cloacal
aperture has longitudinal folds 11 to 13/*. wide, the free border of
each fold with teethlike projections (fig. 2525). Posterior to the
cloacal aperture the ventral surface of body is smooth except for
three transverse rows of small projections posterior to the first pair
of postanal pedunculated papillae. Spicules (fig. 252d and e) un-
equal, 2.1 mm. and 450//, long. The pedunculated papillae number
4 pairs of preanal and 2 pairs of postanal ; the sessile papillae, 9 to
10 toward the end of the tail and 1 large papilla on the anterior edge
of cloacal aperture. Cloacal aperture 200/* from posterior end.

Female 9.2 mm. long by 370//, wide. Vulva 3.4 mm. from pos-
terior end, thus in the posterior part of body. Eggs 47//. by 27.5/x
wide.

NEMATODE PARASITES OF BIRDS

183

Life history. — Unknown; probably involving an insect as inter-
mediate host.

Distribution. — Asia (Siberia.)

HABRONEMA SPINOSA Gendre. 1922

Host. — Primary : Falco tinmuncuhis ; secondary : Unknown.
Location.- — Proventriculus.

Morphology. — Habronema (p. 172) : Body white, cuticle trans-
versely striated and bearing a multitude of fine needlelike spines

Fig. 253. — Habronema SPINOSA. a, Spines ok cuticle ; b, head end; c, median lip;
d, male tail; c, left spicule; f, sight spicule; g, gorgbket; h, female tail.
After GBNDEB, V.V22

(fig. 253a), visible at high magnification (560), their length about
4.4/x. Right cervical papilla anterior to left. One lateral ala, origi-
nating 2G to 30ft posterior to left papilla, which is 130 to 150/* from
the anterior end in the male, 1C>0 to 180/x. in the female. Ala shorter
and narrower than in //. monoptera; its width 8 to 12/x (see below
for length). Head (fig. 253&) very small; 4 lips all of same length.
The lateral lips with 3 or 4 teeth on the internal surface arc not as
large and projecting as in other species of Habronema, while the
dorsal and ventral lips (fig. 253c) are proportionately more highly
3612—27 14

184

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

developed and their lateral lobes again subdivided into second a rv
external lobes, bearing a papilla.

Male 5.47 to 6.86 mm. long by 210 to 260ju wide. Lateral ala 1.82
to 2.55 mm. long. Tail (fig. 253c?) 200 to 230/* long. Internal fold
of caudal alae completely covered with cuticular shields in longi-
tudinal series; they also cover the ventral surface of body to a dis-
tance 650/x anterior to the cloacal aperture. Cloacal lips large.
Eleven pairs of papillae, of which 4 are preanal, 7 postanal ; in addi-
tion a large unpaired papilla on left side of anterior edge of cloacal
aperture. Spicules (fig. 253e and /) unequal, 1.22 to 1.37 mm. and
340 to 410ju, long respectively. Gorgeret (fig. 253^), simple.

Female 8 to 9.37 mm. long by 380 to 410/x wide. Lateral ala 1.77
to 1.94 mm. long. Tail (fig. 253A) 130 to 150^ long. Vulva difficult
to see, a little posterior to middle of body. Vestibule 610/x long.
Eggs 37/x long by 21/x wide, embryonated.

Pig. 254. — Habronema tulostoma. a, Anterior end ; b, lateral lip ; c, dorsal
lip. After Seurat, 1914

Life history. — Unknown; probably involving an insect as inter-
mediate host.

Distribution. — France (Maine-et-Loire (La Chapelle St. Laud)).

HABRONEMA TULOSTOMA (Hcmprich and Ehrenberg, 1866) Gendre, 1922

Synonyms. — Spiroptera tulostoma Hemprich and Ehrenberg in
Schneider, 1866 ; Filaria tulostoma Schneider, 1866 ; Hahronenia uni-
lateralis (Molin, 1860) of Seurat, 1914/ misdet.

Host. — Primary : Neophron percnopterus (Vultur percnopterus )
and N. monachus; secondary : Unknown.

Morphology. — Habronema (p. 172) : Body blood red in color;
cuticle finely striated transversely. Mouth with 4 lips, the 2 lateral
(fig. 254&) quadrangular, trilobed on inner surface, each lobe with
a tooth; dorsal and ventral lips (fig. 254c) have a median keel and
terminate anteriorly in a point. Cervical papillae anterior to nerve
ring, the left slightly more anterior than the right. One large lat-
eral ala (fig. 254a), originating slightly posterior to the left papilla,
finely striated transversely, extending along anterior half of body.

NEMATODE PARASITES OP BIRDS

185

Male 17 mm. long (Schneider) or 13.9 to 16.7 mm. long (Gendre)
or 8 mm. long by 250/x wide (Seurat). Tail (fig. 255a.) straight.
According to Seurat's description, caudal alae very long, 1.23 mm.,
the span 300/*. ; cloacal aperture 300//, from posterior end ; 4 pairs of
long pedunculated preanal and 2 pairs of sessile postanal papillae
and a group of 6 small sessile papillae toward the caudal extremity;
in addition a large impaired papilla on the left side of anterior edge
of cloaca. Spicules dissimilar and unequal, 600/a and 1.9 mm. long
(Seurat), 510 to 550^ and 1.77 to 1.8 mm. long (Gendre). Gorgeret
(fig. 2556) highly developed, according to Seurat, hollowed out and
provided with a strong dorsal spur for the insertion of the muscles.

Female 21 mm. long (Schneider), 19.4 to 22.8 mm. long (Gendre),
11 to 14 mm. long (Seurat). Tail 140/x. long in Seurat's specimens,
260 to 310ju, long in those of Gendre. Vulva 1/3 of body length from

Fig. 255.

-Habeonema tulostoma. a, Male tail; b, cloacal region, showing

GORGERET ; C, OVE.TECTOR. AFTER SEURAT, 1014

anterior end (Seurat) or varying, sometimes a little anterior, some-
times a little posterior to middle of body (Gendre). Eggs 42^ long
by 25^ wide (Seurat), 35 to 39/* wide by 21 to 23/x wide (Gendre).

Life history. — Unknown; probably involving an insect as inter-
mediate host.

Distribution. — Asia (Turkey (Tor)) and Africa (French Guinea
(Labe) and Algeria).

The original description of this species in Schneider's Mona-
graphie is very brief. Seurat (1922/) identified his specimens from
Neophron percnopterus as Habronema unilateralis and made Filaria
tulostoma Schneider a synonym of that species. However, Gendre
notes certain differences between specimens collected by him from
Neophron monachus and the early description of Habronema uni-
lateralis; he considers H. tulostoma and II. unilateralis to be distinct
species and Seurat's description to apply to the former. Although
there are some differences between the descriptions of Seurat and
Gendre as noted above, they are very close together, if not identical,
and have certain features differing from Tlabronema unilateralis,

186

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

namely the relative length of spicules, the longer spicule in H. uni-
lateralis being only 2% times the length of the shorter, in their speci-
mens over 3 times as long; the 2 pairs of postanal papillae situated
between the cloacal aperture and the terminal group of small papillae
are long pedunculated papillae in H. unilateralism whereas they are
sessile papillae in the specimens of Gendre and Seurat. It is there-
fore thought advisable by the present writer to keep as a distinct
species the form from Neophron, species.

HABRONEMA UNILATERALIS (Molin, 1860) Seurat, 1914f

Synonyms. — Spiroptera unilateralis Molin, 1860&/ Spiroptera
unialata Molin, 1860.

Morphology. — Habronema (p. 172) : Body slender anteriorly,
rhynchus and R. mtellinus; secondary : Unknown.

Location. — Esophagus and "ventriculus" (gizzard or proventri-
culus ? ) .

Fig. 256,

-Habronema unilateralis. a, Head, front view ; b, male tail.
Drasche, 1884

Morphology. — Habronema (p. 172) : Head continuous with body;
anterior extremity noticeably attenuated, posterior of increased size.
Cuticle transversely striated. Mouth (fig. 256a) with 2 lateral lips,
the median edge notched; each lip with a large lobe and 3 papillae.
Dorsal and ventral lips tooth-like. Four large submedian papillae.
Body with one lateral ala, moderately wide.

Male 7 to 9 mm. long by 20(V wide. Caudal extremity (fig. 2566)
semispiral; caudal alae moderately wide. Tail with 6 pairs of long
pedunculated papillae, of which 4 pairs are preanal and 2 pairs post-
anal; in addition a terminal group of 3 or 4 pairs of small ses?ile
papillae and an unpaired papilla directly anterior to the cloacal
aperture. Left spicule long, slender, alate, 2y 2 times the length of
the right, which is short and thick.

Female 7 to 20 mm. long by 300 to 500/*. or more wide. Tail
straight, short. Anus near the caudal apex. Vulva prominent, in
posterior part of body.

NEMATODE PARASITES OF BIRDS 187

Life history. — Unknown ; probably involving an insect as inter-
mediate host.

Distribution. — South America (Brazil).

For discussion of the relation of this species to II. tuJostotna, see

page 185.

Genus HADJELIA Seurat, 1916c

Synonym. — Gilsonia Gedoelst, 1919.

Generic diagnosis. — Spirurinae (p. 161) : Head distinct. Cuticle
thick, finely striated transversely. No lateral alae; cervical papillae
at level of nerve ring. Mouth limited by 2 lateral trilobed lips; cer-
vical cuticula prolonged anteriorly to form a discreet collar ("cadre"
of Seurat) which is 6-lobed and provided with 2 pairs of papillae.
Male with unequal spicules and with well-developed caudal alae.
Female with vulva in anterior part of body, opening anterior to
end of esophagus; ovejector tubular, very long. Uteri divergent;
eggs embryonated at maturity.

Parasitic in gizzard of birds, in the hollow tunnels under the
corneous tunic.

Type-species. — Hadjelia Ihuillieri Seurat, 1916c.

KEY TO SPECIES OF HADJELIA

1. Eggs 21(1 long Hadjelia truncata, p. 190.

Eggs 46/x or longer 2.

2. Females not over 9.7 mm. long ; buccal cavity of female 32 to 37/x long.

Hadjelia parva, p. 189.
Females 12 mm. or longer ; buccal cavity of female 50> or longer 3.

3. Vulva salient and limited by 4 or o large nipple-like protuberances formed by

the cuticle and usually arranged cross-wise Hadjelia inermis, p. 188.

Vulva only slightly salient, with no such cuticular formations described.
Male unknown Hadjelia Ihuillieri, p. 187.

HADJELIA LHUILLIERI Seurat, 1916c

Host. — Primary : Gaccabis petrosa; secondary : Unknown.

Location. — Gizzard.

Morphology. — Iladfelia (p. 187) : Blood red in color, slender,
greatly attenuated anteriorly. Head distinct. No lateral ala. Cervi-
cal papillae at level of nerve ring. Mouth with 2 lateral trilobed lips
(fig. 257), the marginal lobes larger than the median. Six lobes
to cuticular collar, of which 2 are lateral, 2 dorsal and 2 ventral; a
large papilla on each dorsal and ventral lobe, thus making a total of
4 papillae. Buccal cavity cylindrical, short.

Male unknown.

Female 19.5 mm. long by 28G> wide. Buccal cavity 60/x, entire
esophagus 3.1 mm. long, muscular esophagus 48G>. Tail short, anus
110/x, from posterior end. Vulva 2.9 mm. from anterior end of body,

188

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

or 335ya anterior to posterior end of esophagus. Ovejector (fig. 258)
tubular, elongated (3 mm. long), directed anteriorly. Uteri diver-
gent.

Life history. — Unknown.

Distribution. — Africa (Bou-Saada, Algeria).

Fig. 257. — Hadjelia lhuillieri. a, Head end, lateral view ; b, head, dorsal view.

After Seurat, 1916

HADJELIA INERMIS (Gedoelst, 1919) Gendre, 1922

Synonym. — Gilsonia inermis Gedoelst, 1919.

Hosts. — Primary : Buchanga atra, var. assimilis, Cranorrhinus cor-
rugatus, Halcyon chelicutensis, Irrisor erythrorhynchus, Lophoceros
semifasciatus, Oriolus auratus, Terpsiphone, species ; secondary : Un-
known.

Location. — In mucosa of gizzard.

Fig. 258. — Hadjelia lhuillieri. Vulva and ovejectok. Aftee Seurat, 1916

Morphology. — Hadjelia (p. 187) : Buccal cavity not cylindrical
but elliptical, its Jong axis being dorso-ventral.

Male 6.1 to 6.7 mm, long by 140 to 160/* wide. Buccal cavity 45
to 55ft, esophagus 2 mm. long. Cloacal aperture 120tt from posterior
end. Caudal extremity tightly curled; ventral surface covered with
longitudinal ridges, extending from the anterior end of the caudal
alae to the position of a transverse fold of cuticle near the posterior
end. Transverse fold obscures a pair of subterminal papillae (fig.
259 a and b). Spicules unequal, the left (fig. 259c) 1.6 to 2.1 mm.
long, the right (fig. 259rf) 200 to 280 /Jt loner.

NEMATODE PARASITES OF BIRDS

189

Female 12 to 21 mm. long by 170 to 260> wide. Buccal cavity 50
to 60/a, esophagus 2.4 to 3.6 mm. long. Anus (fig. 259<g) 90 to
ISO/x from tail end. Vulva 1.8 to 2.9 mm. from cephalic end, salient,
limited by 4 or 5 large nipple-like prominences formed by the cuticle
and usually arranged cross-wise. Eggs 46 to 57/u, long by 30 to 33/x
wide.

Life history. — Unknown.

Distribution. — Africa (Dahomey) .

The host and the character of the cuticular formation around the
vulva appear to be the only differences between the female of this
species and that of H. Ihuillieri, the male of the latter being un-

Fig. 259. — Hadjelia inermis. a, Male tail as usually seen (most posterior

PAPILLAE HIDDEN) ; b, MALE TAIL STRAIGHTENED OUT; C, LEFT SPICULE", d , RIGHT
SPICULE ; e, FEMALE TAIL. AFTER GENDRE, 1922

known. Vulvar prominences or "papillae" have been found to
vary in number and constancy of appearance in species of othei
genera (see Heterakis isolonche, p. 65). There is a possibility that
this species and that of Seurat may prove identical.

HADJELIA PARVA Gendre, 1922

Host. — Primary : Tracltelotis sencgalensis; secondary : Unknown.

Location. — Gizzard.

M ' orphology .—LladjeUa (p. 187) : Male 5.5 inm. long by 180/x
wide. Buccal cavity 33/* long; first part of esophagus 240//,, second
part 1.66 mm. long. Cloacal aperture 110/i from posterior end. Six
pairs of pedunculated papillae (fig. 260r<) on large caudal alae.
Spicules unequal, the left 1.29 mm., the right 240/i. long.

190

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 9.2 to 9.6 mm. long by 200 to 260/a wide. Buccal cavity
32 to 37/u, long; first part of esophagus 280 to 330/u,, second part 1.8(5
to 1.90 mm. long. Anus (fig. 2606) 110/x, from tail end. Vulva
1.52 to 1.97 mm. from head end. Eggs 48 to 49/t long by 29 to 30/*
wide.

Life history. — Unknown.

Distribution. — Africa (Abomey, Dahomey).

Fig. 260. — Hadjelia parva. a, Male tail; b, female tail. After Gendre, 1922
HADJELIA TRUNCATA (Creplin, 1825) Gendre, 1921c

/Synonyms. — Spiroptera upupae Rudolphi, 1819, nomen nudum;
Spiroptera truncata Creplin, 1825; Bispharagus truncatus (Creplin,
1825) Dujardin, 1845; Acuaria upupce-epopis Molin, 18606.

Hosts. — Primary: Coracias garrula and Upupa epops; secondary:
Unknown.

Morphology. — Hadjelia (p. 187): Body slender; color, red.
Mouth (fig. 261a) with 2 large lips, their base concave, their anterior

Fig. 261. — Hadjelia truncata. a, Head; b, male tail. After Mueller, 1897

portion projecting sharply outward; cuticular collar with 6 lobes,
anteriorly directed, 2 of which (the laterals?) are large and rounded,
the other 4 (dorsals and ventrals?) slender and sharply pointed.
Four papillae. Cuticle transversely striated.

Male 5 to 7 mm. long. Caudal extremity (fig. 2616) curled in
semispiral; caudal alae long, the right ala a little longer than the
left, according to Mueller. Six pairs of long caudal papillae, of
which 4 are preanal and 2 postanal. Spicules very unequal, the left
1.6 mm. long, the right 220//, long.

NEMATODE PARASITES OF BIRDS 191

Female 10 to 16 mm. long by 300/x wide. Caudal end obtuse, anus
near the extremity. Vulva 2.6 mm. from head end of a 16 mm.
long specimen, prominent, with 2 large lips. Eggs 27/*. long.

Life history. — Unknown.

Distribution. — Europe (France and Germany).

Genus HARTERTIA Seurat, 1915b

Generic diagnosis. — Spirurinae (p. 164) : Nematodes having the
appearance of an ascarid, with large body, often curved into an arc
with dorsal concavity. Cuticle thick, resistant, finely striated trans-
versely. Lateral alae, when they exist, are limited to the anterior
region (cephalic or esophageal). Mouth limited laterally by 2 large
cuneiform lips, the internal surface deeply divided into 3 lobes or
jaws. Precervical papillae far anterior, situated in the cephalic
region, anterior to origin of esophagus. Male with caudal alae
symmetrical, outspread. Long pedunculated papillae, of which 4
pairs are preanal. Spicules unequal, the left slender, filiform.
Gorgeret present or absent.

Parasitic usually in the intestine, sometimes the gizzard, of birds.

Type- specws. — Hartertia obesa Seurat, 1915&.

KEY TO SPECIES OF HARTERTIA

1. Only 3rd stage larva known (from under skin of carnivore).

Hartertia zorillae, p. 200.
Adults known, in birds 2.

2. Female 12 mm. long ; esophagus 1/3.5 of total body length.

Hartertia zakharowi, p. 200.

Female 20 mm. long or longer ; esophagus 1/6 to 1/17 of total body length- 3.

3. Male 28 to 40 mm., female 60 to 110 mm. long ; esophagus 1/17 of total length,

or shorter ; spicules 800^ and 200m long Hartertia gallinarum, p. 197.

Male not over 37 mm., female not over 66 mm. long; esophagus 1/10 of total
length, or longer ; spicules 1.35 mm. and 450/x, or longer 4.

4. Male 35 to 37 mm. long ; long spicule 1.35 mm. long ; vulva of female posterior

to anterior third of body Hartertia rotundata, p. 198.

Male not over 30 mm. long; long spicule over 2 mm. long; vulva of female
in anterior third of body 5.

5. Male 15.6 mm., female 20.3 mm. long. Tail of male 1/40, of female 1/20 of total

body length ; eggs 39/* long by 26^ wide Hartertia confusa, p. 195.

Male 23.5 mm., female 41 mm. or longer. Tail of male shorter than above
(1/73 of total length) or longer than above (1/31 of total length) ; tail of
female shorter than above (1/27 to 1/56 of total length) ; eggs 50 to 56/u
long by 35p wide 6.

6. Tail of male 1/73 of total body length ; spicules 3.3 mm. and 647//. long ; no

gorgeret. Female with tail 1/56 of total body length ; vestibule not pyri-
form ; sphincter with cul-de-sac projection where it joins the trompe.

Panisitic in gizzard Hartertia annulata, p. 193.

Tail of male 1/31 of total body length; spicules 2.1 mm. and 500jtt long;
gorgeret present. Female with tail 1/27 of total body length ; vestibule
pyriform ; no cul-de-sac of sphincter where it joins trompe. Parasitic in
intestine Hartertia obesa, p. 102.

192 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

HARTERTIA OBESA Seurat, 1915b

Hosts. — Primary: Caccdbis petrosa spatzi, Oedicnemus capensis,
Otis afroides, O. ruftcresta, and Plocepasser mahali; secondary:
Unknown, probably insects.

Location. — Intestine.

Morphology. — Hartertia (p. 191) : Head very small, distinct.
Cuticle finely striated. Lateral fields very large (155/*), opaque.
Polymyarian. No lateral alae. Lips (fig. 262) divided into 3 equal
lobes with very thick edges. Two papillae on each lip.

Male 23.5 mm. long by 1.07 mm. wide. Precervical papillae 145/*
from cephalic extremity. Buccal cavity 160/* long ; esophagus 1/6 of
body length. Tail (fig. 263&) straight. Ventral surface of tail in
region of cloacal aperture covered with longitudinal rows of cuti-

Fig. 262. — Hartertia oeesa. a, Dorsal ; b, lateral ; c, front view of head ;
d, egg. After Sedrat, 1915

cular shields. Cloacal aperture 750/* from posterior end. Caudal
alae 1.3 mm. long; 6 pairs of long pedunculated papillae and in
addition 4 pairs of small sessile papillae near caudal extremity.
Upper lip of cloacal aperture somewhat projecting, triangular.
Spicules unequal, 2.1 mm. and 500/* long, respectively. Gorgeret
present, 200/* long.

Female 40.8 mm. long, by 1.6 mm. wide. Buccal cavity 180/* long ;
total length of esophagus 1/10 of body length. Tail 1.5 mm. long,
rounded at end. Vulva (fig. 263a) not salient, seen with difficulty,
situated at anterior third of body. Ovejector composed of pyriform
vestibule 700/* long, directed anteriorly, a very long sphincter (1.5
mm.) originating at middle of vestibule, and recurved, unpaired
trompe 1.5 mm. long. Uteri divergent. Embryonated eggs (fig.
262c?) peculiar, the thick shell measuring 56 by 35/*, is doubled by a
vitelline membrane, showing in the meridian plane as a thin ring
10/* wide.

NEMATODE PARASITES OF BIRDS

193

Life history. — Unknown; probably similar to that of H. galll-
narum (p. 197).
Distribution. — Africa (Oued N'za and Transvaal).

HARTERTIA ANNULATA, new species

Host. — Primary: Red-headed pheasant (Ptemistes, species);
secondary: Unknown, probably insects.

Location. — Gizzard.

Morphology. — Hartertia (p. 191) : Large thick worms, grayish-
white. Cuticle transversely striated and in addition thrown into
folds or annulations. Mouth (fig. 264 a, b, and c) with 2 lateral lips,
each trilobed, the lateral lobes finely denticulated and all 3 lobes

Via. 263. — Hautertia obbsa. a, Ovejector ; b, male tail. After Seurat, 1915

carrying several large teeth. Head constricted posterior to the lips
in dorso- ventral view. Lateral papillae precervical, situated near the
base of the lips. No other papillae observed on lips. Mouth cavity
very short. Esophagus two-part, the nerve ring situated at the
middle of the anterior part. No lateral alae.

Male 2G to 30 mm. long by 800 to 900/x wide. Anterior part of
esophagus 531/a long by 144/x wide, posterior part 3.G5 mm. long, its
minimum width 183/x, maximum width 3GG/x. Posterior extremity
of body (fig. 2G5) straight or loosely curled. Caudal alae 1.4 mm.
long; they are narrow for a distance of 133ju, from the posterior
end, anterior to that they flare out comparatively wide, their span
being G31ju. Ventral surface densely covered with longitudinal rows
of small rounded protuberances. Four pairs of pedunculated pivanal,
2 pairs of pedunculated postanal papillae, all pairs symmetrical.
Cloacal aperture 382/x from tail end. Two dissimilar spicules (fig.

194

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

265), the one 3.3 mm. long by 17/x wide, its distal end sharply
barbed, there being 3 or more recurrent points or processes; the
other 674/a long by 58ju. wide, its distal end bluntly rounded.

Female 43 to 62 mm. long by 966/x to 1.2 mm. wide. Anterior
part of esophagus 697/a, posterior part about 5.2 mm. long in large
specimens. Anus 1.1 mm. from posterior end of large specimen;

oz mm.

/mm

Fig. 264.— Harteria annulata. a, Lateral ; b, front ; c, dorsal view of head ;

d, EGG ; e, FEMALE TAIL ; /, OVEJECTOR ; g, ENLARGED VIEW OF CUL-DE-SAC AT UNION
OF SPHINCTER AND TRO.MPB. ORIGINAL

rectum 199/* long, cylindrical, with thin chitinous lining (fig. 264e).
A loop of the uterus lies ventral to the intestine just anterior to
the rectum. Vulva 14 mm. from anterior extremity of specimen
62 mm. long, thus a little less than V4 of total length from anterior
end. Vulva difficult to find in the folds of cuticle, a small simple
opening with 2 narrow lips, their length being parallel to the long

Fig. 265. — Hartertia annulata. Male tail and spicules. Original

axis of the worm. Length of vestibule and sphincter 1.4 mm.;
length of unpaired trompe 2 mm. The vestibule is of a very
unusual character; near its union with the trompe it has a pyriform
swelling which continues beyond the point where the trompe opens,
and ends blindly in a cul-de-sac (fig. 264 / and g). The terminal
portion of the latter is a solid mass of circular muscle fibers, no lumen

NEMATODE PARASITES OF BIRDS 195

being: present. The muscular development is pronounced also around
the opening of the trompe into the vestibule, the appearance being
almost that of a valvular arrangement. The uteri are at first parallel
but later diverge; they are very large, attaining a diameter of
500^ and containing an enormous number of eggs. The ovaries
are long and filiform, 83/x in diameter. Eggs 50ju, long by
35/n wide, embryonated when oviposited (fig. 2('Ad).

Life history. — Unknown, probably similar to that of H. galli-
narum (p. 197).

Distribution. — Africa (Union of South Africa (Potchefstroom)).

Type material. — No. 25546, U.S.N.M. (Bureau of Animal Industry
helminthological collection) .

This material was collected by R. O. Wahl, Entomologist at the
School of Agriculture^ Potchefstroom.

HARTERTIA CONFUSA, new name

Synonyms. — Physaloptera rotundata Linstow, 1906 b; Habronema
rotundata (Linstow, 1906) Seurat, 1914 /, not Habronema rotundata
(Linstow, 1883) Seurat, 1914 g and n.

Host. — Primary: Otis houbara; secondary: Unknown, probably
insects.

Location. — Intestine.

Morphology. — Hartertia (p. 191) : Cuticle cross-striated at wide
intervals. Head with 2 large lips, each of which has a prominent
papilla on each side.

Male 15.6 mm. long by 1.1 mm. wide. Esophagus 1/8.6, tail 1/40
of total body length. Caudal alae (fig. 266) narrow; median section
of ventral surface of tail covered with swellings arranged in longi-
tudinal lines. Six pairs of pedunculated papillae, of which 4 are
preanal, 2 postanal; in addition at the posterior end of tail are 5
Sessile papillae arranged as 2 pairs and a median unpaired papilla
anterior to them. Spicules unequal, the left 2.17 mm. long, the
right 510/x long, both with rounded ends.

Female 20.3 mm. long by 1.5 mm. wide. Esophagus 1/9, the narrow
cone-shaped tail 1/20 of total body length. Vulva in anterior third of
body, dividing body length in ratio of 6:15. Eggs 39/* long by 26/x
wide, thick-shelled, embryonated.

LJfe history. — Unknown; probably similar to that of //. galli-
narum (p. 197).

Distribution. — Europe (Germany (Zoological Museum, Konigs-
berg)).

Considerable confusion seems to have existed between the nematode
described by Linstow in 1883 as Fit aria rotundata and that described
by him in 1906 as Physaloptera rotundata. The one described in

196

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

1883 was from Otis macqueeni, collected in Turkestan by Professor
Fedtschenko; the 1906 specimens were described as a new species
from Otis houbara, in the Zoological Museum at Konigsberg.
Seurat (1914;117) found Filaria rotundata Linstow, 1883, in Hou-
bara undulata in Algeria ; he redescribed it and placed it in the genus
Habronema and later (1915 ;14) transferred it to his new genus
Hartertia. Skrjabin (1916 ;501) found this same nematode in Rus-
sian Turkestan in Oedicnemus oedicnemus.

Physaloptera rotundata Linstow, 1906 was transferred to Hab-
loneina by Seurat (1914; 153) but he gives the locality of collection of
the nematode as Algeria and Turkestan, which is not that of the 1906

o/m/n

imm

Figs. 260-268. — 266, Hartertia confusa. Male tail. After Linstow, 1906. 267,
Hartertia gallinarum. a, Lateral ; b, feont view of head ; c, female tail. 268,

OVEJECTOR AND UTERI. ORIGINAL

species but of Linstow's earlier species, Filaria rotundata Linstow,
1883.

In order to straighten out the confusion caused by this use of the
same specific name by the same author at different dates, the present
writer has renamed the species and placed it, at least provisionally,
in the genus Hartertia. The description as given by Linstow makes
no mention of certain characters included in the generic diagnosis of
Hartertia, as the trilobed division of the inner surface of the lips,
but the species seems to be quite similar to Hartertia obesa, the type-
species of the genus.

This species, renamed Hartertia confusa, synonym Physaloptera
rotundata Linstow, 1906, differs from Hartertia rotundata (Linstow,
1883) in the size of the body (H. confusa being less than half the
length of the smallest specimens of H. rotundata), in absence of
lateral alae, in a somewhat more anterior position of the vulva, in
the much smaller size of the eggs, in different spicule lengths, width

NEMATODE PARASITES OF BIRDS 197

of male tail (that of II. confusa being narrow, that of H. rotundata
wide), and number and arrangement of small papillae at caudal ex-
tremity.

HARTERTIA GALLINARUM (Theiler, 1919) Cram, 1927

Synonym. — FUaria gallinarvm Theiler, 1919.

Hosts. — Primary: G alius gallus; secondary: Termites (Hodoter-
mes pretoriensis) .

Location. — Usually in jejunum, occasionally in stomach, of pri-
mary host ; in the body cavity of the secondary host.

Morphology. — Hartertia (p. 191) : Filiform worms, attenuating at
each end. Cuticle transversely striated. Head (fig. 267« and b)
set off by a slight constriction. Two lips, the inner surface of each
lip being divided by 2 ridges into three cup-like grooves, lined by a
thick cuticle. Each lip carries a thorn-like lateral and two circular
submedian papillae. Two cervical papillae a short distance behind
the lateral cephalic papillae. Esophagus 3.5 to 6 mm. long.

Male 28 to 40 mm. long by 672 to 688/* wide, terminating in a
spiral. Esophagus 1/10 of total body length. Cloacal aperture 352
to 498/x (1/80 of total body length) from posterior end. Tail (fig.
269) slightly curved ventrally and with caudal alae supported by 6
pairs of pedunculated papillae, of which 4 are preanal and 2 are
postanal; in addition there is an unpaired sessile papilla situated in
the median line just anterior to the cloacal aperture and 4 pairs of
very small sessile papillae near the caudal extremity. Posterior to
the pedunculated papillae the diameter of the tail decreases sud-
denly. The ventral caudal surface (for a distance of 1.2 mm. from
posterior end in a 40 mm. specimen) covered with oval or rectangular
tubercles in linear arrangement. These tubercles are lacking in the
median region of the tail, from the cloacal aperture to the posterior
end, and on the anterior lip of the cloacal aperture. According to
the original description of this species, the left spicule 800/* long, the
right spicule 200/* long; according to measurements made by the
present writer on 4 specimens of 40 mm. length, the left spicule is
2.3 mm. long, the right spicule 630/* long. No explanation of this
discrepancy in the spicule measurements as made by Theiler and by
the present writer can be offered at this time. The long, slender
spicule (20/* wide), near its sharply pointed end, bears 4 large barbs;
the shorter spicule is 48/* wide and bluntly rounded.

Female 60 to 110 mm. long by 736 to 784/* wide near posterior
end of esophagus. Esophagus 1/17, tail 1/77 of total body length.
Vulva 16 to 24 mm. from the anterior end. inconspicuous, with no
thickening of the cuticle. In specimen 100 mm. long, vestibule and
sphincter 830/* long, unpaired trompe 1.9 mm. long (fig. 268). Eggs

198

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

4(V long by 24/x wide, containing fully developed embryos when
deposited.

Larva, second stage, 12 to 35 mm. long in intermediate host.

Life history. — The eggs from the female worms pass out in the
droppings of the chickens and are ingested by the white ants, or
termites, locally known as the " Houtkapper." Only worker ants are
found infested, the soldier ants not being found infested. The
embryo escapes from the shell and passes to the abdominal cavity,
where the young worm developes to the second larval stage. When
infested ants are fed to chickens, the worms continue their develop-
ment in the chicken intestine and become mature worms in 3 weeks.

Pig. 269. — Hartertia gallinauum. Male tail and spicules. Original

Distribution. — Africa (Orange Free State and Belgian Congo).

Through the courtesy of Sir Arnold Theiler and of Dr. H. O.
Monnig, the present writer has been granted permission to make this
further study and allocation of Filaria gallinarum, from specimens
which had been furnished to the U. S. Bureau of Animal Industry
collections by Doctor Theiler. The nematode appears to agree in all
respects with the generic diagnosis of Hartertia.

HARTERTIA ROTUNDATA (Linstow, 1883) Seurat. 1915b

Synoyins. — Filaria rotundata linstow, 1883; Habronema rotun-
data (Linstow, 1883) Seurat, 1914 g and n, not Habronema rotun-
data (Linstow, 1906) Seurat, 1914 / (see Hartertia confusa, p. 195).

Hosts. — Primary : Houbara undalaia, Oedienemus oedicnemus,
O. vermicularis, Otis afroides, O. nvacqueeni, O. ruficresta and
Plocepasser mahali. Immature forms, probably of this species,

NEMATODE PARASITES OF BIRDS

199

have been reported from numerous other birds by Monnig. Second-
ary: Unknown, probably insects.

Location, — Intestine and cecum.

Morphology. — Hartertia (p. 191) : Color blood-red. Lateral alae
for about 2 mm. length in anterior part of body; this part of body
bent hook-like dorsally. Body robust anteriorly, progressively
slenderer in posterior 2/3. Cuticle with fine transverse striations,
the alae in addition to these transverse striations having a lonm-
tudinal line dividing them in two halves, the outer half more deeply
colored. Two papillae at anterior insertion of alae. Mouth (fig.
270#) with 2 tri-lobed lateral lips, the median lobe having a small
tooth, the lateral lobes a finelv denticulated edge. Length of
esophagus 1/7 that of the body.

Fig. 270. — Hartertia rotundata.

a, Head, ventral view ; b, male tail.
Sbubat, 1914

Aft etc

Male 35 to 37 mm. long by G60/* wide. Caudal alae (fig. 270&)
symmetrical, 1.2 mm. long by 720/u, wide. Six pairs of long pedun-
culated papillae, of which 4 are preanal, in addition a group of 5
pairs of small papillae in the region of the caudal extremity. Ven-
tral surface covered with protuberances. Cloacal aperture 540ti
from tail end. Spicules unequal, 1.35 mm. and 450/* long. Gorgeret
present.

Female 35 to 66 mm. long by 900/* wide at a level of alae. Cephalic
papillae 70/u, from anterior end. Tail long, 1.2 mm. in a 35 mm. speci-
men. Vulva posterior to anterior third of body. Ovejector (fig. 271)
composed of pyriform vestibule 600/* long, sphincter 000//, long,
unpaired trompe 1.5 mm. long. Uteri divergent. Eggs thick-
shelled, 62 to 69/x long by 42 to 56ti wide, embryonated at maturity.

Life history. — Unknown: probably similar to that of //. galli-
naruni (p. 107).

Distribution. — Asia (Turkestan, Russian Turkestan) and Africa
(Algeria and Transvaal).

200 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

HARTERTIA ZAKHAROWI Skrjabin, 1920

Host. — Primary: Lanius minor; secondary: Unknown, probably
insects.

Location. — Intestine.

Morphology.— Hartertia (p. 191) : Body massive, twisted in the
dorsal direction. Mouth with 2 prominent lateral lips, each of which
on close inspection is seen to consist of 3 lobes, a median and 2 sub-
median. Each of the 3 lobes is characterized on the inner surface by
specially formed prominences situated parallel to the outer edge of
the lobe. The prominences appear to be the probable homologue of
those (that is, probably the "pulp structure ") of the lips in several
species of ascarids. The most characteristic feature of this species
consists in the situation of the cephalic papillae. Each lip bears 5
cephalic papillae: in the lateral field, on the median lobe, is 1 large

Fig. 271. — Hartertia rotundata. Ovejector. Aftek Seurat, 1914

papilla and on the submedian lobes are 2 parallel pairs of papillae,
one above the other.

Male unknown.

Female 12 mm. long, the maximum width 1 mm., in middle of
body ; the width of the body at the posterior region of the esophagus
is 900/1,, at the level of the anus 400/x. Esophagus 3.5 mm. long. Anus
550/i from posterior end.

Life history. — Unknown; probably similar to that of H. gallin-
arum (p. 197).

Distribution. — Europe (Russia) .

The present writer is indebted to Dr. Robert Formad for aid in
translating the Russian description of this species. As Doctor For-
mad is not well acquainted with technical terms of helminthology,
the writer has had to supply them by interpretation from the struc-
ture of other species of Hartertia, and the resulting description must
therefore be used with reservation.

HARTERTIA ZORILLAE Seurat, 1919b

Host. — A weasel, Zorilla lybica.
Location. — Under the skin.

NEMATODE PARASITES OF BIRDS

201

Morphology. — Third stage larva (fig. 272) : 13.1 mm. long by
410> wide; tail 158//, long. Precervical papillae 77/x from cephalic
end; buccal cavity 60/*, muscular esophagus 440/*, glandular esopha-
gus 4.3 mm. long.

Distribution. — Africa (Algeria).

This larva had apparently encysted in an aberrant host. Since
the adult is probably a bird form, the above description is included
here.

Genus VIGUIERA Seurat, 1913b

Generic diagnosis. — Spirurinae (p. 164) : Body suddenly narrowed
in anterior region. Cephalic extremity covered by a circular disk
which projects slightly from the underlying part; posterior to this
compression, 2 papillae. Buccal cavity cylindrical, with thick walls,

Fig. 272. — Hartertia zorillab. Third stage larva. «, Lateral view of head;
&, ventral view of head ; c, ventral ; d, lateral view of tail. After Seurat,
1919

30 to 50//. long. No lateral alae. Two precervical papillae situated
far anterior to the nerve ring. Male with caudal extremity rolled
in spiral; bursal alae asymmetrical; 9 preanal papillae (Seurat says
9 pairs but figures them as single in the type-species) on the left
side, 7 on the right side ; 2 pairs of postanal papillae. Spicules very
unequal. No gorgeret. Female with vulva in posterior region of
body, near anus.

Parasitic in proventriculus of birds.

Type-species. — Viguiera eun/optera (Rudolphi, 1819) Seurat,
1913.

VIGUIERA EURYOPTERA (Rudolphi, 1979) Seurat, 1913b

Synonym,. — Spiroptera eumjoptera Rudolphi, 1819.

Host. — Primary: Lanius collurio, L. excubitor, L. minor, L. rufus;
also " pie-grieche a tete rouge " ; secondary : Unknown.

Location. — Proventriculus.

Morphology. — Viguiera (p. 201) : Cuticle thick, transversely stri-
ated. Muscular esophagus very slender, surrounded toward its
posterior third by the nerve ring, posterior to which is situated the
excretory pore.

202

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 5.2 to 6.75 mm. long by 125[/. wide. In the specimens 5.2 mm.
long, the buccal cavity measured 32//,, muscular esophagus 370[jt,, total
esophagus 2.7 mm. long. Tail strongly rolled in spiral. Caudal alae
(fig. 273a) wide, attaining length of 580/*. Left ala longer and
especially wider, than right; a furrow divides the alae throughout
their whole length, at about the middle of their width. Left ala, in
addition, is marked with fine transverse striations in its inner half.
Nine preanal papillae (Seurat says 9 pairs but figures them single)
on the left side, 7 on the right, moderately pedunculated; an un-
paired papilla on the anterior edge of cloacal aperture (Seurat notes
that it is not shown in figure) and 2 pairs of postanal papillae.
Spicules very unequal, 930/* and 192[* long.

Fig. 273. — Viguiera euryoptera.

a, Male tail ; b, ovejector. After Seurat,
1913

Female 9 to 10 mm. long by 290/* wide. Total length of esophagus
is 1/3 body length; muscular esophagus 780/* long. Tail 120/* long.
Vulva situated immediately anterior to anus, a distance of 52f* from
it, not salient. Vagina (fig. 2735) short, with rounded openings
into ovejector, which is a cylindrical tube about 2 mm. long, com-
posed of vestibule and sphincter which together measure 600[*. and
a trompe measuring 1.2 mm. long. Eggs 32/* long by 18[* wide, with
thick shells, embryonated when oviposited.

Life history. — Unknown.

Distribution. — Europe (Austria (Museum, Vienna)) and Africa
(Mascara and Setif).

Seurat has stated that Spiroptera turdi (p. 206) should probably
be transferred to this srenus.

Genus SPIROCERCA Railliet and Henry, 1911b

Generic diagnosis. — Spirurinae (p. 164) : Mouth hexagonal, with 6
teeth projecting into its cavity; a short funnel-shaped vestibule fol-

NEMATODE PARASITES OF BIRDS 203

lows. Male with caudal extremity rolled in spiral, alate, provided
with 4 pairs of preanal papillae and 2 pairs of postanal papillae.
Spicules very unequal. Female with vulva situated far anterior, at
about the anterior 1/20 of body. Eggs cylindrical, thick-shelled,
embryonated at time oviposited.

Parasitic in carnivores, principally in the esophagus and stomach.

Type-species. — Spirocerca sanguinolenta (Rudolphi, 1819) Seurat,
1913.

SPIROCERCA SANGUINOLENTA (Rudolphi, 1819) Seurat. 1913

Synonym. — Spiroptera sanguinolenta Rudolphi, 1819.

Third stage larvae reported by Seurat (19165) as encysted in the
walls of digestive tract and in the mesenteries of the following birds
as accidental hosts: Athene noctua glaux, Corvus corax tingitanus,
Gallus gallus, Lanius excubitor dodsoni, Passer domestica tingitanw,
Upupa epops.

Subfamily Gongyloneminae Hall, 1916

Subfamily diagnosis. — Spiruridae (p. 163) : Body long and filiform,
the anterior cervical portion ornamented with cuticular bosses. In
the median lines, immediately behind the mouth, are 2 semilunar
depressions simulating suckers. The vulva is situated a short dis-
tance anterior to the anus.

Type genus. — Gongylonema Molin, 1857.

Genus GONGYLONEMA Molin, 1857

Synonyms. — Filaria Mueller, 1787, part; Spiroptera Rudolphi,
1819, part; Myzomimus Stiles, 1892.

Generic diagnosis. — Gongyloneminae (p. 203) : Body filiform,
slightly attenuated toward the extremities. Cuticula transversely
striated. Mouth small, elongated dorso-ventrally, and surrounded
by 6 small papillae. The anterior cervical portion of body provided
with longitudinal rows of cuticular bosses. Immediately behind the
mouth are dorsal and ventral semilunar depressions simulating suck-
ers. Esophagus consisting of a slender anterior portion and a thick-
er, muscular posterior portion. Male tail curved ventrally and pro-
vided with 2 asymmetrical alae supported by elongated claviform
papillae, mostly arranged in pairs. Vulva a short distance anterior
to anus. Eggs ellipsoidal, containing well developed embryos when
deposited.

Type species. — Oongylonema minimum Molin, 1857 (synonym G.
musculi (Rudolphi, 1819) Neumann, 1894. Filaria musculi Ru-
dolphi, 1819, is regarded by Railliet as a nomen nudum).

204 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

GONGYLONEMA INGLUVICOLA Ransom 1904b

See Addenda, p. 390, for Gongylonema ingluvicola (?) of Smit
and Notosoediro, 1926.

Host. — Primary: Gallus gallus; secondary: Unknown, probably
coprophagus beetles or other coprophagus arthropods eaten by
chickens.

Location. — Sewn in the mucous lining of the crop and, occasionally,
the undilated portion of the esophagus.

Morphology. — Gongylonema (p. 203) : White or yellowish worms.
Cuticle annulated. At the anterior end of the body (fig. 274a) is
a zone of shield-like markings, few and scattered near the head and
numerous and arranged in longitudinal rows farther back. Around
the excretory pore is a large plate-like shield. On each side of the
anterior end of the body is a narrow lateral membrane, which grad-
ually becomes narrower posteriorly and disappears. The mouth is
small and surrounded by a chitinous collar. The anterior portion

Pig. 274. — Gongylonemna ingluvicola. a, Head ; b, male tail. After Ransom, 1904

of the esophagus is short, slender, and colorless, and the posterior
portion is long, thicker, and yellow.

M ale 17 to 20 mm. long by 224 to 250/* wide. Cuiicular bosses ex-
tend for distance of 575 to 680/* from head end. The tail (fig. 2746)
has two narrow caudal alae, the right 500 to 736/* long and the
left 600 to 800/* long. Genital papillae variable in number and
asymmetrical; preanal papillae are 5 to 7 on the left side (Wharton
finds 2 to 7) and 4 to 5 on the right (Wharton finds to 7) ; post-
anal papillae are 3 to 4 on the left side (Wharton finds 2 to 5) and
4 on the right (Wharton finds 3 to 5). The cloacal aperture 225
to 275/* from the tip of the tail. Left spicule as long as the body
and 7 to 9/* wide, with a barbed point ; right spicule 100 to 120/* long
by 15 to 20/* wide.

Female 32 to 55 mm. long by 320 to 490/* wide. Cuticular bosses
extend for distance of 1.3 to 2.6 mm. from head end. Anus 165 to
288/* from tip of tail. Vulva 2.5 to 3.5 mm. from tip of tail. Vagina
11 to 14 mm. long. Eggs 50 to 57/* long by 36 to 38/* wide, the shell
4/* thick, and containing embryos when deposited.

Embryo 160/* long by 8/* wide, and provided with a spine at the
anterior end.

NEMATODE PARASITES OF BIRDS 205

Life history. — Unknown; probably similar in a general way to
that of G. scutatum. the eggs passing in the droppings and being
eaten by some coprophagns arthropods, probably beetles, the worms
developing to larvae encysted in the intermediate host and the pri-
mary host becoming infected by eating these infected secondary
hosts.

Distribution. — North America (United States (collected at Wash-
ington, D. C, in a chicken from Florida and subsequently collected
in chickens bought at the Washington market and probably originat-
ing either in the District of Columbia, Maryland, or Virginia)),
Asia (Philippines, Formosa), Europe (Roumania) and Australia.

Subfamily Uncertain

The following species are left provisionally in the old and invalid
genus Spiroptera. In certain cases there are some indications show-
ing that some species may belong in valid and established genera,
but the evidence does not appear to be sufficient to warrant a change
in the name at this time with the possibility that further study may
develop evidence not in harmony with the available evidence.

Genus SPIROPTERA Rudolphi, 1819

Generic diagnosis. — Spiruridae (p. 163) : This genus is used here
only to avoid a change in existing names where the change could
serve no useful purpose. Stiles has recently suggested that old and
invalid generic names be used for species which can not be accurately
located generically. While this policy is open to debate as regards
the present day description of new species, it is a good policy to fol-
low as regards species now in these old genera. The status of Spirop-
tera is discussed elsewhere. For the purpose of discussing the follow-
ing species it is only necessary to characterize it as a broad generic
group covering worms having spirurid characters and not capable of
definite assignment to modern and valid genera. For this purpose
no type species is necessary.

SPIROPTERA PENIHAMATA Molin, 1860b

Synonyms. — Spiroptera strigis-atricapillae Molin, 1860; Spirop-
tera strigis-flammeae Molin, I860; Spiroptera strigis-griseatae Molin,
1860; Spiroptera strigis-lineatae Molin, 1860.

Hosts. — Primary : Strix atricapilla, S. flammea, S. griseata, and
S. albomarginata; secondary: Unknown.

Location. — Between the tunics of the gizzard.

Morphology. — Spiroptera (p 205) : Head continuous with body,
not alate. Anterior extremity attenuate. According to Drasche, the
mouth has 2 lateral lips as in Spiroptera bullosa or S. semilimaris.

206

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 5 to 13 mm. long by 100 to 300/* wide. Caudal extremity
twisted in one spiral turn and with wide alae. Drasche states that
the tail has 2 pairs of preanal papillae and has 2 postanal papillae
but his figure (fig. 275) shows 4 pairs of preanal and 2 pairs of post-
anal; the right spicule is short and thick, and the left is long and
alate with its tip barbed like a fishhook.

Female 7 to 15 mm. long by 300/w wide. Anus near caudal ex-
tremity. Vulva in posterior part of body.

Life history. — Unknown ; probably involves intermediate stages
in some other hosts.

Distribution. — South America (Brazil).

Seurat (1915) has suggested that this species is rather close to
Hartertia obesa. It is possible that this species should be transferred
to the genus Hartertia, but in default of further evidence the matter
is left as it is without decision.

275.

Figs. 275-276. — 275, Spiroptera penihamata. Male tail. After Drasche, 1884.
276, Spiroptera turdi. Head, a, Front ; b, lateral view. After Drasche,
1884.

SPIROPTERA TURDI Molin, 1860b

/Synonyms. — Ascaris fissilabium Linstow, 1873; Filaria turdi
(Molin, i860) Linstow, 1877&.

Hosts. — Primary : Turdus musicus, T . iliacus, T . pilaris, T. merula,
Sturnus vulgaris, and Crocidura leucodon; secondary: Unknown.

Location. — In walls or between tunics of gizzard.

Morphology. — Spiroptera (p. 205) : According to Drasche, the
mouth (fig. 276 a and b) is without lips, its aperture circular, and sur-
rounded by a ring-shaped five-cornered ornamentation, with 2 large
lateral papillae and 4 submedian papillae behind this ormentation.

Male unknown.

Female 4.5 mm. long

Life history. — Unknown ; probably involves intermediate stages in
other hosts. Cori has described what he regards as larvae of this
species from the earthworm.

Distribution. — Europe.

NEMATODE PARASITES OF BIRDS 207

Seurat (1913&) suggests that probably this species should be put
in the genus Viguiera. The evidence from the available descriptions
and figures does not seem sufficiently complete to warrant the present
writer in assigning this species to a genus other than the provisional
genus in which it is left here.

SPIROPTERA SAG1NATA (Rudolphi, 1819) Dujardin. 1S15

Synonym. — Physaloptera saginata Rudolphi, 1819.

Hosts. — Primary: Caprimulgus guianensis, C. leucopygeus,Corcus
cajanus, C rotophaga and, Cuculus tinguaeu, FoJco fwcatus, Icterus
crlstatus, Strix atricapUla, S. torquata and Thamnophilus funebris;
secondary : Unknown.

Location. — Intestine.

Morphology. — Spiroptera (p. 205) : Brownish worms with the ex-
tremities somewhat attenuated, white and translucid. Mouth a
rounded oval, widened transversely. Cuticula strongly striated
transversely.

Male unknown.

Female 34 to 45 mm. long, according to Dujardin, or up to 80
mm. long, according to Molin, by 1 to 2.25 mm. wide, the body
curled in a loose spiral. The anus is remote from the caudal ex-
tremity and the tail ends in a bluntly rounded termination.

Life history. — Unknown; probably involves intermediate stages
in some other hosts.

Distribution. — South America (Brazil).

This species has been shifted back and forth between Physaloptera
and Spiroptera. Rudolphi made it Physaloptera, Dujardin Spirop-
tera,, Stossich restored it to Physaloptera, and Qrtlepp (1922) has
recently stated that he has examined specimens of the species and
that it belongs in Spiroptera. As Ortlepp was monographing the
genus Phi/x<il<>ptera, it seems reasonably sure that the species does not
belong in ftlvysdloptera. Its assignment by Ortlepp to Spiroptera
can only be taken to mean that it is a spirurid which is not Phy-
saloptera. Large spirurid worms in the intestine of birds are sug-
gestive of such forms as Hartertia gallinarum, but in default of
further evidence it seems advisable to leave S. saginata in Spiroptera
pending further study of actual specimens by some worker.

SPIROPTERA TENUICOLLIS (Rudolphi, 1819) Dujardin, 1815

Synonym. — Physaloptera tenuicollis Rudolphi, 1819.
LLont*. — Primary: Falco haliaetuSf secondary: Unknown.
Location. — Intestine.
3012—27 15

208 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Spiroptera (p. 205) : Mouth orbicular.

Male unknown.

Female 41 mm. long by 1.15 mm. wide. Anterior extremity of the
body sharply narrowed for a distance of 2.25 mm., a condition
attributed by Dujardin to contraction in alcohol. Tail acute.

Life history. — Unknown.

Distribution. — Europe (Austria) (Vienna)).

The description is inadequate for a transfer of this species to any
other genus. Diesing (1851) states that it is more like an ascarid
than like a Physaloptera. Ortlepp (1922) includes it in his list
of species wrongly attributed to Physaloptera. The description was
based on a single specimen.

Family DESMIDOCERCIDAE, new family

Family diagnosis. — Spiruroidea (p. 162) : Forms with distinctive
larval characters maintained into adult life; both male and female
may be provided at tail end with a cluster of filiform papillae or
modifications of the same, the male not provided with caudal alae
or caudal papillae of usual type. Mouth with 2 lips, each lip
with 4 submedian papillae. Esophagus divided into distinct anterior
and posterior portions.

Parasitic in respiratory tract of birds.

Type genus. — Desmidocerca Skrjabin, 1916.

When Skrjabin made a new genus, Desmidocerca, for his very in-
teresting new forms he did not place it except as in the suborder
Spirurata. The very distinctive characters of the spirurid larva
maintained throughout life, as shown in both the male and female
of his type-species, do not fit into any of the families as previously
made in the Spiruroidea. The present writer has therefore made a
new family for the genus. These forms are perhaps the most prim-
itive of the spirurids, their larval characters, indicative of their
ancestry, not having been lost in adult life, as in the other groups.

Subfamily Desmidocercinae, new subfamily

Subfamily diagnosis. — Characters of the family.

Genus DESMIDOCERCA Skrjabin, 1916b

Generic diagnosis. — Desmidocercinae (p. 208) : Very small nema-
todes. Head with 2 projecting lips and, posterior to each lip, 4
submedian papillae. Esophagus cylindrical, consisting of 2 por-
tions. Caudal extremity in both male and female of type species
provided with a clump of filiform papillae; in the second species
of the genus these are modified to lateral knobs in the female and

NEMATODE PARASITES OF BIRDS 209

;ire lacking in the male but a pair of subterminal papillae are
present on the ventral surface. Male with tail recurved hook-like
and obtusely rounded. Two filiform spicules of unequal size.
Female with the position of vulva variable (in posterior part of
body in type-species).

Parasitic in the respiratory tract of aquatic birds.

Type species. — Desmidocerca aerophila Skrjabin, 1916b.

KEY TO SPECIES OF DESMIDOCERCA

Tail of both male and female provided with terminal clump of filiform papillae;
esophagus 2/3 of body length ; vulva in posterior part of body.

Desmidocerca aerophila, p. 209.

End of male tail bare, of female tail with 2 small knobs on each side ; esophagus
only 1/8 to 1/9 of body length ; vulva in anterior part of body.

Desmidocerca numidica, p. 209.

DESMIDOCERCA AEROPHILA Skrjabin. 1916

Hosts. — Primary: Ardea cinerea, and Phalacrocorax carbo; sec-
ondary : Unknown.

Location. — Air sacs.

Morphology. — Desmidocerca (p. 208) : Cuticle with fine transverse
striations. Head (fig. 277a) with 2 lips projecting forward, and
behind them to each side 2 submedian papillae. A small cylindrical
buccal cavity present. Esophagus very long, 2/3 the total body
length.

Male 3.9 mm. long by 260/x wide (fig. 277&). Caudal extremity
(fig. 277c) recurved hook-like, bearing at its tip a cluster of spine-
like papillae. Spicules 660/* and 270/* long, curved in accordance
with the shape of the caudal extremity. No caudal papillae ob-
served on ventral surface of tail.

Female 4.4. mm. long by 300/* wide. Caudal extremity (fig. 277d)
obtusely rounded, bearing a cluster of spine-like papillae. Vulva
in posterior region of body. Eggs 30/* long by 18/* wide.

Life history. — Unknown.

Distribution. — Asia (Russian Turkestan).

DESMIDOCERCA NUMIDICA Seurat, 1920

Host. — Primary : Heron cendre; secondary : Unknown.

L.ocation. — On inner surface of air sacs.

Morphology. — Desmidocerca. (p. 208) : Body thick-set, slightly
curved. Two lateral hyaline alae; lateral fields large, clearly delim-
ited. Cervical papillae symmetrical, situated at level of nerve ring.
Mouth with 2 lateral trilobed lips, each bearing 2 pairs of large
papillae, the most internal situated near base of interior. Buccal

210

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

cavity short; esophagus 1/8 to 1/9 the total body length, clearly
divided into a colorless muscular region, encircled by a large nerve
ring, and a glandular region.

Male 5.3 mm. long; posterior region curved crosswise. Tail short
(80/*), massive, digitiform, rounded, and bare at its extremity; no
caudal alae. Posterior lip of cloacal aperture salient,, with a pair
of postanal papillae directly posterior to it. Spicules very unequal,
480 and 155/x long.

Female 6.8 mm. long. Tail short (75^), massive, digitiform,
provided at each side with 2 small lateral knobs or buttons. Vulva
with anterior lip prominent, situated directly anterior to end of

Fig. 277.-

PlOSMIDOCERA AEROPIULA. a, HEAD ; b, MALE; C, MALE TAIL; d,
FEMALE TAIL. AFTER S'kRJAB£N, 1916

esophagus ; ovejector rectilinear, directed posteriorly. Uteri parallel
for almost 1/2 their length, then divergent. No mature eggs.

Life history. — Unknown.

Distribution. — Africa ( Algeria ) .

Family ACUARIIDAE Seurat, 1913a

Family diagnosis. — Spiruroidea (p. 162) : Lips triangular, not
expanded. Anterior extremity of body with ornamentations on the
cuticle. Esophagus long, its anterior part differentiated into a
pharynx. Male with caudal alae.

Parasitic in the digestive tract of birds.

Type- genus. — Acuaria Bremser, 1811.

NEMATODE PARASITES OP BIRDS 211

KEY TO SUBFAMILIES OF ACUARIIDAB

1. Cephalic ornamentation consisting of cordons, epaulets or other homologous
structures; mouth with 2 simple lateral lips; usually 4 or 5 pairs of preanal

papillae Acuariinae, p. 211.

Cephalic ornamentations consisting of appendices or festoons; mouth with
2, 4 or 6 lips; preanal papillae variable in number, sometime"' very
numerous Schistorophinae, p. 284.

Subfamily Acuariinae Railliet, Henry and Sisoff, 1912

Subfamily diagnosis. — Acuariidae (p. 210) : Nematodes with cor-
dons, epaulets, or other homologous ornaments at the anterior
extremity. Mouth with 2 simple lateral lips, followed by a vestibule
or pharynx with its wall usually transversely striated, and an
esophagus differentiated into 2 parts. Male with caudal extremity
provided with 2 lateral alae sustained usually by 4 to 6 pairs of
preanal and by a variable number of postanal papillae. Eggs ellip-
soidal and with thick shell, embryonated at time of oviposition.

Parasitic in dicrestn'e tract of birds.

Type-genus. — Acuaria Bremser, 1811.

KEY TO GENERA OF ACUARIINAE

1. Ornamentation of the anterior end in the form of cordons extending post-

teriorly from the head along the cervical region, usually in the sub-
median fields (Acuarui, s.l., p. 212) 2.

Ornamentation of a nature different from the above, confined to cephalic
region 8.

2. Cordons not recurrent, not anastomosing 3.

Cordons recurrent or anastomosing, or both 4.

3. Both spicules thick and only slightly unequal; 6 to 8 pairs of postanal

papillae Acuaria, p. 216.

Spicules very dissimilar and very unequal; 5 to 7 pairs of postanal
papillae Cheilospirura, p. 'J20.

4. Cordons not recurrent, but anastomosing 5.

Cordons recurrent, anastomosing or separate C.

5. Cuticle raised in front of postcervical papillae to form a large collar or

Sheath; cordons anastomose on the free posterior border of the collar.

Chevreuxia, p. 231.
Xo such collar or sheath present Echinuria, p. 244.

6. Cordons recurrent hut not anastomosing Dispharynx. p. 237.

Cordons recurrent and anastomosing— 7.

7. CprdoilS form a loop directly after their origin on the head; cordons not

tl.it against body hut applied to margin of plates or alae; lateral alae

present on body Cosmocephalus, p. 233.

Cordons with loops lacking at their anterior ends; cordons applied directly
to the body; no lateral alae Synhimantus, p. 272.

8. Cephalic ornamentation consists of a narrow denticulated collar which is

the margin of a depression surrounding the head at the base of the lips;
cervical papillae large and crescent-shaped, with numerous teeth.

Streptocara, p. 2G4.

212 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Cephalic ornamentation not in form of collar, but of epaulets or alate

appendages; cervical papillae not as above 9.

9. Cephalic ornamentation consisting of 4 delicate membranous alae directed

posteriorly Sciadiocara, p. 260.

Cephalic ornamentation consisting of 2 crescent-shaped epaulets 10.

10. Free edge of epaulets markedly dentate; precervical papillae enormous,
tricuspid; no lateral alae on body, but in their place 2 double rows of

posteriorly directed hooks Seuratia, p. 262.

Free edge of epaulets not dentate; precervical papillae small, simple, in-
serted in the 2 lateral alae of the body ; no rows of hooks on body.

Rusguniella, p. 258.

Genus ACUARIA Bremser, 1811, sensu lato

Synonyms. — Spiroptera Rudolphi, 1819, part; Anthuris Rudolphi,
1819; Dispharagus Dujardin, 1845.

Generic diagnosis. — Acuariinae (p. 211) : Anterior extremity with-
out vesicular swelling, but bearing 4 cutaneous cordons, salient or
depressed, extending at times directly posteriorly, more often folded
anteriorly, and sometimes uniting in pairs on the lateral surfaces.

Parasitic in esophagus, glandular stomach or gizzard of birds.

Type-species. — Acuaria anthuris (Rudolphi, 1819) Railliet, Henry,
and Sisoff, 1912.

As used in this paper, Acuaria sensu lato is equivalent to its gen-
eric rank as accorded by Railliet, Henry, and Sisoff; the subgenus
Acuaria of these writers is given generic rank in this paper and
regarded as Acuaria sensu stricto. Only those forms which can
not be definitely referred to the subgenera of Railliet, Henry, and
Sisoff, here accorded generic status, are left in Acuaria sensu lato
and covered in the following key. Other species should be traced
through the key to genera of Acuariinae.

K.KY TO SPECIES OF ACUARIA SEXSU LATO

1. Cordons recurrent Acuaria quadriloba, p. 215.

Cordons not recurrent 2.

2. Only larva known ; genitalia unknown ; cordons 145/* long.

Acuaria tarentolae, p. 216.

Adults known ; cordons, where known, more than 145m long ; males

unknown 3.

3. Cordons extend beyond esophagus ; female 21 mm. long.

Acuaria mamillaris, p. 213.

Cordons do not extend beyond esophagus, or, if they do, then female not

over 12.73 mm. long 4.

4. Cordons extend beyond esophagus ; female 8.84 to 12.73 mm. long.

Acuaria ptilopachydis, p. 214.
Cordons do not extend beyond esophagus 5.

5. Lips unequal ; cordons end 330/x from head ; total length of esophagus less

than 1/10 of body length Acuaria muscicapae, p. 214.

Lips apparently equal ; cordons end 800m from head ; total length of esoph-
agus approximately 2/5 of body length Acuaria macrolaima, p. 213.

NEMATODE PARASITES OF BIRDS 213

ACUARIA MACROLAIMA (Linstow, 1906) Railliet. Henry, and Sisoff, 1912

Synonym. — Dispharagus macrolaimus Linstow, 1906a.

Hosts. — Primary : Plotus melanogaster ; secondary : Unknown.

Location. — Stomach ( Gizzard ? ) .

Morphology. — Acuaria sensu lato (p. 212) : Head with 2 small,
conical, rounded lips. Following the mouth is a long vestibule and
then a very long esophagus with a total length of 10/24 to 10/27 of the
body length. Cuticle transversely striated. Cordons extend 800/u.
posteriorly, rather beyond the anterior portion of the esophagus ; not
anastomosing or recurrent. Cone-shaped papillae in lateral fields
immediately anterior to ends of cordons.

Male unknown.

Female 7 to 11 mm. long by 280 to 470/x wide. Tail 1/37 of body
length, terminating in a smaller finger-shaped point. Eggs 31/i
long by 11/u. wide.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Asia (Ceylon (Wirawila)).

This species belongs in Acuaria sensu stricto or G heilosjnr-ura,
apparently, but can not be definitely placed until the male char-
acters are known.

ACUARIA MAMILLARIS (Molin, I860) Railliet, Henry, and Sisoff, 1912

Synonyms. — Dispharagus mcumillaris Molin, 1860c/ Spiroptera
corvi-cajani 3 in Molin, 1860c.

Hosts. — Primary: Corvus cajanus; secondary: Unknown.

Location. — Stomach.

Morphology. — Acuaria sensu lato (p. 212) : Mouth with 2 large
triangular lateral lips. Body densely striated transversely and at-
tenuated anteriorly. Cordons directed posteriorly and gradually
disappearing, not recurrent or anastomosing. According to Gendre,
(1920a), the cordons extend beyond the posterior extremity of the
esophagus and ventricle (the ventricle apparently being the second
part of the esophagus).

Male unknown.

Female 21 mm. long. Posterior extremity conical, with obtuse
extremity.

Life history. — Unknown.

Distribution. — South America (Brazil).

This species belongs in Acuaria (sensu stricto) or Cheilospirura,
apparently, but can not be definitely placed until the male char-
acters are known.

3 Catalojnie of the Vienna Museum.

214 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

ACUARIA MUSCICAPAE (Linstow, 1878) Railliet, Henry, and Sisoff, 1912

Synonyms. — FUaria muscicapae Linstow, 1878; Disphwragus
muscicapae (Linstow, 1878) Stossich, 1891.

Hosts. — Primary : Museicapa atricapilla; secondary : Unknown.

Location. — Stomach (gizzard).

Morphology. — Acuaria sensu lato (p. 212) : Head with 2 conical
lips, of which, according to Gendre, one is smaller than the other.
(Linstow, also quoted without comment by Gendre writes that the
ventral lip is the smaller, but as the lips in spirurids are lateral,
there appears to be some error in this statement.) The cordons ex-
tend 330/* posteriorly from the head, in the female, and hence not
beyond the limits of the esophagus.

Male unknown.

Female 11.5 mm. long by 150/i wide. Mouth followed by a
vestibule 160//, long, and this by an esophagus of which the glandular
portion is 750/x long and the muscular portion 330/x long. The tail
is 130/x long, conical, with a rounded end. Vulva somewhat pos-
terior to middle of body, dividing body in ratio of 7:6. Eggs 29/u,
long by 16//, wide.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Prussia (Hanover)).

This species belongs in Acuaria sensu stricto or Cheilospirura, ap-
parently, but can not be definitely placed until the male characters
are known.

ACUARIA PTILOPACHYDIS Gendre. 1920a

Hosts. — Primary : Ptilopachys fuscus; secondary : Unknown.

L ocation. — Gi zz ar d .

Morphology. — Acuaria sensu lato (p. 212) : Body colorless, filiform
and slender. Cuticle transversely striated. Mouth with 2 large
lateral lips, each bearing on its external surface, near the base, 2
symmetrical papillae, and internally a small median triangular
lobe. Four cordons (fig. 278«), hidden in cuticle, not recurrent or
anastomosing, extend in submedian lines to a length of 2.62 to 3.13
mm., hence beyond the posterior end of esophagus (the esophagus
and ventricle as termed by Gendre). Lateral papillae slightly pos-
terior to anterior end of esophagus.

Male unknown.

Female 8.84 to 12.73 mm. long by 170 to 200> wide. Pharynx 110
to 200^ long. Esophagus 400 to 550/a long; ventricle (evidently
second part of esophagus) 1.07 to 1.43 mm. long. Tail (fig. 2786)
200 to 245/* long, conical, digitiform, rounded at apex. Vulva slightly
salient, bordered with a cuticular margin and slightly posterior to

NEMATODE PARASITES OF BIRDS

215

middle of body, apparently 10/19 of body length from head end.
Ovejector 260/u, long by 46/x, wide, directed posteriorly and without
a distinct vestibule. Eggs thick-shelled, 35 to 39ju long by 21 to 23ju,
wide.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa (Dahomey) .

This species belongs in Acuaria sensu stricto or Cheilospimra, ap-
parently, but can not be definitely placed until the male characters
are known.

ACUARIA QUADRILOBA (Rudolphi, 1819) Railliet. Henry, and Sisoff. 1912

/Synonyms. — Spiroptera quad/ % Uoba Rudolphi, 1819; FUaria quad-
riloba (Rudolphi, 1819) Schneider, 1866; Dispharagus quadrUobus
(Rudolphi, 1819) Dujardin, 1845; Dispharagus crassus Molin, 1860c.

Hosts. — Primary: Dryocopus martini, Picus martins, P. viridis,
Hylomotus pileatus, Gecinus viridis; secondary : Unknown.

Location. — Esophagus.

Z79.

Figs. 278-279. — 278, Acuaria ptilopachydis. a, Anterior end ; b, female tail.
After Gbndre, 1920. 279, Acuaria quadriloha. Head, front view. After
Schneider, 1866

Morphology. — Acuaria sensu lato (p. 212) : Body somewhat thick,
more slender anteriorly. Cordons strongly developed with recurrent
loops extending almost to the head end; whether anastomosing or
not is not stated. Mouth (fig. 279) with 2 thick, papilliform lips
and surrounded by small, slightly salient papillae.

Male unknown.

Female 9 to 11 mm. long by 400/* wide. Caudal extremity taper-
ing quite suddenly; apex obtuse. Anus near caudal apex. Vulva
in posterior part of body, three-fourths of distance from head.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Germany (Vienna Museum and Berlin))
and North America (Philadelphia, Pa.).

3612— 27-

-16

216 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

This species belongs in Dispharynx or Synhimantus, apparently,
but can not be definitely placed until the male characters are known

ACUARIA TARENTOLAE Seurat, 1916f

Tarentola

Hosts. — Primary: Unknown; secondary (aberrant]
mauritanica, and unknown usual secondary hosts.

Location. — In stomach.

Morphology— Acuaria sensu lato (p. 212) : Mouth (fig. 280) with
2 lateral lips, each provided with a conical tooth and a pair
of large lateral papillae situated near the origin of the cordons.
Cervical papillae situated at the level of the posterior edge of the
nerve ring, 175/* from the anterior extremity. Cordons not anasto-
mosing or recurrent.

Male unknown.

Female unknown.

Fig.

280. — Acuaria tap.entolae. Anterior and posterior ends op third stage
larva. After Seurat, 1916

Larva, third-stage, 5.35 mm. long by 120/t wide. Cordons 145/t
long. Tail (fig. 280) 156^ long, ending in a small button with a
smooth surface. Buccal cavity 112^, muscular esophagus 504ju, entire
esophagus 2.2 mm. long.

Life history. — Unknown; probably involves true intermediate
hosts other than lizards and true final hosts which are birds.

Distribution. — Africa (Kouba) .

This species belongs in Acuaria sensu stricto or Cheilospirura,
apparently, but can not be definitely placed until the adult male
characters are known. It is assumed here that a worm in this genus,
Acuaria sensu lato, is probably a parasite of birds when it is
adult and that it is an aberrant parasite of the lizard. This phe-
nomenon of aberrant parasitism on the part of third-stage larvae
of spirurids appears to be fairly common, according to the findings
of Seurat.

Genus ACUARIA Bremser, 1811, sensu stricto

Synomjms.—Spiroptera Rudolphi, 1819 in part; Anthuris Ru-
dolphi, 1819; Dispharagus Dujardin, 1845 in part.

Generic diagnosis.— Acuariinae (p. 211) : Cordons directed pos-
teriorly, not anastomosing or recurrent. Male with 2 short, thick,
slightly unequal spicules and 6 to 8 pairs of postanal papillae.

NEMATODE PARASITES OF BIRDS 217

Parasitic between the tunics of the gizzard or in proventriculus

of birds.

Type species. — Acuaria an t hurts (Rudolphi, 1819) Railliet, Henry
and Sisoff, 1912.

As used here, Acuaria sensu stricto is the equivalent of the sub-
genus Acuaria as defined by Railliet, Henry, and Sisoff, this sub-
genus being here accorded generic rank. Forms which can not be
definitely referred to Acuaria (sensu stricto) or to the other groups
regarded by Railliet, Henry, and Sisoff as subgenera, but here re-
garded as genera, are referred to Acuaria sensu lato (p. 212) in
this paper.

The genus Acuaria has had a complicated and eventful history,
which has been critically discussed in detail by Stiles and Hassall
(1905). Their discussion need not be repeated, but may be briefly
summarized as follows: Bremser proposed Acuaria without a type
species and without naming the included species, but did name the
hosts in which 14 species, supposed by him to belong in this genus,
occurred. Of these Spiroptera antkuris Rudolphi, 1819, is regarded
as type of the genus Antkuris Rudolphi, 1819, by absolute tautonomy
and by Rudolphi's original intentions. But Antkuris is Acuaria
Bremser. 1811, renamed, as Rudolphi admits, and hence the type
species antkuris is type of Acuaria. But since Spiroptera is a
renaming of Acuaria and Antkuris, as Rudolphi admits, it has the
same type, and in default of a valid reason for the renaming falls
into synonomy. There are other features which complicate the case
somewhat, but as these details are carefully discussed by Stiles and
Hassall it is unnecessary to recapitulate them. Dispharagus Du-
jardin, 1845, is a deliberate renaming of a group of species for
which 2 generic names were in existence; it included the type of
Acuaria, and has been dropped into synonomy accordingly.

KEY TO SPECIES OF ACUAKIA SENSU STRICTO

1. Body with 6 cordons, the 4 submedian being double, festooned, and extending

10/23 <d' total body length, the 2 lateral being single and simpler, and ex-
tending the whole body length: spicules 280 and 2." ( () M long.

Acuaria ornata, p. 223.

Body witb 4 cordons, submedian; spicule lengths, where known, differ

from above 2.

2. Male with 9 pairs of caudal papillae; female 22 mm. long.

Acuaria depressa, p. 221.

Male with 10 to 12 pairs of" caudal papillae, where known: if number is not

known, female 18 mm. long (A. attenuata and .1. tenuis) 3.

3. Male 7 mm. long, not otherwise described ; anus of female 160/u from posterior

end; eggs said to be only l5/i long; from Hirundo, species.

Acuaria attenuata, p. 220.

If only male known (,l. paptilifera), is 4.7 mm. long; anus of female 22.V

or more from posterior end. except in A. gracilis; eggs 32 to 4N,u long;

■ • ' :n Hirundo, species _ 4.

218 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

4. Cordons extend 1/4 to more than 1/3 of body length (reported as 8 to 9 mm.

in female) ; spicules at least 220 and 180/* long, may be 287 and 234/* long.

Acuaria anthuris, p. 218.

Cordons, if described, extend for length of not more than GOO/* ; if length

not described (A. tenuis), spicules only 133 and 95/* long 5.

5. Spicules 133 and 95/* long ; cloacal aperture 135/* from posterior extremity ;

from Saxicola rubetra Acuaria tenuis, p. 225.

Spicules longer than above (may be as short as 150 and 98/* long in A. pap-
illifera) ; cloacal aperture 150 to 400m from posterior extremity; not from
Saxicola rubetra 6.

6. Male with 10 pairs of caudal papillae 7.

Male with 12 pairs of caudal papillae 8.

7. Vulva at middle or slightly in front of middle of body ; eggs 32/* long.

Acuaria gracilis, p. 222.
Vulva posterior to middle of body ; eggs 37 to 39/* long.

Acuaria subula, p. 224.

8. Only male known ; 4.7 mm. long ; cordons 190/* long ; cloacal aperture 168/*

from posterior extremity Acuaria papillifera, p. 224.

Both sexes known ; male 10 to 11 mm. long ; cordons of male 280 to 320/*
long; cloacal aperture 300 to 400/* from posterior extremity.

Acuaria cordata, p. 220.

ACUARIA ANTHURIS (Rudolphi, 1819) Railliet, Henry, and Sisoff, 1912

Synonyms. — Spiroptera anthuris Rudolphi, 1819; Dispharagus
anthuris Dujardin, 1845; FUaria anthuris (Rudolphi, 1819)
Schneider, 1866.

Hosts. — Primary: Corvus glandamus, C. cornice, C. frugilegus, C.
corone; secondary : Unknown.

Location. — Proventriculus.

Morphology. — Acuaria sensu stricto (p. 216) : Lateral lips of
mouth strongly developed. Cordons unite at the margin of the
mouth and extend posteriorly 25 to 36 per cent of the body length,
according to Linstow (1873), or in the female 8 to 9 mm. posteriorly,
according to Schneider; they do not anastomose, according to
Schneider (1866) and Linstow (1873); later Linstow (1901), in
describing his Dispharagus invagin&Pw, writes that the cordons
anastomose and are recurrent as in A. anthuris; Railliet, Henry and
Sisoff define Acuaria as a subgenus, type A. anthuris, as with cordons
not anastomosing and not recurrent, so it may be assumed that this
is the case. The margins of the cordons lie in the cuticula.

Male 10 mm. long. Caudal papillae total 10 pairs, with 6 pairs
postanal, according to Schneider (fig. 281), or total 12 pairs, accord-
ing to Linstow (fig. 282a); Schneider finds the relative position of
the fourth, fifth, and sixth pairs from the posterior end to be vari-
able, and the seventh and ninth pairs to be indistinct. The bladder-
like thickening of the cuticle which forms the caudal alae is again
divided longitudinally by a membranous wall which is cross-striated.
Spicules 220 and 180/* long (Linstow, 1873; lie says 22 and 18 mm.,

NEMATODE PARASTTES OF BIRDS

219

but his illustration and magnification show this an error) to 287
and 234/x long (Linstow, 1895), the right 34 and the left 31t* wide,
according to Linstow, or both M/x wide, according to Mueller (1897).
Female 23 mm. long. Vulva (fig. 283) somewhat anterior to mid-
dle of body, or 10 mm. from head end, according to Schneider. Eggs
thick-shelled, elliptical, 92.6^ by 26.2,* (Linstow, 1873), or 39/* by
26ti (Linstow, 1895).

Figs. 281-282. — Acuakia axtiuris. 281, Male tail, a, Cross section ; b, ventral
view. After Schneider, 1SGG. 282, «., Male tail X about 50 ; b, female tail
end. After Linstow, 1873

Life history. — Unknown; Linstow (1895) found an encapsulated
larva in Gam-marus pulex which he regarded as possibly this species.

Distribution. — Europe.

It has unfortunately, but quite naturally, happened that many of
the older genera have poorly described and little known species as
types. This is the case with Acuaria, type A. anthuris, and this is
one reason for the complicated relationships of Acuaria, Spiroptera,
Anthuris, and Dispkaragus ; a well described and commonly known
type species aids in the definition of a genus and in the efforts to

Fig. 283. — Accaria ANTHURIS. Ovejector. After Seurat, 1915

promptly assign the species to a valid genus and retain it there.
Even yet, A. ant hurts is not well described in any work available to
the writer, and there are certain discrepancies that need clearing up
in the available descriptions.

In a discussion of the genus Acuaria sensu stricto (p. 217) the
writer has briefly summarized the somewhat complicated status of
the genus Acuaria as related to Spiroptera, Anthuris, and Disphara-
gus. A detail of that case, not discussed there, is the fact that Ru-
dolphi's Spiroptera anthuris was a composite species and as such
S. anthuris is in part a synonym of several other recognized species.

220 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Schneider (1866) reexamined Rudolphi's material and removed from
it a species from one host, Coraeim garrula, as Filaria capitellata
Schneider, 1866. Under the name of Filaria anthuris he redescribed
Rudolphi's species, restricting it to specimens from Corvus gland-
arius. Mueller (1897) has described a new species, Dispharagus
cordatus, of which A. anthuris is a synonym in part, stating that
D. cordatus is actually the species which Molin and Dujardin have
incorrectly described as A. anthuris; it showed distinct differences
from that described by Schneider and later by Linstow. In addi-
tion, A. anthuris is a synonym in part of Oxyspirura sygmoidea
Diesing, 1851.

ACUARIA ATTENUATA (Rudolphi, 1819) Railliet, Henry, and Sisoff, 1912

Synonyms- — Spiroptera attenuata Rudolphi, 1819; Dispharagus
atten/uatus (Rudolphi, 1819) Dujardin, 1845.

Hosts. — Primary : Hirundo urbica, H. rustica, H riparia; sec-
ondary : Unknown.

Location. — Between the tunics of the gizzard.

Morphology. — Acuaria sensu stricto (p. 216) : Body very slender.
Head 40/x wide. Mouth with 2 lateral papilliform lips. Cordons
thick and sinuous. Buccal cavity narrow, 170/x long. Anterior esoph-
agus 540/t long, posterior esophagus 1.45 mm. long by 52//. wide.

Male 7 mm. long. Caudal extremity twisted once or twice in a
spiral; caudal alae long and with double margin.

Female 18 mm. long by 130/* wide. Vulva with 2 large lips and
situated in posterior part of body, 10.5 mm. from the head and 7.5
mm. from the tail. Anus 160/* from tail end. Eggs only 15/* long,
according to Dujardin, who questions whether they are fully
developed.

Life history. — Unknown: probably involves intermediate stages
in other hosts.

Distribution. — Europe (Austria (Vienna Museum)).

ACUARIA CORD ATA (Mueller, 1897) Gendre, 1920

Synonyms. — Spiroptera anthuris Rudolphi, 1819 of Dujardin,
1845, and of Molin, I860.; Dispharagus cordatus Mueller, 1897.

Hosts. — Primary : Lanius rufus, L. curullio; secondary : Unknown.
Mueller regards his D. cordatus as identical with S. anthuris of
Dujardin ; the hosts for the latter are Corvus glandarius, G. pica, C.
frugilegus, C. corax, C. corone, C. comix, Goracias garrula, Oriolus
galbula, Pyrrhocorax alpinus. Other hosts listed are Gorvus caryo-
catactes, G. corax tingitanus, G. pyrrhocorax, C . americanus, Garru-
lus glandarius, Nucifraga caryocatactes, Pica caudata and Urocissa
occipitalis. Some of these hosts may be regarded as doubtful; the

NEMATODE PARASITES OF BIRDS

221

species is established on the material front Lanius i^ufus and L,
curuTKo.

Location. — In stomach wall (proventriculus?).

Morphology. — Acuaria sensu stricto (p. 216) : Head (fig. 284a)
with 2 lips, cordons joined for a distance of 20/* in the head region,
separating and passing posteriorly, gradually disappearing without
anastomosing.

Male 10 to 11 mm. long by 200 to 220/* wide. Cordons 280 to
320 long. Tail (fig. 2846 and c) recurved, enlarged and cordate,
terminating in a sort of rounded lamina 170/* wide, in front of which
are the 2 lateral alae thickened and heavily striated, the width here
being 350 to 400/*. Twelve pairs of caudal papillae, of which 8 pairs
are postanal. Cloacal aperture 300 to 400/* from tail end. Accord-
ing to Drasche, one spicule is short and thick, 270/* long by 50/* wide,

Fig. 284. — Acuaria cokdata. a, Head end ; b, lateral and c, ventral view
of male tail. Afteb Mueller, 1897

the other more curved, 210/* long, and slenderer; according to
Mueller, the right spicule is 150 to 170/* long and the left 170 to
183/* long by 17/* wide.

Female 22.5 to 40 mm. long by 200 to 320/* wide. Cordons 480 to
600/* long. Anus 270 to 400/* from tail end. Vulva slightly anterior
to middle of body (10.6 mm. from head in worm 22.3 mm. long),
according to Dujardin ; Mueller found it exactly in the middle, 20.5
mm. from head and tail ends. Eggs 43 to 48/* by 27 to 30/*, thick-
shelled.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Portoferrajo), Asia (Russian Turkestan),
Africa (Algeria), and North America (District of Columbia; re-
port by Hassall).

ACUARIA DEPRESSA (Schneider, 1866) Railliet, Henry, and Sisoff, 1912

Synonyms. — Filaria depressa Schneider, 1860; Dispharagus de-
pressus (Schneider, 1866) Gendre, 1912.

Hosts. — Primary: Corvus comix; secondary: Unknown.

222

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Location. — Not given; presumably in proventriculus, gizzard or
esophagus.

Morphology. — Acuaria sensu strict o (p. 216) : The edges of the
cordons sunk in the cuticula.

Male with broad tail (fig. 285) the margins of the caudal alae
strongly developed and the alae with distinct transverse striation.
Nine pairs of caudal papillae, 5 pairs of these postanal. Spicules
short and thick, almost equal in length.

Female 22 mm. long.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Africa ( Egypt ) .

Figs. 285-286. — 285, Acuaria dbpressa. Male tail. After Schneider, 1866.
286, acuakia gracilis. a and b, anterior end; c, left and d, right spicule;
e, male tail ; f, female tail. after gendre, 1912

ACUARIA GRACILIS (Gendre, 1912) Railliet, Henry, and Sisoff, 1912

Synonyms. — Dispharagus gracilis Gendre, 1912; Acuaria similis
Gendre, 1913< .

Hosts. — Primary : Buchanga atra assimilis and (?) Oriolus aura-
tus; secondary: Unknown. The Acuaria from O. auratus shows
certain differences from the A. gracilis from B. a. assimilis.

Location. — Between the tunics of the gizzard.

Morphology. — Acuaria sensu stricto (p. 216) : Head (fig. 286a
and b) with 2 large, conical, lateral lips, each with 2 papillae on its
external surface and with a triangular area, probably for the inser-
tion of muscles, between them. The 4 cordons, not projecting more
than the cuticular striations, extend along the submedian lines 220/*
in male and 38G> in female. Cervical papillae small, situated in

NEMATODE PARASITES OF BIRDS

223

lateral fields between cordons and slightly posterior to middle of
length.

Male 6.26 to 6.96 mm. long by 110 to 130/* wide. Tail 1/39 to 1/42
of total body length. Buccal cavity or pharynx 150 to 170/<, long;
anterior esophagus 350 to 120/*. long; posterior esophagus 770//. to
1.09 mm. long. Caudal alae (fig. 286e) with 2 concentric zones, the
inner thick and transversely striated, the outer thin and often with
longitudinal folds. Four pairs of preanal and 6 pairs of postanal
papillae. Spicules (fig. 286^ and d) 120 and 150/* long.

Female 15.58 to 22.16 mm. long by 130 to 170/* wide. Tail 1/90 to
1/103 of total body length (fig. 286/). Buccal cavity or pharynx 160
to 200/* long; anterior esophagus 530 to 660/* long; posterior esopha-

flc. 2s7. aciaria oenata. o, male tail; b, left spicule; 0, head end: d,

right spicule ; c. cordons of the lateral and f, of the submedian fields.
After Gbndeb, 1912

gus 1 to 1.32 mm. long. Vulva near middle of body, about 10/21 of
body length from head. Eggs 32/t long by 21/* wide, thick-shelled,
embryonated when oviposited.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa ( Abomey ) .

ACUARIA ORNATA (Gendre, 1912) Raillict. Henry, and Sisoff, 1912

Synonyms. — DispJiaragus ornatus Gendre, 1912.

Hosts. — Primary : Gorvus ecapulatus ; secondary : Unknown.

Location. — Between the tunics of the gizzard.

M<jrj)/iolo(fy. — Aouaria sensu stricto (p. 216) : Mouth with 2
large lateral lips (fig. 287<?), each with 2 lateral pedunculated
papillae on its external surface and with a median triangular zone
between them. Six cutaneous cordons, 4 in the submedian lines
(fig. 287/) as usual, extending from the lips almost to the middle
of the body (10/23 of body length), thus beyond the esophagus and
ventricle, these cordons double and festooned, and 2 cordons in the
lateral lines (fig. 287e) originating at the level of the anterior fifth

224

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

of the submedian ones and extending to the base of the tail, these
cordons single and simpler than the others.

Male 9.68 to 11.37 mm. long by 250 to 360/* wide. Tail 10/204 to
10/228 of total body length. Buccal cavity or pharynx 220 to 260/x
long; anterior esophagus 790 to 880/* long; posterior esophagus 2.08
to 2.68 mm. long. Caudal alae (fig. 287a) large and with 2 con-
centric zones, the internal transversely striated and the external
smooth. Four pairs of preanal and 6 pairs of postanal papillae.
Spicules (fig. 2876 and d) robust, 280 and 250/* long.

Female unknown.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution . — Africa ( Abomey ) .

Fig. 288. — Acuaria tapillifera. Head and tail ends of male. After Lin-
stow, 1878

ACUARIA PAPILLIFERA <Linstow, 1878) Railliet. Henry, and Sisoff, 1912

/Synonyms. — Filaria papillifera Linstow, 1878 ; Dispharagus papil-
liferus Stossich, 1891.

Hosts. — Primary : Sylvia palustris; secondary : Unknown.

Location. — In stomach (gizzard?) wall.

Moi^phology. — Acuaria sensu stricto (p. 216) : Cuticle trans-
versely striated. Head with 2 conical lips. Cordons (fig. 288) ex-
tend along submedian lines from lips posteriorly for 190/*. Cervical
papillae 160/*. from head end. According to Gendre, the esophagus
extends posteriorly beyond the limits of the cordons.

Male 4.7 mm. long by 70/* wide. Esophagus 1/12, tail 1/28 of body
length. Caudal alae (fig. 288) wide. Four pairs of preanal and 8
pairs of postanal papillae, very prominent. Left spicule 150/* long,
right 98/* long.

Female unknown.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Not given.

ACUARIA SUBULA (Dujardin, 1845) Railliet, Henry, and Sisoff, 1912

Synonym.— Dispharagus subula Dujardin, 1845.
Hosts.— Primary : Luscinia rubecula (Sylvia rubecula) ; second-
ary: Unknown.

Location. — Between tunics of gizzard.

NEMATODE PARASITES OF BIRDS

225

Morphology. — Acuaria sensu stricto (p. 216) : Body slightly at-
tenuated anteriorly. Mouth with 2 lateral lips, each with 2 papillae
on external surface and between the papillae a triangular area for
the insertion of the muscles of the pharynx. Cordons (fig. 289 a
and b) straight, not recurrent or anastomosing or appreciably pro-
jecting from the cuticular surface, 380/* long and not extending
posterior to the esophagus. Cervical papillae very small, between
the cordons and slightly anterior to their middle.

Male 4.89 mm. long by 120/* wide, according to Gendre, or 6.2 to
7 mm. long by 167/* wide, according to Dujardin. Buccal cavity or
pharynx 150/* long; anterior esophagus 430/* long; posterior esoph-
agus 1.58 mm. long. Caudal alae (fig. 289 c) wide and thick and
of 2 concentric parts, the inner wider and transversely striated, the
outer narrower and granular. Four pairs of preanal and 6 pairs of

Fig. 2S

-Acuaria sdbula. a and b, Head end ; c, male tail ; d, right and e,

LEFT SPICULE. AFTEE GEN-ORE, 1913

postanal papillae. Spicules (fig. 289 d and e) dissimilar, 190/* and
120/* long, respectively.

Female 18 mm. long by 290/* wide. Head 51/* wide. Buccal cavity
or pharynx 170/* long. Tail straight, conical, 225/* long. Vulva
10 mm. from tail end and 8 mm. from head end. Eggs 37 to 39/*
long.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe ( France ) .

ACUARIA TENUIS (Dujardin. 1845) Railliet, Henry, and Sisoff. 1912

Synonyms. — Dispharac/us tenuis Dujardin, 1845.

Hosts. — Primary : Saxicola rubetra; secondary : Unknown.

Location. — Between the tunics of gizzard.

Morphology. — Acuaria sensu stricto (p. 216) : Head with 2 lateral
lips from which the cordons originate. Body very slender.

Male 4.84 mm. long by 94/* wide. Head 20/* wide. Buccal cavity
or pharynx 140/* long; anterior esophagus 250/* long by 20/* Avide;
posterior esophagus 750//, long by 44/* wide. Spicules 133/* and 95/*
long, respectively. Cloacal aperture 135/* from end of tail; tail
curved. Caudal alae 240/* long, with slightly salient papillae.

226 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 18 mm. long by 160/* wide. Tail 230/x long. Eggs 34/*.
long.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe (France (Rennes)).

Genus CHEILOSPIRURA Diesing, 1861

Synonyms. — Acuaria Bremser, 1811, part; Dispharagus Dujardin,
1845, part.

Generic diagnosis. — Acuariinae (p. 211) : Body elongate, tapering
toward extremities. Head with 2 lips. Cordons directed posteri-
orly, not recurrent or anastomosing. Male with tail rolled in spiral,
alate or not alate; spicules very unequal and dissimilar; 5 to 7 pairs
of postanal papillae.

Parasitic between the tunics of the gizzard of birds.

Type-species. — Cheilosph^ura hamulosa (Diesing, 1851) Diesing,
1861.

KEY TO SPECIES OF CHEILOSPIBURA

1. Male 7.9 mm. long, female S.5 mm. long. Vulva in posterior part of body.

dividing body length in ratio of 34: 7 Cheilospirura rotundata, p. 229.

Male 10 mm. long or longer ; female 16 mm. long or longer. Vulva near
middle of body 2.

2. Cordons extend almost to posterior end of body ; 5 pairs of postanal papillae.

Cheilospirura hamulosa, p. 226.
Cordons on anterior region of body ; 6 pairs of postanal papillae 3.

3. Cordons spiny, not extending beyond anterior third of the first part of the

esophagus ; spicules 192^ and 700 to 720/tt long ; eggs 39 to 43/* by 25 to 27/*.

Cheilospirura spinosa, p. 229.

Cordons not spiny, extending to posterior region of first part of esophagus ;

spicules 170fi and 560/* long ; eggs 33/j. by 22/*__ Cheilospirura gruveli, p. 227.

Cheilospirura pavonis, recently described by Ortlepp (see Ad-
denda, p. 389), can be differentiated from the species keyed above by
its size, the male being only 6.5 mm. long, the female 14.7 mm. long,
and by the length of the cordons, which extend into the second
half of the body length but do not approach the posterior end.

An additional species recently described by Ozerska, Cheilo-
spirura skrjabini (see Addenda, p. 389), can be differentiated from
the above species by its short spicules, the one 205 to 239/t long, the
other 125 to 148/x long.

CHEILOSPIRURA HAMULOSA (Diesing, 1851) Diesing, 1861

Synonyms. — Spiroptera hamulosa (Diesing, 1851) ; Dispharagus
hamulosus (Diesing, 1851) Stossich, 1890; Spiroptera perforans Cen-
toscudi, 1911; Acuaria hamulosa (Diesing, 1851) Railliet, 1911.

Hosts. — Primary: Gallus gallus, Meleagris gallopavo, Phasianus
gall us; secondary: Unknown, probably arthropods, perhaps insects.
The turkey is reported as a host by Ransom and Hall.

NEMATODE PARASITES OF BIRDS

227

Location. — In the gizzard, in small fleshy growths on the surface
and in the wall.

Morphology. — Cheilospirura (p. 226) : Head (fig. 2905) with 2
large lateral lips in form of an equal-sided triangle, each lip with 2
lateral papillae on conical lobes; a finger-like median lobe also pres-
ent. The 4 cuticular cordons (fig. 290a) are double, irregularly wavy
and extend almost to the posterior extremity ; they do not anastomose
or recurve anteriorly.

Male 12 to 14 mm. long. Two very unequal and dissimilar spicules,
the left slender and long, six times as long as the right; the right
spicule shaped like a chopping knife. Posterior end bent. Tail with
2 very wide, double-contoured caudal alae. Five pairs of postanal

Fig. 200. — Cheilospirura hami'losa, a, Cordon; b. head, lateral view; c, male
tail. After Drasche, 1883

papillae (fig. 290c), of which 2 are small and near the tail end, 2
are large and at some distance from each other, and the fifth pair is
small and just posterior to the cloacal aperture.

Female 16 to 25 mm. long. Posterior end pointed. Vulva slightly
posterior to middle of body. Eggs 30/u, long by 20^ wide, according
to Centoscudi.

Life history. — Unknown; probably somewhat similar to that of
DispJuvrynx spiralis, p. 238.

Distribution. — North America (United States), South America
(Brazil, Argentina), Europe (Italy, France, Russia (Transcau-
casia)), and Australia.

CHEILOSPIRURA GRUVELI (Gendre, 1913) Cram, 1927

Synonyms. — Dispharagus species Gendre, 1912; Dispharagus gru-
veli Gendre, 1913; Acuaria gruveli (Gendre, 1913) Gendre, 1913;
Acuaria {Cheilospirura) gruveli (Gendre, 1913) Lopez-Neyra, 1923.

Hosts. — Primary: Caccabis rufa and Francolinus bicalcaratus ;
secondary: Unknown.

Location. — Between tunics of gizzard.

228

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology.— Cheilospirura (p. 226) : Mouth with 2 large lateral
lips (fig. 2916), rounded anteriorly, each with 2 pedunculated papil-
lae and a triangular area between them on' the external surface.
Four double cuticular cordons (fig. 291c) extending from the mouth,
which is far anterior, along the margins of the lips and then along
the submedian lines. Cervical papillae small, just posterior to origin
of the esophagus, and hence about at the anterior fifth of the length
of the cordons (fig. 291a).

Male 10.19 to 13.83 mm. long by 160 to 180/* wide. Caudal alae
(fig. 291^) with 2 concentric zones, the inner transversely striated,
the outer more or less scalloped and granular. Four pairs of preanal
and 6 pairs of postanal pedunculated papillae. Spicules (fig. 291 e

Fig. 291. — Cheilospirura gruveli. a, Anterior end; b, head; c, cordon;

d, FEMALE TAIL. AFTER GENDRE, 1912. C, RIGHT AND f, LEFT SPICULE; ff,
MALE TAIL. AFTER GENDBEj 1913

and /) very dissimilar and unequal, the left 560/x and the right
170/a long.

Female 28.9 to 42.74 mm. long by 220 to 270^ wide. Cordons
1.15 mm. long. Tail (fig. 2916?) 1/128 to 1/144 of total body length.
Pharynx or buccal cavity 180 to 280/a long; anterior esophagus
990ju. to 1.24 mm. long; posterior esophagus 2.01 to 2.38 mm. long.
Vulva conspicuous near middle of body, about 10/22 of length from
head. Eggs thick-shelled, 33ju, long by 22/a wide, embryonated when
oviposited.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa (Dahomey) and Europe (Spain).

NEMATODE PARASITES OF BIRDS

229

CHEILOSPIRURA ROTUNDATA (Linstow, 1907) Stiles and Hassall, 1920

Synonym's. — Dispharagus rotundatus Linstow, 1907; Acuaria ro-
tundata (Linstow, 1907) Railliet, Henry and Sisoff, 1912.

Hosts. — Primary: Lanius minor; secondary: Unknown.

Location. — In the lower jaw, according to Linstow (possibly an-
terior end of esophagus?).

Morphology. — Cheilospirura (p. 226) : Cuticle thick, finely striated
transversely. Head attenuated, with 2 papilliform lips. Cervical
papillae 320//. from head end. Cordons extend 480/* posterior to
head, not to posterior end of esophagus, the latter being very long.

Male 7.9 mm. long by 320/* wide. Esophagus 10/27 of body
length. Tail (fig. 292) 1/25 of body length. Four pairs of preanal

Figs. 292-293. — 292, Cheilospirura kotundata. Male tail. After Linstow, 1907.
293, Cheilospihura spinosa. a, Anterior end; b, part of cordon; c, vulva;
d, female tail; c. .spicule. Original

papillae and 5 pairs of postanal papillae. Spicules dissimilar, one
750/x long, slender, the other 180/* long, thick.

Female 8.5 mm. long by 470/* wide. Esophagus 10/24 of body
length. Tail 1/48 of body length. Vulva in posterior part of body,
dividing body length into ratio of 34:7. Eggs thick-shelled, 34/*
by 21,*.'

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Asia (Turkestan ) .

CHEILOSPIRURA SPINOSA. new species

Synonym. — Acuaria {Cheilospirura) species Stafseth and Kothin,
1925.

Hosts. — Primary: Bonasa umbellus; secondary: Unknown.

Location. — Gizzard.

Morphology. — Cheilospirura (p. 226) : Mouth (fig. 293«) with 2
triangular lips; at the base of each and projecting slightly beyond

230

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

its margin is a pair of papillae. Four cordons originating in pairs
between the lips; the 2 cordons of each pair gradually swing to the
submedian lines and do not extend beyond the anterior third of the
anterior esophagus; they do not recur or anastomose. Cordons
double and composed of 2 rows of discrete, sharply pointed elements
(fig. 293 a and b) which project like spines; they may extend for a
length of 10/30 to 10/41 of that of the anterior esophagus, the average
being 10/35. Cuticle with wide transverse striations. Cervical
papillae at the level of the union of pharynx and esophagus.

Male 15 mm. long (14 to 20 mm. in specimens of Stafseth and
Kotian) by 183/j, wide at the level of the posterior end of the esopha-
gus and 232/jt wide in the middle of the body. Cordons 495^. long.
Pharynx or buccal cavity 232ju, long; anterior esophagus 1.1 mm.
long; posterior esophagus 2.6 mm. long. Caudal extremity tightly

Fig. 294. — Cheilospirura spixosa. Male tail. Original.

coiled. Caudal alae (fig. 294) broad and vesicular; a central band
extending lengthwise has delicate transverse striations. Caudal
papillae very slender ; 4 pairs of preanal papillae ; 6 pairs of postanal
papillae arranged as follows : One group of 2 pairs not far posterior
to cloacal aperture, another group of 2 pairs near the caudal ex-
tremity, and between these 2 groups the other 2 pairs, the 2 papillae
of each pair being asymmetrically placed. Cloacal aperture 498//,
from posterior end of body. Spicules unequal and very dissimilar,
the one slender, 700 to 720/* long (660/a according to Stafseth and
Kotian), the other thick, 192 M long (fig. 293c).

Female 34 to 38 mm. long (37 to 40 mm. in specimens of Stafseth
and Kotian) by 315^ wide at posterior end of esophagus and 348/a
wide in middle part of body. Cordons 797 to 813/x long. Pharynx
or buccal cavity 232^ long; anterior esophagus 1.4 mm. long; pos-
terior esophagus 3.4 mm. long. Vulva (fig. 293c) anterior to middle
of body, dividing body length in ratio of 17 : 23. Anus 250 to 300/x
from posterior end. Tail (fig. 293^) slender and digitate. Eggs 39
to 42/a by 25 to 27/x.

NEMATODE PARASITES OF BIRDS 231

Life history. — Unknown; probably involves intermediate stages in
other hosts.

D'/.stiihiiliun. — North America (United States (Michigan)).

Type material. — No. 25538, U.S.N.M. (Bureau of Animal Industry
helminthological collection). Collected from a gizzard sent to the
laboratory by Dr. George R. La Hue, the gizzard having been sent
to him by Dr. Jan Metzeloas of the University of Michigan, Ann
Arbor, Michigan.

The description by Stafseth and Kotlan of a new species of
C heUospirura from the grouse in Michigan appeared after the pres-
ent writer's study of the material submitted by Doctor La Rue was
in manuscript. Doctor Kotlan. learning of this fact, kindly turned
over specimens of his material to the present writer and gave her
the privilege of naming the species. A comparison of the two lots of
material showed them to be the same nematode.

This species is close to C. gruveli in size, number of postanal
papillae, shape of short spicule and the position of vulva and anus
of female. Gendre, however, made no mention of the cordons of 0.
gruveli being spinous in nature and that is the most striking char-
acter of the species under consideration. Other differences are the
length of the spicules, the length of the cordons (in C. gruveli they
extend almost to the posterior end of the first part of the esophagus,
in C. spinosa not beyond the anterior third) the arrangement of
cephalic papillae, the position of the caudal papillae and the egg
sizes. The present writer has therefore made this a new series.

Genus CHEVREUXIA Seurat, 1918c

Synonyms. — Spiroptera Rudolphi, 1811). part; Dispharagus Dujar-
din, 1845, part.

Generic diagnosis. — Acuariinae (p. 211) : Body straight and fili-
form. Cuticle thick, distinctly striated transversely, and raised ante-
rior to the postcervical papillae to form a large collar, encircling this
part of the body as a hyaline ring or sheath. The anterior region of
the body also bears 4 cuticular cordons originating anteriorly in the
median dorsal and ventral lines, extending posteriorly along the sub-
median lines, and uniting on the lateral surfaces in a curve resting
on the free border of the collar; (hey are flanked on their inner sur-
face by a straight cuticular zone of marked striations. Postcervical
papillae symmetrical. Tail short, Caudal pores subterminal. Mouth
with 2 lateral obtusely pointed lips the shape of an inverted V; a
pair of sessile papillae at the angle of insertion of each lip. Buccal
cavity or pharynx long, straight and tubular. Esophagus distinctly
divided into a transparent muscular pari, surrounded directly pos-
terior to its origin by the nerve ring, and a glandular opaque part

232

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

of deepened color. Vulva ventral, slightly salient, near middle of
body. Ovejector short, directed posteriorly; uteri divergent. Eggs
thick-shelled, embryonated when oviposited. Caudal alae of male
hyaline, outspread during life. Nine pairs of pedunculated papillae,
4 of them preanal and 5 postanal. Spicules very unequal.

Parasitic between the tunics of the gizzard of birds.

Type-species. — Chevreuxia revoluta (Kudolphi, 1819) Seurat,
1918c.

CHEVREUXIA REVOLUTA (Kudolphi, 1819) Seurat, 1918c

Synonyms. — Spiroptera revoluta Rudolphi, 1819; Dispharagus
revolutus (Rudolphi, 1819) Molin, I860.

Hosts. — Primary: Gharadrius himantopus, Himantopus himan-
topus, H. melanopterus ; secondary : Unknown.

Location. — Between tunics of the gizzard.

Morphology. — Chevreuxia (p. 231) : Characters of the genus.

Fig. 20-5. — Chevreuxia revoluta. Anterior end. After Seurat, 1918

Male 6.4 mm. long by 140/* wide, straight and colorless. Postcervical
papillae relatively farther away from the nerve ring than in the
female; they are situated at the margin of the collar which is 420/x
from the head end. Tail straight. Caudal alae outspread during
life; in specimens killed by heat the tail is coiled in a spiral; the
alae are hyaline and are united in front of the caudal extremity.
Nine pairs of slender, symmetrical, pedunculated caudal papillae;
the 4 pairs of preanal papillae are equidistant; the 5 pairs of
postanal papillae are arranged in 2 groups, one group of 3 pairs
equidistant behind the cloacal aperture, the other group of 2 pairs
near the caudal extremity. Right spicule 95/x long, wide and with
obtuse end ; left spicule 750/* long, slender, filiform and pointed.

Female 13.2 to 18.3 mm. long by 265 /t wide. Tail 170/* long,
digitiform. Postcervical papillae hidden under the collar at its
anterior third and visible through the collar (fig. 295). Posterior

NEMATODE PARASITES OF BIRDS 233

border of collar 765/z from head end. Vulva near middle of body,
usually slightly posterior to middle, 6/11 of total body length from
head end. Ovejector short, directed posteriorly; vestibule horn-
shaped as in Dispharynsa and Synhimantus ; sphincter rectilinear;
unpaired varnish gland or trompe 35G> long. Eggs 32/x by 18/z,
thick-shelled, embryonated at maturity.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe and Africa (Algeria).

Genus COSMOCEPHALUS Molin, 1858

Synonyms. — Filaria Mueller, 1787, part; /Sp-iroptera Rudolphi,
1819, part; Dispharagus Dujardin, 1845, part; Histiocephalus
Diesing, 1851, part.

Generic diagnosis. — Acuariinae (p. 211) : Cordons recurrent and
anastomosing on the lateral surfaces of the cephalic extremity, not
flat against body cuticle but applied to the margins of membranous
plates or alae and forming a posteriorly directed loop immediately
after their origin at each side of the lips. Lateral papillae post-
cervical, situated posterior to the cordons. Lateral alae present,
originating directly posterior to the lateral papillae.

Parasitic in esophagus of birds.

Type-species. — Cosmocephalus diesingi Molin, 1858, emend.
Cram, 1927.

KEY TO SPECIDS OF COSMOCEPHALUS

1. Cordons not scalloped on inner edge; tail of female ending in a sharp point.

Cosmocephalus diesingi, p. 233.

Cordons scalloped on inner edge; tail of female ending in a buttondike or

suckerdike appendage 2.

2. Head with 4 prominent papillae; length of cordons 2.5 to 3 times the width

of body at level of cordons ; tail of female 10/569 of total length ; vulva 2/5
of total body length from anterior end ; spicules 130/a and 42tV long.

Cosmocephalus obvelatus, p. 235.

Head without prominent papillae; length of cordons less than 2 times the

width of body at level of cordons; tail of female 10/732 of total length;

vulva 20/43 of total length from anterior end : spicules l.SOju. and 700/x long.

Cosmocephalus aduncus, p. 'S'A.

COSMOCEPHALUS DIESINGI Molin. 1858, emended Cram, 1927

Synonym. — Cosmocephalus diesingii Molin, 1858.

Hosts. — Primary: Larus capistranus; secondary :Unknown.

Location. — Esophagus.

Morphology. — Cosmocephalus (p. 233) : Head pointed and subtri-
angular, distinctly set off from body. Four oval plates or alae
(fig. 296 a and b) attached to the head and joined anteriorly; judg-
ing from Molin's figure, these plates are bordered by cordons. Body

234 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

subcylindrical, attenuated posteriorly, slightly eurved, densely stri-
ated transversely. Two lateral alae with numerous transverse stria-
tions in anterior half of body. Two prominent cervical papillae just
posterior to cordons. Buccal cavity or pharynx short and slender.
Esophagus 1/3 as long as body and half as wide as body.

Male unknown.

Female 15 mm. long by 300[jl wide. Tail obliquely truncate and
pointed. Vulva near middle of body; judging from the length of
the esophagus and Molin's figure (fig. 296 a), the vulva is just ante-
rior to middle of body ; it has 2 prominent lips. " Oviduct " passes
posteriorly towards caudal extremity. Anus (fig. 296 c) a short
distance anterior to caudal apex.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Italy (Padua)).

Fig. 29G. — Cosmocephalus dibsingi. a, Anterior and middle parts of worm,

SHOWING POSITION OF VULVA ; b, ANTERIOR END ; C, FEMALE TAIL. AFTER MOLIN,

1861

COSMOCEPHALUS ADUNCUS (Creplin, 1846) Yorke and Maplestone, 1926

Synonyms. — Spiroptera adunca Creplin, 1846 ; Dispharagus adun-
cus (Creplin, 1846) Molin, 1860c.

Hosts. — Primary: Colymbus septentrionalis, Larus argentatus, L.
canus, L. glaucus, L. mari?ius, L. medius, L. ridibundus, Podiceps
auritus; secondary : Unknown.

Location. — Esophagus.

Morphology. — Cosmocephalus (p. 233) : Mouth (fig. 297tf) Avith
2 lips. No cephalic papillae observed. Cordons scalloped on inner
edge as in C. obvelatus, comparatively short, their length about 1.5
times the width of the body at the level of the cordons. Cervical
papillae small and bicuspid, a short distance posterior to the
cordons.

Male 8.6 to 11 mm. long by 280 to 300^ wide. Caudal extremity
(fig. 297 b) obtuse. Caudal alae wide. Nine pairs of long, pedun-
culated papillae, of which 4 are preanal and 5 postanal. Spicules
180[i long and 700/* long, the longer with a barb at its distal end.

Female 10.5 to 15 mm. long by 290 to 400/* wide. Tail 10/732 of
total body length, with a sucker-like enlargement at the end. Vulva

NEMATODE PARASITES OF BIRDS

235

slightly anterior to middle of body, dividing body in ratio of 31 : 39.
Eggs 39,u by 23/i.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distrib ution . — Europe.

Linstow reported this from Lcerw en mis and noted that it differed
from C. obvelatus in the length of the cordons and the presence of
prominent cephalic papillae; other differences such as lengths of
spicules and of female tails support the idea that this is a good
species.

COSMOCEPHALUS OBVELATUS (Creplin, 1825) Seurat, 1919a

Synonyms. — Spiroptera obvelata Creplin, 1825; Filaria obvelata
(Creplin, 1825) Linstow, 18776/ Dispharagus obvelatus (Creplin
1825) Linstow, 1909&/ Histiocephalus spiralis Diesing, 1851; Cosmo-
cephalus papillosus Molin, 1859; Dispharagus papillosus (Molin,
1859) Stossich, 1898; Cosmocephalus alatus Molin, 1860^.

Fig

297.-

-cosmocbphalus aduncds. «, ilead exd ; b, male tail. after
Stossich, 1892

Hosts. — Primary: Actitis hypoleucus, A. macularia, Alca torda,
Gatorrhactes pachyrhynchus (new record), Larus argentatus, L.
argentoides, L. canus, L. fuscus, L. marinus, L. maximus, L. medius,
L. ridibundus, Mergus se?vator, Puffinus kuhli, /Sterna arctica, S.
lisoria, Totanus hypoleucus, T. maculatus, T. fuscus, T tin goides
hy poleucus, Uria grylle; secondary: Unknown.

Location. — Esophagus.

Morphology. — Cosmocephalus (p. 233) : Cordons (fig. 298 a and b)
scalloped on inner edge, the course of the cordons sinuous; originating
in the dorso-ventral lines directly behind the mouth, they first run
posteriorly for a short distance, then loop forward again, then form
a second curve to gain their posteriorly directed course; considerably
farther posteriorly they form a third curve, running anteriorly in
the lateral fields, and then anastomosing in pairs; (he cuticle is
somewhat raised at the level of the second curve. Lateral papillae
bicuspid, at the level of the origin of the muscular esophagus;
posterior to these 2 papillae there are 2 lateral alae, finely striated
transversely, extending to the posterior end of the esophagus. Mouth

236

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

with 2 lateral lips, each bearing a conical tooth and with a pair
of very large papillae at the base. Buccal cavity or pharynx very
long, straight and tubular, extending to the level of the bicuspid
papillae. Nerve ring around esophagus near its origin.

Male 5.7 to 12.2 mm. long by 240 to 255/* wide. According to
Seurat, in specimens 12.2 mm. long the cordons are 400/* long, the
bicuspid papillae are 430/* from the head end, and the tail is 420/*
long; according to Linstow, in specimens 5.7 mm. long the cordons
are 260/* long, the point of anastomosis is 72/* from the head, and the
tail length is 1/21 of the body length. Caudal alae hyaline, long
and wide, uniting anterior to caudal extremity. According to Seurat,

Figs. 298-299. — Cosmocebhalus obvelatus. 298, a, Dorso-ventral view
and b, lateral view of an anterior end; c, female tail, ventral view.
After Seurat, 1919. 299, Male tail. After Linstow, 1877

there are 3 preanal papillae on the right side and 4 on the left ; there
are 5 pairs of pedunculated postanal papillae and in addition there
are 5 small sessile papillae grouped between the 2 papillae of the
most posterior pair. Linstow described and figured 10 pairs of
papillae, of which 4 pairs are preanal and 6 pairs postanal, 1 pair
of the latter lying between the 2 papillae of another pair at the caudal
extremity (fig. 299). Spicules dissimilar and unequal, the right
thick and 130/* long, the left slender and 420/* long in small speci-
mens (Linstow), or 155/* and 540 /* long in large specimens (Seurat).
Female 9.7 to 20 mm. long by 300 to 380/*, wide. According to
Seurat, in a specimen 13.1 mm. long the cordons were 410/* long, the
bicuspid papillae 490/* from the head end, the vulva 2/5 of the total
body length, or 5.5 mm., from the head end, the anus (fig. 298c)
230/* from tail end, the eggs 36/* by 20/*, embryonated when oviposited.
According to Linstow, in a specimen 9.77 mm. long the cordons wer* 1

NEMATODE PARASITES OF BIRDS 237

440/a long, their anastomosis HO/*, from the head end, and the eggs
26/x by 21/x. The tail is conical and ends in a small rounded button.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution, — Europe, Africa (Algeria), and North America
(United States (National Zoological Park, Washington, D. C.)).

Genus DISPHARYNX Railliet, Henry, and Sisoff, 1912

Synonyms. — Acuaria Bremser, 1811, part; Spiroptera Rudolphi.
1819, part; Dispharagus Dujardin, 1845, part.

Generic diagnosis. — Acuariinae (p 211) : Cordons recurrent, not
anastomosing. Lateral papillae postcervical, small. Males with
spicules unequal and dissimilar. Usually with 5 pairs of postanal
papillae.

Parasitic in esophagus, gizzard and proventriculus of birds.

Type species. — Dispharynx nasuta (Rudolphi, 1819) Stiles and
Hassall, 1920.

KEY TO SPECIES OF DISPHARYNX

1. Inadequately described; from Falco tninutus or Rhamphastos vitellinus — 2.
Adequately described; from bosts other than above 3.

2. Lips very small ; cordons short ; from Falco minutus.

Dispharynx capitata, p. 240.
Lips large; cordons long; from Rliamphasios vitellinus.

Dispharynx crassissima, p. 243.

3. Male with 7 pairs of postanal papillae ; female 13 to 16 mm. long.

Dispharynx magnilabiata, p. 241.

Males with less than 7 pairs of postanal papillae; female not over 11 mm.

long 4.

4. Male with 4 pairs of postanal papillae ; longer spicule twice the length of the

shorter one Dispharynx rectovaginata, p. 243.

Male with 5 pairs of postanal papillae ; longer spicule either considerably

more or considerably less than twice the length of the shorter one 5.

.1. Vulva anterior to middle of body Dispharynx nasuta, p. 237.

Vulva posterior to middle of body 6.

6. Postcervical papillae situated posterior to cordons; vulva just posterior to

middle of body ; longer spicule 2GG> long Dispharynx noctuae, p. 242.

Postcervical papillae situated between the cordons ; vulva in posterior third
of body ; longer spicule 400> long or longer 7.

7. Postcervical papillae bicuspid ; tail of male about 27:V. of female about

12<V long; spicules 150/* and 4007* long Dispharynx spiralis, p. 238.

Postcervical papillae tricuspid; tail of male 370ti, of female 17G> long;
spicules 240y, and 865^ long Dispharynx laplantei, p. 241.

See Addenda, p. 391, for a species of Dispharynx recently described
by Smit and Notosoediro.

DISPHARYNX NASUTA (Rudolphi, 1819) Stiles and Hassall, 1920

Synonyms.- — Spiroptera nasuta Rudolphi, 1819; Dispharagus
nasutus (Rudolphi, 1819) Dujardin, 1845; Acuaria {Dispharynx)

238

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

nasuta Railliet. Henry and Sisoff, 1912; C heilospirura nasuta
(Rudolphi, 1819) Ransom, 1916.

Hosts. — Primary : Gallus gallus; secondary : Unknown, but prob-
ably arthropods. PorceUio laevis reported as intermediate host by
Piana ; apparently he was dealing with Dispharynx spiralis.

Location. — Proventriculus and gizzard and connective tissue
between these.

Morphology. — Dispharynx (p. 237) : Two small lateral lips (fig.
300a). Worms white or red. Esophageal bulb with 3 teeth, accord-
ing to Schlegel. Cordons similar to those in D. spiralis (p. 239).

Male 5 mm. long, filiform. Tail end (fig. 3006) spiral, according
to most writers; Schlegel says it is not. According to Schlegel, the
right spicule is long and thin, the left one-third as long, and thicker ;
both are dark brown. Five pairs of postanal papillae.

a.

Fig. 300. — Dispharynx nasi ta. Head and tail of male. After Dujakdin, 1845

Female 5 to 9 mm. long, or 9 to 10 mm. according to Schlegel.
Tail end conical, or, according to Schlegel, awl-shaped. Vulva in
anterior portion of body, according to Schlegel; some writers say
that vulva is in posterior portion ; this species has been confused with
D. spiralis in some cases and some writers would unite the two as
D. nasuta, Eggs thick-shelled.

Life history. — Unknown; Piana reported a life history involving
intermediate stages in the sow-bug, PorceUio laevis, but later writers
believe he was dealing with D. spiralis. The 2 species may have very
similar life histories.

Distribution. — North America (United States), South America,
Africa (Belgian Congo), Australia, and Guam. The records of this
worm from various localities are believed by Kailliet to be records of
D. spiralis (p. 238). Both species need reconsideration before one
may undertake to give their distribution with any certainty.

DISPHARYNX SPIRALIS (Molin, 1858) Skrjabin, 191fib

/Synonyms. — Dispharagus spiralis Molin, 1858; Dispharagus nasu-
tus of Piana, 1897; Dispharagus spiralis columbae Bridre, 1910;
Acuaria spiralis (Molin, 1858) Railliet, Henry, and Sisoff, 1912.

NEMATODE PARASITES OF BIRDS

239

Hosts. — Primary : Gallus gallas, Bonasa umbellus, Gaccabis petrosa,
Columbia. livia\ Mdeagris gailopavo, Phasia/rms gallus, Numida me-
leagris, Quiscalus quiscfula; secondary: .^owbug (Porcellio laevis).

Location. — Esophagus, proventriculus, and intestine, fixed in
mucosa as adults, and encysted in connective tissue about esophagus,
crop, proventriculus and intestine and in the mesentery of primary
host; in anterior portion of digestive canal of secondary host.

Morphology. — Dhphmynx (p. 237) : No vesicular swelling ante-
riorly. Four wavy cutaneous cordons (figs. 301, 302, and 303a) on
anterior end, originating at the 2 papilliform lips, not anastomosing,
but recurrent, the distal extremity of the cordons turning forward
and extending anteriorly a short distance. Postcervical papillae
small, bicuspid, situated between the recurrent branches of the cor-
dons. Body usually rolled in a spiral.

Fig. 301. — Dispharynx spiralis. Male. After Piana, 1897

Male 7 to 8.3 mm. long by 230 to 315/x wide (fig. 301). Cordons
415 to 515/i long. Five pairs of postanal and 4 pairs of preanal
papillae. Cloacal aperture about 275/* from tail end. Long spicule
400/t long, slender and curved; short spicule 150/* long, navicular.
Ventral surface of body in cloacal region and for a certain distance
anterior to cloaca covered with small, elongated shields which give
the appearance of a longitudinal striation.

Female 9 to 10.2 mm. long by 360 to 565/* wide (fig. 302) . Cordons
900/* to 1.06 mm. long. Anus 120/* from tip of tail. Small mucron
on tip of tail. Vulva in posterior portion of body, 2 to 2.45 mm.
from tip of tail. Cylindrical ovejector (fig. 3035) curved, S-shaped
at its origin, and then directed anteriorly; the limiting boundary of
the vestibule and sphincter is clearly marked by an annular cuti-
cular fold; varnish gland (trompc) unpaired, 250/* long. Eggs 36
to 40/t by 21/t, embryonal ed when oviposited.

Life history. — According to Piana, the life history of what he
called Dispharagus nasutus, but which was apparently Disph/trynx
3012—27 17

240

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

spiralis, is as follows : The eggs of the worm pass out in the droppings
and are eaten by sowbugs. The embryos escape and develop to in-
fective larvae in the body cavity. When infested sowbugs are eaten
by suitable birds, the larvae develop in them to adults. This work
is based on comparative studies, not on experimental feedings.

Distribution. — North America (United States and Porto Rico),
Europe (Italy, France, Spain), Asia (Russian Turkestan), Africa
(Tunis), and Australia. In addition to the above localities may be
given Austria, Belgian Congo, and Algeria, as Railliet considers that
the reports of D. nasuta from these places are probably records of
D. spiralis.

Figs. 302-303. — Dispharynx spiralis. 302, Female. After Piana, 1897. 303,
a, Head end. After Seurat, 1916. b, Ovejector. After Set; rat, 1919

DISPHARYNX CAPITATA (Molin, 1860) Cram, 1927

Synonyms. — Dispharagus capitatus Molin, 1860 ; Spiroptera alata 4
in Molin, 1860; Acuaria capitata (Molin, 1860) Railliet, Henry, and
Sisoff, 1912.

Hosts. — Primary : Falco minutus; secondary : Unknown.

L ocation. — Pro vent riculus.

Morphology. — Dispharynx (p. 237) : Head conical, continuous with
body. Cordons markedly flexed, short, strongly recurrent, not
anastomosing. Mouth with 2 very small papilliform lips. Body
filiform and dense, with delicate transverse striations.

Male unknown.

Female 11 mm. long by 300/* wide. Posterior extremity conical,
apex obtuse. Anus not far from caudal extremity.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

* Catalogue of the Vienna Museum.

NEMATODE PARASITES OF BIRDS 241

DISPHARYNX LAPLANTEI Seurat, 1919a

Synonym. — Acuaria {Dispharynx) laplantei Seurat, 1919.

Hosts. — Primary : Garrulus glandarius cervicalis; secondary : Un-
known.

Location. — Proventriculus.

Morphology. — Dispharynx (p. 237) : Body large, attenuated at ex-
tremities. Cordons similar to those of D. noctuae and D. spiralis.
Postcervical papillae tricuspid, subsymmetrical ; Seurat says they are
situated as in D. spiralis, directly in front of the level of the excre-
tory pore, but according to his figures the excretory pore is 590/* and
390/x from the head end in the female and male respectively, whereas
the cervical papillae are 612/* and 672/* from the head end in the 2
cases. Mouth with 2 large triangular lips.

Male 7.7 to 8.4 mm. long, with a maximum width of 300/*; much
slenderer than female. Cordons 455/* long. Caudal extremity
coiled in spiral. Cloacal aperture 370/* from tail end. Four pairs
of preanal and 5 pairs of postanal papillae, arranged as in D. noctuae
and D. spiralis. Right spicule 240/*. long, thick and falciform; left
spicule 865/* long, slender, pointed at tip, and not alate.

Female 9.9 mm. long by 565/* wide. Cordons 805/* long. The
massive body narrows suddenly posterior to the vulva, which is in
the posterior third of the body, 2.87 mm. from the tail end; vulva
not salient. Vestibule (fig. 304) extends 150/* anteriorly, then sud-
denly turns posteriorly, a large unicellular gland being situated at
this point; the descending branch is 320/* long and joins the branches
of the varnish gland (trompe), there being no unpaired varnish
gland. Anus 170/* from tip of tail. Eggs 37/* by 25/*, thick-shelled,
embryonated when oviposited.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Africa (Medea and Ain Ograb).

DISPHARYNX MAGNILABIATA (Molin, 1860) Gendre, 1920d

Synonyms. — Displiaragus magnilabiatus Molin, 1860c; Spiroptera
plataleae-ajajae 5 Molin, 1860; Acuaria (Cheilospirura) magnilabiata
(Molin, 1860) Railliet, Henry and Sisoff, 1912; Cheilospirura magni-
labiata (Molin, 1860) Stiles and Hassall, 1920.

Hosts. — Primary: Ajaja ajaja, Platalea ajaja; secondary:
Unknown.

Location. — Between the tunics of the gizzard.

Morphology. — Dispharynx (p. 237) : Mouth with 2 large papilli-
form lips spread out anteriorly. Cordons straight, not wavy, strongly
recurrent.

"Catalogue of Ihe Vienna Museum.

242

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 7.5 mm. long by 200/z wide. Tail (fig. 305) loosely rolled in
spiral. Caudal alae long and wide. Four pairs of pedunculated
preanal and 7 pairs of postanal papillae.

Female 13 to 16 mm. long by 300/* wide.

Gendre (1920a), on the basis of Molin's description of the cordons
as recurrent, states that this species belongs in Dispharynx and not
in C heilospirura, to which it had been assigned by Kailliet, Henry,
and Sisoff.

Figs. 304-306. — 304, Dispharynx laplantei. Ovejector. After Seurat,
1919. 305, Dispharynx magnilabiata. Male tail. After Drasche,
1884. 306, Dispharynx noctuae. a, Vulva ; 6, anterior end, dorsal
view; c, female; d, anterior end, lateral view; e, ovejector and
vulva. After Seurat, 1913

Life history. — Unknown; probably involves intermediate stages
in other hosts.
Distribution. — South America (Brazil).

DISPHARYNX NOCTUAE (Seurat, 1913a) Stiles and Hassall, 1920

Synonym. — A cuaria noctuae Seurat, 1913; Acuarid {Dispharynx)
noctuae Seurat, 1913a,

Hosts. — Primary : Carine noctua glaux ; secondary : Unknown.

Location. — Esophagus.

Morphology.— Dispharynx (p. 237) : Mouth (fig. 306& and d) with
2 triangular lateral lips, each bearing a large papilla at the base.
Buccal cavity or pharynx narrow and remarkably long. Cervical
papillae posterior to cordons.

NEMATODE PARASITES OF BIRDS

243

Male 6.5 mm. long. Buccal cavity 170 to 200/* long. Caudal
alae very large. Four pairs of preanal and 5 pairs of postanal
papillae. Spicules 260/* and 115/* long.

Female 10 to 11 mm. long by 350/* wide at the level of the vulva.
Buccal cavity 280 to 300/* long. Tail 170/* long. Vulva (fig. 306a
and e) opening on an oval prominence directly behind the middle
of the body. Eggs 43/* by 21/*, thick-shelled, embryonated when
oviposited.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa (Algeria) .

DISPHARYNX RECTOVAGINATA (Molin, 1860) Cram, 1927

Synonyms. — Dispharagus rectovaginatus Molin, 1860; Acuaria
rectovaginata (Molin, 1860) Railliet, Henry, and Sisoff, 1912.
Hosts. — Primary : Faleo ater; secondary : Unknown.
Location. — Pro vent riculus.

-^UhA-%^

-*=&'

fig. .'507. dispharynx rectovaginata. head end and tail of male. after

Dbaschb, 1884

Morphology. — Dispharynx (p. 237) : Head continuous with body.
Mouth with 2 conspicuous triangular lateral lips, each bearing 2
papillae. Body densely striated transversely and twisted in spiral.
Two cordons (fig. 307), sinuous and recurrent, not anastomosing.

Male 4 mm. long by 200/* wide. Posterior extremity (fig. 307)
coiled in 3 spiral turns. Caudal alae conspicuous, the margin thick-
ened like a cord. Papillae thick, 4 pairs preanal and 4 pairs post-
anal, alternately short and long. Left spicule twice as long as right.

Female 6 mm. long by 400/* wide. Tail short and acutely conical,
the anus not far from its extremity.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution, — South America (Brazil).

DISPHARYNX CRASSISSIMA (Molin, I860) Cram, 1927

Synonyms. — Dispharagus crassissimus Molin, 1860; Acuaria cras-
sissima (Molin, 1860) Railliet, Henry and Sisoff, 1912.

244 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Primary: Rhaniphastos vitellinus; secondary: Unknown.

Location. — Proventri cuius.

Morphology. — Dispharynx (p. 237) : Head continuous with body.
Mouth with 2 large papilliform lips. Body with dense transverse
striations. Anterior extremity not attenuated and with obtuse apex.
Two cordons, long and thick, markedly flexed, strongly recurrent, not
anastomosing.

Male unknown.

Female 12 mm. long by 500/*, wide. Tail abruptly acute, conical,
with very sharp tip. Anus not far from caudal extremity. Vulva in
posterior part of body.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil).

Genus ECHINURIA Soloviev, 1912

Synonyms. — Filaria Mueller, 1787, part; Acuaria Bremser, 1811,
part; Spiroptera Rudolphi, 1819, part; Dispharagus Dujardin, 1845,
part; Histiocephalus Diesing, 1851, part; Dispharynx Railliet,
Henry, and Sisoff, 1912, part ; Hamannia Railliet, Henry, and Sisoff,
1912/

Generic diagnosis. — Acuariinae (p. 211) : Cordons not recurrent,
but anastomosing posteriorly in pairs in the lateral fields; their
course and relations to other structures are somewhat variable.
Body sometimes provided with spines disposed in a regular manner.
Spicules unequal and dissimilar. Postanal papillae 4 to 5 pairs, or
lacking. Usually 2 uteri, occasionally only one (E. decorata).
Vulva usually in posterior part of body. Parasitic usually in pro-
ventriculus and gizzard, sometimes in tumors in gizzard, in birds.

Type species. — Echinuria jugadornata Soloviev, 1912.

KEY TO SPECIES OF ECHINURIA

1. Vulva in middle of body ; cordons extend to near middle of body.

Echinuria ardeae, p. 248.

Vulva in posterior portion of body; cordons where length is given, confined

to anterior portion of body 2.

2. Cordons elaborate, their inner part made up of transverse bands, their

outer edge with posteriorly directed scales or teeth; esophagus one-
fourth of total body length or longer 3.

Cordons not as described above; esophagus length, if given, much shorter
than above 4.

3. Cordons narrow and of same width throughout, their anastomosis 1 mm.

from head end; postcervical papillae very large and 3-pointed.

Echinuria squamata, p. 257.

Cordons become progressively wider until they almost cover the body, their

anastomosis 1.9 to 2.4 mm. from head end; postcervical papillae very

small and inconspicuous and with only 1 point_ Echinuria decorata, p. 250.

NEMATODE PAEASITES OF BIRDS 245

4. Vulva posterior, dividing body in ratio of 4: 1.

Echinuria phoenicopteri, p. 257.
Vulva near posterior extremity of body, just anterior to anus 5.

5. Body with 2 double rows of spines extending almost entire length of

body 6.

B«-dy without such rows of spines, or at least none described 8.

6. Male 3.4 mm. long ; female 4 to 11 mm. long ; body surrounded by a cuticular

fold just posterior to anastomosis of cordons ; longitudinal rows of spines

originate just posterior to this fold Echinuria horrida, p. 253.

Male 9 mm. long or longer ; female 15 mm. long or longer ; no cuticular
fold as above ; longitudinal rows of spines originate far anterior to
anastomosis 7.

7. Male 9 mm. long ; spicules 706 /* and 208^, long, respectively.

Echinuria uncinata, p. 24G.
Male 11 to 12 mm. long; spicules 839/i and 140/j. long, respectively.

Echinuria jugadornata, p. 245.

8. Posterior extremity of female ending in a well-developed spur 9.

Posterior extremity of female without such spur 10.

9. Male and female described ; female 19 mm. long ; mouth with 2 small incon-

spicuous lips Echinuria contorta, p. 249.

Only female described ; 8 to 10 mm. long ; mouth with 2 large conspicuous
projecting lips Echinuria calcarata, p. 249.

10. Description incomplete; from Ciconia mayuari.

Echinuria longeornata, p. 255.
From hosts other than above 11.

11. Deft spicule 6 times as long as right Echinuria longevaginata, p. 256.

Left spicule less than 6 times as long as right 12.

12. Male 6.5 mm. long; female 6 to 12 mm. long; right spicule 180/t, left spicule

900/x, long; eggs 32ii by 20*t Echinuria leptoptili, p. 254.

Male 11 to 11.5 mm. long; female 13 to 15 mm. long; right spicule 190 to
210/u, left spicule 650 to 075m, long ; eggs 28/tt by 18^.

Echinuria hargilae, p. 253.

ECHINURIA JUGADORNATA Soloviev, 1912

Hosts. — Primary : Anas boschas; secondary : Unknown,

Location. — In tumor at union of proventriculus and gizzard.

Morphology. — Echinuria (p. 244) : Cuticula smooth, not striated,
provided with 4 longitudinal rows of spines (fig. 2086) each 57/x
long by 32/a wide at the base, originating a little behind the anterior
extremity and extending to the posterior extremity of the body.
Mouth with 2 projecting lips and G papillae, not always readily
visible. Four cordons (fig. 308 a) extend posteriorly from the buccal
aperture and join in pairs at their posterior terminations. Pharynx
103/a long by 47/t wide.

Male 11 to 12 mm. long by G00/t wide. Cordons 761)/*. long by 14/t
wide. Tail (fig. 308c) curved ventrally. Spicules unequal, one 839/x
long by 21//, wide, and curved, the other 140/t long by 33/*, wide, or
wider.

Female 15 to 16 mm. long by 800/t wide. Cordons 792/t long by
23//, wide. Tail pointed. Eggs 35.4/x by 21.8/*, with shells 3//. thick.

246

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown ; probably involves intermediate stages in
other hosts (see E. uncinata, p. 246).

Distribution. — Asia (Russian Turkestan).

There is nothing in the description, given up to this time, of E.
jugadornata and E. uncinata to differentiate the one from the other,
except the slight difference in the size of the male as shown in the
key, whereas the size of the females is the same, the character of
the head and cordons is identical, the hosts of the 2 species are closely
related, and both form nodules or tumors. Since there are several
points on which the 2 descriptions are not comparable, however, due
to scarcity of detailed statement in one or the other (such as caudal

Fig. 308. — Echinuria jugadornata. a, Cordon. After Soloviev, 1912. b, Head
end; c, male tail. From Skrjabin, 1916, after Soloviev, 1912

papillae, spicule lengths, position of vulva, etc.), and since Soloviev
would undoubtedly have made a study of E. uncinata before placing
it as he did in his new genus with E. jugadornata, the present writer
accepts E. jugadornata as a good species on Soloviev's authority in
default of evidence, but wishes to invite attention to this lack of
any adequate evidence on which to differentiate it from E. uncinata.
Since writing the above, the present writer has found specimens
agreeing with the description of E. uncinata and finds that the lengths
of the spicules of the male differ somewhat from those given for E.
jugadornata.

ECHINURIA UNCINATA (Rudolphi. 1819) Soloviev, 1912

Synonyms. — Spiroptera uncinata Rudolphi, 1819; Dispharagus
uncinatus (Rudolphi, 1819) Railliet, 1893; Acuaria (Hamannia)
uncinata (Rudolphi, 1819) Railliet, Henry, and Sisoff, 1912;
Hamannia uncinata (Rudolphi, 1819) Stiles and Hassall, 1920.

Hosts. — Primary: Anas boschas domestica, A. penelope, A.
rubripes, Anser cinereus dom£sticus, Cygnus olor domesticus, Nettion
carolinense; secondary: Small crustaceans (Daphnia puUx).

NEMATODE PARASITES OF BIRDS

247

Location. — Esophagus, proventriculus, gizzard and small intestine,
in the mucosa, and reported once from air sacs, of primary host; in
body cavity of secondary host.

Morphology. — Echinuria (p. 2-1-1) : Mouth with 2 lips and G
papillae. Anterior end (figs. 309 and 310a) not provided with a
vesicular swelling, but bearing 4 cutaneous cordons which anastomose
in pairs on the lateral surfaces. On each side of the body a double
longitudinal series of small spines extending posteriorly almost to
the end of the body and swinging anteriorly to the dorsal surface
between the cordons. Cordons inflected toward the latero-ventral

Figs. 300-310. — Eciiinuria uncinata. 309, Head end; ventral
view. After Seurat, 1919. 310, u, Head end, dorsal view ; b,
cross section, and c, ventral view of male tail. after
Schneider, 1866.

lines where they are joined a short distance beyond the excretory
pore, the change of course being especially noticeable in the latcro-
dorsal cordons which pass beneath the cervical papillae. Cervical
papillae quite far forward, at level of nerve ring, at the height of
the third spine of the external row.

Male 9 to 10 mm. long. Tail (fig. 3105 and c) with straight caudal
alae, somewhat vesicular. Schneider states that there are -1 pairs
of preanal and 4 pairs of postanal papillae, but his figure indicates
that the first pair of postanal papillae are double, suggesting that
there are 5 pairs of postanal papillae; Linstow reports 5 pairs. The
preanal are arranged in 2 groups of 2 pairs each set close together.
The long spicule is slender and alate, with a cuplike widening at its
free end: the other spicule is short and thick. The present writer
3612—27—18

248 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

has recently found specimens agreeing with the descriptions of this
species, in Nettion carolinense and Anas rubripes, and is able to add
the following particulars: Male 8 to 10 mm. long by 300/a to 500/*.
wide. In a male 8 mm. long, cordons 500/* long by 20/* wide;
pharynx 133/*, esophagus 830/*, long ; cloacal aperture 330/* from pos-
terior end ; spicules 706/* and 208/* long, respectively.

Female 12 to 18.5 mm. long. In a female 12 mm. long by 515/*
wide, the pharynx is 150/* long ; cordons 748/* long by 25/* wide ; vulva
1.3 mm., anus 250/*, from posterior end; eggs 37/* by 20/*. Vulva 1
to 1.4 mm. from tip of tail. Ovejector of type of Dispharynx
ovejector, composed of short vagina perpendicular to wall of body
and joining the ovejector proper ; this is directed posteriorly and is
divided into a vestibule and sphincter, the limit of which is marked
by a thickening of the muscular tunic. The sphincter passes directly
to the 2 branches of the trompe ; these extend parallel to the ovejector
to join the uteri ; the latter extend forward at first, twist into a loose
spiral, then separate, one continuing anteriorly while the other turns
and extends posteriorly. Numerous thick-shelled eggs.

Life history. — According to Hamann, the eggs pass out in the
feces of birds and on getting to water may be swallowed by water
fleas, Daphnia pulex. In the intestine the embryo escapes from the
egg and makes its way to the body cavity, where it develops to an
infective larva 1.7 to 2 mm. long. When such infected water fleas
are eaten by suitable bird hosts, the worms develop to maturity.

Distribution. — Europe (Germany and France) and Africa (Al-
geria) and North Anierica (United States).

See the discussion of this species in connection with E. jugadornata
(p. 246).

ECHINURIA ARDEAE (Smith, Fox, and White, 1908) Cram, 1927

Synonyms. — Dispharagus ardeae Smith, Fox, and White, 1908;
Acuaria ardeae (Smith, Fox, and White, 1908) Ward, 1918.

Hosts. — Primary: Ardea herodias; secondary: Unknown.

Location. — Alimentary tract.

Morphology. — Echinuria (p. 244) : Body slightly attenuated an-
teriorly. Cuticula rather coarsely striated transversely. Mouth
(fig. 311 a and b) with 2 prominent lateral lips, each with a pair of
papillae. Cordons originating at the base of the lips on each side,
passing posteriorly along the submedian lines to nearly the middle
of body length, then passing dorsally and ventrally, respectively, to
anastomose with the corresponding cordons from the opposite side.

Male unknown.

Female 17 mm. long by 700/* wide. Head 150/* wide at base of
lips. Anterior esophagus 800/* long by 50 to 90/* wide; posterior

NEMATODE PARASITES OF BIRDS

249

esophagus 1.2 mm. long by 200> wide. Vulva near middle of body
length. Anus 350> from tail end; at this level the body suddenly
narrows to form a conical tail, with a marked latero- ventral rounded
prominence on each side of the anus (fig. 311c). Eggs not found.

Life history. — Unknown; probably involves intermediate stages
in other hosts (see E. uncinata, p. 246).

Distribution. — North America (United States (Zoological Garden,
Philadelphia, Pennsylvania) ).

ECHINURIA CALCARATA (Molin, 1860) Cram, 1927

Synonyms. — Dispharagus calcaratus Molin, 1860; Acuaria cal-
carata (Molin, 1860) Railliet, Henry, and Sisoff, 1912; Hamannia
calcarata (Molin, 1860) Stiles and Hassall, 1920.

Hosts. — Primary : Ibis guarauna; secondary : Unknown.

Location. — Proventriculus.

Morphology. — Echinuria (p. 244) : Characters of the genus.

Male unknown.

Fig. 311.

-Echinuria ardeae. a, Front view and b, lateral view of head; c,

FEMALE TAIL. AFTER SMITH, FOX, AND WHITE, 1908

Female 8 to 10 mm. long by 200/x, wide. Mouth with 2 large con-
spicuous projecting lips (fig. 312 a and b). Cuticula transversely
striated. Anterior extremity of body appreciably attenuated. Four
long cordons, not recurrent but anastomosing posteriorly. Poste-
rior extremity with a spur-like appendage (fig. 312c). Anus near
caudal extremity. Vulva anterior to anus.

Life history. — Unknown; probably involves intermediate stages
in other hosts (see E. uncinate,, p. 246).

Distribution. — South America (Brazil).

This species may be identical with E. contorta, but the description
is not sufficiently complete for a definite decision in regard to this.

ECHINURIA CONTORTA (Molin, 1858) Cram, 1927

Synonyms. — Dispharagus contortus Molin, 1858; Spiroptera fal-
cinelli Rudolphi, 1819; Acuaria falcinelli (Rudolphi, 1819) Molin,
1860; Acuaria contorta (Molin, 1858) Railliet, Henry, and Sisoff,
1912; Hamatnniu contorta (Molin, 1858) Stiles and Hassall, 1920.

250

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Spiroptera falcinelli Kudolphi, 1819, is a nomen nudum, and hence
not available.

Hosts. — Primary: Ibis falcinellus, FalcineUus igneus; secondary:
Unknown.

Location. — In muscular ventriculus (gizzard).

Morphology. — Echinuria (p. 244) : Mouth with 2 small incon-
spicuous lips. Cuticula with dense transverse striations which are
twisted irregularly. Four cordons (fig. 313<z), anastomosing.

Male 7 to 8 mm. long by 200/* wide. Tail (fig. 3135) curved.
Caudal alae wide, semilunar, and transversely striated. Molin states
that there are 7 pairs of caudal papillae, but only figures 3 pairs
of postanal papillae.

Figs. 312-313. — 312, Echinuria calcarata. a, lateral and h, ventral view of
head; c, female tail. After Drasche, 1884. 313, Echinuria contorta. a,
anterior end ; b, male tail ; c, female tail, showing vulva and anus. After
Molin, 1861

Female 19 mm. long by 40/t wide. Caudal extremity (fig. 313c)
with short, obtusely conical lateral appendages. Anus said to be at
caudal apex. Vulva somewhat anterior to anus.

Life history. — Unknown; probably involves intermediate stages
in other hosts (see E. uncinata, p. 24G).

Distribution. — Europe (Italy (Padua) and Austria (Vienna)).

ECHINURIA DECORATA, new species

Hosts. — Primary: Colymbus auritus; secondary: Unknown.

Location. — Beneath lining of gizzard.

Morphology.— Echinuria (p. 244) : No cephalic papillae observed.
Mouth with 2 simple triangular lateral lips, followed by a pharynx
and a two-part esophagus. Esophagus long, more than one-fourth
of body length. Nerve ring at union of pharynx and anterior esoph-
agus. Cuticula transversely striated. Cordons (fig. 314 a and b)
long and becoming progressively wider until they practically cover
the entire body width; they are elaborate in structure, the outer

NEMATODE PARASITES OF BIRDS 251

edge markedly dentate: anastomosis at level of anterior fourth of
posterior esophagus or somewhat posterior to this. Lateral papillae
postcervical, very small, situated just posterior to anastomosis of
cordons.

Male 14 mm. long by 300/* wide. Pharynx 316/* long; anterior
esophagus 714/* long; posterior esophagus 3.4 mm. long. Cordons
extend 880/* along posterior esophagus, their total length 1.9 mm.
Caudal extremity (fig. 314/) tightly coiled, very difficult to unroll;
the present writer was unable to straighten it sufficiently to make
as detailed observation of the caudal structures as was desired.
Cloacal aperture about 340/* from tail end. Caudal alae thick, their
free edges curled ventrally. At least 8 pairs of caudal papillae;
there is a group of 3 pairs of preanal papillae fairly close together,
and there are at least 5 pairs of postanal papillae, of which 4 pairs
are large and pedunculated and the most posterior- pair is relatively
small and close to the tail end. Spicules unequal (fig. 314 g and h)
and dissimilar ; right spicule short and thick, consisting of a strongly
cutinized (or chitinized) part with sharply pointed free end, and
extending beyond this a hyaline portion less sharply pointed, the
spicule length being 191/* without the hyaline distal end, and 207ft
with it, and the maximum width being 58/*; the left spicule is more
than twice as long as the right and more slender, being 457/* long
by 29/* wide, with a blunt distal end. No gubernaculum.

Female 15 to 17.5 mm. long by 315/* wide. Pharynx 332 to 398/*
long; anterior esophagus 714 to 830//. long; posterior esophagus 3.2
to 3.8 mm. long. Cordons extend for 830/* to 1.1 mm. along the
posterior esophagus, their total length 1.9 to 2.7 mm., the former in
a specimen 15 mm. long, the latter in one 17.5 mm. long. Caudal
extremity (fig. 314c) conical, blunt. Anus 199/* from tail end.
Vulva near posterior end of body, just anterior to anus, 350/* from
tail end in small specimens and 448/* in the largest specimen; vulvar
lips (fig. 314</) large and projecting; diameter of body decreases
suddenly below T the posterior lip. Only 1 uterus and 1 ovary. Ove-
jector long and simple, composed of 2 parts, a long, large vestibule.
2.3 to 2.4 mm. long by 133/* wide, extending anteriorly in a straight
line from the vulva, sometimes with a twist about midway (iig.
314<?) ; a narrow sphincter 664/* long by 41.5/* wide connects this
with the uterus; no varnish gland (or trompe) is present if the
present writer interprets these structures correctly. The uterus at-
tains a width of 232/* and is closely packed with eggs; it extends in
a straight course anteriorly to near the posterior end of the eso-
phagus, and at that level it is considerably reduced in width: it
then turns posteriori}^ and soon joins the ovary which has a con-
voluted course. Eggs 36/* by 21/*.

252

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably involves intermediate stages
in other hosts (see E. uncinata, p. 246).

Distribution. — North America (United States (National Zoologi-
cal Park, Washington, D. C.) ).

Type specimens. — United States National Museum (Bureau of
Animal Industry) Helminthological Collection No. 18566; collected
by Dr. Leigh Giltner.

The female genitalia of this species as described above are very
different from those of E. uncinata as described by Seurat. The lat-
ter species has a short ovejector of the type of Dispharynx with a bi-

O-jmm

02 mm

o/mm

o-2/nm

Fig. 314. — Echinuria decorata. a, Anterior end; b, cordon; c, female tail; d,
vulva and vestibule ; e, vulva, ovejector, and uteeus ; f, male tail ; g and 7l,
spicules. Original

partite varnish gland (trompe) which connects with the 2 uteri. As
a transition form between the didelphic E. uncinata and the monodel-
phic E. decorata there is E. hargilae. Baylis and Daubney state
that this species may be considered monodelphic, as it has only 1
functional uterus, the other uterus being represented only by a
blind sac-like structure. . In E. decorata even this remnant has dis-
appeared. The present writer has been unable to find any refer-
ence to any other monodelphic form in the Spiruroidea; whenever
the number of uteri is given there are always two or more. The
absence of any reference to the character of the internal genitalia
in the great majority of forms, however, indicates the need for
closer observation and more extensive description.

NEMATODE PARASITES OF BIRDS 253

The position of the vulva and its prominent lips, and the mono-
delphic nature of this species relate E. decorata to E. hargilae,
whereas the length of the esophagus and the elaborate nature of the
cordons relate it to E. squamata.

ECHINURIA HARGILAE (Baylis and Daubney, 1923) Cram, 1927

Synonyms. — Acuaria {Echinuria) leptoptili (Gedoelst, 1916) of
Baylis and Daubney, 1922; Acuaria {Echinuria) hargilae Baylis and
Daubney, 1923.

Hosts. — Primary: Leptoptilus dubius; secondary: Unkown.

Location. — Not given.

Morphology. — Echinuria (p. 244) : Cuticula finely striated trans-
versely. Cordons 20/x wide, extending posteriorly for 950/*. to 1.1
mm., with a slight forward bend where they anastomose. No cuti-
cular spines. Cervical papillae prominent, slender, situated a little
posterior to the lateral bends of cordons.

Male 11 to 11.5 mm. long by 234/* wide. Caudal extremity coiled
several times in a spiral. Caudal alae 700/* long. Cloacal aperture

Fig. 315. — Echinuria hargilae. a, Anterior end; b, bight spicule. Aftbh
Baylis and Daubney, 1922

115^1 from tip of tail. Four pairs of preanal papillae and 5 pairs
of postanal papillae. Right spicule short, 190 to 210/* long, much
curved (fig. 3155), twisted and flanged; left spicule long, 650 to
675/* long, slender, gently curved, and not twisted.

Female 13 to 15 mm. long by 360/* wide. Vulva 170/* from tail
end, its anterior lip prominent. Vagina extends anteriorly 1.4 mm.
Baylis and Daubney say that this species may be considered mono-
delphic. They state : " The posterior set of organs * * * rep-
resented merely by a blind sac-like uterus which runs back to the
vicinity of the vulva. The anterior uterus runs forward to within
1.75 mm. of the anterior end of the body, the ovary commencing at
this point and running backwards." Anus 50/* from tail end. Eggs
28/* by 18/*.

Life history. — Unknown; probably involves intermediate stages
in other hosts (see E. uncinata, p. 246).

Distribution. — Asia (India (Calcutta)).

ECHINURIA HORRIDA (Rudolphi, 1809) Cram, 1927

Synonyms. — Strongylus horridus Rudolphi, 1809; Spiroptera gdl-
linulae Rudolphi, 1819; Spiroptera aculeata Creplin, 1825; Spirop-

254 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

tera horrida (Rudolphi, 1809) Diesing, 1851; Filaria spinifera
Schneider, 1866; Filaria aculeata (Creplin, 1825) Linstow, 1876;
Dispharagus aculeatus (Creplin, 1825) Stossich, 1891; Echinuria
spinifera (Schneider, 1866) Soloviev, 1912.

Hosts. — Primary : /Scolopax gallinula, Charadnus hiaticula, Tringa
variabilis, T. alpina; secondary: Unknown.

Location. — Proventricnlus.

Morphology.— Echinuria (p. 244) : Mouth with 2 lips, dorso- ven-
tral, according to Stossich, but probably lateral. From these there
originate on each side an arc formed of a row of spines (this does
not agree with Stossich's figure (fig. 317) ) and followed on each side
by a longitudinal row of strongly developed spines, inclined poste-
riorly, which diminish toward the tail and disappear just anterior to
its extremity, according to Stossich. According to Schneider (fig.
316), the cordons are conspicuous; directly under the curve where
they unite the body is surrounded by a cuticular fold, and 2 double
rows of spines originate at this point and extend to the caudal ex-
tremity, the 2 rows on each side starting together and then spreading
out in a bow.

Male 3.4 mm. long. Tail coiled in 3 or 4 spiral turns. Caudal
alae very slender. Four pairs of preanal and 4 pairs of postanal
pedunculated papillae. Large spicule arcuate, enlarged at its base
and hooked at the end ; small spicule bifurcate.

Female 4.02 to 11.25 mm. long. Tail slender, almost straight,
somewhat obtuse at the extremity. Anus close to end of tail. Vulva
very close to anus. Eggs 42/x by 23/a elliptical and thick-shelled.

Life history. — Unknown ; probably involves intermediate stages in
other host (see E. uncinata, p. 246).

Distribution. — Europe (Germany (Greifswald) ).

Schneider (1866) renamed this species, but gave no reasons for
doing so. Soloviev (1912) accepts the specific name spinifera
Schneider, but credits it to Rudolphi.

ECHINURIA LEPTOPTILI Gedoelst, 1916

Synonym. — Acuaiia {Echinuria) leptoptili (Gedoelst, 1916) Bay-
lis and Daubney, 1923, not of Baylis and Daubney, 1922 (see E.
hargilae, p 253).

Hosts. — Primary : Leptoptilus crumenifer; secondary : Unknown.

Location. — Stomach (Gizzard) .

Morphology. — Echinuria (p. 244) : Cuticula transversely striated.
Mouth with 2 lateral lips ending in a blunt point, each provided
with 2 lateral, symmetrical, pedunculated papillae. Cordons (fig.
318a) anastomosing; they are made up of small simple bands with
margins scalloped, or, according to Gendre. denticulate as in E.
squa??iata, projecting 16^ from the cuticula. Cervical papillae coni-

NEMATODE PARASITES OF BIRDS

255

cal, only slightly projecting, in the lateral lines a little posterior to
the curve formed by the anastomosing of the cordons.

Male 6.55 mm. long by 260/* wide. Tail (fig. 318 b) about 160//
long, about 1/40 of total body length. Cordons 610/x long. Caudal
alae membranous and transversely striated ; no subdivision into 2 con-
centric zones as in certain species of Acuaria. Four pairs of preanal
and 5 pairs of postanal papillae. Spicules (fig. 318 c and d) unequal
and dissimilar; the left 900//. long, flexible, the right 180//. long by
19//, wide, arcuate and robust.

Female 6 to 12 mm. long by 360 to 400//. wide. Cordons 900 to
970/x long. Gedoelst describes a membranous expansion of the euti-

Figs. 316-318. — E'chinuuia horrida. 316, Head end. After Schneider, 1S66. 317,
Head end. After Stossich, 1891. 31 S, Echinuria lepxoptili. a, Head end;
b, male tail; c, left spicule; d, right spicule (the small drawing showing

ORIFICE AT FREE END DILATED ON PRESSURE) ; C, FEMALE TAIL. AFTER GeNDKE. 1926

cula along the lateral lines of the caudal region, extending from a
point 50//. from the tail end and measuring 400/*, wide. Gendre does
not mention this; he states that the dilation of the body just ante-
rior to the vulva, as described by Gedoelst, is due to contraction of
the body. Anus 65 to 95//,, according to Gedoelst, or 50/t, according
to Gendre (fig. 318 e) , from tail end. Vulva 80 to 145//., according
to Gedoelst, or 210/*., according to Gendre, anterior to anus. Eggs
32/x by 20/*.

Life history. — Unknown; probably involves intermediate stages in
other hosts; (see E. unrinafa, p. 246).

Distribution. — Africa (Belgian Congo and Dahomey).

ECHINURIA LONGEORNATA (Molin. I860) Cram, 1927

Synonyms. — Dispharagus longeornatus Molin, 1860c; Spiroptera
ardeae-maguari* Molin, 1860; Acuaria longeornata (Molin. 1860)

e Catalogue of Vienna Museum.

256 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Railliet, Henry, and Sisoff, 1912; Hainannia longeomata (Molin,
1860) Stiles and Hassall, 1920.

Hosts. — Primary: Ciconia maguari; secondary: Unknown.

L ocation. — Proventriculus.

Morphology. — Echinuria (p. 244) : Mouth with 2 small inconspicu-
ous lips. Cordons very long, briefly recurrent (brwi regredientes) ,
according to Molin: Gendre (1919) says this may be a retraction due
to fixation, a thing he has seen in E. leptoptili; if the cordons were
truly recurrent the species would belong in Synhimantus ; the cor-
dons anastomose in pairs. Cuticula with thick transverse striations.

Male unknown.

Female 8 mm. long by 200/* wide. Anus near caudal apex. Vulva
small, anterior to anus.

Life history. — Unknown ; probably involves intermediate stages in
other hosts; (see E. uncinata, p. 246).

Distribution. — South America (Brazil).

ECHINURIA LONGEVAGINATA (Molin, 1860) Cram, 1927

Synonyms. — Dispharagus longevaginatus Molin, 1860c; Spirop-
tera ciconiae-maguari' 1 Molin, 1860; Acuaria longevaginata (Molin,
1860) Railliet, Henry, and Sisoff, 1912; Synhimantus longevaginata
(Molin, 1860) Skrjabin, 1924.

Hosts. — Primary : Ciconia maguari; secondary : Unknown.

Location. — Esophagus.

Morphology. — Echinuria (p. 244) : Mouth with 2 projecting tri-
angular, lateral lips (fig. 319a.). Cuticula transversely striated.
Anterior extremity markedly attenuated. Cordons long and
straight, anastomosing. According to Molin, the cordons are briefly
recurrent here as in E. longeomata; they are regarded in this case,
as in that, as not recurrent for the reasons given in discussing that
species.

Male 6 mm. long by 100/* wide. Posterior extremity twisted spir-
ally, excavated ventrally, and with acute apex. Caudal alae (fig.
319 b) long and wide, with thick edges. Four pairs of preanal and
5 pairs of postanal papillae, very small. Right spicule short and
thick; left spicule six times as long as right, slender and alate.

Female 7 to 8 mm. long by 200/i wide. Tail conical, with blunt
tip. Anus near end of tail. Vulva anterior to anus.

Life history. — Unknown; probably involves intermediate stages
in other hosts, (see E. uncinata, p. 246).

Distribution.. — South America (Brazil).

7 Catalogue of Vienna Museum.

NEMATODE PARASITES OF BIRDS 257

ECHINURIA PHOENICOPTERI (Seurat, 1916) Seurat, 1916

Synonym. — Acuaria (Hamannw) phoenicopteri Seurat, 1916;
Hamannia phoenicopteri (Seurat, 1916) Stiles and Hassall, 1920.

Hosts. — Primary: Phoenicopterus roseus; secondary: Unknown.

Location. — In mucosa of proventriculus.

Morphology. — EcMnwria (p. 244) : Anterior extremity (fig. 320 a
and h) with 2 cordons on the lateral and ventral surfaces, the sym-
metry in the anterior body being bilateral, but not radial, in this
respect ; the cordons anastomose a little below the nerve ring. There
are also 4 rows of spines, the 2 latero-dorsal rows originating 60//,
from the head end, and the 2 latero-ventral rows originating poste-
rior to these at the point where the cordons turn.

Male unknown.

Female known only as immature adult at end of fourth larval
stage. This adult 2.2 mm. long by 65/x wide, as viewed through the

Figs. 319-320. — 319, Echixuria longevaginata. a, Head ; b, male tail. After
Drasche, 1884. 320, Echinuria phoenicopteri. Immature female, a and
b, Head end; c, tail; d, vulva and ovbjbctob. After Ski rat, 191(5

cuticula of larva. Vulva not salient, situated at anterior fifth of
body length. Ovejector (fig. 320 d) directed posteriorly; the reser-
voir large, the sphincter short. Tail (fig. 320 c) digitiform, rounded
at tip.

Larva, fourth stage, with slender body, attenuated at both ex-
tremities. Cuticula with fine transverse striations. Mouth with 2
lateral lips, each projecting as a tooth.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa (Algeria ) .

ECHINURIA SQUAMATA (Linstow. 1883) Cram, 1927

Synonyms. — Filaria squamata Linstow, 1883; Dixph<ir<i<pis squa-
matus (Linstow, 1883) Stossich, 1891; Acuaria squamata (Linstow,

258

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

1883) Kailliet, Henry, and Sisoff, 1912; Hannannia squamata (Lins-
tow, 1883) Stiles and Hassall, 1920.

Hosts. — Primary: Phalacrocorax carbo, Carbo cormoranus; sec-
ondary: Unknown.

Location. — Intestine.

Morphology. — Echinuria (p. 244) : Mouth with 2 rounded hyaline
lateral lips, only slightly projecting. Esophagus one-fourth of total
body length. Tail 1/288 of body length. Body attenuated anteriorly
and thickened posteriorly. In the submedian lines are cordons
(fig. 321« and b) made up of peculiar ring-like bands which appear
on the margin as posteriorly directed scales ; the cordons anastomose
1 mm. from the head end; at each point of anastomosis there is a
large 3-pointed cervical papilla from which there extends posteriorly

Fig. 321. — Echinuria squamata. a, Anterior end; b, detail of cordon. After

Linstow, 1883

a simple straight cordon, according to Linstow. There is a ques-
tion as to whether this simple straight cordon as described by Lins-
tow may not be a lateral ala or a prominent lateral line; he states
that these cordons are in the dorsal and ventral lines, but as they
proceed from the lateral cervical papillae they should be in the
lateral lines.

Male unknown.

Female 24 mm. long by 72G> wide. Eggs 39/x by 24/*, thick-shelled.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Asia (Turkestan ) .

Genus RUSGUNIELLA Seurat, 1919a

Synonyms. — Filaria Mueller, 1787, part; Acuaria Bremser, 1811,
part; Spiroptera Rudolphi, 1819, part; Dispharagus Dujardin, 1845,
part.

Gene'i te diagnosis-. — Acuariinae (p. 211) : Body elongate, relatively
slender. Cephalic region with 2 cutaneous ornamentations, crescent

NEMATODE PARASITES OF BIRDS 259

shaped, originating at the angles of insertion of the buccal lips and
extending on the lateral surfaces as 2 epaulets. Two lateral alae
sometimes present, originating slightly posterior to the cordons
(epaulets). Precervical papillae set in the alae when alae are
present. Excretory pore ventral, posterior to nerve ring. .Mouth
with 2 upright lateral lips, each with a pair of large sessile papillae
near their angles of insertion. Buccal cavity or pharynx tubular,
slightly widened anteriorly. Esophagus clearly differentiated into
an anterior, transparent, muscular portion, surrounded anteriorly
by the nerve ring, and a posterior, opaque, glandular portion. Vulva
with projecting lips a short distance anterior to middle of body.
Ovejector cylindrical, directed anteriorly ; uteri and ovaries opposed.
Male unknown. Parasitic in esophagus (and gizzard?) of Chara-
driides, Longipennes, and Pygopodes.

Type species. — Rusguniella elongata (Rudolphi, 1819) Seurat,
1919a.

KEY TO SPECIES OF RUSGUNIELLA

1. Females 4 to 6 mm. long; female tail 1/34 of total body length; no lateral

alae described or figured Rusguniella vanelli, p. 260

Females 24 to 40 mm. long; female tail 1/102 of total body length; lateral

alae present Rusguniella elongata, p. 259

RUSGUNIELLA ELONGATA (Rudolphi, 1819) Seurat, 1919a

Synonyms. — Spiroptera elongata Rudolphi, 1819; Dispharagus
species Wedl, 1856; Dispharagus elongatus (Rudolphi, 1819) Molin,
1860; Filaria elongata (Rudolphi, 1819) Schneider, 1866; Acuaria
elongata (Rudolphi, 1819) Railliet, Henry, and Sisoff, 1912.

Host*. — Primary: ILydrochelidon nigra, Sterna nigra, Podlceps
nigricollis, European gull ; secondary : Unknown.

Location. — Under corneous tunic of gizzard.

Morphology. — Rusguniella (p. 258) : Characters of the genus.

Male unknown.

Female 24 to 40 mm. long; slender. Cuticula thick, finely striated
transversely. Lateral alae (fig. 322 a and b) originate immediately
posterior to cordons (epaulets). Lateral or precervical papillae
asymmetrical; in a specimen 28.2 mm. long by 312/x wide the left
papilla 182/*, the right 192/t, from head end; the origin of lateral alae
105/a, excretory pore 456/*, the vulva 13 mm. from head end. Buccal
cavity or pharynx 135/* long, anterior esophagus 865/* long, posterior
esophagus 3.1 mm. long, tail 275/* long. Vulva with slightly salient
lips. Cuticular ovejector tubular, 450/x long, directed anteriorly.
Eggs oval, 38/i by 24/*, thick-shelled, embryonated when oviposited.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe (Vienna Museum) ami Africa (Algeria).

260 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

RUSGUNIELLA VANELLI (Rudolphi, 1819) Seurat, 1919a

Synonyms. — Spiroptera vanelli Rudolphi, 1819; Acuaria vanelli
(Rudolphi, 1819) Poche, 1912.

Hosts. — Primary : Trvngae vanelli, Vanellus cristatus; secondary :
Unknown.

Location. — Intestine and between tunics of gizzard.

Morphology. — Rusguniella (p. 258) : Cuticula with fine transverse
striations indicated by rows of glittering granules. Mouth with 2
lips with 2 blunt teeth (or projections?) and with pedunculated
papillae. The crescent-shaped cordons (fig. 323) or epaulets
originate at the teeth. Precervical papillae immediately posterior
to cordons and directed anteriorly.

Male unknown.

Female 4 to 6 mm. long, according to Diesing (Linstow's speci-
men 5.3 mm.), by 200^ wide. Esophagus 10/68, tail 1/34 of total

Figs. 322-323. — 322, Rusguniella elongata. Anterior end. a, Lateral view ; &,.
ventral view. After Seurat, 1919. 323, Rusguniella vanelli. Anterior end,
ventral view. After Linstow, 1884

body length. Tail rounded. No lateral alae mentioned or figured.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Vienna Museum).

Genus SCIADIOCARA Skrjabin, 1916b

/Synonyms. — /Spiroptera Rudolphi, 1819, part; Schistorophus
Railliet, 1916, part.

Generic diagnosis.— Acuariinae (p. 211) : Head with 2 small lat-
eral papillae and 4 submedian papillae. Posterior to the lips there
are 4 peculiar, delicate, hemispherical alae (the "Laeppchen" of
Drasche), which are arranged in pairs on each side. Caudal ex-
tremity of male twisted in spiral and provided with alae and with
6 pairs of preanal and several pairs of postanal papillae. Spicules
unequal, the smaller provided with a canal through which the larger

NEMATODE PARASITES OF BIRDS 261

glides, so that the smaller acts as a gubernaculum. Vulva almost
in middle of body. Eggs thick-shelled, containing embryos when
oviposited.

Parasitic under the lining tunic of the gizzard.

Type species. — Sciadiocara umbellifera (Molin, 18G0) Skrjabin,
1916&.

This genus is placed in the Acuariinae on the nature of the ce-
phalic appendages and of the male caudal structures. The append-
ages of S. secunda are particularly like the cordons of the acuarids.
According to Seurat, the cuticular epaulets of Seuratia approach
those of Scladiocara.

KEY TO SPECIES OF SCIADIOCARA

1. Cephalic appendages adhere to the cuticular surface of the body, their edges

forming typical cordons; in Corvus monedula Sciadiocara secunda, p. 262.

Cephalic appendages not adhering to the cuticular surface of the body, but
projecting so as to form a sharp angle with the body ; in hosts other than
above Sciadiocara umbellifera, p. 201.

SCIADIOCARA UMBELLIFERA (Molin, 1860) Skrjabin, 1916b

Synonyms. — Spiroptera umbellifera Molin, 1860&; Spiroptera
iantali rubri 8 Molin, 1860; Spiroptera totani 8 Molin, 1860; Schis-
tor'ophus umbellifera (Molin, 1860) Railliet, 1916.

Hosts. — Primary: Ibis rubra, Tot anus melanoleucus, Scolopax
major; secondary : Unknown.

Location. — Between tunics of the gizzard.

Morphology. — Sciadiocara (p. 260) : Mouth (fig. 324 a and b) with
2 small, conical, lateral lips, posterior to which are 4 submedian
papillae, 2 on each side. Behind these are the cephalic appendages
characteristic of the genus, 4 membranous hemispherical alae directed
backward and forming a sharp angle with the body; in a face view
these organs are seen to form 2 pairs, being united in twos in the
lateral fields.

Male 6 to 6.4 mm. long by 50 to 100/x wide. Two caudal alae (fig.
325 a) with 6 pairs of preanal and 5 pairs of postanal papillae. The
most posterior pair of papillae arc much the smallest. Of the pre-
anal papillae, the most anterior pair stand alone, as do the most
posterior pair; the second and third pairs form one group and the
third and fourth pairs another group. Short spicule peculiar, its
dorsal surface recurved to form a canal ; it is 81^ long with a maxi-
mum diameter of 22/x. The other spicule is 330/* long by 7.5fj. wide,
its posterior end is recurved, and it glides in the dorsal canal of the
smaller one; the smaller spicule, therefore, functions as a guber-
naculum and might be regarded as such.

8 Catalogue of Vienna Museum.

262

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 9 to 10 mm. long by 25 to 102/* wide. Pharynx 37;/. long.
The characteristic caudal extremity (fig. 325 c) consists of a caudal
appendage 85/a long rounded in an obtuse manner and attached to
the body at an obtuse angle. Vulva (fig. 3256) slightly posterior
to the middle of body, 4.08 mm. from tail end. Eggs 44 to 48/x long
by 30yu wide, and thick shelled.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil) and Asia (Russian Turke-
stan).

SCIADIOCARA SECUNDA Skrjabin, 1916b

Hosts. — Primary : Corvus monedula; secondary
Location. — Between tunics of gizzard.

Unknown.

Figs. 324-325. — Sciadiocaea umbellifeba. 324, Head, a, Dobso-ventbal view ; b,
froxt view. After Drasche, 1884. 325, a, Male tail; b, vulva; c, female tail.
Afteb Skrjabin*, 1916

Morphology. — Sciadiocara (p. 260) : In the only available descrip-
tion of this parasite, that by Skrjabin (1916), there is only the state-
ment that the cephalic appendages adhere to the cuticular surface
and that the edges of these appendages are the typical cords corre-
sponding to the cordons of the cervical region of the acuarids.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Asia (Russian Turkestan).

Genus SEURATIA Skrjabin, 1916d

Synonym. — Rictularia Froelich, 1802, part; Acuaria Bremser,
1811, part; Spiroptera Rudolphi, 1819, part; Gnathostoma Owen,
1836, part ; Prionostemma Gendre, 1920&.

Generic diagnosis. — Acuariinae (p. 211) : The cephalic region orna-
mented with short cordons in the form of epaulets, curved as a
handle on the lateral surfaces, situated on the swollen cuticle, and
with free edge markedly dentate. Posterior to epaulets are a pair

NEMATODE PARASITES OF BIRDS 263

of enormous tricuspid hooks. In addition the cuticula is provided
with 2 double rows of hooks, the points of which are directed
backward. Mouth with 2 lateral lips. Buccal cavity tubular. Vulva
directly anterior to middle of body. Uteri divergent. Two unequal
spicules. Four pairs of preanal papillae.

Parasitic in the digestive tract of birds.

Type species. — Seuratia shipleyi (Stossich, 1900) Skrjabin, 1916r/.

KEY TO SPECIES OF SEUKATIA

1. Female 13 to 14 mm. long; from Procellaria anglorum.

Seuratia procellariae, p. 2G3.
Female 21.5 to 35 mm. long; from hosts other than above.

Seuratia shipleyi, p. 263.

SEURATIA PROCELLARIAE (Diesing, 1851) Cram, 1927

Synonyms. — Spiroptera procellariae anglorum Diesing, L851
(based on Bellingham, 1844) ; Spiroptera procellariae Diesing, 1851;
Prionostemma procellariae (Diesing, 1851) Gendre, 19205.

Hosts. — Primary : Procellaria anglorum, secondary : Unknown.

Location. — Attached in crop.

Morphology. — /Seuratia (p. 262) : Mouth orbicular, projecting, sur-
rounded by 4 papillae. Neck armed with recurved hooks and the
anterior third of the body also provided with 4 rows of smaller
spines.

Male unknown.

Female 13 to 14 mm. long. Body thickened posteriorly, very
translucent, the course of the intestine, which enlarges posteriorly,
visible through body wall.

Life history. — Unknown : probably involves intermediate stages
in other hosts.

Distribution. — Europe (Ireland ) .

Seurat states that his new r species, Acuaria pet 'agica (=S. shipleyi),
is close to this species in the character of its epaulets, the cutaneous
spines of the esophageal region, the median position of the vulva.
and the elongation of the cuticular ovejectors.

SEURATIA SHIPLEYI (Stossich, 1900) Skrjabin, 1916d

Synonym. — Gnathostoma shipleyi Stossich, 1900; Rictularia para-
donna Linstow, 19035/ Rictularia shipleyi (Stossich, 1900) Johnston,
1912; Acuaria pelagica Seurat, 1910(2/ Prionostemma pelagicum
(Seurat, 1916) Gendre, 19206/ Prionostemma shipleyi (Stossich,
1900) Gendre, 19206.

Hosts. — Primary: Diomedea exulans, Daption capensis, Larus
canus, Pufflnus kuhli; secondary: Unknown.

Location. — Proventri cuius.

264

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Seuratia (p. 262) : Mouth terminal with 2 trilobed
lips, each lobe with a minute papilla at its extremity. Cuticle at
anterior end of body (fig. 326 a and b) swollen to form a collar
with 2 large disks, posteriorly bilobed, and the free margin with
strong spines. Large tricuspid cervical papillae a little behind the
disks; from these papillae originate 4 longitudinal rows of spines,
extending to about the middle of the body.

Male 15 mm. long. Caudal extremity (fig. 326 c) spirally coiled.
Caudal alae only very slightly developed. Caudal papillae very
delicate, 17 in number; of these, 4 pairs preanal, 4 pairs postanal,
and 1 unpaired at tip of tail, the latter a multicuspidate affair fig-
ured by Stossich in a way resembling the spiny knob of the tail of

Fig. 326. — Seuratia shipleyi. a, Dorso-ventral and b, lateral view of

ANTERIOR END; C, MALE TAIL; d, FEMALE TAIL; e, EGG. AFTER STOSSICH, 1900

the larvae of many spirurids, a character maintained into adult life
in Desmidocerca aerophila (p. 209).

Female 21.5 to 32 mm. (Seurat) or 35 mm. (Stossich) long by
500/* wide. Tail 180/* long. Vulva 14.6 mm. from head end in worm
32 mm. long. Ovejector (fig. 327) very long, 1.8 mm. Eggs (fig.
326e) small, ovoid, numerous, and embryonated at oviposition.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution, — Western Pacific, Australia and Africa (Algeria).

Genus STREPTOCARA Railliet, Henry, and Sisoff, 1912

Generic diagnosis. — Acuariinae (p. 211) : Mouth with 2 conical
projecting lips, originating from a depression, the margin of which
forms a denticulated collar. Esophagus short, swollen, muscular.
Two cervical papillae in form of a crescent with numerous teeth.

NEMATODE PARASITES OF BIRDS

265

Male with 2 membranous caudal alae; 4 pairs of preanal, 5 to G
pairs of postanal papillae, all pedunculate. Two very unequal
spicules, the longer ending in a recurved barb.

Female with vulva usually a little behind the middle of body,
occasionally more anterior or more posterior. Eggs embryonated
when deposited.

Parasitic under the cuticle of gizzard of birds.

Type-species. — Streptocara pectinifera (Neumann, 1900) Skrjabin,
19165.

Fig. 327. — Seuratia shipleyi. a and h, Proximal region of ovejector; c, distal

REGION OF OVEJECTOR. AFTER SEURAT, 1916
KEY TO SFECIES OF STREPTOCARA

1. Cervical papillae with 3 points 2.

Cervical papillae with 5 or more points 6.

2. Female not over 10 mm. long 3.

Female 16 mm. or longer 5.

3. Cervical papillae over 4(XV from anterior extremity ; eggs 27 > long.

Streptocara triaenucha, p. 271.

Cervical papillae not more than 140> from anterior extremity ; eggs 39/*

long 4.

4. Vulva anterior to middle of body (1/2.6 of body length) ; 5 pairs of postanal

papillae; spicules 140/u. and 600> long Streptocara cirrohamata, p. -V>C>.

Vulva in posterior region of body (500ju from posterior end) ; 3 pairs of
postanal papillae; spicules 68^ and 237/n long__ Streptocara decora, p. 269.

5. Male 5.5 mm. long; 6 pairs of postanal papillae; spicules 110> and 400/i

long; from Colymbus arcticus and La run ridibundus.

Streptocara tridentata, p. 271.
Insufficiently described. From Fulmaris glacialis.

Streptocara stellae-polaris, p. L'TO.

6. Male 11 mm. long; female said to be smaller than male; spicules 250^ and

490/* long Streptocara recta, p. lm;'.>.

Male not over 5.5 mm. long; female 6 to 15 mm. long; spicules not over 88m
and 396/a long 7.

7. Five pairs of postanal papillae; long spicule measures 12«>-">m ; in (lalliist gallus.

Streptocara pectinifera, p. 266.

Six pairs of postanal papillae; long spicule measures 314 to 396,u; in other

hosts than above 8.

8. Marked transverse cuticular striations between the collar and cervical papil

lae, and a swelling in the region of the latter. In Charadriiformes.

Streptocara crassicauda charadrii, p. L'in
Marked transverse striations and swelling absent In Antcriformes.

Streptocara crassicauda, p. L'<;7.

266 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

This key does not include Streptocara, species Baylis, 19196, a
single female specimen of which was found in Ui^ia grylle in Yukan-
ski (Arctic Eussia). Baylis states that this is probably /Strepto-
cara pectinifera.

STREPTOCARA PECTINIFERA (Neumann, 1900) Skrjabin, 1916b

Synonym. — Spiroptera pectinifera Neumann, 1900.

Hosts. — Primary : Gallus gallus and Numidameleagris ; secondary :
Unknown, probably arthropod or other small invertebrates.

Location. — In the mucosa of the gizzard.

Morphology. — Streptocara (p. 264) : Whitish worms, attenuated
anteriorly. Cuticle transversely striated. Mouth (fig. 328 a) with 2
conical or triangular lips, the lip extremities slightly curved outward
and terminating in a dentiform process ; they are limited posteriorly
by a denticulated collarette and each bears 2 papillae. About 55/*
posterior to head end, or 42 and 36/* according to Gedoelst and Liege-
ois, are 2 cervical papillae (fig. 328fr) with a semicircular anterior
border and a corresponding posterior border bearing 6 or 7 teeth, ac-
cording to most authors, or 5 to 6, according to Gedoelst and Liegeois.

Male 4 to 5.2 mm. long by 150 to 176/* wide. Esophagus 2 mm.
long. The tail (fig. 328c) has 4 pairs of preanal, the second and
fourth being the longest, and 5 pairs of postanal papillae regularly
diminishing in size from first to fifth. Two very unequal spicules,
the left 265 to 300/* long, ending in recurved barb, the right 75 to 88//.
long, thick and barbed. Caudal alae, 200/* long, unite anteriorly.

Female 6.5 to 9.6 mm. long by 275 to 280/* wide. Esophagus 2.4
mm. long. Vulva slightly posterior to equator of body, about 5.47
mm. from head end. Eggs 33/* long by 20/* wide, according to most
authors, or 37 to 39/* long by 20 to 21/* wide, according to Gedoelst
and Liegeois, containing embryos when deposited.

Life history. — Unknown; probably has intermediate stages in
arthropods or other small invertebrates, which probably eat the eggs
passed in the droppings and in which infective larvae develop, and
such intermediate hosts being eaten by birds, the larval worms
mature.

Distribution. — Europe (France, Belgium, and perhaps Italy and
Arctic Russia).

STREPTOCARA CIRROHAMATA (Linstow, 1888) Skrjabin, 1916b

Synonyms. — Filaria (Spiroptera) cirrohmnata Linstow, 1888;
Spiroptera cirrohamata Stossich, 1897.

Hosts. — Primary: Phalacrocorax verrucosus; secondary: Un-
known, see S. pectinifera, p. 266.

Location. — " Stomach."

NEMATODE PARASITES OF BIRDS

267

Morphology. — Streptocara (p. 2G4) : Head (fig. 329a) with 2 coni-
cal lips and rudimentary neck frills. Cervical papillae 140ft, from
head end, tricuspid.

Male 7.58 mm. long by 250/* wide. Esophagus somewhat less than
1/2 the body length ; tail 1/20 of body length. Posterior extremity
(fig. 3296) rounded; caudal alae broad. Four pairs of preanal, 5
pairs of postanal pedunculated papillae. Spicules very unequal, the
longer 600/n, bearing a small terminal process projecting at right
angles, the shorter spicule 14G> long.

Female 9.72 mm. long by 350/x wide. Esophagus 1/2.6, tail 1/81 of
body length. Vulva just anterior to end of esophagus. Eggs 39/x
long by 19ju, wide, with very thick shell.

Figs. .".28-329. — 328. Streptocara pectinifera. a, Head end; b,

CERVICAL. PAPILLAE ; C, MALE TAIL.. AFTER NEUMANN, 1900. 329,

Streptocaba CIEROHaMata. a, Head; b, male tail. After Lin-

siuiv, 1888

Life history. — Unknown; see S. pectinifera, p. 266.
Distribution. — Not given. Collected on Challenger expedition.

STREPTOCARA CRASSICAUDA (Creplin, 1829) Skrjabin, 1916b

Synonyms. — Spiroptera crassicauda Creplin, 1829; Disp harag us
crassicauda (Creplin, 1829) Molin, L860; Streptocara crassicauda
anseri Skrjabin, 1916.

Hosts. — Primary: Anas clangula, A. glacialis, A. tadorna, A. fusca,
A. bosclias, Alca torda, Bernicla sandwic hen-sis, Bnwephala cZamgula,
Colymbus arcticus, C rufovidgaris, C. septentrional is, Harelda <jl<i-
cialis, Mergus serrator, M. merganser, Nyroca clangula, Oidemda
fusca, Tadorna tadonia; secondary: Unknown: see S. pectimfera,
p. 266.

L ocation . — Gizza rd.

268

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology.— Streptocara (p. 264) : Head (fig. 330 a) with 2
conical salient lips, posterior to which is a finely denticulated col-
larette. Cervical papillae 33/* from head end, with 5 to 9 points of
different sizes.

Male 3 to 5.5 mm. long by 200/* wide. Esophagus 1.4 mm. long.
Four pairs of preanal, 6 pairs of postanal papillae. Spicules un-
equal, the longer 314 to 396/*, the shorter 81 to 88/* long.

Female 5.6 to 15.1 mm. long by 150 to 360/* wide. Buccal cavity
22/*, esophagus 2 mm. long. Tail end (fig. 330 o) rounded, the anus
25/* from end. Vulva (fig. 330 c) just slightly posterior to middle

*<a&

Figs. 330-331. — 330, Streptocara crassicauda. a, Head; b, tail op female;
c, vulva. After Skrjabin, 1916. 331, Streptocara crassicadda chara-
drii. a, Head ; b, cervical papilla. After Skrjabin, 1916

of body, according to Skrjabin. Eggs 26 to 37/* long by 17 to 22/*
wide.

Life history. — Unknown; see S. pectinifera, p. 266.

Distribution. — Europe and Asia (Russian Turkestan).

Skrjabin proposes the variety name anseris for this form to dis-
tinguish it from S. crassicauda charadrii, but this is not allowable
under the rules of nomenclature. He uses the form S. c. anseri but
does not make the combination with anseris.

STREPTOCARA CRASSICAUDA CHARADRII Skrjabin. 1916b

Host. — Primary: Vanellus cristatus; secondary: Unknown, see
S. pectinifera, p. 266.

Location. — Under cuticle of gizzard.

Morphology.— Streptocara (p. 264) : The differences between this
and Streptocara crassicauda (Creplin, 1829) are (1) different host
order, this being in the Charadriiformes and the original species in
the Anseriformes and (2) the cephalic ornamentation: Between the

NEMATODE PARASITES OF BIRDS 269

collar and the cervical papillae in this variety there are very marked
transverse cuticular striations (fig. 331) and a swelling in the region
of the cervical papillae.

Life history. — Unknown ; see S. pectinifera, p. 26G.

Distribution. — Asia (Russian Turkestan).

STREPTOCARA DECORA (Dujardin, 1845) Skrjabin. 1916b

Synonyms. — Dispharagus decorus Dujardin, 1845; Histiocephalus
decorus (Dujardin, 1845) Diesing, 1851; Yseria decora (Dujardin,
1845) Gedoelst, 1919; Prionostemma decorum (Dujardin, 1845)
Gendre, 19206.

Host. — Primary: Alcedo ispida; secondary: Unknown, see S.
pectinifera, p. 266.

Location. — Between the tunics of the gizzard.

Morphology. — Streptocara (p. 264) : Collar (fig. 332 a) in form
of denticulate cordons which surround circularly 2 lateral convex
lobes. Cuticle transversely striated, capable of being swollen be-
hind the head, according to Dujardin. Cervical papillae tricuspid,
situated 120/* posterior to head end.

Male 3.6 mm. long by 110/* wide. Cloacal aperture 130/* from
tail end (fig. 3326). Four pairs of preanal, 3pairsof postanal papil-
lae. Spicules very unequal, the longer measuring 237/« long by 8.3/*
wide, the shorter 68/* long by 14/* wide.

Female 8 mm. long by 200/* wide. Anus 130/* from tail end. Vulva
500/* from tail end. Eggs peculiar; they are 39/* long, flattened in
one direction where they are no wider than 23/* whereas they are
enlarged in the other direction to 31.5/* by two lateral and opposed
rows of 3 to 5 rounded tubercles.

Life history. — Unknown ; see S. pectinifera, p. 266.

Distribution. — France.

Gedoelst placed this species in his new genus Yseria but he evi-
dently had not seen Skrjabin's paper at that time. Subsequently
Gendre stated that the head ornamentations differed from those of
Yseria coronata and Y. calif omica and transferred it to his new
genus Prionostemma, this genus, however, proving to be synonymous
with Seuratia. Although the denticulated collar of S. decora is
somewhat different, in appearance from the majority of species of
Streptocara, it is not as highly developed as in Seuratia and the
body does not have spines as in the latter genus; the present writer
has consequently left it in Streptocara as placed by Skrjabin.

STREPTOCARA RECTA (Linstow. 1879) Skrjabin. 1916b

Synonyms. — FUaria recta Linstow 1879; Spiroptera recta (Lins-
tow, 1879) Mueller, 1897.

270

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Primary : Colymbus cristatus and Podiceps cristatus/ sec-
ondary: Unknown; see S. pectinifera, p. 266.

Location. — Germany.

Morphology. — Streptocara (p. 264) : Head (fig. 333 a and b) with
2 lips and with finely denticulated but inconspicuous collar, not far
posterior to which are 2 cervical papillae, pluridentate (as figured
by Mueller they have at least 5 points).

Male 11 mm. long by 360/x wide. Four pairs of large pedunculated
preanal papillae (fig. 333<?) ; 7 pairs of postanal papillae, of which
5 are large and lateral, the other 2 pairs small, ventral, situated be-

Figs. 332-333. — 332, Streptocara decora, a, Anterior end ; ~b, male
tail. After Dujardin, 1845. 333, Streptocara recta, a, Dorso-
ventral and b, lateral view of head ; c, male tail ; d, spicules.
After Mueller, 1897

tween the papillae of the most posterior lateral pair. Spicules (fig.
333c?) unequal and dissimilar, the one 490/a long, slender, with a
wide lateral membrane in its distal half, the other 250/* long, strongly
developed, " cork-screw " shape, according to Mueller.

Female smaller than male, according to Mueller (neither he or
Linstow give the length). Tail end rounded. Vulva not salient,
dividing the body length in ratio of T : 4. Size of eggs not given.

Life history. — Unknown; see ,6'. pectinifera, p. 266.

Distribution. — Europe ( Germany ) .

STREPTOCARA STELLAE-POLARIS (Parona, 1901) Skrjabin 191Gb

Synonyms. — Histiocephalus stellae-poUaris Parona, 1901; Yseria
stellae-polaris (Parona, 1901) Gedoelst, 1919.

Llost. — Primary: Fulmarus glacialis; secondary: Unknown; see
$. pectinifera, p. 266.

Location. — Not given.

NEMATODE PARASITES OF BIRDS 271

Morphology. — Streptocara (p. 2G4) : Head with 2 large lips and
with a dilation in manner of a hood with denticulate margin. A
tricuspid process a little posterior to this dilation.

Male unknown.

Female 16 mm. long. Anus at caudal extremity, the latter obtuse.
Vulva at about middle of body. Eggs oval, containing an embryo
when oviposited.

Life history. — Unknown ; see S. pectinifera, p. 266.

D istribution. — Arctic region .

There are no figures of this species and the description is scant.
It appears questionable as to whether the species belongs in Strepto-
cara as placed by Skrjabin, or in Yseria as placed by Gedoelst. The
latter author had apparently not seen Skrjabin's assignment; it is
therefore being left for the present in Streptocara by the present
writer.

STREPTOCARA TRIAENUCHA (Wright. 1879) Skrjabin. 1916 b

Synonyms. — Filaria triaenucha Wright, 1879; Acuaria triaenucha
(Wright, 1879) Ward, 1918.

Host. — Primary : Botaurus minor; secondary : Unknown ; see S.
pectinifera, p. 266.

Location. — Proventriculus.

Morphology. — Streptocara (p. 264) : Cuticle densely striated. A
cervical frill present, the tops of the lateral loops being 180/x from
anterior end; it extends posteriorly on the neck for a distance of
405/*. Cervical papillae (fig. 331) tricuspid, the root being 60/*
posterior to the end of the frill ; the papilla itself measures 6G> from
the root to the end of the median fork.

Male unknown.

Female 10 mm. long by 430/* wide. Tail ending in a short rounded
conical projection. Eggs 27/* long by 18/* wide.

Life history. — Unknown.

Distribution. — North America (Canada).

The description of the cervical frill and especially its extent does
not seem to correspond with the species of the genus Streptocara.
It is possible that this species belongs in Yseria, where the cervical
hood or frill is larger and the cervical papillae tridentate. However,
as there are no figures of the head of this species, and as the de-
scription is incomplete, the transfer to another genus is inadvisable.

STREPTOCARA TRIDENTATA (Linstow, 1877) Skrjabin, 191Gb

Synonyms. — Filaria tridentata Linstow, 1877 a and 1>; Spiroptera
tridentata (Linstow, 1877) Neumann, 1900.

Hosts. — Primary: Colymbus arcticus, Larus ridibundus; second-
ary: Unknown; see S. pectinifera, p. 266.

Location. — Esophagus and intestine.
3G12— 27 19

272 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology. — Streptocara (p. 264) : Mouth with 2 short cone-
shaped lips. Linstow states that he could find no cervical frill but
that the mouth is surrounded by indefinite radiating chitinous folds.
Cervical papillae tricuspid (fig. 335&).

Male 5.5 mm. long by 13G> wide. Tricuspid papillae 200/x from
head end. Anterior part of esophagus 140/a long; total esophagus
62G> or 1/9 of total body length ; tail 1/32 of body length. Caudal alae
(fig. 335a) fairly wide. Four pairs of preanal, 6 pairs of postanal
papillae; of the latter 5 pairs are lateral and 1 pair ventral. Spicules
unequal, the left 400//, long, the right 110/i long, the latter with a
bulbous swelling at its proximal end.

Female 16.4 mm. long by 220/a wide. Cervical papillae 16G> from
head end. Esophagus 2.4 mm., tail 300/* long. Vulva divides body

336

33S^^
339

Figs. 334-336. — 334, Streptocara triaenucha. Cervical papilla. After
Wright, 1897. 335, Streptocara tridentata. a, Male tail. After Linstow,
1909. b, Cervical papilla. After Linstow, 1877. 336, Diagram of cordons
(as found in Synhimantus) as they would appear if spread out horizon-
tally. After Schneider, 1866

length in ratio of 54:41. Eggs, according to Skrjabin's tabulated
description, 36/* long by 18/*, wide.

Life history. — Unknown ; see S. pectinifera, p. 266.

Distribution. — Europe.

Genus SYNHIMANTUS Railliet, Henry, and Sisoff, 1912

Synonyms. — Ascaris Linnaeus, 1758, part ; Strongylus Goeze, 1872,
part; Filaria Mueller, 1787, part; Acuaria Bremser, 1811, part;
Spiroptera Rudolphi, 1819, part; Dispharagus Dujardin, 1845, part;
Histiocephalus Diesing, 1851, part; C heilospirura Diesing, 1861,
part.

Generic diagnosis. — Acuariinae (p. 211) : Cutaneous cordons recur-
rent and anastomosing in pairs on each lateral surface (fig. 336).
Cervical papillae tricuspid when evident. Males with spicules un-
equal and dissimilar; usually 5 pairs of postanal papillae.

Parasitic in proventriculus or gizzard of birds.

Type species. — Synhimantus laticeps (Rudolphi, 1819) Railliet,
Henry, and Sisoff, 1912.

key to species of synhimantus

1. Male 5 mm. long; from Falco tridcntatus— Synhimantus sygmoidea, p. 282.
Male (if known) 6 ram. long or longer; from hosts other than above 2.

NEMATODE PARASITES OF BIRDS 273

2. Description incomplete; female 20 nun. long; from Falco subbuteo.

Synhimantus denticulata, p. 27.~.
Female usually less than 20 mm. long; from hosts other than above :;.

3. Vulva anterior to middle of body, dividing body in ratio of 90:103:

spicules 156m and 372m long Synhimantus affinis, p. 273.

Vulva posterior to middle of body (except in 8. laticeps where perhaps
just anterior to middle I : spicules, if male known, differ in length from
above (except possibly in /•>'. sagittata; lengths not given) -1.

4. Only female known, 13 to 15 mm. long; cordons extend posteriorly for %

or more of the length of the muscular esophagus, a distance of S4u> from
the head end; vulva 510/u. anus 170/* from posterior end; from J'clc-

canus species Synhimantus raillieti, p. 283.

Both male and female known; female not agreeing in all respects with
above; from hosts other than PcJccanus 5.

5. Male with 6 to 7 pairs of caudal papillae 0.

Male with 8 to 10 pairs of caudal papillae 7.

6. Cloacal aperture of male only G3m from tail end ; vulva of female not more

than 700m from tail end Synhimantus brevicaudata, p. 274

Cloacal aperture of male much more than G3m from tail end ; vulva described
as posterior to middle of body Synhimantus elliptica, p. 277.

7. Vulva near posterior end of body js.

Vulva not far posterior to middle of body 10.

8. Male with 10 pairs of caudal papillae; G pairs postanal.

Synhimantus recta, p. 280.
Male with 9 pairs of caudal papillae; 5 pairs postanal 9.

9. Male 8.5 to 10.5 mm. long; female 12.2 to 12.8 mm. long; cordons become

very broad in the second half of their length and in their recurrent

portions Synhimantus invaginata, p. 279.

Male 7 mm. long; female 9 mm. long; cordons narrow throughout entire
course Synhimantus sagittata, p. 281.

10. Male with 8 pairs of caudal papillae and in addition 4 small papillae in a

transverse row near the posterior extremity ; spicules 19(>m and 720/x long;

eggs 30m by 19m Synhimantus hamata, p. 27S.

Male with 9 pairs of caudal papillae ; spicule lengths different from above ;
eggs at least 37m long and 22m wide 11.

11. Male G.4 to 6.8 mm. long; female 10.5 mm. long; spicules 200 to 210m and

950 to 960m long; in Asturinula monogrammha.

Synhimantus subrecta, p. 282.

Male 7.2 to 10 mm. long; female 11 to 21 mm. long; spicules 17(>m and 590/u

long; in hosts other than above Synhimantus laticeps, p. 27G.

SYNHIMANTUS AFFINIS (Scurat, 1916) Scurat, 1919

Synony7iiH. — Acuaria affinis Seurat, 1916; FUaria laticeps in part
of Mueller, 1897; Acuaria laticeps, female of Seurat, 1915.

Hosts. — Primary: Strix flam/mea (=z7 1 yto alba); secondary: Un-
known.

Location. — Esophagus.

Morphology. — Synhimantus (p. 272) : Body robust, elongate.
Cuticula thick, transversely striated. Buccal lips hear truncate tooth
and a pair of projecting papillae. Cordons (fig. 3375), longer than
in S. laticeps, descend to level of excretory pore in female, then curve

274 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

and remount along the lateral fields, their anastomosis, when present,
occurring a short distance from the cephalic extremity ; anastomosis
variable, not occurring in many of the males examined by Seurat and
occurring in 1 female only on 1 side and not on the other. Tricuspid
papillae at some distance posterior to cordons.

Male 13.5 mm. long by 290/a wide. Cordons 395/* long. Tri-
cuspid papillae 445/* and 468/* from head end. Tail 360/* long,
slender. Caudal alae narrow, triangular, and pointed at the extrem-
ity, whereas those of S. laticeps are wide and rounded at the extrem-
ity. Four pairs of preanal and 5 pairs of postanal papillae. Right
spicule 156/* long, the left 372/* long.

Female 11.2 to 27.5 mm. long by 300 to 612/* wide. Cordons 660/*
long in worm 19.3 mm. long; tricuspid papillae 410/* and 696/*, and

Fig. 337. — Stnhimantus affinis. a, Female genitalia. After Seurat, 1915.
ft, Head end. After Seurat, 1916

vulva 9 mm. from anterior end in same specimen. Vulva (fig. 337a)
very small, not projecting, situated directly in front of middle of
body. Ovejector directed posteriorly, 420/* long, the cuticular ove-
jector 210/* long. Eggs 38/* by 28/*, thick-shelled, embryonated when
deposited.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe and Africa (Algeria).

SYNHIMANTUS BREVICAUDATA (Dujardin. 1845) Gedoelst, 1919

Synonyms. — Dispharagus brevicaudatus Dujardin, 1845; Spirop-
tera triaenophora Mehlis in Creplin, 1846; Strongylus ardeae-stel-
laris Rudolphi, 1819, nomen nudum; Histiocephalus brevicaudatus
(Dujardin, 1845) Diesing, 1851.

Hosts. — Primary : Ardea stellaris, Botaurus stellaris, Ardetta
minuta; secondary : Unknown.

Location. — Stomach (apparently the gizzard).

Morphology. — Synhimantus (p. 272) : Body filiform. Head 25 to
35/* wide, with 2 conical projecting lips. Cordons (fig. 338a)
transversely striated, extending posteriorly for 230 to 300/*, then
recurrent and anastomosing. Cervical papillae tricuspid, 60/* pos-

NEMATODE PARASITES OF BIRDS 275

terior to cordons in lateral fields. Body cuticula with pronounced
transverse striations, the body margin presenting a serrate appear-
ance. Pharynx (anterior esophagus of Dujardin) 170/* long; an-
terior esophagus (posterior esophagus of Dujardin) 650/* long by
30/* wide; posterior esophagus (ventricule of Dujardin) 2.1G mm.
long.

Mole 10 mm. long by 100/* wide. Cordons extend 228/* from head.
Posterior extremity of body rolled spirally in 3 or 4 turns. Six to 7
pairs of papillae. Cloacal aperture 63/* from tail end. Spicules
indistinct, apparently 60/* and 110/* long.

Female 11.6 mm. long by 290/* wide (Dujardin) or 7.5 to 8.5 mm.
long by 290/* wide (Skrjabin). Cordons extend 310/* from head end
(Dujardin) or 425/* from head end (Skrjabin) then recurrent and
anastomosing, forming a closed arc 230/* from head end, the cordons
gradually increasing in width and attaining a maximum width in the
recurrent part, according to Seurat. Tricuspid cervical papillae
560/* from head end. Pharynx 210/* long, slender; anterior esopha-
gus 850/* long. Caudal extremity thick, in the form of a short obtuse
cone, the anus 68/* (Skrjabin) from tail end (fig. 338&). Vuiva 640
or TOO/i from tail end. Eggs elliptical, 20 to 23/* long (Dujardin) or
30/* by 25/* (Skrjabin).

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe (France and Russia).

SYNHIMANTUS DENTICULATA (Molin, 1860) Skrjabin, 1924

Synonyms. — Dispharagus denticwlatus Molin, 1860c?; Dispharagus
falconis-subbuteonis Diesing, 1851; Spiroptera falconis-subbuteonis
(Diesing, 1851) Diesing, 1851.

Hosts. — Primary : Falco subbuteo; secondary : Unknown.

Location. — Esophagus.

Morphology. — Synhiinantus (p. 272) : Mouth with '2 papilliform
projecting lips (fig. 335)). Body transversely striated, the striations
distinctly denticulate, 5/* apart anteriorly and gradually becoming 8/*
apart posteriorly. Cordons thick, recurrent to midlength, anasto-
mosing in pairs.

Male unknown.

Female 20 mm. long by 200/* wide. Cordons extend posteriorly
520/* from head end; anastomosis 270/* from head end. Eggs elon-
gate oval, 36/* by 16/*.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Europe (France (Kennes)).

276

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SYNHIMANTUS LATICEPS (Rudolphi, 1819) Stiles and Hassall, 1920

Synonyms. — Spiroptera laticeps Rudolphi, 1819; Dispharagus lati-
ceps (Rudolphi, 1819) Dujardin, 1845; Filaria laticeps (Rudolphi,
1819) Schneider, 1866; Spiroptera fallax Siebold, 1837; Filaria in-
voluta Linstow, 1879; Dispharagus spiralis Linstow, 1883, not D.
spiralis Molin, 1858; Dispharagus involutus (Linstow, 1879) Stos-
sich, 1891; Acuaria laticeps (Rudolphi, 1819) Railliet, Henry, and
Sisoff, 1912; Acuaria {Synhimantus) laticeps (Rudolphi, 1819)
Railliet, Henry, and Sisoff, 1912.

Hosts. — Primary : Falco lagopus, Strix bubo, S. alba, S. brachyo-
tus, Falco nisus, F. tinnunculus, Gallus domesticus, Falco cyaneus,

338

339.

Figs. 338-341. — 338, Synhimantus brevicaudata. a, Head end; b,
female tail. after skrjabin, 1917. 339, synhimantus denticu-
lata. Head end. After Dujardin, 1845. 340, Synhimantus lati-
ceps. Male tail. After Seurat, 1916. 341, Synhimantus lati-
ceps. Head. After Schneider, 1S66

Strix flammea, Bubo maximus, Buteo lagopus, Circus cineraceus, G.
cyaneus, Otus braohyotus, Cerchneis tinnunculus, Otus vulgaris, Ac-
cipiter nisus, Aegiolus otus; secondary : Unknown.

Location. — Esophagus and proventriculus.

Morphology. — Synhimantus (p. 272) : Cutaneous cordons (fig. 3-11)
recurrent for one-half their length or more, anastomosing in pairs on
each lateral surface; a little behind their termination there is a tri-
dentate cervical papilla on each side.

Male 7.2 to 10 mm. long by 180/x wide or wider. Body enrolled on
itself. Caudal alae (fig. 340) 1.5 mm. long or longer, with thick
vesicular borders. Spicules unequal and dissimilar, the right 170/*
long by 18/x wide, the left 590/* long, slender, its free portion alate
and its tip slightly dilated and divided into 2 lateral lobes. Six to

NEMATODE PARASITES OF BIRDS 277

8 rows of elevated shields extend 2 mm. anterior to cloacal aperture.
Four pairs of preanal and 5 pairs of postanal papillae.

Female 11 to 21 mm. long, straight. Tail conical, with 2 papillae
near its tip. The small inconspicuous vulva (fig. 342) is just an-
terior to the middle of the body (Seurat) or posterior (Railliet).
Eggs 38 to 42/a by 25/*, thick-shelled, and when oviposited contain an
embryo 225/* long with a sharp slender tail.

Life history. — Unknown; probably involves intermediate stages
in other hosts, which hosts might be small vertebrates, judging from
the fact that the adult worms usually occur in birds of prey.

Distribution. — Europe (Germany, France, Italy), Asia (Russian
Turkestan) and Africa (Algeria). Only reported once from
chicken.

Fig. 342. — Synhimantts laticeps. Female genitalia. Aftek Seurat, 1920
SYNHIMANTUS ELLIPTICA (Molin, 1858) Skrjabin, 1924

Synonyms. — Dispharagus ellipticus Molin, 1858; Acuaria elliptica
(Molin, 1858) Railliet, Henry, and Sisoff, 1912.

Hosts. — Primary : Falco nisus, Accipiter nisus, Astur nisus, Falco
cineraceus, Circus cineraceus, Nisus communis; secondary: Unknown.

Location. — Proventriculus.

Morphology. — Synhlmantus (p. 272) : Mouth with 2 projecting
papilliform lateral lips. Body strongly striated transversely. Cor-
dons thick.

Male 7 mm. long by 200/x, wide. Posterior extremity (fig. 343)
coiled twice in spiral and deeply excavated ventrally. Molin figures
6 pairs of caudal papillae, of which 4 pairs are postanal. Long
spicule with an elliptical dilation at its free end.

Female 20 mm. long by 1 mm. wide, according to Molin, or 10 to
20 mm. long, according to Stossich. Body coiled spirally. Vulva
posterior to middle of body. Eggs 30 to 40^ by 1 ( .) to 27/x.

Gendre (1921a) notes that according to Molin's (1861&) figure the
long spicule is twice as long as the short one; according to the present
writer's interpretation of Molin's figure, one can not be sure where
the short spicule ends, but apparently the long spicule is over four
times as long as the short one. According to Gendre, the presence
of 4 pairs of postanal papillae in this species and of 5 pairs in S.
laticeps is the only point of difference, and he doubts whether S.

278 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

elliptica is a good species since the differences between it and S.
laticeps are minor ones or doubtful.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Europe (Italy (Padua)).

SYNHIMANTUS HAMATA (Linstow, 1877) Skrjabin, 1924

Synonyms. — Dispharague de l'epervier of Dujardin, 1845; Filaria
hamata Linstow, 1877; Dispharagus hamatus (Linstow, 1877) Stos-
sich, 1891; Acuaria hamata (Linstow, 1877) Railliet, Henry, and
Sisoff, 1912.

3M

Figs. 343-344. — 343, Synhimantus elliptica. Male tail. After Molin, 1861.
344, synhimantus hamata. a, head end. aptek linstow, 1879. h, male
tail. After Linstow, 1877

Hosts. — Primary : Falco nisus, Astur nisus, Nisus communis, Buteo
vulgaris; secondary : Unknown.

Location. — Stomach (Gizzard ? ) .

Morphology. — Synhimantus (p. 272) : Mouth with 2 conical lips.
Cordons (fig. 344a) recurrent and anastomosing. Cuticula trans-
versely striated.

Male 6 mm. long by 300/* wide. Buccal cavity 180/* long; esoph-
agus 720/* long. Cordons extend posteriorly 280/* from head end and
anastomose 140/* from head end. Right spicule 720/* long, the free
end shaped like a fishhook; left spicule 190/* long, comparatively
thick and blunt. Four pairs of preanal and 4 pairs of postanal
papillae (fig. 3446), and in addition 4 small papillae in a somewhat
arched transverse row just anterior to tail end. Gendre notes that
these differences in the papillae and in the spicule tip are the only
differences between this species and S. laticeps.

NEMATODE PARASITES OF BIRDS

279

Female 7.4 mm. long (Linstow) or 10 to 11 mm. long (Dujardin)
by 480/* wide. Vulva somewhat posterior to middle of body, the
ratio of part anterior to part posterior being as 4:3. Tail end
rounded, the tail length 1/41 of body length. Esophagus one-third
of body length. Eggs 30/4. by 19/* (Dujardin).

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe (Prussia (Hanover)).

SYNHIMANTUS INVAGINATA (Linstow. 1901) Skrjabin, 1924

Synonyms. — Dispharagus invaginatus Linstow, 1901; Acuaria
{Synh'nnantus) invaginata (Linstow, 1901) Railliet, Henry, and
Sisoff, 1912.

Hosts. — Primary: Babulcus lucidus, Ardeola ibis, purple heron,
and a host unknown; secondary: Unknown.

Location. — Esophagus and under tunic of gizzard.

Fig. 345. — Synhimantus invaginata. a and b, Head end; c, male tail; d,

RIGHT AND C, LEFT SPICULE; f, FEMALE TAIL. AFTER GDNDRE, 1913

Morphology. — Synhimantus (p. 272) : Mouth with 2 triangular.
conical, lateral lips, each bearing 2 very small symmetrical papillae
on the outer surface. Body transversely striated, the striations 5/*
apart anteriorly and 10 to 14/* apart posteriorly. Cordons (tig. 345#
and b) recurrent and anastomosing.

Male 10.45 to 11.1 mm. long by 270 to 290/* wide (Stossich), or
8.46 mm. long by 220/* wide (Linstow). Tail 1/111 to 1/119 of total
body length (fig. 345 c). Pharynx 250 to 270/* long; anterior esoph-
agus 960 to 970/i long; posterior esophagus 3.2 mm. long. Cordons
extend 550/* posteriorly from head and anastomose 300/x from head
end. Tricuspid papillae 630/c from head end. Caudal alae thick,
vesicular, not spread out laterally in leaflike arrangement, but cov-
ering most of lateroventral surface of body; they originate 800/*
anterior to the cloaca. Four pairs of preanal and 5 pairs of postanal
papillae. Left spicule (fig. 345<?) atrophied in appearance, very
slender and flexible, 470/* long by 8/* wide; right spicule (fig. 345c?)
large and robust, 900/* long by 35/x wide (Stossich) or 620/* long

3612—27 20

280 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

(Linstow), its free end peculiar, a digitiform process projecting
from a collarlike expansion.

Female 12.2 to 12.8 mm. long (Stossich) or 9.5 mm. long (Lin-
stow) by 310 to 410ft wide. Pharynx 280 to 310/i long; anterior
esophagus 1 mm. long; posterior esophagus 3.5 to 3.6 mm. long.
Cordons extend 610/* from head end and anastomose 330/u, from head
end. Tricuspid papillae 770/* from head end. Posterior extremity
in a hyaline cylindrical sheath formed from the cuticula (fig.
345 /) . Vulva immediately anterior to anus on a central prominence
which is the real extremity of the body as the small conical tail
projects dorsally from there. Eggs 27 to 29/x by 18 to 19/z.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Germany (Langenburg) and Corsica)
and Africa (Algeria and French Guinea (Labe)).

SYNHIMANTUS RECTA (Molin, 1860) Gendre. 1920

Synonyms. — Dispharfrgus rectus Molin, 1860c/ Spiroptera fal-
conis 9 Molin, 1860; Acuaria recta (Molin, 1860) Railliet, Henry, and
Sisoff, 1912; C heilospirura recta (Molin, 1860) Stiles and Hassall,
1920.

Hosts. — Primary : Falco femoralis, F. unicinctus; secondary : Un-
known.

Location. — Stomacli (gizzard ? )

Morphology. — Synhimantus (p. 272) : Two conspicuous papilla-
like lips. Body straight, transversely striated. According to Molin,
the cordons are equal, flexed, recurrent, and joined in pairs ; Drasche
states that they are not wavy ; Gendre (1921«) states that from Molin's
description of the cordons the species belongs in Synhimantus and
not in C heilospirura where Railliet, Henry, and Sisoff placed it ; the
present writer concurs with Gendre in his interpretation of Molin's
description.

Male 9 mm. long by 100/* wide. Posterior extremity (fig. 346)
rolled spirally. Caudal alae long and wide. Four pairs of peduncu-
lated preanal and 6 pairs of postanal papillae, the latter in 2 groups,
a group of 2 pairs just posterior to cloacal aperture and a group of
4 pairs in the posterior half of the alae; this arrangement constitutes
the chief difference between this species and S. subrecta. Right
spicule short and thick; left spicule four times as long, sharp and
alate.

Female 7 to 10 mm. long by 100 to 300/i wide. Posterior extremity
straight, the anus not far from the tail end. Vulva in posterior
part of body.

• Catalogue of Vienna Museum.

NEMATODE PARASITES OF BIRDS

281

Life history.
in other hosts.
Distribution-.-

-Unknown; probably involves intermediate stages
-South America (Brazil).

SYNHIMANTUS SAGITTATA (Rudolph, 1809) Cram, 1927

Synonyms. — Ascaris sagittata Rudolphi, 1809; /Spiroptera alata
Budolphi, 1819; Dispharagus analis Molin, 1860; FUaria alata
(Rudolphi, 1819) Schneider, 1866; Dispharagus alatus (Rudolphi
1819) Stossich, 1891', Acuaria alata (Rudolphi, 1819) Railliet, Henry,
and Sisoff, 1912; Synhimantus alata (Rudolphi, 1819) Skrjabin,.
1924.

346.

397

Figs. 346-347. — 346, Synhimantus recta. Male tail. After Dbasche,
1SS4. 347, Synhimantus. sagittata. a, Head end ; b, male tail. After
Schneider, 1866

Hosts. — Primary: Ardea nigra, A. purpurea, Ciconia nigra, Butco
borealis, Nycticorax griseus, N. nycticorax, black stork; secondary:
Unknown.

Location,. — In mucosa of stomach (gizzard?).

Morphology.— Synhimantus (p. 272) : Head with 2 small lateral
lips, each provided with a tooth. Cordons (fig. 347«) long, recur-
rent for almost half their length, anastomosing.

Male 7 mm. long. Tail curled in 3 spiral turns. Caudal alae thick,
vesicular. Nine pairs of caudal papillae (fig. 347£>), of which 5
pairs are postanal.

Female 9 mm. long. Vulva very near the anus. Eggs thick-
shelled.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (apparently Germany).

282 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

This species was described by Rudolphi (1809) as Ascaris sagit-
tata, and the name Spiroptera alata Rudolphi, 1819, was a deliberate
renaming of a described species. There is no reason known to the
present writer why sagittata, which is not a nomen nudum, synonym
or homonym, should be dropped and alata used, and the name S.
sagittata is therefore used here in preference to the commonly used
specific name alata in combination with the generic name Synhiman-
tus.

SYNHIMANTUS SUBRECTA (Gendre, 1921) Cram, 1927

Synonym. — Acuaria ( Synhimantus ) subrecta Gendre, 1921a.

Hosts. — Primary: AsturinuJa monogrammica; secondary: Un-
known.

Location. — Stomach ( gizzard ? ) .

Morphology. — Synhimantus (p. 272) : External morphology simi-
lar to S. laticeps. Cordons recurrent and anastomosing at almost the
anterior third of their length, without, however, quite reaching this
level. Tricuspid papillae posterior to cordons.

Male 6.4 to 6.8 mm. long by 250/* wide. Cordons 350 to 360/* long.
Cloacal aperture 380/* from tail end, or 10/169 of total body length.
Four pairs of preanal and 5 pairs of postanal papillae (fig. 348 a),
the latter arranged in 3 groups, one group of 2 pairs just posterior
to cloacal aperture, one group of 2 pairs in middle of caudal alae,
and one group of 1 pair near the tail end. Caudal alae covered with
brilliant granulations, the alae large and thick, extending 800/* an-
terior to cloacal aperture, and continuing even beyond this in the
form of a cuticular pad. Spicules very unequal, the left (fig. 348 c
and d) 950 to 960/* long, the right (fig. 348 b) 200 to 210/* long; these
distinguish this species from S. laticeps.

Female 10.4 to 10.5 mm. long by 330 to 370/* wide. Cordons 420
to 430/* long. Anus 200 to 210/* from tail end, or 10/495 to 10/518 of
total body length from tail end (fig. 348 e) . Posterior extremity coni-
cal, rounded at end. Vulva a little posterior to middle of body, or 10/19
of distance from head. Ovejector directed posteriorly, composed of a
large vestibule 270/* long, followed by a sphincter 200/* long; the
varnish gland or trompe is very short and divides into 2 branches
which immediately diverge. Eggs 37/* by 22/*, embryonated when
oviposited.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Africa (Dahomey) .

SYNHIMANTUS SYGMOIDEA (Molin, 1860) Skrjabin, 1924

Synony?ns. — Dispharagus sygmoideus Molin I860*?; Acuaria syg-
moidea (Molin, 1860) Railliet, Henry, and Sisoff, 1912.

NEMATODE PARASITES OF BIRDS

283

Hosts. — Primary: Falco tridentatus ; secondary: Unknown.

Location. — Xot given; presumably in digestive tract.

Morphology. — Synhimantus (p. 272) : Mouth with 2 very small
papilliform lips. Body transversely striated, the dense striations of
the anterior body forming pseudoannulations, and bent in a sigmoid
curvature. Anterior extremity noticeably attenuated. Cordons
long, flexuous, recurrent, and anastomosing in pairs.

Male 5 mm. long by 300/t wide. Caudal extremity coiled in 2 spiral
turns, its apex obtuse. Alae long and projecting. One spicule short
and recurved; the other long, arcuate and filiform.

Female unknown.

6.

Fig. 348. — Synhimantus subrecta. a, Male tail ; b, right spicule; c, profile

VIEW AND d, VENTRAL VIEW OF LEFT SPICULE; f, FEMALE TAIL. AFTER (JENDRB.
1921

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil).

SYNHIMANTUS RAILLIETI (Skrjabin, 1924) Cram, 1927

Synonym.. — Acuaria raillieti Skrjabin, 1924.

Hosts. — Primary: Pelecanus, species; secondary: Unknown.

Location. — Gular pouch.

Morphology. — Synhimantus (p. 272) : Body slender, cylindrical,
tapering toward both ends. Cuticle delicately striated transversely.
Mouth with 2 lips. Cordons, originating at the base of the lips, ex-
tending posteriorly in 7 symmetrically placed, serpentine coils, re-
current and anastomosing in pairs on lateral surface of body. Width
of cordons varying at different parts, the portion descending from
the lips gradually increasing in width, reaching a maximum in the

284 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

region of the most posterior coil; the ascending branches suddenly
sharply contracting and having approximately the same width
throughout their extent.

Male unknown.

Female 13 to 15 mm. long by 700/* wide. Cordons extending pos-
teriorly for a distance of 840/*; anastomosis 280/* from head end.
Pharynx 425/* long by 30/* wide; muscular esophagus 560/* by 100/*;
glandular esophagus 3.57 mm. by 70/*. Vulva 510/* from the tail
end, a distance of 340/* anterior to the anus. Tail conical, with
rounded end ; anus 170/* from extremity. Eggs 38/* by 20/*, with thick
shells.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Africa (French Somaliland (Jibuti)).

The writer is greatly indebted to Dr. Albert R. Merz, Bureau of
Soils, U. S. Department of Agriculture, for the translation of the
Russian description of this species.

Subfamily Schistorophinae Travassos, 1918c

Subfamily diagnosis. — Acuariidae (p. 210) : Esophagus long, its
anterior part differentiated into a pharynx. Mouth with 2, 4, or
6 lips and cephalic ornamentation consisting of appendices or
festoons.

Parasitic in digestive tract of birds (between the tunics of the
gizzard).

Type-genus. — Schistorophus Railliet, 1916.

KEY TO GBNEKA OF SCHISTOROPHINAE

1. Cephalic ornamentation consisting of 4 sharp cuticular lobes forming a roof

for the head, projecting at sharp angles from the body.

Schistorophus, p. 284.
Cephalic ornamentation different from above 2.

2. Mouth with 6 lips ; cephalic ornamentation of multiple appendices of various

shapes and sizes, some directed anteriorly, some posteriorly ; caudal alae

asymmetrical; vulva in region of anus Serticeps, p. 293.

Mouth with 2 or 4 lips ; cephalic ornamentation different from above ;
caudal alae symmetrical; vulva not in region of anus 3.

3. Mouth with 2 lips ; vulva posterior to middle of body ; lateral papillae

tricuspid Yseria, p. 292.

Mouth with 4 lips ; vulva in anterior part of body ; lateral papillae not
described as tricuspid Histiocephalus, p. 290.

Genus SCHISTOROPHUS Railliet, 1916a

Synonyms. — Tetracanthus Hemprich and Ehrenberg in Schneider,
1866, not Hope, 1835; Ancyracanthus Schneider, 1866, part, not
Diesing, 1838.

NEMATODE PARASITES OF BIRDS 285

Generic diagnosis. — Schistorophinae (p. 284) : Head adorned with
4 sharp cuticular lobes (at times possibly split to give 8 points),
uniting anteriorly with the cuticle, more or less joined at their
origin, especially in the median lines and usually arranged as a
roof over the head. Mouth small, usually with 2 small lips or teeth.
An elongated vestibule. Esophagus composed of 2 parts. Male with
tail blunt, rounded, provided with caudal alae and numerous papil-
lae, the preanal arranged at each side in a long simple series. Two
unequal spicules. Female with short, conical, more or less obtuse
tail; vulva in posterior region or middle of body. Sometimes vivi-
parous.

Parasitic between the tunics of the gizzard of birds.

Type-species. — Schistorophus longicornis (Hemprich and Ehren-
berg, 1866) Railliet, 1916.

KEY TO SPECIES OF SCHISTOROPHUS

1. Description very incomplete ; only host record and size of worm definite — 2.
Description more complete; from other hosts than 2 3.

2. Female 12 mm. long; from Glnreola austriaca.

Schistorophus spinulosus, p. 28S.

Male 4.5 mm., female 9 mm. long. From Tringa helvetica (=Squatarola hvl-

vetica=Squatarola squatarola) Schistorophus bicuspis, p. 2S6.

3. Vulva in middle of body 4.

Vulva in posterior part of body 5.

4. Female 23.5 mm. long. Cuticular processes of head broad. From Haemato-

pus ostralegus ; Russian Turkestan Schistorophus aulieatina, p. 2S7.

Female 14 mm. long. Cuticular processes slender. From Rallus cayennensis;
Brazil Schistorophus laciniatus, p. 288.

5. Head cup-shaped, posterior to which are the cuticular processes, with a total

of 8 points, anteriorly directed Schistorophus bidens, p. 28S.

Head not cup-shaped, cuticular processes with only 4 points, posteriorly

directed 6.

G. Mouth with 2 conspicuous large sharp points; from Sterna, species.

Schistorophus acanthocephalicus, p. 287.

Mouth with 2 small inconspicuous points; from other hosts than Sterna

species Schistorophus longicornis, p. 285.

SCHISTOROPHUS LONGICORNIS (Hemprich and Ehrenberg, I860) Railliet, 1916a

Synonym. — Ancyracanthus longicornis Hemprich and Ehrenberg,
1866.

Hosts. — Primary: Numenius arquatus, Tringa variabilis, Totanus
glottis; secondary: Unknown.

Location. — Between tunics of gizzard.

Morphology. — Schistorophus (p. 284): Mouth (fig. 349 a ami l>)
opening carries laterally a small tooth; around the mouth there are
4 processes, to the dorsal and ventral sides, which project far outward
and backward. Mouth opening leads into a vestibule which is about
twice as long as the distance of the opening from the point of the
head. Esophagus in 2 parts.

286 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 5 to 10 mm. long. Tail with thick swollen walls; 24 pairs
of papillae, 4 of them postanal (Schneider figures (fig. 349<?) only 3
on one side). Two very unequal spicules, one short and thick, the
other long, having a length double that of the space occupied by the
papillae.

Female 8 to 20 mm. long. Vulva 1/3 the total length from the
posterior end; vagina posteriorly directed. Eggs, smooth, thick-
shelled, elliptical.

Life history. — Unknown.

Distribution. — Africa (Egypt).

SCHISTOROPHUS BICUSPIS (Rudolphi, 1819) Railliet, 1916a

Synonyms. — Spiroptera bicuspis Rudolphi, 1819 ; Dispharagus bi-
cuspis (Rudolphi, 1819) Dujardin, 1845; Histiocephalus gracilis
Diesing, 1851; Histiocephalus bicuspis (Rudolphi, 1819) Linstow,
1878.

c. — "-

'Qo.

c

• ee« o eoeooooooo

Fig. 349. — Schistorophus longicobnis. a, Lateral, view and b, front view of
head ; c, male tail. Aftbr Schneider, 1866

Host. — Primary: T ring a helvetica (=Squatarola helvetica); also
reported from Grus cinerea and Vanellus melwiw g aster ; secondary:
Unknown.

Location. — Between tunics of gizzard.

Morphology. — Schistorophus (p. 284) : Body slender at the 2 ex-
tremities, especially the anterior, twice as thick in middle portion.
Head small, continuous with body and armed with papillae ; to each
side, posterior to the head, there is a spine or subulate tooth, directed
sometimes horizontally, sometimes posteriorly. (In this species, as in
S. bidens, the original description of only 2 processes is explained by
the fact that those are the ones seen in profile; the others were evi-
dently overlooked).

Male 4.5 mm. long ; tail forming 2 turns of a spiral ; membranous
alae straight.

Female 9 mm. long; tail ending in a short, flattened, recurved
point, anterior to which is the anus.

Life history. — Unknown.

Distribution. — Europe (Austria (Vienna Museum) ).

Railliet (1916) states that this species is probably identical with
S. longicornis, p. 285.

NEMATODE PARASITES OF BIRDS 287

SCHISTOROPHUS ACANTHOCEPHALICUS (Molin, 1860) Railliet, 1916a

Synonyms. — Spiroptera acanthocephalica Molin, 18606/ S. capil-
lars Molin, 1860; Oheilospirura capillars (Molin. I860) Diesing,
1861; Schistorophus capillaris (Molin, 1860) Railliet, 1916.

It is questionable as to which of the 2 species of Molin was the
first published. Spiroptera acanthocephalica is listed as 18606 in
the catalogue of Stiles and Hassall ; the paper was read in December,
1859, but no month is given on it for its publication. Spiroptera
capillaris is listed as Molin, 1860e; it was published in March of that
year.

Strongylus ambiguus Rudolphi, 1802, Spiroptera sternae Rudolphi,
1819 and Spiroptera sternae hirundinis Deslongchamps, 1824 have
been listed by various of the earlier writers as synonyms of this
species; they would invalidate the present specific name, if they are
identical, but the descriptions are so very inadequate for all of them
as to make them unrecognizable, the same host (Sterna hirundo)
being the only definite grounds for comparison.

Hosts. — Primary : Sterna caspica and S. hirundo; secondary : Un-
known.

Location. — Between the tunics of gizzard.

Morphology. — Schistorophus (p. 284) : Head (fig. 350) continuous
with body. Mouth bilabiate, the lips conspicuous, ending anteriorly
in a sharp point; to each side of the lips there is a lancet-shaped
process (a total of 4), posteriorly directed, strongly developed. Body
filiform, densely striated transversely; anterior extremity noticeably
attenuated.

Male unknown.

Female 15 to 19 mm. long by 200^ wide. Anus not far from caudal
extremity. Vulva prominent, in posterior part of body.

Life history. — Unknown.

Distribution. — Europe (Austria (Museum Vienna)).

SCHISTOROPHUS AULIEATINA Skrjabin. 1916b

Host. — Primary : Haematopus ostralegus; secondary : Unknown.

Location. — Under cuticle of gizzard.

Morphology. — Schistorophus (p. 284) : Anterior part of body (fig.
351) contracted. Cuticle transversely striated. Head identical with
that of S. longicornis (p. 285).

Male unknown.

Female 23.5 mm. long by 170/u, wide. Anus very near to tail end,
the width of body at anus being only 34^. Vulva almost in middle
of body, slightly anterior to middle. Eggs oval, 40/* long by 26/..
wide.

Life histoiy. — Unknown.

Distribution. — Asia (Russian Turkestan).

288 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SCHISTOROPHUS LACINIATUS (Molin. 1860) Railliet, 1916a

Synonym. — Histiocephalus laciniatus Molin, 1860c.

Host. — Primary : R alius cayennensis ; secondary : Unknown.

Location. — Between tunics of gizzard.

Morphology. — Schistorophus (p. 284) : Head discreet, encircled by
a fringe of slender but long, posteriorly directed processes. Mouth
with lips. Body slender in all parts, especially attenuated pos-
teriorly.

Male 7 mm. long by 100//, wide; caudal extremity twisted in spiral,
with long equal alae which have 24 single papillae.

Female 14 mm. long by 200/x wide ; anus near to caudal extremity ;
vulva in median part of body, not prominent.

Life history. — Unknown.

Distribution. — South America (Brazil).

SCHISTOROPHUS SPINULOSUS (Molin, 1860) Railliet. 1916a

Synonyms. — Spiroptera glareolae austriacae 10 / Filaria spinulosa
Molin, 18606?.

Host. — Primary : Glareola austriaca; secondary : Unknown.

Location. — Between the tunics of gizzard.

Morphology. — Schistorophus (p. 284) : Head with a crown of pos-
teriorly directed spinous processes. Body filiform, transversely stri-
ated, spirally twisted ; anterior extremity noticeably attenuated.

Male unknown.

Female 12 mm. long by 100/a wide. Caudal extremity conical,
obtuse.

Life history. — Unknown.

Distribution. — Europe (Austria (Vienna Museum) ).

SCHISTOROPHUS BIDENS (Rudolphi, 1819) Railliet, 1916a

Synonyms. — Spiroptera bidens Rudolphi, 1819; Dispharagus bidens
(Rudolphi, 1819) Dujardin, 1845; Spiroptera denticulata Molin,
1860 b; Ancyr acanthus bidens (Rudolphi, 1819) Schneider, 1866.

Llosts. — Primary: Accipiter nisus, Astur palumbarius, Falco pal-
mnbarius, Merops apiaster; secondary : Unknown.

Location. — Between the tunics of gizzard.

Morphology. — Schistorophus (p. 284) : Body very slender, attenu-
ated at the extremities, especially the anterior extremity; Dujardin
describes the body as macroscopically resembling a simple silk thread.
Head (fig. 353 a) rounded, separated from the body by a constriction,
directly posterior to which there is a wreath of 8 anteriorly directed,
pointed processes. These are usually adherent to the cuticle and at
times difficult to see. This explains the early descriptions and the

""Catalogue of Vienna Museum.

NEMATODE PARASITES OF BIRDS

289

specific name, only the 2 processes shown in profile at the sides of the
body being observed by the earlier authors (Rudolphi and Dujardin).
Mouth cilp-shaped, with 4 papillae. Lepri says that these papillae
are situated on a lip which surrounds the mouth and which can be
retracted into the mouth. Mouth opens into long thin-walled vesti-
bule ; esophagus in 2 parts.

Male 6.75 to 10 mm. long. Posterior extremity rolled twice in
spiral ; alae wide, oval. Lepri says there are 4 pairs of preanal and

35Z

Figs. 350-353. — 350, Sciiistorophus acanthocepiialicus. Head. After .Diuschk,
1884. 351, schistobophos aulieatina. head end. after skejabin, 1916.
352, schistoeophus bidens. male tail. after schneider, 1866. 353,
schistobophus bidens. a, head; b, male tail. after lepri, 1898

2 pairs of postanal pedunculated papillae (fig. 353 b) but Schneider
describes and figures 10 pairs in all (fig. 352).

Female 12 to 19 mm. long. Tail ending in a short, recurved
point anterior to which is the anus. Vulva bilabiate, the lips large
and projecting, situated posterior to (lit! middle of the body; accord-
ing to Schneider mm. from the posterior end in a speciman 15 mm.
long. Eggs 43/x long by 19/* wide.

Life history. — Unkown.

Distribution. — Europe (Austria (Museum, Vienna) ) and Italy
(Province of Rome).

The cup-shaped head, the presence of the processes at the neck
and anteriorly directed instead of on the head and posteriorly di-
rected, are different from those of other species of Schistorophus and
make the status of this species questionable.

290 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Genus HISTIOCEPHALUS Diesing, 1851

Generic diagnosis. — Schistorophinae (p. 284) : Head provided with
4 small lips with submedian papillae. Posterior to the lips 2 tough
lateral appendices, each divided into numerous branches; each
branch may or may not be subdivided into processes at its free end.
Cervical region swollen into a bulla consisting of numerous longi-
tunidal folds. Male with large caudal alae with pedunculated papil-
lae of which 4 pairs are preanal. Spicules equal or unequal. Female
with vulva in anterior part of body.

Parasitic under the tunic of gizzard of birds.

Type species. — Histiocephalus laticaudatus (Rudolphi, 1819) Dies-
ing, 1851.

This genus was placed in the Acuariinae by Skrjabin (19166) on
the basis of the caudal papillae and the ornaments of the head.
Gendre (1921&) states that because of the lip structure, the genus
belongs in the Spiruridae rather than the Acuariidae. Travassos
(19205) has not included the genus in his classification. The present
writer has assigned it to the Schistorophinae in consideration of the
nature of the very striking cephalic ornamentation.

KEY TO SPECIES OF HISTIOCEPHALUS

Head with 12 processes, each ending in a simple point ; spicules unequal, the
left 700m long, the right 155/t long Histiocephalus tridens, p. 291.

Head with 20 to 24 processes, each ending in 2, 3, or 4 branches; spicules equal
(6 mm. long) Histiocephalus laticaudatus, p. 290.

HISTIOCEPHALUS LATICAUDATUS (Rudolphi, 1819) Diesing, 1851

Synonyms. — Spiroptera laticaudata Rudolphi, 1819; Filaria lati-
caudata Schneider, 1866; Dispharagus laticaudatus (Rudolphi, 1819)
Dujardin, 1845.

Hosts. — Primary: Gallus gallus and Otis tetrax; secondary: Un-
known.

Location. — Under the lining of the gizzard.

Morphology.— Histiocephalus (p. 290) : Head (fig. 354 a and b)
with 2 semicircular chitinous rings, each bearing 10 to 12 posteriorly
directed chitinous appendages, which terminate posteriorly in 2, 3,
or 4 branches; these appendices are approximately equal in length
and in the male may measure 55 to 66/*. Posterior to this structure
the neck is swollen by a series of thick longitudinal ridges. There
are 4 small hemispherical lips, the lateral bearing 2 very small
papillae ; there are 4 submedian papillae.

Male 5 to 11 mm. long by 100 to 200/* wide, or to 340/*, according
to Skrjabin (fig. 355). Posterior extremity of body spirally twisted,
according to Molin, slightly enlarged and provided with 2 caudal

NEMATODE PARASITES OF BIRDS

291

alae (fig. 354c). The caudal alae have 6 pairs of ray-like papillae,
of which 4 are preanal and 2 postanal. In a worm 9.3 mm. long,
the 2 slender, equal spicules measure 6 mm. long by 10>n wide. The
cloacal aperture in a worm this size is 110/x from the posterior
extremity.

Female 7 to 14 mm. long by 200 to 400/i. wide. Tail straight and
obtusely conical, with the anus not far from tip. Vulva in anterior
part of body; it is a longitudinal cleft with 2 prominent lips.

Life history. — Unknown.

Distribution. — Europe and Asia (Russian Turkestan).

HISTIOCEPHALUS TRIDENS Gendre, 1921b

Host. — Primary : Traehelotis senegalensis; secondary : Unknown.
Location. — Under lining of gizzard.

Morphology. — Histiocephalus (p. 290) : Red or reddish-yellow
worms when alive, brown after killing so that internal anatomy is

3fl?

Figs. 354-355. — Histiocephalus laticaudatus. 354, a. Front view and b, lateral
VIEW of head; c, male tail. After Drasche, 1S84. 355, Male. After
Skrjabi.nv, 1016

difficult to study. Body filiform; cuticle transversely striated.
Head (fig. 356 a and b) distinctly set off from body by a furrow,
following which is a cuticular collar, 16/* high in the male, 2<V in the
female, formed by narrow projecting ridges. Mouth with 4 lips, the
2 laterals being much larger than the dorsal and ventral. The
former (i. e., the laterals) are trilobed, the lateral lobes of each
bearing a small papilla ; on the inner surface each lateral lip has 4
triangular teeth and on the outer surface 2 membranous fan-shaped
expansions, each divided into 3 divergent, posteriorly directed
branches ending in conical points. The dorsal and ventral lips are
smaller and less distinct than the laterals, reaching only about half
the height of the latter; each has 2 large projecting papillae at its
base.

Male 5.2 mm. long by 20G> wide. Cloacal aperture 96^ from pos-
terior extremity (1/54 of total length). Caudal alae (fig. 356 c) large

292

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

and oval, each composed of 2 membranes, a dorsal and a ventral,,
transversely striated, united at their lateral edges and curved so
that they approach the median line, appearing thus almost as a
sheath. Six pairs of caudal papillae, of which 4 are preanal, 2 post-
anal. Spicules unequal and dissimilar, the left (fig. 356 d, e, and /)
700//, long, cylindrical in its anterior one-fourth, then shaped as a
gutter throughout its length until 42/* from the free end where there
is a hyaline, vesicular membrane enveloping it; the right spicule
(fig. 356 g, h, and i) 155/* long, its appearance different in different
views.

Fig. 356. — Histiocephalijs tridens. a, Front view and 6, profile view of
head ; c, male tail ; d, anterior end ; e, posterior end, lateral view ; f,

VENTRAL VIEW, LEFT SPICULE; g, RIGHT SPICULE, LATERAL VIEW; ll, RIGHT
SPICULE, DORSAL VIEW ; i, SHOWING NOTCHED APPEARANCE WHEN SPICULE RE-
MOVED FROM BODY. AFTER GEiNDRE, 1921

Female 12.1 mm. long by 220/* wide. Anus 87/* from posterior end.
(1/138 of total length) ; posterior extremity conical, rounded at end.
Vulva salient, in anterior region of body, 2.07 mm. from anterior
extremity. Ovejector 1 mm. long, not differentiated into different
parts, directed posteriorly, gradually decreasing in thickness. Egg*
not mature in Gendre's specimens.

Life history. — Unknown.

Distribution. — Africa (French Guinea (Labe)).

Genus YSERIA Gedoelst, 1919

Generic diagnosis. — Schistorophinae (p. 284) : Mouth with 2 lips;
posterior to the lips a hood or crown with festooned edge. Lateral
papillae tricuspid. Female with vulva posterior to middle of body,

Parasitic under mucosa of gizzards of birds.

Type-species. — Yseria calif ornica Gedoelst, 1919.

NEMATODE PARASITES OF BIRDS 293

KEY TO SPECIES OP YSEIUA

Female 12 mm. Ionic by 100/* wide; male 8 mm. long. In Alcedo americana

and Rallus cauennensis Yseria coronata, p. 293.

Female 25.5 mm. long by 416yu. wide; male unknown. In Oidemia deglandi.

Yseria californica, p. 293.

YSERIA CALIFORNICA Gedoelst, 1919

Host. — Primaiy: Oidemia deglandi; secondary: Unknown.

Location. — Under mucosa of gizzard.

Morphology. — Yseria (p. 292) : Color deep brown or almost black;
cuticle finely striated transversely. Mouth with 2 lateral hemispher-
ical lips surmounted by a triangular tooth; posterior to the lips the
head is encircled by a hood with festooned edge. Cervical papillae
tricuspid.

Male unknown.

Female 25.5 mm. long by 416//, wide. Tricuspid papillae 185/u, from
head end. Vulva 14 mm. from head end, thus the ratio of anterior
to posterior part of body is 5 : 4. Eggs not described.

Life history. — Unknown.

Distribution: — North America (United States (California)).

YSERIA CORONATA (Molin, 1860) Gedoelst, 1919

Synonyms. — Spiroptera coronata Molin, 1860; Histioceplialus
coronatus (Molin, 1860) Skrjabin, 19166.

Hosts. — Primary : Alcedo americana and Rallus cayennensis ; sec-
ondary: Unknown.

Location. — Under mucosa of gizzard.

Morphology. — Yseria (p. 292) : Head set off from body, armed witli
crown of sharply pointed, posteriorly directed processes. Mouth
with 2 small tooth-like papillae. Body slender, increasing in width
post! sriorly, the anterior extremity noticeably attenuated.

Male 8 mm. long. Caudal extremity conical, its apex truncate.
Anus not far from tail end. Vulva in posterior part of body.

Life history. — Unknown .

Distribution. — South America (Brazil).

Genus SERTICEPS Railliet, 1916a

Generic diagnosis. — Schistorophinae (p. 284) : Head adorned with
varied and multiple appendices or festoons. Mouth with 6 small
lips each carrying a small papilla. Male with obtuse (ail: caudal
alae asymmetrical; 10 pairs of preanal papillae. Two very unequal
spicules. Female with obtuse tail. Vulva in region of anus.

Parasitic between the tunics of gizzard of birds.

Type-species. — Serticeps vulvoinflata (Molin, 1860) Railliet, 1916a.

294 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

SERTICEPS VULVOINFLATA (Molin, 1860) Railliet, 1916a

Synonym. — Spiroptera vulvoinflata Molin, 18606.

Host. — Primary: Trochilus ochropygus; secondary: Unknown.

Location. — Between the tunics of the gizzard.

Morphology. — Serticeps (p. 293) : Mouth with 6 small lips, each
bearing a papilla. Body densely striated transversely; anterior ex-
tremity noticeably 'attenuated, apex truncate. Molin described the
cephalic appendices simply as a crown of short horizontal spines.
Drasche describes them (figs. 357a and 358) as follows: Posterior to
the lips 2 thin lateral processes, shaped like a bird's tail, projecting
outwards. Farther posterior 2 dorsoventral, arched, outwardly
directed soft processes. Still more posterior 4 submedian, rectangu-
lar flaps directed outwards and turned under on the sides. Lastly
there follows a wreath of numerous processes of different sizes, pos-
teriorly directed. Between the dorsoventral and submedian wings
there project 4 large, long, club-shaped papillae, directed obliquely

358.

Figs. 357-358. — Serticeps vulvoinflata. 357, a, Head; 6, male tail. 358,
Labeled Spiroptera mediospiralis, but apparently incorrectly so. See text
for discussion. after draschd, 1884

outward and forward. (The end face view (fig. 358) given by the
present writer, copied from Drasche, 1884, was labelled by Drasche
as Spiroptera mediospiralis while another figure of the same plate
(fig. 14) was labelled S. vulvoinflata. Since the figure given here,
Drasche's figure 16, agrees with the description of S. vulvoinflata
whereas his figure 14 does not, but on the other hand does agree with
the description of S. mediospiralis, it is evident that Drasche con-
fused the 2 labels.)

Male 10 mm. long by 100/* wide. Caudal alae (fig. 3576) wide
and asymmetrical, the left wider than the right. Twelve pairs of
large pedunculated papillae, of which 10 are preanal, 2 postanal.
The distance between the first 11 papillae (counting from posterior
to anterior) is the same but the distance between the eleventh and
twelfth is twice as great as the former. Right spicule short and
thick ; the left spicule 8 times as long as the right, alate and pointed.

Female 28 mm. long by 200/* wide. Caudal extremity long, acutely

NEMATODE PARASITES OF BIRDS 295

pointed, straight, the apex obtuse. Anus remote from tail end.
Vulva anterior to anus, very prominent, the posterior lip strongly
inflated.

Life history. — Unknown.

Distribution. — South America (Brazil).

Family PHYSALOPTERIDAE Leiper, 1908

Family diagnosis. — Spiruroidea (p. 102) : Mouth with 2 lips pro-
vided with teeth on their inner surface. Head without ornamenta-
tion. Male with large caudal alae, joined anteriorly across the ven-
tral surface; caudal papillae pedunculated. Female with 2, 4, or
more uteri. Vulva anterior to middle of body.

Parasitic in mammals, birds, reptiles, and, rarely, amphibians.

Type-genus. — Physaloptera Rudolphi, 1819.

Subfamily Physalopterinae Stossich, 1898
Subfamily diagnosis. — Characters of the family.
Genus PHYSALOPTERA Rudolphi, 1819

Generic diagnosis. — Physalopterinae (p. 295) : Body robust, mas-
sive. Cuticle thick, finely striated transversely, detached from the
body in the cephalic region where it forms an annular collar which
serves for fixing the parasite. Lateral fields large but with no cu-
ticular expansions. Two sensory papillae in the esophageal region,
situated posterior to the nerve ring. Mouth with 2 large lateral lips
provided with teeth on their internal surface and externally with
papillae near the point of their insertion. Buccal cavity short.
Esophagus clearly divided into 2 parts, a muscular clear portion and
a glandular opaque portion.

Male with wide outspread caudal alae which join each other an-
teriorly across the ventral surface; 4 to 5 pairs of long pedunculated
papillae in cloacal region and a variable number of small ventral
papillae. Two spicules, usually unequal. No gorgeret.

Female with small non-projecting vulva, situated anterior to the
middle of the body; ovejector tubular, very long, usually directed
posteriorly. Uteri 2, 4, or more in number, parallel. Eggs with
thick shells, embryonated at maturity.

Parasitic normally in the digestive tract, generally the stomach, of
mammals, birds and reptiles, very rarely in amphibians.

Type-species. — Physaloptera clausa Rudolphi, 1819.

Travassos (1920<?) divided the genus Physaloptera into .'» genera:
Physaloptera, Olamidonema, Turgida, Abreviata, and Leptosoma.
Ortlepp, however, in an analysis of these genera, points out that this
division is unjustifiable. The present author is therefore using
Physaloptera in its original generic sense.

296 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

KEY TO SPECIES OF PHYSALOPTERA

1. True Physaloptera, i. e., species agreeing with the generic diagnosis 2.

Species inquirenclae, placed in this genus but not agreeing entirely with the

generic diagnosis 16.

2. Normally in reptiles; found as pseudoparasite in Ciconia alba.

Physaloptera abbreviata, p. 307.
Normally in birds 3.

3. Teeth of lips undescribed 4.

Teeth of lips described 0.

4. Description very incomplete; from Lanius minor.

Physaloptera bilabiata, p. 307.
From other hosts than above 5.

5. Male 18, female 30 mm. long; cuticle enlarged posterior to the lips; eggs

81jt long by 26/* wide ; located in head of Micropogon, species.

Physaloptera fusiformis, p. 303.
Male 26, female 26 mm. long ; cuticle reflected over lips ; eggs not described ;
in proventriculus of Falco, species; (probably a synonym of P. alata).

Physaloptera megalostoma, p. 300.

6. Two teeth on each lip, an outer and an inner (tripartite) 7.

One tooth or more than 2 teeth on each lip 15.

7. Relative sizes of teeth not described ; distinctive character consists in the

one species (P. truncata) of the outer tooth being widened or "knobbed"
at the top, in the other species (P. crassa) of the presence of only 1 pair

of central caudal papillae 8.

Relative sizes of teeth described ; outer tooth not as above ; pedunculated
papillae number 11 to 13 9.

8. Male 14 mm., female 23 mm. long ; 5 pairs of lateral pedunculated caudal

papillae; 1 pair of ventral papillae Physaloptera crassa, p. 302.

Male 25, female 33 mm. long ; 4 pairs of lateral pedunculated caudal papil-
lae, 13 ventral papillae Physaloptera truncata, p. 306.

9. Outer tooth large, inner tripartite tooth small 10.

Inner tripartite tooth large, outer tooth small 14.

10. Caudal alae with 4 pairs of lateral pedunculated papillae.

Physaloptera acuticauda, p. 297.
Caudal alae with 5 pairs of lateral pedunculated papillae 11.

11. More than 11 ventral caudal papillae (14 as figured) ; tail of female short

(from 1/40 to 1/70 of total body length) Physaloptera crosi, p. 302.

Eleven ventral caudal papillae; tail of female elongate (1/30 to 1/21 of total
body length) 12.

12. Male 7 mm. long; cervical papillae situated 100/j. posterior to the end of

muscular esophagus Physaloptera alata chevreuxi, p. 301.

Male 17 mm. or longer ; cervical papillae at level of terminal region of
muscular esophagus 13.

13. Male 22 to 28.5 mm., female 33 mm., long ; cloacal aperture of male 1.2 mm.

from posterior extremity; vulva of female anterior to end of esophagus,
at about the anterior eighth of total body length.

Physaloptera alata nouveli, p. 301.
Male 17 to 20 mm., female 19 to 27 mm. long; cloacal aperture of male 650/*
from posterior extremity ; vulva of female posterior to end of esophagus,
at tbe anterior fifth of body length (according to Seurat) or more pos-
terior (according to Schneider) Physaloptera alata, p. 298.

NEMATODE PARASITES OF BIRDS 297

14. Spicules subequal (360 and 380,u long) ; vulva a little anterior to middle of

body Physaloptera galinieri, p. 303.

Spicules unequal (400 and S40fi long) ; vulva at anterior third of body
length Physaloptera subalata, p. 305.

15. Female 7 mm. long; only one simple tooth on each lip.

Physaloptera, inflata, p. 305.
Female 19 mm. long; the large conical tooth of each lip has to each side of
it a smaller tooth ; additional still smaller teeth present internally.

Physaloptera gemina, p. 304.

16. In orbital cavity of Ibis aethiopica; head with wing-like appendages and 2

or more papillae or small lips Physaloptera, species Parona, p. 309.

In alimentary canal (or in P. malleus location not given) of other hosts
than above ; head structure different from above 17.

17. Description very incomplete: alae described as vesicular; in Cuculus, species,

Brazil Physaloptera strongylina, p. 309.

Alae not described as vesicular ; in other hosts than above, in Africa or
Europe 18.

18. Mouth with 2 lips ; 4 pairs of preanal pedunculated papillae 19.

Mouth with 6 lip structures ; 2 pairs of preanal pedunculated papillae 20.

19. In the middle of each lip a papilla and to each side of the papilla a cone-

shaped tooth ; spicules equal ; tail of male 1/69 of body length ; eggs 39/*

long by 2Gfi wide Physaloptera brevicauda, p. 307.

No papillae or teeth described on lips ; spicules unequal ; tail of male 1/33
of body length ; eggs 46,u long by 29fi wide Physaloptera malleus, p. 308.

20. Spicules equal (290^ long) ; tail of male 1/14, of female 1/53 of body length ;

vulva in anterior third of body Physaloptera ovata, p. 309.

Spicules very unequal (SNOm and 2.17 mm. long) ; tail of male 1/46, of female
1/21 of total body length ; vulva near posterior extremity of body. Species
recently transferred to Cyrnea Physaloptera bulbosa, p. 310.

PHYSALOPTERA ACUTICAUDA Molin. 1860

Synonym. — Physaloptera data Rudolphi of Diesing, 1851 in part.

Hosts. — Primary: Elanus caeruleus, Falco atticaplllus (=Spi~l-
■cw+er melanoleucus) , F. cachlnans {=Eerpetotheres cachinans), F.
eayennensis (=zLeptodon cayennensis) , F. cm^onatus (=HarpyIia-
liaetus coronatus), F. dispar (= Elanus leucurus), F. gracilis
{ — Geranosplzlas caerulescens) , F. minutus (=Acdpiter tinus), F.
ornatus (=Splzaetus mauduytlj, F. palustris (—Circus maculosus),
F. species, F. swainsonll (=camps07iyx swal?isotu), F. ruiilans
{=TIeterosplzkis meridio nails) , F. uniclnctus {—Parabuteo unicinc-
ius), F. wubutinga {Vrubitinga urubltlnya) ; secondary: Unknown.

Location, — Mouth, esophagus, gizzard, orbital cavity.

Morphology.- -Physaloptera (p. 295) : Cuticle finely striated trans-
versely, partly reflected over the lips anteriorly. Mouth with 2 lips,
their anterior border semicircular; each lip with a large pointed
outer tooth and internal to it a smaller membranous tooth with 3
denticulations.

Male 18 to 32 mm. long by 620 to 800/x wide. Caudal alae (fig.
359&) long, semilanceolate, supported by 4 pairs of equidistant

298

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

pedunculated papillae, of which 2 are preanal and 2 postanal. Three
preanal ventral papillae; 5 pairs of sessile postanal papillae near
the median line, of which 2 pairs are in a row just posterior to the
anal opening, 2 slightly farther back and the remaining pair about
2/3 of the distance from the cloacal aperture to the tip of tail.
Spicules unequal, sharply pointed, the left 1.89 to 2.17 mm. long
by 45/* wide at the base, the right 420 to 490/* long by 50/* wide.

Female 20 to 43 mm. long by 800/* to 1.2 mm. wide. Posterior ex-
tremity conical, narrowing to a rather acute tip. Anus about 1/56
of total body length from posterior end. Vulva far anterior, open-
ing on a slight elevation anterior to the end of the esophagus. Mus-
cular vagina only about 480/* long by 50/* wide, slightly enlarged
posteriorly; 2 uteri arising from the outer sides (fig. 359«). This
is the only one of the Pkysaloptera from birds known to have this

Figs. 359-360. — 359, Physaloptera acuticafda. a, Terminal part of female;

GENITALIA ; b, MALE TAIL. AFTER ORTLEPP, 1922. 360, PHYSALOPTERA ALATA. C,

Head; 6, lip; c, male tail. After Linstow, 1S77

arrangement of uteri; it is similar to that of P. praeputialis, a par-
asite of mammals. Eggs 51t«, long by 42/* wide, embryonated in
utero.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — South America (Brazil) and Africa (Lake Ny-
assa).

PHYSALOPTERA ALATA Rudolphi, 1819

Synonyms. — Vermis dubius falconis nisi; Physaloptera megalos-
toma Creplin, 1829 (probably a synonym; see page 300) ; Spiroptera
physalura Dujardin, 1845.

Hosts. — Primary: Accipiter nisus, Aquila imperialism A. pennata,
Astur nisus, Buteo vulgaris, Circaetus gallicus, Circus aeruginosus,
C. cineraceus, C. cyaneus, C. rufus, C. pygargus, Falco apivorus, F.
atricapillus, F. biarmicus erlangeri, F. cachinans, F. cayennensis, F.
coronatus, F. dispar, F. gracilis, F. longipennis, F. ornatus, F. palus-

NEMATODE PARASITES OF BIRDS

299

tris, F. pygargm^ F. 7*ufus, F. rutilans, F. species, F. mbbutt us,
F. swainsonii, F. unicincttts, F. urubutinga, Tinnwnculus alaudarius;
secondary : Unknown.

Location. — Gizzard and intestine.

Morphology. — Physaloptera (p. 295) : Cuticle may be completely
reflected over the head, with a large crater-like opening anteriorly,
or in young specimens this may be entirely lacking. Cervical papil-
lae situated at level of terminal region of muscular esophagus.
Mouth (fig. 360a) with 2 lateral lips (fig. 360Z>), each with a
triangular external tooth and 3 smaller, blunter internal teeth; in
addition 3 papillae, near the insertion of each lip.

Male 17 to 20 mm. long. Entire esophagus 1/4 of body length.
Cloacal aperture 650/* from caudal extremity. Circumcloacal region
covered with small tubercules. Caudal alae (fig. 360<?) A'ery elon-
gated. Five pairs of long pedunculated papillae, of which 2 are
preanal. 1 ad anal and 2 postanal. Three papillae on the anterior
edge of cloacal aperture. Four pairs of postanal ventral papillae,
of which 2 pairs form a transverse row on the posterior edge of

Fig. 361. — Physaloptera alata Rudolphi of Ortlepp, 1922. Possibly
equivalent to p. galinieri seuuat. see text for discussion. after ort-
lepp, 1922.

the cloacal aperture, the third pair is at the level of the most pos-
terior pair of lateral pedunculated papillae, and the fourth pair
is situated about midway from cloacal aperture to caudal extremity
(Seurat says 5 pairs of postanal papillae but in describing their
position only lists 4 pairs). Spicules 420/* long, according to Lin-
stow; 280 and 265/t long, according to Seurat.

Female 19 to 27 mm. long. Entire esophagus 1/5 of body length.
Tail 1/21 of body length, conical, attenuated. Schneider states that
the vulva is 7 mm. from the anterior extremity of a 19 mm. long
specimen, but Seurat describes its position as more anterior (250/*
posterior to the end of the esophagus which is at the anterior fifth
of body length). Ovejector tubular, 1.5 mm. long; trompc dilated
as a reservoir, beyond which it narrows again for a course of 300it to
the branching of the uteri. Eggs thick-shelled, embryonated, 4G to
55/x long by 25 to 27/t wide.

300 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Europe (Italy (Rome), and Austria (Museum,,
Vienna)), Asia (Russian Turkestan and India), South America
(Brazil), Africa (Bou-Saada and Biskra), and Australia.

The type host of this nematode is evidently Falco (■=■ Accipiter)
nisus; however, Rudolphi had in addition specimens from F. gallicus.
and F. pennatus, which he lists as this species. Schneider (1866)
and also Linstow (1877a) found this species in Falco nisus, and de-
scribed and figured it; Seurat (1915&) has described it from that
host and from F. biannicus. Ortlepp (1922) examined Rudolphi's-
specimens from F. gallicus and F. pennatus, incorrectly called them
paratypes and noted differences in lip-structure and in caudal pa-
pillae from those described by Schneider, Linstow, and Seurat. Bay-
lis (1925), however, calls attention to the fact that the material dealt
with by the latter authors was from the type host and that as Ort-
lepp states that the material he examined (fig. 361) resembled P.
galinieri Seurat, that part of Rudolphi's material may be the latter
species.

The above description of P. dlata is based on those of Schneider,
Linstow, and Seurat.

PHYSALOPTERA MEGALOSTOMA Creplin, 1829

Synonym. — Spiroptera megalostoma) (Creplin, 1829) Dujardin,
1845.

Hosts. — Primary: Falco nisus and F. palumbarius; secondary:
Unknown.

Location. — Proventriculus.

Morphology. — Physaloptera (p. 295) : Color brownish; head larger
than bod}^. Mouth large, circular, gaping, bare, surrounded by a
swollen margin. Body moderately thick, a little more slender an-
teriorly than posteriorly.

Male about 26 mm. long. Tail curved a single time ; alae inflated,
joined anteriorly on ventral surface, narrowing laterally as they
approach end of tail. Spicules rather short and apparently equal.

Female about 26 mm. long, somewhat thicker than the male. Tail

straight, obtuse, tapering toward the extremity.

Life history. — Unknown ; probably involves intermediate stages in
insects.

Distribution. — Europe (Germany (Greifswald) ).

Various authors (Mehlis, Molin, Stossich, Lepri) consider this
species identical with P. alata Rudolphi, 1819. It is probable that
in the specimens described by Creplin (1 male and 1 female) the
reflection of the cuticle over the lips is accountable for his descrip-

NEMATODE PARASITES OF BIRDS

301

tion of the head, and that he was dealing with liudolphi's species.
Since it is a debatable matter, his description and figures (fig. 362)
are given here for comparison with those of P. alata.

PHYSALOPTERA ALATA CHEVREUXI Seurat. 1914i

Hosts.- — Primary: "Hawk" and Accipiter nisiis; secondary:
Unknown.

Location. — Not given.

Morphology. — Physaloptera (p. 295) : Similar to P. alata except
that this nematode is smaller in size and has the cervical papillae and
excretory pore placed much more posteriorly.

Male 7 mm. long by 515/4 wide. Muscular esophagus 260> long;
entire esophagus 1/4 of body length; cervical papillae 370 and 360/4
from anterior end.

Fig. 362. — Physaloptera megalostoma. After Creplin, 1829

Female (immature), 8 mm. long. Tail long (310/4 ) and conical.
Vulva a short distance posterior to end of esophagus.

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Africa (Sudan (Bone) and Algeria (Bou-Saada) ).

PHYSALOPTERA ALATA NOUVELI Seurat, 1915c

Hosts. — Primary: Accipiter nisus and Aquila chrysaetos; second-
ary: Unknown.

Location. — Esophagus.

Morphology. — (p. 295) : Body robust, usually attenuated anteriorly-
Cervical papillae more or less at the level of the hind end of mus-
cular esophagus. Two lateral lips, each with a large triangular
external tooth and a very small internal tooth with 3 points.

Male 22 to 28.5 mm. long by 900/4 wide. Tail 1.105 mm. long,
slender and pointed; small cuticular tubercles in circumcloacal re-
gion. Caudal papillae as in P. alata. Right spicule robust and
large, 550/* long; left spicule more slender, feebly chitinized and
longer.

Female 33 mm. long by 1.05 mm. wide. Tail 1.1 mm. long, coni-
cal, pointed. Vulva anterior to end of esophagus. 3.8 mm. from
anterior end. Vagina 2.15 mm. long, egg chamber 3.12 mm. long.

302 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

trunk or trompe 500/* long, dividing posteriorly into 2 uteri. Eggs
50 by 25/*, thick-shelled, embryonated.

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Africa (Algeria (Bou-Saada)).

PHYSALOPTERA CRASSA Linstow, 1879b

Host. — Primary : Alauda arvensis; secondary : Unknown.

Location. — Intestine.

Morphology. — Physaloptera (p. 295) : Lips pyramid-shaped, each
with 2 teeth, the inner tripartite, and in addition 2 outer papillae.
Cervical papillae 600/* from anterior end; esophagus 1/4.7 of total
body length.

Male 14 mm. long by 1.5 mm. wide. Tail rounded, its length
about 1/12 of total body length. Six pairs of caudal papillae, the
anterior 5 being pedunculated and situated near the cloacal aper-
ture; the sixth pair non-pedunculated, situated on the inner side of the
fifth pair. Spicules unequal, 660/x and 360/x long.

Female 23 mm. long by 2 mm. wide. Tail rounded, its length 1/22
of total body length. Eggs 49/* long by 26/* wide.

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Europe (Germany (Stuttgart)).

PHYSALOPTERA CROSI Seurat. 1914i

Host. — Primary: Accipiter nisus; secondary: Unknown.

Location. — Proventriculus.

Morphology. — Physaloptera (p. 295) : Body robust. Two lateral
lips (fig. 363 a), each with a large external tooth, strongly chitinized
and an inner tripartite tooth, much smaller and feebly chitinized.
Also 2 pairs cephalic papillae at angle of insertion of lips. Post-
cervical papillae unusually far anterior, immediately posterior to the
nerve ring.

Male 20 mm. long by 660/* wide. Caudal alae (fig. 363 b) well-
developed; circumcloacal region covered with tubercles. Cloacal
aperture 900/* from tail end. Caudal papillae arranged as in P.
galinieri and P. subalata except that in this species there is a sessile
papilla between the 2 papillae of the most posterior ventral pair.
Spicules equal, short, 300/* long.

Female 12 mm. (immature) to 22 mm. long. Tail (fig. 363 c)
short (300/*), conical. Vulva in anterior third of body, 1 mm. pos-
terior to end of esophagus. Vagina shorter than egg-chamber (res-
ervoir), which is 1.2 mm. long by 300/* wide; trunk (trompe) 400/*
long, giving rise to 2 uteri. Eggs 55/t long by 25/* wide, embryo-
nated.

NEMATODE PARASITES OF BIRDS

303

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Africa (Algeria (Mascara and Bordj-Menaiel) ).

PHYSALOPTERA FUSIFORMIS Linstow, 1902

Host. — Primary: Micro pogan, species; secondary: Unknown.

Location. — Head.

Morphology. — Physaloptera (p. 295) : Cuticle thick, enlarged pos-
terior to the lips; narrow cross-striations. Head with 2 large,
lateral, hemispherical lips, each with 3 small papillae on the outer
edge.

Male 18 mm. long by 1.4 mm. wide. Esophagus about 1/4, tail
about 1/23 of total body length. Caudal region (fig. 364) short, heart-
shaped ; laterally 4 pairs of pedunculated papillae; one pair of sessile

Figs. 300-304. — 363, Physaloptera crosi. a, Head end ; b, male tail ; c, female
tail. After Seurat, 1914. 364, Physalopteba fusiformis. Male tail. After
Linstow, 1902

papillae directly anterior and another pair directly posterior to cloa-
cal aperture; near the tail end 3 pairs of sessile ventral papillae and
an unpaired median papilla anterior to them. Linstow's figure
shows longitudinal rows of tubercles in the lateral fields of the tail.

Female 30 mm. long bv 1.97 mm. wide, attenuated at both extremi-
ties. Esophagus about 1/5, tail 1/20 of the total body length. Vulva
in anterior part of body, dividing body length in ratio of 13:56.
Eggs 81/4 long by 26ju, wide, with very thick shells.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — Europe (Germany, Breslau Museum).

PHYSALOPTERA GALINIERI Scurat, 191U

Synonym. — P. alata Rudolphi of Ortlepp, 1922 may be a synonym
of this (see p. 291)).
£$612— 27— 21

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BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hosts. — Primary: Aquila rapax belisaHus and Melierax gdbar;
secondary: Unknown.

Location. — Esophagus and proventriculus.

Morphology. — Physaloptera (p. 295) : Body robust. Mouth (fig.
865#) with 2 lateral lips, each having on its inner surface 3 large
conspicuous, projecting teeth. Two pair of cephalic papillae near
insertion of lips. Postcervical papillae remote from nerve ring,
situated below the end of muscular and glandular esophagus.

Male 21 mm. long by 780/* wide. Caudal alae (fig. 365c) large,
1.5 mm. long, with thick edges. Circumcloacal region covered with
small tubercles (these are not shown in Seurat's figure). Cloacal
aperture 925/* from tail end. Five pairs of lateral pedunculated
papillae, of which 4 pairs are in the circumcloacal region, the other
pair more posterior. Seurat states that there are 11 but figures 13

Fig. 365. — Physaloptera galinieri. a, Head, lateral view ; b, anterior half of

BODY OF FEMALE, SHOWING OVEJECTOR ; C, MALE TAIL. AFTER SEURAT, 1914

ventral papillae near the median line, 3 of them in a transverse row
just anterior to cloacal aperture, 2 pairs just posterior to cloacal
aperture, also arranged in a transverse row, and the other 3 pairs
occurring more posteriorly. Spicules short, subequal, 360/* and
380/i long.

Female 17 to 34 mm. long by 1.04 mm. wide. Tail short* (350/*).
Vulva (fig. 365 b) unusually remote from esophagus, situated a short
distance anterior to middle of body. Vagina 2 mm. long, directed
anteriorly; egg chamber (reservoir) 600/* long; trilnk (trompe) 200/*
long. Two uteri. Eggs 65/* long by 35/* wide.

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Africa (Algeria (Ain-Oussera) and Transvaal).

PHYSALOPTERA GEMINA Linstow, 1899a

Host. — Primary: Gallus gallus and cat (Felis catus doniesticus) ;
secondary: Unknown.

Location. — Stomach and intestine (?) .

Morphology.— Physaloptera (p. 295) : Head end rounded, a ring of
thickened cuticle projecting anteriorly from it. Two large conical

NEMATODE PARASITES OF BIRDS 305

teeth, to each side of these a smaller tooth; internally additional,
still smaller teeth. Four large submedian papillae. Esophagus
about 1/6 of total body length.

Male 11.4 mm. long by 470/* wide. Tail 1/19 of total body length.
Four pairs of long pedunculated papillae (fig. 366), 2 of them
preanal. 2 postanal. Just anterior of the cloacal aperture are 3,
just posterior to it 2, papillae; about midway from cloacal aperture
to tip of tail there are 4 papillae in a transverse row; posterior to
these are 2, making a total of 11 ventral papillae. Circumcloacal
region of bursa covered with longitudinal rows of pointed protuber-
ances.

Female 19 mm. long by 53/t wide. Tail 1/35 of total body length,
conical, rounded at end. Eggs 52/u. long by 32/* wide; thick-shelled.

Life history. — Unknown ; probably involves intermediate stages in
insects.

Distribution. — Africa (Egypt) .

PHYSALOPTERA INFLATA (Molin, 1860) Stossich, 1889

Synonym. — Spiroptera inflata Molin, 18606.

Host. — Primary : Falco unicinctus; secondary : Unknown.

Location. — Esophagus and proventriculus.

Morphology. — Physaloptera (p. 295) : Head continuous with body,
with inflated cuticle. According to Drasche, this species does not
have the 3 small teeth usually found internal to the outer tooth of
the lips, and this character differentiates it from P. acuticauda,
found in the same host. Body densely striated transversely, at-
tenuated posteriorly, truncated anteriorly.

Male unknown.

Female 7 mm. long by 300/* wide. Tail short, conical, with
rounded end. Anus not far from tail end. Vulva in anterior part
of body.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — South America (Brazil).

PHYSALOPTERA SUBALATA Schneider, 1866

Synonym. — L ) hysaloptera alata Iiudolphi, 1819 in part.

Hosts. — Primary : Falco, species and buzzard ; secondary : Un-
known.

Morphology. — Physaloptera (p. 295) : Body large and robust.
Mouth with 2 lips (fig. 367a). According to Seurat each lip with
a small outer tooth, difficult to see, and a large tripartite inner tooth;
Schneider, however, states that the outer tooth is larger than the
inner.

306

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 19 to 32 mm. long by 700/* wide. In a 19 mm. specimen,
caudal alae 1.56 mm. long, cloacal aperture 950/* from caudal ex-
tremity. According to Schneider's figure (fig. 3675), 4 pairs of
lateral pedunculated papillae, 3 preanal ventral and 3 pairs of
postanal ventral papillae. Seurat, however, describes 5 pairs of
long pedunculated papillae, 4 of them in the cloacal region and the
other pair midway between cloacal aperture and caudal end, and 11
median papillae, 3 of them just anterior and 2 pairs just posterior
to the cloacal aperture, the remainder more posterior. Spicules un-
equal, the left 840/* long, slender and pointed, the right not over
400/* long and thicker than the left.

Female 20.5 mm. long by 660/* wide. Tail 240/* long, conical.
Vulva not prominent, in anterior third of body, 3 mm. posterior

Figs. 366-368. — 366, Physalopteka gemina. Male tail. After Linstow,
1899. 367, Physalopteka subalata. «, Lip ; b, male tail. After Schnei-
der, 1866. 368, Physalopteka truncata. a, Lip ; h, male tail. After
Schneider, 1866

to end of esophagus. Vagina 1 mm. long; egg chamber 750/* long;
2 uteri.

Life history. — Unknown ; probably involves intermediate stages in
insects.

Distribution. — South America (Brazil) and Europe (Corsica (San
Martino)).

PHYSALOPTEKA TRUNCATA Schneider, 1866

Hosts. — Primary : Gallus gallus and Phasianus gallus; secondary :
Unknown.

Location. — Proventriculus.

Morphology. — Physaloptera (p. 295) : External median tooth on
each lateral lip is widened anteriorly ; the 3 denticles of the internal
tooth are somewhat heart-shaped, with a cylindrical base and conical
tip (fig. 368«) .

NEMATODE PARASITES OF BIRDS 307

Miile 25 mm. Ion*;. Four pairs of lateral pedunculated papillae
(fig. 3686), of which 2 are preanal, 2 postanal; directly anterior to
cloacal aperture a row of 3 sessile papillae, the central one smaller
than the others; directly posterior to cloaca] aperture is an asym-
metrical cluster of 4 small pedunculated papillae; and posterior to
all the above mentioned papillae are 3 pairs of large sessile papillae
of which the last pair is about equally spaced between the preceding
pair and the tail end. Caudal extremity broadly rounded.

Female 33 mm. long.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — South America (Brazil).

PHYSALOPTERA BILABIATA Creplin, 1829

Synonym. — Spiroptera bilabiata (Creplin, 1829) Dujardin, 1845.

Hosts. — Primary: Lanius minor; secondary: Unknown.

Location. — Intestine.

Morphology. — Physaloptera (p. 295) : Body white; mouth with 2
large projecting lips. Body tapering toward both extremities, espe-
cially the anterior extremity.

Male about 25 mm. long by 1 mm. wide. Tale short, somewhat
recurved, alate, elliptical, with 4 pairs of pedunculated papillae.
One spicule simple, slender, the other undescribed.

Female about 26 mm. long by 1.12 mm. wide. Tail short, conical,
and obtuse.

Life history. — Unknown ; probably involves intermediate stages in
insects.

Distribution. — Europe (Germany (Greifswald) ).

PHYSALOPTERA ABBREVIATA Rudolphi, 1819

This species was described from reptiles and occurs regularly in
reptiles. It has been reported by Linstow (1883) as a pseudo-para-
site in the proventriculus of Ciconia alba, presumably snakes having
formed part of the diet of the bird.

PHYSALOPTERA BREVICAUDA Linstow, 1909b

Host. — Primary: Francolinus adspersus; secondary: Unknown.

Location. — Intestine.

Morphology. — Physaloptera (p. 295) : Cuticle transversely striated.
Head with 2 lips, widened anteriorly, narrower at base, provided in
the middle with a papilla, to the right and left of which' there is a
cone-shaped tooth. Esophagus 1/9 of total body length: tail very
short in both sexes.

Male 27 mm. long by 830/* wide: tail (fig. 309) rounded, its length
1/69 of total body length; caudal alae with chitinous tubercules in

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BULLETIN 140, UNITED STATES NATIONAL MUSEUM

longitudinal rows. Six pairs of pedunculated papillae of which 4
are preanal, 2 postanal, the latter in a transverse row just posterior
to the cloacal aperture. Spicules equal, 1.4 mm. long, their ends
rounded.

Female 42 mm. long by 950/x wide. Tail slender, cone-shaped,
pointed at end. Vulva just posterior to middle of body, dividing the
body length in a ratio of 23 : 21. Eggs 39/a long by 26/a wide, thick-
shelled.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — South West Africa.

Ortlepp notes that this species shows certain differences from
Physaloptera, notably the shape of the lips, the arrangement of
caudal papillae, and the position of the vulva, and he lists it under
Species Inquirendae.

Figs. 371-373. — 371, Physaloptera, species. Headend. After Parona, 1885. 372,
1909. 370, Physaloptera malleus, a, Head; b, male tail. After Linstow,
1883

PHYSALOPTERA MALLEUS Linstow, 1883

Host. — Primary : Corvus comix; secondary : Unknown.

Location. — Not given.

Morphology. — Physaloptera (p. 295) : Head (fig. 370#) with 2 very
large rounded lips, hollowed out on the inner surface. Esophagus
1/6.5 of total length.

Male 20 mm. long by 760/a wide. Tail 1/33.3 of total body length.
Caudal region (fig. 3705) oval; 4 pairs of preanal, 2 pairs of post-
anal, pedunculated papillae ; an unpaired median papilla on anterior
margin of cloacal aperture. Caudal alae covered with round tu-
bercles arranged in transverse rows. Spicules unequal, the larger
having a double barb.

Female 42 mm. long by 1.5 mm. wide. Tail short, only 1/269 of the
total body length. Eggs 46/x long by 29/u, wide, double-shelled, the
outer shell being the thicker.

Life history. — Unknown ; probably involves intermediate stages in
insects.

Distribution. — Asia ( Turkestan ) .

NEMATODE PARASITES OF BIRDS 309

Orlepp notes that this is probably not a Physaloptera, and places
it in Species Inquirendae, the rounded lips without teeth and
the arrangement of the caudal papillae differing from those in
Physaloptera.

PHYSALOPTERA. species Parona. 1885

Host. — Primary : Ibis aethiopica; secondary : Unknown.

Location. — Orbital cavity.

Morphology. — Physaloptera (p. 295).

Male unknown.

Female 32 mm. long. Head (fig. 371) with winglike appendage
and 2 or more papillae or lips.

Life history. — Unknown.

Distribution. — Africa (Abyssinia) .

The so-called " winglike appendages " of the head in the figure of
this species resemble the cuticular sheath which projects forward
over the head of several species of Physaloptera. The description is
too inadequate for any decision as to the status of the species.

PHYSALOPTERA STRONGYLINA Rudolphi, 1819

Synonym. — Spiroptera affinis Dujardin, 1845.

Hosts. — Primary : Cuculus melacoryphus and C. seniculus; second-
ary : Unknown.

Location. — Stomach and intestine.

Morphology. — Physaloptera (p. 295) : Head continuous wtih body.
Mouth with lips. Body much attenuated anteriorly.

Male 6 to 8 mm. long. Caudal alae vesicular. Spicules fairly long
and thick.

Female 15 to 21 mm. long. Posterior extremity straight, rounded.

Life history. — Unknown.

Distribution. — South America (Brazil).

Ortlepp notes that this species is wrongly attributed to this genus
but that the description is too inadequate to warrant its transfer to
any other genus.

PHYSALOPTERA OVATA Linstow. 1907

Hosts. — Primary: Astur melanoleucus ; secondary: Unknown.

Location. — Proventriculus.

Morphology.— Physaloptera (p. 295) : Cuticle with transverse stria-
tions. Mouth opening surrounded by 6 cones, behind them a circle
of 4 large stalked submedian papillae. A thickening of the cuticle
projects forward in a 55(V long spindle-shaped swelling over the
head end.

Male 13.8 mm. long by 66(V wide. Esophagus 1/4.4 of total body
length. Caudal alae (fig. 372) oval, smaller in posterior part; large

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BULLETIN 140, UNITED STATES NATIONAL MUSEUM

longitudinal and smaller transverse striations. Five pairs of long
pedunculated papillae, of which 2 pairs (according to Linstow's fig-
ure the 2 preanal) are longer than the others (postanal). Spicules
equal, 290/* long, curved. Tail 1/14 of total body length.

Female 16.8 mm. long by 1.22 mm. wide. Esophagus 1/3.5 of total
body length. Tail 1/53 of total body length, tapering. Eggs 42//, long
by 13/i wide. Vulva anterior to middle of body, dividing the body
length in ratio of 13 : 31.

Life history. — Unknown; probably involves intermediate stages in
insects.

Distribution. — Africa (Kamerun) .

Ortlepp notes that this species does not belong in Physaloptera,
that it has some resemblance to the genus Cyrnea, but its inadequate
description makes any transfer inadvisable.

Figs. 371-373. — 371, Physaloptera, species. Head end. After Parona, 1885. 372',
Physaloptera ovata. Male tail. After Linstow, 1907. 373, Physaloptera
bulbosa. Male tail. After Linstow, 1906

PHYSALOPTERA BULBOSA Linstow, 1906b

Hosts. — Primary : Pavo cristatus and P. spicifer; secondary : Un-
known.

Location. — Proventriculus.

Morphology. — Physaloptera (p. 295) : Head end rounded, the
mouth surrounded by 6 lips structures. Cuticle smooth. Nerve ring
250/x from head end.

Male 18.4 mm. long by 400/t wide. Esophagus 1/5 of total body
length. Right spicule 880/* long, left spicule 2.17 mm. long; both
spicules with rounded tips. Tail 1/46 of body length. Caudal alae
(fig. 373) oval, with transverse wavelike markings. Four pairs
of pedunculated papillae, of which 2 are preanal, 2 postanal, and
between the two groups on each side a large sessile adanal papilla;
near the tail end there are 5 pairs of smaller pedunculated papillae
forming a row on each side.

Female 27.8 mm. long by 510/* wide. Esophagus 1/7 of total body
length. Anus almost terminal, the tail length being only 1/121 of
total body length. Vulva in posterior part of body, 2/47 or body
length from tail end. Eggs thick-shelled, 44/i. long by 26/t wide.

NEMATODE PARASITES OF BIRDS 311

Life history. — Unknown; probably involves intermediate stages
in insects.

Distribution. — Europe (Germany (Koenigsburg) ).

Ortlepp (1922) states that he has studied specimens of this species
from Pavo cristatus and that the species belongs in the genus Cyrnea.
The detailed description which he says he will give at a later date
has not yet appeared. The lip structure and the position of the
vulva of the nematode is not compatible with Physaloptera but the
above description is inadequate for its assignment to another genus.
Ortlepp's redescription and transfer of this species to Cyrnea ap-
peared after the above had been sent to press. See Addenda, p. 390.

Family THELAZIIDAE Railliet, 1916a

Synonym. — Thelaziidae Skrjabin, 19166. Railliet's paper pro-
posing this new family appeared in March, 1916, and Skrjabin's
paper in October, 1916.

Family diagnosis. — Spiruroidea (p. 162) : Mouth without lips but
provided with papillae; a short buccal cavity present. Male with
or without caudal alae; preanal papillae usually very numerous;
postanal less numerous. Spicules usually very unequal. Female
with tail generally blunt. Situation of vulva variable. Two uteri.
Oviparous, ovoviviparous or viviparous.

Parasitic in orbital region of mammals or birds.

Type genus. — Thelas'm Bosc, 1819.

Railliet included in this family the genera Schistoropkus and
Serticeps, but Travassos subsequently removed them and put them
in the family Schistorophinae, under the family Acuariidae. The
present writer has followed this latter classification ; the above
diagnosis of the Thelaziidae is therefore an emendation of that of
Railliet. Railliet's assignment of the family to the Spiruroidea
is followed here rather than Travassos's assignment of it to the
Filarioidea.

KEY TO GENEKA OF THELAZIIDAE

1. Male with large caudal alae Ceratospira, p. 319.

Male without caudal alae 2.

2. Vulva in anterior part of body Thelazia, p. 311.

Vulva in posterior part of body Oxyspirura, p. 321.

Genus THELAZIA Bosc, 1819

Synonym. — Thelazius Bosc, 1819.

Generic diagnosis. — Thelaziidae (p. 311) : Mouth without lips.
followed by a buccal cavity, the anterior edge of which has 6 pro-
jecting lobes. Two lateral and 4 submedian cephalic papillae. Male
with caudal extremity obtuse, usually recurved, without lateral alae.
:;r,i2— 27— 22

312 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

A large number of preanal papillae of which one is sometimes median,
unpaired, just anterior to the cloacal aperture (the bird forms do
not have this papilla present usually; exceptions are Thelazia lutzi
and T. campanulata) . Postanal papillae number 3 or 4. Two un-
equal spicules. Female with blunt, conical, rounded caudal extrem-
ity, provided with 2 lateral papillae. Vulva in anterior part of
body, usually slightly posterior to the posterior end of esophagus.
Two uteri, directed posteriorly. Embryos hatch in uterus.

Parasitic normally in the ducts of the lacrimal glands of mam-
mals; certain forms occur under the nictitating membrane of the
eye in birds.

Type species. — Thelazia rhodesi (Desmarest, 1828) Blainville,
1828, emend. Raillet and Henry, 1910.

KEY TO SPECIES OF THELAZIA

1. Description incomplete ; mouth large, the 6 lobes of buccal cavity small, each

with a finger-like process extending inward ; from Crax fasciolata.

Thelazia anolabiata, p. 317.

Mouth and lobes of buccal cavity not described as above; from hosts other

than above 2.

2. Male 22 to 24 mm. long ; female 25 to 27 mm. long ; 10 pairs of preanal papil-

lae; cloacal aperture 85m from posterior end Thelazia lutzi, p. 315.

Male smaller than above (up to 20 mm. in T. papulosa) ; female smaller
than above except in T. papulosa (up to 26 mm.) ; in T. papulosa, as
in all other species where number is given, less than 10 pairs of preanal
papillae ; cloacal aperture, if described, less than 85m from posterior
end 3.

3. No cuticular annulations ; no buccal cavity ; no caudal papillae in male.

Position in this genus doubtful Thelazia cirrura, p. 318.

Cuticular annulations, buccal cavity, and caudal papillae present 4.

4. Cervical region with rows of delicate posteriorly directed spines ; left

spicule 6 times as long as right spicule Thelazia papillosa, p. 317.

Cervical region without spines ; left spicule, if lengths given, more than
6 times as long as right spicule 5.

5. Esophagus not over 600/* long ; anus of female 300m or farther from pos-

terior end 6.

Esophagus 750m long or longer; anus of female not over 225m from posterior
end 7.

6. Maximum length of esophagus 340m ; vulva 680m from anterior extremity.

Thelazia stereura, p. 316.
Length of esophagus 460 to 600m ; vulva not over 500m from anterior extremity.

Thelazia campanulata, p. 313.

7. Female with buccal cavity 33m deep ; esophagus 925m long ; anus 85m from

posterior end Thelazia cholodkowskii, p. 313.

Female with buccal cavity not over 28m deep ; esophagus not over 850m
long; anus 120m or farther from posterior end '. 8.

8. Female 18 mm. long; anus 120m from tail end Thelazia digitata, p. 319.

Female not over 14 mm. long ; anus 255m from tail end.

Thelazia dacelonis, p. 314.

NEMATODE PARASITES OF BIRDS

THELAZIA CAMPANULATA (Molin, 1858) Railliet and Henry, 1910

313

Synonyms. — Filaria oami'panulata Molin, 1858; Filaria falco-nis
magnirostns Molin, 1858.

Hosts. — Primary: Falco magnirostris (Rupornis m/ignirostris) ;
secondary : Unknown.

Location. — Under nictitating membrane.

Morphology. — Thelazia (p 311) : Anterior extremity of body atten-
uated, obtuse. Cuticular surface annulated, the annulations compan-
uliform and imbricated. Buccal cavity (fig. 374a) 17 to lit*, deep,
according to Travassos, by 28 to 35/* wide. Esophagus subcylindri T
cal, 460 to GOO/i long.

Male 17 mm. long by 400/t wide. Caudal extremity (fig. 3746)
curved toward the ventral surface; cloacal aperture 140/* from end.

osmm

Fig. 374. — Thelazia campandlata. a, Head end ; b, male tail ; c, female
tail. After Tkavassos, 1918

Seven pairs of preanal papillae and I unpaired median papilla
anterior to cloacal aperture; posterior to the cloacal aperture 1 large
median papilla and 3 pairs of papillae. Spicules very dissimilar
and unequal, one 190/* long by 28/* wide, alate laterally at its distal
extremity, the other 2.67 mm. long by 7/* wide, with a slight swell-
ing 640/1. from the proximal end.

Female 15 to 23 mm. long by 500 to 600/* wide. Tail (fig. 371c)
nearly straight, pointed. Vulva about 350/* from anterior end of
body; anus 300//. from posterior end, in a specimen 23 mm. long.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

THELAZIA CHOLODKOWSKII Skrjabin, 1922

Hosts. — Primary: Caprimulgus europaeus; secondary: Unknown.

Z o cation: — Ey e .

Morphology. — Thelazia (p. 311).

Male unknown.

314 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Female 14 mm. long. Width of body 340/* at level of esophagus
and intestine, 470/* in middle region of body, 200/* at level of anus.
Buccal cavity (fig. 375a) 33/* deep; esophagus cylindrical, 925/* long
by 70/* wide. Nerve ring 220/* from anterior end of body. Vulva
500/* from anterior end of body. Oviduct 90/* wide. Anus 85/* from
tail end (fig. 375c). Eggs 58/* by 30/*.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution: — Europe (European Russia).

As the present writer had great difficulty in translating the Russian
description of this species, the above must be accepted with reser-
vations.

THELAZIA DACELONIS (Breinl, 1913) Travassos, 1918a

Synonym. — Filaria dacelonis Breinl, 1913.

Hosts. — Primary : Dacelo leachii; secondary : Unknown.

Location. — Conjunctival sac.

Fig. 375. — Thelazia cholodkuwskii. Female, a. Head end ; b,

CUTICULAK ANNULATIONS ; C, TAIL. AFTER SKRJABIX, 1022

Morphology. — Thelazia (p. 311) : Body tapering toward both ends ;
f.nterior end bluntly rounded, posterior end pointed. Cuticula with
distinct transverse striations. Mouth (fig. 376a) oval, surrounded
by a chitinous (or cutinous) ring with clefts which give the appear-
ance of 6 lips. No papillae discernible, according to Breinl's descrip-
tion; his figure, however, shows what are apparently 2 cephalic
papillae. Buccal cavity 24 to 28/u. long, cup-shaped. Esophagus 750
to 825/* long, club-shaped.

Male 10 to 11 mm. long by 390 to 400/* wide. Tail (fig. 376c)
curved ventrally, its end bluntly rounded. Cloacal aperture 180/*
from posterior end. Caudal papillae variable, in one specimen 5
pairs of preanal and 2 pairs of postanal papillae; in another speci-
men 4 pairs of large preanal and 1 pair of postanal papillae.
Spicules dissimilar and unequal; one 2.1 mm. long by 21/* wide at
proximal end and 10/* wide at distal end, the other 180/* long by 30/*
wide with its proximal end funnel-shaped and distal end rounded.

Female 11 to 14 mm. long by 500/* wide. Vulva (fig. 376a) 525
to 555/* from anterior end of body. Vagina 1 mm. long. Anus 255/*
from tail end (fig. 3676), and provided with a strong muscle.

NEMATODE PARASITES OF BIRDS

315

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Australia (Queensland) .

THELAZIA LUTZI Travassos. 1918a

Hosts. — Primary: Penelope, species; secondary: Unknown.
Location . — Eye ?

Fig. 376. — Thelazia dacelonis. a, Head end, showing vulva ; b, tail of
female; c, tail of male. After Breinl, 1913

Morphology. — Thelazia (p. 311) : Transverse striations of cuticula
strongly developed (377a). Buccal cavity 35 to 49/* wide by 28/z
deep. Esophagus slightly claviform, about 670 to 740/* long by 42
to 49/* wide.

Male 22 mm. long by 400 to 500/* wide. Caudal extremity (fig.
378) curved ventrally. Ten pairs of preanal papillae, 1 unpaired

Fig. 377. — Thelazia lut/.i. Female. a, Head end, showing vulva; b, tail.
After Travassos, 1918

papilla anterior to clpacal aperture; 2 pairs of postanal papillae
and 1 unpaired papilla. Spicules unequal and dissimilar; one 740/*
long by 7/i wide, the other 190/* long by 35/* wide. Cloacal aperture
85/* from the tail end, the latter obtuse and rounded.

Female 25 to 27 mm. long by 500 to 800/* wide. Vulva (fig. 377a)
530/* from head end. salient. Anus 100/* from tail end (fig. 3776),
the latter obtuse.

316

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

THELAZIA STEREURA (Rudolphi, 1819) Railliet and Henry, 1910

/Synonyms. — Spiroptera stereura Rudolphi, 1819; ? Oxyspirura
stereura (Rudolphi, 1819) Ransom, 1904.

Fig. 378.- — Thelazia lutzi. Male tail. After Travassos, 1918

Hosts. — Primary : Buteo vulgaris, B. vulpinus, Falco naemus
(=AquUa naevia=A. maculata) ; secondary: Unknown.

Location. — Under nictitating membrane and in auditory meatus.

Morphology. — Thelazia (p. 311) : Body thick, attenuated ante-
riorly with the head end truncate. Cuticular annulations with nu-
merous small campanuliform folds. Head (fig. 379 a) without mem-
branous appendages. Mouth orbicular, without lips. According to
Skrjabin's figure, the cuticular striations do not cross the lateral lines

Fig. 379. — Thelazia stereura. Female, u, Head end, showing
vdlva ; b, tail. After Skrjabin, 1922

but leave a smooth area there; the lateral lines end at the level of
the anus.

Male 12 to 15 mm. long by 800/x wide. Tail coiled once spirally ;
no caudal alae. Posterior extremity extended into a short appendage
with knob like end. Two unequal spicules ; the short one styloid,
with a sharp point, and the long one filiform. Schneider states that
there are more than 4 pairs of preanal papillae.

Female 12 to 18 mm. long by 1 mm. wide. Two lateral cervical
spines. Buccal cavity 51/* deep by Q8/x wide in large specimens.
Esophagus 34(V long. Tail (fig. 379&) straight, acutely conical,
prolonged into a styloid appendage with a rounded knoblike end;

NEMATODE PAEASITES OF BIRDS 317

anus at base of the appendage, 370/a from tip of tail. Vulva (fig.
379 a) in anterior part of body, 680> from anterior end in large speci-
mens, at a level just posterior to the middle of the esophagus. Eggs
51/x by 25/x.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Europe (Austria (Vienna Museum), Croatia, and
(?) Russia).

THELAZIA PAPILLOSA (Molin, 1860) Railliet and Henry, 1910

Synonyms. — Spiroptera papillosa Molin, 18606/ Spiroptera fal-
conis hptopodis Molin, 1860; Spiroptera falconis gavi-al redlis Molin,
1860; Spnroptera falconis Molin, 1860, not Rudolphi, 1819; {? Oxy-
spirura) papillosa (Molin, 1860) Ransom, 1904.

Hosts. — Primary: Falco destructor { — Thrasactus harpyia), F.
gracilis {Geranospizias caerulescens) ; secondary: Unknown.

Location. — Under nictitating membrane.

Morphology. — Thelazia (p. 311) : Anterior end abruptly attenuated
with very obtuse rounded apex. Cuticula densely annulated. Neck
with rows of very delicate spines, posteriorly directed, each row
corresponding to a cuticular ring or pseudoannulation. Mouth
(fig. 380) large, orbicular, with tumid border. Lips absent. Two
lateral and 4 submedian cephalic papillae.

Male 8 to 20 mm. long by 300/a wide. Posterior extremity more
attenuated than anterior, the tip of the former obtuse, rounded, and
only slightly curved. Molin describes 7 pairs of preanal papillae,
but Drasche, in redescribing the same material, states that there are
4 pairs of preanal and 2 pairs of postanal papillae. Right spicule
short and broad, with blunt point; left spicule six times as long as
right, slender, filiform and alate.

Female 15 to 26 mm. long by 300 to 500^i wide. Tail gradually
attenuated, tip round, obtuse. Anus remote from posterior end,
prominent, its 2 lips tumid. Vulva in anterior part of body, not
far from mouth.

Life hist&ry. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

THELAZIA ANOLAB1ATA (Molin. 1860) Kailliet and Henry, 1910

Synonyms. — Spiroptera anolabiata Molin, 18606/ Spiroptera
cracis alectoris Molin, 1860; Filaria anolabiata (Molin, 1860) Stos-
sich, 1897; {? Oxyspirura) anolabiata (Molin, I860) Ransom, 1904,

Hosts.. — Primary: Crax fasciolata\ secondary: Unknown.

Location. — Under nictitating membrane and free in the eye.

318

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Morphology.— Thelazia (p. 311) : Anterior end of body abruptly
attenuated, apex truncate. Body encircled with cuticular annula-
tions with sharply cut posterior borders. Mouth (fig. 381) large,
the 6 lobes of the anterior edge of the buccal cavity are small, each
with a fingerlike process extending inward from the periphery of
the head. Buccal cavity short and wide.

Male unknown.

Female 8 to 17 mm. long by 40(V wide. Tail straight, abruptly
conical, with blunt curved tip.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

Figs. 380-382. — 380, Thelazia papillosa. Head ; front view. After
Drasche, 1884. 381, Thelazia anolabiata. Head. After Drasche,
1884. 382, Thelazia ( ?) cirrdra. Head end, showing vulva.
Original

THELAZIA (?) CIRRURA (Leidy. 188G) RaiUiet, 1916

Synonym. — Filaria cirrura Leidy, 1886.

Hosts. — Primary: Quiscalus major ( = Megaguiscualus major)]
!-,econdary: Unknown.

Location. — Orbit of eye.

Morphology. — Thelazia (?) (p. 311) : Body cylindrical. Anterior
extremity conical, rounded. Mouth (fig. 382) a minute funnellike
orifice without papillae or internal armature. No buccal cavity.
No transverse striations of cuticula.

Male 10 mm. long by 375/* wide. Caudal extremity closely rolled
inward, conical, blunt, without alae or papillae. Spicules strongly
curved. Cloacal aperture prominent.

Female 16 mm. long by 500/i wide. Esophagus 863/a long. Cau-
dal extremity slightly curved or nearly straight, conical, obtusely
rounded. Vulva (fig. 382) about 400ju from anterior end of body.

IAfe history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — North America (United States (Florida)).

NEMATODE PARASITES OF BIRDS

319

The present writer has reexamined part of Leidy's original mate-
rial, but on account of the poor condition of the material was unable
to add to the description except as regards the esophageal length;
a figure of the anterior region of the body of the female has also
been made. The position of the species in this genus is very doubt-
ful but a reassignment to another genus is inadvisable with the in-
adequate description.

THELAZIA (?) DIGITATA Travassos, 1918a

Hosts. — Primary: Rampluistus, species; secondary: Unknown.

L ocatio n . — Eye.

Morphology. — Thelazia (p. 311) : Anterior extremity digitiform.
Cuticula transversely annulated. Buccal cavity about 21/a deep by
28/i wide. Esophagus subcylindrical, about 850ju, long by 87/i, wide.

Male unknown.

Fig. 3S. r

-Thelazia ( ?) digitata. Female, a. Head end, showing vulva ;
b, tail. After Tkavassos, 1918

Female 18 mm. long by 500/u wide. Vulva (fig. 383 a) 600> from
anterior extremity, its lips salient. Caudal extremity (fig. 383 5)
obtuse, curved towards the ventral face ; anus 120> from end.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil).

Genus CERATOSPIRA Schneider, 1866

Synonym. — Ancyracanthus Diesing, 1838, in part.

Generic diagnosis. — Thelaziidae (p. 311): Mouth surrounded by
papillae and followed by a short buccal cavity. Male with very short
blunt tail provided with large alae and numerous simple caudal
papillae, of which 9 to 11 pairs are preanal. Two very unequal
spicules. Female with very short blunt tail. Vulva in anterior
part of body. Sometimes viviparous. Parasitic in orbital cavity
of birds.

Type species. — Ceratospira vesiculosa Schneider, 1SG6.

320 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

KEY TO SPECIES OF CERATOSPIRA

1. Male 14.6 mm., female 18 mm., long. Vulva near middle of esophagus. Long

spicule 968m in length Ceratospira ophthalmica, p. 320.

Male 20 mm. long; female length unknown. Vulva near posterior end of
esophagus. Long spicule over 3 mm. in length.

Ceratospira vesiculosa, p. 320.

CERATOSPIRA VESICULOSA Schneider, 1866

Hosts. — Primary: Psittacus sinensis (Eleotus pectoralis) ; second-
ary: Unknown.

Location. — Orbital cavity.

Morphology. — Ceratospira (p. 319) : Head rounded, with (?) pa-
pillae. Buccal cavity short. Cuticula marked by widely separated
annulations with sharp projecting edges.

Male 20 mm. long. Caudal extremity rolled spirally and with
thick vesicular alae. Caudal papillae (fig. 384) asymmetrical, 11
on one side, 12 on the other ; 1 pair is near tip of tail, 3 pairs form a
group on either side of cloacal aperture, and the preanal papillae
(7 on left side, 8 on right side) are distributed at unequal intervals.
Two unequal spicules, one very short and cornet-shaped, the other
very slender and over 3 mm. long.

Female length unknown (specimen incomplete). Tail rounded.
Vulva in anterior part of body, near posterior end of esophagus.
Viviparous.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Europe (Germany (Zoological Garden, Berlin)).

CERATOSPIRA OPHTHALMICA (Linstow, 1898) Ransom, 1904

/Synonym. — Ancyi'acanthus ophthalmicus Linstow, 1898.

Hosts. — Primary: Carpophaga brenchleyi (=Zonoenas breneh-
leyi) ; secondary : Unknown.

L^ocation. — Orbital cavity.

Morphology. — Ceratospira (p. 319) : Head (fig. 385a) surrounded
by 6 cephalic papillae. Buccal cavity wide. Caudal extremity
rounded in both sexes. Cuticula with transverse annulations 13/*
wide, with swollen posterior edges.

Male 14.6 mm. long by 390/x wide. Esophagus 1/18, tail 1/370, of
body length. Caudal end curved hook-like, with broad alae. Nine
to 10 pairs of preanal and 4 pairs of postanal papillae (fig. 3855).
Spicules unequal, the right one short (264/x) and broad, the left long
(968/x) and slender.

Female 18 mm. long by 470/x wide. Esophagus 1/23, tail 1/54, of
body length. Vulva near anterior end of body, about at the middle

NEMATODE PARASITES OF BIRDS

321

of the esophagus, dividing the body length in the ratio of 1:45.
Eggs 23/a by 16/z, very numerous.

Life Mstory. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Bismarck Archipelago.

Genus OXYSPIRURA Drasche in Stossich, 1897

Generic diagnosis. — Thelaziidae (p. 311) : Mouth without lips, sur-
rounded by 2 lateral and 4 submedian cephalic papillae. A short

Figs. 384-385. — 384, Ceratospira vesiculosa. Male tail. After Schneider,
1866. 385, Ceratospira ophthalmica. a, Head end; b, male tail. After
I.instow, 1898

buccal cavity or pharynx present. Membranous alae on head gener-
ally lacking. Tail very slender, acutely pointed. Male with tail
curled or in spiral, without caudal alae; caudal papillae present,
sessile, the preanal of very variable number ('2 to 28), the postanal
(1 to 8) often asymmetrical. Two unequal spicules, one long and
filiform, the other short and thick. Female with straight tail. Vulva
in posterior part of body, a short distance anterior to the anus.

Parasitic under nictitating membrane in birds.

Type species. — Oxyspieura cepfmloptem (Molin, L860) Stossich,
1897.

KEY TO SPECIES en' ciXVSI'IRI'K.V

1. Head with cuticular expansions, at least in the female 2.

Head without cuticular expansions, or none described, or inconstant (in
O. parvovion sometimes present in male) 5.

322 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

2. Cuticular expansions absent in male, 4 in number in female in lateral,

dorsal, and ventral fields ; 2 pairs of postanal papillae in male.

Oxyspirara heteroclita, p. 331.

Cuticular expansions 2 in number in both sexes, situated laterally; more

than 2 pairs of postanal papillae 3.

3. Female 21 mm. long ; left spicule twice as long as right.

Oxyspirura brevisubulata, p. 325.
Female not over 13 mm. long; left spicule 5 times as long as right 4.

4. Male 8 to 10 mm. long; caudal extremity of male coiled in a single turn;

2 (?) pairs of preanal papillae Oxyspirura anacanthura, p. 323.

Male 13 to 15 mm. long; caudal extremity of male coiled twice; 7 pairs of
preanal papillae Oxyspirura cephaloptera, p. 322.

5. Male 8.5 mm., female 8 to 9 mm., long; from Anthochaera carunculata.

Oxyspirura anthochaerae, p. 324.

Male more than 8.5 mm. long (may be as little as 8.9 mm. in O. pwr-

vovum) ; female more than 9 mm. long (may be as little as 9.8 mm. in

0. parvovum) ; from hosts other than above 6.

6. Spicules nearly equal in length and of same shape; position in genus ques-

tionable Oxyspirura brevipenis, p. 332.

Spicules of unequal length and dissimilar shape 7.

7. Female 9.8 mm. long ; male with about 28 pairs of preanal papillae ; spicules

470m and 250m long Oxyspirura siamensis, p. 329.

Female 11 mm. long or longer ; male with not more than 4 pairs of preanal
papillae, or if more numerous (up to 26 pairs), spicules different from
above 8.

8. Female 21 mm. long; male with about 26 pairs of preanal and 1 pair of

postanal papillae; spicules 180m and 290m long.

Oxyspirura ophthalmica, p. 328.

Female less than 21 mm. long (may be as much as 20 mm. in O. parvovum) ;

preanal papillae apparently absent or less numerous than above ; spicules

of lengths different from above 9.

9. Preanal papillae apparently absent; 4 pairs of postanal papillae; left

spicule 2M times as long as right spicule— Oxyspirura sygmoidea, p. 330.

Preanal papillae present, the total number being 5 to 8 : not as many as 4

pairs of postanal papillae ; left spicule either shorter or considerably

longer than 2^ times the length of right spicule 10.

10. Mouth surrounded by a 6-lobed chitinous ring; 4 pairs of preanal papillae;

eggs 50 to 65^ by 40 to 45m Oxyspirura mansoni, p. 325.

Mouth not surrounded by a 6-lobed chitinous ring ; not more than 3 pairs
of preanal papillae; eggs not over 46m by 29m 11-

11. Male 9.2 mm. long ; 3 pairs of preanal and 2 pairs of postanal papillae ; left

spicule 3.4 to 4.1 mm. long ; anus of female 390 to 440m from posterior end.

Oxyspirura parvovum, p. 328.

Male 13 mm. long; caudal papillae very asymmetrical, a total of 5 preanal

and 7 postanal papillae ; left spicule 600m long ; anus of female 510m from

posterior end Oxyspirura tanasijtchuki, p. 330.

OXYSPIRURA CEPHALOPTERA (Molin, 1860) Stossich, 1897

Synonyms. — Spiroptera cephaloptera Molin, 1860; Spiroptera
momoti brasiliensis Molin, 1860; Spiroptera orioli Molin, 1860;
Cheilospirura cephaloptera (Molin, 1860) Diesing, 1861.

NEMATODE PARASITES OF BIRDS 323

Hosts. — Primary: Momotus brasiliensis (=Mo?notus momota)
and Icterus croconotus; secondary : Unknown.

Location.. — Under nictitating membrane.

Morphology. — Oxyspirura (p. 321) : Body gradually attenuated
anteriorly, with rounded apex. Tail awl-shaped, drawn out into a
slender acute point. Cuticula with dense transverse striations. Head
(fig. 386 a) with membranous alae dilated and rounded anteriorly.
Mouth hexagonal, without lips, surrounded by a membranous border;
2 lateral and 4 submedian cephalic papillae.

Male 13 to 15 mm. long by 200/x, wide. Tail (fig. 386 b) coiled in
2 turns. Caudal alae absent. Seven pairs of preanal and 6 pairs
of postanal papillae, the latter more or less inconstant and asym-
metrical. Spicules unequal, the right short, thick, navicular, with
blunt rounded tip, the left filiform, pointed, alate, five times as long
as the right.

Female 10 to 13 mm. long by 300/x wide. Anus remote from end
of tail. Position of vulva not determined.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

OXYSPIRURA ANACANTHURA (Molin, 1S60) Stossich, 1897

Synonyms. — Spiroptera anacanthura Molin, 1860; Spiroptera
crotoph-agae ani Molin, 1860; Spiroptera crotop/iagae majoris Molin,
1860.

Hosts. — Primary : Crotophaga ani, C. major; secondary : Unknown.

Location. — Under nictitating membrane.

Morphology. — Oxyspirura (p. 321) : Body straight and slender,
sharply pointed posteriorly. Head with 2 voluminous lateral cuti-
cular membranes (Molin described 4 membranes, but Drasche, re-
examining the same material, found only 2). Mouth large, circular,
gradually attenuated toward both ends, truncated anteriorly and
surrounded by 6 small membranous lobules and 4 submedian papillae.
Buccal cavity or pharynx short. Cuticula with fine transverse
striations.

Mule 8 to 10 mm. long by 100^ wide. Caudal end (fig. 387«) coded
in a single turn and without alae. Two pairs of preanal papillae and
a variable number, up to 5 pairs of postanal papillae, asymmetrically
arranged. Two unequal spicules, the right (fig. 387 6) one short,
thick, with boat-shaped distal end, the left one slender, sharply
pointed, alate, and five times as long as the right spicule.

Female 11 to 13 mm. long by 200 fi wide. Vulva prominent, a short
distance anterior to anus.

324 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil).

OXYSPIRURA ANTHOCHAERAE (Johnston, 1912) Johnston, 1912

Synonyms. — Ascaris, species Krefft, 1873; Ceratospira antho-
chaerae Johnston, 1912.

Hosts. — Primary: Anthocha-era carunculata; secondary: Un-
known.

3$6.

f/o/nm

Figs. 386-388. — 386, Oxyspirura cephaloptera. a, Male tail ; b, head, front
view. After Drasche, 1884. 387. Oxyspirura anacanthura. a, Right spicule ;

6, MALE TAIL. AFTER DRASCHE, 1884. 388, OXYSPIRURA ANTHOCHAERAE. a, HEAD
END; 1), TAIL END OF FEMALE. AFTER JOHNSTON, 1912

Location. — Eye.

Morphology. — Oxyspirura (p. 321) : Cuticle finely striated trans-
versely. Buccal cavity present. Nerve ring (fig. 388a) surrounds
first part of esophagus, 195/* from anterior end. Specimens studied
were in bad condition, owing to drying, so that complete descrip-
tion was not possible.

Male about 8.5 mm. long by 110/* wide. Cloaca! aperture 80/a from
posterior extremity, which is spirally curled. Caudal papillae not
discernible.

Female 8 to 9 mm. long by 110/a wide. Posterior extremity sharply
pointed. Anus 194/x, vulva 320/*, from posterior end. (Fig. 388 o.)

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Australia.

NEMATODE PARASITES 01 BIRDS 325

OXYSPIRURA BREVISUBULATA (Molin, 1860) Stossich, 1897

Synonyms. — Spiroptera brevisubulata Molin, 1860; Spiroptera
strigis Molin, 1860.

Hosts. — Primary: Strix airicapilla (=Otus choliba) ; secondary:
Unknown.

Location. — Under nictitating membrane.

Morphology. — Oxyspirura (p. 321) : Body filiform, gradually at-
tenuated anteriorly, apex truncate. Two lateral cervical spines (pa-
pillae), directed posteriorly. Molin described 4 membranous alae on
the head, but. Drasche, re-examining the same material, failed to find
these but describes 2 lateral bladderlike expansions of the cuticula
between the head and the cervical papillae. Mouth hexagonal, with-
out lips, surrounded by 2 lateral and 4 submedian papillae.

Male 15 mm. long by 300ft wide. Tail coiled in 2 turns, short,
abruptly subulate, with acute point. Caudal alae absent. Four pairs
of preanal and (?) 8 pairs of postanal papillae. Spicules unequal,
the right short and thick, the left filiform and twice as long as the
right.

Female 21 mm. long by 500/x wide. Tail short, subulate, sharply
pointed. Anus remote from tail end. Vulva very prominent, a short
distance anterior to anus.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

OXYSPIRURA MANSONI (Cobbold, 1879) Ransom. 1904

Synonyms. — Filaria mansoni Cobbold, 1879; Spiroptera emmer-
ezii Emmerez and Megnin, 1901; Spiroptera mansoni (Cobbold,
1879) Gedoelst, 1903.

Hosts. — Primary : Gall us gallus, Meleagris gaJlopavo, Pavo cris-
tatus; secondary : Unknown.

Location. — Under the nictitating membrane and, occasionally, in
nasal cavities and sinuses.

Morphology. — Oxyspirura (p. 321) : Body attenuated at both ends,
the anterior end rounded, the posterior end pointed. Cuticula
smooth. No membranous appendages. A pair of small papillae
near the tip of the tail in both sexes. Mouth (lig. 389 a and b) cir-
cular, surrounded by a 6-lobed cutinous or chitinous ring and with
2 lateral and 4 submedian papillae in relation with the clefts of this
ring; 4 sublateral papillae posterior to these. A pair of cervical pa-
pillae, near the origin of the esophagus. The buccal cavity or phar-
ynx has a short wide anterior portion and a long narrow posterior
portion. Club-shaped esophagus about 1.5 mm. long.

326

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 10 to 16 mm. long by 350/* wide. Tail (fig. 391 a and b)
curved ventrally. Cloacal aperture 320 to 400/* from tip of tail.
Four pairs of preanal and 2 pairs of postanal papillae. Smit says
there are 2 to 4 pairs of postanal papillae, but apparently figures 2
pairs of preanal papillae. Two unequal spicules ; one is 3 to 3.5 mm,
long by 8 to 10/* wide, the other is 200 to 220/* long by 30/* wide.
In Smit's specimens the long spicule was 480/*. long and the short one
150/* long, according to his description, but, taking the width of the
worm as a basis for size in his illustrations, these measurements ap-
pear incorrect, and the relative lengths of the spicules as figured do
not agree with his measurements.

Female 12 to 19 mm. long by 430/* wide. Anus 400 to 530/x from
tip of tail (fig. 390). Vulva 1 to 1.4 mm. from tip of tail. Vagina

/avyu.

Pig. 389. — Oxyspirura mansoni. a, Dorsal view and b, front view of head; c,

EGG; d, EMBRYO ESCAPING FROM THE EGG. (C.O.r., CIRCUMORAL CUTICULAR BING ;
es., ESOPHAGUS ; Lp., AMPHIDS OR SO-CALLED " LATERAL PAPILLAE " ; Jrt, MOUTH ;
ph., PHARYJVX ; S.l.p., SUBLATERAL PAPILLAE; S.m.p., SUBMEDIAN PAPILLAE.) AFTER

Ransom, 1904

1.5 to 2 mm. long. Eggs (fig. 389 c and d) oval, 50 to 65/* by 45/*
(Smit says 40/* long by half as wide; see O. parvovu??i, p. 328).

Larva, first stage, 225 to 250/* long by 12/* wide. Esophagus 50/*
long. Head end rounded. Tail end terminates in a thick, blunt
appendix.

Life history. — Unknown. The eggs produced by the female worm
wash down the tear ducts and are swallowed, passing out in the
droppings. The fact that the eggs will hatch in 2 to 3 days under
favorable conditions suggests that the eggs are not ingested im-
mediately by a secondary host, but that there may be a secondary
host which is infected by larvae, the larvae developing to adults in
the body of the primary host when these secondary hosts are swal-
lowed by it. Emmerez and, later, Ransom were unable to find
either eggs or larvae in the eye, and Ransom and, later, Wilcox and
McClelland fed larvae without obtaining infection in the primary
host. According to the latter authors larvae develop to a length of

NEMATODE PARASITES OF BIRDS

327

850//. to 1 mm. in 4 months, at which time sexual differences appear.
The smallest worm they have seen in the eye was 4 mm. long and
they have found mature worms in chickens 10 days old. They con-
clude that infection takes place by mature worms entering the eye
in soil as chickens dust themselves. Since worms of the genus Oxy-

yooy^.

OV.

vg-

int.

x.

ret.

390.

Figs. 390-391. — Oxyspiruba mansoni. 890, Postkbiob end ov female, {int., Intestine;
ov., OVARY; ret., recti- m ; iff., vagina; nil., vulva; x., cells SUBBOUNDZNG rectum.)
391, Posterior end of male, a, Lateral view. (dct. ej., Ejaculatory duct; int.,
intestine; sp., SPICULES; v.s., seminal vesicle.) b. Ventral view. After Ransom.
1904

spirura probably have life histories comparable to those known for
spirurids and filarids in general, this theory does not seem tenable
and it is probable that an intermediate liost is necessary for their
development. Such hosts would probably be insects, crustaceans,
or similar small animals commonly eaten by chickens and related
birds.

Distribution. — North America (United States (Florida)), South
America (Brazil), Asia (China (Annam)), Africa (Belgian Congo);

328 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

and many islands (Guam, Hawaii, Jamaica, Danish West Indies,
Island of Maurice, Dutch Indies, and Samoa). According to John-
ston, the worms from Australia are probably O. parvovum (p. 328).

OXYSPIRURA OPHTHALMICA (Linstow, 1903) Ransom, 1904

Synonyms. — C heilospirura ophthalmica Linstow, 1903a.

Hosts. — Primary : Turnix taigoor; secondary : Unknown.

Location. — Eye.

Morphology. — Oxyspirura (p. 321) : Cuticula with fine transverse
striations. Mouth (fig. 392a) surrounded by 2 lateral and 4 sub-
median papillae. Buccal cavity short, widened posteriorly. Lateral
fields 42/x wide ; that is, about 1/24 of circumference of body.

Male 14.4 mm. long by 310/* wide. Esophagus 1/16 of body length.
Tail 1/52 of body length finely pointed (fig. 392 b). One pair of
postanal papillae and 26 pairs of preanal papillae (Linstow says 1
pair of preanal and 26 pairs of postanal, but his figure shows that
he confused the two). Spicules unequal, the one 290/* long, the other
180/* long.

Female 21 mm. long by 530/* wide. Esophagus 1/21 of body
length. Tail conical, 1/53 of body length. Vulva near posterior end
of body, dividing the body length in the ratio of 102 : 5. Vagina
anteriorly directed, 400/* long, dividing into 2 uteri. Eggs 39/* by
26/*, with thick shells ; cleavage takes place in the uterus.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — Asia ( Siam ) .

OXYSPIRURA PARVOVUM Sweet, 1910

Hosts. — Primary : Gallus gallus; secondary : Unknown.

Location. — Under the nictitating membrane.

Morphology. — Oxyspirura (p. 321) : Very similar to 0. mansoni
(p. 325), but the head (fig. 393a.) is without a regular cutinous or
chitinous ring with clefts. The 6 cephalic papillae near the mouth
are difficult to see and may not be constant in number. Membranous
cephalic wings sometimes present in males. The lining of the phar-
ynx may project inward and forward at the union of the anterior
and posterior portions, and there may be a somewhat similar pro-
jection from the wall of the anterior portion. The esophagus is 1.13
to 1.3 mm. long.

Male 9.2 to 14.5 mm. long by 260 to SQQ/* wide. Tail (fig. 393Z>)
very sharply curved ventrally. Cloacal aperture 230 to 300/* from
tip of tail. Three pairs of preanal and 2 pairs of postanal papillae.
The long spicule is 3.4 to 4.1 mm. long, 11 to 13/* wide along most

NEMATODE PARASITES OF BIRDS

329

of its length and 24 to 30/* wide at the base; short spicule 180 to
240/* long by 27 to 42/* wide.

Female 13.5 to 20 mm. long by 270 to 390/* wide. Anus 390 to
440/* from tip of tail (fig. 394). Vulva 780/* to 1.07 mm. from tip
of tail. Vagina 2.64 mm. long (judging from the figures this in-
cludes more than the vagina) ; it is dilated at a point 660/* from
the vulva to form a thin-walled portion holding about 10 eggs
in transverse rows; not more than 1 egg at any point in terminal
portion. Eggs 33 to 45/* by 25 to 30/*, rather square-ended, con-
taining embryos when oviposited.

Life history. — Unknown. See remarks on O. mamoni, p. 326.

Figs. 392-393. — 392, Oxyspirura ophthalmica. a, Head; b, male tail. After
Linstow, 1903. 393, Oxyspirura paevovum. a. Head; b, male tail. After
Sweet, 1910

Distribution. — Australia. The worms reported by Smit from the
Dutch Indies have egg measurements in agreement with this species,
40/i by 20/*; he calls them O. mansoni, but they differ in some re-
spects from both O. marisoni and O. parvovum.

OXYSPIRURA SIAMENSIS (Linstow. 1903) Ransom, 1904

Synonym. — (Jheilospirura siamensis Linstow, 1903*7.

Hosts. — Primary: Gentropus siamensis {=Centropus sinensis)',
secondary : Unknown.

Location. — Probably the eye, according to Linstow.

Morphology. — Oxysjrirura (p. 321) : Cuticula with fine transverse
striations. Buccal cavity small.

Male 8.9 mm. long by 260/* wide. Esophagus 1/11, tail 1/34, of
body length, the tail (fig. 395) curved ventrally. About 28 pairs of
preanal papillae, becoming gradually smaller anteriorly. Spicules
unequal, 470/* and 250/* long.

Female 9.8 mm. long by 460/* wide. Esophagus 1/13, tail 1/37, of
body length. Vulva 620/* from tail end, situated a short distance
anterior to anus. Vagina directed anteriorly, 260it long, dividing
into 2 uteri.

330

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — Asia (Siam).

OXYSPIRURA SYGMOIDEA (Molin, 1860) Stossich, 1897

Synonyms. — Spiroptera anthuris Rudolphi, 1819, of Diesing, 1851,
part; Spiroptera sygmoidea Molin, 18605; Filaria anthuris Linstow,
1878, not Filaria anthuris (Rudolphi, 1819) Schneider, 1866.

Hosts. — Primary: Corvus corone, C. frugilegus,' secondary: Un-
known.

Location. — Orbital cavity and under nictitating membrane.

Morphology. — Oxyspirura (p. 321) : Body sigmoidal, attenuated
at both ends, truncated anteriorly, slender and acutely pointed pos-
teriorly. Cuticula with fine transverse striations. Mouth large, or-
bicular, without lips; 4 submedian cephalic papillae.

Pig. 394. — Oxyspirura parvovuji. Posterior end of female. After Sweet, 1910

Male 11 mm. long by 40G> wide. Tail coiled in 2 turns, without
alae. Four pairs of postanal papillae (fig. 396), the papillae of the
third pair from posterior end lateral of the others; preanal papillae
apparently absent. Spicules unequal, the right short, thick, some-
what curved, with very blunt point ; the left filiform, pointed, alate,
and 2^2 times as long as the right.

Female 11 to 15 mm. long bj 7 500/* wide. Anus some distance from
tail end. Position of vulva not determined.

Life history. — Unknown ; probably involves intermediate stages in
other hosts.

Distribution. — Europe (Austria (Vienna) ) and Asia (Russian
Turkestan).

OXYSPIRURA TANASIJTCHUKI Skrjabin, 1916

Hosts. — Primary: Birds of the family Icteridae; secondary:
Unknown.

Location. — Conjunctival sac.

Morphology. — Oxyspwura (p. 321) : Body white. Cuticula with
extremely delicate transverse striations. Anterior extremity (fig.
397a) rounded. Buccal cavity 170^ long, with thick walls. Two
lateral and 4 submedian papillae.

NEMATODE PARASITES OF BIRDS 331

Male 13 mm. long by 220yi wide. Esophagus 518/i long. Caudal
extremity (fig. 397c) tapering to a tine point. Right spicule (fig.
3976) 250/u. long, slightly curved, its dorsal surface convex, its ventral
concave; the ventral surface has a gutter which probably facilitates
the movement of the long spicule in gliding through it, thus serving
as a gubernaculum. Left spicule 600^ long, filiform, the posterior
extremity pointed. Arrangement of caudal papillae unusual ; 5 pre-
anal papillae of which there are 2 symmetrical pairs and a fifth
papilla isolated on the right side with no corresponding papilla on
the left ; 7 postanal papillae, 2 of which form a symmetrical pair
near the caudal extremity, the other 5 having a submedian and very
asymmetrical arrangement and being of unequal size. In addition
to the papillae, there are other asymmetrical formations, the char-
acter of which is undetermined, in the lateral fields posterior to the
cloacal aperture.

Female 14 mm. long by 320^ w T ide. Esophagus 765/a long. Anus
510/* from end of tail. Vulva in posterior part of body. Eggs 46/x
by 29/x.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — South America (Paraguay).

(70XYSPIRURA) HETEROCLITA (Molin, 1860) Ransom, 1904

Synonyms. — Spiroptera heteroclita Molin, 18606/ Spiroptera cracis
Molin, 1860 6.

Hosts. — Primary: Crax urumutum (=Nothocrax urumutum) ;
secondary : Unknown.

Location. — Under nictitating membrane.

Morphology. — Oxyspirwra (p. 321) : Body attenuated anteriorly,
apex truncate. Cuticula with fine transverse striations. Mouth or-
bicular, large, without lips.

Male 12 mm. long by 300/i wide. Head without membranes. Tail
awl-shaped, sharply pointed, curved into a hook, without caudal alae.
Two pairs of postanal papillae. Preanal papillae not described.
Spicules unequal, the one short and thick, the other filiform, half as
long as the body.

Female 55 mm. long by 400/a wide. Head with 4 short, semilunar
membranes, arranged in the form of a cross. Tail gradually attenu-
ated, awl-shaped, with obtuse tip. Anus remote from tail end.
Position of vulva not determined.

Life history. — Unknown; probably involves intermediate stages in
other hosts.

Distribution. — South America (Brazil).

332 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

(70XYSPIRURA) BREVIPENIS (Molin, 1860) Stossich, 1897

Synonyms. — Sjriroptera brevipenis Molin, 18605/ Spiroptera mi-
crodactyli margravii Molin, 18606.

Hosts. — Primary: Dicholophus margravi ( = Cariamu cristata) ;
secondary: Unknown.

Location. — Under nictitating membrane.

Morphology. — fOxyspirura (p. 321) : Body straight, slender, fili-
form. Anterior end attenuated,, apex truncate. Mouth large, orbicu-
lar, without lips or papillae. Buccal cavity absent.

Figs. 395-398. — 395, Oxyspiruka siamensis. Male tail. After Linstow, 1903.
396, Oxyspiruka sygmoidea. Male tail. After Skrjabin, 1916. 397, Oxyspi-
bitra tanasijtchuki. a, head ; b, right spicule ; c, male tail. afteb skrjabin,
1916. 398, Oxyspiruka brevipenis. Male tail. After Drasche, 1884

Male 16 to 22 mm. long by 400/* wide. Tail (fig. 398) coiled in 2
turns, without alae. Six pairs of preanal and 6 pairs of postanal
papillae, the latter inconstant in number and asymmetrical in ar-
rangement. Spicules almost equal, very short, curved, sabre-shaped.

Female 11 to 27 mm. long by 100 to 500/x wide. Tail straight,
slender, conical, with thickened tip. Anus remote from tail end.
Vulva near anus, prominent, with swollen posterior lip.

Life history. — Unknown; probably involves intermediate stages
in other hosts.

Distribution. — South America (Brazil).

Ransom (1904) has noted that the absence of a buccal cavity and
the presence of short spicules make the position of this species in this
genus questionable.

NEMATODE PARASITES OF BIRDS 333

Family TETRAMERIDAE Travassos, 1914

Family diagnosis. — Spiruroidea (p. 162) : Characterized by the
sexual dimorphism of the species. Male with filiform body, white,
unarmed or armed with spines. Cuticle more or less striated trans-
versely. Caudal extremity pointed: usually two unequal spicules.
Caudal papillae or spines may be present. Female red ; body greatly
enlarged in comparison with that of male, its shape either globular,
with the two pointed extremities projecting {Tetrameres), or the
long axis of the body coiled in a more or less complex manner (Mi-
crotetrameres). Digestive tract consists of mouth aperture with 3
small lips, followed by a chitinous mouth capsule, a muscular
pharynx, a muscular esophagus and a thin-walled, wide, sac-like in-
testine which is usually filled with detritus and ends in a narrow
tube opening at the anus. Genital system highly developed, the body
cavity largely filled with the numerous coils of the uteri, ovaries,
and oviducts. Uteri contain an enormous number of eggs in various
stages of development, the embryo being well formed when the egg
is deposited. Vulva in posterior part of body, near anus. A saccu-
lar diverticulum of the ovejector may be present, called by Seurat a
" copulatory bursa " but preferably designated by some term not al-
ready used for a male structure, such as " copulatory receptaculum."

Parasitic in proventriculus of birds, the females in the glands of
Lieberkuehn, the males usually free in the lumen.

Type-genus. — Tetrameres Creplin, 1846.

KEY TO GENERA OF TETRAMEUIDAE

Body of female globular or spindle-shape Tetrameres, p. 334.

Body of female with its long axis spirally coiled Microtetrameres, p. 351.

I have raised the subgenus Microtetrameres Travassos, 19155 to
generic rank and have rewritten the diagnoses for the two genera
Tetrameres and Microtetrameres on the basis of the difference in body
form of the female. The diagnoses of the subgenera of Travassos,
which included with the description of the body form of the female
the proportionate length of the spicules and the presence or absence
of spines in the lateral fields of the male, did not furnish a suitable
basis for the division of Tetrameres {sensu lato), at least in the
present state of knowledge. T. cruzi, the type-species of the sub-
genus Microtetrameres was not consistent with the subgeneric diag-
nosis as given by Travassos, as the long spicule of the male is not
% the body length, but just slightly over half. In four species
the male is unknown; several other species possess certain characters
which are included in the description of one subgenus but at the same
time possess certain other characters which belong to the other sub-

334 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

genus, the result being that these species had to be grouped to-
gether as of uncertain subgeneric classification. It has therefore
been deemed advisable to reduce the descriptions, in raising the sub-
genera of Travassos to generic rank, to the basis of the difference in
female body form, and thus be able to assign all the species of Tetra-
meres (sensu lato) to one of the two genera, Tetrameres or Microte-

Genus TETRAMERES Creplin, 1846

Synonyms. — Tropisurus Diesing, 1835; Tropidurus Wiegmann,
1835; Tropidocerca Diesing, 1851; Astomwn Schlotthauber, 1859;
Acanthophorus Linstow, 1876.

Generic diagnosis. — Tetrameridae (p. 333) : Female body globular
or spindle-shape, with 4 longitudinal furrows corresponding to the
median and lateral lines.

Type-species.— T etrameres paradoxa (Diesing, 1835) Travassos,
1914.

KEIY TO SPECIES OF TETRAMERES

1. From Fulica atra; anus 220/* from posterior extremity; eggs 39/* long by

29fi wide Tetrameres globosa, p. 346.

From other hosts than Fulica atra, or if from that host, anus 71/t from
posterior extremity and eggs 48 to 56/* long by 26 to 30/t wide (Tetra-
meres fissispina) 2.

2. Mouth of female surrounded by 3 projections, each of which bears a thorn

on its outer and on its inner surface ; male unknown, female poorly

described; in Corvus comix Tetrameres unispina, p. 351.

No such structures described on female mouth ; male known in all but two
species and female more fully described than above ; in other hosts
than Corvus comix 3.

3. Female 7 mm. long by 7 to 8 mm. wide ; male 12 to 18 mm. long, its caudal

extremity said to be alate Tetrameres certa, p. 338.

Female smaller than above or if as large (T. gynaecophila), the male is
smaller than that of above and has no caudal alae 4.

4. Female 5 mm. to 13 mm. long by 4 to 7 mm. or more wide, or larger ; male

6.5 to 15 mm. long 5.

Female not over 4.5 mm. long (except in T. confusa and T. fissispina
which may reach 5 and 6 mm. long respectively) ; male, when known,
not over 6 mm. long 7.

5. Female 6.75 to 8 mm. long by 4.5 to 7 mm. wide; male 11 to 15 mm. long;

spicules 3 mm. and 468/t long respectively Tetrameres paradoxa, p. 335.

Female usually smaller or larger than above ; male not over 10 mm. long ;
spicules much smaller than above, or said to be absent 6.

6. Female not over 6 mm. long by 5 mm. wide ; male with 4 longitudinal series

of spines ; spicules 740/* and 16/* long respectively.

Tetrameres gigas, p. 345.

Female 8 mm. long by 7 mm. wide, or larger ; male without spines ; spicules

said to be absent Tetrameres gynaecophila, p. 347.

7. Anus of female 332/*, cloacal aperture of male 232 to 290/* from posterior

end Tetrameres americana, p. 337.

Anus of female and cloacal aperture of male both nearer to the posterior
end than above 8.

NEMATODE PARASITES OF BIRDS 335

8. Female with pharynx 400m, esophagus 2 mm. Long; anus 250/i from posterior

end; cloacal aperture of male 70m from posterior end.

Tetrameres confusa, p. 341.

Both pharynx and esophagus of female shorter than above, if described ;

anus, if described, not more than 175m from posterior end ; eloacal aperture

of male, if descrihed, 100m or farther from posterior end 9.

9. Only female known 10.

Both male and female known 11.

10. Female 2.2 mm. long by 2.5 mm. wide; pharynx 350m, esophagus 1.4 mm.

long; eggs 28 to 30m long by 15 to 18m wide Tetrameres coccinea, p. 339.

Female 3 to 4 mm. long hy 1.5 to 2 mm. wide; pharynx 190^, esophagus 990m
long; eggs 42 to 49/* long by 21ti wide Tetrameres cochleariae, p. 340.

11. Male 2.2 to 2.G mm. long; female not over 1.7 mm. long L2.

Male larger than above or (T. dubia) smaller than above; female 2 mm.

or more in length 13.

12. Eggs 54m by 28m wide, with long filaments at each pole ; one spicule 480m in

length, the other rudimentary Tetrameres nouveli, p. 34S.

Eggs 75 to 78m long by 21m wide, with no filaments described ; spicules 200m
and 22m long respectively Tetrameres tetrica, p. 350.

13. Female not over 2.5 mm. long; male 1.6 mm. long, its esophagus being 300m

long; long spicule 720m in length Tetrameres dubia, p. 342.

Female 2.5 to 6 mm. long; male 3 to 6 mm. long, its esophagus 780m or more
in length ; long spicule not over 490m iu length 14.

14. Ovejector with diverticulum or copulatory receptaculum 400m long ; eggs

not over 56m long; short spicule at least 82m long.

Tetrameres fissispina, p. 343.
Ovejector simple, without diverticulum; eggs 59 to 03m long; short spicule

56m long Tetrameres micropenis, p. 348.

This key does not include Tetrameres zakharoivi Petrow, 1926. See Addenda,
p. 385.

TETRAMERES PARADOXA (Diesing, 1835) Travassos. 1914d

/Synonyms. — Tropisurus paradoxus Diesing, 1835; Tropidocerca
paradoxa (Diesing, 1835) Diesing, 1851.

Diesing's description of 1851 is evidently a composite description
as in 1801 he makes Tropidocerca paradoxa Diesing, 1851 in part a
synonym of his new species Tropvlocerca inflata, and also transfers
the synonomy given originally under Tropisurus paradoxus (that is,
Spiroptera inflata and Tetrameres haemochrous) to Tropidocerca
i7ifl,ata. The original material up to the present time is poorly de-
scribed; it would be advisable therefore that the species be established
on Brazilien material from the type host, Catharte.s urubu. The de-
scriptions of Diesing (1835) and of Drasche (1884) of the same
material are summarized below.

Hosts. — Primary: Cathartes urubu and Strix torquata. Reported
from a large number of other hosts but probably erroneously so, as it
is known that Microtetrameres inflata {Eustrongylides mergorum)
was the species involved in many of the reports and other species
may also have been mistaken for this one, owing to (lie original

3612—27—23

336

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

insufficient description and the later composite one. Diesing (1861)
in his latest discussion of this species limits the hosts to the two given
above, as does also Linstow (1879). Secondary: Unknown; prob-
ably similar in a general way to that of T. fissis-pina (p. 343).

Location. — Glandular stomach or proventriculus.

Morphology. — Tetrameres (p. 334).

Male 5 to 6 lines long by 1/2 line wide (this would apparently be
about 12 to 15 mm. long by 732 to 999/x, wide; see discussion under
size of female) ; according to Linstow (1879) 11 to 13.5 mm. long
by 750/* wide. Body subcylindrical, threadlike, curved to crescent
shape, white, pointed at both ends. Drasche's figure of the posterior
extremity (fig. 400 b) shows spines along the lateral lines. Cuticle
transversely striated. Combined length of pharynx and esophagus
3/4 line (evidently about 1.9 to 2.2 mm. long). The testis extends

C.

Fig. 399. — Tetrameres paradoxa. a, Female, natural size; i, lateral
view, and c, view from above, of same, enlarged ; d, egg. enlarged.
After Diesing, 1835

anteriorly to the posterior extremity of the esophagus. The intestine,
as well as the vas deferens, shows in its posterior portion a pyriform
swelling. Cloacal aperture 1/16 line (apparently about 156 to 187jn)
from posterior extremity. A sudden decrease in dorso-ventral diam-
eter produces a depression immediately posterior to the cloacal aper-
ture. Two spicules of very different lengths; the right one short
(480ju) , the left one 3 mm. or more. In Drasche's figure there is at
least one pair of ventral postanal spines or very small papillae.

Female the size of a pea or larger; 3 lines long by 2 lines wide.
(Diesing gives these measurements; he also gives a figure of the
female (fig. 399 a), " one of the small specimens," natural size, which
measures 8 mm. long by 7 mm. wide ; the measurements in millimeters
are therefore at least 2% to 3 times those given by him in lines.)
Linstow (1879) gives size of female as 6.75 mm. long by 4.5 mm.
wide. Body blood-red, subglobular, with 4 deep, equidistant, longi-

NEMATODE PARASITES OF BIRDS

337

tudinal furrows (fig. 399 b and c). Cuticle transversely striated.
Extremities of body acutely conical. Intestine filled with grayish
material.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

TETRAMERES AMERICANA, new species

Hosts. — Primary: Gallus gallus; secondary: Unknown; probably
similar in a general way to that of T. ftssispina (p. 343).

Fig. 400. — Tetrameues pakaduxa. «, Head (whether of male or female not
stated) ; b, -male tail. after drasche, 1884

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334) : Mouth with 3 small lips; buc-
cal capsule with chitinous walls. Slender muscular pharynx present,
followed by esophagus. In other respects male and female very
dissimilar.

Male 5 to 5.5 mm. long by 11G to 133/* wide. Two double rows of
posteriorly directed spines extend throughout whole body length, in
the submedian lines. Buccal capsule (fig. 402 a) 27/*. deep by 4.5/*

Fig. 401. — Tetrameres Americana. Posterior end of male. Original

wide: pharynx 365/w long; esophagus 996/* long. Cervical papillae,
one slightly higher than the other, 183 and 199/* respectively from
anterior end. Nerve ring just posterior to the latter papilla. Tail
(fig. 401) long and slender, cloaca! aperture 232 to 290/* from
posterior end. Two unequal spicules, KM)/* and 290 to 312/* long
respectively, not heavily chitinized.

Fenwle 3.5 to 4.5 mm. long by 3 mm. Avide; body globulin- (fig.
402/), blood-red in color (when alive), with 4 longitudinal furrows
corresponding to the lateral and median lines. The anterior ex-
tremity (fig. 402c) protrudes from the globular body for a length

338

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

of 913/a ; the protruding posterior part of body is 860/* long. Buccal
capsule 35 to 38/x long by 10/* wide ; pharynx 300 to 315/* long by 27/*
wide; esophagus 1.4 mm. long by 50/* minimum and 125/* maximum
width. Nerve ring 183/* from cephalic extremity. Intestine saccular,
filled with black detritus. Anus 332/*, vulva 631 to 664/* from caudal
extremity. Vestibule of ovejector with diverticulum or copulatory
receptaculum (fig. 402 e) 274/* long. Uteri and ovaries very long,
their numerous coils filling the body cavity. Eggs 42 to 50/* by 25/*.
The American form of Tetrameres from the chicken, as described
above, differs from the two forms reported from other countries from

/mm

oimm

Fig. 402. — Tetrameres Americana, a, Head of male ; b, junction of testis
and vas deferens ; c, head end of female j d, junction of uterus and
ovary; c, copulatoey receptaculum of female; f, cross-section of female.
Original

that host, T. ftssispina and T . confusa, in numerous respects, as
length of buccal capsule, pharynx, and esophagus, distance of anus
and vulva from posterior extremity in the female and of cloacal
aperture in the male, position of cervical papillae, and length of
the diverticulum or copulatory receptaculum of the female.

Life history. — Probabry similar in a general way to that of T.
flssispina (p. 343).

Distribution. — (North America (United States)).

Type material. — No. 26382, U.S.N.M. (Bureau of Animal Industry
Helminthological Collection).

TETRAMERES CERTA (Leidy, 188G) Travassos, 1914

Synonyms. — FUaria dubia Leidy, 1856, not Filaria dubia Creplin,
1846; Tropidocerca certa Leidy, 1886.

Although there is a question as to whether Creplin meant the
combination, Filaria dubia, as a specific name, it is in the interests of

NEMATODE PARASITES OF BIRDS 339

stability of nomenclature to assume that he did. Otherwise Tetra-
meres certa would have to fall if the older specific name dubia of
Leidy is available and in consequence Tetrameres dubia Travassos,
1917, would fall as a homonym. To avoid this needless rearrange-
ment I am interpreting Filaria dubia Creplin as a good combina-
tion and Filaria dubia Leidy as therefore a homonym.

Hosts. — Primary: Albatross (Diomedia ewulans); secondary: Un-
known, probably similar in a general way to that of T. fissispina
(p. 343) ;

Location. — Glandular stomach or proventriculus.

Male 12 to 18 mm. long by 375 to 500/*. wide. Body filiform, at-
tenuated anteriorly. Mouth trilabiate. Caudal extremity strongly
rolled inwardly, sigmoid at the end, mucronate and alate; alae half
oval, narrowing to the end of the mucro. (The character of the tail
of the male is unusual for a Tetrameres and raises a doubt as to the
male specimens described belonging with the female in this genus.)

Female 7 mm. long by 7 to 8 mm. wide. Body subglobular, divided
into zones; tail conical, projecting abruptly from body.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South Atlantic.

TETRAMERES COCCINEA (Seurat, 1914) Travassos, 1914d

Synonym. — Tropidocerca coccinea Seurat, 1914m.

Hosts. — Primary : Phoenicopterus roseus, LJubulcus lucidus, and
Platalea leucorodia; secondary : Unknown, probably similar in a gen-
eral way to that of T . fissispina (p. 343).

Location. — Glandular stomach or proventriculus.

Morphology. — Tetrameres (p. 334).

Mcde unknown.

Female 2.2 mm. long by 2.5 mm. wide. Body globular (tig. 403 a),
cochineal colored, strongly striated transversely and with 4 longi-
tudinal furrows corresponding to the dorsal, ventral, and lateral
lines. Head and tail ends conical, tapering prolongations. Buccal
cavity (fig. 403&) circular. 20/x long: pharynx 350/*, surrounded in
its middle by the nerve ring; cervical papillae in front of nerve ring,
at a distance of 120 and 180/t respectively from the anterior end;
excretory pore ventral, 18/* behind posterior border of nerve ring;
esophagus, 1.4 mm. long, penetrates into the globular mass of the body.
Intestine greatly distended, tilled with brown-black detritus. Anus
at a short distance from tip of tail. Vulva 480/t anterior to anus.
Ovejector (fig. 403c) remarkable for the shortness of vestibule and
sphincter; they form a turnip-shaped organ 450/x long. Sphincter
very straight. Trompe, Y-shaped, almost 1 mm. total length, the

340

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

unpaired branch being 350/* long. Uteri very long, their course
greatly twisted and turned within the distended body; the initial
region of uterus, where it connects with the oviduct, enlarged to form
an enormous seminal receptacle (fig. 403c?), measuring 660/* long by
215/* wide. Diameter of oviduct only 35/*; oviducts and ovaries 8
mm. long. Eggs 28 to 30/* long by 15 to 18/* wide, with thick, smooth
shells and containing well-developed embryo when deposited.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — Africa (Algeria) and Asia (Russian Turkestan
(Lac Kulkainar)).

Figs. 403—404. — 403, Teteamekes coccinea. Female, a, Total; b, anterior ex-
tremity ; e, ovejector ; d, seminal receptacle at juncture of uterus and
oviduct. After Seurat, 1914. 404, Tctrameres cochleariae. Ovejector.
Afteb Travassos, 1919

TETRAMERES COCHLEARIAE Travassos. 1917a

/Synonym. — T etrameres micropenis Travassos, 19156, part.

Hosts. — Cancroma cochlearia; secondary : Unknown, probably sim-
ilar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — T etrameres (p. 334).

Male unknown.

Female 3 to 4 mm. long by 1.5 to 2 mm. wide. Body red color,
with characteristic shape. Buccal capsule barrel-shaped, 24 to 28/*
in height, 14 to 16/* in maximum width ; pharynx 190/* long ; esopha-
gus slightly claviform, 990/* long; nerve ring 140/* from anterior end
of body. Ovejector (fig. 404) very long and muscular and with a
vestibule or proximal portion inserted in the terminal portion at an
angle, measuring 1.9 mm. long, 120/* maximum width and 50/* mini-
mum width at the distal portion. (The length given is evidently the
total length of the ovejector, although Travassos does not make this
clear.) Eggs ellipsoidal, slightly flattened on the sides, 42 to 49/*
long by 21/* wide.

NEMATODE PARASITES OF BIRDS

341

Life history. — Probably similar in a general way to that of T.
ftssispina (p. 343).

Distribution. — South America (Brazil).

TETRAMERES CONFUSA Travassos. 1919b

Synonyms. — Tetrameres fissispina Diesing, 1861 of Travassos,
1911a; Tetrameres travassosi Skrjabin, 1920.

Hosts. — Primary : Columbia livia domestica y Gallus gallus and
Meleagris gallopavo; secondary : Unknown, probably similar in a
general way to that of T. fissispina (p. 343).

Location. — Glandular stomach or proventriculus.

Morphology. — Tetrameres (p. 334).

Male 4 to 5 mm. long by 160/u. wide. Body filiform, with 4 rows
of spines corresponding to the median and lateral lines; spines di-

FiG. 405.

-TETRAMERES CONFUSA. C, FEMALE; b, MALE TAIL. AFTER TRAVAS-
SOS, 1010

rected posteriorly and about 100/u apart in middle portion of body
and 2G> in posterior portion. Tail ends in a hook about 4/u, long.
Buccal capsule cylindrical, 24/x long by 80/u wide; pharynx 250/u. long
by 12/x wide; esophagus 740/z long by 55/* wide. Cloacal aperture
70^ from end of tail (fig. 405 b). Short spicule 68/x long and long
spicule 291/n long. Behind the cloaca there are 2 rows of 5 spines
each on the ventral surface and lateral of these are 2 rows of 3
spines each, extending posterior of the ventral spines.

Female 3 to 5 mm. long by 2 to 3 mm. wide. Body subglobular
(fig. 405a), with deep transverse striations and with 4 furrows
corresponding to the median and lateral lines. The head and tail
ends project as slender conical prolongations, the anterior 1 mm.
long, the posterior 900/u, long. Buccal capsule 20/u. long by 14/x wide;
pharynx 400/* long by 30/* wide; esophagus sinuous, 2 mm. long by
180/x wide; intestine dilated to form a large sac, filled with black
detritus. Anus 250/x from posterior extremity. Vulva in posterior

342 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

portion of body, near anus. At the beginning of the vagina, a
saccular dilatation (copulatory receptaculum) which is filled with
eggs in the adult worm. Eggs 33/a long by 24^ wide.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

TETRAMERES DUBIA Travassos, 1917a

Hosts. — Primary: Gallinago paraguaiae; secondary: Unknown;
probably similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

Male 1.6 mm. long by 85/a wide. Body (fig. 406) attenuated at
the extremities, the greatest width being at the level of the esophagus.
Cuticle transversely striated and with spines in the lateral fields but

Fig. 406. — Tetrameres dobia. Male. After Travassos, 1919

only at the anterior extremity; the spines are few and very small.
Buccal capsule very small (6/* long by 4/* wide) ; pharynx 240/*,
long; esophagus 300/* long. Anus lOOju, from posterior extremity.
Tail sharply pointed with 4 pair of very small spines on the ventral
surface. Spicules of very unequal dimension, measuring respectively
720/* long by 6^ wide and 64/t long by 6/* wide.

Female 2 to 2.5 mm. long by 1 to 1.55 mm. wide. Body globular,
red, with marked transverse striations and with 4 deep longitudinal
furrows corresponding to the median and longitudinal lines. Buc-
cal capsule barrel-shaped, 16/* long by 7/a wide. Anus 75/x from pos-
terior extremity. Ovejector 420/x long, of same type as that of T.
micropenis, the dilated part full of eggs. Eggs 49^ long by 35fi
wide. Mature eggs (that is, containing developed embryos) show
tufts of filaments at the poles; in immature eggs these are absent

NEMATODE PARASITES OF BIRDS 343

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

TETRAMERES FISSISPINA (Diesing, 1861) Travassos, 1914

Synonyms. — Tropidoce?xa fissispina Diesing, 1861 ; Tropisurus fis-
sispinus (Diesing, 1861) Neumann, 1888; Acanthophorus tenuis
Linstow, 1876; Acanthophorus horridus Linstow, 1876; Tropidocerca
tenuis Linstow, 1899 ; FUa7*ia pulicis Linstow, 1894 ; Spiroptera puli-
cis (Linstow, 1894) Linstow, 1909. The nematode described by
Travassos in 1914 as T . fissispina is T. confusa.

Hosts. — Primary: Anas boschas, Anas boschas domestica, Anas
boschas fera, Columba livia domestica, Cygnus melanocoryphus,
Fulica atra, Meleagris gallopavo, Mergus merganser, Nyroca ferina,
Podiceps fluviatilis; secondary : Daphnia, pulex and Gammarus
pulex, the former being reported by Rust and the latter by Linstow
in his description of /Spiroptera (Filaria) pulicis, which Seurat con-
siders to be the fourth stage larva of T. fissispina.

Location. — Proventriculus, the males in the lumen, the females in
the crypts of Lieberkuehn, in birds; in the body cavity of secondary
hosts as larvae.

Morphology. — Tetrameres (p. 334) : Mouth with 3 small lips (figs.
407 and 408), chitinous mouth capsule, and muscular pharynx and
esophagus. In other respects male and female very dissimilar.

Male 3 to 6 mm. long by 140 to 150/* wide; white and slender.
Transverse striations more or less marked. The two median and two
lateral lines are each provided with a longitudinal series of spines;
behind the cloacal aperture there are 5 ventral spines on each and
3 lateral spines on each side (fig. 408). Considerable difference of
opinion has been expressed as to the nature of the spines on the body
of the male, and as to the basis for the specific name, referring to the
cleft spines. Diesing in his original description, after describing 4
longitudinal series of spines on the body, and 2 subbasilar conical
spines on the neck, writes that the head is provided with spines cleft
at their ends. Lieberkuehn (1855) described and pictured double
spines in the submedian lines; Linstow (1876) described and figured
in Acanthophorus tenuis and A. horridus such double spines and
later (1899) made these two species synonyms of T. fissispina; how-
ever, at that time he states " Lieberkuehn described on the head end
double spines; I also have seen this formation. However, they are
always located in the concave bending lines (Beugungslinien) and I
consider them duplicatures of the cuticle." Railliet (1893) and Neu-
mann (1909) describe the spines at the cephalic end, up to the origin
of the intestine, as characteristically double or cleft. Seurat, how-
3612— 27 24

344

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

ever, has more recently (1918a) described alae along each lateral
line, extending from the base of the buccal lips to the posterior end
of the body (175/* anterior to the cloacal aperture) and at the origin
of the alae a long bifid spine (52/* long). (Fig. 4:07a and b.) He
states that it is to the existence of these bifid cephalic spines that
the name fissispina is due and not to the rows of esophageal spines;
they are simple.

According to Travassos, the cervical papillae are 150//, from the
anterior end, buccal capsule 8/* deep by 3/* wide, esophagus 780/*
long by 52/* wide, cloacal aperture 130/* (250/*, according to Wharton)
from posterior end. The tail ends in a spine 4/* long. The spicule

Figs. 407-409. — Tetrameres fissispina. 407, a and ft, Head end

OF MALE, LATERAL VIEW (d), DORSAL VIEW (ft), SHOWING THE TWO
BIFID CEPHALIC SPINES AND THE SIMPLE BODY SPINES ; C, HEAD END
OF FEMALE, DORSAL VIEW. AFTER SEURAT, 1920. 40S, HEAD AND
TAIL OF MALE. AFTER TRAVASSOS, 1919. 409, fl, OVEJECTOR ; ft,
COPULATORY RECEPTACDLUM ; C, FEMALE GENITAL SYSTEM. AFTER

Seurat, 1914. d, Tail of larva, ventral view. After Seurat,
1919

lengths as given in various descriptions are 150 and 320/* (Seurat),
115 and 320/* (Wharton), 82 and 490/* (Travassos), 88 and 280/*
(Railliet).

A comparative study of European material with that from other
parts of the world is needed in order to determine whether or not
it is all the same species.

Female 2.5 to 3 (according to Wharton, 3 to 6) mm. long by 1
to 2 (according to Wharton, 2 to 3.5) mm. wide, oval except for
the head and tail ends which project as conical points, and blood-
red. Buccal capsule (fig. 407c) 21/* long by 10/* wide; pharynx
230/* long; esophagus 1 mm. long by 87/* in maximum diameter.
There are 4 longitudinal furrows which correspond to the 2 median
and 2 lateral lines. The intestine forms a large, piriform sac, ordi-
narily filled with black detritus and visible through the body wall.
The body cavity is largely filled with the numerous coils of the

NEMATODE PARASITES OF BIRDS 345

ovaries, oviducts, and uteri. Anus 71/x from the end of the tail.
The vulva is 310ju from the end of the tail. It connects with a very
short vestibule (fig. 409 a, b, and c) which presents on its ventral
surface a diverticulum closed at its free end, 400/* long, its walls
having the same structure as that of the vestibule. This is not a
receptaculum seminale but a " copulatory bursa " or copulatory re-
ceptaculum, analogous to that of various insects, having a role in
copulation, such as the role played by the vestibule of other nematode
parasites. Sphincter very short, continued by a trompe, which soon
divides into two branches which join the uteri. The uteri measure
21 and 19 mm., respectively, thus are 6 times the total length of the
body; the distal extremity, connected with the oviduct, is enlarged
into an ampoule and differentiated as a receptacle seminale. Oviduct
and ovary represented by a slender tube 6 mm. long. Eggs 48 to 56/x
long by 26 to 30/x wide and containing well developed embryos when
deposited.

Larva, fourth stage, 1.65 mm. long, straight and slender (female) ;
tail (fig. 409d) long with rounded tip bearing 8 ( ?5) long spines
(Seurat says 8 and figures 5) and with 2 latero-ventral spines 70 p
posterior to the anus.

Life history. — The eggs pass out in the feces and are swallowed
by so-called " water fleas " (Daphnia pulex) or " sand fleas " {Gam-
marun pulex) under favorable conditions. In these hosts the larvae
develop to the infective stage in the body cavity, and when such hosts
are swallowed by suitable birds in feeding or drinking, the worms
become mature, mate, and the females then enter the canals of the
glands of Lieberkuehn, the males remaining in the lumen of the
proventriculus. The female lies with the tail in the duct and the
head in the fundus, to facilitate the passage of eggs and feeding.
The body becomes distended with eggs. Travassos has compared this
habit of life with that of the chigoe flea, and the tetramere may well
be regarded as a sort of entoparasitic, verminous chigoe.

Distribution. — North America (United States), Oceania (Guam),
Asia (Russian Turkestan and Philippines), Europe, and Africa
(Algeria). The North American reports probably deal largely, if
not wholly, with T. americana (see p. 337).

TETRAMERES GIGAS Travassos. 1919b

Synonym. — Tetrameres inflata of Zuern, 1882, not Diesing, 1861,
of Travassos.

Hosts. — Primary: Anas boschas domestica; secondary: Unknown,
probably similar in a general way to that of T. fissispina (p. 343).

Location. — Glandular stomach or proventriculus.

Morphology. — Tetrameres (p. 334).

346

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 7.5 mm. long by 180/* wide. Filiform worms. Cuticle finely
striated transversely and with 4 longitudinal series of spines in the
lateral fields. Mouth (fig. 410) with 2 lips, buccal capsule cylindrical,
21/* long by 14/* wide. Pharynx 370/* long. Esophagus 950/* long.
Cloacal aperture 120//, from end of tail. Tail curved dorsally. Cau-
dal papillae absent or apparently very small. Small spicule 16/*
long ; large spicule 740/* long by 14/* wide and acutely pointed.

Female 5 to 6 mm. long by 4 to 5 mm. wide. Transverse striations
of cuticle more prominent in anterior portion of body. Lateral
fields and median lines form four furrows. Ovejector with a sac-
cular diverticulum as in other species. Eggs 50/* long by 21/x wide,
containing embryos when deposited.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

Fig. 410. — Tetrameees gigas. Anterior and posterior ends of male. After

Tkavassos, 1919

TETRAMERES GLOBOSA (Linstow, 1879) Travassos, 1914d

Synonym. — Tropidocerca globosa Linstow, 1879«.

Hosts. — Primary: Fulica atra; secondary: Unknown; probably
similar in a general way to that of T . fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

Male unknown.

Female globular, its two extremities slender projections. Cuticle
transversely striated. Buccal capsule circular with chitinous walls.
Pharynx 280/*; esophagus 780/* long. Anus 220/* from posterior ex-
tremity. Eggs thick-shelled, 39/* long by 29/* wide.

Linstow states that this species differs from Tetrameres -fissispina
in absence of cervical papillae and of a double spine at the tail end.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution, — Locality not given. This species has evidently not
been found since Linstow's original report of it.

NEMATODE PARASITES OF BIRDS

347

TETRAMERES GYNAECOPHILA (Molin, 1858, emend. Diesing, 1861) Travassos, 1914d

Synonyms. — Tropidocerca gynecophila Molin, 1858; Tropidocerca
gynaecophila Diesing, 1861.

Hosts. — Primary : Ardea nycticorax and Nycticorax nycticorax ;
secondary: Unknown; probably similar in a general way to that of
T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

Male 6.4 mm. (Seurat) or 10 mm. long (Molin) by 600/* wide.
Body without spines; curved in a circle. Cervical papillae situated
far in front of the nerve ring, 122 and 133/* respectively from the
anterior extremity of the body. Excretory pore posterior to nerve
ring, in median ventral line, 280/* from anterior extremity. Buccal
capsule 21/* long, conical. Pharynx 265/* long. Nerve ring situated
posterior to middle of pharynx. Esophagus 1 mm. long. Intestine
maroon-colored, forming a cul-de-sac in its posterior portion. Rec-

Fig. 411. — Tetrameres gynaecophila. a, Head op female; b, tail op male.

After Seurat., 1915

turn 50/* long. Cloaca 120/* from posterior extremity; tail conical,
terminating in a small round button. Caudal papillae arranged as
follows: Two pair postanal on short peduncles; 1 pair at level of
anus ; 7 sessile preanal at the right, 6 at the left, disposed as shown
in figure 4116. No spicule present.

Female 8 mm. long by 7 mm. wide (Seurat) or 13 mm. long and
wide (Linstow). Body scarlet-colored, globular, with 4 longitudinal
furrows corresponding to the lateral and median lines. Anterior
and caudal extremities pointed. Cervical papillae situated far in
front of nerve ring, 85/* from the anterior extremity. Buccal (fig.
411a) capsule 28/* long, conical, narrow at entrance, widened behind.
Pharynx 312/*, encircled by nerve ring directly in front of its mid-
dle. Esophagus 2.34 mm. long. Intestine black-colored; rectum
short (180/*). Anus 140/* from posterior extremity. Tail sharply
narrowed behind anus, digitiform, ending in a blunt point. Vulva
in neighborhood of anus. Ovejeetor Y-shape<l : vestibule and sphinc
ter joined to form cylindrical tube 1.26 mm. long, with thick Avails:

348 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

unpaired trompe 600/* long. Eggs 47/* long by 21/* wide, thick-
shelled, embryonated; they are cylindrical and rounded at the ends.

Life history.— Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — Africa (Algeria) and Europe (Italy (Padua) ).

TETRAMERES MICROPENIS Travassos, 1915b

Hosts. — Primary: Nicticorox violacens and Cancroma cochlearia;
secondary: Unknown; probably similar in a general way to that
of T. fissispina (p. 343).

Location. — Proventriculus. Males and young females free in the
cavity of the proventriculus; adult females in the glands of
Lieberkuehn.

Morphology. — Tetrameres (p. 334).

Male 4 to 5 mm. long by 120/* wide. Cuticle transversely striated;
numerous spines along lateral lines and, in addition, 2 pairs of
ventral spines below the anus. Buccal capsule funnel-shaped, 28/*
long; pharynx 355/* long; esophagus more or less cylindrical, 1.3

Fig. 412. — Tetrameres micropenis, a, Tail op male ; b, ovejector, showing

STRIATED APPEARANCE OP MUSCULATURE. AFTER TRAVASSOS, 1919

mm. long by 49/* wide. Cervical papillae about 163/* from anterior
extremity of body; nerve ring 191 to 198/* from anterior extremity.
Posterior extremity slender (fig. 412 a). Anus 184/* from the end of
body. Spicules slender, 355/* and 56/* long, respectively.

Female 3 to 4 mm. long by 1.5 to 2 mm. wide. Body globular,
red, strongly striated transversely and with 4 deep longitudinal
furrows corresponding to the lateral and median lines. Buccal
capsule 21/* long by 14/* wide, oval in form; pharynx 250/* long,
esophagus 1.5 mm. long, cylindrical. Vulva slightly anterior to
anus. Ovejector (fig. 4125) simple, about 710/* long, claviform and
longitudinally striated. Eggs 59 to 63/* long by 39/* wide.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

TETRAMERES NOUVELI (Seurat, 1914) Travassos, 1914d

Synonyms. — Tropidocerca nouveli Seurat, 1914£.

Hosts. — Primary : Himantopus himantopus; secondary : Un-
known; probably similar in a general way to that of T. fissispina
(p. 343) ;

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

NEMATODE PARASITES OP BIRDS

349

Male 2.16 mm. long by 75/t wide. Body slender (fig. 413<? and
413/) and with 2 longitudinal rows of spines, directed backward,
on each side of the lateral lines ; thus there are 2 latero-dorsal and 2
latero-ventral rows. Lateral lines well marked, presenting a cuti-
cular crest, slightly salient, extending the entire length of the body
(Seurat does not show these in his figure). Buccal capsule short
(10/*,) ; pharynx 28G> long, encircled in its posterior third by the
nerve ring. Combined length of pharynx and esophagus 1/3 that
of body. Cervical papillae GO and 9G>, respectively, from an-
terior end. Cloaca 11G> from posterior extremity. Two papillae
toward the posterior third of the tail. Only one spicule present,
slender, filiform, winged, measuring 480/* long. Anterior to the
cloaca there is a feebly chitinized organ, the gorgeret, and at the

SOP,"

Fig. 413. — Tetrameres nouveli. a, Female; b, ovejector ; c, seminal receptacle

AND OVIDUCT ; d, EGO WITH FILAMENTS ; C, HEAD END OF MALE ; f, TAIL END OF MALE.
(500/1 SCALE FOR a TO C ; 100 M SCALE FOR d TO f. ) AFTER SEURAT, 1914

side of this a chitinized part which Seurat regards as the rudiment
of a second spicule.

Female 1.7 mm. long. Body subglobular (fig. 413a). The
strongly distended median region of body measures 1 mm. long by
85G> maximum width. Buccal capsule 15/x long; pharynx 21G>
long, encircled in its posterior third by the nerve ring. Combined
length of pharynx and esophagus 1 mm. Cervical papillae 77/x and
87/x, respectively, from anterior end. Excretory pore ventral, 50>
posterior to nerve ring. Anus 175/* from posterior extremity. Vulva
L20> anterior to anus. Ovejector (fig. 413 b) of type of that of
T. inermis; vestibule 750/* long, cylindrical, ending in a slightly
swollen oval part. Sphincter very clear. Trompe Y-shaped, with a
short unpaired branch and Avith paired branches running parallel to
a length of 1 mm. Uteri parallel, very narrow (50/*), the distal
end swollen into an enormus seminal receptacle (fig. 413c), its trans-
verse diameter being 125//.. Oviducts and ovaries 4 mm. long. Eggs

350 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

54ju, long by 28/a wide, elliptical, slightly flattened on the side, with
thick shells; the embryonated eggs bear at each of the two poles
a tassel of long filaments (70/* in length). The filaments (fig.
•113 d) appear only as the embryo becomes fully developed; when the
egg is enclosed in the uterus the filaments are wrapped around it;
they unfold and spread out when the egg is set free in water.

Life history. — Probably similar in a general way to that of T.
ftssispina (p. 343).

Distribution. — Africa (Algeria) .

Fig. 414. — Tetrameres tetrica. Male. After Travassos, 1919
TETRAMERES TETRICA Travassos, 1917

Host. — Primary: Ar amides cajanea; secondary: Unknown; prob-
ably similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

Male 2.6 mm. long by 130 to 140/z wide (fig. 414). Cuticle trans-
versely striated and with numerous spines along median and lateral
lines. The spines commence 24/* from the anterior extremity where
they reach their greatest size (20^ long by 3/a wide) and then slowly
and progressively diminish to the posterior fourth of the body where
they disappear except on the ventral surface; there one finds 6 pairs
of preanal spines. Postanally there are 4 pairs of lateral spines and
also 4 pairs on the ventral surface of the tail. Buccal capsule 12/x
long by 6/t wide, irregular in shape. Pharynx 230^ long by 42/a wide.
Nerve ring 140/x from anterior extremity of body. Cloacal aperture
200/x from posterior extremity which is long and sharply pointed.
Two spicules of very different lengths, the larger 200/x long by 6/t

NEMATODE PARASITES OF BIEDS 351

wide and the shorter which is only slightly chitinized, 22/i long by
4/i wide.

Female 1.5 mm. long by 1 mm. wide. Body red, globular, with 4
deep longitudinal furrows corresponding to the median and lateral
lines. Buccal capsule oval, 16^. deep by 12ft wide. Tail slender and
very long. Ovaries 21/u, wide. Uterus ending posteriorly in a large
rounded seminal vesicle, about 10/x in diameter. Ovejector similar to
that of T. micropenis. Eggs 75 to 78/*, long by 21yu. wide ; ellipsoidal.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distinbution. — South America (Brazil).

TETRAMERES UNISPINA (Diesing. 1861) Travassos, 1914d

Synonym. — Tropidocerca unispina Diesing, 1861.

Host. — Primary : Corvus comix; secondary : Unknown ; probably
similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Tetrameres (p. 334).

Male unknown.

Female 3 mm. in longitudinal and transverse diameters. Body
globular. Mouth surrounded by 3 swollen protuberances which are
beset internally and externally with 2 points. The tail end is armed
with a spine.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — Not given for original material; reported from
United States by Stiles and Hassall.

Genus MICROTETRAMERES Travassos, 1915b

Generic diagnosis. — Tetrameridae (p. 333) : Long axis of female
body coiled in a simple or a complicated spiral. Body usually with-
out the 4 longitudinal furrows that are present in Tetrameres; if
present, these furrows also spirally coiled.

Type-species. — Microtetrameres crusi (Travassos, 1914) Travas-
sos, 1915.

As noted in the discussion under key to genera of Tetrameridae
(p. 333) I have changed the diagnosis as given by Travassos for
Micro tetrameres.

Kl-Y TO SPECIES OF MICKOTETUAMKIIES

1. Species wrongly placed in this genus, apparently identical with Eustrongy-

tides mergorum Microtetrameres inflata, p. 357.

2. Anns of female 450ft from posterior end; male <> mm. long; longer spicule

5.4 mm. in length Microtetrameres contorta, p. 353.

Anus of female not over 22~>n from posterior end; male not over 5 mm.
long; longer spicule not over '.'>.V> mm. in length 3.

352 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

3. Female 780/* long by 640/* wide, its pharynx 73/* long ; spicules 990/* and 100/i

long, respectively Microtetrameres minima, p. 358.

Female 1.2 to 2.5 mm. long by 1 mm. or more in width ; pharynx 160/* or
longer ; spicule lengths different from above 4.

4. Female 1.2 to 1.3 mm., male 4.9 mm. long; long spicule 3.6 mm. in length.

Microtetrameres helix, p. 355.

Female 2 mm. or more, male not over 4.75 mm. long; long spicule not over

2.3 mm. in length 5.

5. Anus of female 225/* from posterior end; size of male 4.75 mm. by 130/*;

spicules 2.3 mm. and 145/* long, respectively.

Microtetrameres spiralis, p. 360.

Anus of female not over 140/* from posterior end ; size of male not over 4

mm. by 120/*; long spicule not over 1.32 mm., short spicule not over

100/* long 6.

6. Anus of female 74 to 100/* from posterior end ; long spicule 651 to 787/* long.

Microtetrameres cruzi, p. 352.
Anus of female over lOO/i from posterior end ; long spicule over 1 mm 7.

7. Male 3.5 to 4 mm. long Microtetrameres pusilla, p. 359

Male 2.13 mm. long Microtetrameres inermis, p. 355.

MICROTETRAMERES CRUZI (Travassos,, 1914) Travassos, 1915b

Synonym. — T etrameres cruzi Travassos, 1914o\

Hosts. — Primary: Bucco swainsoni and Melanerpes -jlavifrons;
secondary: Unknown; probably similar in a general way to that of
T. -fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Microtetrameres (p. 351).

Male 1.17 to 1.4 mm. long by 85/* wide. Body threadlike, white,
strongly striated transversely but having no spines (fig. 4156).
Buccal capsule 12/t long by 4 to 5/x wide; pharynx 93/*, long by 9/*.
wide ; esophagus 290/*. long by 24/* wide. Anus 132/x from posterior
end. Spicules of very unequal dimension, the small one 82/*, the
larger 651 to 787/* long. Four pairs of papillae at the posterior end
of body, one pair preanal and 3 pairs postanal.

Female 2 mm. long by 1.5 mm. wide. Body red-colored, strongly
striated transversely, and with 4 longitudinal furrows corresponding
to the lateral and median lines; these furrows, along with the axis
of the body, are spirally coiled (fig. 415a). Extremities of body
conical projections. Buccal capsule 16 to 20/* long by 8/* wide;
pharynx 160/* long; esophagus 620/* long. Intestine saclike, filled
with black detritus, narrows toward posterior end and terminates
in a fine canal. Anus 74 to 100/* from posterior end. Eggs 50 to
60/* long by 24 to 28/* wide, many of them containing well-developed
embryos. Vulva 300/* from posterior end.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution. — South America (Brazil).

NEMATODE PARASITES OF BIRDS

353

MICROTETRAMERES CONTORTA (Weidman, 1913) Travassos, 1915b

Synonyms. — Tropidocerca contorta Weidman, 1913 ; Tetrameres
contorta (Weidman, 1913) Travassos, 1914.

Hosts. — Primary: Dichocereus oicornis; secondary: Unknown;
probably similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Mierotetrameres (p. 351'

7 :;v2 r . ;

FIG. 415. MICROTETRAMERES CRDZI. C, FEMALE ; b, MALE. AFTER TRAVASSOS, 1914

Male 6 mm. long by 125/* wide. Body subcylindrical, filiform
(fig. 416 a and b) ; cuticle transversely striated. Anterior extremity
tapers rather abruptly to a rounded end. Posterior extremity
tapers more gracefully; tail strongly curved toward cloaca; sharply
pointed. Cloaca 300^ from posterior extremity, surrounded by
prominent cuticular ring. Spicules unequal; shorter one 150/* long,
the longer one 5.4 mm. long. Four pairs of caudal papillae, two
pairs being preanal and two pairs postanal.

Female 2.1 mm. long by 1.9 mm. Body blood red, tightly coiled
in complex manner (fig. 416 d, e, and j) ; no longitudinal furrows as

354

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

in species of Tetrameres. Cuticle transparent, finely striated trans-
versely, often projecting as the coils tighten. Anterior and posterior
extremities projecting to varying degree, never to a length of more
than half the diameter of the worm, and sometimes retracted into
the center of the coil. Buccal capsule dome-shaped. Esophagus
long. Intestine a black irregular tract, twisting with the coils.
Posterior extremity (fig. 416 /) sharply pointed; anus 450/*, vulva
900/x from the posterior end, cuticle thickened into a rounded swell-
ing between the two openings. Eggs 40 to 45 ju, long by 20 to 25/a
wide, containing coiled embryos when mature; some eggs show
peculiar unilateral bib attached to outside of shell (fig. 416 g).

Fig. 416. — Microtetrameres contorta. a and b, Males ; c, yodngest

FEMALE, SHOWING TENDENCY TO COIL ; d AND e, MATURE FEMALES ;
f, TAIL OF FEMALE ; (J, OVA, UNILATERAL BIB REPRESENTED ON ONE ;
h, MATURE FEMALE, SHOWING COURSE OF ESOPHAGUS AND INTESTINE ;
X, HALF GROWN FEMALE, COILED IN ONE PLANE ONLY. AFTER WEIDMAN,

1913. j, Wax reconstruction of female. After Weidman, 1923

Weidman made a very careful study of females of varying ages
(fig. 416 c to e and h to j) ; one of the wax reconstructions made by
him is shown in figure 416 j. He concludes that the propensity to
coil is of very early development. The arrangement of coils is not
constant, being either clockwise or contraclockwise. Head always
bent dorsally; posterior extremity always twists suddenly in direc-
tion opposite to that of anterior coils. With egg production the
coils broaden out so that the mass appears globular.

Life history. — Probably similar in a general way to that of T.
flssispina (see p. 313).

Distribution. — North America (United States (Pennsylvania
(Zoological Garden, Philadelphia)).

NEMATODE PARASITES OF BIRDS 355

MICROTETRAMERES HELIX, new species

Host. — Primary : Corvus americanus; secondary : Unknown ; prob-
ably similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Microtetrameres (p. 351).

Male 4.9 mm. long by 100/n wide. There are no longitudinal rows
of spines on the body. Buccal capsule 21/* deep; pharynx 274/*,
long; esophagus 531/t long. Nerve ring 191/*, from anterior ex-
tremity. Tail (fig. 417a) slender; cloacal aperture 183/* from pos-
terior end ; cloacal lips prominent. Two small preanal and two small
postanal papillae in the ventral line. Two very unequal spicules,
the longer 3.6 mm. long, extending almost to esophageal region;
distally it ends in two sharp points. Shorter spicule 135/x long,
very feebly chitinized and therefore difficult to see; its distal end
rounded.

Female, when coiled, 1.2 to 1.3 mm. long by 1 to 1.3 mm. wide.
Body loosely coiled; starting from the head end there are approxi-
mately one and one-half turns or coils in one direction, the direction
then reverses itself for approximately one turn (fig. 417 o and c).
No longitudinal furrows as in Tetrameres. Cuticle very loose, pro-
jecting from the body in transparent folds. Head end blunt; buccal
capsule 22.5/* deep; pharynx 225 to 250/* long; esophagus very
thick, soon becoming obscured by the uterine coils so that its distal
end is not observable. Anus 141/*, vulva 216/* from posterior end,
which is finely pointed and projects out from a transparent cuticular
collar (fig. 417 d). Eggs 42/* long by 33/* wide; embryonated.

Life history. — Probably similar in a general way to that of T.
-fissispina (p. 343).

Distribution. — North America (United States (District of
Columbia) ).

Type material. — No. 2064 U.S.N.M. (Bureau of Animal Industry
helminthological collection).

MICROTETRAMERES INERMIS (Linstow, 1879) Travassos, 1915b

Synonyms. — Tropidocerca inermis Linstow, 1879«; Tetrameres
inermis (Linstow, 1879) Travassos, 1914ri.

Hosts. — Primary: Astur nisus, Astur palumbarius, Corvus corax
tingitanus, Passer domesticus, Corvus frugilegus, Corvus corone,
Lanius, species, and sparrow-hawk (Epervier) ; secondary: Un-
known; probably similar in a general way to that of T. -fissispina
(p. 343).

Location. — Proventriculus.

Morphology. — Microtetrameres (p. 351).

356

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 2.125 mm. long hy 90> wide (fig. 418 a to d). Buccal-capsule
19/* long. Pharynx 225/* long. Esophagus 395/* long. Cervical pa-
pillae slightly posterior to nerve ring, which surrounds the middle
of the pharynx. Cloacal aperture 128/* from posterior extremity,
bounded by a chitinous projecting ring. Caudal papillae distributed
as follows : 2 pair preanal ; one small median papilla situated at the
center of the space which they bound ; 2 pair postanal, slightly asym-
metrical. Spicules of unequal length, the left one 1.187 mm. long,
the right one 75/* long. Tail tapering, ends in small button.

Female forming cysts 2 mm. large. Body coiled in complex man-
ner (fig. 419 a to c). No longitudinal furrows as are found in spe-
cies of Tetrameres. Cuticle of anterior extremity folded to give

Q/f7)f7)

Figs. 417-418. — 417, Microtetrameres helix, a, Male tail; b and c, females;
d, female tail. Original. 418, Microtetrameres inermis. a, Male ; b, ven-
tral view and c, lateral view of tail of same ; d, head end of same. (Scale

APPLIES TO b, C, AND d. ) AFTER SEURAT, 1913

appearance of spy-glass (fig. 419d). Buccal capsule 20/* long.
Pharynx 300//, long (Linstow) or 260/* long (Seurat) by 40 /* wide.
Esophagus 1.125 mm. long (Seurat) or 2 mm. long (Linstow) by
24/* wide. Cervical papillae slightly posterior to nerve ring which
surrounds the middle of the pharynx. Intestine dark-brown.
Seurat (1913c) says that in the adult female the intestine is com-
pressed between the uteri, its walls almost pressed together, whereas
in the } 7 oung female the intestine is voluminous and entirely filled
with a maroon-colored mass which is undoubtedly a reserve supply of
material to be used later during egg formation. Anus 135/* from
posterior extremity. Vulva (fig. 420 6?) 95/* anterior to anus. Ves-
tibule of ovejector (fig. 420 a and 5) 875/* long. Trompe in the form
of a large reservoir 375/* long by 110/* wide, bifurcated at its end
where it joins the uteri. Seminal receptacle (fig. 420 c) piriform, of

NEMATODE PARASITES OF BIRDS

357

characteristic appearance in that the lining cells are wider than high
whereas those of the oviduct are higher than wide. Eggs (fig.
420 e and /) 36,u. long by 20> wide (Linstow) or 52/a long by 37ju.
wide (Seurat), embryonated, thick-shelled, cylindrical except for
operculated ends.

Seurat found that the eggs hatched after being kept in water at
room temperature for 48 hours. Larvae emerged at one of the
poles of the eggs, either the head or tail end emerging first. They
died as soon as they were free of the shell. Larva 235/x long, cuticle
finelv striated.

Fig. 419. — Microtetrameres inermis. Female, a, Adult ; b, adult examined
perpendicularly to determine manner of coiling; c, immature; d, anterior
end. After Seurat, 1913

Life history. — Probably similar in a general way to that of T.
flssispina. Seurat points out that the short life of the newly hatched
larva indicates that it passes through an intermediate host. He has
found a fourth-stage larva in the proventriculus of a crow.

Distribution.— Asia (Russian Turkestan) and Africa (Algeria
(Bou-Saada) ).

MICROTETRAMERES INFLATA (Mehlis, 1816) Travassos, 1915

Synonyms. — Spiroptera inflata Mehlis, 1846; Tropidocerca para-
doxa Diesing, 1851, part; Tropidocerca inflata (Mehlis, 184G) Dies-
ing, 18G1 ; Tropidocerca paradoxa Linstow, 1877 (not Tropidocerca
paradoxa Diesing, 1835) ; Tropisurus inflatus (Mehlis, 1846) Neu-
mann, 1892; Tetrameres inflata (Mehlis, 1846) Travassos, 1914.
Tetrameres haemochrous Creplin, 1846 is probably a synonym of
T . inflata; if not, it is a nomen nudum.

Mehlis (in Creplin, 1846) gave no description; his Spiroptera
inflata was a nomen nudum. Diesing gave the first descriptiou

358

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

under Tropidocerca inflata in 1861 but it was a one-line description
which is not recognizable. Linstow (1879a) says the material he
described in 1877 as Tropidocerca paradoxa is Diesing's Tropido-
cerca inflata. Since Linstow probably had access to the collections
in making his comparisons, and since we have no evidence one way
or the other, we must assume he is correct in saying he was dealing
with the same species as that of Mehlis, and accept his description
of it.

However, at a later date (1899) Linstow states that T. paradoxa is
a synonym of Hystrichis papillosus (Eustrongylides papillosus)
while Jaegerskiold (1909) has listed both T. paradoxa Linstow, 1877,
and Hystrichis papillosus Linstow, 1899 in part as synonyms of

Fig. 420. — Microtetramerbs inermis. a, Ovejector. After Seurat, 1914.
b, Part of same ; c, ovary, oviduct, and initial region of dterus ; d,
vulva ; e and f, hatching of eggs. After Seurat, 1913

Eustrongylides elegans. Linstow's description and figures are un-
questionably of a species of Eustrongylides and agree with E. elegans
in all particulars except the length of the spicule. It follows there-
fore, from all the available evidence, that Microtetrameres inflata is
in reality a species of Eustrongylides and the present writer will not
include the description of it in the Tetrameridae but under E. ele-
gans, the latter, however, apparently also being synonymous with
E. mergorum (p. 372).

MICROTETRAMERES MINIMA (Travassos, 1914) Travassos, 1915b

Synonym. — Tetrameres minima Travassos, 1914.

Hosts. — Primary: T achy phonics cristatus brunneus; secondary:
Unknown ; probably similar in a general way to that of T. flssispina
(p. 343).

L ocation. — Pro ventriculus.

Morphology. — Microtetrameres (p. 351).

NEMATODE PARASITES OF BIRDS 359

Male 1.4 mm. long (fig. 421). Cuticle cross-striated. Buccal cap-
sule very small. Spicules of very unequal lengths, 990/x and 100/i
long, respectively.

Female 780/* long by 640/x wide. Body red, strongly striated trans-
versely ; long axis coiled (this is assumed to be true as Travassos in-
cluded this species in Microtetrameres). Buccal capsule 12/i long;
pharynx 73/i long by 10/i wide; esophagus 490/a long by 50//, wide.
Intestine saclike, becomes thinner posteriorly. Anus 68/1 from pos-
terior extremity. Vulva slightly anterior to anus. Eggs 45/t long by
24/i wide, many of them embryonated.

Life history. — Probably similar in a general way to that of T .
fissispina (p. 343).

Distribution. — South America (Brazil).

MICROTETRAMERES PUSILLA Travassos, 19156

Synonym. — Tetrameres pusilla Travassos, 19155.

Hosts. — Primary: Turdus rufiventris and Platycichla flavipes;
secondary : Unknown ; probably similar in a general way to that of
T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Microtetrameres (p. 351).

Male 3.5 to 4 mm. long by 100 to 120/t wide. Body (fig. 422) fili-
form, white, cuticle transversely striated and without spines. Buccal
capsule cylindrical, 17/a deep by 7/* wide. Pharynx 300//, long. Eso-
phagus yellow-colored, slightly claviform. Travassos says it is 42/*
long but he evidently means width ; judging its length as compared
with that of the pharynx, in his figure, the esophagus measures about
600//. long. Two slender spicules of very different lengths, the smaller
about 85/* long by 5/i wide, the larger approximately 1.32 mm. long
by 7/i wide, having the basal part dilated while the distal end has a
rounded point. Cloaca 1 aperture 170/* from posterior extremity.
Five pairs of caudal papillae, asymmetrical, of which 2 are preanal,
1 adanal, and 2 postanal.

Female 2 mm. long by 1.5 mm. wide. Body of red color and with
its long axis spirally coiled (assumed to be true as Travassos places
this species in Microtetrameres). Culticle with marked transverse
striations and with 4 furrows corresponding to the median and lat-
eral lines. Buccal capsule barrel-shaped, 9/i long and 10/i in maxi-
mum width. Pharynx 273/t long. Esophagus cylindrical, 974/x long
(in his description in 1919 Travassos says 530/* long by 90/t wide).
Anus 140/x from posterior extremity. Eggs ellipsoidal, 42 to 49/*
long by 28/i wide.

Life history. — Probably similar in a general way to that of T.
fissispina (p. 343).

Distribution.— South America (Brazil).

360

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

MICROTETKAMERES SPIRALIS (Seurat, 1915) Travassos, 1915b

Synonyms. — Tropidocerca spiralis Seurat, 1915a; Tetrameres
spiralis (Seurat, 1915) Travassos, 1915&.

Hosts. — Primary : Bubulcus lucidus; secondary : UnknoAvn ; proba-
bly similar in a general way to that of T. fissispina (p. 343).

Location. — Proventriculus.

Morphology. — Microtetrameres (p. 351).

Male 4.75 mm. long by 130/x wide. Body filiform, white. Buccal
capsule (fig. 423a) 30/t long. Pharynx narrow, its length 1/4 that
of the esophagus. Combined length of pharynx and esophagus 1/3

&%

Figs. 421-423.— 421, Microtetrameres minima. Male. After Travassos, 1014. 422,
Microtetrameres pusilla. Male. After Travassos, 1919. 423, Microtetrameres
spiralis. Head and tail of male. After Seurat, 1915

that of body. Nerve ring encircles middle of pharynx. Cervical
papillae symmetrical, behind the nerve ring, 300^ from the anterior
extremity of the body. Excretory pore very small, 25ju in front of
the level of these papillae ; it is connected with a cuticular canal 50/i
long. Cloacal aperture (fig. 423 b) with 2 projecting lips, the lower
one more strongly developed. Four pairs of sessile genital papillae
near the cloacal aperture, 2 preanal and 2 postanal. Two spicules
of very different lengths, the right short (145/x) and curved, the left
2.3 mm. long, slender and filiform.

Female 2.5 mm. long by 2 mm. wide. Body red color, rolled spirall}'
(3^2 times). No longitudinal furrows as are present in species of
Tetrameres. Cuticle transversely striated. Buccal capsule bottle-
shaped, 30/* long. Pharynx 175/* long. Intestine brown or black.

NEMATODE PARASITES OF BIRDS 361

The cuticle is inflated in the posterior part of the body, into a muff-
like formation, from which projects the caudal extremity, with a
length of 180y. Anus 225/* from posterior extremity; tail digitiform,
rounded at the end and carrying at the point a small rounded spike.
Vulva a short distance in front of anus. Ovejector similar to that of
M. inermis; vestibule turnip-shape, 1.6 mm. long, containing a large
number of eggs ; sphincter short ; trompe Y-shaped ; combined length
of vestibule, sphincter and unpaired trompe 2.6 mm. Uterus 95^t in
diameter, filled with eggs disposed in 3 linear rows; receptaculum
seminis not delimited. Oviducts short (300/*) ; ovaries slender, fili-
form, 6.5 mm. long. Eggs 50/x long by 30/* wide, embryonated at
maturity, thick-shelled, oval, flattened on one face.

Larva 2.3 mm. long by 70/* wide. Buccal capsule 20/x long;
pharynx 320/i. long, surrounded in the middle by nerve ring ; esopha-
gus 600/x long. Cervical papillae 210/x from the anterior extremity.
Tail pointed, 170/a long.

Life history. — Probably similar in a general way to that of T. /?.<?-
sispina (p. 343).

Distribution. — Africa (Algeria) .

Family ANCYRACANTHIDAE Railliet, 1916

Family diagnosis. — Spiruroidea (p. 162) : Head with 4 pinnate,
posteriorly directed appendages, set crosswise. Male with caudal
extremity coiled in spiral; caudal alae present, provided with papil-
lae. Female with vulva posterior to middle of body ; 2 uteri.

Parasitic in digestive tract of fish, reptiles, and birds.

Type-genus. — Ancyracanthus Diesing, 1838.

Genus ANCYRACANTHOPSIS Diesing, 1861

Generic diagnosis. — Ancyracanthidae (p. 361) : Body capillary.
Head continuous with body, armed with 4 pinnate, posteriorly di-
rected appendages, set crosswise. Mouth terminal, with 2 small
papillae or lips. Male with caudal extremity coiled twice in spiral,
with 2 alae provided with papillae. Female with caudal extremity
which may be spirally twisted, apex obtusely conical. Vulva situated
posterior to middle of body.

Parasitic between the tunics of the gizzard of birds.

Type-species. — Ancyracanthopsis bilabiata (Molin, 1860) Diesing,
1861.

Diesing created this genus to remove from Ancyracanthus the one
species at that time reported from a bird, all the others being fish or
reptile parasites. At a later date Mueller, in describing a second
species from a bird, states that the bird forms do not have the com-
plicated " immersionssystem " that is present in the species found in
reptiles and fish (see discussion under A. bihamata. p. 362).

362 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

KEY TO SPECIES OP ANCYRACANTHOPSIS

Male 3.5 to 4 mm., female 5 to 6 mm. long; vulva in posterior part of body;

in Sterna risoria Ancyracanthopsis bihamata, p. 362.

Male 7 mm., female 9 mm. long; vulva slightly posterior to middle of body,

dividing body length in ratio of 4 : 3 ; in Eurypyga helias.

Ancyracanthopsis bilabiata, p. 362.

ANCYRACANTHOPSIS BILABIATA (Molin, 1860) Diesing, 1861

Synonym. — Ancyracanthus bilabiatus Molin, 1860<i.

Host. — Primary : Eurypyga helias; secondary : Unknown.

Location. — Between tunics of gizzard.

Morphology. — Ancyracanthopsis (p. 361) : Body attenuated an-
teriorly, densely striated transversely. Head with 4 pinnate append-
ages which are larger in the male than in the female.

Male 7 mm. long. Caudal extremity twisted twice in spiral. Cau-
dal alae wide, papillae short. One spicule long and filiform, the
other short, thick, navicular.

Female 9 mm. long. Caudal extremity twisted in spiral, apex
obtuse, depressed in center. Anus not far from posterior extremity.
Vulva in posterior part of body, prominent, bilabiate.

Life history. — Unknown.

Distribution. — South America (Brazil).

ANCYRACANTHOPSIS BIHAMATA (Mueller, 1897) Cram, 1927

Synonym. — Ancyracanthus bihamata Mueller, 1897.

Host. — Primary: Sterna risoria; secondary: Unknown.

Location. — Between tunics of gizzard.

Morphology. — Ancyracanthopsis (p. 361) : Body threadlike, with
fine transverse striations. Anterior part of body (fig. 424 a and b)
diminished gradually or sometimes suddenly to one-fifth the average
thickness. Mueller states that this anterior part of body is appar-
ently capable of extension and retraction ; the esophagus in this sec-
tion is very convoluted. Head with 4 posteriorly directed pointed
processes, 23 to 40/x long, similar to those of A. bilabiata. In addi-
tion, at the ba*se of the large processes the head bears a crown of
smaller papillae or projections, apparently 8 in number, sometimes
resembling the larger ones in form. Esophagus in 2 parts.

Male 3.5 to 4 mm. long by 170 to 180/* wide. Caudal extremity
(fig. 424 c and d) rolled in loose spiral. Caudal alae long and nar-
row, but thick; 6 pairs of preanal, 5 pairs of postanal papillae along
the margin and, in addition, in the median field at the posterior ex-
tremity a pair of small papillae and anterior and posterior to this
pair, there is a pair of small knobs. Spicules unequal, the right (fig.
424 e) IOO/a long, ending in 3 fine points, the left 34/x long and, ac-

NEMATODE PARASITES OF BIEDS

363

cording to Mueller, serving not only as a gubernaculum for the
longer spicule but also as a clasping organ.

Female 5 to 6 mm. long by 200/i wide. Vulva (fig. 424 g) slightly
prominent, situated slightly posterior to middle of body, dividing
the body length in ratio of 4 : 3. Anus 100/* from posterior extremity ;
body narrowed into a conical point posterior to anus (fig. 424 /').
Eggs 40/a long by 33/*. wide, embryonated, bluntly oval, the shell
10/* thick.

Life history. — Unknown.

Distribution. — Not given (Germany?).

This species should certainly be in the same genus with A. bilabiata.
Mueller evidently did not know that Diesing had made a new genus
for the species bilabiatus as he calls it Ancyracanthus bilabiatus and

Fig. 424. — Ancyracanthopsis bihamata. a and b, Head end ; c and d, male
tail; e, spicule; f, female tail; g, vulva. After Mueller, 1897

says his species is very close to it. As in it, his species does not have
the " immersionssystem," a complicated system of connections be-
tween the pinnate appendages and cervical sacs; Mueller made
numerous longitudinal sections of the appendages. This system is
present in the genus Ancyracanthus but apparently lacking in
Ancyracanthopsis. The host difference between the two genera would
also throw bihamatus into Ancyracanthopsis, the latter having been
made for bird forms and Ancyracanthus left for fish and reptile
forms.

Family GNATHOSTOMIDAE R. Blanchard, 1895

Synonyms. — Cheiracanthidea Diesing, 1861 ; Oxyuridae Railliet
and Henry, 1916, in part; Heterakidae Seurat, 1918, in part.

Family diagnosis. — Spiruroidea (p. 162) : Mouth with 2 large tri-
lobed, lateral lips, a longitudinal tooth-like ridge on their inner sur-
face, meeting the one on the opposite side. Male with caudal alae
more or less well-developed ; two spicules. Female with vagina an-

364 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

teriorly directed, giving off 2 to 4 uterine branches. Eggs with thin
shells, ornamented externally with fine granulations.

Parasitic, usually in the digestive tract, of reptiles, fish, mammals;
rarely, as larvae, encysted in birds.

Type-genus. — Gnathostoma Owen, 1836.

Subfamily Gnathostominae Baylis and Lane, 1920

Sub family diagnosis. — Gnathostomidae (p. 363) : A cuticular head
bulb present, provided with transverse striations or rows of hooks,
and containing 4 (or 6?) membranous ballonets, the cavity of each
communicating with an elongated, blind, cervical sac hanging freely
in body cavity and functioning as glandular apparatus.

Type-genus. — Gnathostoma Owen, 1836.

Genus GNATHOSTOMA Owen, 1836

Synonym. — G heir acanthus Piesing, 1838.

Generic diagnosis. — Gnathostominae (p. 364) : Head-bulb armed
with simple hooks. Anterior part or entire body covered with spines,
the anterior spines incised into points of varying number and shape.
Male with unequal spicules; 4 pairs of large lateral and 2 pairs of
small ventral caudal papillae. Female with vulva posterior to mid-
dle of body.

Parasitic normally in gastric wall, usually of carnivorous mam-
mals; occasionally as larvae in subcutaneous tissue of birds.

Type-species. — Gnathostoma spinigerum Owen, 1836.

KEY TO SPECIES OF GNATHOSTOMA

Glandular apparatus of head consisting of 6 tubes; length of esophagus equal
to almost 1/2 that of body ; as encysted larvae and immature adults in
Pelecanus onocrotalus Gnathostoma pelecani, p. 365.

Glandular apparatus of head consisting of 4 tubes ; length of esophagus equal
to 1/3 that of body ; as encysted larvae in Aquila impenalis.

Gnathostoma accipitri, p. 364.

GNATHOSTOMA ACCIPITRI Skrjabin, 1916b

Host. — Aquila imperialis. The parasite is probably aberrant so
that this does not represent either primary or secondary host.

Location. — Encysted in subcutaneous connective tissue.

Morphology. — Gnathostoma (p. 364) : Larval form. Body curled
in circle in nodule; body length 2.8 mm., width 510/x. Head (fig.
425a) set off from body by depression and encircled by 4 parallel
transverse rows of chitinous hooks or spines (fig. 425 b), 44 spines
in each circle. Head 150> long by 340/* wide. Two valve-like lips,
each with a strongly-developed papilla. Cuticle of body covered

NEMATODE PARASITES OF BIRDS

365

with small chitinous spines, posteriorly directed, arranged in trans-
verse rows. Posterior extremity obtusely rounded. Esophagus 1/3
the total body length, its anterior part cylindrical, posterior part
bulbous. Around the anterior part of esophagus 4 cylindrical cervi-
cal glands. Intestine large, cylindrical, ending at anus at posterior
extremity of body.

Life history. — Unknown.

Distribution, — Asia (Russian Turkestan).

GNATHOSTOMA PELECANI (Chatin, 1S74) Skrjabin, 1916b

Synonym, — Sclerostoma pelecani Chatin, 1874.
Host, — Pelecanus onocrotalus. The parasite is probably aberrant
so that this does not represent either primary or secondary host.

M00^r^?--

Figs. 42. r >-42f>. — 425, Gnathostoma accipitri. a, General appearance : 6, hooks
of cephalic REGION. After Skrjabin, 1916. 426, Gnathostoma pelecani. a,
General appearance ; h, glandular apparatus ; c. vulva. After Chatin, 1874

Location, — As larvae in cysts in the subcutaneous connective tissue
and as immature adults in cysts in the respiratory sacs.

Morphology. — Gnathostoma (p. 3G4) : Length 3 to 4 mm. long.
Denticulations very pronounced in the anterior region of the body.
Head (fig. 426«) rounded, flattened, with 4 concentric and super-
posed series of chitinous hooks. Glandular apparatus (fig. 42GZ>)
in conjunction with the mouth consists of 3 pairs of tubes, 40^ in
diameter, of unequal length, the largest ones being external, the
others internal. Digestive tract consists of esophagus and intestine;
esophagus swollen posteriorly, its length equal to almost 1/2 that
of the body. Anus at posterior extremity of body, which ends in a
small mucronate point 7(V long. The immature adults showed a
characteristic vulva (fig. 42G c) in the median region of the body.

Life history. — Unknown.

Distribution. — Europe (France (Paris Museum)).

366 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Chandler (1925) has recently studied Gnathostome larvae which
he found encysted in snakes and in burrows in the liver and peri-
toneal walls of cats, in India. He concludes that these must be
classed for the present as Gnathostoma pelecani. They differ in
certain respects from Chatin's description, however, in that they
have 4 cervical sacs of equal length, and the oviduct which can be
faintly traced from the vulva leads forward, whereas G. pelecani is
described as having 6 cervical sacs of unequal length and the oviduct
leading backward.

Superfamily DIOCTOPHYMOIDEA Railliet, 1916

Superfamily diagnosis. — Spirurata (p. 162) : Nematodes of median
or very large size. Cuticle relatively transparent, at the anterior
and posterior extremities transversely striated, with or without
spines. Mouth without lips but with 6, 12, or 18 papillae forming
1 or 2 circles. Esophagus very long, without bulb; esophageal
glands well-developed, all three of about equal size and opening at
almost the same level. Male with closed caudal bursa or as the
present writer prefers to call it " bursal cup " (an attempt having
been made in this paper to confine the term " bursa " to the structure
found in the strongyles), bell-shaped, with muscular walls and with-
out rays. Female with anus at the very posterior end of body, in
middle of obtuse tail. Vulva either in neighborhood of anus or
in anterior part of body (1/10 of body length from head end). Va-
gina very long. Eggs with thick shell, at the poles of different
construction than elsewhere; surface of shell pitted or, rarely, non-
pitted but with ridges {Hystrichis acanthocephalicus) .

Parasitic as adults in digestive tract of birds or kidneys and body
cavity of mammals.

Type- family. — Dioctophymidae Railliet, 1915.

Various writers refer this superfamily to Railliet, 1910, but the
present writer has been unable to verify this earlier date ; moreover,
it is unlikely that the superfamily would be created before the
family.

Family DIOCTOPHYMIDAE Railliet, 1915

Synonym. — Eustrongylidae Leiper, 1908.

Family diagnosis. — Dioctophymoidea (p. 366) : Characters of the
superfamily.

Type-genus. — Dioctophyme Collet-Meygret, 1802.

KEY TO GENERA OP DIOCTOPHYMIDAE

Body without spines ; head with 12 to 18 papillae Eustrongylides, p, 367.

Body (in at least the anterior region) with spines; head with 6 papillae.

Hystrichis, p. 375.

NEMATODE PARASITES OF BIRDS 367

Genus EUSTRONGYLIDES Jagerskibld, 1909

Generic diagnosis. — Dioctophymidae (p. 366) : Body equally thick
throughout or thickened in median part. Head not enlarged, pro-
vided with 12 to 18 papillae, arranged as 2 circles. Cuticle of an-
terior and posterior extremities transversely striated, without spines.
Male with closed bell-shaped caudal bursal cup, the structure of
which is very typical for the individual species. One spicule, very
long. Female with anus in middle of rounded extremity; vulva
directly adjacent to anus.

Parasitic in glands of proventriculus or other parts of digestive
tract of birds (for the most part aquatic birds feeding on fish).

Type-species. — Eustrongylides tubifex (Nitzsch, 1819) Jaegers-
kiold, 1909.

KEY TO SPECIES OF EUSTRONGYLIDES

1. Unrecognizable species Eustrongylides papillosus, p. 373.

Recognizable species 2.

2. The inner crown of round, relatively low or very low papillae, more dif-

ficult to see than the outer ; body proportionately short and thick, the

female especially with much thickened middle region 3.

Inner crown of papillae as high or higher than those of outer crown, easily
seen ; body slender, no thickening of middle region even in female 4.

3. Papillae of outer circle relatively short, not digitiform. Mouth cavity com-

paratively short, scarcely half as long as the width of the body at that

level Eustrongylides tubifex, p. 367.

Papillae of outer circle long (36 to 44/*), digitiform. Mouth cavity compara-
tively long, its length almost as great as the width of body at that level.

Eustrongylides mergorum, p. 372.

4. Six papillae in inner circle, 12 in outer, making a total of 18.

Eustrongylides perpapillatus, p. 374.
Six papillae in each circle, making a total of 12 5.

5. Papillae of both circles of same size and with narrow bases; mouth cavity

very short, its length only % to % as great as body width at that level.

Eustrongylides africanus, p. 369.

Papillae of the outer circle of different size than those of the inner ; none

of the papillae with narrow bases. Mouth cavity long, its length at

least % as great as body width at that level 6.

6. Margin of bursal cup entire, without incision. Eustrongylides ignotus, p. 371.
Margin of bursal cup with a deep incision on the ventral side.

Eustrongylides excisus, p. 370.

This key is a translation, with slight modifications, of the one
given by Jaegerskiold.

EUSTRONGYLIDES TUBIFEX (Nitz.ch in Rudolphi. 1819) Jac*erskiold. 1909

Synonyms. — Strongylus tubifex Nitzsch, 1819, part; Eustrongylus
tubifex (Nitzsch, 1819) Diesing, 1851, part; Hystrichis tubifex
(Nitzsch, 1819) Molin, 1861, part.

Hosts. — Primary: Anas boschas dom-estica, Colymbus arcticus, C.
septentrionalis; secondary : Unknown, but fish probably (see below
3612—27 25

368

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

(life history) and see also E. ignotus, p. 371), according to Jaeger-
skiold and to Ciurea.

(Location. — Intestine.

Morphology. — Eustrongylides (p. 367) : The middle of the worm
is much thickened, the extremities are slenderer, the anterior por-
tion longer and slenderer than the posterior. The 6 papillae near
the mouth (fig. 4276) very small and those outside of these are
larger but inconspicuous. Cuticle grossly annulated, the annulation
almost disappearing in the middle portion of the body. The mouth
aperture is usually triangular, but may be 6-angled and give the
impression of being round. The mouth cavity is triangular in cross-
section and is 100 to 160/x, long.

Male 34 mm. long by 2 mm. wide. Bursal cup (fig. 427«) trumpet-
shaped, its margin with 2 incisions on the ventral surface.

Fig. 427. — Eustrongylides tubifek. a, Male tail; 5, heiad
end; c, female tail. Aftek Jaegerskiold, 1909

Female 35 to 44 mm. long by 2.5 to 3 mm. wide. Vulva near
anus (fig. 427 c). Vagina 11 mm. long. Eggs 65 to 75/x. long by 44ju
wide, oval with blunt ends and thick shells, the latter pitted.

Life history. — Unknown; probably involves intermediate stages
in fish, according to Jaegerskiold and to Ciurea (1924). Larvae
described as Filaria cystica by Rudolphi from under the peritoneum
and in the muscle of certain fish (Symbranchus laticaudatus and
Galaxias scriba) were regarded by Jaegerskiold as the larva of a
species of Eustrongylides; see E. ignotu-s, p. 371. Recently Ciurea
has found similar larvae in certain fish (Ba.rhus fluviatilis, Lota
lota, Esox lucius, and Perca fluviatilis) in the Danube and regards
them as larval forms of some species of this genus. As E. tubifex
is the type species of the genus, Ciurea's larvae are figured here
(fig. 428 a to d) to show the probable nature of the larval Eus-
trongylides, the specific identity of the larvae not being ascer-

NEMATODE PARASITES OF BIRDS

369

tained and the life history still lacking experimental continuation.
The larvae are relatively large, 28 to 70 mm. long by 264 to 539/u
wide, rose-red or brown-red in color. On each side of the body
near the anterior end is a row of small lateral papillae. The month
aperture has the form of a cleft and has about it 3 small pointed
papillae on each side and beyond these 3 large papillae on each
side. The larvae have tails of 2 types, one enlarged near the end
and regarded as that of the male, and the other rounded off without
enlargement and regarded as that of the female. See also E. ignotus,
p. 371.

Distribution. — Europe.

plg. 428. eusteongylides laevae. species not determined. (1 , front view of

iiead ; b, anterior end; c, outline of male tail,; (t, outline op female tail.
After Ciubea, 1924

EUSTRONGYLIDES AFRICANUS Jaegcrskiold, 1909

Hosts. — Primary: Anhinga rufa, Ardea) goluith, Leptoptilus cru-
menifer, Pelecanus rufescen^, Platalea leucorodia; secondary: Un-
known, probably fish; see E. tubifea;, p. 367, and E. ignotus, p. 371.

Location. — Proventriculus.

Morphology. — Eustrong glides (p. 367) : Body almost equally thick
throughout, without noticeable swelling in the middle region. Head
(tig. 429«) with 12 papillae in 2 circles of 6 each, those of the inner
circle are somewhat taller but otherwise not as large as those of
the outer. The papillae of the inner circle are 56/*. long by 40 to
56/7. wide; those of the outer circle 48^ long and 72 to 96/x wide.
Cuticle with coarse transverse striatums. Mouth cavity distinct
though not large, of triangular cross-section; its length is 80 to lOOp.

Male specimen imperfect.

Female 90 to 166 mm. long; maximum width 1.5 to 2.5 mm., width
at anterior end of body 430 to 575/1, at tail end (fig. 429 b). Esopha-
gus about 16.5 to 18 mm. long. Nerve ring 160/* from anterior end.
Eggs (fig. 429 c to e) 70 to 76/x long by 36 to 42/t wide.

Life history. — Unknown ; see E. tubifecc, p. 367.

hixtHbution— Africa (Sudan) and Asia (Russian Turkestan (lac
Kul-Kainar) ).

370 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

EUSTKONGYLIDES EXCISUS Jaegerskiold, 1909

Synonyms. — Strongylus tubifex Rudolphi, 1819, part; Eustrongy-
lus papilloma Diesing, 1851, part; Hystrichis papillosus Molin,
1861, part; Hystrichis elegans Stossich, 1899, part.

Fig. 429. — Eustrongylides afiucanus. a, Head end ; b, female tail; c, ex-
terior OF EGG ; d, riTTING OF SHELL ; e, INTERIOR OF EGG. AFTER JAEGERSKIOLD,

1909

Hosts. — Primary : Phalacrocorax carbo and P. pygmaeus; second-
ary : Unknown, probably fish ; see E. tubifex, p. 367, and E. ignotus,
p. 371.

Location. — Gizzard.

Fig. 430. — Eustrongylides excisus. a, Head; b and c, tail of male.

Jaegerskiold, 1909

After

Morphology. — Eustrongylides (p. 367) : Head (fig. 430a) with 12
papillae, those of the inner circle longer and more slender than
those of the outer, the inner ones measuring 25 to 30/* long, spine-
like with a sharp point, the outer ones 20 to 25/* long, wart-like,
with broad bases. Mouth cavity moderately large, 110 to 150/* long.

NEMATODE PARASITES OF BIRDS

371

Male 28 to 35 mm. long; maximum width 630 to 800/*, width at an-
terior end of body 168 to 190^., at posterior end (just anterior to
bursal cup) 370 to 480/u.. Esophagus 7.3 to about 12 mm. long.
Bursal cup 480 to 560/x long by 560 to 700/i. wide, trumpet-shaped
(fig. 430 b and c) with a distinct, fairly deep incision on the ventral
side.

Female evidently imperfect; length not given. Maximum width
1.2 mm., width at head end 240^; esophagus 12 mm. long.

Life history. — Unknown ; see E. tubifex, p. 367.

Distribution. — Europe (Austria (Museum, Vienna)) and Asia
( Russian Turkestan ) .

Fig. 431. — Eustrongylides ignc-tus. a, Head; b, female tail; c and d, male

TAIL; e AND f, EGG. AFTER JAEGERSKIOLD, 1909
EUSTRONGYLIDES IGNOTUS Jaegerskiold. 1909

Synonyms. — Filana cystica Rudolphi, 1819; Agamonema cysticuni
(Rudolphi, 1819) Diesing, 1851; Eustrongylus papillosus Diesing,
1851, part; Hystrichis papillosus Molin, 1861, part; Eustrongylus
tubifex Schneider, 1866, part; Spiroptera bicolor Linstow, 1899.

Hosts. — Primary: fAnMnga anhinga, Ardea cocoi, A. herodias,
liotaurus pinnatus; secondary : Fish ; larvae in Symbranchus laticau-
datus and Galaxias scriba resemble this species, according to Jae-
gerskiold, and Chapin has found the preadult stage of this species
in Fundulus diaphanus.

Location. — In fat around gizzard of primary host, in body cavity
of secondary host.

Morphology. — Eustrongylides (p. 367) : Body equally thick
throughout. Head (fig. 431«) witli 12 papillae in 2 circles, those
of the inner circle being larger and more conspicuous than those of
the outer circle, which are low and wart-like. Mouth opening hexag-
onal. Mouth cavity 100 to 160/a long.

Male 36 mm. long; maximum width 1 to 1.4 mm., width at head
end 250^, at tail end (just anterior to bursal cup) 350 to 460/*.

372 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Esophagus 8 mm. long. Bursal cup (fig. 431 c and d) 290 to 350.x
long by 400 to 550/*. wide; no incision of margin on ventral surface

Female 55 to 96 mm. long; maximum width 1.5 to 2 mm. ; width at
head end 250 to 400/a. Esophagus 11.6 to 16.4 mm. long. Eggs 58
to 66ti long by 35 to 44/* wide; shell 4.5ii thick, with pittings rela
tively sparse (fig. 431 e and /).

Life history. — Unknown, involving intermediate stages in fish.
Jaegerskiold states that larvae found in Brasilian fishes by
Schneider and by Leuckart resemble this species and Chapin recently
(1926) has found the preadult stage of this species in Fundulus
diaplianus at Washington, D. C. In a large number of fish exam-
ined by Chapin, each fish contained one to three specimens of the
nematode, the adult characters of which could be seen within the
last cuticle, corresponding exactly to those of adult worms found by
him in Ardea herodias, also at Washington, D. C. See also E.
tubifex, p. 367.

Distribution. — Europe (Germany (Berlin) and Austria (Museum,
Vienna) ).

EUSTRONGYLIDES MERGORUM (Rudolphi, 1809) Cram, 1927

Synonyms. — Strongylus mergorwnv Rudolphi, 1809; Strongylus
papillosus Rudolphi, 1809, part; Strongylus elegans Olfers, 1816;
Strongylus tubifex Rudolphi, 1819, part; Tropidocerca paradoxa
Linstow, 1877 not T. paradoxa Diesing, 1851 ; Hystrichis elegans
(Olfers, 1816) Railliet, 1893, part; Eustrongylides elegans (Olfers,
1816) Jaegerskiold, 1909; Microtetrameres inflata (Mehlis, 1846)
Travassos, 1915 and its synonyms (p. 357).

It is regrettable that the well-known specific name elegans should
have to be dropped but Rudolphi's name clearly antedates it and is
not a nomen nudum, having a brief description by him, based on a
description and figure by Redi.

Hosts. — Primary : Alca torda, Anas hoschas domestica, A. glacialis,
A. mollissima, Charadrius pluvialis, Ciconia nigra, Colymbus septen-
trionalis, Harelda glacialis, Merganser serratus, Mergus albellus, M.
merganser, M. serrator, Numenius arquatus, Phalacrocorax carbo,
Prodiceps cristatus, P. minor, Somateria molisswia, TJria troile;
secondary : Unknown ; probably fish, according to Jaegerskiold and
to Ciurea. See E. tubifex, p. 367, and E. ignotus, p. 371.

Location. — In tubercles in the esophagus and pro vent riculus.

Morphology. — Eustrongylides (p. 367) : White, thick fusiform
worms, attenuating at the 2 extremities. The head end (figs. 432 and
433) is elongate piriform, connecting by a slender, neck-like part
with the middle portion of the body, in some adults. About the
mouth is a circlet of 6 lateral and submedian papillae, and outside of

NEMATODE PARASITES OF BIRDS

373

this circlet is another of 6 larger papillae, 36 to 44//, long. Cuticle
grossly annulated except in the middle of the body. Head end
rounded: mouth opening round; mouth cavity narrow and tubular,
110/x to 175//. long (fig. 434a).

Male 18 to 53.5 mm. long by 1.5 to 2 mm. wide. Esophagus 6 to
10 mm. long. Bursal cup (fig. 434 c and (/) almost trumpet-shaped,
275 to 320//. long by 260 to 360//, wide; it is often not in the direct
body line but curved dorsally; its margin is more or less scalloped.
Spicule almost 8 mm. long (in Microtetrameres inflata 3.6 mm. long,
according to Linstow).

Figs. 432-433. — Eustrongylides mergorum. 432, a, Female ; b, male ; natural
size ; c, head. After Linstow, 1877. 433, a, Front view and 5, lateral
view of head; c, egg. After. Jaegerskiold, 1909

Female 25 to 36 mm. long by 2 to 2.75 mm. wide (it probably be-
comes as long as male or longer). Esophagus 8 to 12 mm. long.
Posterior end bluntly rounded (fig. 4345). Vulva close to anus.
Eggs 60 to 70//. long by 33 to 38//, wide, with blunt ends and pitted
shells.

Life history. — Unknown; see E. tubifex, p. 367.

Distribution. — Europe.

EUSTRONGYLIDES PAPILLOSUS (Rudolphi, 1802) Jaegerskiold, 1909

Synonyms. — Strongyl/us papillosus Rudolphi, 1802, part ; Eustron-
gyhis papillosus (Rudolphi, 1802) Diesing, 1851, part; Ilystnchis
papillosus (Rudolphi, 1802) Molin, 1861, part.

Hosts. — Primary : Anas boschas domestica, Anser cinereus domes-
ticus, and Nuecifraga caryocatactes ; secondary : Probably fish ; see
E. tubifex, p. 367, and E. igrnotw, p. 371.

Location. — In tubercles in the esophagus.

Morphology. — Eustrongylides (p. 367) : Body not enlarged in the
middle portion, according to Jaegerskiold; Linstow says it is thick-
ened. Mouth with the usual 2 circles of 6 papillae each about it. as
in most other species of this genus.

374

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Male 19 to 30 mm. long by 1 to 2 mm. wide. Bursal cup (fig.
435) trumpet-shaped, cloacal aperture at bottom of bursal hollow;
the bursal margin is fringed or papillate. Spicule long and slender.

Female 29 mm. long by 2.6 mm. wide, Eggs 68/x long by 38^ wide,
with 2 shells, the outer with pitted markings, and with opercula at
the ends.

This species is not well described, according to Jaegerskiold. The
above description is from Linstow.

Life history. — Unknown; probably involves intermediate stages
in fish. See E. tubifex, p. 367.

Distribution. — Europe.

FIG. 434. EUSTRONGYLIDES MEEGORUM. a, HEAD END ; I), FEMALE TAIL ; AND d,

MALE TAIL. AFTER JAEGERSKIOLD, 1909

EUSTRONGYLIDES PERPAPILLATUS Jaegerskiold, 1909

Synonyms. — Eustrongylus papillosus (Rudolphi, 1802) Diesing,
1851, part; Hystrichis papillosus (Rudolphi, 1802) Molin, 1861, part.

Hosts. — Primary: Ardea leuca and Herodias egretta; secondary:
Unknown, probably fish. See E. tubifex, p. 367.

Location. — Not given.

Morphology. — Eustrong glides (p. 367) : A total of 18 cephalic
papillae (fig. 436 a and b) instead of the usual 12; of these, 6 form
an inner circle and 12 an outer. Those of the inner circle are small
round knobs with a small point ; of the outer circle 6 are larger than
the others, the small alternating with the large. Mouth opening
triangular. Mouth cavity 96 to 150/a long.

Male 47 mm. long by 1 mm. wide ; maximum width about 1 mm.,
width at head end 175 to 200/*,, at posterior end (just anterior to

NEMATODE PABASITES OF BIRDS

375

bursal cup) 270 to 290jt. Esophagus 10.4 mm. long. Bursal cup
(fig. 436c) 290 to 340/u long by 320 to 350/x wide; it is compressed so
that the shape is almost spherical, and has on the ventral side 2 dis-
tinct rounded indentations between which is a wide tongue-like pro-
jection.

Female 17 mm. long; maximum width 1.2 to 1.5 mm., width at
head end 300/*, at tail end (fig. 436 d) 250/x. Esophagus about 13.6
mm. long. Eggs 53 to Olju. long by 31 to 33ju. wide, the poles very
wide, the shells thin, with closely-set pittings (fig. 436 e and f).

Life history. — Unknown; see E. tubifex, p. 367.

Distribution: — South America (Brazil).

435.

Figs. 435-436. — 435, Eustrongylides papillosus. Male tail. Afteu
jaegerskiold, 1909. 436, eustrongylides perpapillatus. (i, front
view and b, lateral view op head ; c, male tail - , d, female tail;
e and f, eggs. After Jaegerskiold, 1909

Genus HYSTRICHIS Dujardin, 1845

Generic diagnosis. — Dioctophymoidea (p. 366) : Body usually
equally thick throughout but may at times be much swollen in middle
part. Head with a circle of 6 relatively small papillae, 2 of which are
lateral, 4 submedian. Head more or less swollen. Cuticle of ante-
rior and posterior parts of body with coarse cross-striations. Head,
usually the anterior part of body and sometimes the whole body
covered with spines, the head usually being more thickly covered
and its spines sometimes of different shape than those of other
parts. Esophagus long, without bulb. Male with a bursal cup more
or less bell-shaped. One very long spicule. Female with anus in
middle of rounded posterior end; vulva with same situation as that
of anus or in its vicinity.

Parasitic usually in the glands of proventriculus of birds, chiefly
water birds but those which do not, or at least not exclusively, live
on fish.

Type-species. — Hystrichis tricolor Dujardin, 1845.
3(512—27 26

376 BULLETIN 140, UNITED ST/ ES NATIONAL MUSEUM"

KEY TO SPECIES O^' HYSTKICHIS

1. Only one circle of spines present ; it is directly posterior to mouth, the spines

very large Hystrichis coronatus, p. 378.

Numerous circles of spines, on head and cervical region if not on entire
body length 2.

2. Spines of head and of cervical region of approximately the same size, or

the latter smaller than the former 3.

Spines of head and those of anterior part of body of different size, the latter
being longer and larger G.

3. Head end with marked bulbous swelling Hystrichis cygni, p. 378.

Swelling of head end not of marked bulbous nature 4.

4. Spines of head comparatively sparce, only about 30 in each circle ; vulva

not at tail end but 175/u. anterior to it ; eggs not pitted but with network

of ridges Hystrichis acanthocephalicus, p. 377.

Spines of head thickly set, about 50 to 57 in each circle ; vulva situated di-
rectly at anus; eggs pitted 5.

5. Middle of body, at least in adult female, strongly swollen.

Hystrichis tricolor, p. 376.
Entire body, even that of adult female, very slender.

Hystrichis neglectus, p. 379.

6. Spines, especially those of head, but also to a lesser extent those of cervical

region, bent or crooked Hystrichis varispinosus, p. 387.

Spines, as far as can be told from incomplete descriptions, straight 7.

7. Female 100 mm. long by 3 mm. wide ; 17 spines in each circle on head ;

from Fulica atra Hystrichis wedli, p. 381.

Female 25 to 44 mm. long by 500/* to 1 mm. wide ; 23 spines in each circle
on head ; from Cygnus olor and litis faleincllus.

Hystrichis orispinus, p. 381.

The above key is a translation, with several modifications, of one
given by Jaegerskiold.

HYSTRICHIS TRICOLOR Dujardin, 1845

Synonyms. — Spiroptera tricolor (Dujardin, 1845) Diesing, 1851;
Spiroptera tadornae Bellingham, 1844.

Hosts. — Primary: Anas boschas domestica, A. b. fera, A. tadorna,
Tadorna bellonii, T. tadorna; secondary : Probably fish, according to
Jaegerskiold.

Location. — Glands of proventriculus.

Morphology. — Hystrichis (p. 375) : Filiform worms, having the
anterior extremity of the body (fig. 437) armed with spines directed
posteriorly and arranged in 40 to 42 rows; on the head, where they
are the thickest, the spines number as high as 50 to 55 in each circle;
the largest spines are 40 to 50/a long. The exterior of the body is
white, the intestine black, and the intermediate portion and esopha-
geal region red. The mouth is round and somewhat protractile,
and there are six small buccal papillae.

Male 25 mm. long.

NEMATODE PARASITES OF BIRDS

377

Female 27 10 40 mm. long by 350 to 500/* wide. The vulva is at
the posterior end, just anterior to the anus (fig. 437). Eggs 85 to
88/* long by 36 to 40/* wide, oblong, somewhat truncated at the ex-
tremities, covered with slightly salient granules or tubercles, com-
paratively large and thickly set.

Life history. — Unknown. Apparently the worms undergo at least
part of their molts in the tissues of the host and may die there, leav-
ing their borrows filled with eggs as the worms themselves decom-
pose or are absorbed. This habit is somewhat similar to that of
Hepaticola hepatica, a hairworm in the liver of rats and not a re-
mote relative of Hystrichis. There are probably intermediate larval
stages in fish, according to Jaegerskiold.

Distribution. — Europe (France, Germany, Italy, Hungary and
Ireland).

Fig. 437. — Hystrichis tricolor. Head and tail ends of female. After

Jaegerskiold, 1909

HYSTRICHIS ACANTHOCEPIIALICUS Molin, 18Gla

Synonym. — Strongylus tubifex tantali. 11

Hosts. — Primary : Ibis nudifrons, Phimosus infuscatus; second-
ary : Probably fish, according to Jaegerskiold. Ibis tubifex, re-
ported as a host in the index-catalogue of Stiles and Hassall, is a
lapsus, the generic name of the host (Ibis nudifrom) having been
combined with the specific name from the synonym.

Location. — Glands of proventriculus.

Morphology. — Hystrichis (p. 375) : Swelling of head end (fig.
438 a and b) slight or lacking. Head with 7 to 9 circles of spines, the
spines relatively sparce, not closely set, the largest 25 to 30/* long by
15 to 17/* wide. Mouth opening triangular; mouth cavity 130/* long.

Male 23 to 15 mm. long; maximum width 1.5 to 2 mm., width at
head end 270 to 200/*, at tail end (just anterior to bursal cup) 1G0/*.
Spines on anterior part of body to a distance of 800/i from head end.
Esophagus about 8 mm. long. Bursal cup 100/* long by 240/x wide,
a round bell-shape, its walls thick, muscular (fig. 438 c and d).

Female 35 to 47 mm. long; maximum width 2 to 3 mm., width at
head end 300 to 3G0/*, at tail end 320 to 350/*. Spines on anterior part

11 Catalogue of the Vienna Museum.

378

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

of body for a distance of 1.1 mm. from head end. Esophagus 9.6
mm. long. Vulva 175/z from posterior end (fig. 438e). Eggs 75
to 79jm long by 40 to 44/x wide; shells not pitted, as in other species,
but with an irregular network of ridges (fig. 438/).

Life history.— Unknown ; see H. tricolor, p. 376.

Distribution. — South America (Brazil).

HYSTRICHIS CORONATUS Molin, 1861

Synonyms. — Hystrichis, species Molin, 1860; H. mergi-mergan-
sei^is Diesing, 1861.

Host. — Primary: Mergus merganser; secondary: Unknown, prob-
ably fish, according to Jaegerskiold.

Location. — Glands of proventriculus.

Fig. 438. — Hystrichis acanthocephalicus. a, Side view ; b, front view of

HEAD; C AND d, MALE TAIL; e, FEMALE TAIL; f, EGG. AFTER JAEGERSKIOLD,

1909

Morphology. — Hystrichis (p. 375) : Head (fig. 439a) not set off
from body. Only one circle of spines; it is directly posterior to
mouth and, according to figure, made up of 18 large spines. Pos-
terior region of body with coarse cross-striations or annulations.

Male unknown.

Female 27 mm. long ; maximum width 3 mm. According to figure
(fig. 4396), body considerably swollen in middle region. Vulva near
to anus, at the rounded posterior end of body (fig. 439c).

Life history. — Unknown; see H. tricolor, p. 376.

Distribution. — Europe (Italy (Padua)).

HYSTRICHIS CYGNI (Molin, 1858) Diesing. 1861

Synonyms. — Echinocephalus cygni Molin, 1858 ; Hystrichis pachi-
cephalus Molin, 1861a.

NEMATODE PARASITES OF BIRDS

379

Hosts. — Primary: Cygnus olor; secondary: Probably fish, ac-
cording to Jaegerskiold.

Location. — In vesicles between the external coats of the proven-
triculus.

Morphology. — Hystrichis (p. 375) : Body irregularly swollen in the
middle and posterior portions. The head end (fig. 440) also has a
marked bulbous swelling which is wider than long and provided
with 20 rows of large spines, directed posteriorly and thickened at
the base. In addition the anterior portion of the body is armed with
small spines which gradually disappear posteriorly. The circular
mouth is protractile and surrounded by a crown of small spines.

Male unknown.

£'','.' "'•'.'.',v3 :■■$

a

Fig. 439. — Hystrichis COBONATUs. a, Head end; b, female, natural size;

C, FEMALES TAIL. AFTER MoLIN, 1SG0

Female 30 mm. long. Anus terminal, large; vulva at posterior
end of body, near anus.

Life history. — Unknown; possibly similar to that of H. tricolor
(p. 376).

Distribution. — Europe (Italy) .

HYSTRICHIS KEGL&CTUS Jaegerskiold. 1909

Synonyms. — Eustrongylus papillosum (Rudolphi, 1802) Diesing,
1851, part; Hystrichis papillosus (Rudolphi, 1802) Molin, 1861a,
part.

In addition to the above, Jagerskiold lists Hystrichis, species Wedl
as a possible synonym and states that it is very close and possibly
identical with his new species. HystricMs, species Wedl, however,
was named //. wedlii by Linstow, emended by Jaegerskiold to //.
wedli and if the two species, H. neglectus and //., species Wedl are
identical, Jaegerskiold's species will fall as a synonym of //. wedli.
The fact that the larva described by Linstow at the same time that

380

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

he named Wedl's species may not be identical with it, as he thought
(see H. wedli, p. 381) does not alter the status of H. wedli as that is
the name given to Wedl's species.

Hosts. — Primary : Numenius arquatus, Querquedula circia; sec-
ondary: Probably fish, according to Jaegerskiold.

Location. — Esophagus.

Morphology. — Hystriohis (p. 375) : Body long and slender. Mouth
opening small, triangular, surrounded by 6 small papillae, wart-
shaped with small points. Head (fig. 441a) with closely-set spines,
larger than those found on anterior cervical region. Largest spines

ng$m

Figs. 440-441. — 440, Hystbichis cygni, Head end. After Molin, 1861. 441,
Hystbichis neglectus. a, Head end ; b, male tail ; c, female tail ; d, egg.
Aftek Jaegerskiold, 1909

40 to 50/a long by 20/* wide. Spines extend along body for distance
of 1.5 mm.

M ale specimens imperfect; length more than 33 mm. Bursal cup
(fig. 4415) not prominent, scarcely wider than the portion of body
preceding it, of thick muscle, the hollow inner part shallow. An-
terior to this on each side of body a row of papillalike structures,
12 or more in number.

Female about 111 mm. long; maximum width 1.5 mm., width at
head end 450 to 500^, at tail end (fig. 441<?) 350/x. Each circle of
the head contains 55 to 57 spines. Esophagus 8.25 mm. long. Eggs
79 to 84/a long by 42 to 44/x wide, with large but not closely-set pit-
tings (fig. 441a*). Shell about 4.5^, thick; poles rounded.

NEMATODE PARASITES OF BIRDS 381

Life history. — Unknown; possibly similar to that of H. tricolor,
p. 376.

Distribution. — Europe (Italy (Museum, Genoa and Cagliari) and
Austria (Museum, Vienna)).

HYSTRICHIS ORISPINUS Molin, 1858

Hosts. — Primary: Cygnus olor, Ibis falcinellus; secondary: Un-
known, probably fish, according to Jaegerskiold.

Location. — Glands of pro vent riculus.

Morphology. — Hystrichis (p. 375) : Head (fig. 442) distinctly
though gradually marked off from cervical region. Mouth at the
apex of a rectractile cone, large, with 4 short spines set crosswise.
Head end with closely-set straight triangular spines, 23 spines in
a circle, gradually becoming larger and more sparse posterior to
cervical region.

Male unknown.

Female 25 to 44 mm. long, 500y to 1 mm. wide. Posterior part of
body without spines but with dense transverse annulations. Anus
terminal. Vulva just slightly anterior to anus. Eggs not described.

Life history. — Unknown; possibly similar to that of H. tricolor,
p. 376.

Distribution. — Europe (Italy (Padua)).

HYSTRICHIS VARISPINOSUS Jaegerskiold, 1909

Most. — Primary : Mergus serrator; secondary: Probably fish, ac-
cording to Jaegerskiold.

Location. — Not given; probably proventriculus.

Morphology. — Hystrichis (p. 375) : Head (fig. 443a) only slightly
swollen, covered with 15 circles of small, closely-set strongly-bent
spines, the smallest of which are 27/* long, the largest 48/x long;
the highest number of spines in a circle is 52. Spines (fig. 443&)
of cervical region 88 to dO/x long, larger and less closely-set than
those of head; the highest number in a circle is 14 to 17, becoming
more scarce posteriorly and disappearing at a distance 7.2 mm. from
the head end.

Specimen incomplete, sex undetermined, only the anterior region,
11 mm. long, present; maximum width 800/a, the width of head 400/*.
Esophagus 6 mm. long.

Life history. — Unknown; possibly similar to that of II. tricolor,
p. 376.

Distrib utiori. — Not given.

HYSTRICHIS WEDLI Linstow, 1879 emended Jaegerskiold. 1909

Synonyms. — Hystrichis, species Wedl, 1856; H. wwdlii Linstow,
187i)«. //. neglectus is possibly a synonym of this (see page 379).

382

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Host. — Primary : Fulica atra; secondary : Probably fish, according
to Jaegerskiold.

Location. — Esophagus.

Morphology. — Hystrichis (p. 375) : Head a knob-like thickening
with rounded summit surrounded by a horny border, with a crown
of conical, straight smooth spines, 17 to 18 in each row. According
to Wedl T the spines disappear 3 mm. from the head end.

Male unknown.

Female 100 mm. long by 3 mm. wide.

mm

Km

I

W/Mf'/H
7/iffi/r

tinum i
[nrwj:

Figs 442-444. — 442, Hystrichis orispinus. Head end. From Jaeger-
skiold, 1909, after Molin. 443, Hystrichis varispinosus. a, Head

END ; b, SPINE OF ANTERIOR CERVICAL REGION. AFTER JaEGEIRSKIOLD,

1909. 444, Hystrichis wbdli (Linstow's material), a, Anterior
portion of body ; b, one of the most anterior spines, lateral view ;

C, SAME, FRONT VIEW; (I, ONE OF THE MOST POSTERIOR SPINES; €, POSTERIOR
END OF BODY (X, PROBABLE PRIMORDIUM OF VAGINA). AFTER JAEGERSKIOLD,
1909

Larva (found by Linstow) 24 mm. long by GOOju. wide; anus
terminal; head and tail ends rounded; esophagus 1/4 of body length.
Cuticle thick, made up of 3 layers, with transverse striations, with
posteriorly directed conical spines 69^. long by 29/x wide at the base,
very thick at head end (fig. 444 a to d), smaller and less numerous
posteriorly but extending to tail end of body (fig. 444e). Outer
cuticle about to be cast off, the head end showing new structure as
described by Wedl for adult.

Life history. — Unknown; possibly similar to that of H. tricolor,
p. 376.

Distribution. — Not given.

NEMATODE PARASITES OF BIRDS 383

Jaegerskiold considers it doubtful that the larva which Linstow
had was the same species as the adult described by Wedl. He says
that in several respects Linstow's form resembles H. varispirwsus
more closely than it does Wedl's species.

ADDENDA

Since the present paper has been in press there have appeared
several notable contributions to the study of nematodes parasitic in
birds, of which brief mention should be included here.

The following new species or redescriptions of former species have
been published :

AMIDOSTOMUM SKRJABINI Bouleneer. 1926

Boulenger 12 described as a new species an amidostome found in
Anser albifrons in Egypt. This species appears to be identical with
Amidostomum chevreuxi Seurat, 1918, Boulenger apparently over-
looking Seurat's species. The only differences to be noted in the
two descriptions refer to the swelling at the posterior end of the
esophagus; Seurat says there is such a swelling or bulb, which,
however, is of the same width as the part of the esophagus anterior
to it, and thus is not differentiated externally; in A. skrjabini, Bou-
lenger says that the swelling is absent.

Genus PSEUDAMIDOSTOMUM Boulenger, 1926

Boulenger 13 has made a new genus based on three female speci-
mens of nematodes found along with specimens of Amidostomum
raillieti in Fulica atra, the head structures of these three nematodes
being especially different from those in Amidostomum. Boulenger
gave no diagnosis for his new genus but, based on the type species,
the generic diagnosis may be given as follows: Head with cuticle
slightly expanded in form of mouth collar and with four conspicu-
ous submedian papillae. Buccal capsule broad and very short, with-
out teeth. Vulva posterior to middle of body; diameter of body
sharply diminished posterior to anus.

Type species. — Pseudamidostomum loosi Boulenger, 1926.

PSEUDAMIDOSTOMUM LOOSI Bouleneer, 1926

Boulenger 14 describes this species from stomach wall of Fulica
atra; Egypt.

Females only, 7.3 to 7.8 mm. long by 16(V to 17G> wide. Cuticle
of head slightly expanded; four conspicuous submedian papillae;

15 1026 : Parasitology, Cambridge, Eng., vol. 18, pp. 94-06, figs. 18-22.

13 Idem, pp. 05-97.

M Idem, p. 06, figs. 23-25.

384 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

mouth opening small, circular. Buccal capsule broad (12//,) and very
short, without teeth. Esophagus 800/* to 930/* long. Anus 70/* to
110/* from posterior end; body narrows suddenly behind anus, as in
Epomidiostomum. Vulva 1.7 to 1.75 mm. from posterior end. Com-
bined length of ovejectors (including sphincters) about 325/*; uteri
divergent. Eggs 57/* to 69//. long by 37// to 42// wide.

EPOMIDIOSTOMUM QUERQUETULAE Boulenger, 1926

Boulenger 15 also describes this species from stomach wall of Quer-
quedula crecca {=Nettion crecca) ; Egypt.

Morphology. — E pamidiostomum (p. 26) : Body slender, of yellow-
ish color. Head very small, provided laterally with a pair of cuticu-
lar expansions, corresponding to, though less developed than, the
"epaulettes" of the type species (E. unclnatum) . Mouth with
six (four submedian, two lateral) papillae. Mouth cavity funnel
shaped; esophagus elongated, club shaped. Cuticle with conspicu-
ous cross striations, highly developed in cervical region into folds.
Male 8.8 to 10.5 mm. long by 180// to 200// in diameter. Head only
35// in diameter; esophagus, 1 to 1.05 mm. long. Bursa trilobed;
dorsal lobe very small. Dorsal ray divided at distal end into four
very small branches. Externo-dorsal rays short and thick; lateral
rays of approximately equal size. Genital cone well developed and
provided with a pair of conspicuous papillae at its apex, as in type
species. Spicules 180//, to 200// long, the distal portion divided into
two unequal branches. Gubernaculum absent. Female 13 to 14.5
mm. long by 220//. wide. Head 38// in diameter; esophagus 1 to
1.1 mm. long. Anus 130// from posterior end of body; diameter of
body sharply diminished posterior to anus. Vulva a transverse slit
on slight elevation, 2.7 to 3.2 mm. from posterior end. Ovejector,
including sphincter, 460/* long. Eggs 80// to 90/* long by 50// wide.

This species, as described by Boulenger, is very similar to Epo-
midiostomum orispinurn from the same host, tc which species Bou-
lenger makes no reference. Since, however, the descriptions differ
in the two cases with reference to the number of branches of the
spicules and with reference to the head, and since there is no illus-
tration of the head of E. orispinurn to compare with that of E. quer-
quetulae, the latter must be considered a distinct species.

EPOMIDIOSTOMUM SKRJABINI Petrow, 1926

Petrow 1G reports this species from Anser anser domestieus and
Anser albifrons; Don district, Russia.

Morphology. — Epomidiostomum (p. 26) : Male 9.5 to 11.5 mm.
long by 210/* to 240// wide. Esophagus 1.02 mm. long. Spicules

15 1926 : Parasitology, vol. 18, pp. 91, 92, figs. 8-10.

16 1926 : Trudy gossudarstvennogo instituta eksperimentalnoi veterinaril ; Moskow, vol.
3 (1), pp. 103-105; 4 figs.

NEMATODE PARASITES OF BIRDS 385

206/* to 210// long, dividing into three branches distally. General
shape of bursa similar to that of E. uncinatum, with two large
lateral lobes and a small dorsal lobe; the dorsal ray, however, is
pictured as dividing only into two branches, not four as in other
species. Bursa strongly striated, a character described for E. orl-
spinum also. Female 15 to 17 mm. long by 270//, to 290//, wide.
Esophagus 1.29 mm. long. Anus 156// to 175/i, vulva 3.32 mm., from
posterior end of body. Eggs 101/* to 105/t long by 58// to 62/x wide.
The measurements given above for this species correspond very
closely to those of E. orispinum (p. 28), of which species Pet row
makes no mention; the hosts of the two species are identical also.
However, the present writer has not translated all of the Russian
text, and since there are differences in the characters of the heads
of the two species (see key, p. 27) and differences in the distal ter-
mination of the dorsal ray, the two species must be left distinct for
the present.

TETRAMERES (TETRAMERES) ZAKKAROWI Pctrow, 1926

This species Petrow 17 also describes from Anser albifrons; Don
district, Russia. The present writer has translated the Russian text
only to the extent of securing the following brief description : Male
onty, 9.47 mm. long by 170//. wide. Body armed with posteriorly
directed spines throughout its whole length ; additional caudal spines
present. Spicules slender and very unequal, the one 1.02 mm. long,
the other 195// long.

Chandler 1S has described four new species of Heterakis, two new
species of Pseudaspidodera* and a new subulurid.

HETERAKIS HASTATA Chandler, 1926

Chandler 19 reports this species from cecum and large intestine of
Lophura rufti; Zoological Gardens, Calcutta, India.

Morphology. — IJeterakis (p. 50) : Large stout worms. Esophagus
1/7, or a little less, of body length. Lateral alae well developed, in
female 80// wide, in male a little less; they extend about 2/3 the
length of the body. Lips prominent, papillae smaller than in H.
lanei. Mcde 10.5 to 11.5 mm. long by 375/x wide. Esophagus 1.5
mm. long by 80// wide (narrow part) ; bulb 385// long by 240// wide.
Tail 550/x long; ventral surface roughened with numerous small
granulations, which obscure the papillae. Papillae said to be of
typical number and arrangement, the second lateral paracloacal
being the largest. Sucker, 150// anterior to cloacal aperture, large,

17 1926 : Trudy gossudarstvennogo instituta experimentalnoi veterinarii ; Moskow, vol.
3 (1), p. 108, 2 figs.

18 Indian J. M. Res., Calcutta, vol. 13 (3), pp. 617-624.
18 1926 : Idem, p. 619, fig. 3.

386 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

its antero-postero diameter 145/*. Spicules slightly unequal, the
left 1.75 mm. long, with broad alae, which are slightly asymmetrical
near tip. Eight spicule, 1.65 mm. long, with narrow alae. Female
12 to 13 mm. long by 420/* to 460/* wide. Esophagus 1.6 mm. long;
bulb 420/* long by 270/* wide. Vulva approximately at middle of
body. Anus 1.2 to 1.3 mm. from posterior end. Eggs 70/* by 40/*.

HETERAKIS LANEI Chandler, 192G

Chandler 20 describes this second new species also from the cecum
and large intestine of Lophura rufa; Zoological Gardens, Calcutta,,
India.

Morphology. — Heterakis (p. 50) : Small, slender worms. Esoph-
agus 1/7 to 1/8 of body length. Lips of moderate size, with
unusually conspicuous papillae. Lateral alae narrow. Male 9 to
9.6 mm. by 290/* wide. Esophagus 1.25 mm. long by 75/* wide, in
front of bulb, which is 300/* long by 190/* wide. Tail 400/* to 45G>
long; caudal alae narrow. Caudal papillae said to be normal in
number, but the third and fourth lateral paracloacals (=adanals)
have common root and are large and coarse, the first and second
smaller. Ventral paracloacals small and inconspicuous. Sucker
large, 140/* in antero-postero diameter; its posterior border about
175/* anterior to cloacal aperture. Spicules almost equal but of
different shape, the left spicule 2 to 2,2 mm. long, alate; the right
spicule nonalate, about 2 mm. long. Female 9 to 10 mm. long by
280/* to 300/* wide. Esophagus 1.15 to 1.4 mm. long; bulb 360/* long
by 200/* wide. Vulva very slightly anterior to middle of body;
pre- and postvulvar protuberances present. Tail about 950/* long.
Eggs 65/* to 68/* by 40/*.

HETERAKIS VARIABILIS Chandler, 1926

Chandler 21 reports the discovery of this species in the cecum and
large intestine of Poly plectrum bicalcaratum ; Zoological Gardens,
Calcutta, India.

Morphology. — Heterakis (p. 50) : Rather large and slender worms.
Esophagus 1/8 to 1/7 of body length. Lips large, papillae not prom-
inent. Lateral alae very narrow along head, suddenly widening
about 150/* from anterior end to a width of 40/*; gradually narrow
posteriorly, extending 2/3 of body length in female, but disappearing
anterior to middle of body in male. Mais 9 to 10 mm. long by 310/*
wide. Esophagus 1.5 mm. long; bulb 320/* long by 230/* wide, Alate
tail end narrow and long, 480/* long by 220/* wide. Caudal papillae
said to be normal in number; the first lateral paracloacals (adanals)

20 1926: Indian J. M. Ros. Calcutta, vol. 13 (3). p. 618, figs. 1 and 2.
21 1926: Idem., pp. 619, 620, fiff. 4.

NEMATODE PAEASITES OF BIRDS 387

are shifted forward and are asymmetrical both in position and devel-
opment, the right being a single stout papilla, the left slit into two
slender papillae. Sucker 85/* in diameter, its posterior edge 180/*
anterior to anus. Tail about 400/* long. Spicules unequal, the right
720/* to 880/*, the left 1.04 to 1.18 mm. long, similar in form and
narrowly alate. Female 11.5 to 12.5 mm. long by 385/* wide. Esoph-
agus 1.6 to 1.05 mm. long; bulb 300//. long by 200/* wide. Vulva pos-
terior to middle of body, dividing body length in ratio of 0:5 or 5.5.
Posterior third of tail tapers rapidly. Eggs 00/* to 05/* by 38/* to 40/*.

HETERAKIS VULVOLABITA Chandler, 1926

Chandler -- describes this species from cecum and large intestine of
Arborlcola torqueola; Zoological Gardens, Calcutta, India.

Morphology. — Ilete'rakis (p. 50) : Small, stout worms. Esophagus
1/8 to 1/7 of body length. Lateral alae narrow. Male to 7 mm.
long by 250/* wide. Esophagus about 870/* long; bulb 210/* long by
105/* wide. Tail 230/* long, the slender part posterior to papillae
only 70/* long. Sucker small, 50/* in diameter, situated 90/* anterior
to cloacal aperture. Alate tail end short, 330/* long by 145/* wide.
An extra pair of caudal papillae present, apparently due to the
splitting of the second lateral paracloacal (adanal). Spicules un-
equal and dissimilar, the left 535/* long and very slender (3/* to 9/*
wide), the right 290/* to 300/* long by 18/* wide. Female 7 to 8 mm.
long by 205/* wide. Esophagus 875/* long; bulb 235/* long by 105/*
wide. Vulva prominent, situated well behind the middle of the
body, dividing the body length in a ratio of 7 : 5.5. Ovejector makes
a prominent loop near vulva. Tail about 540/* long. Eggs small,
55/* to 00/* by 35/*.

PSEUDASPIDODERA VOLUPTUOSUS Chandler, 1926

Chandler 23 describes this species from the cecum and large intes-
tine of Argitsianus argus; Zoological Gardens, Calcutta, India.

Morphology. — Pseudaspidodera (p. 102) : Small, stout worms.
Lateral alae narrow, 25/* wide, originating 150/* behind the head,
extending about 3/5 of the body length in the female, about 3/4 in
the male. Subcuticular cephalic cordons conspicuous. Esophagus
1/7 to 1/0 of body length. Male 0.25 to 9.25 mm. long by 325/* to
380/* wide; head about 100/* broad. Esophagus 1.3 mm. long; bulb
300/* long by 250/* wide. Alae of tail well developed, their length
520/*, total width 270/*. Tail 300/* to 390/* long. Sucker, 100/* in
diameter, is 125/* to 100/* anterior to the cloacal aperture. Spicules

»1926: Indian J. M. Res. Calcutta, vol. 3 (3), pp. G20, 621, figs. 5-8.
23 1!)26 : Idem., pp. 621, 622, figs. 9, 10.

388 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

unequal, the right 1.44 to 1.7 mm. long, the left always almost ex-
actly twice that length, 2.7 to 3.45 mm. long. Slender part of tail
posterior to papillae 200/* long. Female 9.35 to 10.15 mm. long by
460/* wide. Esophagus 1.6 to 1.7 mm. long; bulb 320/* long by 250*t
wide. Vulva well behind the middle of body, dividing body length
in ratio of 6:4. Sometimes several postvulvar papillae present.
Tail slender, 1.05 to 1.25 mm. long. Eggs oblong, 60/* to 68/* by 35//.
to 38/*.

PSEUDASPIDODERA VOLUPTUOSUS MINOR Chandler, 1926

This species is reported by Chandler 2i as having been found in
large intestine and cecum (?) of Rollulus roulroul; Zoological Gar-
dens, Calcutta, India.

Morphology. — Pseudaspidodera (p. 102) : Cephalic cordons less
conspicuous than in the typical form but of same shape and ar-
rangement. Male 5 to 5.5 mm. long by 225/* to 250/* wide. Esopha-
gus 1/5 of body length. Spicules not reduced in size as is the body,
but are of almost the same length as in the typical form, the right
being about 1.5 mm., the left 3 to 3.4 mm. long. Tail relatively
longer, 360/* to 400/*. Female 6.75 to 7.85 mm. long by 310/* wide.
Esophagus a little less than 1/5 of body length. Vulva a little
farther forward than in typical form, dividing body length in ratio
of about 6 : 5, or even more nearly in half. Tail about 1 mm. long.

SUBULURA MULTIPAPILLATA (Chandler, 1926) Cram, 1927

/Synonym. — Allodapa multipapiUata Chandler, 1926. 25 In cecum
and large intestine of Rollulus roulroul; Zoological Gardens, Cal-
cutta, India.

Morphology. — Subulura (p. 104): Small, slender worms; cervical
alae short and narrow. Buccal cavity in two portions, the anterior
with very thick walls; three small teeth at base of cavity. Entire
esophagus 1.25 mm. long; posterior bulb about 240/* long by 250/*
wide, in female. Male 6 to 7 mm. long by 260/* to 280/* wide.
Esophagus 1.12 mm. long. Tail 230/* long. Sucker 430/* anterior
to cloacal aperture, poorly developed. Fifteen pairs of papillae,
of which seven are preanal, eight postanal. The most anterior of
the papillae are well anterior to the sucker and 590/* anterior to the
cloacal aperture; the preanal papillae are all rather ventral; of the
postanals, six pairs are ventral, two lateral. Spicules similar and
equal, 780/* long and 22/* wide. Gubernaculum in two parts, a nar-
row stout piece 145/* long, dorsal to the spicules, and a flat piece
170/* long ventral to the spicules. Female 8 to 10 mm. long by

^1926: Indian J. M. Res. Calcutta, vol. 3 (3), p. 622, figs. 11, 12.
25 Idem., pp. 622, 623, figs. 13-15.

NEMATODE PARASITES OF BIRDS 389

350,1* wide. Vulva inconspicuous, anterior to middle of body, divid-
ing latter in ratio of about 3 : 5. Ovejeetor directed forward. Tail
about 2 mm. long. Eggs 56/* to 63/* by 34/* to 42/*.

CHEILOSPIRURA SKRJABINI (Ozerska, 1926) Cram, 1927

Synonym. — Acuaria (Oheilospirura) skrjabini Ozerska, 1926. 20
From the house sparrow ; Don district, Russia.

Morphology. — Oheilospirura (p. 226) : Male 8.7 to 9.2 mm. long
by 140/* to 190/c wide. Pharynx 160/* to 194/* long; first part of
esophagus 399/* to 456/* long; second part of esophagus 930/* to 1.08
mm. long. Cordons 250/* to 285/* long. Spicules unequal, the one
205/* to 239/* long, the other 125/* to 148/* long. Cloacal aperture
200/* from posterior end. Ten pairs of caudal papillae, of which
four are preanal and six postanal; of the postanal, the distance
between pairs 3 and 4 is slightly greater than between the other pairs,
the spacing between the others being almost equal. Female 23 to
27 mm. long by 130/* to 200/* wide. Pharynx 182/* to 228/* long;
first part of esophagus 593/* to 707/* long; second part of esophagus
1.25 to 1.7 mm. long. Cordons 285/* to 387/* long. Vulva 11.11
to 13.11 mm. from head end of worm; anus 170/* to 250/* from
posterior end. Eggs 43/* to 48/* by 28/* to 29/*.

CHEILOSPIRURA PAVONIS (Ortlepp, 1925) Cram, 1927

Synonym. — Acuaria (Cheilospirwa) pavonis Ortlepp, 1925. 27
From under lining of gizzard of Pavo muticus; locality not given.

Morphology. — Cheilosplrura (p. 226) : Head with two lateral tri-
angular lips, each with a large blunt tooth; no external papillae
noted. Cuticle with coarse cross striations. Cordons in their pos-
terior parts appear corrugated, as if made up of a series of elongated
bosses. Total length of esophagus about 1/3.2 of body length.
Male 6.5 mm. long by 180/* wide. Cordons 4.8 mm. long. Pharynx
157/* long by 30/* wide; muscular part of esophagus 654/*. long;
glandular part of esophagus 1.38 mm. long. Posterior end of body
coiled slightly; caudal papillae well developed; ventral surface with
coarse cross striations. Ten pairs of caudal papillae, of which four
pairs are preanal, six pairs postanal. Spicules unequal and dissimi-
lar, the left 1.56 mm. long by 16/* wide, with chisel-like tip; the right
215/* long and very broad. Gubernaculum absent. Female 14.7
mm. long by 358/* wide. Pharynx 305/* long by 60/* wide ; muscular
part of esophagus 1.14 mm. long; glandular part 3.25 mm. long.
Dorso-lateral cordons extend slightly posterior to vulva; ventro-

28 Trudy Gossud. Inst. Eksper, Vet., Moskow, vol. 2 (2), pp. 103, 104, figs. 1. 2.
"Journ. Helminth., London, vol. 3, pp. 177-179, 3 figs.

390 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

lateral cordons terminate at level of vulva. Vulva 8.4 mm. from
anterior end, dividing body length in ratio of 3:2. Two divergent
uteri. Anus 407/* from tail end. Eggs not mature in specimens
described.

CYRNEA BULBOSA (Linstow, 190G) Ortlepp, 1925

Ortlepp 28 gives the following redescription of Linstow's species,
transferring it from Physaloptera (see p. 310) to Cyrnea : Mouth with
two lateral lips, each of which is subdivided into a large median and
two smaller lateral lobes. Cervical papillae small, spikelike, situated
just anterior to or at level of nerve ring. Mouth cavity well devel-
oped. Esophagus made up of anterior, muscular and posterior,
glandular parts, the anterior being about 1/10 of the total length
of the esophagus. Male 10.4 mm. long by 325/* wide. Esophagus
just less than 1/10 of total body length. Right spicule 945/* long;
left spicule 2.45 mm. long. Immediately anterior to cloacal aperture
the ventral surface of body covered with irregular tubercles. Caudal
papillae as in original description except that Ortlepp says the
adanal papillae are pedunculated, rather than sessile, and the group
of papillae near the tail end consists of four pairs, rather than five
pairs. FemaU 24 mm. long by 455/* wide. Tail length 1/129 of total
body length; vulva 1.25 mm. from tail end, dividing body in ratio
of 18 : 1. Vagina 546/* long by 137/* wide at base, from which laterally
arises the thick-walled ovejector, 590/* in length, from which the two
uteri branch off and run parallel anteriorly.

GONGYLONEMA, new species?

Synonym. — Gongylonema ingluvicola 1 of Smit and Notosoediro,
1926. 29 From the crop of the chicken ; Java ( ? ) .

Morphology. — Gongylonema (p. 203) : Cuticular bosses extend for a
distance of about 500/* from the head end. Mouth cavity long (34/*)
and slender (7/*). Male 18 to 19 mm. long by 215/* wide. Caudal
papillae asymmetrical, on the left side four preanal, one adanal,
and five postanal ; on the right side five preanal, one adanal, and four
postanal. Spicules very unequal, the left spicule long (10.5 mm.),
slender and sharply pointed ; the right spicule short and thick, meas-
uring 580/*. Female 40 to 56 mm. long by 330/* wide. Anus 332/*
from tail end. Vulva 2.5 mm. anterior to anus. Eggs 55/* by 35/*.

The spicule lengths of Gongylonema ingluvicola Ransom being
very different from the above, Smit and Notosoediro are apparently
dealing with a new species.

28 1925: Journ. Helminth., London, vol. 3, p. ISO. 4 figs.

30 Nedrl.-Iud. Blad. v. Diergeueesk., Buitenzorg, vol. 38, pp. 92-94, 7 figs.

NEMATODE PAKASITES OF BIRDS 391

DISPHARYNX, new species?

Synonym. — Dispharagus new species ? of Smit and Notosoediro,
1926. 30 From the gizzard of the Java cock.

Morphology. — Dispharynx (p. 237) : Mouth with two lips. Mouth
cavity becomes narrower at base. Mouth cavity 100/x to 116/a deep;
length of esophagus 750/*, (this the length of only the first part of
the esophagus ? In the illustration the cordons extend about 2/3
the length of the anterior or muscular esophagus). Cordons extend
posteriorly for a distance of 500^, then recurve and extend anteriorly
almost to the mouth. Male 5 mm. long. Eight pairs of caudal
papillae, of which four are preanal, four postanal. Spicules unequal,
the left 335/* long and thin, the right short (120/1, long) and thick.
Female 6 mm. long. Anus 166/t from posterior end of body ; vulva
1.22 mm. anterior to anus. Eggs 37/a by 20/*.

This is apparently a new species of Dispharynx ,' it resembles most
closely Dispharynx rectovaginata in size and in having only four
pairs of postanal papillae, but in D. rectovaginata the left spicule is
twice as long as the right, in this species more than that. The hosts
and distribution in the two cases are also very different.

Yorke and Maplestone have recently published a volume 31 which
will prove of inestimable aid in systematic work with nematodes.
This comprehensive study gives diagnoses and keys from orders
down to genera, with text figures illustrating one species, usually the
type species, of each genus. The classification used by these authors
differs in certain respects from that followed in the present study of
bird nematodes. Yorke and Maplestone have not employed the
orders Myosyringata and Trichosyringata, but have used Eunema-
toda Ward, 1916, and Gordiacea Siebold, 1848, as the orders; the
superfamilies within each order are then considered, suborders not
having been employed as they have been in the present study. Yorke
and Maplestone have made a new family Subuluridae, whereas I have
followed Travassos in using the subfamily Subulurinae in the family
Heterakidae. They have put Heterakidae and Subuluridae in the
Oxyuroidea, whereas I considered them as Ascaroidea, as did Hall
(1916). In Yorke and Maplestone's classification Yseria has been
made a synonym of fitreptocara, and this genus put in the Physa-
lopterinae; I have put Yseria in Schistorophinae, Gedoelst having
indicated that it is closely related to HistiocephaJus, and have not
considered its festoons as analogous to cordons, whereas the denticu-
lated collar of Streptocara is considered an analogous structure and
this genus accordingly allocated to Acuariinae. Yorke and Maple-
stone list only Yseria califomica, the second species which I included

80 Nedrl.-Ind. Blad. v. Diergeneesk., Buitonzorp. vol. 38, pp. 95-07. 4 figs.

81 102C : The Nematode Parasites of Vertebrates. 536 pp., 307 fi^s. London.

392 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

in that genus, Yseria coronata, having been left by them in Histio-
cephalus, which was Skrjabin's tentative assignment. They have
put Schistorophinae in the family Ancyracanthidae ; I have placed
it in Acuariidae, as Travassos had done formerly. A new subfamily,
Ancyracanthinae, has been made for Ancyracanthus by Yorke and
Maplestone; Ancyracanthopsis they place in Schistorophinae, the
only species listed by them in this genus being A. bilabiata, and the
only species in Ancyracanthus being the type species, A. pvmiatvfidus.
In addition to Ancyracanthopsis, they have included in Schistoro-
phinae the genera Sciadiocara, which I put in Acuariinae, and
Viguiera, which I put in Spirurinae, family Spiruridae. With regard
to the genus Spirura these authors and also Baylis and Daubney,
whose Synopsis of the Families and Genera of Nematoda (1926,
277 pp., London) appeared soon after Yorke and Maplestone's book,
limit the genus to forms having a cuticular boss near the anterior
end of the body. Hall (1916) did not make this a generic character,
nor did Railliet and Henry (1911) when they placed the two species
uncinipenis and zschokkei in this genus, neither of these species pos-
sessing this cuticular boss. Yorke and Maplestone apparently do
not list these two species in any genus. The nematodes appear to
be closely related to Spirura talpae, the type species, and at the
present time it would appear advisable to leave them in that genus,
as I have done, and consider the presence or absence of the cuticular
boss a specific rather than a generic character.

SUMMARY

The present work gathers together the descriptions of nematodes
of the suborders Strongylata, Ascaridata, and Spirurata found in
birds, the great majority of these descriptions until now having been
available only as written in foreign languages, and many of them in
obscure publications to which most workers would not have access.
The purpose of the paper is to facilitate the identification and study
of nematodes parasitic in birds. The author recognizes the two
orders of Ward, the Myosyringata and the Trichosyringata, and the
six suborders as made by various other workers : Ehabdiasata, Oxy-
urata, Strongylata, Ascaridata, Spirurata, and Trichurata. Of these
the first two, the Rhabdiasata and Oxyurata, as used by the present
writer, contain no forms found in birds. The next three suborders,
the Strongylata, Ascaridata, and Spirurata are dealt with in the
present paper. Although the author includes under the Spirurata
the Filarioidea, in order to unite the latter with the other closely-
related superfamily of heteroxenous nematodes, the Spiruroidea, the
present paper does not deal with the Filarioidea or with the sixth

NEMATODE PARASITES OF BIRDS 393

suborder, the Trichurata. A study of these groups is much needed
but was not considered possible in the scope of the present paper.

The author has made several new superfamilies and families in
order to coordinate the groups below them. The nematodes treated
in detail fall into 5 superfamilies; there are approximately 50 genera,
containing a total of about 500 species. About two-thirds of this
material falls in the Spiruroidea and the emphasis has accordingly
been placed on that superfamily. Due to the heteroxenous nature of
the spirurids and the food habits of many birds, which insure the
latter coming in contact with the intermediate hosts, an opportunity
is afforded for a rich variety of parasitic nematodes of this type.
The usual intermediate host of the spirurid is an arthropod; these
arthropods are eaten by insectivorous birds in the case of insects and
by water birds in the case of entomostracans, and a completion of
the life cycle ensured. Birds of prey may possibly derive their
spirurid parasites from rodents or other small animals in which the
larvae have encysted as so-called "aberrant " forms. Doubtless many
insect hosts are eaten incidentally and accidentally by birds which
are not classed as insectivores, when these feed on plant or animal
food in which insects are present, and some of these cases may be
of a nature suitable for the common transmission of parasitic worms.

Diagnoses and keys are given in the present paper to all groups
from species to orders, inclusive. The descriptive material of each
species includes s^ynonyms, hosts, location, morphology, life history
and distribution. There are several hundred illustrations copied
from the publications of other authors. Several species, the data
concerning which were previously incomplete, have been redescribed
and figured by the author ; new names have been given to two species
(in the genera Heterakis and Hartertia) which were misdetermina-
tions by the original describer or have in other ways become con-
fused with other species; various species have been allocated to
genera other than those in which they had previously been placed.
Seven new species have been added ; six of these are spirurids, falling
in the genera Cyrnea, C heilospincra, Hartertia, Echinuria, Tetra-
meres, and Mierotetrameres ; the sixth is a heterakid.

It is hoped that the collecting of the widely scattered material,
published previously in many different countries and many different
languages, after having been translated, systematically arranged and
as far as possible critically analyzed, will prove of value to other
workers in the field of parasitology.

LIST OF HOSTS AND THETR PARASITIC NEMATODES 32

The writer is greatly indebted to Dr. H. C. Oberholser of the Bio-
logical Survey, United States Department of Agriculture, for deter-

32 See also Addenda (pp. 383-392) for recent reports not included in this list.

394

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

mining the scientific names in present usage for the hosts that have
been dealt with in this paper. As mentioned previously, the hosts
have been recorded throughout the text by the names given them at
the time they were reported. In the following host list, the nema-
todes will be listed only under the scientific names which are con-
sidered correct at the present time; other names will be indicated
as synonyms, the correct name indicating the place where the list
of nematodes from that host may be found.

Accipiter bicolor:

Porrocaecum depressant.
Accipiter melanolcucus:

Physaloptera ovata.
Accipiter nisus:

Porrocaecum depressum.

Habronema leptoptera.

Schistorophus bidens.

Synhimantus elliptica.

Synhimantus Jiamata.

Synhimantus laticeps.

Physaloptera alata.

Physaloptera alata chevreuxi.

Physaloptera alata nouveli.

Physaloptera crosi.

Physaloptera megalostoma.

Microtetramcres inermis.
A ccip it er s u percil toys us:

Dispharynx capitata.

Physaloptera acuticauda.
Accipiter firms:

Physaloptera acuticauda.
Aerocephalus arundinaceus:

Porrocaecum ensicaudatum.
Acroeeph a lit s pal ustris :

Acuaria papillifera.
Actitis hypoleuca:

Cosmoeephalus obvelatus.
Actitis macularia:

Cosmoeephalus obvelatus.
Aeyialitis dubia ; see Charadrius

dubius.
Aeyialitis hiaticola ; see Charadrius

hiaticula.
Acgolius brachyotus; see Agio flam-

mcus.
Aegolius otus; see Asio otus.
Aegypius monachus:

Porrocaecum depressum.

Habronema tulostoma.
Afrotis afra afraoides:

Hartertia obesa.

Hartertia rotundata.

Agamia agami:

Porrocaecum serpentulus.
Aithyia ferina-:

Tetrameres fisshpina.
Aix sponsa:

Heterakis caudata.
Ajaia ajaja:

Dispharynx magnilabiata.
Ajaja ajaja; see Ajaia ajaja.
Alauda arvensis:

Physalcptera crassa.
Alauda, species :

Porrocaecum ensicaudatum.
Alea torda:

Contraeaeeum spiculigcrum.

Cosmoeephalus obvelatus.

Streptoeara crassicauda.

Eustrongylides mergorum.
Alcedo americana; see Chloroccryle

amerieana.
Alcedo ispida:

Streptoeara decora.
Alcctoris barbara:

Heterakis tenuicauda.

Subulura sen rati.

Cymea eurycerca.

Hadjclia Ihuillieri.

Dispharynx spiralis.
Alcctoris barbara spatzi:

Hartertia obesa.
Alectoris gracea:

He tera kis ten u ica uda.

Subulura curvata.
Alectoris graeca chukar:

Ascaridia eompar.

Subulura, curvata.
Alectoris graeca saxatilis:

Heterakis gallinae.

Heterakis tenuicauda.

Ascaridia eompar.
Alectoris rufa:

Subulura allodapa.

NEMATODE PARASITES OF BIRDS

395

Alectoris rufa — Continued.

Subulura seurati.

Cheilospirura gruveli.

Cymea eurycerca.

Cyrnea seurati.
Alectura latJiami:

Heterakis bancrofti.

Ascaridia catheturina.
Amazona aestiva:

Ascaridia hermaphrodita.
Amazon® auropalliata:

Habronema incerta.
Amazona barbadensis:

Habronema incerta.
Amazona farinosa:

Ascaridia hermaphrodita.
Amazona farinosa guatemalae:

Habronema incerta.
Amazona f estiva:

Ascaridia hermaphrodita.
Amazona leucocephala:

Ascaridia hermaphrodita.

Habronema incerta.
Amazona ochrocephala:

Ascaridia hermaphrodita.
Amazona vinacea:

Ascaridia hermaphrodita.
Amazona vittata:

Ascaridia hermaphrodita.
Anas acuta; see Daflla acuta.
Anas albifrons; see Anser albifrons.
Anas anser domcsticus; see Anser an-
ser domesticus.
Anas anser fera; see Anser anser.
Anas bosehas; see Anas platyrhyncha.
Anas bosehas domcstica; see Anas

platyrhyncha domcstica.
Anas bosehas fera; see Anas platy-
rhyncha.
Anas canadensis; see Branta cana-
densis.
Anas clangula; see Glaucionetta

clangula.
Anas crecca; see Nettion crecca.
Anas fuligula; see Fuligula fuligula.
Anas fusca; see Melanitta fusca.
Anas glacialis; see Clangula hyemalis.
Anas leucops; see Branta leucopsis.
Anas leucopsis; see Branta leucopsis.
Anas mollissima; see Somateria Mollis-

sima.
Anas moschata; see Cairina moschata.
Anas nigra; see Oidemia nigra.
Anas penelope; see Marcca penelope.

Anns platyrhyncha :

Trichostrongylus tenuis.

Syngam us trachea.

Heterakis dispar.

Ascaridia galli.

Porrocaecum crassum.

Porrocaecum ensbcaudaium.

( 'ontracaccum microcephalum.

atrcptocara crassicauda.

Tetrameres flssispina.

Hystrichis tricolor.
Anas platyrhyncha domcstica:

Trichostrongylus tenuis.

Epomidiostomum uncinatum.

Cyathostoma brinchialis.

Heterakis dispar.

Heterakis gallinae.

Ascaridia galli.

Ascaridia lincata.

Porrocaecum crassum.

V ontracaccum microcephalum.

Echinuria jugadornata.

Echinuria uncinata.

Tetrameres flssispina.

Tetrameres gigas.

EustrongyUdes mcrgorum.

Eustrong glides papillosus.

EustrongyUdes tubifex.

Hystrichis tricolor.
Anas qucrquedula ; see Querquedula

querqucdula.
Anas rubripes:

Echinuria uncinata.
Anas segetum; see Anser fahalis.
Anas spon sa ; see Aix sponsa.
Anas tadorna; see Tadorna tadorna.
Anhinga anhinga:

Contraca ecu m sp icu ligcru in .

EustrongyUdes ignotus.
Anhinga melunogastris ;

Contracaeeiim spiculigrruin.

Con tracaccum tricuspe.

Acuarin. macrolaima.
A n h inga novaeh ollan d iae :

Contracaecum spiculigerum.
Anhinga rufa:

Contracaecum rodh a in i.

Contracaecum spiculigerum.

Subulura plotiua.

EustrongyUdes african us.
A user acuta,- see Daflla acuta.
Anser albifrons:

Trichostrongylus tenuis.

Am idostom um anseris.

396

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Anser albifrons — Continued.

Epomidiostomum orispinum.

Epomidiostomum uncinatum.
Anser anser:

Trichoslrongylus tenuis.

Amidostomum anseris.

Epomidiostonium orispinum.

Syngamus trachea.

Heterakis dispar.

Hetcrakis gallinae.
Anser anser domesticus:

Trichostrongylus tenuis.

Amidostomum anseris.

Epom idiostomum orispinum.

Cyathostoma bronchialis.

Heterakis dispar.

Heterakis gallinae.

Ascaridia anseris.

Probably Ascaridia lineata; see
" goose."

Echinuria uncinata.

Eustrongylides papillosus.
Anser cinereus; see Anser anser.
Anser cinereus domesticus; see Anser

anser domesticus.
Anser clangula; see Glaucionetta clan-

gula.
Anser crecca; see Nettion crecca.
Anser domesticus; see Anser anser

domesticus.
Anser fabalis:

Amidostomum anseris.

Epomidiostomum orispinum.

Hetcrakis dispar.
Anser ferus; see Anser anser.
Anser ferus domesticus ; see Anser an-
ser domesticus.
Anser fuligula; see Fuligula fuligula.
Anser fusca; see Melanitta fusca.
Anser leucops; see Branta leucopsis.
Anser marila; see Fulix marila.
Anser mollissima; see Somateria mol-
lis sima.
Anser nigra; see Oidcmia nigra.
Anser penelope; see Mareca pcnelope
Anser segetum; see Anser fabalis.
Anthochacra carunculata; see Crea-

dion carunculatus.
Anthropoides virgo:

Porrocaccum serpentulus.
Antrostomus vociferus; see Setochal-

cis vocifera.

Aquila albicilla; see Haliaeetus albi-

cilia.
Aquila chrysaetos:

Porrocaccum depressum.

Physaloptera alata nouveli.
Aquila fasciata; see Hieraaetus fas-

ciatus.
Aquila heliaca:

Porrocaecum depressum.

Porrocaccum kirghisensis.

Physaloptera alata.

Tctramcrcs, species.

Gnathostoma accipitri.
Aquila imperialis; see Aquila helica.
Aquila maculata:

Porrocaecum depressum.

Thclasia stereura.
Aquila naevia; see Aquila pomarina.
Aquila pennata; see Hieraaetus peiv-

natus.
Aquila pomarina:

Porrocaecum depressum.

Thclazia stereura.
Aquila rapax belisarius:

Physaloptera galinieri.
Ara ararauna:

Ascaridia hermaphrodita.
Ara macao:

Ascaridia hermaphrodita.
Ara m ides cajanea:

Tctramcrcs tetrica.
Aratinga leucophtha Ima :

Ascaridia hermaphrodita.
Aratinga pertitnax:

Ascaridia hermaphrodita.
Arceuthornis musicus ;

Porrocaccum ensicaudatum.

Spiroptera turdi.
Arceuthornis philomclos ;

Porrocaccum ensicaudatum.

Porrocaccum heteroura.

Spiroptera turdi.
Arceuthornis pilaris ;

Porrocaecum ensica uda tu m .

Spiroptcra turdi.
ArccuUiornis torquatus:

Porrocaccum ensicaudatum.
Arceuthornis viscivorus ,•

Porroca ccum ensica u datum.
Archibuteo (Buteo) lagopus; see

Buteo lagopus.
Archibuteo vulgaris; see Buteo buteo.

NEMATODE PARASITES OF BIRDS

397

Ardea agami; see Agamia agami.
Ardea caerulea; see Florida caerulea.
Ardca cinerca ;

Porrocaecum rcticulatum.

Porrocaecum serpentulus.

Contraeaecum microccphalum.

Desmidoccrca acrophila.

D esm ido cerca nun ildica .
Ardea cocoi ;

Porrocaecum rcticulatum.

Eu strong glides ignotus.
Ardea comata ; see Comatibis eremita.
Ardea garzetta; see Garzetta garzetta.
Ardea goliath;

Eustrongylides afrieanus.
Ardea grus; see Megalornis grus.
A rdea herodia s ;

Con tracaeeum microeepha him.

Eehinuria ardeae.

EustrongyUdes ignotus.
Ardea leuee; see Casmerodius egretta.
Ardca major; see Ardea cinerea.
Ardea manillensis; see Phoyx pur-
purea.
Ardea melanocephala:

Porrocaecum serpentulus.
Ardca minor; see Botaurus lentigi-

nosus.
Ardea nigra; see Ciconia nigra.
Ardea nycticorax ; see Nycticorax

nycticorax.
Ardca pilcata ; see Pilherodius pilea-

t us.
Ardea purpurea; see Phoyx purpurea.
Ardea seapularis; see Butorides

striatum.
Ardea, species:

Contracaccum microccphalum.

Contracaccum multipapillatum.

Contraeaecum tricuspe.
Ardca stellaris; see Botaurus stellaris.
Ardea violacca ; see Nyctanassa vio-
la cea.
Ardeola grayii;

Contraeaecum microccphalum.
Ardeola ibis; see Bubulcus ibis.
Ardeola ralloides;

Contracaccum microcephaluin.
Ardetta minuta; sec Ixobrychus

minutus.
Arenaria interpres morinella •

Porrocaecum eusicaudatum.

Porrocaecum hctcroura.

Asio flammeus:

Porrocaecum depression.

Porrocaecum spirale.

Synhimantus laticeps.
Asio otus:

Porrocaecum depression.

Porrocaecum spirale.

Ilabronema leptoptera.

Synhimantus laticeps.
Astur gentilis:

Porrocaecum deprcssum.

Ha bronema leptoptera.

Schistorophus bidens.

Physaloptera megalostoma.

Microtetrameres inermis.
Astur melanoleueus ; see Accipiter

melanoleucus.
Astur nisus; see Accipiter nisus.
Astur palumbarius; see Astur gen-
tilis.
Astur trivirgatus:

Ilabronema leptoptera.

Synhimantus sygmoidea.
Asturinula monogrammica; ste Kau-

pifalco monogrammicus.
Athene noctua; see Carine noctua.
Athene noctua glaux; see Carine noc-
tua glaux,
Babulcus lueidus; see Bubulcus ibis.
Balearica pavonina:

Ascaridia cristata.

Porrocaecum serpentulus.
Balearica rcgulorum;

Ascaridia cristata.
Balearica regulosum; see Balearica

rcgulorum.
Barnard ius barnardi;

Habronema incerta.
Bernicla saudirichcnsis ; see Nesochen

sandvicensis.
Bolborh ynch us lineola:

Habronema incerta.
Bolborhynchus monachus; see Myiop-

sitta monachus.
lionasa sylvestris; see Tetrastes bon-
us i a.
lionasa umbclliis;

Heterakis bon a m e.

Ascaridia lincata.

Contracacciun, species.

Cheilospirura spinosa.

D ispharynx sp ira I is.

398

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Bota u rus lent ig inosus :

Contracaecum microcephalum.

Streptocara triaenucha.
Botaurus minor; see Botaurus lentigi-

nosus.
Botaurus mugitans; see Botaurus

lentiginosus.
Botaurus pinnatus:

Eustrongylides ignotus.
Botaurus stcllaris:

Contracaecum microcephalum.

Synhimantus brevicaudata .
Branta canadensis:

Heterakis dispar.
Branta leucopsis:

Amidostomum anseris.

Epom idiostomum orispinum.

Heterakis dispar.
Brotogeris virescens:

Habronema ineerta.
Bubo bubo:

Porrocaecum depressum.

Porrocaecum spirale.

Synhimantus laticeps.
Bubo maximus; see Bubo bubo.
Bubo virginianus:

Porrocaecum depressum.

Porrocaecum spirale.
Bubulcus ibis:

Habroncma ficlieuri.

Synhimantus invaginata.

Tetrameres coccinea.

Microtetrameres spiralis.
Bubulcus lucidus; see Bubulcus ibis.
Bucco capensis:

Subulura forcipata.

Subulura strongylina.

Subulura travassosi.
Bucco ehacuru:

Subulura travassosi.
Bucco collaris; see Bucco capensis.
Bucco macrorhynchus:

Subulura travassosi.

Subulura forcipata.
Bucco melanolcucos ; see Bucco tectus.
Bucco rufiventris ; see Bucco swainsoni.
Bucco, species :

Subulura forcipata.
Bucco striolatus:

Subulura forcipata.

Subulura strongylina.

Subulura travassosi.

Bucco swainsoni:

Subulura forcipata.

Subulura strongylina.

Subulura travassosi.

Microtetrameres cruzi.
Bucco tamatia:

Subulura forcipata.

Subulu7'a strongylina.

Subulura travassosi.
Bucco tamatuia; see Bucco tamatia.
Bucco tectus:

Subulura forcipata.

Subulura strongylina.

Subulura travassosi.
Buccphala clangula; see Glaucionetta

clangula.
Buchanga atra assimilis; see Dicrurus

adsimilis.
" Ijiise " ;=Buteo buteo.
Buteo borealis:

Synhimantus sagittata.

Cyathostoma americana.
Buteo buteo:

Porrocaecum angusticolle.

Porrocaecum depressum.

Habroncma leptoptera.

Habronema mansioni.

Spirura talpae (probably acci-
dental).

Synhimantus hamata.

Thelazia stereura.

Physaloptera alata.
Buteo buteo vulpinus:

Thelazia stereura.
Buteo lagopus:

Syngamus coelebs (nomen nudum).

Porrocaecum angusticolle.

Porrocaecum depressum.

Synhirnantus laticeps.
Buteo vulgaris; see Buteo buteo.
Buteo vulpinus; see Buteo buteo vul-
pinus.
Butorides striatus:

Porrocaecum scrpentulus.
Butorides virescens virescens:

Contracaecum microcephalum.
" Buzzard " :

Physaloptera subalata.
Caccabis chucar; see Alectoris graeca

chukar.
Caccabis petrosa; see Alectoris bar-
bar a.

NEMATODE PARASITES OF BIRDS

399

Caccabis pctrosa spatsi; see Alectoris

barbara spatzi.
Caccabis rufa; see Alectoris rufa.
Caccabis saxatilis; see Alectoris

graeca saxatilis.
Cairina moschata:

Heterakis dispar.

Ascaridia galli.

Porrocaecum crassum.
Callipcpla squamata:

Ascaridia cordata.
Calonectris kuhlii:

Cosmocephalus obvelatus.

Seuratia shipleyi.
Cuneroma cochlearia; see Cochlearius

cochlearius.
Capella media:

Sciadiocara um belli f era.
Capito collaris; see Bucco capensis.
Capito macror; see Bucco macrorhyn-

chus.
Capito melanoleucus ; see Bucco tectus.
Capito rufiventris ; see Bucco swain-

soni.
Capito, species :

Physaloptera fusiformis.
Capito striolatus; see Bucco striolatus.
Capito tamatia; see Bucco tamatia.
Caprimulgus aegyptius saharae:

Subulura leprincei.

Subulura subulata.
Caprimulgus bacaurau; see Nycti-

dromus guianensis derby anus.
Caprimulgus campcstris ; see Podager

nacunda.
Caprimulgus candicans; see Thermo-

chalcis candicans.
Caprimulgus cortopan; see Setochalcis

rufa.
Caprimulgus diurnis; see Podager

nacunda.
Caprimulgus curopaeus:

Subulura subulata.

Subulura suctoria.

Thelazia cholodko irskii.
C a prim u Igus fossii :

Subulura leprincei.
Caprimulgus guianensis; see Nycti-

dromus albicollis.
Caprimulgus leucopygeus; see Nycti-

progne leucopyga.
3612—27 27

Caprimulgus mercurius; see Eleo-

threptus anomalus.
Caprimulgus nacandua; see Podager

nacunda.
Caprimulgus nattereri; see Lurocalis

semitorquatus.
Caprimulgus nigrescens; see Nycti-

polus nigrescens.
Caprimulgus ruficollis:

Subulura forcipata.

Subulura strongylina.

Subulura subulata.
Caprimulgus rufus; see Setochalcis

rufa.
Caprimulgus scaphiuris ; see Lurocalis

semitorquatus.
Caprimulgus semitorquatus ; see Luro-
calis semitorquatus.
Caprimulgus, species :

Subulura suctoria.
Caprimulgus trifurcus; see Hydrop-

salis climacocerca.
Caprimulgus urutas (or urutau) ; see

Nyctibius jamaicensis.
Caprimulgus vociferus; see Setochal-
cis vocifera.
Carbo brasiliensis ; see Phalacrocorax

vigua.
Carbo cormoranus ; see Phalacrocorax

carbo.
Carbo cristatus ; see Phalacrocorax

aristotelis.
Carbo dilophus ; see Phalacrocorax

auritus.
Carbo graculus; see Phalacrocorax

aristotelis.
Carbo pygmaeus; see Phalacrocorax

pygmaeus.
Cariama cristata:

Ascaridia pterophora.

Subulura allodapa.

Subulura forcipata.

Subulura strongylina.

Subulura suctoria.

Oxyspirura brevipenis.
"Cariama hupp6 " ;=Cariama cris-
tata.
Carine noctua:

Syngamus trachea.

Heterakis dispar.

Subulura suctoria.

Porrocaecum spirale.

400

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Carine noctua glaux:

Subulura noctuae.

Spirocerca sanguinolenta (larva).

Dispharynx noctuae.
Carpophaga brenchleyi; see Zonoenas

brenchleyi.
Cascara cascara; see Cascara ferru-

ginea.
Cascara ferruginea:

Cyathostoma bronchialis.
Casmerodius alba egretta:

Contracaecum microcephalum. _

Eustrongylides perpapillatus.
Casmerodius egretta; see Casmerodius

alba, egretta.
Casuarius casuarius:

Cyathostoma boularti.
Ca,suarius galeatus; see Casuarius

casuarius.
Catarractes pachyrhynchus:

Cosmocephalus obvelatus.
Cathartes urubu; see Coragyps urubu.
Catheturus lathami; see Alectura

lathami.
Catorrhactes pachyrhynchus; see Ca-
tarractes pachyrhynchus.
Celeus grammicus:

Habronema mansioni.
Celeus jumana:

Habronema longistriata.
Centropus monachus:

Subulura similis.
Centropus »iame>nsis; see Centropus

sinensis.
Centropus sinensis:

Ascaridia circularis.

Ascaridia trilabium.

Subulura rimula.

Oxyspirura siamensis.
Centurus flavifrons:

Microtetrameres cruzi.
Cephalopterus ornatus:

Habronema unilateralis.
Cepphus grylle:

Contracaecum spiculigerum.

Cosm ocephalu$ o bvelatus.

Streptocara, species (IS. pectini-
fera ) .
Cerchneis naumanni:

Habronema seurati.

Tetrameres, species.

Cerchneis tinnunculus:

Porrocaecum depressum.

Habronema leptoptera.

Habronema spinosa.

Synhimantus laticeps.

Physaloptera alata.
Ceriornis satyra; see Tragopan satyra.
Chaemepelia talpacoti:

Ascaridia columbae.
Charadrius dubius:

Porrocaecum ensicaudatum.
Charadrius hiaticula:

Porrocaecum ensicaudatum.

Echinuria horrida.
Charadrius himantopus ; see Himan-

topus himantopus.
Charadrius morirvellus; see Arenaria

interpres morinella.
Charadrius ocdicnemus; see Oedicne-

mus oedienemus.
Charadrius pluvialis; see Pluvialis

apricaria.
Chelicutia chelicuti:

Hadjelia inermis.
Chelidonaria urbica:

Acuaria attenuata.
Chelidoptera tenebrosa:

Subulura forcipata.

Subulura. strongylina.

Subulura travassosi.
Chenonetta jubata:

Heterakis chenonettae.
Chenopis atrata:

Heterakis circumvallata.

Heterakis gallinae.
Chenopsis atrata; see Chenopis atrata.
Chlidonias nigra:

Rusguniella elongata.
Chloephaga poliocephala :

Amidostomum anseris.

Heterakis acuticaudata.
Chloroceryle amcricana:

Yseria coronata.
Chordeiles semitorquatus ; see Luro-

calis semitorquatus.
Chro'wocephalus ridibundus :

Contracaecum spiculigerum.
Chroocephalus ridibundus ; see Chroi-

cocephalus ridibundus.
Chrysolophus amherstiae :

Heterakis isolonche.

NEMATODE PARASITES OF BIRDS

401

Chrysoloph us pictus:

Syngamus trachea.

Hctcrakis gallinae.

Hetcrakis isolonche.

Hcterakis neoplastica.
Chrysolophus pictus obscurus:

Hetcrakis neoplastica.
Chrysotis auripalliata ; see Amazona

auropalliata.
Chrysotis fcstiva; see Amazona fes-

tiva.
Chrysotis guatemalac; see Amazona

farinosa guatemalae.
Chrysotis leucocephalus ; see Amazona

leucoccphala.
Chrysotis ochroptera; see Amazona

barbadcnsis.
Ciccaba huhula:

Syiroptera penihamata.
Ciconia alba; see Ciconia ciconia.
Ciconia alba asiatica; see Ciconia

ciconia.
C iconia cicon ia :

Syngamus trachea.

Cyathostoma variegatum.

Contracaecum microcephalum.

Cyrnea excisa.

Physaloptera abbreviata (as pseudo-
parasite).
Ciconia maguari; see Euxenura ma-

yuari.
Ciconia nigra:

Syngamus trachea.

Cyathostoma variegatum.

Contracaecum engonium.

Contracaecum microcephalum.

Synhimantus sagittata.

Eustrongylides mcrgorum.
Circaetus gallicus:

Porrocaecum depressum.

Physaloptera alata.
CircactUtv pcctoralte:

Porrocaecum depressum.
Circcstus pectoralis ; see Circaetus pec-
toralis.
Circus aeruyinosus :

Porrocaecum angusticolle.

Porrocaecum depressum.

Ilabronema leptoptera.

Physaloptera alata.
Circus buffoni:

Physaloptera acuticauda.

Physaloptera alata.

Circus cineraceus ; see Circus pygar-

gus.
Circus cyancus:

Porroca ee u m deprcssu m.

Ilabronema leptoptera.

Synhimantus laticeps.

Physaloptera alata.
Circus maculosus ; see Circus buffoni.
Circus pygargus:

Porrocaecum. depressum.

Porrocaecum spirale.

Ilabronema leptoptera.

Synhimantus elliptica.

Synhimantus laticeps.

Physaloptera alata.
Circus rufus ; see Circus acruginosus.
Circus spilonotus:

Ascaridia dolichocerca.
Circus spilothorax ; see Circus spilo-
notus.
Clangula hy emails:

Streptocara crassicauda.

Eustrongylides mcrgorum.
Coccyzus melaeoryphus :

Subulura acutissima.

Subulura forcipata.

Subulura strongylina.

Subulura subulata.

Physaloptera strongylina.
Coccyzus melanocoryphus ; see Coc-
cyzus melaeoryphus.
Coccyzus minor:

Subulura acutissima.

Subulura forcipata.

Physaloptera strongylina.
Cochlearius cochlearius:

Tetrameres cochleariae.

Tetrameres micropenis.
Colaptes auratus luteus:

Ilabronema colaptes.
Colaptes eampestrin:

Ilabronema longistriata.
Col inns virg inianus:

Trichostrongylus per gracilis.

Hcterakis gallinae.

Ascaridia compar.

Cyrnea colini.
Coloeus moncdula:

Syngamus trachea.

Sciadiocara secunda.
Columba arquatrix:

Ascaridia columbae.

402

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Columba domestica; see Columba livia

domestica.
Columba domestica laticauda; see Co-
lumba livia domestica.
Columba gutturo&a ; see Columba livia

domestica.
Columba livia:

Ascaridia columbae.

Dispharynx spiralis.
Columba, livia domestica:

Ornith ostrongylus quadriradiatus.

Ascaridia columbae.

Telrameres confusa.

Tetrameres fissispina.
Columba picui; see Columbina picui.
Columba risoria; see Streptopelia ri-

soria.
Columba speciosa; see Lepidoenas

speciosa.
Columba talpacoti; see Chaemepelia

talpacoti.
Columbina picui:

Ascaridia columbae.
Colijmbus arcticus; see Gavia arctica.
Colymbus atrigularis; see Gavia arc-
tica.
Colymbus auritus:

Porrocaccum praelongum.

Contracaecum spiculigerum.

Cosmocephalus aduncus.

Echinuria decorata.
Colijmbus cristatus:

Contracaeum ovale.

Contracaeum spiculigerum.

Streptocara recta.

Eustrong glides mergorum.
Colymbus dominions :

Contracaecum spiculigerum.
Colymbus nigricans; see Colymbus

auritus.
Colymbus nigricollis:

Contracaecum nigricollis.

Rusgiiniella elongata.
Colymbus ruficollis:

Contracaecum spiculigerum.

Tetrameres fissispina.

Eustrongylides mergorum.
Colymbus ruficollis capensis:

Contracaecum praestriatum.
Colymbus rufovulgaris ; see Gavia

stellata.
Colymbus septentrionalis ; see Gavia

stellata.

Coma t ibis cron it a :

Contracaecum microcephalum.
Conurus leucotis; see Pyrrhura leu-

cotis.
Conurus pavua; see Aratinga leu-

cophthalma.
Conurus pertinax ; see Eupsittnla pcr-

tinax.
Conurus solstitlalis; see Eupsittula

solstitialis.
Coprotheres pomarinus:

Contracaecum spiculigerum.
Coracias abyssinicus:

Snbulura similis.
Coracias garrulus:

Hadjelia truncata.

Acuaria anthuris.

Acuaria cordata.
Cora gyps urubu:

Tetrameres paradoxa.
Corone cornix ; see Corvus comix.
Corone corone; see Corvus corone.
Corvus americanus; see Corvus bra-

cliyrhynchos.
Corvus brachyrhynchos :

Syngamus gracilis.

Acuaria anthuris.

Microtetrameres helix.
Corvus cajanus; see Cyanocorax cay-
anus.
Corvus caryocatactes ; see Nucifraga

caryocatactes.
Corvus corax:

Acuaria anthuris.

Acuaria cordata.
Corvus corax tingitanus:

Spirocerca sanguinolcn ta ( la rya ) -

Acuaria anthuris.

Microtetrameres inermis.
Corvus cornix:

Porrocaecum semitcres.

Acuaria anthuris.

Acuaria cordata.

Acuaria depressa.

Tetrameres unispina.

Physaloptera malleus.
Corvus corone:

Syngamus trachea.

Acuaria anthuris.

Acuaria cordata.

Microtetrameres inerm is.

Oxyspirwa sygmoidea.

NEMATODE PARASITES OF BIRDS

403

Corvus frugilegus ; see Trypanoeorax

frugilegus.
Conns glandarius ; see Garrulus glan-

darius.
Corvus monedula; see Coloeus mone-

dula.
Corvus pica; see Pica pica.
Corvus pyrrhocorax; see Pyrrhocorax

pyrrhocorax.
Corvus scapulatus:

Acuaria ornata.
Coscoroba coscoroba:

Cyathostoma coscorobae.
Cotinus virginianus; see Colinus vir-
gin ianus.
Coturnix communis ; see Coturnix co-
turnix.
Coturnix coturnix:

Hetcrakis gallinac.

Ascaridia compar.

Cyrnea eurycerca.
Coturnix dactylisonans ; see Coturnix

coturnix.
Coturnix delagorguei ; see Coturnix

dclagorguei.
Coturnix delagorguei:

Subulura suctoria.
Cranorrh in us corrugatus:

Hadjelia inermis.
Crax blumenbaeliii:

Hetcrakis nattereri.
Crax fasciolata; see Crax sclateri.
Crax sclateri:

Thclazia anolabiata.
Crax uriimutum; see Notliocrax uru-

m u turn.
Crcadion carunculatus:

Oxyspirura anthochaerae.
Creciscus viridis:

Sch istoroph us lacinia t us.
Y.scvia coronata.
Crocopus phoenicoptcrus:

Ascaridia columbae.
Crossoptilon manehurianum; see

Crossoptilon mantchurieum.
Cross opt Hon mantchu ricu m :

Hetcrakis gallinac.

Hetcrakis isolonche.
Crotophaga ani:
Hub ul ura reclinata.

Syiroptera saginata.
Oxyspirura anacanthura.

Crotophaga major:

Subulura reclinata.

Cyrnea semilunaris.

Oxyspirura anaca nthura.
Cryptoglaux funerea:

Porrocaecum deprcssum.

Po)vocaecum spiralc.
Cryptoglaux tengmahm; see Crypto-
glaux funerea.
Crypturornis noctivagus:

Hetcrakis valvata.

Subulura strong ylina.
Crypt urornis, species :

Hetcrakis alata.

Subulura strong ylina.
Crypturornis tataupa:

Subulura strongylina.
Crypturus cupreus (this host uniden-
tifiable) :

Hetcrakis arquata.

Hetcrakis valvata.
Crypturus noctivagus; see Crypturor-
nis noctivagus.
Crypturus parvirostris; see Micro-

crypturus parvirostris.
Crypturus, species; see Crypturornis,

species.
Crypturus tatuapa; see Microcryp-

turus tataupa.
Cuculus cayanus; see Piaya cayana.
Cuculus melanorhynchus ; see Coccy-

zus mclacoryphus.
Cuculus naevius; see Tapcra nacvia.
Cuculus seniculus; see Coccyzus

minor.
Cuculus tingazu; see Piaya cayana.
Cuculus tinguacu; see Piaya cayana.
Cu nc uma leu cog as t ris :

Contracaccum, haliacti.
Cupidonia cupido; see Tympanuchus

cupido.
Curucujus mclanurus:

Cyrnea semilunaris.
Cyanocorax cayanus:

Hetcrakis gallinac.

Subulura papHlosa.

Spiropterd saginata.

A cu a ria ma millaris.
Cygnus atratus; see Chenopis atrata.
Cygnus mclaiwcoryphus; see Sthctw-
Udes melancoriphus.

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BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Cygnus olor; see Sthenelides olor.
Cygnus olor domesticus ; see Stheneli-
des olor domesticus.
Cypselus apus; see Micropus apus.
Dacelo leachii:

Thelazia dacelonis.
Dafila acuta:

Amidostomum anseris.

Epomidiostomum uncinatum.

Ascaridia galli.
Dandalus rubecula; see Erithacus ru-

becula.
Daption capensis:

Scuratia shipleyi.
Dichocercus bi-cornis; see Dichoceros

bicornis.
Dichoceros bicornis:

Microtetrameres contorta.
Dicholophus cristatus; see Cariama

cristata.
Dicholophus margravi; see Cariama

cristata.
Dicrurtis adsimilis:

Iladjelia inermis.

Acuaria gracilis.
Diomedea exulans:

Seuratia shipleyi.

Tetrameres cert a.
Diomedea melanophrys ; see Thalas-

sarchc melanophrys.
Diomedia exulans; see Diomedea ex-
ulans.
Diplopterus naevius; see Tapera

naevia.
Dryobates major:

Syngamus mucronatus (nomen nu-
dum ) .
Dryocopus martins:

Acuaria quadriloba.
Eclcctus pectoralis:

Ceratospira vesiculosa.
Eclcctus roratus:

Habronema incerta.
"Egret" (see also Casmerodius and
Garsetta) :

Porrocaecum reticulatum.
Elanus cacruleus:

Physaloptera acuticauda.
Elanus leucurus:

Physaloptera acuticauda.
Elcctus pectoralis; see Eclectus pec-
toralis.

Eleothreptus anomalus:

Subulura subulata.

Subulura suctoria.
Emberiza pecoris ; see Holothrus ater.
" Engoulevents " ( goatsuckers ) =Cap-
rimulgidae :

Subulura leprincei.

Subulura subulata.
" Epervier " ;= Accipiter nisus.
Erithacus rubecula:

Acuaria subula.
Eupodotis senegalensis:

Hadjelia parva.

Histiocephalus tridens.
Eupsittula pertinax:

Habronema incerta.
Eupsittula solstitialis:

Ascaridia hermaphrodita.
Eurypyga helias:

Ancyracanthopsis bilabiata.
Eurystomus afer:

Subulura recurvata.

Subulura similis.
Euxenura maguari:

HeteraMs valdemucronata.

Cyrnea excisa.

Echinuria longeornata.

Echinuria longevagina ta.
Falcinellus igneus; see Plegadis fal-

cinellus.
Falco aesalon; see Tinnunculus acsa-

lon.
Falco albioilla; see Haliaeetus albi-

cilla.
Falco albicollis; see Leucopternis al-

bicollis.
Falco apivorus; see Pernis apivorus.
Falco ater; see Milvus migrans.
Falco atricapillus ; see Spizastur me-

lanoleucus.
Falco aurantius ; see Falco deiroleucus.
Falco biarmicus erlangeri:

Physaloptera alata.
Falco bidentatus; see Harpagus bi-

dentatus.
Falco brachydactylus; see Circaetus

gallicus.
Falco brasiliensis ; see Polyboru*

plancus.
Falco buteo; see Buteo buteo.
Falco cachinans; see Herpetotheres

cachinnans.
Falco cayennensis ; see Leptodon pal-

liatus.

NEMATODE PARASITES OF BIRDS

405

Falco cenchris; see Cerchneis nau-

manni.
Falco chcrrug:

Porrocaecum depressum.

Habronema leptoptera.
Falco chrysactos ; see Aquila chry-

saetos.
Falco cineraceus; see Circus pygar-
gus.
Falco corona t us; see Harpyhaliaetus

coronatus.
Falco cyaneus; see Circus cyaneus.
Falco dcgener; see Milvago chima-

chima.
Falco deiroleucus:

Habronema leptoptera.
Falco destructor; see Harpia harpyja.
Falco dispar; see Elanus leucurus.
Falco femoralis; see Rhynchofalco

coerulescens.
Falco fnrcatus; see Elanoides forfi-

catus.
Falco gallicus; see Circaetus gallicus.
Falco gracilis; see Geranospiza caeru-

lescens.
Falco haliactus; see Pandion haliae-

tus.
Falco imperialis; see Aquila heliaca.
Falco lagopus; see Buteo lagopus.
Falco lanarius; see Falco chcrrug.
Falco lithofalco; see Tinnunculus reg-

ulus.
Falco longipcnnis:

Physaloptera alata.
Falco magnirostris; see Rupornis mag-

nirostris.
Falco milvus; see Milvus milvus.
Falco minutus; see Accipiter super-

ciliosus.
Falco nacvius; see Aquila maculata.
Falco nisus; see Accipiter nisus.
Falco ornatux; see Spizactus ornatus.
Falco palumbarius ; see Astur gentilis.
Falco palustris ; see Circus buffonu
Falco pcnnatus; see Hieraaetus pcn-

natus.
Falco pcregrinus; see Rhynchodon per-

egrinus.
Falco pygargus ; see Circus pygargus.
Falco rufus; see Circus aeruginosus.
Falco rutilans; see Hcterospizias mer-

idionalis.

Falco, species :

Physaloptera acuticauda.

Physaloptera alata.

I'hysaloptera subalata.
Falco subbuteo:

Habronema leptoptera.

Synhimantus denticulata.

Physaloptera alata.
Falco subbutcus; see Falco subbuteo.
Falco sivainsonii; see Oampsonyjo

swainsoni.
Falco tinnunculus; see Cerchneis tin-
nunculus.
Falco tridentatus; see Astur trivir-

gatus.
Falco unicinctus ; see Parabuteo uni-

cinctus.
Falco urubutinga; see Urubutitiga

urubutinga.
Falco xanthothorax ; see Micastur rufi-

collis.
Ficedula ficedula:

Acuaria, muscicapac.
Flammea flammca; see Tyto alba.
Florida caerulea:

Porrocaecum serpentulus.

Contracaccum andersoni.
Francolinus ad^persus :

Subulura poculum.

Physaloptera brevicauda.
Francolinus biealcaratus:

Heterakis brerispiciilum.

Ascaridia francolina.

Subulura diffcrens.

Subulura suctoria.

Cheilospirura gruveli.
Francolinus gularis:

Heterakis longccaudata.
Francolinus scphaena:

Subulura suctoria.
Francolinus, species :

Subulura gracilis.
Frcgata aquila:

Contracaccum spiculigerum.
Fulica atra:

Amidostom um anscris.

A midos t omit m fu licae.

A midostomu m raillieti.

Epomidiostomum orispinum.

Tetrameres fissispina.

Tetramcres globosa.

Hystrichis icedli.

406

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Fulica leucoptera;

Contracaecum spiculigerum.
Fuligula cristata; see Fuligula fuli-

gula.
Fuligula ferina; see Aithyia ferina.
Fuligula fuligula:

Amidostomum acutum.

Amidostomum anseris.

Epomidiostomum orispinum.
Fuligula fusca; see Melanitta fusca.
Fuligula marila; see Fulix marila.
Fuligula mollissima; see Somateria

mollissima.
Fuligula nigra ; see Oidemia nigra.
Fuligula nyroca; see "Nyroca nyroca.
Fulix marila:

Amidostomum anseris.
Fulmar is glacialte:

Streptocara stellae-polaris.
Gallinago paraguaiae:

Tetrameres dubia.
Gallinula cliloropus:

Amidostomum anseris.

Porrocaecum ensicaudatum.
Galloperdix spadicea:

Eeterakis longecaudata.

Su bulura gallopcrdieis.
Gallus gallus:

Trichostrongylus tenuis.

Syngamus trachea.

Heterakis beramporia.

Heterakis brevispiculum.

Heterakis gallinae.

Heterakis pusilla.

Heterakis put austr alls.

Ascaridia brasiliensis.

Ascaridia compar.

Ascaridia co-mpressa.

Ascaridia galli.

Ascaridia granulosa.

Ascaridia lineata.

Ascaridia styphloccrca.

Subulura brumpti.

Subulura differ ens.

Subulura strongylina.

Subulura suctoria.

Hartcrtia gallinarum.

Spirocerca sanguinolenta (larva).

Gongylonema ingluvicola.

Cheilospirura hamulosa.

DispJiarynx nasuta.

Dispharynx spiralis.

Gallus ga Uus- — Continued.

Streptocara pectinifera.

Syn himantus la t iceps.

Histiocephalus laticaudatus.

Physaloptcra gemina.

Fhysaloptera truncata.

Oxyspirura mansoni.

Oxyspirura parvovum.

Tetrameres americana.

Tetrameres confusa.

Tetrameres fissispina.
Gallus lafayettii:

Heterakis pusilla.
Gampsonyx swainsoni:

Physaloptera acuticauda.

Physaloptera alata.
Garrulus glandarius:

Acuaria anthuris.
Garrulus glandarius cervicalis:

Dispharynx laplantei.
Garzetta garzetta:

Ascaridia aegyptiaca.
Gavia arctica:

Contracaecum spiculigerum.

Streptocara crassicauda.

Streptocara tridentata.

Eustrongylides tubifex.
Gavia stellata:

Contracaecum spiculigerum.

Cosmocephalus aduncus.

Streptocara crassicauda.

Eustrongylides mergorum.

Eustrongylides tubifex.
Gecinus inridis; see Picus viridis.
Gelochclidon nilotica:

Cosmoccph alus obvelatus.

Ancyracanthopsis bihama ta.
Gennaeus nycthemerus:

Heterakis gallinae.
Geopelia, species :

Ascaridia longccirrata.
Geotrygon montana; see Oreopelia

montana.
Geranospiza cacruleseens :

Physaloptera acuticauda.

Physaloptera alata.

Thclazia papulosa.
Geranospizias caerulescens ; see Ger-
anospiza cacruleseens.
Glarcola austriaca; see Glareola prat-

incola.
Glareola pratincola:

Schistorophus spinulosus.

NEMATODE PARASITES OF BIEDS

407

Ola ii c id in m passerinum :

Heterakis dispar.
Olaucionetta clangula:

A )>i idostomum anseris.

E pomidiostomum orispinum.

Contracaecum micropapillatum.

Contracaecum spicutigerum.

Streptocara crass icon da.
" Goose " :

Ascaridia lineata.
Orossiptodon manschuricum; see

Crossoptilon mantchuricum.
" Grouse " :

Trichostrongylus pergracilis.
Grits antigone; see Mathcwsena anti-
gone.
Grus australiasiana ; see Mathewscna

rubicunda.
Grus cinerea; see Megaiornis grus.
Grus communis ; see Megaiornis grus.
Grus grus; see Megaiornis grus.
Grus paradisea; see Tetrapteryx

paradisea.
Grus pavonina; see Balearica pavo-

nina.
Grus viridirostris; see Megaiornis

japonensis.
Guara rubra:

Sciadiocara umbellifera.
Gulra guira:

Subulura forcipata.
Gy pact us barbatus:

Porrocaecum depression.
Gyps fulvus:

Porrocaecum depressum.
Hacmatopus ostralegus:

Con tracaecum spicutigerum.

Sch istorophus aulieatina.
Halcyon chelicutensis; see Cheticutia

cheiicuti.
Haliaeetus albicilla:

Porrocaecum angusticotle.

Porrocaecum depressum.
Haliaeetus Icueocephalus :

Contracaecum haliacti.
Haliaitus leucogaster ; see Cuncuma

leucogastris.
Halieus brasilicnsis ; see Phalacrocor-

ax vigu a.
Harelda glacialis; see Clangula liye-

malis.
Hurpagus bidentatus:

Habronema leptoptcra.

3612—27 28

Harpia harpy ja:

Tlielazia papulosa.
Harpyhaliaetus coronatus:

Physaloptera acuticauda.

Physaloptena alata.
" Hawk ";

Physaloptera alata chevreuxi.
Heliotreptus anomalus ; see Eleothrep-

tus anomalus.
Helotarsus albicilla; see Haliaeetus

albicilla.
Helotarsus ccaudatus ; see Tcrathopius

ecaudatus.
Yerodias egretta: see Casmerodius alba

egretta.
Herodias tricholor; see Hijdranassa

tricolor.
"Heron cendre' ,= Ardea cinerea.
"Heron pourprc" (purple heron) =

Phoyx purpurea.
Herpetotheres cachinnans :

Ha bronema leptoptcra.

Physaloptera acuticauda.

Physaloptera alata.
Heterospizias meridionalis:

Physaloptera acuticauda.

Porrocaecum depressum.
Hieraaetus fasciatus:

Porrocaecum depressum.
Hieraaetus pennatus:

Porrocaecum depressum.

Physaloptera alata.
Jlirnantopus h imantopus:

Amidostomun ch evreuxi.

Porrocaecum ensicaudatum.

Porrocaecum hetcroura.

Chevreuxia re vol hit a.

Tetramcres nouvcli.
Himanfopus mclunopterus; see Himan-

topus himantopus.
Hirundo riparia ; see Riparia riparia.
Hirundo rustica:

Acuaria attenuata.
Hirundo urbica ; see Chelidonaria

urbica.
Houbara macquecni:

S ubulura suctoria.

Hartertia rotundata.
Houbara undulata:

Subulura rima.

Subulura suctoria.

Hartertia confusa.

Hartertui rotundata.

408

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Hydranassa tricolor:

Contracaecum microcephalum.
Hydrochclidon nigra; see Chlidonias

nigra.
Hydroprogne caspia:

Schistorophus acanthocephalicus.
Hydropsalis climacocerca:

Subulura subulata.

Subulura suctoria.
Hylotomus pileatus; see Phloeotomus

pileatus.
Hypotriorchis subbnteo; see Falco

subbuteo.
Ibis aethiopica; see Threskiornis

aethiopicus.
Ibis falcinellus ; see Plegadis falcinel-

lus.
Ibis guarauna; see Plegadis guarauna.
Ibis ibis:

Contracaecum punctatum.
Ibis nudifrons; see Phimosus nudi-

frons.
Ibis rubra: see Guara rubra.
Icteridae :

Oxyspirura tanasijtchuki.
Icterus cristatus; see Ostinops decu-

manus.
Icterus croconotus:

Oxyspirura cephaloptera.
Irrisor erythrorhynchus ; see Phoeni-

culus purpureus.
Ithagenes cruentus; see Ithaginis

cruentus.
Ithaginis cruentus:

Heterakis isolonche.

Ascaridia galli.
Ixobrychus minutus:

Synhimantus brevicaudata.
Kakatoe sulphurea:

Ascaridia hermaphrodita.
Kaupifalco monogrammicus :

Synhimantus subrecta.
Lagopus lagopus:

Heterakis gallinae.

Ascaridia borealis.

Ascaridia compar.
Lagopus mutus:

Heterakis gallinae.

Ascaridia borealis.
Lagopus scoticus:

Trichostrongylus per gracilis.

Syngamus trachea.

Heterakis gallinae.
Lampronessa sponsa; see Aix sponsa.

Lanius collurio:

Viguiera euryoptera.

Acuaria cordata.
Lanius excubitor:

Viguiera euryoptera.
Lanius excubitor dodsoni:

Spirocerca sanguinolenta (larva).
Lanius minor:

Hartertia zakharoici.

Viguiera euryoptera.

Cheilospirura rotundata.

Physaloptera bilabiata.
Lanius rufus; see Lanius senator.
Lanius senator:

Viguiera euryoptera.

Acuaria cordata.
Lanius, species :

Microtetrameres inermis.
Larus argent atus ; see Larus varius.
Larus argentoides; see Larus varius.
Larus canus:

Contracaecum spiculigerum.

Seuratia shipleyi.

Cosmocephalus aduncus.

Cosmocephalus obvelatus.
Larus capistranus; see Larus ridi-

bundus.
Larus fuscus:

Cyathostoma lari.

Contracaecum spiculigerum.

Cosmocephalus obvelatus.
Larus glaucus; see Larus hyperboreus.
Larus hyperboreus:

Cosmocephalus aduncus.

Cosmocephalus obvelatus.
Larus marinus:

Contracaecum spiculigerum.

Cosmocephalus aduncus.

Cosmocephalus obvelatus.
Larus maximus; see Larus marinus.
Larus medius; see Larus hyperboreus.
Larus ridibundus:

Cyathostoma lari.

Contracaecum spiculigerum.

Cosmocephalus aduncus.

Cosmoccp ha lu 8 d ies in g i .

Cosmocephalus obvelatus.

Streptocara tridentata.
Larus, species :

Cyathostoma lari.

Rusguniella elongata.
Larus tridactylus; see Rissa tridac-

tyla.

NEMATODE PABASITES OF BIRDS

409

Larus rarius:

Contracaecum spiculigerum.

Cosmoccphalus aduncus.

Cosmocephalus obvelatus.
Lepidocnas speciosa:

Ascaridia columbae.
Leptodon cayennensis; see Leptodon

palliatus.
Leptodon palliatus:

Physaloptera acuticauda.

Physaloptera alata.
Leptoptila rufaxilla; see Leptotila

ruf axilla.
Leptoptilos crumeniferus:

Echinuria leptoptili.

Eustrongylides africanus.
Leptoptilos dubiub.

Echinuria hargilae.
Leptoptilus crumenifer; see Leptop-
tilos crumeniferus.
Leptoptilus dubius; see Leptoptilos

du bins.
Leptotila rufaxilla:

Ornithostrongylus fariai.
Lestris parasitica; see Stercorarius

parasiticus.
Lestris pomarinus; see Coprotheres

pomarinus.
Leucoptcrnis albicollis:

Habronema. leptoptera.
Limnoeryptcs gallinula:

Echinuria horrida.
Lophoceros scniifasciatus :

Ha djelia inerm is.
Lophophorus impejanus:

Heterakis isolonche.

Heterakis longecaudata.
Lophophorus impeyanus; see Lopho-
phorus impejanus.
Lophortyx calif amicus:

Habronenw incerta.
Lophos t rix crista ta:

Spiroptera penihamata.
Lophotis ruflcrista:

Eartertia obc.sa.

Hartertia rotundata.
Luroca lis semitorqua tus :

Subulura subulata.

Su bulura suctoria.
Luscinia luscinia:

Porrocaecum ensicaudatum.
Luscinia philomcla; see Luscinia lus-
cinia.

Luscinia rubeeula; see Erithacus rube-

cula.
Lyrurus tetrix:

Ornithostrongylus hastatus.

Ascaridia compar.

Ascaridia magnipapilla.
Mackenziaena leachii:

Spiroptera saginata.
Macrodipteryx longipennis:

Subulura leprincei.
Macrodipteryx macro dipt er us ; see

Macrodipteryx longipennis.
Macropygia, nigrirostris:

Ascaridia australis.
Malacoptila torquata:

Subulura forcipata.

Subulura strongylina.

Subulura travassosi.
Mareca penelope:

Amidostomum anseris.

Epomidiostomum ortepinum.

Epomidiostomum uncinatum.

Echinuria uncinata.
MatJiewsena antigone:

Ascaridia cristata.

Ascaridia stroma.
Mathewsena rubicunda:

Porrocaecum serpentulus.
Megacephalon maleo:

Heterakis longecaudata.
Megaloperdix himalayanus:

Heterakis macroura.
Megaloperdix nigelii; see Megaloper-
dix himalayanus.
Megalopterus hauaidensis; see Micra-

nous hawanensis.
Megalornis grus:

Porrocaecum serpentulus.

Ascaridia stroma.

Schistorophus bicuspis.
Megalornis japonensis :

Cyathostoma variegatum.
Mcgaquiscalus major:

Thelazia cirrura.
Melancrpes flavifrons; see Centurus

flavifrons.
Mclanitta dcglandi:

Yscria califomica.
Mclanitta fusca:

Amidostomum acutum.

Amidostomum anseris.

Epomidiostomum orispinum.

Strcptocara crassicauda.

410

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Meleagris gallopavo:

Syngamus trachea.

Hetcrakis gallinae.

Subulura brumpti.

Subulura suctoria.

Cheilospirura hamulosa.

D ispharyn x spiralis.

Oxyspirura mansoni.

Tetrameres confusa.

Tetrameres fissispina.
Mclierax gabar:

Physaloptera galinieri.
Merganser castor; see Mergus mer-
ganser.
Merganser serratus; see Mergus ser-

raior.
Mergellus albellus:

Eustrongylides mergorum.
Mergus albellus; see Mergellus albel-
lus.
Mergus merganser:

Contracaecum spiculigerum.

Streptocara crassicauda.

Tetrameres fissispina.

Eustrongylides m ergorum.

Hystrichis coronatus.
Mergus scrrator:

Cosmocephalus obvclatus.

Streptocara crassicauda.

Eustrongylides mergorum.

Hystrichis varispinosus.
Merops apiastcr:

Schistorophus bidens.
Merula merula ; see Turdus merula.
Merula nigra; see Turdus merula.
Micro nous hawaiiensis :

Contracaecum magnipapillatum.
Micrastur ruficollis:

Habronema leptoptera.
Microcarbo pygmaeus; see Phalacro-

corax pygmaeus.
Microcrypturus parvirostris:

Subulura olympioi.
Microcrypturus tataupa:

Subulura strongylina.
Microdactylus cristatus; see Cariama

cristate/-.
Micropogon, species ; see Capito, species.
Microptts apus:

Syngamus trachea.
Milvago chimachima:

Porrocaecum deprcssum.
Milvus ater; see Milvus migrans.

Milvus govinda; see Milvus migrans

govinda.
Milvus ictinus; see Milvus milvus.
Milvus korschum ; see Milvus migrans.
Milvus migrans :

Porrocaecum deprcssum .
Habronema leptoptera.
Habronema mansioni.
D ispharynx rectovag inn ta .
Milvus migrans govinda:

Porrocaecum angusticollc.
Milvus milvus:
Porrocaecum depressum.
Habronema leptoptera.
Milvus regalis; see Milvus milvus.
Mimus polyglottos:

Porrocaecum ensicauda turn.
Mimus polyglottus; see Mimus poly-
glottos.
Molothrus ater:

Habronema leptoptera.
Momotus brasiliensis; see Momotus

momota.
Momotus momota:

Oxyspirura cephaloptera.
Monacha morpheas; see Monasa mor-

ph oeus.
Monacha nigra,- see Monasa nigra^
Monasa leucops; see Monasa mor-

phoeus.
Monasa morphocus:
Subulura forcipata.
Subulura strongylina.
Subulura travassosi.
Monasa nigra:

Subulura forcipata.
Subulura strongylina.
Subulura travassosi.
Monasa ?iigrifrons:

Subulura travassosi.
Monasa tenebroso; see Chclidoptera

tenebrosa.
Monasa torquata; see Malacoptila tor-

quata.
Monasa tranquilla; see Monasa nigra.
Monticola saxatilis:

Porrocaecum ensicaudatum.
Motacitla alba:

Porrocaecum ensicaudatum.
"Mouette" (European gull) ;=Larus,

species.
Muscicapa atricapilla ; see Ficcdula
ficcdula.

NEMATODE PARASITES OF BIRDS

411

Mycteria americana:

Contracaecutn multipapillatum.
Myiopsitta monachus:

Ilabronema incerta.
" Nandou "—Rhea rothschildi.
Necrosyrtcs monachus:

Porrocaecum depression.
Neophron monachus; see Neerosyrtes

monachus.
Neophron perenopterus ; see Neophron

perenopterus.
Neophron perenopterus:

Habroncma tulostoma.
Nesochen sandvicensis:

Heterakis dispar.

Streptocara crassicauda.
Nettion carolinense:

Echinuria uncinata.
Nettion crecca:

Amidostomum acutum.

Amidostomum anseris.

Epomidiostom um orispinum.
Nicticorax violacens; see Nyctanassa

violacea.
Nisaetus fasiatus; see Hieraaetus fas-

ciatus.
Nisus communis; see Accipiter nisus
Nonnula rubecula:

Subulura strongylina.

8u b u ' u ra travassos i.
Nothocrax urumutum:

Oxyspirura heteroclita.
Nothura maculosa:

Subulura olympioi.
Nucifraga. caryocutactes:

Syngamus parvus.

Acuaria anthuris.

Eustronyylides papillosus.
Numenius arquatus:

Schistorophus longicornis.

Eustronyylides mergorum.

Hystrichis neglectus.
Numida meleagris:

Heterakis brevispiculum.

Heterakis gallinae.

Ascaridia compar.

Ascaridia galli.

Ascaridia numidae.

Subulura diffcrens.

Dispharynx spiralis.

Streptoca ra p&ct in if era.
Numida mitrata rikwae:

Subulura acuticauda.

Numida papulosa transvaalensis:

Ascaridia numidae.

Subulura suctoria.
Numida ptilorhyncha; see Numida me-
leagris.
Numida rikwae; see Numida mitrata

rikwae.
Nyctalc tengmalmi; see Cryptoglaux

funert a.
Nyctanassa violacea:

Porrocaecum serpentulus.

Tetrameres micropenis.
Nyctea nyetea:

Porrocaecu m depressu m .

Porrocaecum spirale.
Nyctiardea grisea; see Nycticorax

nycticorax.
Nyctibius aethercus:

Subulura subulata.

Subulura suctoria.
Nyctibius grandis:

Subulura subulata.

Subulura suctoria.
Nyctibius griseus jamaicensis:

Subulura forcipata.

Subulura strongylina.

Su bulura suctoria.
Nyctibius jamaicensis: see Nyctibius

griseus jamaicensis.
Nycticorax europaeus ; see Nycticorax

nycticorax.
Nycticorax griseus; see Nycticorax

nycticorax.
Nycticorax nycticorax:

Porrocaecum reticulatum.

Porroca ecu m serpentulus.

Contracaecum microcephalum.

Contracaecutn rosarium.

Sytihimantus sa git tat a.

Tetrameres yymiccophila.
Nycticorax, species:

Contracaecum rosarium.
Nyctidromus albicollis:

Subulura subulata.

Subulura suctoria.

Spiroptera saginata.
Nyctidromus guianensis derbyan us:

Subulura forcipata.
N yclipol u s n igresccn s ;

Subulura suctoria.
Nyctiprognc leueoyyya :

Spiroptera saginata.

412

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Nyroca clangula; see Glaucionetta

clangula.
Nyroca ferina; see Aithyia ferina.
Nyroca fuligula; see Fuligula fxili-

gula.
Nyroca marila; see Fulix marila.
Nyroca nyroca:

Tetramcres, species.
Odontophorus capueira:
Heterakis fariai.
Subulura strongylina.
Odontophorus guianensis:

Subulura strongylina.
Oedicnemus capensis:

Hartertia obesa.
Oedicnemus crepitans; see Oedicnemus

oedicnemus.
Oedicnemus oedicnemus:
Porrocaecum cnsicaudatum.
Porrocaecum heteroura.
Hartertia rotundata.
Oedicnemus vermicularis ; see Oedic

nemus vermiculatus.
Oedicnemus vermiculatus:

Hartertia rotundata.
Oidemia deglandi; see Melanitta de

glandi.
Oidemia fusca; see Melanitta fusca.
Oidemia nigra:
Amidostomum acutum.
Amidostomum anscris.
Amidostomum monodon.
Epomidiostomum orispinum.
Epomidiostomum uncinatum.
Oreopelia montana:

Ascaridia magalhaesi.
Orioius auratus:
Had) elm inermis.
Acuaria gracilis.
Orioius galbula; see Orioius orioius.
Orioius orioius:

Acuaria cordata.
Ortyx virginianus; see Cotinus vir-

ginianus.
Ostinops decumanus:

Spiroptera saginata.
Otis afroides; see Afrotis afra afra-

oidcs.
Otis houbara; see Houbara undulata.
Otis macqueeni; see Houbara mac-

queeni.
Otis ruficresta; see Lophotis rufi-
crista.

Otis tarda:

Trichostrongylus tenuis.
Ornithostrongylus papillatus.
Syngamus trachea.
Heterakis gallinae.
Heterakis papulosa.
Otis tetrax:
Heterakis gallinae.
Heterakis papillosa.
Subulura forcipata.
Histiocephalus laticaudatus.
Otus brachyotus; see Otus flammeus.
Otus choliba:

Subulura acutissima.
Spiroptera penihamata.
Spiroptera saginata.
Oxyspirura brevisubulata.
Otus flammeus:
Porrocaecum spirale.
Synhimantus laticeps.
Otus leucotis:

Subulura similis.
Otus vulgaris; see Asio otus.
Palaeornis fasciatus; see Psittacula

fasciata.
Pandion haliaetus:

Porrocaecum angusticolle.
Spiroptera tenuicollis.
Parabuteo unicinctus:
Habronema leptoptera.
Cheilospirura recta.
Physaloptera acuticauda.
Physaloptera alata.
Physaloptera inflata.
Passer domesticus:

Microtetrameres inermis.
Passer domesticus tingitanus:

Spirocerca sanguinolenta (larva).
Pavo cristatus:
Syngamus trachea.
Pseudaspidodera pavonis.
Heterakis gallinae.
Heterakis hamulus.
Cyrnea bulbosa.
Oxyspirura mansoni.
Pavo muticus:
Heterakis hamulus.
Pseudaspidodera pavonis.
Cyrnea bulbosa.
Cheilospirura pavonis.
Pavo spicifer; see Pavo muticus.
Pelecanus americanus; see Pelecanus
crythrorhynchos.

NEMATODE PARASITES OF BIRDS

413

Pelecanus carbo; see Phalacrocorax

carbo.
Pelecanus conspicillat u s :

Contracaecum spicul igerum.
Pelecanus crispus:

Contracaecum micro papillat am.
Pelecanus crythrorhynchos :

Contracaecum spiculigerum.
Pelecanus fuscus; see Pelecanus Oc-
cident alls.
Pelecanus occidentalis:

Contracaecum spiculigerum.
Pelecanus onocrotalus:

Syngamus trachea.

Contracaecum spiculigwum.

Gnathostorna pelecani.
Pelecanus pygmaeus; see Phalacroco-
rax pygmaeus.
Pelecanus rufescens:

Eustrongylides africanus.
Pelecanus, species :

Contracaecum micropapillatum.

Contracaecum spiculigerum.

Synhimantus raillieti.
Pelecanus trachyrhynchus ; see Pele-
canus erythrorhynchos.
Pelidna alpina :

Schistorophus longicornis.

Echinuria horrida.
Penelope humeralis (this host uniden-
tifiable) :

Ascaridia serrata.
Penelope, species :

Thelazia lutzi.
Perdix cinerea ; see Perdix perdix.
Perdix coturnix; see Coturnix cotur-

nix.
Perdix dentata; see Odontophorus

guianensis.
Perdix graeca; see Alectoris graeca.
" Perdix grecque " ; see Alectoris

graeca.
Perdix perdix:

Trichostrongylus tenuis.

Syngamus trachea.

Heterakis gallinae.

Ascaridia compar.
" Perdix rouge "= Alectoris rufa.
Perdix saxatilis; see Alectoris graeca

saxatilis.
Perdix, species:

Heterakis brasiliana.
" Perdrix de roche "=Alectoris graeca
saxatilis.

Pernis apivorus:

Porrocaecum depressum.
Physaloptera alata.
Pernis, species :

Porrocaecum angusticolle.
Phalacrocorax aristotelis:

Contracaecum spiculigerum.
Phalacrocorax atcr:

Contracaecum spiculigerum.
Phalacrocorax auritus:

Contracaecum spiculigerum.
Phalacrocorax carbo:

Syngamus microspiculum.
Contracaecum spiculigerum.
Desmidocerca aerophila.
Echinuria squamata.
Eustrongylides mergorum.
Eustrongylides excisus.
Phalacrocorax fuscicollis:

Contracaecum spiculigerum.
Phalacrocorax graculus; see Phalacro-
corax aristotelis.
Ph a lacrocorax javaniscus :

Contracaecum spiculigerum.
Phalacrocorax pelagicus:

Contracaecum spiculigerum.
Phalacrocorax pygmaeus:
Contracaecum spiculigerum.
Eustrongylides excisus.
Phalacrocorax sulcirostris ; see Phala-
crocorax ater.
Phalacrocorax urile:

Contracaecum spiculigerum.
Phalacrocorax verrucosus:
Contra ca ecu m spieu ligerum.
Streptocara cirrohamata.
Phalacrocorax vigua:

Contracaecum spiculigerum.
Phasianus chrysomelas ; see Phasianus

eolchicus chrysomelas.
Phasianus eolchicus:
Trichostrongylus tenuis.
Syngamus trachea.
Heterakis gallinae.
Heterakis isolonche.
Heterakis ncoplastica.
Phasianus eolchicus chrysomelas:

Heterakis isolonche.
Phasianus eolchicus satscheuensis:

Heterakis neoplastica.
Phasianus gallus; see Gallus gallus.
Phasianus nycthemcrus ; see Gennaeus
nycthemerus.

414

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Phasianus pictus; see Chrysolophus

pictus.
Phasianus reevesi; see Syrmaticus

reevesii.
Phasianus satscheunensis; see Phasi-
anus colchicus satscheuensis.
Phasianus veneratus; see Syrmaticus

reevesii.
Phasianus versicolor:
Heterakis gallinae.
Phimosus infuscatus; see Phimosus

nudifrons.
Phimosus nudifro-n/s :

Hystrichis acanthocephalicus.
Phlegoenas luzonica:

Ascaridia columbae.
Phloeotomus pileatus:
Acuaria quadriloba.
Phlogocnas luzonica; see Phlegoenas

luzonica.
Phoetiicopterus roseus:
Porrocaecum serpentulus.
Echinuria phocnicopteri.
Tetrameres coccinea.
Phoethomis pretrei:

Serticeps vulvoinfiatus.
Phoyx purpurea:
Porrocaecum reticulatum.
Porrocaecum serpentulus.
Contracaecum microcephalum.
Synhimantus invaginata.
Synhimantus sagittata.
Piaya cajanea; see Piaya cay ana.
Piaya cayana:
Subulura carlosi.
Subulura forcipata.
Subulura reclinata.
Subulura strongylina.
Subulura subulata.
Spiroptera saginata.
Pica caudata; see Pica pica.
Pica pica:

Syngamus pugionatus (nomen nu-
dum).

Syngamus trachea.
Porrocaecum ensicaudatum.
Acuaria anthuris.
Acuaria cordata.
Picus campestris; see Colaptes campe-

stris.
Picus canus:

Syngamus mucronatus (nomen nu-
dum).
Syngamus trachea.

Picus grammicus; see Celeus gram-

micus.
Picus jumana; see Celeus jumana.
Picus major; see Dryobates major.
Picus martws; see Dryocopus mar-

tius.
Picus viridis:

Syngamus trachea.

Acuaria quadriloba.
"Pie grieche a Tete rouge "= La nius

senator.
Pilherodius pileatus:

Porrocaecum serpentulus.
Pionus fuscus:

Ascaridia hermaphrodita.
Pionus menstruus:

Ascaridia hermaphrodita.
Pionus (Psittacus) aestivus; see Ama-
zona aestiva.
Pionus (Psittacus) aracanga; see Ara

macao.
Pionus (Psittacus) ararauna; see Ara

ararauna.
Pionus (Psittacus) dominicensis ; see

Amazona vittata.
Pionus (Psittacus) festivus; see Ama-
zona f estiva.
Pionus (Psittacus) leucocephala ; see

Amazona leucocephala.
Pionus (Psittacus) leucotis; see Pyr-

rhura leucotis.
Pionus (Psittacus) menstruus; see

Pionus menstruus.
Pionus (Psittacus) pertinax; see Ara-

tinga pertinax.
Pionus (Psittacus) phoenicurus; see

Pyrrhura molinae.
Pionus (Psittacus) pulverulentus ; see

Amazona farinosa.
Pionus (Psittacus) purpureus; see

Pionus fuscus.
Pionus (Psittacus), species; see Psit-
tacus, species.
Pionus (Psittacus) sulfureus; see Ka-

katoe sulphurca.
Pionus (Psittacus) vinaceus; see Ama-
zona vinacea.
Pisorhina atricapilla; see Otus choliba.
Plata-lea ajaja; see Ajaia ajaja.
Platalca leucorodia:

Tetrameres coccinea.

Eustrong glides afriean us.
Platycercus adscitus:

Ilabroitema incerta.

NEMATODE PARASITES OF BIRDS

415

Platycercus barnardi ; see Barnard ins

barnardi.
Platycercus eximius:

Habronema incerta.
Platycercus palladiceps; see Platycer-
cus adscitus.
Platyeichla flavipes:

Mierotctrameres pusilla.
Plegadis falcinellus:

Echinuria contorta.

Hystrichis orispinus.
Plegadis guara una:

Echinuria calcarata.
Ploccpasser mahali:

Hartertia obesa.

Hartertia rotundata.
Plotus anhinga; see Anhinga anhinga.
Plot us lavaillanti; see Anhinga rufa.
Plotus mclanogastcr; see Anhinga me-

lanogastris.
Plotus novae-hollandiae ; see Anhinga

novaehollandiae.
Plotus rufus; see Anhinga rufa.
Pluvialis apricaria :

Porrocaccum ensicaudatum.

Porrocaecuni hctcroura.

Porrocaccum semiteres.

Eustrongylides mergorum.
Podager nacunda:

Ascardia amblymoria.

Subulura forcipata.

Sit b ulura strongyllna.

Subulura subulata.

Subulura suctoria.
Podiceps auritus; see Colymbus auri-

tus.
Podiceps capensis; see Colymbus rufi-

collis capensis.
Podiceps cristatus; see Colymbus cris-
ta t us.
Podiceps dominicensis ; see Colymbus

dominicus.
Podiceps fluviatilis; see Colymbus rufi-

collis.
Podiceps minor; see Colymbus rufi-

collis.
Podiceps nigricollis; see Colymbus

nigricollis.
Poeoeephalus senega! ns; see Poicepha-

lus senegalus.
Poicephalus senegalus:

Habronema incerta.

Polyborus pluncus:

Porroca ecu m depress u m.
Proccllaria anglorum; see Puffin us

puffinus.
Prodiceps cristatus; see Colymbus

cristatus.
Prodiceps minor; see Colymbus rufi-

collis.
Protogerys virescens; see Brotogeris

virescens.
Pseudotantalus ibis; see Ibis ibis.
Psittaeula fasciata:

Habronema incerta.
Psittacus sinensis; see Eclectus pec-
toral is.
Psittacus, species :

Ascaridia h ermaph rodita.
Psophia viridis:

Heterakis arquata.

Heterakis psophiae.
Pternistes, species; see Pternistis,

species.
Pternistes swainsoni; see Pternistis

swainsoni.
Pternistis, species :

Hartertia annulata.
Ptcrnis t is s wainsoni :

Subulura suctoria.
Ptilopachus petrosus:

Acuaria p t ilopaeh ydis.
Ptilopachys fuscus; see Ptilopachus

petrosus.
Puffinus kuhli; see Calonectris kuhlii.
Puffinus puffinus:

Seuratia procellariae.
Pu Isatriw perspicillata :

Spiroptera saginata.

Tetramcres paradoxa.
Pyrrhocorax alpinus; see Pyrrhocorax

graculus.
Pyrrhocorax graculus:

Syngamus trachea.

Acuaria cordata.
Pyrrh ocorax pyrrhocorax :

Acuaria anthuris.
Pyrrh ura leucotis:

Ascaridia hermaphrodita.

Habronema incerta.
Pyrrhura molinae:

Asea rid ia h ermaph rodita.
Querquedula circia; see Qucrqucdula
querquedula.

416

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Querquedula crecca; see Nettion

crecca.
Querquedula querquedula:

Amldostomum anseris.

Hystrichis neglectus.
Quiscalus major; see Megaquiscalus

major.
Quiscalus quiscula:

Dispharynx spiralis.
Rallus caycnnensis ; see Creciscus viri-

dis.
Ramphastos monilis:

Habronema unilateralis.
Ramphastos, species :

Thelazia digitata.
Ramphastos vitellinus:

Haoronema unilateralis.

D isph arynx crassissvma.
" Rapace nocturne" (owl):

Haoronema monoptera.
Ramphastus, species ; see Ramphastos,

species.
Rhamphastos erythrorhynchus ; see

Ramphastos monilis.
Rhamphastos vitellinus ; see Ramphas-
tos vitellinus.
Rhea americana; see Rhea rothschildi.
Rhea rothschildi:

Dcletrocephalus dimidiatus.

Hetcrakis parisi.

Ascaridia orthocerca.

Spirura imcinipenis.

Spirura zschokkei.
Rhizothcra longirostris:

Hetcrakis interlabiata.
Rhynchodon peregrinus:

Porrocaecum depressum.
Rhynchofalco coerulescens :

Synhimantus recta.
Rhynchotus rufescens:

Hetcrakis brasiliana.

Subulura olympioi.
Riparia riparia:

Acuaria attenuata.
Rissa tridactyla:

Contracaccum spiculigerum.
Rupomis magnirostris-'

Haoronema leptoptera.

Thelazia campanulata.
Salicaria turdoides; see Acroccphahts

arundinaceus.
Saxicola rubetra:

Acuaria tenuis.
Scolopax gallinula; see Limnocryptcs

gallinula.

Scolopax major; see Capella media.
Scops leucotis; see Otus leucotis.
Sephina francolinus ; see Francolinus

sephaena.
Setochalcis rufa:

Subulura subulata.

Subulura suctoria.
Setochalcis vocifera:

Subulura forcipata.

Subulura suctoria.
Somateria dresseri:

Amidostomum anseris.
Somateria mollissima :

Amidostomum acutum.

Amidostomum mollissima.

Epomidiostomum orispinum.

Eustrongylid-es mergorum.
Spizaetus mauduyti; see Spizaetus or-

natus.
Spizaetus ornatus:

Physaloptera acuticauda.

Physaloptcra alata.
Spizastur mclanoleucus:

Physaloptera acuticauda.

Physaloptera alata.
Spiziaster melanoleucus; see Spizas-
tur mclanoleucus.
Squatarola helvetica; see Squatarola

squatarola.
Squatarola squatarola:

Porrocaecum ensicaudatum.

Porrocaecum heteroura.

Porrocaecum semiteres.

Schistorophus bicuspis.
Stenopsis candicans; see Thermochal-

cis candicans.
Stcrcorarius parasiticus :

Contracaecum spiculigerum.
Sterna arctica; see Sterna paradisaea.
Sterna caspica; see Hydroprogne cas-

pia.
Sterna hirundo:

Schistorophus acanthocephalicus.
Sterna nigra; see Chlidonias nigra.
Sterna paradisaea:

Cosmocephalus obvelatus.
Sterna risoria; see Gelochelidon nilo-

tica.
Sthenelide*s melancoriphus:

Tetrameres fissispina.
Sthenclides olor:

Heterakis circumvallata.

Hystrichis cygni.

Hystrichis orispinm.

NEMATODE PAEASITES OF BIRDS

417

Sthenelides olor domcsticus:

Echin uria uncinata.
Stictoenas arquatrix; see Columba

arquatrix.
" Stork, black "=Ciconia nigra.
Streptopelia risoria:

Ascaridia columbae.
Streptopelia turtur:

Ascaridia columbae.
Strix alba; see Tyto alba.
Strix albomarginata; see Ciccaba hu-

hula.
Strix aluco:

Porrocaecum dcpressum.

Porrocaecum spirale.
Strix atricapilla; see Otas choliba.
Strix brachyotus ; see Asio flammeus.
Strix bubo; see Bubo bubo.
Strix dasypus; see Cryptoglaux fu-

nerea.
Strix flammea; see Tyto alba.
Strix griseata; see LopJiostrix cris-

tata.
Strix nivea; see Nyctea nyctea.
Strix noctua; see Carine noctua.
Strix nyctea; see Nyctea nyctea.
Strix otus; see Asio otus.
Strix passerina; see Carine noctua.
Strix, species :

Subulura lutzi.

Spiroptera penihamata.
Strix stridula; see Strix aluco.
Strix tangmalmi; see Cryptoglaux fu-

nerca.
Strix torquata ; see Pulsatrix perspicil-

lata.
Struthio australis; see Struthio ca-

melus.
Struthio camelus:

Ornithostrongyhis douglasi.

Codiostomum struthionis.
Struthio camelus molybdophanes :

Codiost 6m um 8 1 ru t hionis.
Struthio molybdophanes; see Struthio

camelus molybdophanes.
Sturnus vulgaris:

Syngamus pugionatus {nomcn nu-
dum).

Syngamus trachea.

Porrocaecum ensicaudatum.

Porrocaecum hcteroura.

Spiroptera turdi.
Surnia passerina; see Glaucidium pas-

serinum.

Sylvia palustris; see Acrocephalus

2>alustris.
Sylvia rubecula; see Erithacus rube-

cula.
Sylvia turdoides; see Acrocephalus

arundinaceus.
Syrmaticus reevesii:

Syngamus trachea.

Heterakis gallinae.
Syrnia aluco; see Strix aluco.
Syrnia nyctea; see Nyctea nyctea.
Tachypetes aquila; see Fregata aquila.
Tachyphonus cristatus brunneus:

Microtetrameres minima.
Tadorna beloni; see Tadorna tadorna.
Tadorna tadorna:

Cyathostoma tadornae.

Heterakis dispar.

Heterakis gallinae.

Streptocara crassicauda.

Hystrichis tricolor.
Tadorna vulpanser; see Tadorna ta-
dorna.
Talegallus lathami; see Alectura la-

thami.
Tantalus loculator; see M y ct er i a

americana.
Tapera naevia:

Subulura forcipata.

Subulura subulata.
Terathopius ecaudatus:

Porrocaecum angusticolle.
Terpsiphone, species :

Hadjelia inermw.
Tctrao bonasia; see Tetrastes bonasia.
Tetrao lagopus; see Lagopus lagopus.
Tctrao tetrix; see Lyrurus tetrix.
Tetrao urogallus:

Syngamus trachea.

Heterakis gallinae.

Ascaridia compar.

Ascaridia cylindrica.

Ascaridia galli.

Subulura' forcipata.

Subulura strongylina.
Tetrao uru; see Tetrao urogallus.
Tetrapteryx paradisea:

Ascaridia stroma.
Tetrastes bonasia:

Heterakis gallinae.

Ascaridia galli.
Thalassarche melanophrys:

Contracaecum scotti.

418

BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Thamnophilus funebris; see Mackcti-

ziaena leachii.
Thaumalea amherstiae; see Chryso-

lophus amherstiae.
Thaumalea obscwus; see Chryso-

lophus pictus obscurus.
Thaumalea. picta; see Chrysolophu<

pictus.
Thermochalcis candicans:

Subulura subulata.

Subulura suctoria.
Thrcslciornis aethiopicus :

Physaloptera, species I'arona.
Tinamus, species :

Heterakis alata.

Heterakis skrjabini.

Ascaridia strclnikowi.

Subulura strongylina.
Tinamus tataupa; see Microcrypturus

tat a u pa.
Tinnunculus aesalon:

Porrocaecum depressum.
Tinnunculus alaudarius ; see Cerchneis

tinnunculus.
Tinnunculus regulus:

Porrocaecum depressum.
Totanus fuscus; see Totanus macu-

latus.
Totanus glottis; see Totanus nebu-

larius.
Totanus hypoleucus ; see Actitis hypo-

leuca.
Totanus maculatus:

Cosmocephalus obvelatus.
Totanus melanoleucus :

Sciadiocara umbellifera.
Totanus nebularius:

Schistoroph us longicornis.
Trachclotis scnegalensis ; see Eupo-

dotis sencgalensis.
Tragopan satyra:

Heterakis bosia.

Heterakis gallinae.

Heterakis isolonche.

Heterakis longccaudata.
Tringa alpina; see Pelidna alpina.
Tringa helvetica; see Sqtiatarola

squatarola.
Tringa vanellus; see Vanellus vanellus.
Tringa variabilis; see Pelidna alpina.
Tringae vanelli; see Vanellus vanellus.
Tringoidc.s hypoleucus ; sec Actitis

hypoleuca.

Trochilus ochropygus; see Phoethornis

pretrei.
T rogon collaris; see Trogonurus col-

laris.
Trogon mellanurus; see Curucujus

melanurus.
Trogon, species :

Subulura bentocruzi.
Trogon variegatus:

Subulura bentocruzi.
Trogon viridis:

Subulura trogoni.
Trogonurus collaris:

Cyrnea semilunaris.
Trypanocorax frugilegus:

Syngamus trachea.

Acuaria anthuris.

Acuaria cor data.

Oxyspirura sygmoidea.

Microtetrameres inermis.
Turdus iliacus; see Arccuthornis:

musicus.
Turdus merula:

Porrocaecum ensioaudatum .

Spiroptera turdi.
Turdus musicus; see Arccuthornis

philomelos.
Turdus pilaris; see Arceuthornis pi-
laris.
Turdus rufi ventri-s:

Microtetrameres pusilla.
Turdus saxatilis; see Monti cola saxa-
tilis.
Turdus torquatus; see Arceuthornis

torquatus.
Turdus riscirorns ; see Arccuthornis

viscivorus.
Turnix javanica taijoor:

Oxyspirura ophthalmica.
Turnix, species :

Subulura, species Baylis and Daub-
ney.
Tumix taigoor; see Turnix javarUca

taijoor.
Turtur sylvatious ; see Strcptopclia

turtur.
Tympanuchus cupido:

Heterakis gallinae.
Tyto alba:

Porrocaecum depressum.

Porrocaecum spirale.

NEMATODE PARASITES OF BIRDS

419

Tyto alba — Continued.

Rpiroptcra pen ih ama ta.

Sjfti h im ant us affin is.

Synhimantus laticeps.
Ulula aluco; see Strix a1uc#.
Upupa epops:

Hadjelia truncata.

Spirocerca sanguinolenta.
Uria aalge:

Contracaecum spiculigerum.

Eustrongylides mcrgorum.
Uria grylle; see Cepphus grylle.
Uria troile; see Uria aalge.
Urocissa erythrorhyncha occipitalis:

Acuaria anthuris.
Urocissa occipitalis; see Urocissa ery-
throrhyncha occipitalis.
Urubitinga urubitinga:

Physaloptcra acuticauda.

Physaloptcra alata.
Utnmania torda; see Alca torda.

Vanellus cristatus; see Vanellus va-
nellus.
Vanellus melanogastcr ; see Squatarola

squatarola.
Vanellus vanellus:

Amidostomum henryi.

Porrocaecum ensica udat nm.

Porrocaecum semiteres.

Rusguniella vanelli.

Streptocara crassicauda charadrii.
Vinago delalandii:

Ascaridia fascia ta.
Vultur cinereus; see Aegypius mbna-

chus.
Vultur fulvus; see Gyps fulvus.
^'1tltl<r monachus; see Aegypius mona-

chus.
Vultur percnopterus ; see Neophron

perenopterus.
Zonocnas brenchlcyi:

Ccratospira oph thai mica.

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422 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

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NEMATODE PAEASITES OF BIRDS 423

Gendre, E. — Continued.

1920a — Sur une espece nouvelle <T " Acuaria," parasite de " PtUopachys
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1922. — Sur quelques espece d' " Hahronema," parasites des oiseaux. Actes
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Gmelin, J. Fr.

[1790] — Caroli a Linne . . . Systema naturae per regna trianaturae, secun-
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Horst, R.

1885 — A new entozoon from Struthio molybdophanes, Rchw. Notes Ley-
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Irwin-Smith, Vera.

1920 — Nematode parasites of the domestic pigeon (Columbia livia domes-
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Jagerskiold, L. A.

1909 — Zur Kenntnis der Nematoden-Gattungen Eustrongylides und Hyst-
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48 pp., 12 figs., 5 pis.
Johnston, T. Harvey.

1912 — Notes on some Entozoa. Proc. Roy. Soc. Queensland, vol. 24, pp.
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Kowalewski, M.

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424 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Kraemeb, A.

1853 — Fragmentarisehe Notizen und Abbildungen zur Helminthologie and
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1914 — Suckered roundworms from India and Ceylon. Indian J. M. Re-
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Leidy, Joseph.

1S56 — A synopsis of Entozoa and some of their ectocongeners observed

by the author. Proc. Acad. Nat. Sci. Philadelphia, vol. 8, pp. 42-58.

1904 — Researches in helminthology and parasitology. With a bibliography

of his contributions to science arranged and edited by Joseph

Leidy, jr. 281 pp., figs. Washington.

Leipeb, R. T.

1908 — An account of some helminthes contained in Dr. Wenyon's collec-
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1915 — Parasitic worms, with a note on a free-living nematode. Brit. Ant-
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Lepbi, Giuseppe.

1898— Elminti in rapaci della provincia di Roma. Boll. Soc. rom. per
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1855 — Beitriige zur Anatomie der Nematoden. Arch. f. Anat., Physiol, u.
wissensch. Med., Berlin, pp. 314-336, pis. 12-13, figs. 1-11.

VON LlNSTOW, O.

1873 — Einige neue Nematoden, nebst Bemerkungen fiber bekannte Arten.
Arch. f. Naturg., Berlin, vol. 39, pp. 293-306, pi. 13, figs. 1-16.

1875 — Beobachtungen an neuen und bekannten Helminthen. Arch. f.
Naturg., Berlin, vol. 41, pp. 183-207, pis. 2-4, figs. 1-32.

1876 — Helminthologische Beobachtungen. Arch. f. Naturg., Berlin, vol. 42.
pp. 1-18, pis. 1-2, figs. 1-38.

1877a — Helminthologica. Arch. f. Naturg., Berlin, vol. 43, pp. 1-18, pi. 1,
figs. 1-26.

1877b— Enthelminthologica. Arch. f. Naturg., Berlin, vol. 43, pp. 173-198,
pis. 12-14, figs. 1-30.

1878a — Compendium der Helminthologie. Ein Verzeichniss der bekann-
ten Helminthen, die frei oder in thierischen Korpern leben, geord-
net nach ihren Wohnthieren, unter Angabe der Organe, in denen
sie gefunden sind, und mit Beifiigung der Litteraturquellen.
xxii -f 382 pp. Hannover.

1878b — Neue Beobachtungen an Helminthen. Arch. f. Naturg., Berlin,
vol. 44, pp. 218-245, pis. 7-9, figs. 1-35.

1879a — Helminthologische Studien. Arch. f. Naturg., Berlin, vol. 45, pp.
165-188, pis. 11-12, figs. 1-39.

NEMATODE PARASITES OF BIRDS 425

ton Linstow, O. — Continued.

18796 — Helininthologische Untersuchungen. Jahresb. d. Ver. f. vaterl.

Naturk. in Wiirttemb., Stuttgart, vol. 35, pp. 313-342, figs. 1-24.
1882 — Helminthologische Studien. Arch. f. Naturg., Berlin, vol. 48, pp.

1-25, pis. 1-2, figs. 1-29.
1883 — Nematoden, Trematoden und Aeanthocephalen, gesaimnelt von Prof.

Fedtschenko in Turkestan. Arch. f. Naturg., Berlin, vol. 49, pp.

274-314, pis. 6-9, figs. 1-52.
1S84— Helminthologisches. Arch. f. Naturg., Berlin, vol. 50, pp. 125-145,

pis. 7-10, figs. 1-35.
1888 — Report on the Bntozoa collected by H. M. S. Challenger during

the years 1873-76. Rep. Voyage H. M. S. Challenger (1873-76),

London, vol. 23, Zool., pp. 1-18, pis. 1-2.
1889 — Compendium der Helminthologie. Nachtrag. Die Litteratur der

Jahre 1878-18S9. xvi + 151 pp. Hannover.
1S90 — Beitrag zur Kenntniss der Vogeltanien, nebst Bemerkungen fiber

neue und bekannte Helminthen. Arch. f. Naturg., Berl., vol. 56,

pp. 171-188, pi. 10, figs. 1-20.
1892 — Beobachtungen an Helminthenlarven. Arch. f. mikr. Anat., Bonn.,

vol. 39, pp. 325-343, pi. 15, figs. 1-37.
1894 — Helminthologische Studien. Jenaische Ztschr. f. Naturw., Jena,

vol. 28, pp. 328-342, pis. 22-23, figs. 1-21.
189S — Nemathelminthen gesamnielt von Herrn Prof. Dr. F. Dahl in Bis-

marck-Archipel. Arch. f. Naturg., Berlin (1897), vol. 63, pp. 281-

291, pis. 21-22, figs. 1-21.
1899a — Nematoden aus der berliner zoologischen Sammlung. Mitt. a. d.

zool. Samml. d. Mus. f. Naturk. in Berlin, vol. 1, pp. 3-28, pis.

1-6, figs. 1-7S.
1899ft — Zur Kenntniss der Genera Hystrichis und Tropidocerca. Arch.

f. Naturg., Berl., vol. 65, pp. 155-164. pis. 13-14, figs. 1-16.
1901 — Helminthen von den Ufern des Nyassa-Sees, ein Beitrag zur Hel

minthen-Fauna von Siid-Afrika. Jenaische Ztschr. f. Naturw.,

Jena, vol. 35, pp. 409-428, pis. 13-14, figs. 1-34, A-E.
1902 — Beobachtungen an neuen und bekannten Nemathelminthen. Arch.

f. mikr. Anat., Bonn, vol. 60, pp. 217-232, pi. 13, figs. 1-34.
1903a — Parasiten, meistens Helminthen, aus Siam. Arch. f. mikr. Anat.,

Bonn, vol. 62, pp. 10S-121, pi. 5, figs. 1-23.

19036 — Entozoa des zoologischen Museums der Kaiserlichen Akademie der

Wissenschaften zu St. Petersburg. 2. Ann. Mus. zool. Acad. imp.

d. sci. de St. Petersb., vol. 8, pp. 265-294, pis. 17-19, figs. 1-36.

1904 — Nematoda in the collection of the Colombo Museum. Spolia Zey-

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Zeylanica, Colombo, pt. 11, vol. 3, Jan., pp. 163-188, pis. 1-3,
flgS. 1-55.
19066 — Nematoden des zoologischen Museums in Konigsberg. Arch. f.

Naturg., Berl., vol. 72, Dec, pp. 249-258. pis. 16-18, figs. 1-20.
1906c— Neue helminthen. Central bl. f. Bakteriol. [etc.], Jena, vol. 41,

Orig., pp. 749-752, 1 pi., figs. 1-6.
1906a" — Ostpreussische Nematoden. Schrift. d. phys.-bkonora. Gesellsch.
z. Konigsberg i. Pr., vol. 47, pp. 111-114, 1 pi. 7 figs.

426 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

von Linstow, O. — Continued.

1907 — Nematoden aus dem Koniglichen Zoologischen Museum in Berlin.
Mitt. a. d. zool. Mus. in Berlin, vol. 3, pp. 251-259, pis. 6-7,
figs. 1-22.
1909a — Parasitische Nematoden. Siisswasserfauna Deutschlands (Brauer),

Jena, Heft 15, pp. 47-83, figs. 1-80.
19096 — Neue Helminthen aus Deutsch-Siidwest-Afrika. Centralbl. f. Bak-
teriol. [etc.], Jena., vol. 50, Orig., pp. 448-451, figs. 1-4.
Linton, Edwin.

1892 — Notes on a nematode parasite from the chipping sparrow (Spisilla
sociaUis). Am. Naturalist, Philadelphia, (308), vol. 26, pp. 705-
707, figs. 1-5.
Looss, Arthur.

1905 — Das Genus Trichostronyylus n. g., mit zwei neuen gelegentlichen
Parasiten des Menschen. Centralbl. f. Bakteriol. [etc], Jena, vol.
39, Orig., pp. 409-422, pis. 1-2, figs. 1-14.
L6pez-Neyra, Carlos Rodriguez.

1918 — Sur un nouveau Cyrnea de la Perdix. Compt. rend. d. sc, Pars., vol.

166, pp. 79-82, figs. 1-2.
1922— Notes helmintologicas (cuarta serie) con dos especies nueva del
genero "Alladapa." Bol. real soc. Espanola de hist. nat.Madrid,
vol. 22, pp. 402-418.
Lucet, A. ; and Henry, A.

1911 — La typhlite verruqueuse des faisans et son parasite (Heterakis iso-
lonche v. Linstow). Bull. Soc. centr. de m£d. veL, Paris, vol.
65, pp. 320-333, figs. 1-15.
Lundahl, Carl.

1848 — Bemerkungen iiber zwei neue Strongylus- Arten. Notis. Slillsk. pro
Fauna et Flora Fenn. Forh., Helsingfors, Haft. 1, pp. 283-287.
de Magalhaes, P. S.

1892— Notes d'helminthologie bresilienne. (Deuxieme note). Bull. Soc.
zool. de France, Paris, vol. 17, pp. 219-221, 1 fig.
Megnin, J. P.

1884 — Memoire sur un nouvel helminthe, le Sclerosloma boularti, qui vit
dans la trach§e du casoar. J. de l'anat. et physiol. [etc], Paris,
vol. 20, pp. 455-461, pi. 30, figs. 1-6.
Mehlis, E.

1831 — Novae observationes de entozois. Auctore Dr. Fr. Chr. H. Creplin.
Isis (Oken), Leipzig, vol. 1, pp. 6S-99, pi. 2, figs. 1-18; vol. 2,
pp. 166-199.
Molin, R.

(1857) — Notizie elmintologiche. Atti. r. 1st. veneto di sc, lett. ed art.,
Venezia (1856-1857), vol. 2, pp. 146-152; pp. 216-223, 1 pi., fig.
1-15.
1858 — Prospectus helminthum, quae in prodromo faunae helminthologicae
Venetiae continentur. Sitzungsb. d. k. Akad. d. Wissensch., Wien,
math.-naturw. CI., vol. 30, pp. 127-158.
1859 — Prospectus helminthum, quae in parte secunda prodromi faunae hel-
minthologicae venetae continentur. Sitzungsb. d. k. Akad. d. Wis-
sensch., Wien, math.-naturw. CI. (1858), vol. 33, pp. 287-302.
1860a — Sulla metamorfosi regressiva di alcuni vermi rotondi. Sitzungsb.
d. k. Akad. d. Wissensch., Wien, math.-naturw. CI., vol. 3S, pp.
706-716, 1 pi., figs. 1-4.

NEMATODE PARASITES OF BIRDS 427

Molin, R. — Continued.

1860& — Una monografia del genera Spiroptera. Sitzungsb. d. k. Akad. d.

Wissensch., Wien, inath.-naturw. CI., vol. 38, pp. 011-1005.
1860c — Una monografia del genera Dispharagus ed una monografia del
genere Histiocephalus. Sitzungsb. d. k. Akad. d. Wissensch.,
Wien, math.-naturw. CI., vol. 39, pp. 479-516.
1860d — Trenta specie di neniatoidi. Sitzungsb. d. k. Akad. d. Wissensch.,

Wien, inath.-natury. CI., vol. 40, pp. 331-358.
[lS61fl] — II sottordine degli acrofalli ordinato scientificamente secondo i
risultamenti delle indagini anatomiche ed erubriogeniche. Mem.
r. 1st. Veneto di sci., lett. ed arti, Venezia (1800). vol. 9, pp.
427-633, pis. 25-33.
18616 — Prodromus faunae helminthological venetae adjectis disquisitioni-
bus anatomicis et criticis. Denkschr. d. k. Akad. d. Wissensch.,
Wien, math.-naturw. CI., vol. 19. pp. 1S9-33S, pis. 1-15.
Monnig, H. O.

1923 — On some new South African parasitic nematodes. Tr. Roy. Soc. S.

Africa, Cape Town, vol. 11, pt. 2, pp. 105-117, figs. 1-12.
1924 — South African parasitic nematodes. 9. and 10. Rep. Dir. Vet. Educ.
and Research, Dept. Agric, Union of South Africa, Pretoria, Apr.
1923, pp. 435-478, figs. 1-46.
Montagu, George.

1811 — Account of a species of Fasciola which infests the trachea of
poultry with a mode of cure. Mem. Werner. Nat. Hist. Soc,
Edinburgh (1S0S-10), vol. 1. pp. 194-198. pi. 7, fig. 4; note by
editors, pp. 199-200.

MUEHLIG.

1S84 — Syngamus broach talis. Deutsche Ztschr. f. Thiermed., Leipzig,
vol. 10, pp. 265-271, pi. 13, figs. 1-4.
Mueller, Arthur.

1787 — Verzeichniss der bisher entdeckten Eingeweidewurmer, der Thiere.
in welchen sie gefunden worden, und besten Schriften, die dersel-
ben erwahmen. Naturforscher, Halle, vol. 22, pp. 33-86.
1897 — Helminthologische Mittheilungen. Arch. f. Naturg., Berlin, vol. 63,
pp. 1-26, pis. 1-3.
Neumann, L. G.

1900 — Stir une nouvelle espece de spiroptere (Spiroptera pectinifcra) du
gesier de la poule. Rev. vet., Toulouse, vol. 57, pp. 513-515,
figs. 1-2.
1909 — Parasites et maladies parasitaires des oiseaux domestiques. viii +
230 pp.. 89 figs., Paris.
Ortlepp, R. J.

1922 — The nematode genus Physaloptera Rud. Proc. Zool. Soc. Lond.,

pp. 999-1107, figs. 1-44.
1923 — Two new nematodes, collected in the Zoological Gardens of London.
J. Helminth., London, vol. 1, pp. 61-64, 2 figs.
Owen, Richard.

1835 — Description of a microscopic entozoon infesting the muscles of the
human body. Tr. Zool. Soc. London., vol. 1, pp. 315-324, pi. 41,
figs. 1-9.
Parona, Corrado.

1887 — Elmintologia sarda. Contribuzione alio studio dei vermi parassiti in
animali di Sardegna. Ann. mus. civ. di storia nat. di Genova
(1886-87), vol. 24, pp. 275-384, pis. 5-7, figs. 1-58.

428 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Pebroncito, Edoardo ; and Tomiolo, Aristide

1901 — Sulla elinintiasi mortale dei fagiani. Studi ed osservazioni. Ann.
r. Accad. d'agric. di Torino (1000), vol. 43, mein.. pp. 63-72.

PlANA, G. P.

1879a — Osservazioni sul Dispharaffiis nasutus Rud., dei polli e sulle larve
nematoelmintiche deel mosche e dei porcellioni. Atti Soc. ital.
di sc. nat. (etc.), Milano. vol. 36, pp. 239-262, figs. 1-21.
1S79& — Idem. Mod. zooiatro, Torino, vol. 8, pp. 152-154, fig. 1 ; pp. 170-174,
figs. 2-14; pp. 189-193, figs. 15-21.
Pol6nio, Antonio F.

1S60 — Novae helminthuni species. Lotos, Prag, vol. 10, pp. 21-23.
Ratt.liet, A.

1893 — Traite de zoologie m£dicale et agricole. ed. 2. [fasc. 1]. 736 pp.,

494 figs., Paris.
1895— Idem. [fasc. 2]. xv + 737-1303 pp., figs. 495-892. Paris.
1915 — L'emploi des medicaments dans le traitement des maladies causees
par des nematodes. Rep. 10. Internat. Vet. Cong., London (Aug.,
1914), vol. 3, pp. 733-749.
1916a — La famille des Thelaziidae. J. Parasitol., Urbana, 111., vol. 2, pp.

99-105.
1910b — Nematodes parasites des rongeurs. Rec. de med. vel., Paris, vol.
92, pp. 517-521.
Railliet, A. ; and Henry, A.

1909— Sur la classification des Strongylidae: 1. Metastrongylinae. 2.
Ankylostomfnae. Compt. rend. Soc. de biol., Paris, vol. 66, pp.
85-88; pp. 168-171.
1911a — Les helmintbes du nandou. Bull. Soc. nat. d'acclimat. de France,

Paris, vol. 58, pp. 538-541 ; pp. 573-582, figs. 1-6.
1911& — Helmintbes du pore recueillis par M. Bauche en Annam. Bull. Soc.

de patb. exot., Paris, vol. 4, pp. 693-699.
1912 — Quelques nematodes parasites des reptiles. Bull. Soc. de patb. exot.,

Paris, vol. 5, pp. 251-259, figs. 1-3.
1913 — Un Haemostrongylus des bronches du leopard. Bull. Soc. de path.

exot., Paris, vol. 6, pp. 451^54, figs. 1-2.
1914 — Essai de classification des " Heterakidae." [Compt.=rend.] 9.

Cong, internat. de zool., Rennes (Monaco, 1913), pp. 674-682.
1915 — Sur les nematodes du genre Goezia Zeder. Bull. Soc. de patb. exot.,
Paris, vol. 8, pp. 270-275.
Railliet, A. ; Henry, A. ; and Sizoff, P.

1912— Sur les affinites des dispharages (Acuaria Bremser), nematodes
parasites des oiseaux. Compt. rend. Soc. de biol., Paris., vol. 73,
pp. 622-624.
Ransom, Brayton EL

1904a — Manson's eye worm of chickens (Oxyspirura mansoni), with a
general review of nematodes parasitic in the eyes of birds. Bull.
60. Bureau of Animal Indust, U. S. Dept. Agri., Washington,
pp. 1-54, figs. 1-40, pi. 1, 3 figs.
1904& — A new nematode (Gongylonema ingluvicola) parasitic in the crop
of chickens. Circ. 64, Bureau Animal Indust., U. S. Dept. Agri.,
Washington, 3 pp., figs. 1-2.
RErEiscn, Johannes.

1893 — Trichosomum strumosum n. sp., ein Parasit aus dem Epithel des
Oesophagus von Phasianus eolchicus. Arch. f. Naturg., Berl., vol.
59, pp. 331-340, pi. 13, figs. 1-7.

NEMATODE PARASITES OF BIRDS 429

Rietz, J. H.

1924 — Paralysis in a rooster due to parasites. Jour. Am. Vet. Med. Assoc.,
Detroit, vol. 66, pp. 104-105, figs. 1-3.
Rudolphi, Carl A.

1803 — Neue Beobaehtungen liber die Eingeweidewiirmer. Arch. f. Zool. u.

Zoot.. Braunschweig, vol. 3, pp. 1-32.
1809 — Entozoorum sive vermium intestinaliuin historia naturalis, vol. 2,

457 pp., pis. 7-12. Amstelaedami.
1814 — Erster Nachtrag zu meiner Naturgeschichte der Eingeweidewiirmer.
Mag. f. d. n. Entdeck. in d. ges. Naturk., Berlin, vol. 0, pp. 83-113.
1819 — Entozoorum synopsis cui accedung mantissa duplex et indices locu-
pletissimi. x + 811 pp., 3 pis. 8°. Berolini.
Rust.

1908 — Entenerkrankung durch Tropidocerca fissispina. [Abstract]. Ver-
offentl. a. d. J. Vet. Ber. d. beamt. Tieriirzte Preuss., Berlin (1905),
vol. 6, p. 30.

SCHLOTTHAUBEK.

1860 — Beitriige zur Helminthologie. Amtl. Ber. ii. d. 31. Versamml. deutsch.
Naturf. u. Aerzte, Gottingen (1854), pp. 121-133.
Schneider, Anton.

1866 — Monographie der Nematoden. viii + 357 pp., 122 figs., 28 pis., 343
figs. Berlin.
Schrank, F.

1788 — Verzeichniss der bisher hiniinglich bekannten Eingeweidewiirmer,
nebst einer Abhandlung fiber ihre Anverwandtschaften. 5 p. 1, 116
pp., 1 table. Miinchen.
1790 — Fortekning, pa nagra hittils obeskrifne intestinal-krak. K. Vetensk.
Acad. n. Handl., Stockholm [ners. ser.], vol. 11, pp. 118-126.
Schwartz, Benjamin.

1925 — Ascuridia lineata, a parasite of chickens in the United States. Jour.
Agri. Research, vol. 30, pp. 763-772, figs. 1-18.
Skurat, L. G.

1913n — Sur un Dispharage de la chevf-che et les affiuites du genre Acuaria
Bremser. Compt. rend. Soc. de biol., Paris, vol. 74, pp. 103-106,
figs. 1-6.
1913b — Sur le spiroptere des pies-grieches. Bull. Soc. d'hist. nat. de

TAfrique du nord, Alger, vol 5, pp. 223-225, figs. 1-2.
1913c — Observations sur le Tropidocerea inermis Linst. Bull Soc. d'hist.

nat. de TAfrique du nord, Alger, vol. 5, pp. 191-199, figs. 1-11.
1914a — Sur YHabronema (Spiroptera) Irptoptrra (Rud.). Compt. rend.

Soc. de biol., Paris, vol. 76, pp. 21-24, figs. 1-5.
1914b — Sul la morphologie de I'appareil genital femelle des Spiruridae.

Compt. rend. Acad. d. <w&., Paris, vol. 159, pp. 1016-1018.
1914c — Sur un nouveau parasite de la perdrix rouge. Compt. rend. Soc. de

biol., Paris, vol. 76, pp. 390-393, figs. 1-5.
1914rf— Sur un nouvel habitat et sur la morphologie du Subulura allo&apa
(Crepltn). Compt. rend. Soc. de biol., Paris, vol. 77, pp. 154-157,
figs. 1-4.
1914c — Sur quelques Hittrakis d'oiseaux. Bull. Soc. d'hist. nat. de

TAfrique du nord, Alger, vol. 6. pp. 195-202, figs. 1-5.
1914/ — Sur un nouveau parasite du percnoptere. Bull. Soc. d'hist. nat. de

TAfrique du nord, Alger, vol. 6, pp. 149-153, figs. 1-6.
1914 ff — s ur un nouveau parasite de l'outarde houbara. Bull. Soc. d'hist.
nat. de TAfrique du nord, Alger, vol. 6, pp. 117-119, fig. 1.

430 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Setjbat, L. G. — Continued.

1914/i — Sur deux nouveaux Hete'rakis du Sud-algerien. Bull. Soc. d'hist.

nat. de l'Af rique du nord, Alger, vol. 6, pp. 222-225, figs. 1-4.
1914J — Sur les physalopteres des rapaces. Bull. Soc. d'hist. nat. de

l'Afrique du nord, Alger, vol. 6, pp. 244-253, figs. 1-3.
1914; — Sur la morphologie de l'ovejecteur des Tropidocerca. Coinpt. rend.

Soc. de biol., Paris, vol. 76, pp. 173-176, figs. 1-3.
1914A- — Sur un nouveau spiroptere des rapaces. Compt. rend. Soc. de biol.,

Paris, vol. 76, pp. 427-429, figs. 1-3.
1914/ — Sur un Tropidocerca parasite d'un echassier. Compt. rend. Soc. de

biol., Paris, vol. 76, pp. 778-781, figs. 1-8.
1914m — Sur un nematode parasite du flammant rose. Compt. rend. Soc. d*»

biol., Paris, vol. 76, pp. 814-817, figs. L-4.
1914n — Sur un nouveau spiroptere du chat gante. Compt. rend. Soc. de

biol., Paris, vol. 77, pp. 344-347, figs. 1-5.
1915a — Sur deux Tropidocerca des Ardeidae. Coinpt. rend. Soc. de biol.,

Paris, vol. 78, pp. 279-2S2, figs. 1-4.
1915& — Expedition de AIM. Walter Rothschild, E. Hartert et C. Hilgert dans
le Sud Algerien (mars-mai 1914). Nematodes parasites. Novi-
tates zool., vol. 22, pp. 1-25, figs. A-C, 1-27.
1915c — Un nouveau physaloptere des rapaces. Bull. Soc. d'hist. nat. de

l'Afrique du nord, Alger, vol. 6, pp. 157-159.
1916a — Sur un nouveau dispharage des palmipedes. Compt. rend. Soc. de

biol., Paris, vol. 79, pp. 785-788, figs. 1-5.
19166 — Contribution a l'etude des formes larvaires des nematodes parasites
heteroxenes. Bull. sci. de la France et de la Belg., Paris and
London, ser. 7, vol. 49, pp. 297-377, figs. 1-14.
1916c — Sur un nouveau type de Spiruridae. Compt. rend. Soc. de biol.,

Paris, vol. 79, pp. 517-519, figs. 1-3.
1916d — Sur un nnuvel Habronema du Bubulcus lucidus Raf. Compt. rend.

Soc. de biol., Paris, vol. 79, pp. 295-297, figs. 1-3.
1916e — Sur la quatrieme mue d'un dispharage du flammant. Compt. rend.

Soc. de biol., Paris, vol. 79, pp. 439-441, figs. 1-4.
1916/ — Dispharages d'Algerie. Compt. rend. Soc. de biol., Paris, vol. 79,

pp. 934-938, figs. 1-4.
1917 — Nematodes de la perdrix de roche. Bull. Soc. d'hist. nat. de l'Af-
rique du nord, Alger, vol. 8, pp. 208-215, figs. 1—4.
1918a — Nematodes du Caccabis petropa Gmel. (2e note). Bull. Soc. d'hist.

nat. de l'Afrique du nord, Alger, vol. 9, pp. 50-60, fig. 1.
19186 — Contribution a l'etude de la faune parasitaire de la Tunisie. Ne-
matodes. Arch, de l'lnst. Pasteur de Tunis, vol. 10, pp. 243-275,
figs. 1-4.
1918c — Sur le dispharage de l'echasse. Bull. Soc. d'hist. nat. de l'Afrique

du nord, Alger, vol. 9, pp. 106-109, figs. 1-2.
1918<Z — Sur un nouveau strongyle (Trichostrongylide) de l'echasse. Bull.

du mus. d'hist. nat., no. 2, pp. 113-115, figs. 1-4.
1918c — Sur les strongles du gesier des palmipedes. Bull, du mus. d'hist.

nat. no. 5, pp. 345-351.
1919a — Dispharages (Nematodes) de l'Afrique mineure. Novitat. zool.,

London & Aylesbury, vol. 26, pp. 179-189, figs. A-D.
19196 — Contributions nouvelles a l'etude des formes larvaires des nema-
todes parasites heteroxenes. Bull. biol. de la France et de la
Belg., Paris and London (1918), vol. 52, pp. 344-378, figs, i-xii.

NEMATODE PARASITES OF BIRDS 431

Seubat, L. G. — Continued.

1920 — Sur une filaire du heron cendre. Bull. Soc. d'hist. nat. de l'Afrique
du nord, Alger, vol. 11, p. 142.
Shiplet, A. E.

1911 — "Grouse disease" — continued — strongylosis. Part 1. The thread-
worms (Nematoda). (In The grouse in health and in disease.
London, vol. 1, pp. 207-218, pi. 33, figs. 1-5).

VON SlEBOLD, C.

1836 — Helminthologische Beitrage. Zweiter Beitrag Syngamus trachealis.
Ein doppelleibiger Eingeweidewurin. Arch. f. Naturg., Berlin, vol.
2, pp. 105-116, pi. 3, figs. 1-2.
1837 — (Zusatz to Nathusius, H. : Helminthologische Beitrage, pp. 52-65)
Arch. f. Naturg., Berl., vol. 3, pp. 66-68.
Skbjabin, K. I.

1915ff — Syngamus'y turkestanskikl ptits (Syngamus in birds of Turke-
stan). (Russian text.) Vestnik Obsh. vet., Petrograd, vol. 27,
pp. 645-658, figs. 1-5.
19156 — Strongilidy mysechnago zheludka turkestanskikh pitts. (Vidy
roda Amidostomum Raill. et Henry 1909.) [Strongylidae in the
gizzard of birds of Turkestan. (Variety Amidostomum . . .)]
[Russian text.] Vestnik Obsh. vet., Petrograd, vol. 27, pp. 693-
700, figs. 1-8.
1916a — K karakteristikie Ascaridia cylindrica — vozbuditelia kishechno-
glistnoi boliezniglukharei. [Characteristics of Ascaridia cylin-
drica, the agent causing intestinal parasitism in the moor-hen]
[Russian text.] Vestnik. Obsh. vet., Petrograd, vol. 28, pp. 432-
435, 1 fig.
19166 — Nematodes des oiseaux du Turkestan russe. Ann. Mus. zool.
Acad. imp. d. sci. de Petrograd (1915), vol. 20, mem., pp. 457-
557, figs. 1-58, pi. 8, figs. 40, 41, 54, 59.
1916c — Contribution a l'etude de la faune helminthologique du Paraguay.
J. russe de zool., Petrograd, vol. 1, pt. 4, pp. 736-757, pis. 24-26,
figs. 1-27.
1916rf — Seuratia u. g., nouveau genre de nematodes d'oiseaux. Compt.

rend. Soc. de biol., Paris, vol. 79, pp. 971-973.
1917 — Sur quelques nematodes des oiseaux de la Russie. Parasitology.
Cambridge [Eng.], vol. 9, pp. 460-481, fig. 1, pis. 18-19, figs. 1-19.
1920 — Hartertia zakharowi n. sp. Novaia nematoda kishechnika ptits. Iz-

viest. Donsk. Vet. Inst., Novocherkassk, vol. 2, 4 pp.
1922 — Materialy k monografi ptichikh nematod. II. Rod Thelazia Bosc
1819. [Materiaux pour servir a une monographie des Nematodes
d'oiseaux. II. Le genre Thelazia Bosc 1819.] Ezhegodnik Zoo-
log. Muzeia rossiiskoi Akad. Nauk., (Ann. Mus. zool. de l'Acad.
d. sci. de Russie), 1917-1921, Petrograd, pp. 236-246, figs. 1-4,
pis. 1-2, figs. 1-5.
1923 — Nematody domaschnich ptiz. Opyt monografitscheskoi rasrabotki.
1. Kruglyje tscherwi kurizy, indjeki, zjesarki, pawlina i golubja.
[Abstract] Berl. tieriirztl. Wchnschr., vol. 39, pp. 297-299. From:
Iswestija Donskogo Wet. Inst., vol. 1 and 2, 1920.
19241 — K faune paraziticheskikh chervei turkestanskikh pelikanov (Pele-
canus onocrotalus L.) Trudy Gossud. Inst. Eksper. Vet., vol. 2,
pp. 149-154.
3612—27 29

432 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Smith, Allen J. ; Fox, H. ; and White, C. Y.

1908 — Contributions to systematic helminthology. Univ. Penn. M. Bull.,
Philadelphia, vol. 20, pp. 283-294, pis. 2-10.
Soloviev, P. F.

1912 — Parazitieheskie ehervi ptits Turkestana. [Parasitic worms of the
birds of Turkestan] [Russian text.] Ann. Mus. zool. Acad. imp. d.
sci. de St.-Petersb., vol. 17, pp. 86-115, figs. 1-15.

SONSINO, P.

[1890] — Un nuovo Heterakis del Gallus domesticus, Heterakis differens
mini. Atti Soc. tosc. di sc. nat., Pisa, proc. verb., vol. 7, pp.
136-137.
Stafseth, H. J. and Kotlan, A.

1925 — Report of Investigations on an alleged Epizootic of Ruffed Grouse
in Michigan. Journ. Am. Vet. Med. Ass., new ser., vol. 20, pp.
260-267, figs. 1-5.
Stevenson, E. C.

1904 — A new parasite (Strongylus quadriradiatus n. sp.) found in the
pigeon. (Preliminary report.) Circ. 47, Bur. Anim. Indust.,
U. S. Dept. Agri., Washington, 6 pp., figs. 1-10.
Stiles, C. W.

1892 — Preliminary note on Myzomimus gen. nov., type-species M. scutatus
Mueller, '69, a parasite in cattle. (Note on parasites. 4.) Jour.
Comp. Med. and Vet. Arch., New York, vol. 13, pp. 65-67, 1 fig.
Stiles, C. W. ; and Hassall, A.

1894 — A preliminary catalogue of the parasites contained in the collec-
tions of the United States Bureau of Animal Industry, United
States Army Medical Museum, Biological Department of the Uni-
versity of Pennsylvania (Coll. Leidy) and in Coll. Stiles and Coll.
Hassall. Vet. Mag., Philadelphia, vol. 1, pp. 245-253 ; pp. 331-354.
1905 — The determination of generic types, and a list of roundworm genera,
with their original and type species. Bull. 79, Bur. Anim. Indust.,
U. S. Dept. Agri., Washington, pp. 1-150.
1920 — Index-catalogue of medical and veterinary zoology. Subjects :
Roundworms (Nematoda, Gordiacea and Acanthocephali) and
the diseases they cause. Bull. 114, Hyg. Lab., U. S. Public Health
Serv., Washington, 886 pp.

gTOSSICH, MlCHELE.

18S7 — II genere Heterakis Dujardin. Glasnik hrv. nar. druztva. Zagreb,

vol. 2, pp. 277-301, pis. 3-9, figs. 1-45.
1889 — II genere Pht/saloptera Rudolphi. Lavoro monografico. Boll. Soc.

adriat. di sci. nat. in Tieste, vol. 11, pp. 36-59, pis. 1-3, figs. 1-42.
1S90 — Elminti della Croazia. Glasnik hrv. nar. druztva, Zagreb, vol. 5.

pp. 129-136, pis. 4-5, figs. 1-15.
1891 — 11 genere Dispharagus Dujardin. Lavoro monografico. Boll. Soc.

adriat. di sci. nat. in Trieste, vol. 13, pp. 81-108, 3 pis., figs. 1-27.
1892 — Osservazioni elmintologiche. Glasnik hrv. nar. drustva, Zagreb,

vol. 7, pp. 64-73, pis. 1-2, figs. 1-10.
1898 — Filarie e spiroptere. Lavoro monografico. Boll. Soc. adriat. di

sci. nat. in Trieste, vol. 18, pp. 13-162.
(1898) — Saggio di una fauna elmintologica di Trieste e provincie conter-

mini Program, d.civ. scuola r. sup., Trieste, 162 pp.
1899 — Strongylidae. Lavoro monografico. Boll. Soc. adriat. di sci. nat

in Trieste, vol. 19, pp. 55-152.

NEMATODE PARASITES OF BIRDS 433

Stossich, Michele — Continued.

1900 — Contribute) alio studio degli elminti. Boll. Soc. adriat. di sci. nat.

in Trieste, vol. 20, pp. 1+8+1, pis. 1-2, figs. 1-18.
1902a — Sopra alcuni nematodi della collezione elmintologica del Prof. Dott.
Corrado Parona. Boll. mus. di zool. [etc], Genova (116), 16
pp., pis. 3-5, figs. 1-38.
1902b — Idem. Atti Soc. Ligust. di sci. nat. e geogr., Genova, vol. 13 guigno,

pp. 61-76, pis. 3-5, figs. 1-38.
1904 — Sopra alcuni nematodi. Ann. Mus. zool. d. r. Univ. di Napoli, n. s.,
vol. 1, pp. 1-4, pi. 1, figs. 1-11.
Sweet, Georgina.

1910 — Some new and unrecorded endoparasites from Australian chickens.
Proc. Roy. Soc. Victoria, Melbourne, new ser., vol. 23, pp. 242-
256, pis. 31-35, figs. 1-21.
Theileb, Arnold.

1919 — A new nematode in fowls, having a termite as an intermediary
host. [Filaria gallinarum (new species)]. 5 and 6. Rep.
Director Vet. Research, Dept. Agri., Union South Africa, Pretoria
(1918), pp. [695]-707, 1 pi., fig. 1.
Theiler, A. ; and Robertson, W.

1915 — Investigations into the life history of the wireworm in ostrichs.
3 and 4. Rep. Dir. Vet. Res. Dept. U. S. Africa, Pretoria, pp.
291-345, 9 pis.
Tbavassos, Lauro.

1913 — Sobre as especies brazileiras da subfamilia Heterakinae Railliet &

Henry. Mem. Inst. Oswaldo Cruz, Rio de Janeiro-Manguinhos,

vol. 5, pp. 271-318, pis. 27-31, figs. 1-38.

1914a — Contribuicao para o conhecimento da fauna helmintolojica brazi-

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Contribution to the study of brazilian helminthology. A new

genus of the family Heterakidae, Railliet & Henry. Mem.

Inst. Oswaldo Cruz, Rio de Janeiro-Manguinos, vol. 6, pp. 137-

142, pi. 15, figs. 1-3.

19146 — Tricostrongylinae brazileiras. (Nota previa). Brazil-med., Rio de

Janeiro, vol. 28, p. 163.
1914c — Tricostrongylideos brazileiros. (3. Nota previa). Brazil-med., Rio

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1914d — Contribuicoes para o conhecimento da fauna helmitolojica brazi-
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Creplin, 1846. Mem. Inst. Oswaldo Cruz, Rio de Janeiro-
Manguinhos, vol. 6, pp. 150-162, pis. 16-23, figs. 1-21.
1915a — Contribuicoes para o conhecimento da fauna helmintolojica brazi-
leira. Mem. Inst. Oswaldo Cruz, Rio de Janeiro-Manguinhos,
vol. 7, pp. 146-172, pis. 23-26, figs. 1-14.
1915& — Sobre as especies brazileiras de genero "Tetrameres Cliplin,"
[sic] 1846. (Nota previa). Brazil-med., Rio de Janeiro, vol.
29, pp. 297-298.
1917a — Tetrameridae brazileiras. (2. Nota previa). Brazil-med., Ri<» de

Janeiro, vol. 31, pp. 65-66.
1917& — Contribuicao para o conhecimento da fauna helminthologica Sul-

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1918a — Contribuicao para o conhecimento da fauna helminthologica bra-
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16 pp., 3 pis., figs. 1-13. 8. Sao Paulo.

434 BULLETIN 140, UNITED STATES NATIONAL MUSEUM

Travassos, Lauro — Continued.

19186 — Observacoes sobre os Heterakidae. Rev. Soc. brazil, de sci., Rio
de Janeiro, No. 2, pp. 93-97, 2 pis., figs. 1-2.

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Rio de Janeiro, vol. 9, pp. 79-89, figs. 1-9.

19196 — Contribuicoes para o conbecimento da fauna helmintolojica bra-
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Creplin, 1846. Mem. Inst. Oswaldo Cruz, Rio de Janeiro-Man-
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pp. 63-71.

1920a — Tricbostrongylidae brazileiros. Rev. Soc. brazil, de sci., Rio de
Janeiro, pp. 191-205.

19206 — Esboco de uma cbave geral dos nematodes parasitos. Rev. de vet.
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1920c — Contribuicao a sistematica dos Pbysalopterinae. Rev. de sci., Rio
de Janeiro, vol. 4, pp. 57-58.

[1921] — Contribuicoes para o conbecimento da fauna helmintolojica bra-
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Manguinhos, vol. 13, pp. 1-135, pis. 1-56, figs. 1-250.

1923. — Infromacoes sobre a fauna helminthologica de Matto Grosso. Folha
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Vevebs, G. M.

1923 — Some new and little known helminths from British Guiana. J. Hel-
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Ward, H. B.

1917 — On the structure and classification of North American parasitic
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1-14.

Walter, H.

1866 — Helminthologische Studien. 7 Ber. . . . d. Offeubacher Ver. f.
Naturk., (1865-66), pp. 51-79, pi. 1, figs. 1-6.
Walton, A. C.

1923 — Some new and little known nematodes. J. Parasitol., Urbana, 111.,

vol. 10, pp. 59-70, pis. 6-7, figs. 1-25.
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A-B, pis. 8-11, figs. 1-118.
Weidman, Fred D.

1913— A study of metazoan parasites found in the Philadelphia Zoological
Gardens. Proc. acad. Nat. Sc. Philadelphia, vol. 65, pp. 126-151,
pi. 4, figs. 1-9.
Weinland, David Friedrich.

1858 — Human cestoides (etc.). To which is added an appendix, containing
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Wharton, Lawrence D.

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219-221.

BIRD PARASITES OF NEMATODES 435

Weight, R. R.

1879 — Contributions to American helminthology. No. 1. Proc. Canad. Inst.,
Toronto, n. s., vol. 1 (1), pp. 54-75, pis. 1-2, figs. 1-22.
Zedeb, J. G. H.

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pp., 4 pis. Bamberg.

INDEX

[Nematode synonyms and principal page references are printed in italics]

Page

abbreviata, Physaloptera... 296,507

Abreviata 295

abyssinicus, Coracias 127

acanthocephalica, Spiroptera 287

acanthocephalicus, Hystrichis _ xvn,

366, 376, S77-S7S

Schistorophus xiv, 285, 287

Acanthophorus 334

horridus 343

tenuis 343

Accipiter bicolor 138

nisus 138,

176, 276, 277, 288, 298, 299, 301, 302

tinus __ 297

accipitri, Gnathostoma xvi, S64-S65

Acrocephalus arundinaceus 139

Actitis hypoleuca 235

macularia 235

Acuaria 210,

211, SIS, 213, 214, 215, S16-S17, 218, 219, 220, 221
222. 223, 224, 225, 226, 237, 244, 255, 258, 262, 272

affinis 273

alata 281

anthuris xn, 212, 217, S18-SS0

ardeae 248

attenuata 217,220

calcarata 249

capitata 240

contorla 249

cordata xn, 218, 220-22/

crassissima. ._ 243

depressa xn, 217, 220-22/

elliptica 277

elongata 259

jalcinelli 249

gracilis xn, 217, 218,222-225

gruveli 227

hamala 278

hamulosa 226

laticeps 273,276

longeornata 255

longevaginata.. 256

macrolaima. 212,2/5

mamillaris 212, 2/5

muscieapae 212, SI 4

noctuae 242

ornata xn, 217, SSS-SSi

papillifera xn, 217, 218-224, 394

pelagica. 263

ptilopachydis xn, 212, SI4-SI6

quadriloba xn, 212, S15-S16

raillieti 283

recta 280

rectovaginata 243

rotundata 229

Page

Acuaria similis 222

spiralis 238

squamata 257

subula xn, 218, 224-225

sygmoidea. 282

tarentolae xn, 212, S16

tenuis 217,218,225-226

triaenucha 271

upupae-epopis 190

vanelli.. 260

Acuaria (Cheilospirura) gruveli 227

magnilabiata 241

pavonis 389

skrjabini 389

species 229

Acuaria (Dispharynx) laplantei 241

nasuta 237

noctuae 242

Acuaria (Echinuria) hargilae 253

leptoptili 253

Acuaria (Hamannia) phoenicopteri. 257

uncinata 246

Acuaria (Synhimantus) invaginata 279

laticeps 276

subrecta 282

Acuariidae 163,2/0,211,284,290,311,392

Acuariinae 2//, 212, 216, 226, 231,

233, 237, 244, 258, 261, 262, 264, 272, 290, 391, 392

aculeata, Filaria 254

aculeata, Spiroptera 253

aculeatus, Dispharagus 254

acuta, Anas 27,81

Anser 20

acuticauda, Heterakis vn,52,7ff,77

Oxysoma 133

Physaloptera xiv, 296, S97-S98, 305

Subulura IX, 106, ISS-ISi

acuticaudatus, Strongylus 77

acutissima, Heterakis 107

Subulura vm, 106, 107-108, 109

acutum, Amidostomum 20,24-25

acutus, Strongylus. 24

adspersus, Francolinus 123,307

adunca, Spiroptera 234

aduncus, Cosmocephalus xn, 254-255

Dispharagus. 234

Aegolius brachyotus 144

otus 144,276

aegyptiaca, Ascaridia vn, 78, 85

Heterakis. 83

aegyptius saharae, Caprimulgus -- 118,129

aerophila, Desmidocorca xn, 209, 2/0, 264

aeruginosus, Circus 137,138,176,298

aesalon, Falco.. 138

aestivus, Pionus (Psittacus) 80

437

438

INDEX

Page

aethereus, Nyctibius.- 129,130

aethiopica, Ibis - 297,309

afer, Eurystomus 124,127

affinis, Acuaria 273

Spiroptera 309

Synhimantus Xiv, S73-S74

africanus, Eustrongylides xvn, 367, S69, S70

afroides, Otis 192,198

agami, Ardea 142

Agamonema cysticum 371

Ajaja ajaja 241

ajaja, Ajaja 241

Platalea 241

alata, Acuaria 281

Filaria 281

Heterakis vi, 52, 56, 66

Physaloptera xiv,

296, 297, m-300, 301, 305, 394

Spiroptera 165, 240, 281, 282

Synhimantus.. 281

•lata chevreuxi, Physaloptera 296, S01, 394

alata nouveli, Physaloptera 296, S01-S0S, 394

alatut, Cosmocephalus.- 235

Dispharagus 281

Alauda arvensis 302

species 139

alaudarius, Tinnunculus 299

alba, Ciconia 34,44,152,168,296,307

Motacilla 139

Strix 276

Tyto 273

alba asiatica, Ciconia 168

albellus, Mergus 372

albicilla, Aquila 138

Falco 138

Haliaeetus 138

Helotarsus 137

albicollis, Nyctidromus 130

albifrons, Anas 28

Anser. 10, 20, 22, 27, 383, 384, 385

albomarginata, Strix 205

Alcatorda 147,235,267,372

Alcedo americana 293

ispida 269

Allodapa 104

allodapa.. 108, 126

brumpti 112

leprincei 118

multipapillala 338

noctuae.. 119

suctoria 112, 130

typica 108

allodapa, Allodapa 108,126

Heterakis 108

Oxyuris : 108

Subulura vin, 105, 108-110, 126, 394

alpina, Tringa 254

alpinus, Pyrrocorax 35,220

aluco, Strix 138, 144

Syrnia 144

Ulula .. 144

Amazona ochrocephala 80

ambiguus, Strongylus 287

amblymoria, Ascaridia vn, 78, 8S-8i

amblymoria, Heterakis 83

americana, Alcedo 293

Rhea 32,69,97,165

Page

americana, Tetrameres x v, 334, SS7-SS8

americanum, Cyathostoma vi, 42, -{5-47

americanus, Corvus 220,355

Pelecanus 148

amherstiae, Thaumalea 64

Amidostomidae 7,/7

Amidostominae 18, 19,26

Amidostomum 17, 18, 19, 20, 22, 23, 24, 25, 26, 383

acutuin 20, H-15

anseris v, 19, £0-2/, 29

chevreuxi v, 19, 20, SI-14, 25, 383

fulicae '. 20, t5

henryi v, 20, «

monodon v,20, IS,16

nodulosum 20

raillieti v, 19,20, tl,U, 383

skrjabini 22,383

anacanthura, Oxyspirura xv, 322, SlS-Sti

Spiroptera 323

analis, Dispharagus 281

Anas acuta 27,81

albifrons 28

anser domesticus 28,62

anser fera 28

boschas 10, 34, 62, 136, 139, 152, 245, 267, 343

boschas domestica 10, 27, 42, 53, 62,

81, 93, 136, 152, 246, 343, 345, 367, 372, 373, 376

boschas fera 81,343,376

canadensis 62

clangula 28,147,267

crecca 24,28

fuligula 24,28

fusca 24,28,267

glacialis 267,372

leucops 28,62

mollissima 24,28,372

moschata 62,81,136

nigra 24,27,28

penelope 27,28,246

querquedula 20

rubripes 246,248

segetum 28

sponsa 59

tadorna 53,62,267,376

anatinum, Epomidiostomum 27

Ancylostoma 29

Ancyracanthidae 163,36/, 392

Ancyracanthinae 392

Ancyracanthopsis 36/, 362, 363, 392

bihamata xvi, S6!-S6S

bilabiata 361,562,363,392

Ancyracanthus 284,319,361,363,392

bidens 288

bihamata 362

bilabiatus 362,363

longicornis 285

ophthalmicus 320

pinnatifidus 392

andersoni, Contracaecum x, 147, 11,9-150

anglorum, Procellaria 263

Angullla vulgaris 146

anguillae, Ascaris 146

Porrocaecum ix, 136, 146

angusticolle, Porrocaecum ix, 136, 1S7, 1S8

angusticollis, Ascaris 137

Anhinga anhinga 371

rufa 369

INDEX

439

Page

anhinga, Anhinga 371

Plotus 148

ani, Crotophaga 124,207,323

Anisakinae 135, 146

Anisakis 135

annulata, Hartertia xi, 191, 193-195

anolabiata, Filaria 317

(JOxyspirura) 317

Spiroptera 317

Thelazia... xv, 312, 3/7-3/8

anomalus, Heliotreptus 130

Anser acuta 20

albifrons 10, 20, 22, 27, 383, 384, 385

anser 10,20,28,34,53,62

anser domesticus 20,384

cinereus 10,20,62

cinereus domesticus 10, 11, 42, 53, 246, 373

clangula 20

crecca 20

domesticus 10,101

fabalis 20,62

ferus 10

ferus domesticus 10

fuligula 20

fusca 20

leucops 20

marila 20

mollissima 20

nigra 20

penelope 20

segetum 20,62

anser, Anser - 10,20,28,34,53,62

anser domesticus, Anas 28,62

Anser 20,384

anser fera, Anas 28

Anseriformes 265,268

anseris 268

Amidostomum v, 19,«0-2/,29

Ascaridia vni,79, 101-10t

Strongylus 19,20,28

Anthochaera carunculat3 322

anthochaerae, Ceratospira 324

Oxyspirura. xv,322, 324

Anthropoides virgo... . 142

Anthuris 212,216,217,219

anthuris, Acuaria xii, 212, 217,2/ 8-£?0

Dispharagus 218

Filaria 218,220,330

Spiroptera 217,218,219,220,330

antigone, Orus 89,99

Antrostomus vociferus 129

apiaster, Merops 288

apivorus, Falco 138,298

Pernis 137

apricaria, Pluvialis 139,141

apus, Cypselus 34

Aquila albicilla 138

chry saetos 138, 301

fasciata 138

lmperialis 138,145,298,364

maculata 316

naevia 138,316

pennata 138, 293

rapax belisarius 304

aquila, Tachypetes 138

aquiline, Ascaris 150

aracanga, Pionus (Psittacus) 80

3612—27 30

Pago

Aramides cajanea 350

ararauna, Pionus ( Psittacus) 80

Arboricola torqueola 387

Archibuteo (Buteo) lagopus 137,138

Archibuteo vulgaris 138

arctica, Sterna 235

arcticus, Colymbus 148,205,267,271,367

areuata, Kathleena 152,153

Ardea agami 142

caerulea 142

cinerea 141,142,152,209

cocoi - 141,371

comata . 152

garzetta 83

goliath 369

grus 142

herodias 141,152,248,372

major . 14 2

manillensis 14 1

melanocephala 142

minor 152

nigra 281

nycticorax 142,152,347

pileata. . 142

purpurea 142

scapularis -. 142

species. 152, 154, 160

stellaris. .... 152,274

violacea.. 142

ardeae, Acuaria 248

Ascaris.... 141,142

Dispharagus.. 248

Echinuria-. xra, 244, 248-249

ardcae-stellaris, Strongylus 274

ardea-maguari, Spiroptera 255

Ardeola grayii --- 152

ibis 279

ralloides.. 152

Ardetta minuta,. - 274

Arduenna 4

argentatus, Larus.— 148,234,235

argentoides, Lams 235

argus, Argusianus 387

Argusianus argus 387

arquata, Ascaridia — 55

Heterakis vi, 51, 55, 66, 71, 72

arquatrix, Columba - 87

Stictoenas 87

arquatus, Numenius - 285,372,380

arundinaceus, Acrocephalus... 139

arvensis, Alauda -- 302

Ascaridao - 48, 49, 136

Ascaridata - li 5, 1,3, 392

Ascaridea.. - 48

Ascaridia 49, 50, 55, 77, 78, 80, 81, 83, 84, 85, 86, 87,

88, 89, 90, 91, 92, 93, 95, 96, 97, 98, 99, 100, 101

aegy ptiaca vn, 78, 83

amblymoria vn, 78,83-84

anseris vm,79, 101-102

arquata 55

australis vu, 78, 84,86

borealis. vit, 79, 84,85

brasiliana.. 58

brasillensis - vn, 79, 84-80

catheturina vn, 78, 86-86

circularis vn, 79,85

columbae vn, 79, 86-88, 89, 90

440

INDEX

Page

Ascaridia compar vn, 78,86,88

compressa... 78, 88

cordata vn, 79, 88-89

cristata. vn, 79, 89-90

cylindrica vn, vm, 78, 90, 91

doliehocerca vra, 78,90-9/

fasciata.. vra, 80,91-92

francolina vra, 78,92,94

galli vn,

49, 54, 79, 80, 81-82, 83, 84, 85, 86, 87, 88,
89, 90, 91, 92, 93, 94, 95, 96, 97, 98, 99, 100,
101, 102, 108, 110, 111, 112, 113, 114, 115,
116, 117, 118, 119, 121, 122, 123, 124, 125,
126, 128, 129, 131, 133, 134, 135.

granulosa vra, 78, 92-93, 94

hamia 93,94

hermaphrodita vn, 77, 79, 80, 81

in.fl.exa. 81

lineata vm, 79, 80, 82, 93-94

longecirrata vra, 78, 94-95

magalhaesL vra, 79,95,96

magnipapilla vm, 78, 95-96

numidae vm, 79, 96-97

orthocerca vm, 79,97,98

perspicillum.. 49,81,87

pterophora vm, 79, 97-98

serrata vm, 79,98-99

strelnikowi vra, 79, 99, 100

Stroma vra, 79, 99-100

styphlocerca vra, 78, 100-101

trilabium vra, 79, 101

truncata 80

Ascaris . 49,104,135,146,272

anguillae.. 146

angusticollis. 137

aquillae. 150

ardeae 141,142

columbat— 86

compar 88

crassa.. 136

depressa.. 137

dispar.. 61

ensicaudata 139

fissilabium 206

forciparia 115

forcipata 115,131

galli 81

gallinae 52

gallopavonis. 81

gibbosa 81

hermaphrodita 80

heteriira 144

heteroura 144

hirghisensis . . 145

incisa... 139

inflexa 81

kirghisensis __ 136,145,147

lagopodis... 88

laticauda 97

lumbricoides _ 82

maculosa.- 86

microcephala 152

micropapillata. 153

mucronata 20

multipapillata. . 154

oralis 155

papulosa 67,68,122

Page

Ascaris perspicillum 81

praelonga 140

pterophora 97,98

reclinata 124

reticulata.. 141

rosarius 158

sagittata 281

semiteres 141

serpentulus 142

species 324

spiculigerum 147

spiralis 143

strongylina 77, 128

subulata 128

truncata 80

valdemucronata 74

vesicularis 52

Ascaroidea. 48, 49, 135, 391

Aspidodera 102

Astomum _ 334

Astur melanoleucus. _ 309

nisus 176, 277, 278, 298, 355

palumbarius 138,176,288,355

Asturi nula monogrammica 273, 282

ater, Falco 138,177,243

Milvus 138

Athene noctua... 139

noctua glaux 203

atra, Fulica.... 20, 24, 25, 28, 334, 343, 346, 376, 382, 383

atra assimilis, Buchanga 188,222

atrata, Chenopis. 53

atratus, Cygnus 52,61

atricapilla, Muscicapa _ 214

Pisorhina 107

Strix 107,205,207,325

atricapillus, Falco 297,298

atrigularis, Colymbus 148

attenuata, Acuaria.. 217,220

Spiroptera 220

altenuatus, Dispharagus 220

aulieatina, Schistorophus .. xiv,2$7

aurantius, Falco 177

auratus, Oriolus.. 188,222

auratus luteus, Colaptes 173

auripalliata, Chrysotis 175

auritus, Colymbus... 139,250

Podiceps 148,234

australiasiana, Grus 142

australis, Ascaridia.. vn, 78, 84, 85

Heterakis.. 84

Struthio 31

austriaca, Glareola 285,288

Babulcus lucidus 279

bacaurau, Caprimulgus 115

bancrofti, Heterakis. vi, 50,66

barbatus, Gypaetus. 138

Barbus fluviatilis 368

barnardi, Platycercus 175

Belearica pavonina.. 89

regulosum 89

bellonii, Tadorna 376

bentocruzi, Subulura... vra, 106, 110-111

beramporia, Heterakis... vi, 52, 56-57

Bernicla sandwichensis. 62,267

biarmicus erlangeri, Falco 298,299

bicalcaratum, Polyplectrum. 386

bicalcaratus, Francolinus.. 59,92,111,130,227

INDEX

441

Page

bicolor, Accipiter 138

Spiroptera 371

bicornis, Dichocercus 353

bicuspis, Dispharagus.. 286

Hisliocephalus. 286

Schistorophus 285, 286

Spiroptera 286

bidcns, Ancyracanthus 288

Dispharagus 288

Schistorophus xiv, 285, 286, t88-t 89

Spiroptera.. 288

bidentatus, Falco 177

Harpagus 177

bihamata, Ancyracanthopsis xvi, 562-565

Ancyracanthus 362

bilabiata, Ancyracanthopsis 361,562,363,392

Physaloptera 296,507

Spiroptera 307

bilabiatus, Ancyracanthus 362,363

blumenbachii, Crai 75

Bolborhynchus lineola 175

monachus 175

Bonasa sylvestris 53

umbellus. 75,93,229,239

bonasae, Heterakis vn, 51,75,76

bonasia, Tetrao 53,81

borealis, Ascaridia vn, 79, 84, 86

Buteo 42,45,281

Heterakis 84

boschas, Anas 10,34,62,136,139,152,245,267,343

boschas domestica, Anas 10,27,42,53,62,81,

93, 136, 152, 246, 343, 345, 367, 372, 373, 376

boschas (era, Anas 81,343,376

bosia, Heterakis vi, 51,57-58

Botaurus minor 271

mugitans 152

pinnatus 371

stellaris 152,274

boularti, Cyathostoma vi, 41, 44~46

Sclerostorna 44

Syngamus. . 44

brachydactylus, Falco 138

brachyotus, Aegolius 144

Otus 144,276

Strix.. 138,276

brachyrhynchos, Corvus 39

brasiliana, Ascaridia 58

Heterakis vn, 52,58-59

brasiliensis, Ascaridia vn, 79,8^-85

Carbo 147

Falco 138

Halieus 148

Heterakis 84

Momotus 323

brenchleyi, Carpophaga 320

Zonoenas 320

brevicauda, Physaloptera Xiv, 297,507-508

brevicaudata, Synhimantus... Xiv, 273, 2 74-876, 176

brevicaudatus, Dispharagus 274

Histiocephalus 274

brevipenis, Oiyspirura xv, 322,552

Spiroptera 332

brevispiculum, Heterakis vn, 52,59,60

brevisubulata, Oxyspirura... 322,525

Spiroptera 325

bronchialis, Cyathostoma vi, 35, 41,42, 47

Syngamus 42

Page

brumpti, Allodapa 112

Subulura vm, 106,//2-//5, 131

Bubo inaximus 138,144,276

virginianus 138,144

bubo, Strix 138,144,276

Bubulcus lucidus 174,339

Bucco capensis 128

cbacuru 131, 132

collaris 131

macrorhynchus 131

melanoleucos 128

ruflventris 128, 131

species 115

striolatus 128, 131

swainsoni. 218,131,132,352

tamatia 131

tamatina 128

tectus 131

Bucephala clangula 267

Buchanga atra assimilis 188,222

bulbosa, Cymea 390

Physaloptera xv, 167, 297, 5/0-5/ 1

bullosa, Spiroptera 205

Buteo borealis 42,45,281

buteo 137

lagopus 276

vulgaris 137, 164, 176, 177, 278, 298, 316

vulpinus 316

buteo, Buteo 137

Falco 138, 177

Butorides virescens virescens 152

"Buzzard" 305

Caccabis chucar 88

chukar 114

petrosa 73,126,167,171,187,239

petrosa spatzl 192

rufa.... 126,172,227

saxatilis 88

cachinans, Falco 177,297,298

cachinnans, Herpetotheres 177,297

caerulea, Florida 149

caerulescens, Geranospizias 297,317

caeruleus, Elanus 297

Cairina moschata... 81

cajanea, Aramides -- 350

cajanus, Corvus 53,122,207,213

calcarata, Acuaria 249

Echinuria xin, 245, $49

Hamannia 249

Heterakis... - 96

calcaratus, Dispharagus 249

californica, Yseria 269,292,295,391

callfornicus, Lophortyx 175

Callipepla squamata 88

camelus, Struthio 14,31

campanulata, Filaria 313

Thelazia xv, 312,5/5

campestris, Caprimulgus 78,83,130

Picus 178

canadensis, Anas 62

Cancroma cochlearia 340,348

candicans, Caprimulgus 129

Stenopsis 130

canus, Larus 148,234,235,263

Picus 34

capensis, Bucco 128

Daption 263

442

INDEX

Page

capensis, Oedicnemus 192

Podiceps 156

capillaria, Cheilospirura 287

Schistorophus 287

Spiroptera 287

caplstranus, Larus 233

capitata, Aeuaria 240

Dispharyni 237,«40,394

capitatus, Dispharagus 240

Capito collaris 115

macror 115

melanoleucus 115

ruflventris 115

striolatus 115

tamatia ^ 115

Caprimulgldes 115

Caprimulgus aegyptius saharae 118, 129

bacaurau 115

campestris 78,83,130

candicans 129

cortopan 129

diurnis 129

europaeus 129,130,313

fossii 118

guianensis _ 129,207

leucopygeus 207

mercurius. 129

naeandua 115,128

nattereri 129

nigrescens 130

ruflcollis 115, 128, 129

rufus 130

scaphiuris 129

semitorquatus 129

species 130

trifurcus 129

urutas.. 128

urutau- '. 115

vociferus 130

capueira, Odontophorus 63,128

Carbo brasillensis 147

cormoranus.. 147,258

cristatus 147

dilophus 147

graeulus 147

pygmaeus 147

carbo, Phalacrocorai 38, 148, 209, 258, 370, 372

Cariama cristata 97,108,332

"Cariama huppe". 108

Carine noctuaglaux 119,242

carlosi, Subulura vin, 105, US-llb

carolinense, Nettion 246,248

Carpophaga brenchleyi 320

carunculata, Anthochaera 322

caryocatactes, Corvus 220

Nucifraga 39,220,373

Cascara cascara 42

cascara, Cascara 42

caspica, Sterna 287

castor, Merganser 148

Casuarius galeatus 41,44

Cathartes urubu 335

catheturina, Ascaridia vu, 78,85-58

catheturinus, Heterakis 85

Catheturus lathami... 56,85

Catorrhactes pachyrhynchus 235

Page

catus domesticus, Felis 304

caudata, Heterakis vii, 52, 59-60, 61, 394

Pica 34,139,220

cayana, Piaya 113,115,124

cayanus, Ouculus 129

cayennensis, Falco 297,298

Leptodon 297

Rallus 285,288,293

cenchris, Falco 182

cendre, Heron . 209

Centropus monachus 127

siamensis 329

sinensis 86,101,125,329

cephaloptera, Cheilospirura 322

Oiyspirura XV, 321, 322-323, 32b

Spiroptera 322

Cephalopterus ornatus 186

Cephalostrongylus 11

quadriradiatus 12

Ceratosplra 311,5/9.320

anthochaerae 324

ophthalmica xv, 320-321

vesiculosa xv, 319, 320, 321

Cerchneis tinnunculus. 276

Ceriornis satyra 53,57

certa, Tetrameres 334,358-359

Tropidocerca 338

chacuru, Bucco 131, 132

Charadriiformes 265,268

Charadrius dubius 139,394

hiaticula 139,254,394

himantopus 232

morinellus 139, 144

oedicnemus 144

pluvialis 139,144,372

Cheilospirura 211,

213, 214, 215, 216, 226, 227, 228, 229,
231, 272, 280, 389, 393.

capillaris 287

cephaloptera 322

gruveli XH,226,«27-.&?8,231

hainulosa xa, 226-227

longistriata 178,179

magnilabiata 241

nasuta 238

ophthalmica 328

pavonis 226, 389

recta 280

rotundata xn, 226,229

siamensis 329

skrjabini 226,589

spinosa xn, 226, 229-231

uncinipenis 165

Cheiracanthidea 363

Cheir acanthus 364

chelicutensis, Halcyon 188

Chelidoptera tenebrosa 128, 131

Chenonetta Jubata 60

chenonettae, Heterakis vii, 50,60-61

Chenopis atrata 53

chevreuxi, Amidostomum. ... v, 19, 20, 22-2b, 25, 383

Ch»vreuiia 211,231-232

revoluta .- xu,232-23S

Chloephaga poliocephala 20,77

chloropus, Qallinula 20,139

choliba, Otus 325

INDEX

443

Page

cholodkowskii, Thelazia xv, 312, 3/3-3/4

Chordeiles semitorquatus 129

Chroocephalus ridibundus 148

chrysaetos, Aquila 138,301

Falco 138

Chrysolophus pictus 53

chrysomelas, Phasianus 64

Chrysotis auripalliata 175

festiva 80

guatemalae 175

leucocephalus 175

ochroptera 175

chucar, Caccabis 88,114

Ciconiaalba 34,44,152,168,296,307

alba asiatica 168

ciconia 168

maguari 50,74,168,245,256

nigra 34,44,150,152,281,372

ciconia, Ciconia 168

eiconiae-maguari, Spiroptera 256

cineraceus, Circus 138,176,276,277,298

Falco 177,277

cinerea, Ardea 141,142,152,209

Grus 142,286

Perdii 10,34,53,88

cinereus, Anser 10,20,62

Vultur 138

cinereus domesticus, Anser 10, 11, 42, 53, 246, 373

Circaetus gallicus 138,298

Circestus pectoralis 138

clrcia, Querquedula 380

circularis, Ascaridia vu, 79,86

Heterakis 86

circumvallata, Heterakis vn, 52, 61

Circus aeruginosus 137,138,176,298

cineraceus 138,176,276,277,298

cyaneus 138,176,276,298

maculosus 297

pygargus 298

rufus 138,177,298

spilothorai 90

cirrohamata, Filaria (Spiroptera) 266

Spiroptera 266

Streptocara xm, 265, 266-X67

cirrura, Filaria 318

Thelazia xv, 312, S18-S19

Clamidonema 295

clangula, Anas 28,147,267

Anser 20

Bucephala 267

Qlaucionetta 153

Nyroca - 20,267

clausa, Physaloptera 295

cllmacocerca, Hydropsalis 130

Coccigyforme - 127

coccinea, Tetrameres xvi, 335, SS9-S40

Tropidocerca 339

Coccyzus melacoryphus 107

melanocoryphus 115

minor 115

cochlearia, Cancroma 340,348

cochleariae, Tetrameres xvi, 335,340-34/

cocoi, Ardea - 141,371

Codiostomum 30-31

struthionis v. 81-St

coelebs, Syngamus 34

Page

Colaptes auratus luteus 173

colaptes, Habronema x, 173-174

colchicus, Phasianus 10,34,53,64,65

colini, Cyrnea x, 168-171

Colinus virginianus. 9,53,88, 168

collaris, Bucco 131

Capito 115

Trogon 171

collurio, Lanius 201,220,221

Columba arquatrix ... 87

domestica 87

dom. laticauda 87

gutturosa 87

livia 87,239

livia domestica 12,341,343

picui 87

risoria 87

speciosa 87

talpacoti 87

columbae, Ascaridia vn, 79,86-88,89,96

Ascaris 86

Heterakis 86

Colymbus arcticus 148,265,267,271,367

atrigularis 148

auritus--. 139,250

cristatus 270

nigricans 148

rufovulgaris 148,267

septentrionalis 148, 234, 267, 367, 372

comata, Ardea 152

communis, Coturnix 53,88

Grus 99, 143

Nisus 277,278

compar, Ascaridia vn, 78,86,88

Ascaris 88

Heterakis 88

compressa, Ascaridia..." 78,88

Heterakis 88

confusa, Hartertia xi, 191,105-/97

Tetrameres. . . xvi, 334, 335, 338, 341-34*, 343

conspicillatus, Pelecanus 148

contorta, Acuaria 249

Echinuria xni, 245, 249-? 50

Hamannia 249

Microtetramercs xvi, 351, 353-354

Tetrameres 353

Tropidocerca 353

contortus, Dispharagus 249

Contracaecuro 135, 145, 146, 147, 148, 149, 150, 151,

152,153, 155, 156,157,158,159, 160, 161

andersoni v, 147, 11,9-150

engonium x, 146, 150

haliaeti x, 146, 150-151

magnipapillatum x, 147, 161-162

microcephalum.. x,15t-l 63, 159, 160
micropapillatum. x, 146, 147, 163-164

multipapillatum x, 146, 154-155

multipapillosa 154

ovale X.147, 165-166

praestriatum x, 147, 156-157

punctatum x, 146, 167

ijuadricuspe 152, 153

rodhaini x, 147, 167-168

rosarium X, 147, 158-169

scotti x,147, 159-160

spiculigcrum x, 141, 146, 147-149

444

INDEX

Page

Contracaecum tricuspe x, 147, 160-161

Conurus leucotis 175

pavua 80

pertinax... 175

solstitialis... 80

Coracias abyssinicus 127

corax, Corvus 220

corax tingitanus, Corvus 203,220,355

cordata, Acuaria . _ xu, 218, 220-221

Ascaridia vn, 79,88-89

Heterakis 88

cordatus, Dispharagus 22o

cormoranus, Carbo 147,258

cornix, Corvus 34,141,218,220,221,308,334,351

coronata, Spiroptera 293

Yseria 269,293,392

coronatus, Falco 297,298

Harpyhaliaetus 297

Histiocephalus 293

Hystrichis xvn, 376, 578, 379

corone, Cor\us 34,218,220,330,355

corrugatus, Cranorrhinus 188

cortopan, Caprimulgus 129

corvi-cajani, Spiroptera 122,213

Corvus americanus... 220,355

brachyrhynchos 39

oajanus 53, 122, 207, 213

caryocatactes 220

corax 220

corax tingitanus 203,220,355

cornix 34, 141, 218, 220, 221, 308, 334, 351

corone 34,218,220,330,355

frugilegus 34,218,220,330,355

glandarius 218, 220

monedula.. 34,261,262

pica 34,220

pyrrhocorax 220

scapulatus 223

species 35

Coscoroba coscoroba- 42,47

coscoroba, Coscoroba 42,47

coscorobae, Cyathostoma vi, 42, 47-48

Cosmocephalus 211,233, 234, 235

aduncus xn, 234-235

alatus 235

diesingl xn, 2SS-2S4

dtesingii... 233

obvelatus XU, 234, 235-237

papillosum 235

Coturnix communis 53,88

coturnix 167

dactylisonans 53,88

delagorgnei 130

coturnix, Coturnix 167

Perdix 53

cracii, Spiroptera 331

cracis alectoris, Spiroptera 317

Cranorrhinus corrugatus 188

crassa, Ascaris. 136

Physaloptera 296,302,394

crassicauda, Dispharagus 267

Spiroptera. 267

Streptocara xm, 265, 268-269

crassicauda anseri, Streptocara 267,268

charadrii, Streptocara. xm, 265, 268-269

crassissima, Acuaria 243

Disphary nx 237, 2 43-2 44

Page

crassissimus, Dispharagus.. 243

crassum, Porrocaecum 136-1S7

crassus, IMspharagus 215

Orax blumenbachii 75

fasciolata... 312,317

urumutum 331

crecca, Anas : 24,28

Anser 20

Nettion 384

Querquetula 384

crepitans, Oedicnemus.. 139

crispus, Pelecanus 153

cristata, Ascaridia vn, 79, 89-90

Cariama 97,108,332

Fuligula 24

Heterakis 89

cristatus, Carbo 147

Colymbus 270

Dicholophus 115

Icterus 207

Microdactylus 97, 128, 130

Pavo 34,53,63,103,310,311,325

Podiceps 148, 155,270

Prodiceps 372

Vanellus 22, 139, 141, 260, 268

cri status brunneus, Tachyphonus 358

Crocidura leucodon 206

croconotus, Icterus 323

Crocopus phoenicopterus 87

crosi, Physaloptera xrv, 296, 302-303, 394

Crossoptilon manchuricum 64

Crotophaga ani 124,207,323

major 124, 171, 323

crotophagae ani, Spiroptera 323

majoris, Spiroptera 323

cruentus, Ithagenes 64,81

crumenifer, Leptoptilus 254,369

eruzi, Microtetrameres xvi, 351, 352, 353

Telrameres 333,352

Crypturus cupreus 55,72,74

noctivagus 74,128

parvirostris 121

species 55,128

tatuapa.. 128

Cuculus cayanus 129

melacoryphus 129,309

melanorhynchus 128

naevius 115, 129

seniculus 107,115,309

species 297

tingazu 115, 128, 129

tinguacu 207

cupido, Cupidonia 53

Cupidonia cupido 53

cupreus, Crypturus 55,72,74

curvata, Heterakis 114

Subulura vm, 105, 114-116, 116

cyaneus, Circus 138,176,276,298

Falco 138,177,276

Cyathostoma.. 33, 34, 41, 42, 43, 44, 45, 47

americanum VI, 42, 45-47

boularti vi, H, 44-45

bronchialis vi, 35, 41, 42, 47

coscorobae vi, 42,47-48

lari vi, 41, 42, 43-44

tadornae vi, 41, 42-43

variegatum vi, 41,44

INDEX

445

Page

cyathostomtim, Sclerostoma 43

Cyclistomeae 30

eygni, Echinocephalus 378

H ystrichis x vii, 376, S78-379, 380

Cygnus atratus 52,61

melanocoryphus 343

olor 61,376,379,381

olor domesticus 246

cylindrica, Ascaridia vh, vin, 78, 90, 91

Heterakis 90

Cypselus apus 34

Cyrnea. ... 1 64, 166, 167, 168, 171, 172, 310, 311, 390, 393

bulbosa 590

colini x, 167, 168-171

eurycerca x, 167, 168

excisa x, 167-168, 169

parroti 167, 171

scurati 172

semilunaris 167, 170

seurati x, 167, 169, 171

seuratii 172

cystica, Filaria 368,371

cysticum, Agamonema 371

Dacelo leachii 314

dacelonis, Filaria _. 314

Thelazia xv, 312, 3/4-3/5

daetylisonans, Coturnix 53,88

Daphniapulei 246,248,343,345

Daption capensis... 263

dasypus, Strix 138

decora, Streptocara xin, 265, £69, S70, 394

Yseria. . 269

decorata, Echinuria xni, 244, £50-2 53

decorum, Prionostemma 269

decorus, Dispharagus 269

Histiocephalus 269

degener, Falco 138

delagorgnei, Coturnix 130

deglandi, Oidemia 293

delalandii, Vinago... 91

Deletrocephalae 30

Deletrocephaleae SO

Deletrocephalus 30, St

dimidiatus \,S£-33

dentata, Perdix . 128

denticulata, Spiroptcra 288

Synhimantus xiv, 273, £75, £76

denticulatus, Dispharagus 275

depressa, Acuaria XII, 217, ££0-££l

Ascaris . - - 137

Filaria 221

Fusaria - 137

depressum, Porrocaecum - ix,

136, 1S7-1S8, 1S9, 144, 145

deprestut, Dispharagus 221

Desmidocerca £08-209

aerophila xn, £09, £10, 264

numidica.. £09-£10

Desmidoecrcidae - 163, £08

Desmidocercinae 208

destructor, Falco 317

Diaphanocephalidae 6

diaphanus, Fundulus 371,373

Dichocercus bicornis 353

Dicholophus cristatus 115

margravi 97, 108, 130, 332

diesingi, Cosmocephalus xn, £33-234

Page

diesingii, Cosmocephalus 233

differens, Heterakis Ill

Subulura vm, 106, 108, 11 1-1 If

digitata, Thelazia (?) xv,312,3/9

dilophus, Carbo 147

dimidiatum, Sclerostoma 32

dimidiatus, Deletrocephalus v,S£-33

Dioctophyme 360

Dioctophymidae 366,367

Dioctophymoidea 162,366,375

Diomedea exulans 263,339

melanophrys 159

Diplopterus naevius 115

dispar, Ascaris 61

Falco 297,298

Heterakis vn, 52, 6/, 62

Dispharagus 212,216,

217, 219, 226, 231, 233, 237, 244, 258, 272

aculeatus 254

aduncus 234

alatus 281

analis 281

anthuris 218

ardeae 248

attenuatus 220

bicuspis 286

bidens 288

brevicaudatus - 274

calcaratus 249

capitatus 240

contortus 249

cordatus 220

crassicauda 267

crassissimus 243

crassus 215

decorus 269

denticulatus 275

depressus 221

ellipticus 277

elongatus 259

falconis-subbuteonis 275

gracilis 222

gruveli 227

hamatus 278

hamulosus 226

invaginatus 218,279

involutus 276

laticaudatus -- 290

laticeps 276

longeornatus 255

longevaginatus - 256

macrolaimus 213

magnilabiatus 241

mamillaris 213

muscicapae -- 214

nasutus 237, 238

obvelatus - 235

ornatus - 223

papilliferus 224

papillosus 235

quadrilobus 215

rectovaginatus 243

rectus 280

revolutus 232

rotundatus 229

species 227, 259

species new ? 391

446

INDEX

Page

Dispharagus spiralis 238,276

tpiralit eolumbae 238

squamatus 257

sygmoideus 282

tubula.. 224

tenuis 225

truncatus 190

uncinatus 246

Dispharynx 211, 126, 233, 237, 238,

239, 240, 241, 242, 243, 244, 248, 252, 391

capitata 237,^0

crassissima 7X1,243-244

laplantei xn,237, 241,242

magnilabiata xn, 237, 241-242

nasuta xn,«37-238

noctuae xn, 237, 241, tit-US

rectovaglnata xin, 237, 243, 391

species new? 391

spiralis xn, 237, 238-240, 241

diurnus, Caprimulgus 129

dolichocerca, Ascaridia vm, 78, 90-91

Heterakis 90

domestica, Columba 87

domestica laticauda, Columba 87

domesticus, Anser 10, 101

Passer.. 355

domesticus tingitanus, Passer 203

dominicensis, Pionus (Psittacus) 80

Podiceps. 148

douglasi, Ornithostrongylus v, 1 1, 14-16

Strongylus . 14

Trichostrongylus 14

douglassii, Strongylus . 14

dresseri, Somateria . 20

Dryocopus martius 215

dubia, Filaria 338

Tetrameres x vi, 335, 342-343

dubius, Charadrius 139,394

Leptoptilus 253

ecaudatus, Helotarsus.. 137

Echinocephalus cygni 378

Echinuria 1, 211, «U, 245, 247, 248,

249, 250, 253, 254, 256, 257, 258, 393

ardeae xra, 244, 2 48-249

calcarata xra, 245, 249

contorta xm, 245, 249-250

decorata xm, 244, 250-253

hargilae xm, 245, 252, 253, 254

horrida xm, 245, 268-264

jugadornata - xm, 244, 246-246

leptoptili xm, 245, 254-256, 256

longeornata 245, 266-256

longevaginata xm, 245, 256

phoenicopteri xm, 245, 267

spinifeTa 254

squamata xm, 244, 253, 254, 257-258

uncinata xm, 245,

246-248, 249, 250, 252, 253, 254, 255, 256

Eclectus pectoralis 320

roratus 175

"Egret" 141

egretta, Herodias 152

Elanus caeruleus 297

leucurus.. 297

elegans, Eustrongylides 358,372

Hystrichis 370,372

Strongylus 372

Page

elliptica, Acuaria 277

Synhimantus xiv, 273, 277-278

ellipticus, Dispharagus 277

elongata, Acuaria.. 259

Filaria.. 259

Rusguniella xm, 259

Spiroptera 259

elongatus, Dispharagus 259

Emberiza pecoris 177

tmmerezii, Spiroptera 325

engonium, Contracaecum x, 146,150

ensicaudata, Ascaris 139

Fusaria 139

ensicaudatum, Porrocaecum ix,

136, 1S9, 140, 142, 145

"Epervier" 355

Epomidiostomum 18, 19, 26-27, 28, 384

anatinum 27

orispinum v, 28-29, 384, 385

querquetulae 27,38-4

skrjabini 27, S84-S85

uncinatum v, 27-28, 384, 385

epops, Upupa 190,203

erythrorhynchos, Pelecanus 148,153,154

erythrorhynchus, Irrisor 188

Rhamphastos 186

Esox lucius 368

Eunematoda - 391

europaea, Talpa. 138

europaeus, Caprimulgus 129,130,313

Nycticorax 152

eurycerca, Cyrnea x, 167,168

euryoptera, Spiroptera 201

Viguiera.. XI, 201-202

Eurypyga helias. 362

Eurystomus afer 124, 127

Eustrongylides 358, 366, S67, 368, 369, 370, 371, 372

africanus x vn, 367, 369, 370

elegans 358,372

excisus X vn, 367, 370-371

ignotus. . . xvn, 367, 368, 371-372, 373

mergorum xvn,

335, 351, 358, 367, 372-373, 374
papillosum, xvn, 358, 367, 373-374, 375

perpapillatus xvn, 367, 374-376

species xvn, 369

tubifex xvi, 367-369,

370,371,372,373,374,375

Eustrongylus papillosum 370, 371, 373, 374, 379

tubifex 367,371

Euxenura maguari 50, 74

excisa, Cyrnea x, 167-163,169

Spiroptera 167

excisus, Eustrongylides xvn, 367, 370-37/

excubitor, Lanius 201

excubitor dodsoni, Lanius... 203

eximius, Platycercus 175

exulans, Diomedea 263,339

fabalus, Anser 20,62

falcinelli, Acuaria 249

Spiroptera. 249, 250

Falcinellus igneus 250

falcinellus, Ibis 250,376,381

Falco aesalon 138

albicilla 138

albicollis 177

apivorus .- 138,298

INDEX

447

Page

Falco ater 138,177,243

atricapillus 297,298

aurantius 177

biarmicus erlangeri 298,299

bidentatus 177

brachydactylus 138

brasiliensis 138

buteo. 138,177

cachinans 177,297,298

cayennsis 297,298

cenchris 182

chrysaetos 138

cineraceus 177, 277

coronatus 297,298

cyaneus 138, 177,276

degener 138

destructor 317

dispar 297,298

femoralis 280

furcatus 207

gallicus 138,297,299,300

gracilis 298,317

haliaetus 137,207

imperialis 138

lagopus 34,138,276

lanarius 138,177

lithofalco 138

longipennis 298

magnirostris..- 177,313

milvus 138

minutus 237,240,297

naevius 138,316

nisus 138, 177, 276, 277, 278, 299, 300

ornatus 297,298

palumbarius 138,177,288,300

palustris 297,298

pennatus 299,300

peregrinus 138

pygargus 144,299

rufus... 138,177,299

rutilans 138,297,299

species 297,299,305

subbuteo 177,273,275

subbuteus 299

swainsonii 297,299

tinnuncuhis 138,177,183,276

tridentatus 177,272,283

unicinctus 177,280,297,299,305

urubutinga 297,299

xanthothorax 177

falconis, Splroptera 280,317

falconis gavial realis, Spiroplera 317

leptopodis, Spiroptera 317

magnirostris, Filaria 313

falconis-subbuteonis, Dispharagus 275

Spiroptera 275

Jallaz, Spiroptera 276

fariai, Heterakis vii, 51,63

Ornithostrongylus v, 11, 12, 13-1J,, 15

fasciata, Aquila 138

Ascaridia vm, 80, 91-9t

fasciatus, Nisaetus 138

Palaeornis 175

Fasciola trachea 34

fasciolata, Crax 312,317

Felis catus domesticus 304

Page

femoralis, Falco.. 280

ferina, Nyroca 343

ferus, Anser 10

ferus domesticus, Anser 10

festiva, Chrysotis 80

festivus, Pionus (Psittacus).. 80

ficheuri, Habronema x.172, 174-176

Filaria... 203,233,244,258,272

aculeata 254

alata 281

anolabiata 317

anthuris 218,220,330

ca mpanulata 313

capitellata 220

cirrura 318

cystica 368,371

dacelonis.. 314.

depressa 221

dubia 338

elongata 259

falconis magnirostris 313

gallinarum.. 197,198

hamata 278

laticaudata 290

laticeps 273,276

leptoptera 176

mansoni... 325

muscicapae... 214

musculi _ 203

obvelata 235

papillijera 224

pulicis 343

quadriloba 215

recta 269

rotundata 195, 196, 198

spinijera 254

spinulosa 288

squamata 257

strumosa 164

triaenucha 271

tridentata 271

tulostom