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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
‘
PERE INCRE
ae ae Re '
Ke DIFF MiG MRE OD
: NGEANS NE
. x 5 SS MATIONNI 1S
aya
UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON
1927
Vy
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
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.
WasuineTon, 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
asa 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.
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14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25;
26.
27.
28.
29.
30.
OL.
oe
. Deletrocephalus dimidiatus. Head; a, lateral; 06, front view. After
LIST OF FIGURES OF NEMATODES
Trichostrongylus tenuis. Male bursa. 150. After Railliet, 1893.
. Trichostrongylus pergracilis. Bursa; lateral view. Original.
Trichostrongylus pergracilis. Dorsal and externo-dorsal rays of bursa.
Original.
Trichostrongylus pergracilis. Terminal genitalia of female. After Ship-
ley, 1909.
Ornithostrongylus quadriradiatus. Male bursa. After Stevenson, 1904.
Ornithostrongylus quadriradiatus. Telamon. After Stevenson, 1904.
Ornithostrongylus quadriradiatus. Tail of female. After Stevenson, 1904.
Ornithostrongylus fariai. Male bursa. After Travassos, 1921.
Ornithostrongylus dougiasi. Male bursa. After Theiler and Robertson,
1915.
Ornithostrongylus douglasi. Tail of female. After Theiler and Robert-
son, 1915.
Ornithostrongylus papillatus. Male bursa. After Linstow, 1882.
Ornithostrongylus hastatus. Male bursa. After Linstow, 1905.
Amidostomum anseris. Anterior end. After Railliet, 1893.
Amidostomum anseris. Male bursa. After Railliet, 1893.
Amidostomum raillieti. Anterior end. After Skrjabin, 1916.
Amidostomum raillicti. Bursa of male. After Skrjabin, 1916.
Amidostomum henryi. Anterior end. After Skrjabin, 1916.
Amidostomum henryi. Male bursa. After Skrjabin, 1916.
Amidostomum henryi. Tail of female. After Skrjabin, 1916.
Amidostomum chevreuvi. Anterior end. a, Dorsal view; b, lateral view.
After Seurat, 1918.
Amidostomum chevreuci. Male bursa. After Seurat, 1918.
Amidostomum chevreuri. Outline of bursa; lateral view. After Seurat,
1918.
Amidostomum monodon. Anterior end. After Linstow, 1882.
Epomidiostomum uncinatum. Anterior end. After Skrjabin, 1916.
Epomidiostomum uncinatum. Male bursa. After Skrjabin, 1916.
Bpomidiostomum uncinatum. Female tail. After Skrjabin, 1916.
Epomidiostomum orispinum. Male bursa. After Molin, 1861.
Codiostomum struthionis. Wead end. After Horst, 1885.
Codiostomum struthionis. Buccal capsule. After Horst, 1885.
Codiostomum struthionis. Male bursa, spread out. After Horst, 1885.
Codiostomum struthionis. Male bursa, lateral view. After Monnig, 1923.
Codiostomum struthionis. Female tail. After Monnig, 1923.
Diesing, 1857.
. Deletrocephalus dimidiatus. Male bursa. After Diesing, 1857.
Deletrocephalus dimidiatus. Female tail. After Diesing, 1857.
VI
36.
~
38.
39.
40.
41.
42.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Syngamus trachea. Outline. After Chapin, 1925.
7. Syngamus trachea. Head. a, Teeth in 8-toothed (normal) form; b, front
view of head; c, teeth in 9-toothed form. After Chapin, 1925.
Syngamus trachea. Spicules. After Chapin, 1925.
Syngamus trachea. Male bursa. After Chapin, 1925.
Syngamus microspiculum. a, Male and female; b, head end; c, male
bursa. After Skrjabin, 1916.
Syngamus parvus. Outline of pair. After Chapin, 1925.
Syngamus parvus. Spicules. After Chapin, 1925.
. Syngamus parvus. Dorsal portion of male bursa. After Chapin, 1925.
. Syngamus gracilis. Male and anterior portion of female. After Chapin,
1925.
. Syngamus gracilis. Male bursa. After Chapin, 1925.
. Sungamus gracilis. Female tail. After Chapin, 1925.
. Cyathostoma bronchialis. Head end. a, Side view; b, front view. From
Chapin, 1925, after Muehlig, 1884.
8. Cyathostoma bronchialis. Male bursa. From Chapin, 1925, after Muehlig,
1884.
. Cyathostoma tadornae. Male. From Chapin, 1925, after Chatin, 1874.
. Cyathostoma tadornae. Female tail. From Chapin, 1925, after Chatin,
1874.
. Cyathostoma lari. Female; entire worm. After Blanchard, 1849.
. Cyathostoma lari. Female; head end. After Blanchard, 1849.
. Cyathostoma lari. Vulva. After Blanchard, 1849.
54. Cyathostoma variegatum. Male bursa. From Chapin, 1925, after Linstow,
1890.
5. Cyathostoma variegatum. Head. a, Front view; 0b, side view. From
Chapin, 1925, after Linstow, 1890.
. Cyathostoma boularti. Head; side view. From Chapin, 1925, after Megnin,
1884.
. Cyathostoma boularti. Male bursa. From Chapin, 1925, after Megnin,
1884.
. Cyathostoma boularti. Head; front view. From Chapin, 1925, after
Megnin, 1884.
59. Cyathostoma americanum. Outline of female and male worms. After
(als
72.
73.
Chapin, 1925 (#=extent of uteri).
. Cyathostoma americanum. Head. a, Side view, male; 0, front view; ¢,
side view, female. After Chapin, 1925.
. Cyathostoma americanum. Male. a, Bursa; 06, gubernaculum. After
Chapin, 1925.
. Cyathostoma americanum. a, Egg; 6, vulva; e¢, tail (female). After
Chapin, 1925.
. Cyathostoma coscorobae. Buceal eapsules of female and male. After
Chapin, 1925.
. Cyathostoma coscorobae. Bursa. After Chapin, 1925.
66, and 67. Heterakis gallinae. 65, tail, lateral view; 66, tail, ventral
view ; 67, female tail. After Lane, 1917.
. Heterakis alata. Male tail. After Schneider, 1866.
and 70. Heterakis arquata. Male tail; 69, after Schneider, 1866; 70, after
Travassos, 19138.
Heterakis bancrofti. Male tail. After Johnston, 1912.
Heterakis beramporia. Male tail. After Lane, 1914.
Heterakis bosia. a, Male tail and point of right spicule; b, left spicule
from dorsum; ec, right spicule from side. After Lane, 1914.
99.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
110.
114.
112.
1138.
114.
115.
116.
LLG
118.
119.
120.
121.
422.
123.
Heterakis
5. Heterakis
Heterakis
Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
2. Heterakis
. Heterakis
. Heterakis
86. Heterakis interlabiata.
and
LIST OF FIGURES OF NEMATODES Vil
After Travassos, 1918.
After Gendre, 1911.
brasiliana. Male tail.
brevispiculum. Male tail.
brevispiculum. Spicule. After Gendre, 1911.
brevispiculum. Region of sucker. After Gendre, 1911.
gallinae. Region of sucker. After Gendre, 1911.
caudata. Male tail. After Linstow, 1906.
chenonettae. Male tail. After Johnston, 1912.
circumvallata. Male tail. After Linstow, 1906.
dispar. Male tail. After Railliet, 1898.
fariai. Male tail. After Travassos, 1913.
hamulus. Male tail. After Linstow, 1906.
85, anterior end; 86, male tail. After
Ortlepp, 1923.
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
. Heterakis
venter ;
Heterakis
Heterakis
Heterakis
Heterakis
Heterakis
Heterakis
inal.
Heterakis
Heterakis
Heterakis
Ascaridia
Ascaridia hermaphrodita.
Ascaridia galli.
Ascaridia
Ascaridia
Ascaridia
Ascaridia
Ascaridia
Ascaridia
Ascaridia
Ascaridia
Ascaridia
isolonche. Male tail. After Linstow, 1906.
neoplastica. Male tail. After Wassink, 1917.
longecaudata. a, Cuticle; 6, male tail. After Linstow, 1879.
macroura. Male tail. After Linstow, 1883.
papillosa. Male tail, ventral view. After Linstow, 1907.
papillosa. Male tail, lateral view. After Stossich, 1892.
parisi. Male tail. After Blanc, 1914.
psophiae. Male tail. After Travassos, 1913.
psophiae. Vulva. After Travassos, 1913.
pusilla. Male tail. After Linstow, 1906.
putaustralis, Male tail. After Lane, 1914.
putaustralis. a, Left spicule from side; 6, left spicule from
ec, right spicule. After Lane, 1914.
skrjabini. Male tail. After Skrjabin, 1916.
tenuicauda. Male tail. After Linstow, 1883.
tenuicauda. Male tail. After Seurat, 1918.
valvata. Male tail. After Travassos, 1913.
valvata. Male tail. After Schneider, 1866.
bonasae. Male tail. a, Ventral view; b, lateral view. Orig-
Vulva. Original.
Male tail. After Cobbold, 1861.
Female tail. After Cobbold, 1861.
Male tail. After Schneider, 1866.
Male tail. After Skrjabin, 1917.
After Smit, 1922.
After Baylis and Daubney, 1922.
Male tail. After Linstow, 1902.
Male tail. After Drasche, 1882.
After Linstow, 1898.
After Linstow, 1884.
After Magalhaes, 1892.
After Johnston, 1912.
After Linstow, 1903.
a, Ventral view; 3b,
bonasae.
acuticauda.
acuticauda.
hermaphrodita.
Male tail.
gall. Male tail.
aegyptiaca.
amblymoria.
australis. Male tail.
borealis. Male tail.
brasiliensis. Male tail.
catheturina. Male tail.
circularis. Male tail.
columbae. Male tail.
side view. After
Johnston, 1918.
Ascaridia compar.
Ascaridia cordata.
Ascaridia
Ascaridia
Male tail. After Linstow, 1899.
Male tail. After Linstow, 1906.
a, Head; 6, male tail. After Linstow, 1901.
Male tail. After Blome, 1909.
cristata.
cylindrica.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
. Ascaridia cylindrica. Spicule. After Blome, 1909.
. Ascaridia dolichocerca. Male tail. After Stossich, 1902.
. Ascaridia fasciata. Head. After Baylis, 1920.
. Ascaridia fasciata. Cuticular rings of body. After Baylis, 1920.
. Ascaridia fasciata. Male tail. After Baylis, 1920.
. Ascaridia francolina. Male tail. After Linstow, 1899.
. Ascaridia granulosa. Male tail. After Linstow, 1906.
. Ascaridia lineata. Male tail. After Boulenger, 1923.
. Ascaridia longecirrata. Male tail. After Linstow, 1879.
. Ascaridia magalhaesi. Male tail. After Travassos, 1913.
. Ascaridia magnipapilla. a, Head; b, male tail. After Linstow, 1906.
. Ascaridia numidae. Male tail. After Gendre, 1909.
. Ascaridia orthocerca. Male tail, ventral view. After Stossich, 1902.
. Ascaridia orthocerca. Male tail, lateral view. After Stossich, 1902.
. Ascaridia pterophora. Male tail. After Travassos, 1918.
. Ascaridia serrata. Male tail. After Schneider, 1866.
. Ascaridia strelnikowi. a, Head; b, male tail. After Skrjabin, 1916.
. Ascaridia stroma. Male tail. After Linstow, 1899.
. Ascaridia styphlocerca. Male tail. After Stossich, 1904.
. Ascaridia trilabium. Male tail. After Linstow, 1904.
. Ascaridia anseris. Male tail. After Schwartz, 1925.
. Pseudaspidodera pavonis. Head, lateral view. After Baylis and Daub-
ney, 1922.
. Pseudaspidodera pavonis. Male tail. Left, lateral view; right, ventral
view. After Baylis and Daubney, 1922.
. Pseudaspidodera pavonis. Vulva and vagina in lateral view, showing
cement plug in vulva. Arrow points in direction of head. After Baylis
and Daubney, 1922.
8. Subulura differens. Anterior end. Original.
9. Subulura differens. a, Vulva; b, female tail; ec, ovejector; d, egg. (Scale
the same for a and 6b.) Original.
. Subulura differens. Male tail. 37.5. After Barreto, 1918.
. Subulura acutissima. Head. a, Side view; b, front view. After Drasche,
1882.
. Subulura acutissima. Male tail. After Drasche, 1882.
3. Subulura allodapa. Anterior end. After Barreto, 1918.
. Subulura allodapa. Head, front view. After Drasche, 1882.
. Subulura allodapa. Male tail. After Drasche, 1882.
. Subulura allodapa. a and b, Ovejector. After Barreto, 1918.
. Subulura bentocruzi. Male tail, lateral view. After Barreto, 1918.
. Subulura bentocruzi. Male tail, ventral view. After Barreto, 1918.
. Subulura brumpti. a, Anterior end; b, male tail. After Lopez-Neyra,
1922.
. Subulura brumpti. Male tail. After Gedoelst, 1916.
. Subulura brumpti. Male tail. After Gendre, 1909.
. Subulura carlosi. Male tail. a, Side view; b, ventral view. After Bar-
reto, 1918.
. Subulura carlost. Vulva and ovejector. After Barreto, 1918.
. Subulura curvata. Male tail. After Linstow, 1883.
. Subulura forcipata. Male tail. After Barreto, 1918.
. Subulura forcipata. Vulva and ovejector. After Barreto, 1918.
. Subulura galloperdicis. Male tail. After Baylis and Daubney, 1922.
. Subulura halli. a, Male tail; 6, ovejector ; c, buccal cavity. After Seurat,
1914.
169.
170.
TT.
172.
175.
174.
175.
176,
GG
178.
179.
180.
. Subulura reclinata. Male tail. After Barreto, 1918.
. Subulura recurvata. Male tail. After Linstow, 1901.
. Subulura rima, Male tail. After Linstow, 1906.
. Subulura rimula. Male tail. After Linstow, 1903.
. Subulura seurati. a, Anterior end; 6b, male tail; ¢, ovejector. After
190.
191.
192
194.
195.
196.
197.
198.
199.
200.
201.
202.
203.
204.
205.
206.
207.
208.
209.
210.
211.
212.
213.
214.
LIST OF FIGURES OF NEMATODES Ix
Subulura leprincet. a, Buccal cavity; b, ovejector. After Seurat, 1914.
Subulura leprincei. a, Male tail, ventral view; 0, lateral view; c¢, spicule,
d, gubernaculum; e, egg. After Gendre, 1909.
Subulura lutzi. Anterior end. After Barreto, 1918.
Subulura lutzi. Male tail, ventral view. After Barreto, 1918.
Subulura lutzi. Male tail, lateral view. After Barreto, 1918.
Subulura lutzi. Vulva and ovejector. After Barreto, 1918.
Subulura noctuae. Male tail. After Seurat, 1914.
Subulura olympiot. Male tail. a, Ventral view; b, lateral view. After
Barreto, 1918.
Subulura olympioi. Ovejector. After Barreto, 1918.
Subulura papillosa. Male tail. From Barreto, 1918 after Drasche.
Subulura plotina. Malte tail. After Baylis, 1919.
Subulura poculum. Male tail. After Linstow, 1909.
Seurat, 1914.
. Subulura similis. Male tail. After Gendre, 1909.
7. Subulura strongylina. Male tail. After Travassos, 1913.
8. Subulura suctoria. Male tail. From Barreto, 1918 after Drasche, 1882.
. Subulura suctoria. Head, front view. From Barreto, 1918 after Drasche,
1882.
Subulura suctoria. Ovejector. After Barreto, 1918.
Subulura travassosi. Ovejector. After Barreto, 1918S.
and 193. Subulura travassosi. Male tail. After Barreto, 1918. (Figs. 191,
192, and 193 from Buceco chacuru.)
Subulura travassosi. From Bucco swainsoni. Male tail. After Barreto,
1918.
Subulura trogoni. Anterior end. After Barreto, 1918.
Subulura trogoni. Male tail. After Barreto, 1918.
Subulura acuticauda. Male tail. After Linstow, 1901.
Subulura gracilis. Head. After Linstow, 1899.
Subulura gracilis. Male tail. After Linstow, 1899.
Porrocaecum angusticolle. Dorsal lip. After Baylis and Daubney, 1922.
Porrocaecum angusticolle. Male tail. After Baylis and Daubney, 1922.
Porrocaecum depressum. Dorsal lip. After Baylis and Daubney, 1922.
Porrocaecum depressum. Male tail. After Linstow, 1875.
Porrocaecum ensicaudatum. a, Dorsal lip; b, male tail; c, egg. After
Linstow, 1884.
Porrocaecum reticulatum. Male tail. After Baylis and Daubney, 1922.
Porrocaecum reticulatum. Dorsal lip. After Linstow, 1899.
Porrocaecum semiteres. Anterior end (dissected). After Creplin, 1829.
Porrocaecum serpentulus. Male tail. After Monnig, 1923.
Porrocaecum serpentulus. Head. After Linstow, 1899.
Porrocaecum serpentulus. Male tail. After Baylis and Daubney, 1922.
Porrocaecum spirale. Dorsal lip. After Linstow, 1875.
Porrocaecum spirale. Egg. After Linstow, 1875.
Porrocaecum spirale. Male tail. After Linstow, 1875.
Porrocaecum heteroura. Anterior end (dissected). After Creplin, 1829.
215a. Porrocaecum anguillae. Anterior end (dissected). After Linstow, 1899.
3612—27. 2
215.
216.
217.
218.
219.
221.
222.
223.
224.
225.
226.
239.
240.
241.
242.
243.
244.
245.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
btoe. Porrocaecum kirghisensis. 0b, Lip; c, interlabium; d, lateral and e,
ventral view of male tail. After Skrjabin, 1916.
Contracaecum spiculigerum. Head. After Linstow, 1909.
Contracaecum spiculigerum. a, Esophagus and appendices; 6, male tail.
After Schneider, 1866.
Contracaecum andersoni. a, Dorsal lip; b, interlabium; c, vulva; d, male
tail. After Vevers, 1923.
Contracaecum engonium. a, Head; b, male tail. After Baylis and Daub-
ney, 1922.
. Contracaecum haliaéti. a, Lateral view; b, front view of head; ¢, egg.
After Smith, Fox, and White, 1908.
Contracaecum haliaéti. a, Inner surface; b, outer surface of lip. After
Smith, Fox, and White, 1908.
Contracaecum microcephalum. a, Front view of head; 6b, dorsal view of
head; c, male tail; d, free end of spicule; e, cross section of spicule.
After Gedoelst, 1916.
Contracaecum micropapillatum. Anterior end (dissected). Original.
Contracaecum micropapillatum. a, Male tail; 6, lip and interlabia. After
Stossich, 1890.
Contracaecum multipapillatum. Digestive organs of male. After Skrjabin,
1916.
Contracaecum multipapillatum. a, Head; 6b, male tail. After Drasche,
1882.
. Contracaecum ovale. Head. After Linstow, 1907.
. Contracaecum praestriatum. Anterior end. After Monnig, 1923.
. Contracaecum praestriatum. Male tail. After Monnig, 1923.
. Contracaecum punctatum. Male tail. After Gedoelst, 1916.
. Contracaecum rodhaini. Head. After Gedoelst, 1916.
. Contracaecum rosarium. Anterior end. After Connal, 1912.
. Contracaecum rosarium. Male tail. After Baylis and Daubney, 1922.
. Contracaecum scotti. a, Esophagus and appendices; 6, head; c, lateral
view and d, ventral view of male tail. After Leiper and Atkinson, 1915.
. Contracaecum tricuspe. a, Male tail; b, dorsal view and e¢, front view of
head. After Gedoelst, 1916.
. Contracaecum magnipapillatum. Male tail. After Chapin, 1925.
. Spirura uncinipenis. a, Lateral view; b, front view of head; c, male
tail. After Drasche, 1884.
. Spirura uncinipenis. a, Head; b, egg; c, free end of short spicule. After
Linstow, 1899.
Cyrnea eurycerca. a, Female tail, showing ovejector; 6, male tail; e,
male (length 7.6 mm); @, lateral view and e, ventral view of head.
After Seurat, 1914.
Cyrnea excisa. a, Dorsal view; 6b, lateral view of head end; c, female;
d, male tail. After Seurat, 1914.
Cyrnea seurati. Tail end of male and of female. After Lopez-Neyra, 1918.
Cyrnea colini. Head. a, Ventral; b, front; c¢, oblique lateral view.
Original.
Cyrnea colini. a, Male tail; b, female tail; c, ovejector. (Scale for b and
ce the same.) Original. A
Habronema colaptes. a, Dorsal; 6, lateral; c, front view of head; d,
spicules and gubernaculum; e, male tail; f, female tail. After Walton,
1928.
Habronema ficheuri. a, Lateral; b, dorsal view of head; ¢c. male tail.
After Seurat, 1916.
246.
247.
248.
249.
250.
251.
252.
253.
254.
255.
256.
258.
259.
260.
261.
262.
263.
264.
265.
266.
267.
268.
269.
270.
271.
272.
LIST OF FIGURES OF NEMATODES XI
Habronema inecrita. 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.
Habronema leptoptera. a, Anterior end, dorsal view; 6, male tail. After
Seurat, 1914.
Habronema leptoptera. a, Head, lateral view; b, receptacle seminale, join-
ing oviduct with uterus; c, ovejector. After Seurat, 1914.
Habronema longistriata. a, Head, ventral view; b, head, front view; ¢,
male tail. After Drasche, 1884.
Habronema mansioni. a, Anterior end, ventral view; 6, male tail; c, ove-
jector. After Seurat, 1914.
Habronema monoptera. a, Anterior end; b, lateral lip; c, median lip; 4d,
male tail; e, left spicule; f, right spicule; g, female tail. After Gendre,
1922,
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.
Habronema spinosa. a, Spines of cuticle; 0, head end; c, median lip; @,
male tail; e, left spicule; f, right spicule; g, gorgeret; h, female tail.
After Gendre, 1922.
Habronema tulostoma. a, Anterior end; 6, lateral lip; c, dorsal lip.
After Seurat, 1914.
Habronema tulostoma. a, Male tail; b, cloacal region, showing gorgeret ;
c, ovejector. After Seurat, 1914.
Habronema unilateralis. a, Head, front view; vb, male tail After
Drasche, 1884.
. Hadjelia lhuillieri. a, Head end, lateral view; b, head, dorsal viery.
After Seurat, 1916.
Hadjelia lhuillierit. Vulva and ovejector. After Seurat, 1916.
Hadjelia inermis. a, Male tail as usually seen (most posterior papillae
hidden) ; 6, male tail straightened out; ¢, left spicule; d, right spicule;
e, female tail. After Gendre, 1922.
Hadjelia parva. a, Male tail; b, female tail. After Gendre, 1922.
Hadjelia truncata. a, Head; b, male tail. After Mueller, 1897.
Hartertia obesa. a, Dorsal; b, lateral; c, front view of head; d, egg.
After Seurat, 1915.
Hartertia obesa. a, Ovejector; b, male tail. After Seurat, 1915.
Hartertia annulata, a, Lateral; b, front; c, dorsal view of head; d, egg;
e, female tail; f, ovejector; g, enlarged view of cul-de-sac at union of
sphincter and trompe. Original.
Hartertia annulata. Male tail and spicules. Original.
Hartertia confusa. Male tail. After Linstow, 1906.
Hartertia gallinarum. a, Lateral; b, front view of head; ec, female tail.
Original.
Hartertia gallinarum. Ovejector and uteri. Original.
Hartertia gallinarum. Male tail and spicules. Original.
Hartertia rotundata. a, Head, ventral view; b, male tail. After Seurat,
1914,
Hartertia rotundata. Ovejector. After Seurat, 1914.
Harteria zorillae. Third stage larva. a, Ventral view of head; Db, lateral
view of head; c, ventral; d, lateral view of tail. After Seurat, 1919.
. Viguiera euryoptera. a, Male tail; 6b, ovejector. After Seurat, 1913.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
. Gongylonema ingluvicola. a, Head; b, male tail. After Ransom, 1904.
. Spiroptera penihamata. Male tail. After Drasche, 1884.
. Spiroptera turdi. Head. a, Front; b, lateral view. After Drasche, 1884.
. Desmidocerca acrophila. a, Head; b, male; ¢, male tail; d, female tail.
After Skrjabin, 1916.
8. Acuaria ptilopachydis. a, Anterior end; b, female tail. After Gendre,
1920.
. Acuaria quadriloba. Head, front view. After Schneider, 1866.
. Acuaria tarentolae. Anterior and posterior ends of third stage larva.
After Seurat, 1916.
. Acuaria anthuris. Male tail. a, Cross section; b, ventral view. After
Schneider, 1866.
. Acuaria anthuris. a, Male tail X about 50; b, female tail end. After
Linstow, 1873.
3. Acuaria anthuris. Ovejector. After Seurat, 1915.
. Acuaria cordata. a, Head end; b, lateral and c, ventral view of male tail.
After Mueller, 1897.
5. Acuaria depressa. Male tail. After Schneider, 1866.
. Acuaria gracilis. a and b, Anterior end; c, left and d, right spicule; e,
male tail; f, female tail. After Gendre, 1912.
. Acuaria ornata. a, Male tail; b, left spicule; ec, head end; d, right
spicule; e, cordons of the lateral and /f, of the submedian fields. After
Gendre, 1912.
. Acuaria papillifera. ead and tail ends of male. After Linstow, 1878.
. Acuaria subula. a and b, Head end; ec, male tail; d, right and e, left
spicule. After Gendre, 1913.
. Cheilospirura hamulosa. a, Cordon; b, head, lateral view; c, male tail.
After Drasche, 1888.
. Cheilospirura gruveli. 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.
2. Cheilospirura rotundata. Male tail. After Linstow, 1907.
293. Cheilospirura spinosa. a, Anterior end; 6, part of cordon; c, vulva; d,
294.
295.
296.
299.
300.
301.
302.
303.
304.
305.
306.
female tail; e, spicule. Original.
Cheilospirura spinosa. Male tail. Original.
Chevreuxia revoluta. Anterior end. After Seurat, 1918.
Cosmocephalus diesingi. a, Anterior and middle parts of worm, showing
position of vulva; 6, anterior end; c, female tail. After Molin, 1861.
. Cosmocephalus aduncus. a, Head end; 0b, male tail. After Stossich,
1892.
. Cosmocephalus obvelatus. a, Dorso-ventral view and Db, lateral view of an-
terior end; c, female tail, ventral view. After Seurat, 1919.
Cosmocephalus obvelatus. Male tail. After Linstow, 1877.
Dispharyne nasuta. Head and tail of male. After Dujardin, 1845.
Dispharyne spiralis. Male. After Piana, 1897.
Dispharyne spiralis. Female. After Piana, 1897.
Dispharyna spiralis. a, Head end. After Seurat, 1916. 6, Ovejector.
After Seurat, 1919.
Dispharyneg laplantei. Ovejector. After Seurat, 1919.
Dispharynze magnilabiata. Male tail. After Drasche, 1884.
Dispharynx noctuae. a, Vulva; b, anterior end, dorsal view; c, female;
d, anterior end, lateral view; e, ovejector and vulva. After Seurat,
1913.
309.
310.
312.
813.
314.
315.
516.
317.
318.
319.
520.
321.
322.
“ojo
var.
824.
328.
329.
350.
LIST OF FIGURES OF NEMATODES XIII
. Dispharyne rectovaginata. Head end and tail of male. After Drasche,
1884.
. Echinuria jugadornata. a, Cordon. After Solovievy, 1912. b, Head end;
c, male tail. From Skrjabin, 1916, after Soloviev, 1912.
Echinuria uncinata. Head end; ventral view. After Seurat, 1919.
Echinuria uncinata. a, Head end, dorsal view; b, cross-section and e,
ventral view of male tail. After Schneider, 1866.
. Bchinuria . a, Front view c eral view of head; ec, femal
Echinuria ardeae. a, Front view and b, lateral view of head; c¢, female
tail. After Smith, Fox, and White, 1908.
Echinuria calearata. a, Lateral and 0b, ventral view of head; ec, female
tail. After Drasche, 1884.
Echinuria contorta. a, Anterior end; 6, male tail; ¢, female tail, showing
vuiva and anus. After Molin, 1861.
Echinuria 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.
Echinuria hargilae. a, Anterior end; 6, right spicule. After Baylis and
Daubney, 1922.
Echinuria horrida. Head end. After Schneider, 1866.
Echinuria horrida. Head end. After Stossich, 1891.
Echinuria leptoptili. a, Head end; 6, male tail; ec, left spicule; d, right
spicule (the small drawing showing orifice at free end dilated on
pressure) ; e, female tail. After Gendre, 1920.
Echinuria longevaginaia, a, Head; 6, male tail. After Drasche, 1884.
Lehinuria phoenicopteri. Immature female. @ and b, Head end; ¢, tail;
d, vulva and ovejector. After Seurat, 1916.
Lehinuria squamata, a, Anterior end; b, deta’l of cordon. After Linstow,
1883.
Rusguniella elongata. Anterior end. a, Lateral view; b, ventral view.
After Seurat, 1919.
Rusguniella vanelli. Anterior end, ventral view. After Linstow, 1884.
Sciadiocara unibellifera. Wead. a, Dorso-ventral view; b, front view.
After Drasche, 1884.
. Sciadiocara umbellifera. a, Male tail; b, vulva; c, female tail. After
Skrjabin, 1916.
. Seuratia shipleyi. a, Dorso-ventral and b, lateral view of anterior end; ©,
male tail; @, female tail; e, egg. After Stossich, 1900.
. Seuratia shipleyi. a and b, Proximal region of ovejector ; ¢, distal region
of ovejector. After Seurat, 1916.
Streptocara pectinifera. a, Head end; b, cervical papillae; ¢, male tail.
After Neumann, 1900.
Streptocara cirrohamata. a, Head; 6, male tail. After Linstow, 1888.
Streptocara crassicauda. a, Head; 6, tail of female; ¢c, vulva. After
Skrjabin,1916.
. Streptocara crassicauda charadrii. a, Head; b, cervical papiila. After
Skrjabin, 1916.
. Streptocara decora. a, Anterior end; b, male tail. After Dujardin, 1845.
. Streptocara recta. a, Dorso-ventral and b, lateral view of head; ¢, male
tail; d, spicules. After Mueller, 1897.
Streptocara triaenucha. Cervical papilla. After Wright, 1897.
. Streptocara tridentata, a, Male tail. After Linstow, 1909. b, Cervical
papilla. After Linstow, 1877.
. Diagram of cordons (as found in Synhimantus) as they would appear
if spread out horizontally. After Schneider, 1866.
339.
340.
341.
342.
343.
344.
345.
35D.
356.
357.
358.
359.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
. Synhimantus affinis. a, Female genitalia. After Seurat, 1915. 6, Head
end. After Seurat, 1916.
8. Synhimantus brevicaudata. a, Head end; b, female tail. After Skrjabin,
1917.
Synhimantus denticulata. Head end. After Dujardin, 1845.
Synhimantus laticeps. Male tail. After Seurat, 1916.
Synhimantus laticeps. Head. After Schneider, 1866.
Synhimantus laticeps. Female genitalia. After Seurat, 1920.
Synhimantus elliptica. Male tail. After Molin, 1861.
Synhimantus hamata. a, Head end. After Linstow, 1879. 6, Male tail.
After Linstow, 1877.
Synhimantus invaginata. a and b, Head end; c, male tail; d, right and
e, left spicule; f, female tail. After Gendre, 1913.
. Synhimantus recta. Male tail. After Drasche, 1884.
. Synhimantus sagittata. a, Head end; 0b, male tail. After Schneider,
1866.
. Synhimantus subrecta. a, Male tail; b, right spicule; c, profile view and d,
ventral view of left spicule; e, female tail. After Gendre, 1921.
. Schistorophus longicornis. a, Lateral view and b, front view of head; ce,
male tail. After Schneider, 1866.
. Schistorophus acanthocephalicus. Head. After Drasche, 1884.
. Schistorophus aulieatina. Head end. After Skrjabin, 1916.
. Schistorophus bidens. Male tail. After Schneider, 1866.
3. Schistorophus bidens. a, Head; b, male tail. After Lepri, 1898.
. Histiocephalus laticaudatus. a, Front view and b, lateral view of head; ec,
male tail. After Drasche, 1884.
Histiocephalus laticaudatus. Male. After Skrjabin, 1916.
Histiocephalus tridens. a, Front view and b, profile view of head; c, male
tail; d, anterior end; e, posterior end, lateral view; /, 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.
Serticeps vulwoinjflata. a, Head; 6b, male tail. After Drasche, 1884.
Serticeps vulvoinflata (probably). Labelled Spiroptera mediospiralis but
apparently incorrectly so. See text for discussion. After Drasche,
1884.
Physaloptera acuticauda. a, Terminal part of female genitalia; b, male
tail. After Ortlepp, 1922.
. Physaloptera alata. a, Head; b, lip; c, male tail. After Linstow, 1877.
. Physaloptera alata Rudolphi of Ortlepp, 1922. Possibly equivalent to
P. galinieri Seurat. See text for discussion. After Ortlepp, 1922.
. Physaloptera megalostoma. After Creplin, 1829.
. Physaloptera crosi. a, Head end; b, male tail; ec, female tail. After
Seurat, 1914.
. Physaloptera fusiformis. Male tail. After Linstow, 1902.
. Physaloptera galinieri. a, Head, lateral view; b, anterior half of body
of female, showing ovejector; c, male tail. After Seurat, 1914.
. Physaloptera gemina. Male tail. After Linstow, 1899.
. Physaloptera subalata. a, Lip; b, male tail. After Schneider, 1866.
. Physaloptera truncata. a, Lip; b, male tail. After Schneider, 1866.
. Physaloptera brevicauda. Male tail. After Linstow, 1909.
. Physaloptera malleus. a, Head; b, male tail. After Linstow, 1883.
. Physaloptera, species. Head end. After Parona, 1885.
372.
373.
ot.
375.
376.
384.
385.
386.
387.
388.
389.
390.
391.
392,
393.
394.
395.
396.
397.
398.
399.
400.
401.
402.
LIST OF FIGURES OF NEMATODES XV
Physaloptera ovata. Male tail. After Linstow, 1907.
Physaloptera bulbesa. Male tail. After Linstow, 1906.
Thelazia campanulata. a, Head end; b, male tail; c, female tail. After
Travassos, 1918.
Thelazia cholodkowskii. Female. a, Head end; 0b, cuticular annulations;
ec, tail. After Skrjabin, 1922.
Thetazia dacelonis. a, Head end, showing vulva; 0, tail of female; ec, tail
of male. After Breinl, 1913.
. Thelazia lutsi. Female. a, Head end, showing vulva; 0b, tail. After
Travassos, 1918.
. Thelazia tutzi. Male tail. After Travassos, 1918.
. Thelazia stereura. Female. a, Head end, showing vulva; }b, tail. After
Skrjabin, 1922.
. Thelazia papillosa. Wead; front view. After Drasche, 1884.
. Thelazia anolabiata. Head. After Drasche, 1884.
. Thelazia (?) cirrura. Head end, showing vulva. Original.
. Thelazia (?) digitata. Female. a, Head end, showing vulva; b, tail.
After Travassos, 1918.
Ceratospira vesiculosa. Male tail. After Schneider, 1866.
Ceratospira ophthalmica. a, head end; b, male tail. After Linstow, 1898.
Oxryspirura cephaloptera. a, Male tail; 6, head, front view. After
Drasche, 1884.
Oxryspirura anacanthura. a, Right spicule; 6, male tail. After Drasche,
1884.
Oxryspirura anthochaerae. a, Head end; b, tail end of female. After
Johnston, 1912.
Oxryspirura mansoni. a, Dorsal view and b, front view of head; ec, egg;
d, embryo escaping from the egg. (c.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.
Oxyspirura mansoni. Posterior end of female. (int., intestine; ov.,
ovary; ret., rectum; vg., vagina; vul., vulva; @., cells Surrounding rec-
tum.) After Ransom, 1904.
Oxyspirura mansoni. 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.
Oxyspirura ophthalmica. a, Head; 6b, male tail. After Linstow, 1903.
Oxyspirura parvovum, a, Head; b, male tail. After Sweet, 1910.
Oxryspirura parvovum. Posterior end of female. After Sweet, 1910.
Oxryspirura siamensis. Male tail. After Linstow, 1903.
Oxyspirura sygmoidea. Male tail. After Skrjabin, 1916.
Oxryspirura tanasijtchuki. a, Head; 6b, right spicule; c, male tail. After
Skrjabin, 1916.
Oxyspirura brevipenis. Male tail. After Drasche, 1884.
Tetrameres paradora. a, Female, natural size; 6, lateral view, and ec,
view from above, of same, enlarged; d, egg. Enlarged. After Diesing,
1835.
Tetrameres paradora. a, Head (whether of male or female not stated) ;
b, male tail. After Drasche, 1884.
Tetrameres americana. Posterior end of male. Original.
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
403.
404.
405.
406.
407.
408.
409.
410.
411.
412.
413.
414,
415.
416.
417.
418.
419.
420.
421.
422.
423.
424.
425.
426.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Tetrameres coccinea. Female. a, Total; b, anterior extremity; c, ovejec-
tor; d, seminal receptacle at junction of uterus and oviduct. After
Seurat, 1914. ,
Tetrameres cochleariae. Ovejector. After Travassos, 1919.
Tetrameres confusa. a, Female; b, male tail. After Travassos, 1919.
Tetrameres dubia. Male. After Travassos, 1919.
Tetrameres fissispina. a and b, Head end of male, lateral view (a), dorsal
view (0b), showing the two bifid cephalic spines and the simple body
spines; c, head end of female, dorsal view. After Seurat, 1920.
Tetrameres fissispina. Head and tail of male. After Travassos, 1919.
Tetrameres fissispina. a, Ovejector; b, copulatory receptaculum; c, fe-
male genital system. After Seurat, 1914. d, Tail of larva, ventral
view. After Seurat, 1919.
Tetrameres gigas. Anterior and posterior ends of male. After Travassos,
1919.
Tetrameres gynaecophila. a, Head of female; 0b, tail of male. After
Seurat, 1915.
Tetrameres micropenis. a, Tail of male; b, ovejector, showing striated
appearance of musculature. After Travassos, 1919.
Tetrameres nouveli. a, Female; b, ovejector; ¢, seminal receptacle and
oviduct ; d, egg with filaments; e, head end of male; f, tail end of male.
(500u scale for a to ¢; 100u scale for d to f.) After Seurat, 1914.
Tetrameres tetrica. Male. After Travassos, 1919.
Microtetrameres cruzi. a, Female; b, male. After Travassos, 1914.
Microtetrameres contorta. a and b, Males; ec, 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. j, Wax reconstruction of female. After
Weidman, 1923.
Microtetrameres helix. a, Male tail; b and c, females; d, female tail.
Original.
Microtetrameres inermis. a, Male; b, ventral view and e, lateral view of
tail of same; d, head end of same. (Scale applies to b,c, and d.) After
Seurat, 1913.
Microtetrameres inermis. Female. a, Adult; 0, adult examined perpen-
dicularly to determine manner of coiling; c, immature; d, anterior end.
After Seurat, 19138.
Microtetrameres inermis. a, Ovejector. After Seurat, 1914. b, Part of
same; ¢, ovary, oviduct, and initial region of uterus; d, vulva; e and f,
hatching of eggs. After Seurat, 1913.
Microtetrameres minima. Male. After Travassos, 1914.
Microtetrameres pusilla. Male. After Travassos, 1919.
Microtetrameres spiralis. Head and tail of male. After Seurat, 1915.
Ancyracanthopsis bihamata. a and b, Head end; ec and ad, male tail;
e, spicule; f, female tail; g, vulva. After Mueller, 1897.
Gnathostoma accipitri. a, General appearance; b, hooks of cephalic
region. After Skrjabin, 1916.
Gnathostoma pelecani. a, General appearance; b, glandular apparatus;
c, vulva. After Chatin, 1874.
. Eustrongylides tubifex. a, Male tail; b, head end; c, female tail. After
Jaegerskiold, 1909.
428.
429.
430.
431.
432.
433.
434.
435.
436.
440.
441.
442.
443.
444.
LIST OF FIGURES OF NEMATODES XVII
Eustrongylides larvae. Species not determined. a, Front view of head:
b, anterior end; ¢, outline of male tail; d, outline of female tail. After
Ciurea, 1924.
Eustrongylides africanus. a, Head end; b, female tail; ¢, exterior of egg;
d, pitting of shell; e, interior of egg. After Jaegerskiold, 1909.
Eustrongylides excisus. a, Head; b and ¢, tail of male. After Jaegers-
kiold, 1909.
Eustrongylides ignotus. a, Head; b, female tail; ¢ and d, male tail; e
and f, egg. After Jaegerskiold, 1909.
Eustrongylides mergorum. a, Female; 6, male; natural size; c, head.
After Linstow, 1877.
EBustrongylides mergorum. a, Front view; b, lateral view of head; c, egg.
After Jaegerskiold, 1909.
Eustrongylides mergorum. a, Head end; b, female tail; ¢ and d, male
tail. After Jaegerskiold, 1909.
Eustrongylides papillosus. Male tail. After Jaegerskiold, 1909.
Bustrongylides perpapillatus. a, Front view; 6, lateral view of head;
c, male tail; d, female tail; e and f, eggs. After Jaegerskiold, 1909.
. Hystrichis tricolor. Head and tail ends of female. After Jaegerskiold,
1909.
. Hysirichis acanthocephalicus. a, Side view; b, front view of head; c and
d, male tail; e, female tail; f, egg. After Jaegerskiold, 1909.
. Hystrichis coronatus. a, Head end; b, female, natural size; c, female
tail. After Molin, 1860.
Hystrichis cygni. Head end. After Molin, 1861.
Hystrichis neglectus. a, Head end; b, male tail; c, female tail; d, egg.
After Jaegerskiold, 1909.
Hystrichis orispinus. Head end. From Jaegerskiold, 1909, after Molin.
Hystrichis varispinosus. a, Head end; b, spine of anterior cervical region.
After Jaegerskiold, 1909.
Hystrichis wedli (Linstow’s material). a, Anterior portion of body; Bb,
ohe of the most anterior spines, lateral view; c, same, front view;
d, one of the most posterior spines; e, posterior end of body (., probable
primordium of vagina). After Jaegerskiold, 1909.
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BIRD PARASITES OF THE NEMATODE SUBORDERS
STRONGYLATA, ASCARIDATA, AND SPIRURATA
By Exotsr B. Cram
Of the Bureau of Animal Industry, United States Department of Agriculture
INTRODUCTION
The present work 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-cel:
sarcoma of pheasants has been attributed to species of Weterakis,
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
1
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
avaliable 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 new 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 3
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 layer 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), ovoviviparous, 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 in or otherwise im
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
MOUE See soe SE ee 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 SUBGRDERS OF MYOSYRINGATA
1. Heterogamie 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).
Homogamie forms, the parasitic generation consisting of males and
TEMiales SAAS STR) NEE SEE Pe ee ee eee 2...
:
NEMATODE PARASITES OF BIRDS 5
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.
8. Heteroxenous forms, the larval stages occurring in various intermediate
hosts; mouth usually with 2 lips, or a larger number of lips or lobes
eapable of resolution into 2 basic lips, or without lips; oviparous, vivi-
DALOUS; OL: OVOVIVIPATOUS:—. ==! .2---s* 22-8 Spirurata, p. 162.
Monoxenous forms as a rule, exceptional forms (seal ascarids) with inter-
mediate hosts; mouth usually with 3 lips, exceptionally without lips:
OVI DATO US Ets FE AER ENT ee ee as Se etd ee AD Mee Se Oe 4.
. Meromyarian; males with 1 spicule, at times reduced, imperfectly chitinized,
or absents)—ssseiecusl ues 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 1918 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
8 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 (Syngamus), circulatory system (Strongylus), or tissues
(Stephanurus) as adults or agamic forms, and often in such locations as
larvae or agamiec individuals; bursa typical, relatively large, and with the
dorsal ray approximately as wide as other members of the bursa rays. 2.
2. Bueeal capsule present; usually relatively thick forms; usually in digestive
tract, sometimes in respiratory system or tissue as adults.
Strongyloidea, p. 29.
Buceal eapsule absent or present as a much-reduced structure (Amidos-
tomum) ; usually relatively slender and sometimes hair-like forms; always
INV GiZeStive SV SUCM ae oe Sas Se eee re 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 buecal 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
Superfamily 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
Family diagnosis——Metastrongyloidea (p. 7): Characters of the
superfamily.
Type genus.—Metastrongylus Molin, 1861.
It has seemed advisable to create this new superfamily to balance
tlie 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.
KEY T6 FAMILIES OF TRICHOSTRONGYLOIDEA
1. Body frequently spirally curled. Females with 1 ovary.
Heligmosomidae, p. &.
Body. .not spirally. curled. Females with 2 ovaries____._._.-__----------_- 2:
2. A small buccal capsule present, or if rudimentary (absent?) the head has 4
papillaey and, epauletso-4!.5- 2224.- 22-14 a- Amidostomidae, p. 17.
Head simple; no buceal capsule and cephalic structures as above.
Trichostrongylidae, p. 8.
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 Railliet and Henry, 1909.
Subfamily HELIGMOSOMINAE Travassos, 1914¢
Subfamily diagnosis.—Heligmosomidae (p. 8): Characters of the
family.
Type genus.—Heligmosomum Raillet 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
ubsent. 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
and simple. Parasites of the digestive system.
Type genus.—Trichostrongylus Looss, 1905.
Subfamily TrrcHostroncyiiIna® Leiper, 1908
Subfamily diagnosis——Trichostrongylidae (p. 8): Characters of
the family.
Type genus.—T richostrongylus Looss, 1905.
Tt will undoubtedly be necessary in the near future to break up
the new superfamily Trichostrongyloidea into a number of families
and 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 OF TRICHOSTRONGYLINAB
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 acspiculescuides=o 2a ee eee a 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 6 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. /emale 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.
KBY TO SPECIES OF TRICHOSTRONGYLUS
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 4 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
1/, of body length from tail end___.______ Trichostrongylus tenuis, p. 10.
TRICHOSTRONGYLUS PERGRACILIS (Cobbold, 1873) Railliet and Henry, 1909
Synonym.—sStrongylus pergracilis Cobbold, 1873.
Host.—Colinus virginianus and Lagopus 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 6 to 8 mm. long. Bursa (figs. 2 and 3) with 2 lateral lobes
and a small sharply pointed median lobe. The latero-ventral ray
curves 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. EExterno-
dorsal rays (50 in length) are longer than the dorsal ray (36 in
10 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
length). The dorsal ray spht 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) 14 to 14 of body length from posterior end, the vulva
a transverse slit with crenelated edges. Eggs 72 to 75u long by 46u
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 360p long by 15p 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 4602 long, which
Fics. 1-4.—1, TRICHOSTRONGYLUS TENUIS. MALE BURSA. 150. ArrTHuR RAIL-
LIbT, 1893. 2, TRICHOSTRONGYLUS PERGRACILIS. BURSA; LATERAL VIEW.
ORIGINAL. 3, SAME. DORSAL AND EXTERNO-DORSAL RAYS OF BURSA. ORIGINAL.
4. SAMBP, TERMINAL GENITALIA OF FEMALE. AFTER SHIPLEY, 1909)
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 (Mehlis, 1846) Railliet and Henry, 1909
Synonyms.—Strongylus tenuis Mehlis, 1846 (in Creplin, 1846) ;
Strongylus serratus Linstow, 1876.
Hosts —Anas boschas, A. boschas domesticus, Anser albifrons, A.
anser, A. cinereus, A. cinereus domesticus, A. domesticus, A. ferus,
NEMATODE PARASITES OF BIRDS 11
A. ferus domesticus, Gallus gallus, Otis tarda, Perdix cinerea, and
Phasianus colchicus.
Location.—Cecum and small intestine.
Morphology —Trichostrongylus (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 margin. 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 shghtly 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 (112p long), twisted, and slightly dissimilar; there
is an elongate lemon-shaped gubernaculum, 63, lone.
Female 7.3 to 7.8 mm. long. The tail terminates in a sharp cone.
The vulva is near the posterior end, about + of the body length from
the tip of the tail. The eggs are elliptical, 65 to T5y long by 35 to
42 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 North America (collected and determined by Foster
from A. cinereus domesticus, Washington, D. C., in 1910, but not
reported until the present time).
Genus ORNITHOSTRONGYLUS Travassos, 1914b
Synonym.—C ephalostrongylus Irwin-Smith, 1920.
Generic diagnosis —YTrichostrongylinae (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
the 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 distally. ‘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, 19140.
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 long. Anus 80u and vulva 800u from tip of tail. In ostrich.
Ornithostrongylus douglasi, p. 14.
Males more than 6 and females more than 8 mm. long_______ -_----_--_--- as
. Males Jess than 7 mm. and females less than 9 mm. long. Spicules 140
long, with 2 spines at their anterior third. Eggs 80u long. In Otis tarda.
Ornithostrongylus papillatus, p. 16.
Males more than 8 mm. and females more than 16 mm. long. Spicules 150 to
285u lone... DegsinotoOvieriw oaulomg see og bee ce ee 3
. Spicules 220u long. Eggs 30u by 21u. In Europe.
Ornithostrongylus hastatus, p. 17.
Spicules less than 200u or more than 2754 long. Beggs not less than 7T0u by
38u. In Western Hemisphere and Australia____--~--~------------s---= 4
4. Male up to 12 mm. and female up to 24 mm. long. Anus 400 from tip of
tail. Spicules 150 to 160u long. From pigeon; United States and
AUSbralias so-so eee ee Ornithostrongylus quadriradiatus, p. 12.
Male not over 10 and female not over 20 mm. long. Anus 250 to 2844 from
tip of tail. Spicules 285 long. From Leptotila rufazilla; Brazil.
Ornithostrongylus fariai, p. 13.
bo
Oo
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—Columba 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 freshly 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 160u 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 70p
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.
Fics. 5—-7.—ORNITHOSTRONGYLUS QUADRIRADIATUS. 5, MALE BURSA. 6,
TELAMON, 7, TAIL OF FEMALE, ALL AFTER STEVENSON, 1904
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 1404 from the end of the tail. Eggs 70 to
75p long by 38 to 40n wide, and apparently may be segmenting or
contain embryos when deposited. In normal saline eggs may hatch
in the uterus. ,
Larvae, first stage, 500p 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. from the end of body. Mouth with a slight dis-
tinct chitinous armature. Excretory pore 260. from head end.
Esophagus short, 350 to 450p long, slightly dilated posteriorly.
Male 9 to 10 mm. long by 110» 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» 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 T1p long by 21p 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. from tip to tail. Tail terminates in a very fine spine-
like point 21 to 224 long. Eggs T1p by 42u, 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 douglassti Cobbold, 1882; Strongylus
douglast (Cobbold, 1882, emend. Gedoelst, 1911) Gedoelst, 1911;
Trichostrongylus douglasit (Cobbold, 1882, emend. Gedoelst, 1911)
Theiler and Robertson, 1915.
Host—Struthio camelus.
Location.—Proventriculus.
Morphology.—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 27p
in diameter. Mouth with 3 small lips, each having a small pro-
tuberance; the mouth aperture triradiate. our submedian papillae
and 2 lateral structures, either lateral papillae or amphids, in shallow
grooves. Esophagus filariform, 480 to 500~ long. Excretory pore
about 300 behind head end.
Male \ength averages 4.65 mm.; width about 93 to 95u. 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-
lateral 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,
Fics, 8-10.—8. ORNITHOSTRONGYLUS FARIAI MALE BURSA. AFTER TRAVASSOS, 1921. 9,
ORNITHOSTRONGYLUS DOUGLASI. MALE BURSA. 10, SAME. TAIL OF FEMALE. Nos. 9-10
AFTER THEILER AND ROBERTSON, 1915
easily overlooked. Spicules 140 to 158 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 5.63 mm.; width about 105 to 109». Body
terminates in a blunt point 80, long, its tip slightly curved (fig. 10).
Vulva a transverse or oblique slit about 800n from the tip of the tail.
A short vagina leads to the muscular ovejector, which is about
3612—27——3
16 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
300u long. Posterior uterus smaller than anterior. Eggs 59 to T4p
by 36 to 44p.
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 750u 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 745p long and
without it 530. long. Molting occurs in the host in 4 to 5 days; the
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.—sStrongylus 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 740n 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 correctly 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 7604 long. Vulva in posterior body, dividing body in ratio of
41:7, or about 1.45 mm. from tail end. Eggs colorless, 80n by 36y.
Life history.—Unknown; probably simple and direct.
Distribution.— Europe.
NEMATODE PARASITES OF BIRDS Lé
GRNITHOSTRONGYLUS HASTATUS (Linstow, 1905) Travassos, 1920a
Synonym.—sStrongylus 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 784 deep. Esophagus 5/32 of body
length or about 100 to 300u 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 postero-lateral set off from the medio-lateral and ex-
terno-lateral, which are close together and parallel; the externo-
Fies. 11—12.—11, ORNITHOSTRONGYLUS PAPILLATUS. 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 5304 wide. Vulva near the posterior
extremity. Eggs thick-shelled, 302 by 21p in diameter.
Life history—Unknown; probably simple and direct.
Distribution.—Kurope.
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, Amédostomum 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. L'po-
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 Amédostomum 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, ictularia, has a buccal
capsule.
KEY TO GENERA OF AMIDOSTOMINAB
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
OL telam on presenmb ee = ose eA eee ee eae Amidostomum, p. 19.
Mouth with a much reduced buceal capsule (Seurat) or without buccal eap-
sule (Skrjabin). Head with 4 outwardly directed papillae and 2 pos-
teriorly directed epaulets. Spicules divided into 3 branches. Gorgeret or
telamon absent: 2G eues Pree Ea it eee 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 Amddostomum 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 Henry, 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. chevrewxi) at the base of the dorsal ray in
a common stem, but in other cases (A. radllieti) 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 Strongylus nodulosus Rudolphi as type, stating that Zeder
confused worms belonging in 2 genera (Amidostomum and E'pomi-
diostomum) 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. 26.
Female not over 13 mm. long, or, if longer, then with a buccal capsule having
3 teeth, or vulva with prominent anterior lip (A. acuwtum) ~~ ~_________ a
. Males 11.25 to 13.5 mm. long; females 13.5 to 15.75 mm. long; vulva 1% 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 1204 long; 1 tooth in buccal capsule; externo-dorsal ray originates
inj Common’ with: dorsal ray—-.--- #2 Amidostomum chevreuxi, p. 22
Spicules over 150u long; 3 teeth in buccal capsule and externo-dorsal ray
originates in common with other rays, not with dorsal, where known
(unknown for iA. fulicae@ye Hees. AE Ee TEES PGANIRS ES ORES EE 4,
4. Males 10 to 17 mm. long; females 12 to 23.7 mm. long; spicules 200 to 300u
long; female tail 487 long; buccal capsule with 3 teeth and 6 paired
bo
TL OOS 8 oe i ae ae ei a Amidostomum anseris, p. 20.
Males less than 9 mm. long; females up to i4.5 mm. long; spicules not over
204u long; paired ridges ot reported tor buceal capsuleses. = 2s 5.
5. Spicules 1p long; telamon 7Ou long__________ Amidostomum fulicae, p. 25.
Spicules 166 or 204 long; telamon 90 to 100u long___________-_--_-- 6.
6. Spicules 166 long; telamon 90u long; buccal capsule 27.54 wide in male, with
1 large and 2 very small teeth ____________.__ Amidostomum henryi, p. 22
Spicules 204u long; telamon 100u long; buceal capsule 154 wide in male,
with 1 large and 2 small teeth_2_________ Amidostomum raillieti, p. 24.
AMIDOSTOMUM ANSERIS (Zeder, 1800) Railliet and Henry, 1909
Synonyms.—A scaris 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. fuligula, A. fusca, A. leucops, A. marila, A. mollissima,
A. nigra, A. penelope, A. segetum, Chloephaga poliocephala, Fulica
atra, Fuligula cristata, F. marila, Gallinula chloropus, Nyroca clan-
gula, NV. fuligula, N. marila, and Somateria dressert.
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 ae
Male 10 to 17 mm. long by 250 to 350p wide. Bursa (fig. 14) with
2 large lateral lobes and a small median lobe. A pair of large con-
- tiguous 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
Fies. 18-16.—13, AMIDOSTOMUM ANSERIS. ANTERIOR END. 14, SAME. MALE
. BURSA. NOS, 153-14 AFTER RAILLIET, 1893. 15, AMIDOSTOMUM RAILLIETI. AN-
TERIOR END. 16, SAME. Bursa OF MALE. NOS. 15-16 AFTER SKRJABIN, 1916
maroon-colored spicules are 200u 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 long.
Female 12 to 24mm. long. The body is slender anteriorly, widens
at the vulva to 300 or 400y, and thins abruptly behind the anus; the
tail is long and straight and bluntly pointed. The vulva is a trans-
verse slit, 160 long, % of the body length from the tip of the tail,
and is sometimes covered with a projecting appendix. The eggs are
thin-shelled, 85y long by 50, wide, or, according to Skrjabin, 110p
long by 82» wide, (or 100u by 66» 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 18.54 wide, with 1 large tooth and 2 very small,
scarcely perceptible, teeth.
Male 8 mm. long by 130 to 1504 wide. Buccal capsule 15p
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 1664 long, irregular in shape and each cleft dis-
‘9.
Fics. 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, 90u 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
850n long. Buccal capsule 18.54 wide. Vulva 2.72 mm. from
tail end. Eggs oval, 92.5 to 103.64 by 70 to 80u, 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 skrjabinit Boulenger, 1926 (see Ad-
denda, p. 383).
Host—Himantopus himantopus and (A. skrjabini) 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 (fig. 20) 7 to 8p
deep and 10» 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 80u wide, and terminating in an un-
cinate tail concave ventrally. Esophagus and bulb 636y long.
Fics. 20-23.—20, AMIDOSTOMUM CHEVxKe£UXI. ANTERIOR END. @, DORSAL VIEW; B,
LATERAL VIEW. 21, SAME. MALE BURSA. 22, SAME. OUTLINE OF BURSA; LATERAL
view. Nos. 20-22 arrpr SeurarT, 1918. 23, AMIDOSTOMUM MONODON. ANTERIOR
END. AFTER LINSTOW, 1882
Bursa (figs. 21 and 22) with 2 large lateral lobes, 105p 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, 60p long.
Female 12.8 mm. long by 120, wide, terminating in an elon-
gate digitiform tail 265» long. Nerve ring 3/7 of esophagus
length from head. Esophagus and bulb 840u long. Vulva a trans-
verse slit, 90» long, slightly salient, in the posterior fourth of the
3612-—27——4
24 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
body and 2.835 mm. from anus. Ovejector branches divergent; vesti-
bule 510% 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.
long; sphincter 105p long. Uteri opposed; ovaries parallel. Eggs
SOpn by 45p, thick-shelled, and in blastula stage as oviposted.
Life history—Unknown; probably simple and direct.
Distribution.—Africa (Bone).
AMIDOSTOMUM RAILLIETI Skrjabin, 1915
Host.—F ulica 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 2554 wide. Buccal capsule 27.5p
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 290n. Eggs 90 to 105
by 50 to 65p.
Life history—Unknown; probably simple and direct.
Distribution—Asia (Russian Turkestan) and Egypt.
AMIDOSTOMUM ACUTUM (Lundahl, 1848) Seurat, 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 fulsque (Anas
fuligula) and Fallcauda 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 2254 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 were
close together and parallel, a thing which would not accord with the
assignment of this species to the genus Amiédostomum. 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. chevreuui, all rays except the dorsal ray have
such an origin where this point is covered or illustrated. YLinstow
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 1% 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
Synonyns.—s piroptera fulicae Rudolphi, 1819.
Host.—Fulica atra.
Morphology.—Amidostomum (p. 19): Characters of the genus.
Male 8.58 mm. long by 1554 wide. Tail 100n long. Buceal cavity
164 wide. Esophagus 884 long. Spicules 175. long. Telamon
(gorgeret) 70 long.
Female 9 mm. long by 175p wide. Buccal cavity 144 wide. Esoph-
agus 9724 long. Vulva 1.56 mm, from end of tail. Eggs 1054 by
o2u.
This description is taken from Seurat’s table without critical con-
sideration of the works of Creplin, Stossich, 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 Lainstow, 1882; Sclerostoma
monodon (Linstow, 1882) Stossich, 1889.
Host.—Oidemia 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 140 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 92u long by 62» wide.
Life history—Unknown; probably simple and direct.
Distribution.—Kurope.
Genus EPOMIDIOSTOMUM Skrijabin, 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 latera] 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 24
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 uncinatum (Lundahl, 1848)
Seurat, 1918 (=. anatinwm 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
ASOp longish. ee) ee Epomidiostomum uncinatum, p. 27.
Males 8.8 mm., or more, in length; females 18 mm., or more, in length.
Spiculese!SO:to 2100 Yong Fes a ee oe
2. Spicules dividing distally into 2 branches.
Epomidiostomum querquetulae (Addenda, p. 384).
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-
TOCHCC i es 9 ns eS Epomidiostomum skrjabini (Addenda, p. 384).
EPOMIDIOSTOMUM UNCINATUM (Lundahl, 1848) Seurat, 1918e
Synonyms.—Strongylus uncinatus Lundahl, 1848; EL pomidiosto-
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.—k pomidiostomum (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 150u wide, yellowish. Cuticle trans-
versely striated. Esophagus cylindrical, 8004 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 bifureating at its tip. There are a
pair of large caudal papillae near the base of the dorsal ray. Two
equal spicules, brown, 120 to 180u long, divide to form 3 terminations
posteriorly. No gubernaculum.
Female 10 to 11.5 mm. long by 250n wide. Tail (fig. 26) forms a
conical appendix 140 to 170u 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.
Fics. 24-27.—24, EPOMIDIOSTOMUM UNCINATUM.. ANTERIOR END. 25, SAME. MALE
BURSA. 26, SAME. FRrMALE TAIL. Nos. 24-26 arrra SKRJABIN, 1916, 27, EPo-
MIDIOSTOMUM ORISPINUM. Mae BURSA, AFTER MOULIN, 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; Stronigylus
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.—E pomidiostomum (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 2104 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 Amédostomum anseris
(p. 21), and similar papillae on the posterior edge of the cloacal
aperture. The spicules are 200 long, divided proximally into 3
branches, of which the dorsal is longitudinally striated.
Female 16.7 mm. long by 275» 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, Y5 of the body
length from the tail end. Eggs ovoid, thick-shelled, 952 long by
55u wide, segmenting when deposited.
Life history—Unknown; probably simple and direct.
Distribution.—Europe and Africa (Algeria).
Superfamily STRONGYLOIDEA Weinland, 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
(Syngamus). 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 cireu-
latory system or tissues as agamic individuals (Strongylus).
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 ((’so-
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 (1913). The superfamily diagnosis given by Hall (1916) 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--i.-iL----_=.—-=~.-----=. 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 body. 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 SrroneyiinaE Railliet, 1893
Subfamily 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.—Deletrocephalus 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 DELETROCEPHALEAE
1. Corona radiata present. Buccal capsule without teeth or longitudinal sus-
LEED OU GY olay eh FSF) ely W215 fea aa yo at oe a Codiostomum, p. 30.
Corona radiata absent. Buccal capsule with longitudinal sustaining ribs or
rays and usually“with teeth: 3 ss ee Deletrocephalus, p. 32.
Genus CODIOSTOMUM Reailliet 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 at
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.
MAAAAMUIE ER Lay
Neg
A aM ANNA EAA
My ey ia its
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Fics. 28-30.—CoDIOSTOMUM STRUTHIONIS. 28, HEAD END. 29, BUCCAL CAPSULE.
30, MALE BURSA, SPREAD OUT. AFTER Horst, 1885
CODIOSTOMUM STRUTHIONIS (Horst, 1885) Railliet and Henry, 1911
Synonym.—sSclerostoma struthionis Horst, 1885.
Hosts.—Struthio molybdophanes, S. camelus, S. australis.
Location.—Ceca, large intestine, and stomach.
Specific diagnosis.—C odiostomum (p. 30) : White or yellow, rather
stiff worms. Buccal capsule (fig. 29) 2802 deep by 1002 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 450n 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
to 1 mm. long, alate, transversely striated, curved, and widened in
the proximal third of their length. Gubernaculum boat-shaped,
a2 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
84 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 700n wide. Esophagus length
\ of body length. Lateral cervical papillae 960, from anterior end,
excretory pore 800, nerve ring 660n. Vulva (fig. 32) 250» anterior
to anus. Anus 572u 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, long. Ovejectors parallel,
the cuticular portions 620» long and the musculo-epithelial portions
1.5mm. long. Eggs oval, 63 by 35p.
Life history—Unknown; probably simple and direct.
Distribution.—A frica.
Figs. 381-32.—CoDIOSTOMUM STRUTHIONIS. 931, MALE BURSA, LATERAL VIEW. 382,
FEMALB 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.—Sclerostoma 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
laterally, 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 6 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, 900u to 1.025 mm. long.
Female 17 to 24 mm. long. Vulva (fig. 35) close to the anus and
often covered by a cuticular expansion. Eggs thin-shelled, 120 to
125p by 70 to T5y, segmenting when deposited.
Life history—Unknown; probably simple and direct.
Distribution —South America (Brazil) and Europe (in museum
material).
Fics. 33—-35.—DELETROCEPHALUS DIMIDIATUS. 33, HEAD; da, LATERAL; DJ, 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 (Syngamus) 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 postmedian. Eggs operculated after
deposition.
Type genus.—Syngamus von Siebold, 1836.
KEY 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,
150u or less. Vulva in anterior half of body---------~-- Syngamus, p. 34.
Worms not permanently joined in copula. Buccal capsule with 6 or 7 teeth
rarely 2 (error?). Bursa rays slender. Spicules long, over 400u. 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 25u. 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 150u long.
Syngamus microspiculum, p. 38.
Each branch of dorsal ray trifurecated or simple; spicules less than 100u
2. Each branch of dorsal ray trifurcate; spicules 60y long.
Syngamus trachea, p. 34.
Each branch of dorsal ray simple, not subdivided_______________________ oe
3. Spicules subequal, 50u long. Parasites of Nucifraga.
Syngamus parvus, p. 39.
Spicules distinctly unequal, the right 79u, the left 694 long. Parasites of
COP Rn Es a a See or ae Syngamus gracilis, p. 39.
The following species are nomina nuda and are omitted from the
above key; Syngamus coelebs from Falco lagopus, 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 primitivus Molin, 1860; Strongylus primi-
tivus (Molin, 1860) Hutyra and Marek, 1910; S. trachealis (Siebold,
1836) Nathusius, 1837; Sclerostoma 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 by Chapin.
Hosts—Meleagris gallopavo and Gallus gallus. Also reported
from Anas boschas, Anser anser, Ciconia alba, C. nigra, Corvus
cornix, 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. viridis,
NEMATODE PARASITES OF BIRDS SD
Pyrrocorax alpinus, Strix noctua, Sturnus vulgaris, Tetrao urogal-
dus, “thrush.” Chapin notes that the specimens from birds other
than the Galliformes need further study; those from Corvus, spp.
may be 8. 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 (Mégnin).
Morphology—Syngamus (p. 34): Cylindrical red worms, the
color more pronounced in the female. Head (fig. 37) enlarged and
FIG. 36.—SYNGAMUS TRACHEA. OUTLINE. AFTER CHAPIN, 1925
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).
1004
ic. 37.—SYNGAMUS TRACHEA. HEAD. a, TEETH IN 8-TOOTHED (NORMAL) FORM;
b, FRONT VIEW OF HEAD; C, TEETH IN 9-TOOTHED FORM. AFTER CHAPIN, 1925
Male 2 to 6 mm. long by 200” 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 64 long
according to Chapin, who states that the reports of 140, 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 (w« 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 long by 50n
wide, or, according to Ortlepp, 78 to 100u long by 43 to 46 wide.
Embryo 280p 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 94 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
a5 vulva under the margin of the male
Fics. 38-39.—Syneamus tracuea. 38, bursa and get to the lumen of the
SFICULDS.. 39, MALE BURSA, AFTER: trachea.qand,bronehin. Lhese|,eesRs
CHAPIN, 1925 : : 3
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
nim. 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 17 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. Apparently, 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 1806, 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, 19lia
Host.—Phalacrocorax carbo.
Location.—Trachea.
Morphology.—Syngamus (p. 34): Base of buccal capsule (fig.
406) armed with 3 small triangular teeth.
Male 3.9 to 4mm. long by 270 to 290» wide, brown in color. Buccal
capsule 170 deep by 220» wide; its basal teeth are 70u long. Ksoph-
agus 5804 long and is widened posteriorly. Spicules 150, long, ac-
cording to Skrjabin (1915), or 115 long, according to Skrjabin
Fic. 40.—SYNGAMUS MICROSPICULUM. @, MALE AND FEMALE; 0, HEAD END; C, MALE
BURSA. AFTER SKRJABEN, 1916
(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 250u at the anus.
Tail pointed and curved, according to Skrjabin’s figure. Buccal cap-
sule 2502 deep by 3404 wide. Esophagus 765n long, widened pos-
teriorly. Vulva 4.45 mm. from head end. Eggs 75y by 48, of char-
acteristic shape, flattened on one side.
Life history—Unknown; probably similar to that of S. 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% wide, cylindrical, with neck
region slightly constricted. Buccal capsule heavily chitinized, 178
in depth, 207, in its greatest inside diameter; walls 29» thick. Buc-
cal teeth and circumoral papillae as in S. trachea. Esophagus about
326 long; nerve ring and excretory pore not located. Bursa (fig.
43) 350 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
(004
Fics. 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, similar in shape to those of S. trachea.
Female about 7.8 mm. long by 6502 maximum width; width just
posterior to head 350u. Buccal capsule 300n in depth, 440p 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 (# in fig. 41) ;
anus subterminal; tip of tail very blunt. Eggs 74u by 44y, 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 45y thick. Buccal teeth
as in S. trachea, the largest 45. high. Esophagus nearly 600» long,
clavate, its greatest diameter 135y. Nerve ring, excretory pore, and
cervical papillae not seen. Bursa (fig. 45) narrow anq deep. As
in S. parvus, the dorsal ray branched, the branches simple; however,
whereas in S. 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. shghtly shorter, measuring 132u. Lateral rays
mutually contiguous, parallel, and about the size of either branch
10044
Fics. 44-—46.—SYNGAMUS GRACILIS. 44, MALE AND ANTERIOR
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, 69 long, is
nearly straight.
Female 8 to 11 mm. long and proportionately stouter than the
male. Buccal capsule relatively enormous, its internal dimensions
525y wide by 300% deep; wall 50n thick. Teeth as in S. trachea.
Esophagus 825p long, attaining its greatest diameter, 225y, 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
300u from posterior end, which is very blunt. Eggs 79 to 83 by
40 to 46p.
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 S.
NEMATODE PARASITES OF BIRDS 4]
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 4002) and filiform. Vulva anterior,
iC
Fies. 47-48.—CYATHOSTOMA BRONCHIALIS. 47, HEAD END. 4, SIDE VIEW; 0, FRONT
view. 48, MALF 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 EK. Blanchard, 1849.
KEY TO SPECIES OF CYATHOSTOMA
ia Vulva ‘at or near the anterior third of the body ___-.---_..__<«....~--.!1_4. 2.
Viulvannear the middle ofthe body_. 1 -2.+=2 24 5.
2. Spicules 500u long; eggs 56u long; in Tadorna tadorna,
Cyathostoma tadornae, p. 42.
Spicules 6004 long or longer, where known; eggs 80u long or longer__--—_-~ 3.
3. Eggs SOu by 40u or 924 by 53x; spicules 690u long (Skrjabin) ; in ciconiform
DE Ck Soto eg eer a es eS Cyathostoma variegatum, p. 44.
Eggs 80 to 90u long by 55 to 60u wide or wider; spicules 600 to 650u long_— 4.
4. Eggs 90u by 604; spicules 600u long; in anseriform birds.
Cyathostoma bronchialis, p. 42.
Eggs 80u by 55u; spicules 650u long; in Casuarius galeatus.
Cyathostoma boularti, p. 44.
42 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
5. Male unknown; female small, up to 18 mm. long, in orbital cavity of Larus
LIA DUIS 1A) fs ease ae a ULE yee VE Di mage ee ee Cyathostoma lari, p. 43.
Male known; female up to 20 mm. long or longer________________________ 6.
6. Spicules 490u long; gubernaculum 67u long; female up to 30 mm. long; in
thoracic air sac of Buteo borealis_______~ Cyathostoma americanum, p. 45.
Spicules 660 to 720u long; gubernaculum 92u long; female up to about 20 mm.
long; in trachea of Coscoroba coscoroba__. Cyathostoma coscorobae, p. 47.
CYATHOSTOMA BRONCHIALIS (Muehlig, 1884) Chapin, 1925
Synonym.—Syngamus bronchialis Muehlhg, 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 6004 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. 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 90u long by 60 wide and provided,
according to Railliet, with a single polar operculum.
Life history—sSimilar to that of Syngamus trachea (p. 36). Rail-
let 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.—sSclerostoma 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 or 1/15 of body length. Bursa sup-
ported by 10 rays of which 4 are simple and 6 are bifurcated at their
apices. Spicules 500% long; brown in color.
Female 23 mm. long, brighter red than male. Head 900p 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 564 long (Chatin says 0.56 mm., but this is evidently
an error and Skrjabin has given the figures as 56n).
Life history—Unknown; probably similar to that of Syngamus
trachea (p. 36).
Distribution.—Europe.
Tics. 49-53.—CYATHOSTOMA TADORNAE. 49, MALBP. 50, FEMALE TAIL. FROM CHAPIN,
1925, arreR CHATIN, 1874. 51, CYANHOSTOMA LARI. FEMALE; ENTIRB WORM. 52,
FEMALE; HEAD END. 53, VULVA. AFTER BLANCHARD, 1849
CYATHOSTOMA LARI E. Blanchard, 1849
Synonyms.—Strongylus, species Siebold, 1837; Sclerostoma cya-
thostomum Diesing, 1851; Sclerostoma lari (EE. Blanchard, 1849)
Molin, 1861a; Syngamus lari (KE. Blanchard, 1849) Railliet, 1898.
Hosts —Larus 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 500u 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 1% 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, C. 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 260n long by 140, wide.
Fies. 54-55.—CYATHOSTOMA VARIEGATUM. 54, MALE BURSA. 55, HBAD. @, FRONT
VIEW ; Db, SIDE VIEW. FROM CHAPIN, 1925, arreR 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, 6904 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 80p by
40u (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 (Mégnin, 1884) Chapin, 1925
Synonyms.—Sclerostoma boularti Mégnin, 1884; Syngamus bou-
larti (Mégnin, 1884) Railliet, 1898.
Host.—Casuarius galeatus.
Location.—Trachea.
NEMATODE PARASITES OF BIRDS 45
Morphology.—Cyathostoma (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 4504 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 850u wide. Head 500u wide. Buc-
cal capsule (or aperture?) 3504 wide. Body shaped lke 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 55n, operculated at small end.
Life history—Unknown; probably similar to that of Syngamus
trachea (p. 36).
Fics. 56-58.—CYATHOSTOMA BOULARTI. 56, HEAD; SIDE VIEW. 57, MALE BURSA.
58, HEAD; FRONT VIEW. FROM CHAPIN, 1925, AFTER MEGNIN, 1884
Mistribution—Australia and Europe (in zoological garden,
Paris).
CYATHOSTOMA AMERICANUM Chapin, 1925
Host.—Buteo borealis.
Location.—Posterior thoracic air sacs.
Morphology.—C yathostoma (p. 41): Cuticle smooth, without
transverse striations.
Male 12 mm. long; body (fig. 59) somewhat attenuated in ante-
rior fifth. Buccal capsule about as wide as deep, its internal trans-
verse diameter 185y, the depth of its chitinous portion 1704; wall of
capsule 25y 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
57m high. Head with circle of 6 papillae, situated 60u 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, 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, 7504 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
x
Wa
Vv Gar eceliate videdess?
Fie. 59.—CYATHOSTOMA AMERICANUM. OUTLINE OF FEMALE AND MALE WORMS. AFTER
CHAPIN, 1925 (@=EXTENT OF UTERI)
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 490n 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 370n, the depth of its chitinous portion 280y;
60,
Tip “oop
Fic. 60.—CYATHOSTOMA AMERICANUM. HEAD. a, SIDE VIEW, MALE; D,
FRONT VIEW; C, SIDE VIEW, FEMALE. AFTER CHAPIN, 1925
wall 20u thick. Buccal teeth shorter and blunter than in male;
largest tooth 60. high. Head papillae as in male, slightly more
anterior in location. Esophagus 960, long, clavate, its diameter 130p
at anterior end, 250 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. 62) just anterior to
middle of body, its lips prominent. Anus (fig. 62c) just anterior
to slender caudal appendage. Eggs (fig. 62a) 72u by 42, 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.
Location.—Trachea.
Morphology—Cyathostoma (p. 41):
Male 5.5 mm. long (estimated) ; anterior portion attenuate. Buce-
cal capsule (fig. 63) with straight sides, 834 wide by 100 deep;
lateral walls 13, thick. Six buccal teeth, the alternate teeth (about
35p high) are about twice the size of the others. Head papillae ap-
parently asin C. americanum. Esophagus about 700 long, attaining
its greatest diameter (150) at its posterior end. Nerve ring just
200 ys.
Ries. 61-62.—CYATHOSTOMA AMERICANUM. 61, MALE. a, Bursa; Db,
GUBERNACULUM. 62, a, EGG; b, VULVA; C, TAIL (FEMALE). AFTER
CHAPIN, 1925
anterior to middle of esophagus; position of cervical papillae and
excretory pore not determined. Bursa (fig. 64) well developed, 720
in transverse diameter when outspread. Ventral rays similar, slen-
der and approximate, about 105py long; lateral rays parallel and con-
tinuous throughout their length; externo-lateral ray the shortest
(135 long); medio-lateral and postero-lateral rays 150n and 210,
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
C. bronchialis. Spicules filiform, from 660p to 720% long, united at
tips and each bearing a finely striated wing. Gubernaculum present,
92u long.
Female about 20 mm. long. Buccal capsule (fig. 63) trapezoidal
in optical section, 210» deep by 225 wide at base, 285 wide at apex:
3612-—27T—_5
48 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
walls 30y thick. Six teeth, their height proportionately less than in
the male, the highest teeth being 60p. Esophagus 900, long, attain-
ing its greatest diameter (225) near its posterior end. Nerve ring
at anterior 24 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, 80n long by 50 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.).
00x
Fics. 63-64.—CYATHOSTOMA COSCOROBAP. 63, BUCCAL CAPSULES OF FEMALH 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.
Ty pe-family —Ascaridae Baird, 1853.
NEMATODE PARASITES OF BIRDS 49
KEY TO FAMILIES OF ASCAROIDEA
Mouth with 8 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_—+.—4_- + .-5_ ++ .-_._-_- 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.
Ty pe-genus.—H eterakis Dujardin, 1845.
The Heterakidae, including Heterakis, Ascaridia, and Subulura,
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 of
Ascaridia galli (=A. perspicillum) are identical with those of the
heterakids, showing the affinities of Ascaridia with Heterakis.
KBY TO SUBFAMILIES OF HDTERAKIDAD
Mouth with 3 well-defined lips. Esophagus with or withdut 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)
Ca a pase er a ee a Subulurinae, p. 104,
Subfamily Hereraxrnaz 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.
Ty pe-genus.—H eterakis 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.
SEEMS ea: S01 HO Ua SSC ray TN Sa aS a Heterakis, p. 50.
Genus HETERAKIS Dujardin, 1845
Synonyms.—Heteracis Molin, 1858; Ganguleterakis Lane, 1914.
Generic diagnosis.—Heterakinae (p. 49): Mouth with 3 lips.
Ksophagus subcylindrical, progressively swollen toward its posterior
extremity and with a distinct bulb. Two lateral membranes present
or absent. J/ales with caudal alae well developed and sustained by.
papillae of ray-like appearance. Spicnies 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 Ciconia (Hurenura) maguari.
Heterakis valdemucronata, p. 74.
Prom. othervhosts than sabovest)t owe week ab pee ein bee Ae De
2. Male with not more than 9 pairs of caudal papillae_____-___-» 3.
Male with more;than 9 pairs of caudal papillae__--__~ 2 4.
3. Male with 6 pairs of caudal papillae; sucker 35u anterior to cloacal aperture;
tail of female: /6:8"of total lensth_ 2 #247. 4s Heterakis bancrofti, p. 56.
Male with 9 pairs of caudal papillae; sucker 194” anterior to cloacal aper-
ture; tail of female 1/14 of total body length. Heterakis chenonettae, p. 60.
4. -Spicules wnequale 228 22 eRe oe SS ER eae Res eS Oa ne Re 9 5D.
Spicules equal or subequal (not more than 50u difference in their lengths)_ 13.
Se LOnger SpLeulev Oye, Acme Dae Ceti se a ee ee 6
Honger spicule less*than: i mm, iinslength 2s eee ees ae ee 12,
6. woneer spicule over: 2 min. in Leng bhi se sen eee ee ee Th
Eonver;splculewess than: 2) mmssin leneth =: 233 ee ee 10.
fersOrt SPICuten O4 Oui LOM Pees cee wie Ses ee eet oo Eee Heterakis parisi, p. 69.
SHOntsSDICUlLe: ROOM OL) LONGO = ee anata tees nie ee 8.
8. Tail short in both sexes, the anus of female being 780, the cloacal aperture
or malert On; from posterior ends tse Heterakis valvata, p. 74.
Tail long compared with above, the anus of female being not less than 988u,
the cloacal aperture of male not less than 4164, from posterior end____ 9.
9. Preanal sucker 60 to 75x in diameter; right spicule 2 to 2.17 mm. long;
female 10 to 15 mm. long; eggs 63 to 71u long by 88 to 48u wide.
Heterakis gallinae, p. 52.
NEMATODE PARASITES OF BIRDS ol
Preanal sucker SO to 90u in diameter; right spicule 2.8 mm.. long; female
7.9 to 9.6 mm. long; eggs 75 to TSu long by 43 to 484 wide.
Heterakis longecaudata, p. 66.
. Preanal sucker 250% in diameter; cloacal aperture 1.3 mm. from posterior
extremity; female 9.5 mm. long; anus 1.5 mm, from posterior extremity.
Heterakis bosia, p. 57.
Preanal sucker not over 124u in diameter; cloacal aperture not over 488u
from posterior extremity; female 7 to 8 mm. long; anus not more than
Mm: from: POSLELIOL extremity =~ ee ee A wk Ne tt,
11, Preanal sucker S83 internal diameter, 1244 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 Papiulae=_____ = __--. = == Heterakis bonasae, p. 75.
Preanal sucker only 36u in diameter; 38 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 5404 from posterior
end; vulva at about posterior third of body, dividing body length in ratio
Dike le fe nee eee ee Se eee SIS Heterakis pusilla, p. 70.
Short spicule 260u long; female 8.2 mm. long, its anus S830u from pesterior
end; vulva slightly anterior to middle of body.
Heterakis putaustralis, p. 71-
SP SILCULGS. COVED: Tl IIT, LOIN es are erent a eee eer a Ae VA 14.
Spicules less) Ghamat mr, Tome teeet thls Sten ee AS RK AEN co Reh Pe oA 17.
14. Male 14 to 22 mm. long; female 22 to 31 mm. long; eggs 52u 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 T0u or more
Dy +2e-0r More" spicules not over 1-S-mm: long==--2<--* ae 15.
15. Head with 3 lips and 3 small interlabia; no papillae deseribed 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
Iii OF total lengthe ls et aes petted shee Heterakis isolonche, p. 64.
Lateral alae extend almost the whole body length; male 15 mm. long, its
tail 1/23 of total length; female 17 mm. long, its tail 1/14 of total length.
Heterakis neoplastica, p. 65.
bed
~
Cc
7. spicules-over 600 Tong: Y 4? wal! il Gs ON Ok Sh ek Oe 2 OU 18.
NDLCUICS NOtLOVER OU mL LOMe see 2s Te Deak ee ne) 2) St ate Ae ae SY 23.
18. Male 4 mm. long; female 4.2 mm. long _________ Heterakis nattereri, p. 75.
Male 9 mm. or longer; female 14 mm. or longer_—_u_1-_- 19.
19. Preanal sucker 290 in diameter; the most anterior pair of preanal eaudal
papillae very small and situated a considerable distance anterior to the
sucker ; spicules at least 890” long_______-___ Heterakis tenuicauda, p. 73.
Preanal sucker not over 2304 in diameter; the most anterior preanal
papillae not situated as above; spicules not over S860u long ____-__— 20.
20. Length of females 25 mm. or more, the tail short (anus not over 460 from
POSTEHLOL VEG) 227 ees i Wh EE OAS eS OE) Oe VE 21.
Females not over 15 mm. long, the tail comparatively long (anus 800u or
FATEUEH LOM, POStETION, GN!) oo. Se 2 ee es ee 22.
52 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
21. Male with cloacal aperture 270u from posterior end; preanal sucker 180u in
diameter ; spicules 630 to 6504 long; female with anus 460y from posterior
end; vulva at anterior third of body__.-_--__---- Heterakis alata, p. 55.
Male with cloacal aperture 4204 from posterior end; preanal sucker 2304 in
diameter; spicules 850” long; female with anus 220, from posterior end;
vulva at about anterior fifth of body_____-_____ Heterakis skrjabini, p. 71.
22. Tail of male 1/28 of total body length; preanal sucker 904 in diameter; 10
pairs -of, caudal papillae: ss e Heterakis psophiae, p. 69.
Tail of male 1/55 of total body length; preanal sucker very pyromi-
nent, 220% in diameter; 8 pairs of caudal papillae.
Heterakis papillosa, p. 67.
re OMAle NOt OVIER si cGeTe. | LOIS eS as re Se a ee ee 24,
HemalerTaiG mms toys LO es a a ee see 26.
24, Preanal sucker broken on each side, aS well as posteriorly, by a papilla;
spicules equal, 400u long; eggs 63u long by 35y wide.
Heterakis brevispiculum, p. 59.
Preanal sucker not broken on each side, but only posteriorly, by a papilla;
spicules subequal, there being 50 difference in their lengths; the longer
NOE GOVT Ss Leer Tay Vey a we aes a 25.
. Male 5.5 mm. long by 2104 wide; preanal sucker 554 in diameter; female
6.6 mm. long by 300” wide, its tail ro of total body length; eggs 50u long
Dy SSO pe Wye ees ees See oe Se eas fe eee ca eh Heterakis beramporia, p. 56.
Male 7.5 mm. long by 410u wide; preanal sucker 7ly in diameter; female
7.6 mm. long by 4604 wide, its tail 144 of total body length; eggs 57 long
no
OU
DY pes Wal LG a te spe eS ee Heterakis hamulus, p. 63.
26. Spicules 110 and 130u long respectively; tail of female 1/7.8 of total body
Teer Ga ok nig eg ai Ear eat Heterakis macroura, p. 67.
Spicules 260u long or longer; tail of female not as long as above except
possibly in H. dispar where length not given__________-___________- PAG
27. Female 34 mm. long; vulva in anterior part of body; male 19 to 20 mm.
long; spicules not over 390u 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” long____-_-_____-_____-~ 28.
28. Male 7.8 mm., female 11.6 mm. long; eggs 70u long.
Heterakis caudata, p. 59.
Male 11 mm. or longer; female 14.8 mm. or longer; eggs not more than
Go lOne sa. oot ome ere ae en ee eee 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 vesicularis
Froelich, 1791, part; Heterakis vesicularis (Froelich, 1791) Dujar-
NEMATODE PARASITES OF BIRDS D3
din, 1845; Heterakis papillosa Railliet, 1885, misdet., not Ascaris
papillosa Bloch, 1782 (=Heterakis monticelliana Stossich, 1892)
from bustard.
Hosts—Anas boschas domestica, A. tadorna, Anser anser, A.
cinereus domesticus, Bonasa sylvestris, Ceriornis satyra, Chenopsis
atrata, Chrysolophus pictus, Colinus virginianus, Corvus cajanus,
Coturnix communis, C. dactylisonans, Cupidonia cupido, Gallus
gallus, Grossiptodon manschuricum, Lagopus mutus, L. scoticus,
Meleagris gallopavo, Numida meleagris, Ortyx virginianus, Otis
tarda, O. tetrax, Pavo cristatus, Perdix cinerea, P. coturnix, P.
perdix, P. saxatilis, Phasianus colchicus, P. gallus, P. nycthemerus,
P. pictus, P. veneratus, P. versicolor, Tadorna tadorna, Tetrao
Lonasia, T. lagopus, T. urogallus.
The specimens from Otis tarda are probably Heterakis papillosa
(Bloch, 1782).
Fias. 65-67.—HETERAKIS GALLINAE. 65, MALH TAIL, LATERAL VIEW; 66, VENTRAL
VIEW ; 67, FEMALE TAIL. AFTER LANE, 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. from caudal extremity. There are 12 pairs of
papillae and a well-developed preanal sucker 60 to 75y in diameter
with strongly chitinized walls (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. 67) 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 45 mm. Eggs thick-shelled,
ellipsoidal, 63 to 71p long by 388 to 48 wide, according to most
authors, or 68 to 75p long by 36 to 38 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 days under favorable conditions of temperature (18 to 29° C.)
and moisture to the point where each contains an infective embryo.
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 by suitable birds, the embryos are
released from the shell and develop to adult worms. Within 41%
hours after ingestion of the eggs, the larvae are found to have
emerged into the small intestine. At the end of 24 hours they are
present in the ceca, and sometimes in the colon, and are 250u long by
i8 wide. In 10 days they are 790» long by 45h wide. The entire
development in the bird takes place in the digestive tract, the worms
being mature in 24 days. There is no wandering of the larvae to
the lungs as is possibly the case in Ascaridia 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.
_
Or
NEMATODE PARASITES OF BIRDS
HETERAKIS ALATA Schneider, 1866
Hosts —Crypturus, 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 300p; the slender part
measures about 1.2 mm. long.
Male 17 to 20 mm. long. Candal alae (fig. 68) wide; preanal
sucker about 180» in diameter, with a papilla on its posterior border.
Schneider described and figured only 9 pars of caudal papillae but
Travassos states there are 14; his figure seems to show 13 pairs.
Spicules slender, about 630 to 6504 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 4604 from posterior end. Eggs 59u long by 37p wide.
Life history—-Probably similar to that of H. 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
Ascaridia evidently was based on Schneider’s inadequate descrip-
tion; Travassos has shown, in a more complete study, that it belongs
in the genus [Teterakis.
Hosts.—Crypturus 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 1.85 to 1.44 mm. long, with a small bulb and a slight dilation in
the anterior part.
Mate 14 to 22mm. long. Preanal sucker 90» in diameter, a papilla
on its posterior border. In addition (fig. 69) 12 pairs of caudal
papillae, 3 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), sea he says the cloacal aperture is 270p
from the posterior end.
Female 22 to 31 mm. long. Vulva salient, situated a little an-
terior to the middle of the body. Anus about 1.12 mm. from pos-
terior end. Eggs 52y long by 29n wide.
Life history—Probably similar to that of H. gallinae (p. 54)
Distribution —South America (Brazil).
3612—27 6
56 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
HETERAKIS BANCROFTI Johnston, 1912
Host.—Catheturus latham.
Location.—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, 860n long, sharply pointed.
15 (0
Fics. 68—71.—68, HETPRAKIS ALATA. MALB TAIL. AFTER SCHNEIDER, 1866. H®TERAKIS
ARQUATA. MALB TAIL. 69, AFTER SCHNEIDER, 1866. 70, ArTER TRAVASSOS, 1913. 71,
HETERAKIS BANCROFTI. MALB TAIL. AFTER JOHNSTON, 1912
Female 6.25 mm. long by 3304 wide. Tail rather short (anus 910p
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 8004 from head
and extend almost entire length of body.
Male 5.5 mm. long by 210% wide. The circular sucker (fig. 72) is
554 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” 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, from the tip
of the tail. The eggs are 50u long by 30p wide.
Life history —Probably similar to that of H. gallinae (p. 54).
Distribution—Asia (Irtlia (Bengal) and Philippines).
HETERAKIS BOSIA Lane, 1914
Hosts.—Ceriornis satyra and Tragopan satyra.
Location.—Intestine, probably ceca.
Morphology—Heterakis (p. 50): Cuticle with cross-striations.
O:Imm.
Fic. 72.—HETERAKIS BERAMPORIA. 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 81 mm. long. Cloacal aperture 1.3 mm. from caudal ex-
tremity; long slender appendage (that is, part of tail posterior to
the papillae) is 700n long. Sucker 250u in diameter, its posterior
rim 350n from the cloacal aperture. Twelve pairs of caudal
papillae (fig. 732) of which 4 are preanal, 2 adanal, 6 postanal.
Spicules unequal, the right (fig. 73c) 1.6 mm. long, its proximal end
wide (75y) like a spear head, its distal end a long sharp point; left
spicule (fig. 736) 900» long, with a one-sided expansion at about a
quarter of its length from the point.
Female 9.5 mm. long by about 4002 wide. Anus 1.5 mm., caudal
papillae 800p, from posterior extremity. Vulva slightly posterior
to middle of body; posterior to it there are occasionally 5 or 6
cuticular tubercles. Eggs 75y long by 35y wide.
58 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history.—Probably similar to that of H. 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 rufescens.
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 590u wide. Tail 1/57
of total length, according to Linstow; 450 long, according to
O17
Fic. 73.—HETERAKIS BOSIA. da, MALE TAIL AND POINT OF RIGHT SPICULE; Db, LEFT
SPICULE FROM DORSUM; C, RIGHT SPICULE FROM SIDE. AFTER LANE, 1914
Travassos. Preanal sucker 200» 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
260u long, according to early descriptions; 390» long, according to
Travassos, 1918.
Female 34 mm. long by 790” wide. Vulva in anterior part of
body. Tail 1/30 of total length. Eggs 65y 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
Life history—Probably similar to that of H. gallinae (p. 54).
Listribution—South America (Brazil).
HETERAKIS BREVISPICULUM Gendre, 1911
Hosts —Francolinus bicalearatus, Gallus gallus, and Numida
meleagris.
Location.—Ceca.
Morphology—Heterakis (p. 50): These worms are very similar
to H. 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 5.34 to 8.35 mm. long by 250 to 300” wide. The spicules
(fig. 76) 4004 long and equal; sucker as described above. The
O/T
ic. 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 H. 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).
Female 6.38 to 10.6 mm. long by 260 to 860u wide. The vulva is
in the middle of the body. The eggs are elliptical, 632 long by 36,
wide, and deposited before the formation of the embryo.
Life history.—Probably similar to that of HZ. gallinae (p. 54).
Distribution.—Africa (Dahomey and Belgian Congo) and South
America (Brazil).
HETERAKIS CAUDATA Linstow, 1906
Hosts —Anas sponsa and Lampronessa sponsa.
Location.—Ceea.
Morphology. — Heterakis (p. 50): Three semicircular lips.
Esophagus with bulb.
60 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 7.8 mm. long by 2904 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 1804 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 70u long by 44 wide.
©) ©
® ®
SP, ».
78.
Fics. 75-78.—HETERAKIS BREVISPICULUM. 75, MALE TAIL. 76,
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.—C 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, anterior to it. Johns-
ton’s description of the spicules is confusing. He states: “The
spicules are 1.17 mm. in length and 0.018 mm. in breadth. The
ionger 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.—FProbably similar to that of H. gallinae (p. 54).
Distribution—Australia (Sydney).
cy
la
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=
= i
=a}
i
50,
ODT?
Fics. 79-82.—79, HerprakIS CAUDATA, MALE TAIL. AFTER LINSTOW, 1906. 80,
HPTERAKIS CHENONETTAR. MALP TAIL. AFTER JOHNSTON, 1912. 81, Hprera-
KIS CIRCUMVALLATA. MALP TAIL. AFTER LiINSTOW, 1906. 82, HETERAKIS
DISPAR. MALB 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 @. 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 H. gallinae (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. moschata, A. tadorna, Anser anser,
A. cinereus, A. fabalis, A. segetum, Bernicla sandwichensis, Surnia
passerina, Strix 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 cuticular
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). Tt 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.—Odontophorus capueira.
Location.—Intestine.
Morphology —Heterakis (p. 50): Mouth with3 equal lips. Esoph-
agus about 900u 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 36y 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 810u and 170» 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 9002 and the longer spicule over 1
mm. (roughly 1.3 mm.) in length.
Fic. 83.—HETERAKIS FAaRIaAl. MALE TAIL. AFTER TRAVASSOS, 1913
Female 7 mm. long. Vulva near middle of body. Anus about 1
mm. from the posterior end. Eggs 74p long by 33 wide.
Life history.—Probably similar to that of H/. gallinae (p. 54).
Distribution —South America (Brazil).
HETERAKIS HAMULUS Linstow, 1906
Hosts.—Pavo 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% wide. Esophagus length 1/9 of body
length. Right spicule (fig. 84) slender, 370 long, ending in a hook
distally; left spicule straight, 320 long, surrounded by a broad cor-
neous sheath from which only the point projects. Sucker 71 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 460u wide. Esophagus 4% of body length.
Tail length 14 of body length. Vulva almost in equator of body.
Eggs 57» long by 42u. wide.
Life history.—Probably similar to that of H. gallinae (p. 54).
Distribution —EKurope (Germany).
HETERAKIS INTERLABIATA Ortlepp, 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 300 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 400p wide. Tail long and
pointed. Vulva at about middle of body. Eggs oval, 66» long by
37 wide.
Life history.—Probably 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 1/138 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 150u 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 1/7.8 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~ long by 42
to 46 wide.
Life history.—Probably similar to that of H. gallinae (p. 54).
TTR
JIA
CT
85,
:
4177/77
TT
TT
———
2/7177
Fics, 84-86.—84, HrTpraAKIS HAMULUS. MALE TAIL. AFTER LINSTOW,
1906. 85-86, HETERAKIS INTERLABIATA. 85, ANTERIOR END; 86, MALE
TAIL. AFTER 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 H. isolonche are identical with his species, HZ. 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 7’. pictus.
Location.—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 5502 maximum width. Esophagus, including
bulb, 1.9 mm. long; esophagus 79» wide; bulb 370u long by 210» wide.
Preanal sucker 950u 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 6002 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 40p.
Yorn
Fics. 87—-88.—87, HETERAKIS ISOLONCHE. MALP TAIL. AFTER LINSTOW, 1906. 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. zsolonche, 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. tsolonche have not been
shown to be of that nature.
Since the above was written, Baylis (1925) has examined Was-
sink’s specimens and has come to the conclusion that this species is
identical with ZZ. zsolonche (p. 64).
HETERAKIS LONGECAUDATA Linstow, 1879
Hosts—Francolinus gularis, Galloperdix spadicea, Lophophorus
impeyanus, Megacephalon maleo, Tragopan satyra.
Location.—Ceca.
NEMATODE PARASITES OF BIRDS 67
Morphology—fHeterakis (p. 50): Mouth with 3 lips, followed by
a short vestibule. Esophagus 1/8.5 of total Jength, with well de-
veloped bulb with valve teeth. Lateral alae well developed, extend-
ing for greater part of length of body.
Male 7.9 to 9.1 mm. long by 240n wide. Tail (fig. 89) very long
and sharply pointed, 1/19 of total length. Caudalalae wide. Preanal
sucker 80 to 90u in diameter, 100 to 1504 from cloacal aperture.
Twelve pairs of caudal papillae, of which 8 are postanal and 4 pre-
anal. Left spicule (720% 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 sight anterior to middle of body, dividing body length
in ratio of 73:75; according to Baylis and Daubney slightly posterior
to middle, 3.7 i 4.75 mm. from tail end. Tail 1g of total length.
According to Baylis and Daubney, there is a conspicuous pair of
caudal papillae at about 6802 from posterior end. Eggs 75 to
78» long by 43 to 48u wide.
Life history —Probably similar to that of ZZ. gallinae (p. 54).
Distribution.—Asia (India (Zoological Garden, Calcutta) ).
HETERAKIS MACROURA Linstow, 1883
Host —Megaloperdix nigelii.
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 600p 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, 150u and 110, long.
Female 12.6 mm. long by 660% wide. Tail 1/7.8 of total length.
Eges 72 long by 48, wide. Position of vulva not given.
Life history.—Probably similar to 7. gallinae (p. 54).
Distribution—Asia (Turkestan).
HETERAKIS PAPILLOSA (Bloch, 1782) Railliet, 1885
Synonyms.—Ascaris papillosa Bloch, 1782; Heterahis monticel-
liana Stossich, 1892; Heterakis stylosa Linstow, 1907.
This species has been greatly confused with Meterahis gallinae
(H. vesicularis). Railliet and Henry have consequently used the
name Heterakis monticelliana to differentiate it but that is not
allowable under the rules of nomenclature.
Hosts —Otis tarda and O. tetrav.
Location.—Colon and ceca.
68 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Morphology—Heterakis (p. 50): The original descriptions of
both Ascaris papillosa and Heterakis monticelliana are inadequate ;
that of H. stylosa is more complete. Baylis has examined speci-
mens of both 7. 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 6202 wide. Esophagus 1% of total
length. Caudal alae (figs. 91 and 92) wide, elliptical. Preanal
Fics. 89-92.—89, HETERAKIS LONGECAUDATA. @, CUTICLE; b, MALE TAIL.
AFTER LINSTOW, 1879. 90, H®TERAKIS MACROURA. MALE TAIL, AFTER
Linstow, 1883. 91, H®TERAKIS PAPILLOSA, MALE TAIL, VENTRAL VIEW.
AFTER LINSTOW, 1907. 92, HETERAKIS PAPILLOSA. MALB TAIL, LATERAL
VIEW. AFrTHPR STOSSICH, 1892
sucker very prominent, 220 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,
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 BIRDS 69
Life history—Probably similar to that of H. gallinae (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.
4 P=
One
ales Oss LY
U
CO
Fic. 93.—HETERAKIS PARISI. MALE TAIL. AFTER BLANC, 1914
Male 7 to 9 mm. long by 350 to 400 wide. Caudal extremity (fig.
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 6402 long, provided with large membranous alae 45, in trans-
verse diameter.
Female 10 mm. long by 340 to 3604 wide. Tail very slender.
Vulva slightly anterior to middle of body. Eggs 68 long by 45.
wide.
Life history—Probably similar to that of H. gallinae (p. 54).
Distribution —KEurope (France (Dijon) ).
HETERAKIS PSOPHIAE Travassos, 1913
Synonym.—Ganguleterakis psophiae (Travassos, 1913) Lane, 1917.
Host—Psophia viridis.
Location.—Intestine.
70 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
>
Morphology.—Heterakis (p. 50): Mouth with 3 small equal lips.
Esophagus 1.13 to 1.26 mm. long, with small bulb.
Male 10 mm. long. Cloacal aperture (fig. 94) 352, from posterior
end. Preanal sucker circular, about 90u 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, long.
Female 12 mm. long. Anus 800 to 9002 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 52p
long by 39 wide.
Life history—Probably similar to that of H. gallinae (p. 54).
Distribution—South America (Brazil).
Fics. 94-95.—HETERAKIS PSOPHIAE. 94, MALE TaIL. 95, Vunva. AFTER TRAY-
ASSOS, 1913
HETERAKIS PUSILLA Linstow, 1906
Hosts —Gallus gallus and Gallus lafayettii. This species is listed
from the chicken by Gedoelst. Linstow described it from G. lafay-
ettiz and there appears to be no definite record from the chicken.
Location.—Ceca.
Morphology.—Heterahis (p. 50): Head with small rounded lips.
Male 5 mm. long by 190u 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 2404 wide. Esophagus length equal to
5/29 of body length. Tail length equal to 2/19 of body length; long
and pointed. Vulva 12/17 of body length behind head end. Eggs
thick-shelled, 65» long by 31 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 PARASITES OF BIRDS 71
HETERAKIS PUTAUSTRALIS Lane, 1914
Host.—Gallus gallus.
Location.—Ceca.
Morphology.—eterakis (p. 50): Small worms, curved dorsally
at the cephalic end. Lateral membranes present, extending almost
the entire length of body.
Male 7.6 mm. long by 800% 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
!
il
|
li
Fics. 96-98.—96, HETPRAKIS PUSILLA. MALe TAIL. AFTER Linstow, 1906. 97, Herrra-
KIS PUTAUSTRALIS. MALE TAIL. 98, a, LEFT SPICULE FROM. SIDE; 0, LEFT SPICULE FROM
CENTER ; C, RIGHT SPICULE. 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. 98¢) 550.
long and tapering to a blunt conical point. Left spicule (fig. 98 a
and 6) 2604 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 the
middle of the body. Anus 830, from tip of tail. Eggs 65p long by
37m wide.
Life history.—Probably similar to that of WZ. gallinae (p. 54).
Distribution —aAsia (India (Bengal) ).
HETERAKIS SKRJABINI Cram, 1927, new name
Synonym.—Heterakis arquata Schneider, 1866, of Skrjabin, 1916.
Host.—Tinamus, species.
Location —Abdominal cavity.
ie 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 700u wide. Esophagus 1.125 mm. long,
its cylindrical part 2384 wide; bulb 272y long by 300 wide. Tail
(fig. 99) pointed, provided with caudal alae; cloacal aperture 420.
from posterior end. Preanal sucker 230n in diameter, its posterior
border 6304 from the caudal extremity. Thirteen pairs of caudal
papillae of which 2 are preanal, 7 adanal, 4 postanal. Spicules equal,
850n 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 255y long by 272 wide. ‘Tail
conical, anus 2204 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 Z'inamus,
species of Paraguay as Heterakis arquata, he had not seen Travassos’s
paper (1913) but based his identification orf 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 cwpreus.
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) _| 90u_-..-.1-_-._-__2 2 . 230p.
Cloaeal aperture, from end} 270p. #22 eee 420u.
of tail.
Spicuiess: 67). SUMU y= Very long (according to | Relatively short (850,).
figure over 2 mm.)
Caudal papillae.-...____- 12 Dain ee eo ee 13 pairs.
eee female from tail, |" 1.12) mm. 2 220z.
end.
Wer eg a8 ge hs en ysl A little anterior to mid- | In anterior fifth of body.
dle of body.
Dee Aer ss seer 52p by 29p02 222.20 Aape 76.54 to 85u by 47x.
<i
og
NEMATODE PARASITES OF BIRDS
HETERAKIS TENUICAUDA Linstow, 1883
Hosts —Caccabis petrosa and Perdix graeca.
Location.—Intestine.
Morphology.—Heterakis (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 5524 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
a2
Figs. 99-101.—99, HrrreraKIS SKRJABINI. MALE TAIL. AFTER SKRIABIN, 1916. 100,
HETERAKIS TENUICAUDA.. MAL TAIL. AFTER LINSTOW, 1883. 101, MALE TAIL. AFTER
SEuRAT, 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 960» long, according to Lin-
stow, or 890» according to Seurat.
Female 14 mm. long by 600» 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 672 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 H. gallinae (p. 54).
Distribution.—Asia (Turkestan) and Africa (Algeria (Medea) ).
HETERAKIS VALVATA Schneider, 1866
Hosis.—Crypturus cupreus and C. noctivagus.
Location.—Ceca.
Morphology.—fHeterakis (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 alae small. Preanal
sucker 90u 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.
Fies. 102-103.— HETERAKIS VALVATA. 102, MALE TAIL. AFTER TRAVASSOS, 1913. 103,
MALE TAIL. AFTER SCHNEIDER, 1866
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 74u long by 37 wide.
Life history.—Probably similar to that of H. gallinae (p. 54).
Distribution—South America (Brazil).
(?) HETERAKIS VALDEMUCRONATA (Molin, 1860) Stossieh, 1887
Synonym.—A scaris valdemucronata Molin, 1860.
Hosts.—Ciconia maguart and EHuxenura 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 795
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.—Heterakis (p. 50): Head with 8 lips. Buccal cay-
ity present. Esophagus with bulb at posterior end. Lateral alae
present, very narrow (16.64), extending from a point about 153,
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 mm. long by 365» maximum width. Buccal cavity 41.5
deep ; esophagus, including bulb, 1.0 mm. long; buib 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
sucker 83, external diameter 100 to 120n. 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 3984 wide. Buccal cavity 50,
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 83p long by 42 wide.
Life history —Probably similar to that of 4. gallinae (p. 54).
Distribution —North America (United States (Pennsylvania) ).
Type material —No. 26376, U.S.N.M. (Bureau of Animal Industry
Helminthological Collections).
T am indebted for this material from the ruffed grouse to Dr. K. L.
Brunett, Cornel! University.
HETERAKIS NATTERERI Travassos, 1923
Host—Crax blumenbachi.
Location.—Ceca.
Morphology—Heterakis (p. 50): Head with 3 lips, vestibule 25y.
to 28. deep; total length of esophagus 6002; pyriform bulb 170» in
diameter, with heavily chitinized valves.
76 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 4mm. 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
a.
a
O-3 17/77
Fics. 104-107.—HETERAKIS BONASAD. 104, MALE TAIL. @, VENTRAL VIEW; J,
LATPRAL VIEW. 105, VULVA. HBETERAKIS ACUTICAUDA. 106, MALm TAIL.
107, FEMALE TAIL. Nos. 106-197 arrpR 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 4002 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
300n from the anus, anterior branch to 300n from the esophageal
bulb. Eggs 634 long by 354 wide. Tail conical, elongated; anus
300u from extremity.
Life history—Probably similar to that of H. gallinae (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. Yemale about 12 mm. long by 3383p wide.
Tail (fig. 107) straight, pointed.
Distribution—Europe (London Zoological Garden).
No further study 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
nN
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
KEY TO SPECIES OF ASCARIDIA
. Inadequately described; female 85 mm., male 53 mm, long; from Gallus
Calis, MUSE a lia =e ae te aie oo eae Ascaridia compressa, p. 88.
Description more.adequate fhan above: 2292s) ae eee 2:
. Male with 18 pairs of caudal papillae, in 2 unusually closely-set rows;
spicules equal, 6804 long by 2.44 wide; from Macropygia nigrirostris,
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 aboye__________________ ae
3:4Npicules (deseribed (asi Um ec ees a ey Ee aaa ae ee ae 4,
SPiculeSsequal Orss ue aay sss ae eee ee ee 5.
4. Female 39 mm., male 40 mm. long; 9 pairs of caudal papillae; from Capri-
MULGOUS-COMPESUVISLLEVO ie Los) oe) ais Ascaridia amblymoria, p. 83.
Female 55 mm., male 43 mm. long; caudal papillae number 10 pairs plus
an occasional unpaired papilla; from Vetrao 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_2 22s Tosi 16.
6. No preanal caudal papillae but 8 pairs of postanals.
Ascaridia dolichocerca, p. 90.
Preanal jcaudal,,papillae, present—s+——_+.—.---_s-__ plc ae re Gs
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.
8. Esophagus 1/14 of total body length; 4 pairs of preanal caudal papillae;
co
10.
LAR
13.
14.
tail of female 1/65 of total body length _____ Ascaridia compar, p. 88.
Esophagus 1/12.8 to 1/12.8 of total body length; 3 pairs of preanal caudal
papillae; tail of female not described in one species, 1/133 of total body
Tenet pin SOPOT SPC CHES ess Te fas ac eral ped elec eee 9,
. Female 90 to 100 mm. long; male 50 mm. or longer; ventral surface of
male tail covered with conspicuous granulations.
Ascaridia styphlocerea, p. 100.
Female 62 mm. long; male 40 mm. long; male tail without granulations.
Ascaridia francolina, p. 92.
Male’ with 6 or Tépairs,of.postanal spapillae=«- - de
Male with not more than 5 pairs of postanal papillae__________________ 14.
Spicules 570u 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
SUT GI 2 25 Ne se WI aM 2d Ds ental lh OP on Di ee a A?
. Body not described as having lateral alae; spicules 3.63 mm. long; tail
of male 1/77 of total length; eggs 96u long by 57u 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__________--_ 3.
Only male known, 80 mm. long; papillae scattered over the whole body;
spicules subequal, 2.1 and 1.9 mm. long___Ascaridia longecirrata, p. 94.
30th male and female known. Male 21 mm. long; no papillae described
on body as above; spicules equal, 1.05 mm. long.
Ascaridia catheturina, p. 8d.
Vulva posterior to middle of body, dividing body length in ratio of 15:11;
SHICUIES (90m. OMS ss eee Ascaridia aegyptiaca, p. 83.
Vulva slightly anterior to middle of body; spicules 1.7 mm. or longer__ 15.
15.
16.
ty
-I
NEMATODE PARASITES OF BIRDS 79
Female: tail 1/35, male tail 1/70 of total body length; eggs 91yu long by
57u wide; spicules subequal, 2.06 and 2.37 mm. long.
Ascaridia cordata, p. S88.
Female tail 1/67, male tail 1/48 of total body length; eggs 73u long by
42u wide; spicules equal, 1.74 mm. long___-__ Ascaridia circularis, p. 86.
From Gallus gallus; male only, 24 mm. long by 600u% wide; preanal sucker
110% in diameter 201) fe siee es Dubie Ascaridia brasiliensis, p. 84.
From other hosts than above or, if from Gallus gallus, total length of male
30 mm. or more and sucker over 200u in diameter_________________ 1%.
. 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 3404 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 ;302 pairs jof cervical: papillae: sk 20.
Body without numerous cervical papillae, or none described___________ ails
20. Tail of female 1/78 of total body length; spicules 950u long by 42u wide.
Ascaridia cristata, p. 89.
Tail of female 1/30 of total body length, or longer; spicules 1.2 to 1.9
minwlong tsi omies Oke Ascaridia columbae, p. S86.
21. Male only known; 9 pairs of postanal papillae; spicules 820u and 827u
| ae SR NE eee ee ee ee Ascaridia anseris, p. 101.
Both male and female known; male with less than 9 pairs of postanal
papillae; spicules, if length given, different from aboye______________ woe
22. Tail of female 1/117 of total length; eggs 172u long by 146u wide__________
Ascaridia stroma, p. 99.
Tail of female, if length given, longer than above; eggs considerably
Smoller than nbove=. 252 228 ese fe a 23.
Poa Vilva. POSLErlOr. cto middle of bodys. ==. - 2-2...
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.__iiine so 25.
25. Spicules equal, 3 mm. long; eggs 98u by 538u--____ Ascaridia numidae, p. 96.
Spicules subequal, 1.95 and 1.97 mm. long; eggs Sluw long by 47m wide___-_
Ascaridia trilabium, p. 101.
26. Male and female both 41 mm. long. Vulva at anterior third of body. In
Penclope humeralis____..._-._____--___ Ascaridia serrata, p. 98.
Total length usually greater than above; vulva usually more posterior than
aboye,;:-in sother ‘hosts;than;; above wos.) sons ef ee ee 2H.
2f.. Preanal sucker; not. over 162y, in, diameter....--.5-{oi2-c5. sb a 28.
Preanal sucker 200u or more in diameter _____-_________________ 29.
28. Male with tail 1/80 of body length and with 12 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 6094 long__Ascaridia pterophora, p. 97.
29. Female not over 52 mm. long; eggs 68u long; preanal sucker of male 340u
in diameter; spicules 3.4 mm. long__________ Ascaridia strelnikowi, p. 99.
Female 60 mm. or longer; eggs 754 or longer, except possibly in A. lin-
eata; preanal sucker not over 250u in diameter; spicules shorter than
3.4 mm. except in A. galli where they may possibly be 4 mm. long____30.
3612—27 os
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
netioven:2djeinvdiameter.. 2 Ue Ascaridia lineata, p. 93.
Body with lateral alae of usual type; male with 10 to 18 pairs of caudal
papillae; preanal sucker 220 to 2504 in diameter_______--_--___-__- ole
31. Cuticle with marked annulations which may be 100u wide; lateral alae 3
to 4 mm. long by 300u 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. dominicensis, P. festivus, P. leucoc., P. leucotis, P. menstruus, P.
pulverulentus, P. pertinax, P. phoenicurus, P. purpureus, 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,
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; Pusaria infleva Zeder, 1800; Ascaris inflera (Zeder,
1800) Rudolphi, 1809; Ascaridia infleca (Zeder, 1800) Dujardin,
1845; Heterakis inflexa (Zeder, 1800) Schneider, 1866; H. perspicil-
lwm (Rudolphi, 1803) Railhet, 1893; Ascaridia perspicillum (Ru-
dolphi, 1803) Dujardin, 1845.
Hosts.—Anas acuta, A. boschas domestica, A. b. fera, A. moschata,
Catrina moschata, Gallus gallus, Ithagenes cruentus, Numida mele-
agris, Phasianus gallus, Tetrao bonasia, T. urogallus, and, rarely, cat.
ATA AW
TO
QUO tii et aes
108.
ir
4 f/
> \ /
“SA
\-: “A MED =
Ke By eS
ht , ee ee
Figs. 108-109.—AScCARIDIA HERMAPHRODITA. 108, MALP TAIL. AFTER SCHNEIDER,
1866. 109, MaLw TAIL. AFTER SKRIJABIN, 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 are 10 pairs of caudal papillae (figs.
110 and 111), of which 3 pairs of pedunculated papillae are near the
sucker; this sucker (220, 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 80n long by 45 to 50u 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 lumbricoides 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.
NEMATODE PARASITES OF BIRDS 83
ASCARIDIA AEGYPTIACA (Linstow, 1902) Railliet and Henry, 1914
Synonym.—Heterakis aegyptiaca Linstow, 1902.
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 5904 wide. Esophagus 1/138.6, 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 78 long by 48» wide.
beat
03MM.
Fics. 110-113.—110, ASCARIDIA GALLI. MALE TAIL. AFTER SMIT, 1922. 111, MALD TAIL.
AFTER BAYLIS AND DAUBNEY, 1922. 112, ASCARIDIA AEGYPTIACA. MALE TAIL. AFTER
LINSTOW, 1902. 1138, ASCARIDIA AMBLYMORIA. MALE TAIL. AFTER DRASCHH, 1882
Life history—Probably similar to that of A. galli (p. 82).
Distribution —Africa (Egypt).
ASCARIDIA AMBLYMORIA (Drasche, 1883) -Railliet and Henry, 1914
Synonym.—LH eterakis amblymoria Drasche, 1883a.
Host.—Caprimulgus campestris.
Location.—Intestine.
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 cutinous (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—aAscaridia (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, 75u long by 42u wide.
Life history —Probably similar to that of A. galli (p. 82).
Distribution.—Oceania (Bismarck-Archipelago (Ralum)).
ASCARIDIA BOREALIS (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, 340u 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 600n 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
Figs. 114-117.—114, ASCARIDIA AUSTRALIS. MALE TAIL. AFTER LINSTOW, 1898. 115,
ASCARIDIA BORBALIS. MALE TAIL. AFTER LINSTOW, 1884. 116, ASCARIDIA BRASILIENSIS,
MALH 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) Railliet and Henry, 1914
Synonym.—Heterakis catheturinus Johnston, 1912.
Host.—Talegallus (Catheturus) lathami.
Location.—Intestine.
Morphology.—Ascaridia (p. 77): Anterior end rounded, narrow;
posterior end pointed in both sexes. Lateral alae extend for about
2mm. posterior to mouth. Three equal lips.
Male 21 mm. long. Caudal alae narrow. (Fig. 117.) Cloacal
aperture 400, from posterior end. Preanal sucker 275, 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 6 paired postanal
papillae, thus a total of 18 caudal papillae. Spicules slightly curved
and pointed, 1.05 mm. long by 22 wide.
Female 28 mm. long. Vulva at about middle of body, shghtly
salient. Anus 860. from posterior end.
Life history.—Probably similar to that of A. galli (p. 82).
Distribution—Burnett River (Australia ?).
ASCARIDIA CIRCULARIS (Linstow, 1903) Railliet and Henry, 1914
Synonym.—Heterakis circularis Linstow, 1903.
Host.—Centropus sinensis.
Location —Not given.
Morphology.—Ascaridia (p. 77): Mouth with 3 large semicir-
cilar lips, a dorsal with 2 papillae, the other 2 ventro-lateral.
Figs. 118-120.—-118, ASCARIDIA CIRCULARIS. MALE TAIL. AFTER LINSTOW, 1903. 119
ASCARIDIA COLUMBAE. MALE TAIL. @, VENTRAL VIEW; Db, SIDE VIEW. AFTER JOHNSTON
1918. 120, ASCARIDIA COMPAR. MALE TAIL. AFTER LINSTOW, 1899
Male 31 mm. long by 7904 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 126:129. Eggs 73 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.—Asearis columbae Gmelin, 1790; Ascaris maculosa
Rudolphi, 1802; Heterakis maculosa Schneider, 1866; Heterakis
columbae (Gmelin, 1790) Railliet, 1885.
NEMATODE PARASITES OF BIRDS 87
Hosts—Columba arquatrix, C. domestica, C. d. laticauda, C.
gutturosa, C. livia, C. picui, C. risoria, C. speciosa, C. talpacoti,
Crocopus phoenicopterus, Phlogoenas luzonica, Stictoenas arquatrix,
Turtur sylwaticus.
Chatin has reported this species from the pheasant and Sweet from
the chicken; Skrjabin thinks these writers probably had Ascaridia
galli (A. perspicillum).
Location.—Small intestine, usually; in esophagus, proventriculus,
cizzard, 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 Baylis 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
&00 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, possibly corre-
lated with host variation. Tail end obliquely truncated and
mucronated at the tip. Circular sucker, with chitinous walls, 150 to
200 long by 150 to 160u 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 mm. long; Castejon says usually 50 to 60 mm.
long in adult birds; Baylis and Daubney say 70 to 95 mm, long by
2.5 mm. wide. Vulva near the middle of body. Tail straight,
conical, mucronated. Anus 1.2 mm. from end of tail; posterior to
anus a longitudinal depression. Eggs 68 long, according to
Travassos; 80 to 90» long, according to Railliet, by 40 to 50” wide;
72 long by 48 wide, according to Irwin-Smith; 60 to 70u 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 that when such eggs
were fed to pigeons the young worms developed to adults in 3 weeks.
The occurrence of larval worms in the liver, reported by Bedel,
saggests 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 compar Schrank, 1790; Heterakis compar
(Schrank, 1790) Stossich, 1887; Ascaris lagopodis Froelich, 1802.
Hosts—Caccabis chucar, C. sawatilis, Colinus virginianus, Coturnix
communis, C. dactylisonans, Gallus gallus, Numida meleagris, Ortyx
virginianus, Perdix cinerea, Tetrao lagopus, 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, 3404 by 300u.
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
91p long by 57p wide, according to Linstow; 80n by 60n, 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.—G@allus 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.—H eterakis cordata Linstow, 1906c.
Host—Callipepla squamata.
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 8804 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 91p long by 57p wide.
Life history—Probably similar to that of A. galli (p. 82).
Distribution.—North America (Mexico).
a.
Fics. 121-122.—121, ASCARIDIA CORDATA. MALm TAIL. AFTER LINSTOW, 1906. 122,
ASCARIDIA CRISTATA. @, HEAD; 0, MALE TAIL. AFTER LINSTOW, 1901
ASCARIDIA CRISTATA (Linstow, 1901) Railliet and Henry, 1914
Synonym.—Heterakis cristata Linstow, 1901.
Hosts.—Belearica pavonina, 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 900p
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 1.84 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,
+ postanal; according to Baylis and Daubney, 13 pairs, of which
6 are preanal, 7 postanal. Spicules equal, 950n long by 424 wide,
alate.
Female 57 mm. long by 1.76 mm. wide, according to Linstow;
£8 to 40 mm. long by 1.1 to 1.2 mm. wide according to Baylis 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 91p long by 58 to 62 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 cylindrica 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.
Female 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 (1916a@) 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, 1914
Synonym.—tH eterakis dolichocerca Stossich, 1902.
Host.—Circus spilothorax.
Location.—Not given.
Morphology.—Ascaridia (p. 77): Head distinct from body, with
3 large lips. Lateral membranes wide. Length of specimens 25
to 50 mm.
NEMATODE PARASITES OF BIRDS Qi
Male with preanal 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 cloacal 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).
tos
Figs. 123—-125.—123, ASCARIDIA CYLINDRICA. MALE TAIL. 124, SPICULE. AFTER BLOME,
1909. 125, ASCARIDIA DOLICHOCERCA. MALE TAIL. AFTER STOSSICH, 1902
ASCARIDIA FASCIATA Baylis, 1920a
Host.—Vinago delalandii.
Location.—Not given.
Morphology.—Ascaridia (p. 77): Mouth (fig. 126) with 38 large
semicircular lips, the dorsal with 2 large papillae, the others with a
single median papilla. Cuticle of body (fig. 127) with thickened
transverse rings resembling annulations, their maximum width, in
middle of body, being 100u. Lateral alae well developed, 3 to 4
mm. long by 3002 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 (700u). Cuticle raised in 2 longitudinal ridges at the
sides of the sucker; sucker 2502 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
58 long by 454 wide.
Life history.—Probably similar to that of A. galli (p. 82).
Distribution—Kast Africa (Dar-es-Salaam).
ASCARIDIA FRANCOLINA (Linstow, 1899) Railliet and Henry, 1914
Synonym.—Heterakis francolina Linstow, 1899.
Host—F rancolinus bicalcaratus.
Location.—Not given.
Morphology.—Ascaridia (p. 77) : Esophagus 1/12.3 of total length.
j27 SN
Fics. 126—-128.—ASCARIDIA FASCIATA. 126, Heap. 127, CUTICULAR RINGS OF BODY. 128,
MALE TAIL, AFTER BAYLIS, 1920
Male 40 mm. long by 870» 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. gallz (p. 82).
Distribution—Africa (Togo and Bismarckburg).
ASCARIDIA GRANULOSA (Linstow, 1906) Railliet and Henry, 1912
Synonym.—Heterakis granulosa Linstow, 1903.
Host— Gallus gallus.
Location.—Small intestine.
Morphology—Ascaridia (p. 77): Large, thick worms.
Male 27 mm. long by 590 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 PARASITES OF BIRDS 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. galli (p. 82).
Distribution.—Asia (Ceylon (Colombo) ).
ASCARIDIA LINEATA (Schneider, 1866) Railliet and Henry, 1912
Synonyms.—Heterakis lineata Schneider, 1866; Ascaridia hamia
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 boschas 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 shght 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% wide.
Anal sucker 200 to 250u 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 540u 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;
Fies. 129-1382.—129,AscarIpDIA FRANCOLINA. MALE TAIL. AFTER LINSTOW, 1899. 130
ASCARIDIA GRANULOSA. MALH TAIL. AFTER LINSTOW, 1906. 131, ASCARIDIA LINEATA.
MALE TAIL. AFTER BOULENGER, 1923. 132, ASCARIDIA LONGECIRRATA. MALE TAIL.
AFTER LINSTOW, 1879
according to Lane, that of A. hamia is 1.8 mm. long and there are
lateral papillae 500 from the tip. Eggs elliptical, 804 long by 50s
wide; in A. hamia Lane says they are thick-shelled and 65« long
by 40« 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. haméa)), 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.—tHeterakis longecirrata Linstow, 1879.
Host.—G'eopelia, species.
Location.—Intestine.
NEMATODE PARASITES OF BIRDS 95
Morphology.—Ascaridia (p. 77): Cuticle cross-striated with pe-
euliar 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 8 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 1904 wide. Esophagus 1/11, tail 1/46 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 1.9 mm. long.
Female apparently unknown, as not described.
Life history.—Probably similar to that of A. galli (p. 82).
Distribution.—Not given.
ASCARIDIA MAGALHAESI Travassos, 1913
llost.—Geotrygon montana.
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 500n 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, 1.66 mm. long.
Female 41 to 48 mm. long. Vulva slightly anterior to middle of
body. Anus 1.07 mm. from caudal extremity. Eggs 74, long by
44u wide.
Life history.—Probably similar to that of A. ga/li (p. 82).
Locality.—South America (Brazil).
ASCARIDIA MAGNIPAPILLA (Linstow, 1906) Railliet and Henry, 1914
Synonym.—Heterakis magnipapilla Linstow, 1906d.
Host—Tetrao tetrix (Lyrrurus tetrix).
Location.—Intestine.
Morphology.—Ascaridia (p. 77): Cuticle with cross-striations and
additional deep contractions. Mouth with 3 wide low lips (fig.
1342) which measure 180u deep by 3704 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. 1345) wide. Nine pairs of large pedunculated
papillae of which 2 are preanal, 7 postanal. Preanal sucker 280,
wide by 310y long. Spicules 3.63 mm. long, with 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, 96u long by 57 wide, with
thick shells.
Life history—Probably similar to that of A. gall (p. 82).
Distribution.—Kurope (Prussia (Friedland) ).
ASCARIDIA NUMIDAE (Leiper, 1908) Travassos, 1913
Synonyms.—Heterakis 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,
Fies. 188-185,—1338, ASCARIDIA MAGALHAESI. MALE TAIL. AFTER TRAVASSOS, 19138. 134,
ASCARIDIA MAGNIPAPILLA. @, HEAD; b, MAL TAIL. AFTER LINSTOW, 1906. 135,
ASCARIDIA NUMIDAE, 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 sahent transverse slit, posterior of
the middle of the body. The eggs are 98» long by 53» 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. 186 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 24 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-
cauda Molin, 1860; Heterakis laticauda (Molin, 1860) Stossich, 1887.
Hosts—Cariama cristata (Dicholophus marcgraft) and Micro-
dactylus 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 (71. laticauda) or 20 mm. long by 500p
wide (A. pterophora). Caudal extremity (fig. 138) broad; cloacal
aperture 360. from posterior end in a 20 mm. specimen. Caudal
alae narrow. Preanal sucker large (100p in diameter), with a pa-
pilliform nodule in the posterior rim; it is 5202 from the posterior
end of body. Ten pairs of caudal papillae, 4 of which are preanal
(3 pairs ventral, 1 pair lateral). Spicules equal, 6002 long by 2ipn
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).
O
Ae?
os
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oS d SONS
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SFT Tid
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ie
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Kies, 156—-139.—136,
ASCARIDIA ORTHOCERCA.,
TAIL, LATERAL VIEW.
MALE TAIL, VENTRAL VIEW. 137, MALE
AFTER STOSSICH, 1902. 138, ASCARIDIA PTEROPHORA. MALE TAIL.
AFTER TRAVASSOS, 1918. 139, ASCARIDIA SERRATA. MALDP TAIL.
AFTER SCHNEIDER, 1866
ASCARIDIA SERRATA (Schneider, 1866) Railliet and Henry, 1914
Synonym.—H eterakis 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. 189), 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. galli (p. 82).
Distribution.—South America (Brazil).
ASCARIDIA STRELNIKOWI Skrjabin, 1916
Host.—Tinamus, species.
Location.—Intestine.
Morphology—Ascaridia (p. 77): Cuticle with fine transverse
striations. Lips (fig. 1402) 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 136y long
by 250n wide. Esophagus 2.88 mm. long by 220u wide. Caudal ex-
tremity (fig. 1406) conical, pointed; cloacal aperture 6004 from
posterior end. Preanal sucker circular, 340u 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 187y 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 (Tetrapteryx) paradisea, G. antigone and G. com-
munis.
Location.—Not given.
Morphology—Ascaridia (p. TT): 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 56 mm. long by 1.7 wide. Tail 1/117 of total length,
with digitiform prolongation. Eggs 172y long by 146 wide.
100 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history.—Probably similar to that A. galli (p. 82).
Distribution —Kurope (Germany (Museum Berlin)) and Asia
(India).
ASCARIDIA STYPHLOCERCA (Stossich, 1904) Railliet and Henry, 1914
Synonyms.—H eterakis styphlocerca Stossich, 1904.
Hosts—This worm was described from a domestic bird (“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.
Figs. 140-142.—140, ASCARIDIA STRELNIKOWI. @, HEAD; b, MALY TAIL. AFTHR 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
eranulations. (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 PARASITES OF BIRDS 101
as an asymmetrical papilla, was a symmetrical pair, making 6 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
232u 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
yiomm
Fies. 143-144:—148, ASCARIDIA TRILABIUM. MALE TAIL. AFTER LINSTOW, 1904. 144,
ASCARIDIA ANSERIS. MALE TAIL. AFTER SCHWARTZ, 1925
Synonym.—HLHeterakis 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 26 mm. long by 6902 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 81p 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.—Anser domesticus.
Location.—Small intestine.
Morphology.—Ascaridia (p. 77).
Male 32 mm. long by 600 wide. Head, separated from body, 172
wide near base. Esophagus 1.75 mm. long by 2854 in maximum
width. Nerve ring about 3504 from head end. Preanal sucker
circular, 1384 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, 8202 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, nof 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.
Ty pe-species.—P seudaspidodera 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 PAVCNIS Baylis and Daubney, 1922
Hosts.—Pavo cristatus and Pavo muticus.
Location.—Not given; presumably ceca.
Morphology—Pseudaspidodera (p. 102): Small worms. Head
(fig. 145) with cordons as described in generic diagnosis. Diameter
of head at posterior limit of cordons, about 100u. 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 1904 wide and containing the usual valves.
Nerve ring 400 to 4604 from head end. Excretory pore 600 to 650
from head end.
Male 6 mm. long by 250% wide. Esophagus 1.4 to 1.48 mm. long.
[76
0-3 MM.
03mm.
Figs. 145-147.—PSEUDASPIDODERA PAVONIS. 145, H@®AD, LATERAL VIEW. 146, MALE
TAIL. LEFT, LATERAL VIEW; RIGHT, VENTRAL VIEW. 147, VULVA AND VAGINA IN
LATERAL VIEW, SHOWING CEMENT PLUG IN VULVA. ARROW POINTS IN DIRECTION OF
HEAD. AFTER BAYLIS AND DAUBNEY, 1922
Tail (fig. 146) 380 to 430u 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 170, anterior to cloacal aperture; the
greatest diameter (antero-posterior) of sucker aperture is 70x.
Spicules unequal and dissimilar. Right spicule slender and simple,
(80p 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 300n 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
tailend. 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, 70u long by 40 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 Susunurrnar Travassos, 1914
Synonym.—Kathlaniinae Lane, 1914.
Subfamily 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
Synonyms.—Ascaris Linnaeus, 1758, part; Weterakis Dujardin,
1845, part; Oxyuris Rudolphi, 1803, part; Al/odapa Diesing, 1861.
Generic diagnosis—Subulurinae (p. 104) : Head rarely 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. J/ale 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 SUBULURA
1. Only female known; tail short (38202) : from Turnix, species.
Subulura, species Baylis and Daubney, p. 132.
Male known; tail of female longer than above (4434 or longer except pos-
sibly in S. forcipata where length not given and in S. papillosa 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 aboye____________ 3.
em US CUMS UIT CUT SR Bae ee ee ee 4,
Sy ULC SCOUT SR eee eee et ee BPA.
4. Longer spicule 840u in length; 9 pairs of caudal papillae; female 9.3 mm.
ANG oY eS Se cee a ee ee ee Subulura rima, p- 125.
Longer spicule over 1 mm. in length; 10 or 11 pairs of caudal papillae;
female: OMe ORAM LOR. =. ee ee ee 5.
5. Lateral alae said to be absent; 10 pairs of caudal papillae; gubernaculum
PAOLO) ci 1 scare aa ae a 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, 144u
STV ONS oe co a a ke ee 6.
6. Long spicule 2.54 mm. in length; lateral alae about 930 long.
Subulura carlosi, p. 115.
Long spicule not over 1.52 mm. in length; lateral alae 1.27 mm. or longer
(possible exception S. curvata where not described) __~___-_-____-_--_-_-_- i
. 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 443u long________________- Subulura lutzi, p. 11S.
ail OL temeleski mm. or loOn@er.-22 222.228 Os feo ee ee 9.
9. Lateral alae extend to middle of body in male, 144 of body length in
female, thus for a length of at least 3 mm.; cloacal aperture of male
169%, from. posterior end__....--.-- 2-25... 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 850u long; eggs 55u long by 45u wide.
Subulura seurati, p. 126.
Shorter spicule 1 mm. or longer; eggs 76u long by 50u wide or 83u long by
ESTELLE ee eee th ot Om 11.
106 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Atk
13:
14.
15.
16.
ore
19.
20.
24.
Preanal sucker of male 2804 long, its posterior edge 8834 from tail end;
ovejector of female 899 long__________=___ Subulura bentocruzi, p. 110.
Preanal sucker of male 1438 long, its posterior edge 508 from tail end;
ovejector of female 1.28 mm. long________ “_ Subulura reclinata, p. 124.
2 Male with *8*or"9" pairsor CauGgal papillae’ eee eee ee TSE
Male with 10 or more pairs ‘of ‘caudal papillae___-=_4) 2 =f tt ee 15.
Spicules 590 long; tail of female 1.16 mm. long_-_ Subulura gracilis, p. 134.
Spicules 880 long; tail of female not over S00u long_---_----____--_-- 14.
Male 10.5 mm., female 14.8 mm. long; male tail 1/38, female tail 1/18.7 of
COLA DOUY: LON SEN ek aes See Subulura acuticauda, p. 133.
Male 6.78 mm., female 9.26 mm. long; male tail 1/25.7, female tail 1/14 of
total. bodyrlenoth: t=. se te ee ee eee Subulura recurvata, p. 124.
Male with 10 pairs‘of'caudal papillae.) -s2- 2242) S+24_ 22s een gee 16.
Male with 11 or more pairs of caudal papillae-______~_-______-~__=--=.- 23.
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
OE EDO Ny a ea ee er ee se en ee AG
Spicules*noettover 1902 long. ees Sr a ee ee ee a eee 18.
Spicules 880; or longer 2. = 2 425 Me ae ee ee eee ee eee 20.
. Male 4.5 mm., female 8.5 mm. long; spicules 790u long; eggs 654 long by
49 LA WwAdesuees 2606 Os IA eS eee Mee SUbitlura poculums, pyckas:
Male 8.2 mm. or longer; female 13.6 mm. or longer; spicules not over
700u long; eggs not over 50u long by 40n wide____--_----__---_---__ £9:
Male 8.2 mm., female 13.6 mm. long; gubernaculum rectilinear; ovejector
short, its total length 1.28 mm., the trompe being 385y 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.
Spicales! Mot OVI GOO ts LOM ee ae a ee ee Pill.
Spiculées* over a” mim? Tong a re ee eee a a 22:
. Cloacal aperture of male 3184, anus of female 600u from end of tail; eggs
AD OOM DY? Ala) WIGea= 2 Se aes eee ee ee Subulura rimula, p. 125.
Cloacal aperture of male 200u, anus of female SOOu from end of tail; eggs
65. to 5p lone by 520 00m .WiGe=- =e Subulura plotina, p. 122.
. Cloacal aperture of male 170z, posterior end of sucker 5004 from tail end;
gubernaculum 100u long; eggs 59u long by 50u wide.
Subulura differens, p. 111.
Cloacal aperture of male 250 to 315, posterior end of sucker 590 to S15z
from end of tail; gubernaculum 175 to 210u long; eggs 75 to 80x long
by 65 to 70u wide_______ at 2 eee eee HORST ULULA, DIUM tl Deus
. Lateral alae extend entire length of body; male with 13 pairs of caudal
papillae; female 21 to 35 mm. long, its tail only +; of total body length.
° Subulura leprincei, p. 118.
Lateral alae extend not farther than first part of intestine (except possibly
in S. halli where alae not described); male with 11 pairs of caudal
papillae; female not over 19 mm. long (except possibly in S. suctoria
where may reach 23 mm.), its tail being longer than above_---------- 24.
Posterior end of preana] sucker 860y, or farther, from caudal extremity_ 25.
Posterior end of preanal sucker not over 50u from caudal extremity_-__ 26.
NEMATODE PARASITES OF BIRDS 107
25. Cloacal aperture of male 5004 from end; spicules 1.5 mm. long; tail of
female 1/8.5 of total length; eggs SOu long by 60u wide.
Subulura halli, p. 117.
Cloaecal aperture of male 210u from end; spicules 760 to S800” long; tail
of female 1/11 to 1/12 of total length; eggs 65u long by 35u wide.
Subulura galloperdicis, p. 116.
26. Preanal sucker 385 long; spicules 1.71 mm. long Subulura travassosi, p. 131.
Preanal sucker not over 1S86u 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 __~___ 29.
28. Preanal sucker S4u long; spicules S899u long by 16u wide; gubernaculum
127 long; tail of female 976 long__-_________ Subulura olympioi, p. 121.
Preanal sucker 1864 long; spicules 5584 long by 38xe wide; gubernaculum
228u long; tail of female 1.2 mm. long________ Subulura trogoni, p. 131.
29. Lateral alae extend to median part of bulb; preanal sucker 169u long;
ovejector of female 950u Jong, the trompe equivalent to 24 of its length.
Subulura strongylina, p. 128.
Lateral alae extend to middle of esophagus; preanal sucker 135 long; ove-
jector of female 779u long, the trompe equivalent to less than 14 of
its length
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
fromthe Second palrse—-—- #7 bese 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 multipapillata recently
described by Chandler (see Addenda, p. 388).
SUBULURA ACUTISSIMA Molin, 1860
Synonyms.—P hysaloptera saginata strigis brasiliensis !; P. stron-
gylina cuculi-senicult*; Heterakis acutissima (Molin, 1860) Stossich,
1887.
Hosts —Cuculus seniculus, Coccyzus melacoryphus, Pisorhina
atricapilla, Strix atricapilla,
Location.—Proventriculus and intestine.
Morphology —sSubulura (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 mm. long by 300 wide. Caudal extremity (fig. 152)
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.
1Catalogue of the Vienna Museum.
108 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Female 12 mm. long by 300 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-
\ la
i
\ Dy
%,
: fy
tf
(i
mITAL
A
iS
O5 S717
Fics. 148-150.—SuBULURA DIFFERENS. 148, ANTERIOR END. ORIGINAL. 149, a, VULVA;
b, FEMALF TAIL; ¢c, OVEJECTOR; d, EGG. (SCALH THE SAME FOR @ AND 0.) ORIGINAL.
150, MALE TAIL. X37.5. AFTER BAaRRETO, 1918
paria Schneider, 1866, part; Heterakis allodapa (Creplin, 1853)
Seurat, 1914, part; Al/odapa 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.—SUBULURA ACUTISSIMA. 151, HEAD. a, SIDE VIEW; b, FRONT VIEW. 152,
MALE TAIL. AFTER DRASCHE, 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 1694 from end. Preanal
sucker 169 long, without rim, 719» from posterior end of body.
Eleven pairs of papillae, of which 8 are preanal and ventral, 2
A. SC
~~ © 3
Q ———— (*)
Q O wy
NS INS
r\\
—_—
or titya
» xy PTY A
ESRI OTe
Fics. 153-156.—SuBuLuRA ALLODAPA. 153, ANTERIOR END. AFTER BARRETO, 1918. 154,
HPAD, FRONT VIPW. AFTER DRASCHE, 1882. 155, MALp TAIL, AFTER DRASCHE, 1882.
156, a@ AND Db, OvEsECTOR. AFTER BArRETO, 1918
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 152, long.
Female 10 to 14 mm. long by 440n 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 slightly salient, of irregular shape, a little anterior to middle
of body. Ovejector (fig. 156) anteriorly directed, about 762» long;
vestibule short, 3134; sphincter, 2114; trompe 338p 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 7. variegatus.
Location.—Intestine.
Morphology.—sSubulura (p. 104): Cuticle with transverse stria-
tions. Lateral alae extend slightly beyond esophageal bulb, their
Fies. 157-158.—SUBULURA BENTOCRUZI. 157, MALE TAIL, LATERAL Virw. 158, MAL TAIL,
VENTRAL VIEW. AFTER BARRETO, 1918
length about 1.97 mm. Mouth hexagonal, with 6 small equal papil-
Jae. 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, 280u long, its lower limit 8834 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. (44 the length of the for-
mer.) Gubernaculum 169, long.
NEMATODE PARASITES OF BIRDS Lit
Female 13 to 21 mm. long by 6434 wide. Tail with appendage
140n long. Anus 1.16 mm. from posterior end. Vulva salient, ante-
rior to middle of body. Ovejector relatively long (599p); vestibule
334 long; sphincter small; trompe 541p long. Two uteri divergent.
Kggs elliptical, 834 long by 49» wide.
Life history.—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—South America (Brazil).
SUBULURA DIFFERENS (Sonsino, 1890) Railliet and Henry, 1912
Synonym.—Heterakis differens Sonsino, 1890.
Hosts.—Francolinus bicalearatus, Gallus gallus, and Numida mele-
agris. 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 wide. Cloacal aperture 166 to 190.
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, long.
Female 11.3 to 19 mm. long by 335 wide. Anus (fig. 1496) 631
to 770p from tailend. Vulva (fig. 149@) 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
3612—27-——_9
PZ BULLETIN 140, UNITED STATES NATIONAL MUSEUM
species. Barreto says the ovejector of S. differens 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. sewrati (p. 126)
and S. leprinct (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, 594 by 50., thin-shelled, containing an embryo when
oviposited (fig. 149d).
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.—G@allus 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
14 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 420u wide. Tail ends in a pro-
longation about 14 of length from cloacal aperture to end of tail.
Caudal sucker 170 to 220n 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. 1595), 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 14 of the distance from the posterior border
of the sucker to the cloacal aperture, and a fifth pair is at the side
NEMATODE PARASITES OF BIRDS 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, 650n to 1 mm. long, terminating in a point
100 long. Vulva slightly salient, 4.3 to 5.4, or 6.8 mm. in Seurat’s
specimens, from the head end, or slightly anterior to the middle of
the body. Ovejector directed anteriorly, 9802 to 1 mm. long, the
vestibule 600u long, and the sphincter 380p long. Eggs subspherical,
SOOM
(9G fi 0
Figs. 159-161.—SuBULURA BRUMPTI. 159, a, ANTERIOR END; 0, MALE TAIL,
AFTER LOPEZ-NEYRA, 1922. 160, MALE TAIL. AFTER GEDOELST, 1916. 161,
MALE TAIL. AFTER GPNDRE, 1909
with smooth shell, 75 to 804 long by 65 to 70 wide, and containing
an embryo when deposited.
Life history—Unknown; probably similar to that of Asearidia
galli (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 alae 930 long by 51p wide, extending to level of
posterior end of esophagus. Mouth with small lips armed with 6
papillae, 2 of which are large and the other 4 small. Mouth cavity
short; esophagus about 1.1 mm. long; bulb subspherical.
114 BULLETIN 140, UNITED’ STATES NATIONAL MUSEUM
Male 9 to 10 mm. long by 290» wide. Caudal extremity (fig. 162)
curved ventrally; caudal alae much reduced; cloacal aperature 186p
from posterior end. Preanal sucker without chitinous ring, 212»
long, its posterior end 465. 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» long by 25.
wide. Gubernaculum 169y 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 24 of body length from head). Ovejector (fig.
cr
SIS.
~~, UA
Cc)
Vw
ga TIER oe SETH
RODS YW
Zoi a ye
ZARMINGAS a
ASTIN
D Beh
ex ep
I'ies. 162-163.—SvuBULURA CARLOSI. 162, MALE TAIL. @, SIDE VIEW; 0, VENTRAL VIEW.
163, VULVA AND OVEJECTOR. AFTER BARRETO, 1918
163) long (1.14 mm.); vestibule 2544 long, made up of 2 charac-
teristic parts, with chitinous lining; sphincter short (135). Two
divergent uteri. Eggs elliptical, 84u long by 674 wide, embryonated
when deposited.
Life histery—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution —South America (Brazil).
SUBULURA CURVATA (Linstow, 1883) Railliet and Henry, 1914
Synonym.—H eterakis curvata Linstow, 1883.
Hosts.—Caccabis 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 5402 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. 164). Of the postana!
papillae, 4 are ventral, 3 lateral. Spicules 1.2 mm. and 900p long
respectively.
Female 12.8 mm. long by 540» wide. Tail 144 of body length,
pointed. Vulva slightly posterior to middle of body, dividing body
length in ratio of 14:13. Eggs 39p by 26u wide.
Life history.—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution —Asia (Turkestan (by Linstow) and Russian Turk-
estan (Aoulie-ata; by Skrjabin) ).
SUBULURA FORCIPATA (Rudolphi, 1819) Raillict and Henry, 1914
Synonyms.—A scaris forcipata Rudolphi, 1519, part; Ascaris foret-
paria Schneider, 1866, part; Heterakis forciparia Schneider, 1866,
part.
Hosts —Bucco, species, Capito collaris, C. macror., C. melanoleucus,
C. rufiventris, C. striolatus, C. tamatia, Coccyzus melanocoryphus, C.
minor, Cuculus naevius, C. seniculus, C. tingazu, Caprimaulgus
bacaurau, C’. nacandua, C. ruficollis, C. urutau, Dicholophus eristatus,
Diplopterus naevius, Guira guira, Monasa leucops, M. tranquiila, M.
tenebrosa, M. torquata, Piaya cayana, Tetrao wi.
Railliet and Henry say that the reports from Bucco and the
Caprimulgides are probably misidentifications.
Location.—Intestine and ceca.
Morphology. Subulura (p. 104) : Cuticle with transverse striations.
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 80g long at its
hase. Esophagus 1.03 mm. long, bulb spherical, 170 in diameter.
Male 8.2 mm. long by 2822 wide. Tail (fig. 165) strongly recurved,
its alae rudimentary ; cloacal aperture 211, fromend. Preanal sucker
without chitinous ring, elliptical, 177» long, 5924 from caudal ex-
tremity. Ten pairs of papillae of which 3 are preanal, 2 adanal, and
5 postanal. Spicules equal, 677 long. Gubernaculum 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. 166) short (1.28 mm.), vestibule pyriform, strongly
chitinized, 514p 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.—G@alloperdix spadicea.
Location.—-Intestine.
Morphology.—Subulura (p. 104): Head small, buccal cavity 60p
deep by 234 wide anteriorly, with 3 teeth at base. Esophagus 1.5
mm. long, exclusive of bulb; bulb 200» in diameter. Lateral alae
narrow, extend for about 1 mm. from anterior end of body.
Male 9.5 to 10 mm. long by 300% wide. Tail 210» long, drawn
cut to fine point. Sucker spindle-shape, 650, anterior to cloacal
W/)
(
\
Sry D
Fics. 164—166.—164, SuBuLuraA cURVATA. MALE TAIL. AFTER LINSTOW, 1883. 165,
SUBULURA FORCIPATA. MALE TAIL. 166, VULYA AND 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 800u long, alate,
tapering. Gubernaculum 180» long with a spur at about 60u from
its anterior end.
Female 11.5 to 12.5 mm. long by 4004 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 654 long by 35 wide, em-
bryonated at time of deposit.
Life history—Unknown; probably similar to that of Ascaridia
galli, (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.—sSubulura (p. 104): Pharyngeal teeth (fig. 168¢)
cuneiform, their free ends pointed.
Male 10.5 to 12 mm. long by 5404 wide. Esophagus, including
bulb, 1/6.5 of total length. Tail (fig. 168@) slender, ending in a long
fine point. Cloacal aperture 5004 from posterior end. Caudal alae
narrow. Eleven pairs of pedunculated papillae, 5 of which are
rx Err
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Z
H
i
!
ro ALM
Fics. 167-168.—167, SUBULURA GALLOPERDICIS. MALE TAIL. AFTER BAYLIS AND DAUBNEY
1922. 168, SUBULURA HALLI. a, MALB TAIL; b, OVEJECTOR; C, BUCCAL CAVITY. AFTER
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% 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 25 of its length. Ovejector
(fig. 1686) with remarkably short sphincter. Eggs 80p long by
60u wide.
Life history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—Africa (Algeria (Maison-Carrée) ).
118 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
SUBULURA LEPRINCEI (Gendre, 1909) Travassos, 1913
Synonyms.—Heterakis leprincet Gendre, 1909b; Allodapa leprin-
cei (Gendre, 1909) Seurat, 1914.
Hosts —Caprimulgus aegyptius saharae; C. fossii; Macrodipteryx
macrodipterus.
Location.—Ceca.
Morphology—Subulura (p. 104): Head without lips and not
marked off from body; but with 6 papillae, 4 of them submedian
and 2 lateral; buccal cavity with a plate of chitinous teeth (fig.
1692). 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 420u 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 }), 7 of which are preanal and 6
postanal. Spicules (fig. 170c) equal, 1 mm. to 1.4 mm. long. Gu-
bernaculum (fig. 170d) triangular, 130, long.
Female 21 to 35 mm. long by 400 to 600n 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. 1696) very long, similar to
that of S. allodapa. Eggs (fig. 170e) 60p 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 Ascaridia
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, 250n in diameter.
Male 14 mm. long by 8702 wide. Caudal extremity strongly
curved ventrally (fig. 173), with a sharply pointed appendage 279
long. Cloacal aperture 1692 from posterior end. Preanal sucker
elliptical, about 2134 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.
Gubernaculum 144» long.
Female 14 to 22.5 mm. long.by 460 wide. Tail with slender
appendage similar to that of male, 110z long. Anus 4483p 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
:
Q
| Z%
7"
— =
Figs. 169-17
169, a, BuccaL CAVITY; b, OVEJECTOR.
AFTER SEeuRAT, 1914. 170, a, MALE TAIL, VENTRAL VIEW; D, LATERAL VIEW, ¢,
SPICULE; d, GUBERNACULUM; e, EGG. AFTER GENDRE, 1909
anterior third and posterior two-thirds of body. Ovejector long
(about 1.156 mm.) ; vestibule short, 2754; sphincter small; trompe
elongate, 899u. Two uteri. Eggs 834 long by 55u wide, elliptical,
embryonated at time of deposit.
Life history —Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution—South America (Brazil).
SUBULURA NOCTUAE (Seurat, 1914) Barreto, 1918
Synonym.—Allodapa noctuae Seurat, 1914.
Host—Carine noctua glaux.
Location.—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.
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Fics. 171-174.—SuBuLURA LUTZI.
VENTRAL VIEW.
JECTOR,
171, ANTERIOR END. 172, MALE TAIL
173, MALE TAIL, LATERAL VIEW.
AFTER BarRETO, 1918
174, VULVA AND ovE-
Male 14.5 mm. long by 250n wide. Esophagus, including bulb,
1/10 of total length. Tail short, ending in a point 85y long.
aperture 3004 from posterior end. Preanal sucker in form of a
longitudinal slit.
Cloacal
Caudal alae present.
Ten pairs of papillae (fig.
175), of which 3 are preanal, 2 adanal, 5 postanal.
The caudal
Yi
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SY
a AN
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ap—_—
So) ane
—_————s Oo
Fie. 175.—SvuBuLURA NOCTUAB.
MALE TAIL. AFTER SEURAT, 1914
glands open between the second and third pairs of papillae (num-
bering from the posterior end). Spicules equal, 700, long, filiform.
Gubernaculum triangular, elongated, prolonged anteriorly by 2 small
horns.
Female slender, 22 mm. long by 380 wide.
1/14.5 of total length.
Esophagus short,
Tail short (6302). 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 50 long by 40u wide, embryonated.
Life listory.— Unknowns: probably similar to that of Ascaridia
galli (p. 82).
Distribution —Africa (Algeria (Bou-Saada) ).
SUBULURA OLYMPIOI Barreto, 1918
Hosts—Crypturus parvirostris, Nothura maculosa, Rhynchotus
rufescens.
Location.—Intestine.
Morphology—Subulura (p. 104): Cuticle with transverse stria-
tions. Lateral alae extend to point a little posterior to the bulb.
Fics. 176-177.—SuBuLuraA OLYMPIOL 176, MALE TAIL. a, VENTRAL VIEW; b,
LATERAL VIEW. 177, OvesECTOR, AFTER Barrero, 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 411p wide. Buccal cavity 414 deep, 29p
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 465y from caudal extremity. Eleven pairs of papillae, of which
3 are preanal, 2 adanal, and 6 postanal. Spicules equal, strongly
chitinized, 899» long by 16u wide. Gubernaculum 127,» long.
Female 7.7 to 15.6 mm. long by 4624 wide. Buccal cavity 67p deep
by 83 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 long; trompe ee (3382). Two uteri, with many circum-
volutions. Eggs elliptical, 67» long by 50p 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*, in Molin, 1860; Ascaris pa-
pillosa 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 Baylis, 1919a
Host —Plotus rufus.
Location.—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 3404 wide. Esophagus 1.25 mm. long. Tail
(fig. 179) 200. lone; caudal alae absent. Preanal sucker elongated,
without chitinous border, 400% anterior to anus. Ten pairs of
papillae of which 3 are preanal, 1 adanal, and 6 postanal. Spicules
equal, 9002 long. Gubernaculum 150, long.
Female 14.2 mm. long by 4602 wide. Esophagus 1.5 mm. long.
‘Tail 800» long. Vulva in middle third of body, 5.8 mm. from an-
terior end (thus anterior to middle). Eggs 65 to 75y long by 52.5
to 554 wide. Uterus extends posterior to anus into cavity of tail.
2 Catalogue of the Vienna Museum.
NEMATODE PARASITES OF BIRDS 123
Life history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—Africa (Uganda).
SUBULURA POCULUM (Linstow, 1909) Railliet and Henry, 1914
Synonym.—Heterakis poculum Linstow, 1909.
Host.—F rancolinus adspersus.
Location. —Intestine.
Morphology.—Subulura (p. 104): Cuticula with fine cross-stria-
tions. Lateral membranes present, wide anteriorly. Mouth cavity
small; mouth with 6 papillae. Esophagus with bulb.
oO
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Wy
IK
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Zi
ax
Qo
Oo
Zor
Se
179,
Fics. 178-180.—178, SuBULURA PAPILLOSA. MALE TAIL. FROM Barreto, 1918, ArrbrR
Drascup. 179, SUBULURS PLOTINA. MaALe TAIL. AFTER Bay Lis, 1919. 180, SupuLUra
POCULUM. MAL TAIL. AFTER LINSTOW, 1909
Male 4.5 mm. long by 280u wide. Esophagus 1/5.8, 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 6 postanal. Spicules 790» long, sharply pointed.
Female 8.5 mm. long by 430p 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 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 C. 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 3594 wide. Tail strongly curved ventrally
(fig. 181); caudal alae atrophied; cloacal aperture 2114 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 25h 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, 764 long by 50% wide, embryonated when deposited.
Life history—Unknown; probably similar to that of Ascaridia
galt (p. 82).
Listribution—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 6 papillae set cross-wise. Tail pointed. Esophagus
with bulb.
Male 6.78 mm. long by 2302 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 430n 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 36y wide.
Life history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—Africa (Langenburg, Nyassa See).
NEMATODE PARASITES OF BIRDS 125
SUBULURA RIMA (Linstow, 1906) Travassos, 1913
Synonym.—H eterakis 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 400n 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
Reo
ISN
BESANT
B/S
VAIN
Fies. 181-184.—181, SuBULURA RECLINATA. MALE TAIL. AFTOR BaRRETO, 1918.
182, SUBULURA RECURVATA. MAL®b TAIL. AFTER LINSTOW, 1901. 183, SUBULURA
@IMA. MAL® 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 840y long, the left 700 long.
Female 9.3 mm. long by 430n 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.—Centropus 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, 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
49u long by 41u wide.
Life history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—Asia (Siam).
SUBULURA SEURATI Barreto, 1917
Synonyms.—Subulura 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 by 6
papillae. Three small teeth (fig. 185@) at entrance to esophagus.
Esophagus with bulb.
Male 14.5 mm. long by 500p wide. Cloacal aperture 430. from
posterior end. Preanal sucker elliptical, elongated, without chitinous
ring. Caudal alae narrow; 11 pairs of papillae (fig. 1850), 5 of
which are preanal. Spicules unequal, 1.35 mm. and 850, long re-
spectively, thus their relative lengths as 3:2. Gubernaculum triangu-
lar, 150 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 185¢c) remarkably long (over 5 mm.) ;
vestibule pyriform; sphincter and trompe very long, the latter over
4mm. The ovejector of Subulwra leprinceé is identical with that of
this species; in S. forcipata and S. subulata it is very different.
Eggs 55p long by 45p wide.
Life history —Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution.—Africa (Bou-Saada, Algeria).
NEMATODE PARASITES OF BIRDS ]
<7
ho
SUBULURA SIMILIS (Gendre, 1909) Travassos, 1913
Synonym.—Heterakis similis Gendre, 19096.
Hosts—Centropus monachus, Coracias abyssinicus, Eurystomus
afer, Scops leucotis.
Railliet and Henry suggest that the reports from Scops leucotis
(Strigiforme) and Centropus monachus (Coccigyforme) may be
confused. Barreto considers that Gendre was dealing with several
species from this wide variety of hosts and suggests that the material
from Lurystomus 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
106 Ld
Te TEs} pS
curr eos
Cocets Ce
os
v/o01
Fic. 185.—SUBULURA SEURATI. @, ANTERIOR END; Db, MALE TAIL; C, OVEJECTOR. AFTER
SeuraT, 1914
is much more slender in S. 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 600u, whereas that of
S. similis as given by Gendre is 300 to 400» in the male and 420 to
500z 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 wide.
Baers es. male 13.1 mm. long by 400y wide.
female 18.8 mm. long by 500 wide.
S. leucotis ____- male 12.6 to 15 mm. by 300 to 360x.
female 20.1 to 22.5 mm. by 420 to 500..
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.—Ascaris strongylina Rudolphi, 1819; Strongylus spicu-
latus Cobbold, 1861; Heterakis spiculatus (Cobbold, 1861) Travassos,
1923.
Hosts.—Bucco capensis, B. melanoleucos, B. rufiventris, B. striola-
tus, B. swainsoni, B. tamatina, Caprimulgus ruficollis, C. nacandua,
C. urutas, Chelidoptera tenebrosa, Crypturus noctivagus, C’. species,
C. tatuapa, Cuculus melanorhynchus, C. tinguacu, Gallus gallus,
Malacoptila torquata, Microdactylus cristatus, Monasa leucops, M.
tranguilla, Nonnula rubecula, Odontophorus capueira, Perdix den-
tata, Podager nacunda, Tetrao uru, Tinamus, species, T. tataupa.
Location.—Intestine.
Morphology.—Subulura (p. 104): Cuticle finely striated trans-
versely. Lateral alae well developed, 1.37 mm. long by 65» 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 481 deep; 3 rectangular teeth, 19. high, at entrance to
esophagus. Esophagus 1.04 mm. long. Bulb spherical, 2054 in
diameter.
Male 4.36 to 12 mm. long by 3084 wide. Tail (fig. 187) conical,
with straight chitinous appendage 102y long. Anus 186 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 411p 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 9502 long; vestibule sinuous, sphincter small; trompe
equivalent to 24 the length of ovejector. Two divergent uteri.
Eggs 84y 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 nun Ee 1819; Heterakis subulata
Schneider, 1866.
NEMATODE PARASITES OF BIRDS 129
Hosts —Caprimulqus aegyptius saharae and C. ruficollis. Other
reports, at least part of which are probably confused with other
species, are: Antrostomus vociferus, Caprimulgus candicans, C.
cortopan, C. diurnis, C. europaeus, C. guianensis, C. mercurius, C.
nattereri, C. scaphiuris, C. semitorquatus, C. trifurcus, Chordeiles
semitorquatus, Cuculus cayanus, C. melacoryphus, C. naevius, C.
tinguazu, Nyctibius aethereus, 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 mm. long;
according to Seurat, 7.5 mm. long. Tail slender, without caudal
Fics. 186-190.—186, SUBULURA SIMILIS. MALW TAIL. AFTER GENDRE, 1909.
187, SUBULURA STRONGYLINA. MALE TAIL, AFTER TRAVASSOS, 1913. 188,
SUBULURA SUCTORIA. MALE TAIL. 189, HEAD, FRONT VIEW. Fics. 188-189
FROM BARRETO, 1918 AFTER DRASCHE, 1882. 190, OveJECTOR. AFTER
BARRETO, 1918
alae. Cloacal aperture 220. 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» long.
Female, according to the original description, 22 to 22.5 mm. long;
according to Seurat, 12 mm. long by 310p 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 85y long by 56 wide.
130 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history —Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution—Europe (Austria (Vienna Museum), Spain, and
Corsica. )
SUBULURA SUCTORIA (Molin, 1860) Railliet and Henry, 1912
Synonyms.—H eterakis suctoria Molin, 1860; Allodapa suctoria,
(Molin. 1860) Seurat, 1914.
Hosts—Athene noctua, Caprimulgus campestris, C. europaeus, C.
nigrescens, C. rufus, C. species, C. vociferus, Coturnix delagorgneéi,
Dicholophus margrafi, Francolinus bicalearatus, Gallus gallus, Helio-
treptus anomalus, Hydropsalis climacocercus, Lurocalis semitor-
quatus, Meleagris gallopavo, Microdactylus cristatus, Numida melea-
gris, N. papillosa transvaalensis, Nyctibius aethereus, N. grandis, N.
jamaicensis, Nyctidromus albicollis, “ Otis houbara” (Houbara mac-
queent or H. undulata), Podager nacunda, Pternistes swainsona,
Sephina francolinus, Stenopsis candicans.
Location.—Ceca.
Morphology.—Subulura (p. 104): Filiform worms, the anterior
portion curved with the curvature toward the dorsal face. Lateral
cephalic alae 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 25p
long. Esophagus dilated slightly in the posterior portion and ending
in a piriform bulb.
Male 11.8 to 13.8 mm. long by 3594 wide. Posterior extremity in-
fundibular, straight, and terminating in a short appendix. Cloacal
aperture 211» from posterior end. Sucker ellipsoidal, without chi-
tinous wall, 135y long; it is 296 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 1504 long; Seurat says triangular and
175y long.
Female 20 to 23 mm. long by 5392 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
body. 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 45y wide.
For the descriptions given for this worm by Gendre, by Seurat
and by Gedoelst see Sublura brumpti (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 TRAVASSOSI Barreto, 1918
Synonyms.—Ascaris forcipata Rudolphi, part; Heterakis forci-
paria Schneider, 1866, part; Subulura strongylina Railhet 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. mgra,
Monasa 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 360u wide. Esophagus 1.03 mm.
long, bulb 231p in diameter. Cloacal aperture 127, from posterior
extremity. Preanal sucker elliptical, 3854 long, 5084 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, long.
Female 6 to 10 mm. long by 400 to 410u 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,) ; vesti-
bule 406y, sphincter 169y, 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 2231; width 554. Mouth with 6 small equal papillae. Buccal
cavity (fig. 195) 45 long by 384 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 2324 from posterior end. Preanal sucker 186y long, without
chitinous ring, 4234 from caudal extremity. Caudal alae rudimen-
tary. Eleven pairs of papillae, of which 8 are preanal, 2 adanal, 6
postanal. Spicules equal, 558. long by 38 wide. Gubernaculum
228 long.
/94F.
Figs. 191—-194.—SUBULURA TRAVASSOSI. 191, OVEJECTOR. 192-193, Map TAIL. (FIGS.
191, 192, anD 193 FROM Bucco cHacurRU.) 194, FrRoM BuccCO SWAINSONI. MALE TAIL.
ALL AFTER BARRETO, 1918
Female 15.5 mm. long by 8584 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 history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution—South America (Brazil).
SUBULURA, species Baylis and Daubney, 1922
Host.—Twurnix, 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 4102 wide. Head 80, in diameter; buccal
cavity 35u deep by 20n wide; esophagus 1.1 mm. long; bulb 150 in
diameter. Tail acutely pointed; anus 320, from posterior end.
Vulva 6.1 mm. from anterior end. Eggs 85» long by 56y wide, em-
bryonated.
I
il
MI i\||
196,
Fics. 195—-196.—StUBULURA TROGONI. 195, ANTERIOR END, 196, MALE TAIL. AFTER
BArRRETO, 1918
Life history.—unknown: probably similar to that of Ascaridia
gali (p. 82).
Distribution.—Asia (India).
SUBULURA (7) ACUTICAUDA (Linstow, 1901) Railliet and Henry, 1914
Synonym.—O cysoma 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 390p wide. Esophagus 1/10.7, tail 1/38 of
votal 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 510n 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 47p long by 34 wide.
Life history—Unknown; probably similar to that of Ascaridia
galli (p. 82).
Distribution —AMfrica (Usanga, Rukwa-See).
Fics. 197-199.—SusuLura acuTicAuDA. 197, MaLe TatL. AFTER Linstow, 1901.
198, SUBULURA GRACILIS. Heap. 199, MaLe Tain. AFTER LINSTOW, 1899
SUBULURA (?) GRACILIS (Linstow, 1899) Railliet and Henry, 1914
Synonym.—Ouxysoma gracile Linstow, 1899a.
Host.—F rancolinus, 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 310n wide. Esophagus 1/8, tail 1/40 of
total length. Spicules 5904 long, sword-shaped. Nine pairs. of
caudal papillae (fig. 199) of which 3 are preanal, 6 postanal.
Female 9.48 mm. long by 390» 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, long by 44» wide.
NEMATODE PARASITES OF BIRDS 135
Life history.—Unknown: probably similar to that of Ascaridia
galli (p. 82).
Distribution—Europe (Germany (Berlin Zoological Garden) ).
Family ASCARIDAE Baird, 1853
Family diagnosis —Ascaroidea (p. 48): Polymyarian. Mouth with
3 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). J/ale 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.
Ty pe-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 GENERA OF ANISAKINAB
Intestinal cecum and esophageal appendix both present___ Contracaecum, p. 146.
Intestinal cecum present; esophageal appendix absent____ Porrocaecum, p. 135-
Genus PORROCAECUM Railliet and Henry, 1912
Synonym.—Terranova Leiper and Atkinson, 1914.
Generic diagnosis —Anasakinae (p. 185) : 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
Ty pe-species.—P orrocaecum crassum (Deslongchamps, 1824) Rail-
het and Henry, 1912.
KEY TO SPECIES OF PORROCAECUM
1. Larva only known, in eel, Anguilla vulgaris__ Porrocaecum anguililae, p. 146.
Adults; knowns in pind S226 see = es ee ee ee eee 2.
2. Lateral) membranes; present: ~22- 22) see Bec ea el ea 3.
Lateral membranes absent or undescribed.——.+.._.----~---+~---_-.--- 5.
3..Cecum long (about 8 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 91u long by 854 wide___-__-_________ Forrocaecum semiteres, p. 141.
Lips with 2 large oval papillae; female tail 1/46 of total length; eggs 110u
long by Sdw wide==_ 22 == Seas roe Porrocaecum ensicaudatum, p- 139.
5. Species unrecognizable; possibly identical with P. ensicaudatum.
Porrocaecum heteroura, p. 144.
Species recogmiza le ees ee el ee 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. #21 ts te 7.
i Vulya-posterior to: middle of body. 2-222. 2 tt, ee es ee 8.
Mulvacanterior to middle Of body. 52 22 2 2a ae ee ee ee 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 digitiform projections of pulp; male up to 48 mm.
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
CUPTNSL2 OCT IG TATGES Aid hy bts Porrocaecum praelongum, p. 140.
Female not over 112 mm. long; male without caudal alae; from other hosts
ED SUT D7 2p ND Vee a a a TS 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 moschata, Numida mele-
agris.
Location.—Small intestine.
NEMATODE PARASITES OF BIRDS 137
Morphology.—Porrocaecum (p. 185): Mouth with 3 convex lips.
Cuticle with marked transverse striations. Muscular esophagus
claviform, followed by a ventriculus 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, 520. 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 100,» 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 haliaetus, Helotarsus albicilla, H.
ecaudatus, 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, +.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, 9502 long, not alate.
Female 40 to 90 mm. long by 500 to 1.5 mm. wide. Tail 700
long, blunt; caudal papillae 2002 from end. Vulva in anterior half
of body, dividing body length in ratio of 3:5. Eggs 85 to 93y
long by 58 to 74 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, UNIFED 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. virginianus, Cir-
caetus gallicus, Circestus pectoralis, Circus aeruginosus, C. cinera-
ceus, C. cyaneus, C. rufus, Falco aesalon, F. albicilla, F. apivorus, F.
ater, F. brachydactylus, F. brasiliensis, F. buteo, I’. chrysaetos, F.
cyaneus, F. degener, F. gallicus, F. imperialis, F. lithofalco, F.
lanarius, F. lagopus, F. milvus, F. naevius, F. nisus, F. palumbarius,
F. pennatus, F. peregrinus, PF. rufus, F. rutilans, F tinnunculus,
Gypaetus barbatus, Gyps fulvus, Haliaetus albicilla, Milvus ater,
Milvous milous, M. regalis, Nisaetus fasciatus, Strix aluco, S. brachy-
otus, S. bubo, S. dasypus, S. flammea, 8. nyctea, S. otus, S. stridula,
S. tangmalmi, Vultur cinereus, V. fulvus, V. monachus. Larvae in
Sorex tetragonurus and Talpa europaea.
Location.—Intestine.
ae
O/177 177
Figs. 200—202.—200, PoRROCAECUM ANGUSTICOLLE. DorsaL Lip. 201, MALE TAIL.
202, PORROCAECUM DEPRESSUM. DORSAL LIP, AFTER BAYLIS AND DAUBNEY, 1922
Morphology.—Porrocaecum (p. 185): 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 encapsuled
in the peritoneum of the shrew and mole (see above, under //osts).
Distribution —Europe, Asia (Russian Turkestan and India),
South America (Brazil and British Guiana) and Africa (Trans-
vaal).
PORROCAECUM ENSICAUDATUM (Zeder, 1800) Baylis, 1920b
Synonyms.—F usaria ensicaudata Zeder, 1800; Ascaris ensicaudata
(Zeder, 1800) Rudolphi, 1809.
Hosts.—Acrocephalus arundinaceus, Alauda species, Anas boschas,
Charadrius dubius, C. hiaticula, C. morinellus, C. pluvialis, Gallinula
chloropus, Himantopus melanopterus, Luscinia philomela, Merula
nigra, Mimus polyglottus, Motacilla alba, Oedicnemus crepitans,
“rooks,” Pica caudata, Pluvialis apricarius, Salicaria turdoides,
Squatarola helvetica, Sturnus vulgaris, Sylvia turdoides, Turdus
iliacus, T. musicus, T. merula, T. pilaris, T. sawatilis, T. torquatus,
T. viscivorus, Vanellus cristatus, V. melanogaster.
Location.—Intestine.
Fic. 208.—PorRROCAECUM DBPRESSUM. MALE TAIL. AFTER LINSTOW, 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, 1204
wide. Intestinal cecum small, almost rudimentary.
Male 28 to 32 mm. long by 1.02 mm. wide. Esophagus 1/9.9, tail
1/86 of total body length. Body narrows suddenly posterior to
cloacal aperture. Six pairs of postanal papillae (fig. 2040), 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. Wulva somewhat anterior to middle of
body, dividing body length in ratio of 4:5. Eggs (fig. 204c) 110y
long by 85 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—Colymbus auritus.
Location.—Intestine.
Morphology.—Porrocaecum (p. 135): Body white, filiform, very
elongate. Head proportionately large, 400% wide. Esophagus 4.3
mm. long by 4004 wide, followed by a ventriculus 400n long and
accompanied by a thick cecum 3 mm. long by 600, wide, joining
with the intestine. Cuticle with very conspicuous transverse stria-
tions.
Fic. 204.—PORROCAECUM BNSICAUDATUM. a, DORSAL LIP; b, MALE TAIL; ¢, EGG.
AFTER LINSTOW, 1884
Male 90 mm. long by 9004 wide. Tail suddenly narrowed and end-
ing in a short conical point; cloacal aperture, with a large tubercule,
3304 from posterior end. Membranous caudal alae present, anterior
to cloacal aperture, supported by 5 pairs of papillae. Spicules equal,
9004 long by 584 wide.
Female 154 mm. long by 1.8 mm. wide. Tail straight, conical.
pointed; anus 6002 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.
Life history—Unknown.
NEMATODE PARASITES OF BIRDS 141
Distribution —KEurope (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 cinerea, A. cocot, A. manillensis, Nycticorax 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 280u 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 1.6 mm. wide. Eggs 110u long by 91p
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 cornix, Pluvialis apricarius, Tringa vanellus,
Vanellus cristatus, V. melanogaster.
Location.—Intestine.
142 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Morphology.—Porrocaecum (p. 135): Body white. Head (fig.
207) 230 to 8004 wide, with 8 large convex lips each carrying a pa-
pilla in the middle of the convexity. Hsophagus 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
ee 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 65 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
206.
Figs. 205—-208.—205, PorrocaECUM RETICULATUM. Maur 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, 1928
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, 914 long by 84 wide, with thick reticulated shell.
Life 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) Baylis, 1920b
Synonyms.—A scaris serpentulus Rudolphi, 1809; dscaris ardeae
Froelich, 1802.
Hosts.—Anthropoides virgo, Ardea agami, A. cinerea, A. cerulea,
A. grus, A. major, A. melanocephala, A. nycticorax, A. pileata, A.
purpurea, A. scapularis, A. violacea, Grus australasiana, 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 210u, width at base 78p, at level
of the 2 papillae 180u. Interlabia large, ear-shaped, 2/3 the size of
the large lips. Esophagus 3.4 mm., ventriculus 400», 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 by Monnig measured 55 mmm. long by 1 mm. wide.
(
/ 209
EuGs. 209-210.—PorROCAECUM SPRPENTULUS. 209, Heap. 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” 2604 long in his specimen), with wide alae.
Female (see above for size). Eggs 1044 long by 78 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.—Aegolius otus, A. brachyotus, Bubo maximus, B. virgui-
anus, Falco pygargus, Flammea flammea, Nyctale ten gmalma, Otus
Faachuptis, O. vulgaris, Strix aluco, S. bubo, S. flammea, S. nivea, S.
noctua, S. stridula, Syrnia nyctea, S. aluco, U lula aluco.
Location.—Intestine.
Morphology.—Porrocaecum (p. 185): Large lips (fig. 211) six-
sided, 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. 218) 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,
inost anterior, 2 pairs and the cloacal aperture. Spicules sickle-
shaped.
Fics. 211-213.—PorROCAECUM SPIRALP. 211, DORSAL LIP. 212, Eee. 213,
MALE TAIL. AFTER LINSTOW, 1875
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, 1024 long by 604 wide, with double shell, the
outer thickened at the ends and with small glittering prominences.
Life history—Unknown.
Distribution.—Kurope.
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; Ascaris heteroura
Mehlis, 1831.
Hosts.—Charadrius morinellus, C. oedicnemus, OC. pluvialis,
Himantopus melanopterus, Squatarola helvetica, Sturnus vulgaris,
Turdus musicus.
Location —Intestine.
Morphology.—Porrocaecum (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.
Jnstribution—Europe (Ireland, Germany, and Italy (Portofer-
rajo)).
This species is given as a synonym of P. ensicaudatum by numerous
authors but P. ensicaudatum 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.
PORROCAECUM KIRGHISENSIS (Skrjabin, 1916) Cram. 1927
Synonym.—Ascaris kirghisensis Skrjabin, 1916; Ascaris hirghi-
sensis Skrjabin, 1916 (evidently misspelled).
Host.—Aquila imperialis.
Location.—Small intestine.
Morphology. — Porrocaecum Cy
(p. 185): Cuticle transversely nay
striated. Head with lips and LPF,
interlabia (fig. 215c). Lips
(fig. 2156) made up of 3 parts,
the central one with denticulate “| f
edge and with the pulp divided
into 2 equally large lobes. Two ¢ i
small papillae situated far an- TV we
terior on lips. = eA
Male 75 to 99 mm. long by 1.3 \
to2 mm. wide. Tail (fig. 215 d Lie
and e) conical, pointed; cloacal ics. 214-215.—214, Porrocancum Hers-
‘ : ip i F s ROURA. ANTERIOR END (DISSECTED). AFTER
aperture 5900p from end. Spic- CREPLIN, 1829. 215a, PORROCAECUM AN-
ules equal, 1.18 mm. long. Five GUILLAE, ANTERIOR BEND (DISSECTED).
abe . eel. AFTER LINSTOW, 1899. 215 6b TO €, Por-
pairs of postanal papillae, the ROCAECUM KIRGHISENSIS. 0, LIP; ¢, IN-
most anterior pair of these com- TERLABIUM; @, LATERAL AND @€, VENTRAL
: LEW > MALE. TAIL. 'TER SKRJABIN,
posed of double papillae. Four- eT PF AR A AYZIE SERA
teen pairs of preanal papillae.
Female (immature) 80 mm. long by 1.8 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 species to Porrocaecum is made tentatively
by the present writer on the basis of Skrjabin’s statement that it is
very close to P. depressum. The main differential characters of the
two genera Porrocaecum and Contracaecum, that is, the nature of
the esophageal appendages, not being known in connection with this
species, the allocation can not be made with certainty but it is.
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 970p.
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.
Ty pe-species.—C ontracaecum spiculigerum (Rudolphi, 1809)
Railliet and Henry, 1912.
KEY TO SPECIES OF CONTRACAECUM
TANT Ae alone“ GesGribe Gasset a
Both male and female, or female alone, deseribed______________________ 3.
2. Male 13 mm. long; esophageal appendix 700u long; spicules 1.8 mm. Jong.
Contracaecum engonium, p. 150.
Male 16.5 to 32.5 mm. long; esophageal appendix 9604 to 1.2 mm. long;
Spiculés*4: ‘mim: slong’ scviseawey ay Contracaecum punctatum, p. 157.
3. Vulva posterior to middle of body dividing body length in ratio of 2: 1.
Contracaecum multipapillatum, p. 154.
Vulva, anterionste middle. of podye 5 jt yo 4.
4. Female alone known; eggs 80 to 904 long by 60u wide; from Haliaétus
SDOCIOG oe ene Meh ree eee Contracaecum haliaéti, p. 150-
Both male and female known; eggs different from above; from other hosts
than above oo sant oS elt ew ee Baie eee 5.
5. Pulp of lips with 2 anteriorly directed bipartite processes; interlabia
strongly curved, the anterior ends split into 2 rounded processes ; male
with 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.
6. Esophageal appendix not, moeke, than-1 mm. lenge... jen cfin deen 7
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 26u long by 21p
TG) Ott ee ee os * 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 55y long
by tomy widewts oo Aw cae eg Se ee 8.
8. Esophageal appendix about 500u, cecum 1.8 mm. long; spicules unequal,
2.7 and 3 mm. long; eggs 100u long by 60x wide.
Contracaecum scotti, p. 159.
Esophageal appendix 720u or longer, cecum 2.6 mm. or longer; spicules
equal and either shorter or longer than those above; eggs not over 694
henge byc45., Wider ste sw lan SN teow ema AA ee ee Nee eke 9.
9. Tail of female 130u long; vulva divides body length in ratio of 9.5:10.5; tail
of male 85u long; spicules 1 mm. long___ Contracaecum andersoni, p. 149-
Tail of female 300u or longer; vulva more anterior than in C. andersoni;
tail of male 175y or longer; spicules 3.6 mm. or longer___________-_- 10.
10. Dorsal lip with 2 single papillae; esophageal appendix 720% 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 7380u 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 gubernaculum present, 120u long; spicules 1.9 mm. long; eggs 40u long
DY See wie 28 =e NS Contracaecum praestriatum, p. 156.
Gubernaculum absent or undescribed; spicules jonger than above; eggs
larger than above (except possibly in C. tricuspe where not de-
ETC ee aE EN a eee a eR EN ee re es Es 12.
12. Male 13.8 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 abeve_______________-______ 13.
13. Esophageal appendix 3 to 4.5 mm. long; interlabia low and rounded; anus
of female 5644 from caudal extremity_____-_ Contracaecum ovale, p. 150d.
Esophageal appendix not over 1.9 mm. long; interlabia different from
above; anus of female not over 500u 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.
VON SS Soe 2 5 wet 22ers ee Contracaecum microcephalum, p. 152.
In connection with species of Contracaecum, Ascaris kirghisensis
(placed by the present writer in Porrocaecum) should be kept in
mind as possibly belonging in this genus (see discussion, p. 145).
oD ?
CONTRACAECUM SPICULIGERUM (Rudolphi, 1809) Raillet and Henry, 1912
Synonym.—A searis spiculigerum Rudolphi, 1809.
Hosts—Alea torda, Anas clangula, Carbo brasiliensis, C.
cormoranus, C. cristatus, C. dilophus, C. graculus, C. pygmaeus,
148 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Chroocephalus ridibundus, Colymbus articus, C. atrigularis, C.
nigricans, C. rufogularis, C. septentrionalis, Fulica leucopt.,
Haematopus ostralegus, Halieus brasiliensis, Larus argentaius, L.
canus, L. fuscus, L. marinus, L. ridibundus, L. tridactylus, Lestris
parasitica, L. pomarinus, Merganser castor, Mergus merganser,
Microcarbo pygmaeus, Pelecanus americanus, P. conspicillatus, 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. lavaillanti, P. melanogaster, P. novae-hollan-
diae, Podiceps auritus, P. eristatus, P. dominicensis, P. minor, P.
nigricollis, Tachypetes aquila, Uria grylle, U. troile, Utamania torda.
Location.—Intestine and, in Pe/ecanus, the gular pouch.
Fes. 216-217.—ConTrRACAECUM SPICULIGERUM. 216, Heap, ArtTER LINSTOW,
1909. 217, a, FSSOPHAGUS AND APPENDICES: 0, MALE TAIL. AFTER
SCHNEIDER, 1866
Morphology.—C ontracaecum (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.
LLinstow 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. 2177).
Male 32 to 45 mm. long by 800 to 900u wide. Tail curled, end-
ing in conical point. Cloacal aperture 250» from posterior end
in small specimens. Spicules, according to Dujardin, 2 mm. long
by 38302 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 pa‘rs of postanal papillae (fig. 2170),
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 4002 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 52y
in diameter, according to Dujardin, 72 in outer diameter and
42u as diameter of the yolk, according to Linstow. The latter author
describes the shell as 4.94 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) ).
QQ
» °
° :
° ° “
SE WS
2 S
Oil we
“7 8 8
> of .
aN
e%
OS7 of
Fic. 218.—CONTRACAECUM ANDERSONI. @, DORSAL LIP; b, INTERLABIUM ; €, VULVA;
d, MALE TAIL. AFTER VEVERS, 1923
CONTRACAECUM ANDERSONI Vevers, 1923
Host.—F lorida caerulea.
Location.—Small intestine.
Morphology.—tContracaecum (p. 146): Cuticle transversely stri-
ated. Head 4002 wide by 160» long. Three fleshy lips and 3
large interlabia; the former (fig. 218@) each bear 2 papillae; the in-
terlabia (fig. 218)) are triangular, 1502 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% wide. Cloacal aperture 85
from posterior end. Five pairs of postanal papillae (fig. 2187), the
most anterior pair being double; 5 pairs of adanal, and 20 to 25
pairs of preanal papillae. Spicules equal, 1 mm. long by 10y wide.
Female 19 to 22 mm. long by 750» wide. Anus 130, from
caudal end which is conical and sharply tapering. Vulva (fig. 218¢)
just anterior to middle of body, 9.5 mm. from head end in a 20
mm. specimen. Vagina short (1504), muscular, with 3 pairs of
150 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
leaf-like valves. Eggs oval, 55 long by 45h wide, with thick shells
of mosaic appearance.
Life history—Unknown.
Distribution—South America (British Guiana (Georgetown) ).
CONTRACAECUM ENGONIUM Baylis and Daubney, 1922
Host—Ciconia nigra.
Location —Not given.
Morphology—Contracaecum (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» wide. Muscular esophagus 2.75 mm.
long, ventriculus 140, long; posteriorly directed appendix 700 long
4
OLMMT7177
O17 (77
Fic. 219.—CONTRACAECUM ENGONIUM. a, HEAD; 0b, MALE TalIL, AFTER Baylis
AND DAUBNEY, 1922
by 150. wide. Intestinal cecum broad, reaching to within 690, of
head end. Cloacal aperture 1254 from caudal extremity. Ten
pairs of postanal papillae (fig. 2196), 4 of them being latero-ventral
and the other 6 lateral and pedunculated. No preana! papillae de-
scribed. Spicules equal, 1.8 mm. long by 22” wide.
Female unknown.
Life history —Unknown.
Distribution.—Asia (India).
CONTRACAECUM HALIAETI Baylis and Daubney, 1923
Synonym.—Ascaris aquillae Smith, Fox, and White, 1908, not
Ascaris aquilae Gmelin, 1790.
Hosts.—Haliaétus leucocephalus and H. leucogaster. .
Location.—Intestine.
Morphology—Contracaecum (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 150n
wide at base, 140u 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.
Cc.
/0u 20 30 40 50 $0 70 30 $0 10s
Fic. 220.—CoNYVRACAECUM HALIAETI. @, LATERAL VIEW; Db, FRONT VIEW OF HEAD:
c, EGG. AFTER SMITH, Fox, AND WHITS, 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, 250» wide at
base of lips. Tail acutely conical, anus subterminal. Vulva an-
p40 30 0H a
Fic. 221.—ConTRACAECUM HALIABTI. @, INNER SURFACE; Bb, OUTER SURFACE OF
Lip. AFTER SMITH, FOX, AND WHITE, 1908
terior to middle of body, small and inconspicuous. Vagina long,
slender. Eggs (fig. 220c) of varying shape, from 80 to 90p 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).
3612—27T 12
152 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
CONTRACAECUM MICROCEPHALUM (Rudolphi, 1869) Baylis, 1920b
Synonyms.—Ascaris microcephala Rudolphi, 1809; Aathleena are-
uata Gedoelst, 1916; Contracaecum quadricuspe Walton, 1923.
Baylis and Daubney have examined the type specimens of Kath-
leena arcuata and find that species identical with Rudolphi’s. The
present writer considers C. qguadricuspe identical also, the descrip-
tion given by Walton agreeing closely with that of Gedoelst in all
particulars except that the esophageal appendix of C. guadricuspe
is somewhat shorter and the vulva slightly more anterior.
Hosts—Anas boschas, Anas domestica, Ardea cinerea, A. comata,
A. herodias, A. minor, A. nycticorax, A. purpurea, A. species, A.
stellaris, Ardeolo grayi, A. ralloides, Botauras mugitans, B. stel-
laris, Butorides virescens virescens, Ciconia alba, C. nigra, Herodias
Fic. 222.—CoNTRACABCUM MICROCEPHALUM. @, FRONT VIEW OF HEAD; b, DORSAL
VIEW OF HHAD; C, MALE TAIL; d, FREE END OF SPICULBD; €, CROSS SECTION OF SPICULE.
AFTER GEDOELST, 1916
egretta, H. tricolor, Nyctiardea grisea, Nycticorax europaeus, N.
nycticorax.
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. 2226) 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, wide, with
an appendix 1.1 to 1.25 mm. long by 145. wide; cecum extends
anteriorly along esophagus for 3/4 the latter’s length. According to
NEMATODE PARASITES OF BIRDS 153
Walton (C. qguadricuspe), muscular esophagus 3.2 to 3.6 mm. long,
ventriculus 175y long, esophageal appendix 800 to 8164 long by 80,
wide, intestinal cecum 2.8 to 2.9 mm. long.
Male 18 to 26 mm. long by 900z to 1 mm. wide (4. arcuata) ; 15 to
45 mm. long (C. microcephalum). Tail (fig. 222c) curved ventrally ;
no caudal alae. Cloacal aperture 230 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 paurs of large papillae; anterior to the cloacal
aperture at least 20 pairs in more or less regular rows (. arcuata),
or 381 pairs (C. microcephalum). Spicules equal, 2.3 to 2.8 mm. long,
with curved alae.
Female 23 to 387 mm. long by 1.1 mm. or less wide (A. arcuata) ;
45 to 70 mm. long (C. microcephalum). Tail conical; anus about
440 to 500u from posterior end; 2 lateral papillae, 2404 from tail
end. Vulva anterior to middle of body, dividing body length in
ratio of 2:3 (in Walton’s specimen, which was 37.9 mm. long, vulva
9.6 mm. from the head). Eggs of variable shape, globular or ellip-
tical, averaging 68» long by 58.5 wide (the range being 72. by 48 to
64u down to 644 by 56 to 64) ; shell thin, with irregular surface.
Life history —Unknown.
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 micropapillata Stossich, 1890,
Host.—Pelecanus, species, P. crispus, P. eruthvorhynchos, and
Glaucionetta clangula.
Location.—Intestine.
Morphology.—Contracaecum (p. 146). The present writer has
recently identified this parasite from P. erythrorhynchos and G.
clangula, 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 26 mm. long. Three pairs of double postanal papillae
(fig. 224a). Preanal papillae numerous, in 2 regular rows.
Female 35 to 40 mm. long.
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 theesophagus. Ma/e
15 to 20 mm. long by 950» wide. In a specimen 18 mm. long, esopha-
ous 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. /Hemale 20 to 30 mm. long
by 1.2 mm. wide. In a specimen 23 mm. long, esophagus 3.1 mm.,
esophageal appendix 500y, intestinal cecum 2 mm. long. Vulva 1/4
of body length from anterior end. Eggs 58 by 54p.
Fig, 223-224.—ConTRACABCUM MICROPAPILLATUM. 223, ANTERIOR END (DIS-
SECTED). ORIGINAL. 224, a, MALE TAIL; b, LIP AND INTERLABIA. AFTER
Srossicu, 1890
Life history.—Unknown.
Distribution—Kurope (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. erythrorhyn-
chos.
CONTRACAECUM MULTIPAPILLATUM (Drasche, 1882) Baylis, 1920b
Synonyms.—A scaris multipapillata Drasche, 1882 (1888a) Con-
tracaecum multipapillosa (Drasche, 1882) Skrjabin, 1916 (evidently
a misspelling).
fTosts.—Ardea, species and Tantalus loculator.
Location.—Esophagis and proventriculus.
NEMATODE PARASITES OF BIRDS 155
Morphology.—C ontracaecum (p. 146): Lips (fig. 226a) with ear-
hike projections and with 2 papillae; dentigerous ridges absent. Ac-
cording to Drasche, interlabia 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 600 to 900% wide. Esophagus 3.23 mm.
long, its appendix small, dactyliform (fig. 225). Intestinal cecum
7/8 the length of esophagus. Tail (fig. 2266) 150y 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 500n to 1 mm. wide. Posteriorly
directed esophageal appendix 1/3 the length of the anteriorly directed
Ne tt
QS A'"O"(o o
S©coeg
ec
So
FIGs, 225—226.—CoONTRACAECUM MULTIPAPILLATUM. 225, DiGhsSTIVE ORGANS OF MALE.
AFTER SKRJABIN, 1916. 226, a, HEAD; b, MALE TAIL. AYFTER DRASCHE, 1882
intestinal cecum, the latter being 3/4 the length of the esophagus.
Tail pointed, anus 370n from end. Vulva in posterior part of body,
dividing body length in ratio of 2:1. Eggs oval, 681 long by 60,
wide.
Life history —Unknown.
Listribution.—South America (Brazil and Paraguay).
CONTRACAECUM OVALE (Linstow, 1907) Baylis, 1920b
Synonym.—Ascaris ovalis Linstow, 1907.
Host.—Podiceps cristatus.
Location.—Proventriculus and intestine.
Morphology.—C ontracaecum (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 140, 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 880p 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 68y long by 57» wide.
Life history —Unknown.
Distribution—Kurope (Germany (Berlin Museum) ).
CONTRACAECUM PRAESTRIATUM Monnig, 1923
Host.—Podiceps capensis.
Location.—Not given.
Morphology —Contracaecum (p. 146): Color white. Cuticnlar
striations, 88» wide, very marked in anterior 1/3 of body, then sud-
Zar
Pies. 227—229.—227, CONTRACAECUM OVALE. HrAD. AFTER LINSTOW, 1907. 228,
CONTRACABCUM PRAESTRIATUM. ANTERIOR END. 229, MALE TAIL. AFTER MONNIG,
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 650n 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 £57
Female 33 mm. long by 800% wide. Tail 3604 long, sharply
pointed. Vulva in anterior third of body, dividing body length in
ratio of 5:12. Eggs thin-shelled, very round, 40u long by 32» wide.
Life history —Unknown.
Distribution.—Africa (Transvaal).
CONTRACAECUM PUNCTATUM (Gedoelst, 1916) Baylis, 1920b
Synonym.—Kathleena punctata Gedoelst, 1916.
Host.—* Grand Heron a bec de Pelican,” probably Pseudo-
tantalus rbis.
Location.—Proventriculus.
Morphology.—Contracaecum (p. 146): Color white. Cuticle with
cross-striations. Head small, 148 in dorso-ventral diameter. Three
lips, the dorsal with 2 papillae, the ventro-laterals each with a single
papilla. Three interlabia present. Precervical papillae 470» from
head end in small specimens. Esophagus 2.24 to 4.5 mm. long, with
ie! nas
2350 ZO.
Fics. 230—231.—230, CONTRACARCUM PUNCTATUM. MALE TAIL. AFTHR 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 665p 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 42.5y
wide, with curved alae.
Female—Unknown.
Life history —Unknown.
Distribution —Africa (Belgian Congo (Bukama, Katanga) ).
CONTRACAECUM RODHAINI (Gedoelst, 1916) Baylis, 1920b
Synonym.—K athleena rodhaini Gedoelst, 1916.
Host.—Plotus rufus.
Location.—Proventriculus.
Morphology. — Contracaecum (p. 146): Color yellow - white.
Cuticle with fine cross-striations. Head (fig. 231) 300~% wide by
158 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
150 to 160p 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» from anterior end. Esophagus 3.3 to 4 mm. long, with post-
eriorly directed appendix 720 to 1.04 mm. long; anteriorly directed
cecum 2.6 to 2.8 mm. long.
Male 18.5 to 20 mm. long by 830,» 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.
BOS:
01/7/77
Pies. 232—233.—CoNTRACAECUM ROSARIUM. 232, ANTERIOR END. AFTER CON-
NAL, 1912. 233, MALE TAIL. AFTER BAYLIS AND DAUBNEY, 1922
Female 22 to 25 mm. long by 940 to 1.2 mm. wide. Tail conical,
with a papilla on each side, 120% from posterior end. Anus 336p
from end. Vulva not salient, at the anterior 3/10 of body length.
Eggs oval or globular, 694 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 Nycticoraaz, 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 cuticular collar around base of lips. Esophagus
3.9 mm. long (in the male), its appendix 700u 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 600n 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
smalier, 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.
7.
Fig. 2384.—CONTRACAECUM SCOTTI. @&, ESOPHAGUS AND APPENDICES; 0b, HEAD; ¢,
LATERAL VIEW AND @, VENTRAL VIEW OF MALE TAIL. AFTER LEIPER AND ATKIN-
SON, 1915
Female 30.8 mm. long by 700% wide. Vulva 1.59 mm. from head
end. Vagina about 170» long, dividing into 2 parallel uteri. Eggs
oval, 26 long by 21 wide, with thick shells.
Life history —Unknown.
Distribution —Africa (Lagos) and Asia (India).
CONTRACAECUM SCOTTI (Leiper and Atkinson, 1914) Baylis, 1920
Synonym.—K athleena scotti Leiper and Atkinson, 1914.
Host.—lDiomedea melanophrys.
Location.—Intestine.
Morphology.—C ontracaecum (p. 146): Three lips (fig. 2346) 135.
long by 130» wide. Interlabia large, pentagonal, 802 long by 50.
wide near base. Esophagus (fig. 234a) 2.53 mm. long by 400p wide,
posterior part slightly differentiated into muscular bulb and with a
posteriorly directed appendix 550 long by 220 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 400u. Cuticle transversely striated.
Male 15.3 mm, long by 9002 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. Four pairs of papillae
near tail end (fig. 234d), 4 pairs about halfway from end to cloacal
aperture; in addition, starting somewhat posterior to the cloacal
aperture and running se anterior, on each side of body, is a double
row of papillae, 20 or more apables pairs.
Female 15 to 25 mm. long. Tail ending in digitiform process.
In a specimen 25 mm. long, anus 400u from posterior end, vulva 4
mm. from head end. Eggs 100 long by 60y wide.
Life history.—Unknown.
Distribution. Antarctic (in the southern Pacific (Campbell
Islands) ).
CONTRACAECUM TRICUSPE (Gedoelst, 1916) Baylis 1920b
Synonym.—K athleena tricuspis Gedoelst, 1916.
Hosts.—Ardea, species and Plotus melanogaster.
Location.—Proventriculus.
Morphology.—C ontracaecum (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 6 and ¢) and
5 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 8 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 160u wide.
with an appendix 1.6 to 2 mm. long by 190 to 2254 wide. Cecum
originating from intestine extends to the anterior fourth of the
esophagus.
Male 13.8 mm. long by 7854 wide. Tail conical (fig. 235a), ending
in a pointed appendage, curved ventrally. Cloacal aperture 140u
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 (. microcephalum.
Female 12.1 to 17.5 mm. long by 960u 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
180n, 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.
Morphology—t ontracaecum (p. 146): Lips subequal, the width
of each at base greater than its height. Interlabia slightly less high,
subtriangular. Dorsal hp 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, wide near middle of body. Lips T5y
high ; cervical constriction 75 behind base of lips. Cervical papillae
645, posterior to base of lips, oval, shghtly prominent. Esophagus
Fics. 2385—236.—235, CoNnrrRaCaECUM TRICUSPE. a, MALE TAIL; 0b, DORSAL VIEW;
C, FRONT VIEW OF HEAD. AFTER GEDOELST, 1916. © 236, CONTRACAECTM MaAG-
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» from posterior
extremity of body, which ends in an acuteiy conical tip 15 lone.
Caudal papillae (fig. 236) arranged as follows: A pair of very
large double papillae, situated 26 posterior to the cloacal aperture,
on two bosses, each boss measuring 43» 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 sublateral 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» 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
came asinmale. Vulva ona slight prominence, 8 mm. from anterior
extremity, or at about the anterior third of the body. Vagina short
(120), straight, opening into a muscular sinuous oviduct about 2
mm. long. Oviduct bifurcates to form two posteriorly directed
uteri. iAnus 300p from posterior end of body, which is conical with
a terminal spike 10n long. Eggs 68 by 45», 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. J/ale with
1 or 2 spicules. Tail usually provided with papillae, usually curved
spirally, caudal alae present or absent or a closed muscular, bell-
shaped “ bursal cup” present. /emale 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-
roxenous, the larval stages occurring in various intermediate hosts.
KEY TO SUPERFAMILIES OF SPIRURATA
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
Superfamily 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).
Ly pe-family.—Spiruridae Oerley, 1885. .
NEMATODE PARASITES OF BIRDS 163
KEY TO FAMILIES OF SPIRUROIDEA
1. Mouth with 2 large trilobed lips with a longitudinal ridge on their inner
surface meeting the one on the opposite sides; adults in gastrie traet of
earnivorous mammals; larvae may oceur in birds_. Gnathostomidae, p. 363.
Mouth without lips or with lips different from above; adults (except in
Sptrocerca): May OCCUL In "DITdS 22-5 2 ee te ae
2 Inorbital région or respiratory trach...=2. --— =... 8,
Ree eStiVe CiaACh= == =.= Soa 2 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 ef
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______________________ §.
Anterior region lacking cuticular ornaments___________________________ ie
6. Head with 4 pinnate, posteriorly directed processes; mouth without lips or
with 2 small papilliform lips; bird forms found in gizzard.
Ancyracanthidae, p. 361-
Cuticular ornaments of anterior region different from above; 2, 4, or 6
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+. 4,165 15. ott te Fh et ce Spiruridae, p. 163.
Family SPIRURIDAE Oerley, 1885
Family diagnosis —Spiruroidea (p. 162): Mouth with 2 or 4 ex-
panded lips, or without lips. Ma/e 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, generally in the diges-
tive tract.
Ty pe-genus.—Spirura F. Blanchard, 1849.
KEY TO SUBFAMILIES OF SPIRURIDAE
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 or with circular projecting disk; cervical region without
cuticular bosses; vulya usually remote from posterior extremity.
Spirurinae, p. 164.
164 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Subfamily SPIRURINAE Railliet, 1915
Subfamily 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.
Ty pe-genus.—Spirura EK. Blanchard, 1849.
KEY TO GENERA OF SPIRURINAB
1. Larvae encysted, probably accidentally, in birds; adults in other hosts.
Spirocerca, p. 202.
AGES ane ir See ee ae ee ee ee eee 2:
2. Cephalic extremity covered by a circular projecting disk__-__ Viguiera, p- 201.
Cephalic extremity possessing no such disk but with the usual lip structures
ssid SN en, Se Ee ea A CR ae 3.
S Mouth. swith 4) slipse 2. see a eh a Se ee pees 4,
VEO ULE WUE UNS se ae ST oR as a 5.
4. No lateral alae on body; sensory papillae postcervical; postanal papillae
SV TTUENYG Bel Gell see secre era a catcied Neth aw nc vai ee eee Cyrnea, p. 166.
Lateral alae usually present; sensory papillae precervical; postanal papillae
ELS AVIIINC ETL Cate sk a A 2 lh As Habronema, p. 172.
5. Mouth surrounded by chitinous ring; lips small, integral____ Spirura, p. 164.
No chitinous ring around mouth; lips comparatively large, trilobed, at
IES Su Stee NM TSN SAT Dyess pe ee A 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
Generic 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. Jfale with unequal spicules and with caudal alae which,
however, are not continuous across the ventral surface; 4 pairs of
preanal papillae. /emale with vulva anterior or posterior of middle
of body; two uteri.
Parasitic in stomach and intestine of mammals and birds.
Type-species—Spirura 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 Spirwra
talpae (Filaria strumosa) 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. falpae, and that
the parasite was accidental in the bird.
SPIRURA ZSCHOKKEI Railliet and Henry, 191la
Synonym.—sS piroptera alata Zschokke, 1889, not Rudolphi, 1819,
renamed.
Host.—Primary: Rhea americana; secondary: Unknown; prob-
ably an insect.
Location.—Proventriculus.
Morphology—sSpirura (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
lp 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 gastrophila have been found en-
cysted in cockroaches.
Nistribution—South America (Brazil) and Europe (Switzer-
Jand (Zoological Garden, Basel) ).
SPIRURA UNCINIPENIS (Molin, 1860) Railliet and Henry, 19lla
Synonyms.—s piroptera uncinipenis Molin, 1860: Cheilospirura
uncinipenis (Molin, 1860) Diesing, 1861.
Host.—Primary : Rhea americana; secondary : Unknown, probably
an insect.
Location.—Proventriculus.
Morphology.—S pirura (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. 237a and 6 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. wide. Cloacal aperture
1/22 of total length from posterior end. Caudal region (fig. 237e),
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,
(Linstow) or 180, long (Railliet and Henry), thick and ending
in a hook with rounded point (fig. 238¢c).
Fies. 237-238.—SPIRURA UNCINIPENIS. 237, a, LATDRAL VIEW; b, FRONT VIEW OF
HEAD; C, MALE TAIL. AFTER DRASCHE, 1884. 238, a, HEAD; b, EGG; ¢, FREE END
OF SHORT SPICULE. AFTER LINSTOW, 1899
Female 18 to 26 mm. long by 700 to 8004 wide. Tail 1/100
total length. Vulva almost at middle of body, salient. Eggs (fig.
238b) 43 to 47p long by 26 wide, embryonated; shell thick, rein-
forced at each pole.
Life history—Unknown; probably involving an insect as inter-
mediate host, as larvae of Spirura 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. /emale with vulva
usually situated considerably posterior to middle of body. Ovejec-
tor with vestibule differentiated into an organ for storage of eegs.
Uteri parallel.
NEMATODE PARASITES OF BIRDS 167
Parasitic usually under corneous tunic of gizzard of birds.
Ty pe-species.—C yrnea eurycerca Seurat, 1914e.
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 _____-_-------__ me
BP VaVe, PLOMIMCNE-- S.— e ne ee en Cyrnea semilunaris, p. 171.
Velva not prominbntii isis esis ce Cegees se bie Be) ssh ee Sef 3.
8. Left spicule 2.0 mm., right spicule 3654 long; male with 10 pairs of caudal
papillae; tail) of female 332u long___--__-_--+---___- Cyrnea colini, p. 168.
Left spicule not over 1.7 mm., right spicule 3804 or longer; male with 9
pairs of caudal papillae; tail of female not over 285u long____--___--_ 4.
4. No papillae directly in front ef cloacal aperture; caudal alae 410 to 550, long,
ther Span 275 tO, co0 ns 22 ees Boa ee Cyrnea seurati, p. 172.
A pair of papillae directly in front of cloacal aperture; caudal alae about
290u long, their tga yO peers Be Bek a ee eS ees iy
5. Left spicule 1.7 . right spicule 380u long; tail of female 2854 long; vulva
720u anterior a anus; eggs 42u long by 18% wide_ Cyrnea eurycerca, p. 167.
Left spicule 1.8 mm., right spicule 4904 long; tail of female 140u long; vulva
1.14 mm. anterior to anus; eggs 52u 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 945u 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 250
wide (fig. 239c). Buccal cavity 554 long; muscular esophagus 285.
long; total length of esophagus 2.5 mm. Excretory pore and sensory
papillae 280 from anterior end. Caudal alae (fig. 2396) outspread,
short, the semilunar wings transversely striated, 290u long, a litile
greater than the span from margin to margin, 250u. Nine pairs of
long pedunculated papillae, of which 3 are preanal; in addition 2
pair of very small papillae directly anterior to the cloacal aperture.
Left spicule 1.68 mm., right spicule 880n long. Gorgeret 70, long.
Female 14.7 to 20.5 mm. long. Tail (fig. 239@) attenuated, rounded
at end, 285 long. Vulva not salient, 720u anterior to anus. Vesti-
bule 1.05 mm. long. Eggs 424 long by 184 wide, with thick shell.
Life history—Unknown.
Distribution.—Europe (Corsica) and Africa (Algeria).
CYRNEA EXCISA (Molin, 1860) Seural, 19i4c
Synonyms.—Spiroptera excisa Molin, 1860; Physaloptera striata
Linstow, 1883.
168 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Hosts—Primary: Ciconia alba, C. a. asiatica, C. ciconia and C.
maguari. Winstow has also reported this species from the snake,
Tropidonotus hydrus. Secondary: Unknown.
Location.—Esophagus, proventriculus, and gizzard.
Morphology —Cyrnea (p. 166) : Blood-red in color. Sensory pap-
illae far posterior to nerve ring. The lateral lips (fig. 240a and 6)
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
Fie. 239.—CYRNEA EURYCERCA. @, FEMALE TAIL, SHOWING OVEJECTOR ; 0, MALE TAIL;
C, MALE (LENGTH 7.6 MM); d, LATERAL VIEW; €, 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 620u long. Gorgeret present.
Female 19 to 26 mm. long by 670u wide (fig. 240c). Tail short
(300). Vulva not salient, little visible, immediately in front of
middle of body. Vestibule 1.6 mm. long, sphincter 800z long. Eggs
42u long by 21p wide.
Life history —Unknown.
Distribution—South America (Brazil), Africa (Algeria), and
Asia (Turkestan).
CYRNEA COLINI, new species
Hosts—Primary: Colinus virginianus,; secondary: Unknown.
Location.—In wall of proventriculus, at its junction with gizzard.
Morphology. Cyrnea (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
Fic. 240.—CyYrNEA EXCISA. a, DORSAL VIRW; 0b, LATERAL VIEW OF HEAD END; C,
FEMALE; d, MALE TAIL. AFTER SEURAT, 1914
(fig. 2426 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
Se reece ee Fite ey
OO.u
Fic. 241.—CyrNEA SbURATI, TAIL END OF MALE AND OF FEMALE. AFTER LOPEHZ-
NEYRA, 1918
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 2504 wide. Buccal cavity 58. deep;
total length of esophagus 2mm. Caudal alae (fig. 248a) outspread,
280u long, their span 230u; they bear coarse transverse striations.
fo
0-051] @
Fig. 242.—CyRNBA COLINI. HBAD. ad, VENTRAL; b, FRONT; 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» wide, the right 365» long by 24y
HIG, 243..-CYRNEA COLINI. @, MALE TAIL; 0, FEMALE TAIL; C, OVEJECTOR. (SCALP
FOR b AND @ 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 754 deep;
first part of esophagus about 2804 long; second part of esophagus
about 2.5 mm. long. Tail (fig. 243b) 3324 long. Vulva (fig. 2480)
NEMATODE PARASITES OF BIRDS 171
only slightly salient, 915, anterior to anus. Vestibule, including
sphincter, 1.2 mm. long (fig. 243¢). Eggs 40.54 by 22.5u.
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 yen-
tral hip deeply notched in the middle.
Male 11.7 mm. long by 265» wide. Tail 160» long. Spicules
nnequal, 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 1404 long. Vulva
1.14 mm. from caudal extremity, not salient. Ovejector like that of
some species of Habronema of birds; vestibule very short (300,) ;
unpaired trompe 660p. Eggs 52 long by 24 wide.
Life history——Unknown.
Distribution.—Africa (Algeria (MacMahon) ).
CYRNEA SEMILUNARIS (Molin, 1860) Seurat, 1914f
Synonyms.—S piroptera semilunaris Molin, 1860; Spiroptera lan-
ceolata Molin, 1860.
Hosts—Primary: Crotophaga major, Trogon collaris, and 7. mel-
anurus; secondary: Unknown.
Location.—Between the tunics of the gizzard.
Morphology.—C yrnea (p. 166): Male 6 to 10 mm. long by 150.
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 4002 wide. Caudal ex-
tremity straight, obtuse; anus remote from caudal apex. Vuiva
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. yrnea (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 18.6 mm. long by 300 to 360 » wide. Tail not rolled;
caudal alae outspread, strongly striated transversely; their length
410 to 550, the span of the 2 alae 275 to 390y. 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 cloacal aperture. Left spicule 1.25 to
1.39 mm., right spicule 450 to 500u long. Gorgeret 65 to 70p long.
Female 13 to 17 mm. long by 320 to 420 » wide. Tail (fig. 241)
140 to 1804 long. Vulva not salient, 700 to 950. anterior to anus.
Ovejector made up of small pyriform vestibule 300 to 420y long and
a tubular sphincter. Eggs 45 to 48y long by 25 to 26n wide.
Life history—Unknown.
Distribution —Europe (Spain).
Genus HABRONEMA Diesing, 1861
Generic 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 eggs.
Uteri divergent.
Parasitic in digestive tract, usually stomach, of birds and mammals.
Intermediate host, where known, arthropods (flies).
Ly pe-species—Habronema muscae (Carter, 1861) Diesing, 1861.
KEY TO SPECIES OF HABRONEMA
i. No lateral alae presente 2 Fp ey eg Habronema ficheuri, p. 174.
One or, More lateral’ alae: present! ce Tt ee as ye ee oe
a Only one lateral "ata resent. 2 oars Liar aon eee Seen enya nde oe
Two lateral. alae present although they may be poorly developed (1.
$euradt) <e- teae cyt starr sue 3 oetiehs bebo meee ee 6.
3. Long slender spicule 214 times the length of the short broad one; vulva
PLOMIN CM GY eee Ae Rae eae erg Habronema unilateralis, p. 186.
Long slender spicule over 3 times the length of the short broad one; vulva
Mot Wrominen tc as FS ee ee Dele tea 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 600u long; gorgeret highly devel-
oped and of complex structure, being hollowed out and having a strong
Gorell (SUT as as oe Se Habronema tulostoma, p. 184.
Lateral ala does not extend posterior to termination of glandular esophagus 3
right cervical papilla anterior to the left; short spicule not over 460u;
gorgeret simple in structure or poorly developed and difficult to see__ 5.
5. Male 5.5 to 6.9, female § to 9.4 mm. long; cuticle with many fine needle-like
spines, 4.44 long; lateral ala 8 to 124 wide; long spicule 1.2 to 1.4 mm.
IGS Gone ee rs ead Go ee eh eh Habronema spinosa, p. 183.
Male 7.6 to 9.8, female 12.5 to 13.6 mm. long; no spines on cuticle; lateral
ala attains width of 18 to 22u; 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.
. Dorsal and ventral lips very small and apparently lacking a median pro-
jecting crest; internal to the lips 2 rows of 8 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 ;
a)
nowrows OL. papillae: as above —_..~_..-~-=- =—-===—~.-— 2 SS.
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-
10. Lateral alae of unequal length, one about 1 mm. long, the other 500, Jong,
their margins strongly crenulated; anus of female about 200u from
DOSEELION ExtremitVas2-—- tS L ae: Habronema incerta, p. 175.
Lateral alae of equal length, extending 1/4 of body length; margin not
crenulated ; anus of female about 360u from posterior extremity.
Habronema leptoptera, p. 176.
HABRONEMA COLAPTES Walton, 1923
Host.—Primary: Colaptes awratus luteus; 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 a, 6 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 alae
narrow.
174 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 10 to 15 mm. long by 300n wide. Caudal alae (Fig. 244¢)
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. 244d), 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
Vig. 244.—HABRONEMA COLAPTES. @, DORSAL; b, LATERAL; C, FRONT VIEW OF HEAD;
Soe ee AND GUBERNACULUM; €, MALE TAIL; f, FEMALE TAIL. AFTER WALTON,
40u 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, TI-
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. 245) with keel-like projection.
Male 8.4 mm. long by 3364 wide. Tail (fig. 245¢) 1202 long. Two
long caudal alae, transversely striated. Ventral region of body also
transversely striated for 450 posterior to cloacal aperture. Four
NEMATODE PARASITES OF BIRDS 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 350p long. Gorgeret 70u long.
Female 12.8 mm. long by 3854 wide. Tail 170» long. Vulva 6.6
mm. from anterior end of body, slightly salient. Vestibule 300,
long; sphincter 240u long; unpaired trompe 120 long, dividing into
2 branches 720z in length. Eggs 47» long by 234 wide, embryonated
at maturity.
Fic. 245.—HABRONEMA FICHEURI. a, LATERAL; b, DORSAL VIBW OF HEAD; C, MALB
TAIL. AFTER SEURAT, 1916
Life history—Unknown; probably involving insects as interme-
diate hosts.
Distribution—Africa (Algeria).
HABRONEMA INCERTA (Smith, 1908) Gendre, 1922
Synonym.—s piroptera incerta Smith (A. J.), 1908.
Hosts —Primary: Bolborhynchus lineola, B. monachus, Chrysotis
auripalliata, C. guatemalae, C. leucocephalus, C. ochroptera, Conurus
leucotis, C. pertinax, Eclectus roratus, Lophortyx californicus,
Palaeornis fasciatus, Platycercus barnardi, P. eximius, P. palladi-
ceps, Poeocephalus senegalus, Protogerys virescens; 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 310. wide, the maximum 12 mm. long by 350, wide. Caudal
alae slightly asymmetrical. Ventral surface of tail (fig. 246q)
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 3504 from posterior end. Spicules unequal.
Fic. 246.—HABRONEMA INCERTA. @, MALE TAIL; 0, EGGS; c, FEMALE TAIL; 4,
TRANSVERSE SECTION OF ANTERIOR END; €, ANTERIOR END; f, FEMALE, SHOWING
POSITION OF VULVA; g, FRONT VIEW} h, DORSO-VENTRAL VIEW OF HEAD. AFTER
SMiItH, Fox, aND WHITE, 1908
Female size varies in different birds, the average being 14 mm.
long by 450p. wide, the maximum 20 mm. long by 600z wide. Anus —
(fig. 246¢) about 200. from posterior end. Vulva (fig. 246f) 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. 2460) 34 to 42u long by 16 to 20n 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, Ff. cineraceus, F. cyaneus, F.
lanarius, F. magnirostris, F. nisus, F. palumbarius, F. rufus, F. sub-
buteo, F. tinnunculus, F. tridentaius, F. xanthothorax, F. unicinctus,
Harpagus bidentatus, Herpetotheres cachinans, Hypotriorchis sub-
buteo, Milvus 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
Fic. 247.—HABRONEMA LEPTOPTERA. a, ANTERIOR END, DORSAL VIEW; b, MALE TAIL.
AFTER SpurAT, 1914
far anterior to nerve ring. Head (figs. 247a and 248@) 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 4 papillae at the point of insertion of these
two lips. Muscular esophagus long and slender.
Male 71 mm. long by 720% wide. Spicules unequal, the left long
and slender, 640 in length; the right 200p 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 325y wide. Tail 360n long. Vulva (fig.
248c) 8 mm. from posterior extremity. Eggs elliptical, 32» long by
20 wide, embryonated at maturity.
Life history—Unknown; probably involving insects as interme-
diate hosts.
Distribution—Kurope and Africa (Algeria).
Fig. 248.—HABRONEMA LEPTOPTERA. @, HEAD, LATERAL VIEW; b, RECEPTACLE
SEMINALE, JOINING OVIDUCT WITH UTERUS; ¢, OVEJECTOR. AFTPR SEURAT,
1914
HABRONEMA LONGISTRIATA (Molin, 1859) Gendre, 1922
Synonyms.—S piroptera longistriata Molin, 1859, in part; Cheilo-
spirura longistriata (Molin 1859) Diesing, 1861, in part.
Hosts—Primary: Picus campestris and P. jwmana; secondary:
Unknown.
Location —Proventriculus.
Morphology.—H abronema (p. 172) : Body rolled in spiral. Head
(fig. 249a and 6) 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 200n wide. Caudal alae (fig. 249¢) wide,
symmetrical. No unpaired papilla anterior to cloacal aperture. In
other respects bursa apparently like that of 1. mansioni (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 179
HABRONEMA MANSIONI Seurat, 1914
Synonyms.—S piroptera longistriata Molin, 1859 in part.; Cheilo-
spirura longistriata (Molin, 1859) Diesing, 1861 in part.
Hosts—Primary: Buteo vulgaris and Picus grammicus. Possibly
also in Milvus korschum,; secondary: Unknown.
Location.—Proventriculus.
Morphology. abronema (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 @) 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 315h wide. Tail (fig. 250 6) straight.
An unpaired papilla on left side of body 500 anterior to origin of
Fic. 249.—HABRONEMA LONGISTRIATA. @, HEAD, VENTRAL VIEW; 0, HEAD, FRONT
VIEW ; ¢, MALE TAIL. AFTER DRASCHE, 1884
caudal alae. Alae slightly asymmetrical, elongate (840n). 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, 680p and 315y long. Gorgeret present.
Female 13 to 17 mm. long by 400u wide. Tail 240n long. Vulva
(fig. 250 ¢) very small, situated slightly anterior to middle of body.
Ovejector like that of 17. leptoptera (p. 176). 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 korschwm 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, 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 280u from the anterior extrem-
——
Fic. 250.—HABRONEMA MANSIONI. @, ANTERIOR END, VERTICAL VIEW; b, MALE
TAIL; c, OVEJECTOR. AFTER SEURAT, 1914
ity and extending to the level of the termination of the glandular
esophagus, its greatest width 18 to 224. Cervical papillae in inverse
position from those of H. tulostoma, the right one being anterior to
the left. Head (fig. 251 6 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 350u wide. Lateral ala about
2.58 mm. long. Tail (fig. 251 d) 230 to 2604 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 610, 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 f) unequal, the
mae |
yt 4
s
Fic. 251.—HABRONPMA MONOPTHRA. a, ANTERIOR END; D, LATERAL LIP; C€, MEDIAN
LIP; d, MALE TAIL; e, LEFT SPICULE; [f, RIGHT SPICULD; g, FEMALE TAIL. AFTER
GENDRE, 1922
Q\
ea
{/ oo
an)
one 1.56 to 1.68 mm. long, the other 380 to 460u long. Gorgeret diffi-
cult to see.
Female 12.3 to 13.6 mm. long by 4380 to 440, wide. Lateral ala
about 2.98 mm. long. Tail (fig. 251 7) 180u long. Vulva near middle
of body, sometimes a little anterior, sometimes a little posterior to
middle, not salient, difficult to find. Vestibule 810n long. Eggs 38
to 42n long by 17 to 19 wide.
182 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history—Unknown; probably involving an insect as inter-
mediate host.
Distribution —Africa (French Guinea (Labé) ).
HABRONEMA SEURATI Skrjabin, 1917
Host.—Primary: Falco 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 1444 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. mansioné (p. 179).
yh
es
=
r~
—
()
Fig. 252.—HABRONEMA SEURATI. @, TRANSVERSE STRIATION OF BODY; 0b, CUTICULAR
STRUCTURD OF VENTRAL SURFACE OF MALE TAIL; C, MALE TAIL; d, LEFT SPICULB; €,
RIGHT SPICULE. AFTER SKRJABIN, 1917
Male 5.5 mm. long by 300 wide. Caudal alae (fig. 252¢c) 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. 2526). 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 4504 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 2004 from posterior end.
Female 9.2 mm. long by 370u wide. Vulva 3.4 mm. from pos-
terior end, thus in the posterior part of body. Eggs 47» by 27.5
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 tinnunculus,; secondary: Unknown.
Location.—Proventriculus.
Morphology.—Habronema (p. 172): Body white, cuticle trans-
versely striated and bearing a multitude of fine needlelike spines
A,
“h
J
Fig. 253.—HABRONEMA SPINOSA. a, SPINES OF CUTICLE; D, HEAD PND; C, MEDIAN LIP ;
d, MALE TAIL; e€, LEFT SPICULP; f, RIGHT SPICULE; g, GORGERET; h, FEMALP TAIL.
AFTER GENDRE, 1922
=
(fig. 253a), visible at high magnification (560), their length about
4.44. Right cervical papilla anterior to left. One lateral ala, origi-
nating 26 to 30n posterior to left papilla, which is 130 to 150p from
the anterior end in the male, 160 to 180 in the female. Ala shorter
and narrower than in /7. monoptera; its width 8 to 12 (see below
for length). Head (fig. 2536) very small; 4 lips all of same length.
The lateral lips with 3 or 4 teeth on the internal surface are not as
large and projecting as in other species of Habronema, while the
dorsal and ventral lips (fig. 253¢) are proportionately more highly
3612—27 14
184 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
developed and their lateral lobes again subdivided into secondary
external lobes, bearing a papilla.
Male 5.47 to 6.86 mm. long by 210 to 260 wide. Lateral ala 1.82
to 2.55 mm. long. Tail (fig. 253d) 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 6504 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 f) unequal, 1.22 to 1.87 mm. and
340 to 410» long respectively. Gorgeret (fig. 253g), simple.
Female 8 to 9.37 mm. long by 380 to 410n wide. Lateral ala 1.77
to 1.94 mm. long. Tail (fig. 253) 180 to 1504 long. Vulva difficult
to see, a little posterior to middle of body. Vestibule 610, long.
Eggs 37 long by 21n wide, embryonated.
Fic. 254.—HABRONEMA TULOSTOMA. a, ANTERIOR END; 0, 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 (Hemprich and Ehrenberg, 1866) Gendre, 1922
Synonyms.—S piroptera tulostoma Hemprich and Ehrenberg in
Schneider, 1866; /ilaria tulostoma Schneider, 1866; Habronema uni-
lateralis (Molin, 1860) of Seurat, 1914f misdet.
Host.—Primary: Neophron percnopterus (Vultur percnopterus }
and NV. monachus, secondary: Unknown.
Morphology.—Habronema (p. 172): Body blood red in color;
cuticle finely striated transversely. Mouth with 4 lips, the 2 lateral
(fig. 2546) 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 OF BIRDS 185
Male 17 mm. long (Schneider) or 13.9 to 16.7 mm. long (Gendre)
or 8 mm. long by 250% wide (Seurat). Tail (fig. 255@) straight.
According to Seurat’s description, caudal alae very long, 1.23 mm.,
the span 300,; cloacal aperture 800 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 unpaired papilla on the left side of anterior edge
of cloaca. Spicules dissimilar and unequal, 6002 and 1.9 mm. long
(Seurat), 510 to 550n and 1.77 to 1.8 mm. long (Gendre). Gorgeret
(fig. 2555) 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),
ii to 14 mm. long (Seurat). Tail 140» long in Seurat’s specimens,
260 to 310n long in those of Gendre. Vulva 1/3 of body length from
Fic, 255.—HABRONEMA TULOSTOMA. (d, MALD TAIL; 0b, CLOACAL REGION, SHOWING
a
GORGERET; ¢C, OVESECTOR. APrreR SEURAT, 1914
anterior end (Seurat) or varying, sometimes a little anterior, some-
times a little posterior to middle of body (Gendre). Eggs 42u long
by 254 wide (Seurat), 35 to 394 wide by 21 to 234 wide (Gendre).
Life history—Unknown; probably involving an insect as inter-
mediate host.
Distribution.—Asia (Turkey (Tor)) and Africa (French Guinea
(Labé) 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 H. 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 Habronema unilateralis,
186 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
namely the relative length of spicules, the longer spicule in H. uni-
lateralis being only 21% 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. unilateralis, 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, 18600; Spiroptera
unialata Molin, 1860.
Morphology—Habronema (p. 172): Body slender anteriorly,
rhynchus and FR. vitellinus; secondary: Unknown.
Location —KEsophagus and “ventriculus” (gizzard or proventri-
culus?).
Fic. 256.—HABRONEMA UNILATERALIS. @, HEAD, FRONT VIEW; Db, MALE TAIL. AFTER
DRASCHE, 1884
Morphology abronema (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 200n wide. Caudal extremity (fig. 2560)
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 sessile
papillae and an unpaired papilla directly anterior to the cloacal
aperture. Left spicule long, slender, alate, 214 times the length of
the right, which is short and thick.
Female 7 to 20 mm. long by 300 to 5004 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 7. tulostoma, see
page 185.
Genus HADJELIA Seurat, 1916c¢
Synonym.—Gilsonia Gedoelst, 1919.
Generic diagnosis —Spirurinae (p. 164): 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.
Ty pe-species.— Had jelia lhuilliert Seurat, 1916c.
KEY TO SPECIES OF HADJELIA
EDD teri (in OWN 2 oe ot Hadjelia truncata, p. 190.
Ween 400 0r lone eras 8 at ee ee 2.
2. Females not over 9.7 mm. long; buccal cavity of female 32 to 37y long.
Hadjelia parva, p. 189.
Females 12 mm. or longer; buccal cavity of female 504 or longer_________.. 3.
3. Vulva salient and limited by 4 or 5 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 —..___ 2.54 este Hadjelia lhuillieri, p. 187.
HADJELIA LHUILLIERI Seurat, 1916c
Host.—Primary: Caccabis petrosa; secondary: Unknown.
Location.—Gizzard.
Morphology—Hadjelia (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
sarge 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 280% wide. Buccal cavity 60y, entire
esophagus 3.1 mm. long, muscular esophagus 480y. Tail short, anus
110, from posterior end. Vulva 2.9 mm. from anterior end of body,
188 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
or 335p 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).
= Te MT
c
el Heh
ot hi
Z UT
Ligh
Fic. 257.—HADJELIA LHUILLIERI. dad, HEAD END, LATERAL VIEW ; 0, HEAD, DORSAL VIEW.
AFTER SHURAT, 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.
Hic. 258.—HADJELIA LHUILLIPRI. VULVA AND OVEJECTOR. AFTER 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 554, esophagus 2 mm. long. Cloacal aperture 120, 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 6). Spicules unequal, the left (fig. 259¢) 1.6 to 2.1 mm.
long, the right (fig. 259d) 200 to 280, lone.
NEMATODE PARASITES OF BIRDS 189
Female 12 to 21 mm. long by 170 to 260 wide. Buccal cavity 50.
to 60n, esophagus 2.4 to 3.6 mm. long. Anus (fig. 259e) 90 to
130 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» long by 30 to 33p
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. lhuillieri, the male of the latter being un-
Fic. 259.—HADJELIA INERMIS. a, MALB TAIL AS USUALLY SEEN (MOST POSTERIOR
PAPILLAE HIDDEN); 6, MALE TAIL STRAIGHTPNED OUT; ¢, LEFT SPICULE; d, RIGHT
SPICULE; €, FEMALE TAIL. AFTER GENDRD, 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: Trachelotis senegalensis; secondary: Unknown.
Location.—Gizzard.,
Morphology.—Hadjelia (p. 187): Male 5.5 mm. long by 180,
wide. Buccal cavity 33 long; first part of esophagus 240,, second
part 1.66 mm. long. Cloacal aperture 110» from posterior end. Six
pairs of pedunculated papillae (fig. 2607) on large caudal alae.
Spicules unequal, the left 1.29 mm., the right 240, long.
190 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Female 9.2 to 9.6 mm. long by 200 to 2602 wide. Buccal cavity
32 to 37. long; first part of esophagus 280 to 330p, second part 1.86
to 1.90 mm. long. Anus (fig. 260b) 110. from tail end. Vulva
1.52 to 1.97 mm. from head end. Eggs 48 to 49 long by 29 to 30,
wide.
Life history—Unknown.
Distribution.—Africa (Abomey, Dahomey).
£.
Fic. 260.—HADJELIA PARVA. @, MALB TAIL; b, FEMALE TAIL. AFTER GENDRE, 1922
HADJELIA TRUNCATA (Creplin, 1825) Gendre, 1921c
Synonyms.—s piroptera upupae Rudolphi, 1819, nomen nudum;
Spiroptera truncata Creplin, 1825; Dispharagus truncatus (Creplin,
1825) Dujardin, 1845; Acuaria upupe-epopis Molin, 18600.
Hosts —Primary: Coracias garrula and Upupa epops; secondary :
Unknown.
Morphology—Hadjelia (p. 187): Body slender; color, red.
Mouth (fig. 261@) with 2 large lips, their base concave, their anterior
Fic. 261.—HADJELIA TRUNCATA. 4d, HEAD; 0, 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 T 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% 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 are
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. J/ale 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.
Ty pe-species.—H artertia obesa Seurat, 1915b.
KEY TO SPECIES OF HARTERTIA
1. Only 3rd stage larva known (from under skin of carnivore).
Hartertia zorillae, p. 200.
ACS TRMOWIe IN DITdS! 2052s Kee ee eee a 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 200u 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 450u, or longer_______________ 4.
4. Male 35 to 37 mm. long; long spicule 1.35 mm. long; vulva of female posterior
to. anterior} third’ of; vodyas-sa tse Hartertia rotundata, p. 198.
Male not over 30 mm. long; long spicule over 2 mm. long; vulva of female
in) anterior: third of Wed yeu hee 8 iri tees fe Jocuri ey lt ee 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 39u long by 264 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 56u
long; by ;pour widee teailes jac VaR ort) gets gg ta Mab 6.
6. Tail of male 1/73 of total body length; spicules 3.3 mm. and 647u 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.
PAPASitiC Il PIZZAT Oe Se ee Hartertia annulata, p. 193.
Tail of male 1/31 of total body length; spicules 2.1 mm. and 500u 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
Mmtestinemio be wees Fe EA bee Hartertia obesa, p. 192.
192 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
HARTERTIA OBESA Seurat, 1915b
Hosts —Primary: Caccabis petrosa spatzi, Oedicnemus capensis,
Otis afroides, O. ruficresta, and Plocepasser mahali; secondary :
Unknown, probably insects.
Location.—Intestine.
Morphology—Hartertia (p. 191): Head very small, distinct.
Cuticle finely striated. Lateral fields very large (1554), 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 145p
from cephalic extremity. Buccal cavity 160u long; esophagus 1/6 of
body length. Tail (fig. 263b) straight. Ventral surface of tail in
region of cloacal aperture covered with longitudinal rows of cuti-
{mm soe
LATERAL; €, FRONT VIEW OF HEAD,
a, DORSAL; D,
d, mGG. AFTER SEURAT, 1915
Fic. 262.—HARTERTIA OBESA.
cular shields. Cloacal aperture 7504 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 500n long, respectively. Gorgeret
present, 200n 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.
262d) peculiar, the thick shell measuring 56 by 35n, is doubled by a
vitelline membrane, showing in the meridian plane as a thin ring
10u wide.
NEMATODE PARASITES OF BIRDS 193
Life history—Unknown; probably similar to that of H. galli-
narum (p. 197).
Distribution—Africa (Oued N’za and Transvaal).
HARTERTIA ANNULATA, new species
Host—Primary: Red-headed pheasant (Pternistes, 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, 6, and c) with 2 lateral lips,
each trilobed, the lateral lobes finely denticulated and all 3 lobes
Fig. 263.—HARTERTIA OBESA. ad, OVEJECTOR; Db, MALE TAIL. AFTER SHURAT, 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 26 to 30 mm. long by 800 to 9002 wide. ‘Anterior part of
esophagus 531p long by 144» wide, posterior part 3.65 mm. long, its
minimum width 183», maximum width 366y. Posterior extremity
of body (fig. '265) straight or loosely curled. Caudal alae 1.4 mm.
long; they are narrow for a distance of 133» from the posterior
end, anterior to that they flare out comparatively wide, their span
being 6381p. Ventral surface densely covered with longitudinal rows
of small rounded protuberances. Four pairs of pedunculated preanal,
2 pairs of pedunculated postanal papillae, all pairs symmetrical.
Cloacal aperture 382, from tail end. Two dissimilar spicules (fig.
194 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
265), the one 3.3 mm. long by 17m wide, its distal end sharply
barbed, there being 3 or more recurrent points or processes; the
cther 674 long by 58, wide, its distal end bluntly rounded.
Female 43 to 62 mm. long by 966 to 1.2 mm. wide. Anterior
part of esophagus 697, posterior part about 5.2 mm. long in large
specimens. Anus 1.1 mm. from posterior end of large specimen;
—<$___—_—_—_— a
02MM. O2 17/77.
Fic. 264.—HARTERIA ANNULATA. @, LATERAL; b, FRONT; C, DORSAL VIEW OF HEAD;
d, EGG; @, FEMALE TAIL; f, OVEJECTOR; g, ENLARGED VIEW OF CUL-DE-SAC AT UNION
OF SPHINCTER AND TROMPER. 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 14 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
Fie. 265.—HARTERTIA ANNULATA. MALP 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 f 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
5002 and containing an enormous number of eggs. The ovaries
are long and filiform, 832 in diameter. Eggs 50» long by
354 wide, embryonated when oviposited (fig. 264d).
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 7, not Habronema rotundata
(Linstow, 1883) Seurat, 1914 g and n.
Host—Primary: Otis houbara; secondary: Unknown, probably
insects.
Location.—Intestine.
Morphology.—tHartertia (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 510u long, both with rounded ends.
Female 20.3 mm. long by 1.5mm. 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
wide, thick-shelled, embryonated.
Life history——Unknown; probably similar to that of ZZ. 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 Filaria 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 macqueenz, collected in Turkestan by Professor
Yedtschenko; the 1906 specimens were described as a new species
from Otis houbara, in the Zoological Museum at Ko6nigsberg.
Seurat (19143117) 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-
ronema by Seurat (19143153) but he gives the locality of collection of
the nematode as Algeria and Turkestan, which is not that of the 1906
Figs. 266—-268.—266, HarTerRTIA CONFUSA. MALE TAIL. AFTER LINSTOW, 1906. 267,
HARTERTIA GALLINARUM. @, LATERAL; 0, FRONT VIEW OF HEAD; C, FEMALE TAIL. 268,
OVEJECTOR AND UTERI. ORIGINAL
species but of Linstow’s earlier species, taria 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 7. 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
over
NEMATODE PARASITES OF BIRDS 197
of male tail (that of H. 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.—Filaria gallinarum Theiler, 1919.
Hosts —Primary: Gallus 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. 267a@ and 6)
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 498u (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 (20u 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
40y long by 24» 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.
OOM
Wests eenrgsess BESS
REE Ca
SRS SEE SEY
LESEENY a
ay serra ie :
209
O-OF/TUN
Fic. 269.—HARTERTIA GALLINARUM. 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
Synoyms.—Filaria rotundata Linstow, 1883; Habronema rotun-
data (Linstow, 1883) Seurat, 1914 g and n, not Habronema rotun-
data (Linstow, 1906) Seurat, 1914 f (see Hartertia confusa, p. 195).
Hosts —Primary: Houbara undalata, Oedicnemus oedicnemus,
O. vermicularis, Otis afroides, O. macqueeni, 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 longi-
tudinal line dividing them in two halves, the outer half more deeply
colored. Two papillae at anterior insertion of alae. Mouth (fig.
270a) with 2 tri-lobed lateral lips, the median lobe having a small
tooth, the lateral lobes a finely denticulated edge. Length of
esophagus 1/7 that of the body.
Fig. 270.—HARTERTIA ROTUNDATA. @, HEAD, VENTRAL VIEW; Db, MALE TAIL. AFTER
SHURAT, 1914
Male 35 to 37 mm. long by 660u wide. Caudal alae (fig. 2706)
symmetrical, 1.2 mm. long by 720 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 540u
from tail end. Spicules unequal, 1.35 mm. and 450 long. Gorgeret
present.
Female 35 to 66 mm. long by 900p wide at a level of alae. Cephalic
papillae 70u 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 6004 long, sphincter 900, long,
unpaired trompe 1.5 mm. long. Uteri divergent. Eggs thick-
shelled, 62 to 69» long by 42 to 56n wide, embryonated at maturity.
Life history—Unknown; probably similar to that of 77. gal/i-
narum (p. 197).
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
Fic. 271.—HARTERTIA ROTUNDATA. OVEJECTOR. AFTER 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 900u, at the level of the anus 400un. Esophagus 3.5 mm. long. Anus
5504 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—tThird stage larva (fig. 272): 13.1 mm. long by
4102 wide; tail 158, long. Precervical papillae 77» from cephalic
end; buccal cavity 60x, 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,
Fic, 272.—HARTERTIA ZORILLAB. THIRD STAGH LARVA. 4, LATERAL VIEW OF HEAD;
b, VENTRAL VIEW OF HEAD; ¢, VENTRAL; d, LATERAL VIEW OF TAIL. AFTER SEURAT,
1919
30 to 50u 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—Viquiera euryoptera (Rudolphi, 1819) Seurat,
1913.
VIGUIERA EURYOPTERA (Rudolphi, 1979) Seurat, 1913b
Synonym.—s piroptera euryoptera Rudolphi, 1819.
Host—Primary : Lanius collurio, L. excubitor, L. minor, L. rufus;
also “ pie-griéche a téte 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 125p wide. In the specimens 5.2 mm.
long, the buccal cavity measured 324, muscular esophagus 370g, total
esophagus 2.7 mm. long. Tail strongly rolled in spiral. Caudal alae
(fig. 273a) wide, attaining length of 580u. 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, 9304 and 192y. long.
Fic. 273.—VIGUIERA EURYOPTERA. 4, ee TAIL; 0B, OVEJECTOR. AFTER SEURAT,
Female 9 to 10 mm. long by 290u wide. Total length of esophagus
is 1/3 body length; muscular esophagus 780p long. Tail 120 long.
Vulva situated immediately anterior to anus, a distance of 52u. from
it, not salient. Vagina (fig. 273) 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 32u long by 18y 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 genus.
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.
Ty pe-species.—S pirocerca sanguinolenta (Rudolphi, 1819) Seurat,
1913.
SPIROCERCA SANGUINOLENTA (Rudolphi, 1819) Seurat, 1913
Synonym.—S piroptera sanguinolenta Rudolphi, 1819.
Third stage larvae reported by Seurat (19164) 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 tingitanus,
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; Sptroptera 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—Gongylonema minimum Molin, 1857 (synonym @.
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
cr Wh
Se 2 A a A br
Tie 6 es
Fig. 274.—GONGYLONEMNA INGLUVICOLA. @, HpaD; b, MALE TAIL. AFTER RANSOM, 1904
of the esophagus is short, slender, and colorless, and the posterior
portion is long, thicker, and yellow.
Male 17 to 20 mm. long by 224 to 250u wide. Cuticular bosses ex-
tend for distance of 575 to 6804 from head end. The tail (fig. 2740)
has two narrow caudal alae, the right 500 to 736 long and the
left 600 to 800u 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 0 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 275y on the tip of the tail. Left spicule as long as the body
and 7 to 9u 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 4902 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 14mm. long. Eggs 50 to 57p long by 36 to 38, wide, the shell
4u thick, and containing embryos when Me eed
Embryo 1604 long by 84 wide, and provided with a spine at the
anterior end.
ee
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 coprophagus 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. Tor this purpose
no type species is necessary.
SPIROPTERA PENIHAMATA Molin, 1860b
Synonyms.—S piroptera strigis-atricapillae Molin, 1860; Spirop-
tera strigis-flammeae Molin, 1860; Spiroptera strigis-griseatae Molin,
1860; Spiroptera strigis-lineatae Molin, 1860.
Hosts—Primary: Striv 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. semilunaris.
206 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 5 to 13 mm. long by 100 to 300u 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 lke a fishhook.
Female 7 to 15 mm. long by 300n 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 1s without decision.
Fics. 275—276.—275, SPIROPTERA PENIHAMATA. MALE TAIL, AFTPR DRASCHE, 1884.
276, SPIROPTPRA TURDI. HEAD. a, FRONT; 6, LATERAL VIEW. AFTER DRASCHE,
1884.
SPIROPTERA TURDI Molin, 1860b
Synonyms.—Ascaris fissilabium Linstow, 1873; Filaria turdi
(Mohn, 1860) Linstow, 18770.
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—sSpiroptera (p. 205): According to Drasche, the
mouth (fig. 276 a@ and 6) 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 (19136) 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 SAGINATA (Rudolphi, 1819) Dujardin, 1845
Synonym.—Physaloptera saginata Rudolphi, 1819.
Hosts—Primary : Caprimulgus guianensis, C. leucopygeus, Corcus
cajanus, Crotophaga ani, Cuculus tinguacu, Falco furcatus, Icterus
eristatus, Strix atricapilla, 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 Ortlepp (1922) has
recently stated that he has examined specimens of the species and
that it belongs in Spiroptera. As Ortlepp was monographing the
genus Physaloptera it seems reasonably sure that the species does not
belong in Physaloptera. 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 //artertia 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, 1845
Synonym.—P hysaloptera tenuicollis Rudolphi, 1819.
Hosts —Primary: Falco haliaetus; secondary : Unknown.
Location.—Intestine.
3612—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 DESMIDOCERCIDAH, new family
Family diagnosis —Spiruroidea (p. 162): Forms with distinctive
tarval 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, Vesmidocerca, 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 DESMIDOCERCINAB, new subfamily
Subfamily diagnosis —Characters of the family.
Genus DESMIDOCERCA Skrijabin, 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
are 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.—Alr 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. wide (fig. 2776). Caudal extremity
(fig. 277¢) 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 300p wide. Caudal extremity (fig. 277¢)
obtusely rounded, bearing a cluster of spine-like papillae. Vulva
in posterior region of body. Eggs 30u long by 18» wide.
Life history.—Unknown.
Distribution —Asia (Russian Turkestan).
DESMIDOCERCA NUMIDICA Seurat, 1920
TTost—Primary: Héron cendré; secondary : Unknown.
Location.—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 5.8 mm. long; posterior region curved crosswise. ‘Tail short
(S0u), 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 long.
Female 6.8 mm. long. Tail short (75u), 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.—DESMIDOCERA AHROPHILA. ad, HEAD; Db, MALE; ¢, MALE. TAIL; d,
FEMALE TAIL. AFTER SKRJABIN, 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.
Ly pe-genus.—A cuaria Bremser, 1811.
NEMATODE PARASITES OF BIRDS AE
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
PAN aes sete eens Oe ee ee Acuariinae, p. 211.
Cephalic ornamentations consisting of appendices or festoons; mouth with
2, 4 or 6 lips; preanal papillae veriable in number, sometimes very
MUDIET OUSt tan ee ee ee ee Schistorophinae, p. 254.
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. Jale 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 digestive tract of birds.
Ty pe-genus.—Acuaria Bremser, 1811.
KEY TO GENERA OF ACUARIINAB
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. (A.cuaria. Sil. Dp: 212) 22222 = 2 Be 2,
Ornamentation of a nature different from the above, confined to cephalic
PEG 1 Olean wee An keene es LAER Se Re ee ee ee 8.
2) ;Cordons mot 'recurrent,: not-/anastomosing?:_.- 21 its4s_ 3.
Cordons réecurrent or. anastomosing, or both. =~ -----__---+-_+u_+-_-+-=+ 4.
3. Both spicules thick and only slightly unequal; 6 to 8 pairs of postanal
POSURE eh eos ee ge ee Acuaria, p. 216.
Spicules very dissimilar and very unequal; 5 to 7 pairs of postanal
Te LU oem nae ei ad we eens ea ok EYL ee Cheilospirura, p. 226.
4. Cordons: not recurrent, but anastomosing____-=-~---~-=2+—2=-_=_______ 5.
Cordons recurrent, anastomosing or separate+2__--_------+____-_-___-_ 6.
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. 251.
No such collar or sheath present__-__-___---_-_-__--_- Echinuria, p. 244.
6. Cordons recurrent but not anastomosing__-----------~- Dispharynx, p. 237.
Cordons recurrent? ‘and: “ahastomosing «sn Me
7. Cordons form a loop directly after their origin on the head; cordons hot
flat against body but applied to margin of plates or alae; lateral alae
presen t-on: bod yerns sre Cosmocephalus, p, 283.
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. 264.
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
TOSUCTAOT M2 ae ee ee eee 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:=22 =+- << 2a es eee 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.—S piroptera 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.
Ty pe-species—Acuaria anthuris (Rudolphi, 1819) Railhet, 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 Acuwaria 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 Acwaria sensu lato
and covered in the following key. Other species should be traced
through the key to genera of Acuariinae.
KEY TO SPECIES OF ACUARIA SENSU LATO
ies COLOONS TECULTCNE ssa RRs ae ee ae Acuaria quadriloba, p. 215.
Cordons;Net Tecurme tee ee eee 2.
2. Only larva known; genitalia unknown; cordons 145y long.
Acuaria tarentolae, p. 216.
Adults known; cordons, where known, more than 1454 long; males
URTUIK INO OT a aL a Re ae 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. Cordons extend beyond esophagus; female 8.84 to 12.73 mm. long.
Acuaria ptilopachydis, p. 214.
a)
Gordons: do. not. extend. beyond esophagus. +—. ee ee 5.
5. Lips unequal; cordons end 330u from head; total length of esophagus less
than) 1/10 of:body lengthje2 25.42 Acuaria muscicapae, p. 214.
Lips apparently equal; cordons end 800u 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
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 470u wide. Tail 1/37 of body
length, terminating in a smaller finger-shaped point. Eggs 31
long by 11p wide.
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution —Asia (Ceylon (Wirawila) ).
This species belongs in Acuaria sensu stricto or Cheilospirura,
apparently, but can not be definitely placed until the male char-
acters are known.
ACUARIA MAMILLARIS (Molin, 1860) Railliet, Henry, and Sisoff, 1912
Synonyms.—Dispharagus mamillaris Molin, 1860c; Spiroptera
corvi-cajani* 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,
(19202), 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 Catalogue of the Vienna Museum.
214 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
ACUARIA MUSCICAPAE (Linstow, 1878) Railliet, Henry, and Sisoff, 1912
Synonyms.—Filaria muscicapae Linstow, 1878; Dispharagus
muscicapae (Linstow, 1878) Stossich, 1891.
Hosts—Primary: Muscicapa 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 1504 wide. Mouth followed by a
vestibule 160 long, and this by an esophagus of which the glandular
portion is 750n long and the muscular portion 330y long. The tail
is 1302 long, conical, with a rounded end. Vulva somewhat pos-
terior to middle of body, dividing body in ratio of 7:6. Eggs 29p
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 Chedlospirura, ap-
parently, but can not be definitely placed until the male characters
are known.
ACUARIA PTILOPACHYDIS Gendre, 1920a
Hosts.—Primary: Ptilopachys fuscus; secondary: Unknown.
Location.—Gizzard.
Morphology.—A cuaria 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 3.18
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 200n wide. Pharynx 140
to 2002 long. Esophagus 400 to 5502 long; ventricle (evidently
second part of esophagus) 1.07 to 1.43 mm. long. Tail (fig. 2780)
200 to 245u long, conical, digitiform, rounded at apex. Vulva slightly
salient, bordered with a cuticular margin and slightly posterior to
NEMATODE PARASITES OF BIRDS “B15
middle of body, apparently 10/19 of body length from head end.
Ovejector 260u long by 46, wide, directed posteriorly and without
a distinct vestibule. Eggs thick-shelled, 35 to 39 long by 21 to 23u
wide.
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution —Africa (Dahomey).
This species belongs in Acuaria sensu stricto or Cheilospirura, 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 quadriloba Rudolphi, 1819; Faria quad-
riloba (Rudolphi, 1819) Schneider, 1866; Dispharagus quadrilobus
(Rudolphi, 1819) Dujardin, 1845; Dispharagus crassus Molin, 1860e.
Hosts—Primary: Dryocopus martius, Picus martius, P. viridis,
Hylomotus pileatus, Gecinus viridis; secondary: Unknown.
Location.—Esophagus.
Fies. 278-279.—278, ACUARIA PTILOPACHYDIS. 4, ANTERIOR END; 0, FEMALE TAIL.
AFTER GBENDRE, 1920. 279, ACUARIA QUADRILOBA. HEAD, FRONT VIEW. APBTER
SCHNPIDER, 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—Kurope (Germany (Vienna Museum and Berlin) )
and North America (Philadelphia, Pa.).
3612—27——16
216 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
This species belongs in Dispharyna or Synhimantus, apparently,
but can not be definitely placed until the male characters are known
ACUARIA TARENTOLAE Seurat, 1916f
Hosts.—Primary: Unknown; secondary (aberrant): Tarentola
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.
Uf AMT
cy A MT
AA 0 LU
Fig. 280.—ACUARIA TARENTGLAE. ANTERIOR AND POSTERIOR ENDS OF THIRD STAGD
LARVA. AFTER SEURAT, 1916
Larva, third-stage, 5.35 mm. long by 1204 wide. Cordons 145,
long. Tail (fig. 280) 1564 long, ending in a small button with a
smooth surface. Buccal cavity 112», muscular esophagus 504, 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 Acuarta 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
Synonyms.—Spiroptera Rudolphi, 1819 in part; Anthuris Ru-
dolphi, 1819; Dispharagus Dujardin, 1845 in part.
Generic Times — Aouininne (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 peep illees
NEMATODE PARASITES OF BIRDS 217
Parasitic between the tunics of the gizzard or in proventriculus
of birds.
Type species —Acuaria anthuris (Rudolphi, 1819) Railhet, 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 anthuris Rudolphi, 1819, is regarded
as type of the genus Anthuris Rudolphi, 1819, by absolute tautonomy
and by Rudolphi’s original intentions. But Anthuris is Acucria
Bremser, 1811, renamed, as Rudolphi admits, and hence the type
species anthuris is type of Acuaria. But since Spiroptera is a
renaming of Acuaria and Anthuris, 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 ACUARIA SENSU STRICTO
1. Body with 6 cordons, the 4 submedian being double, festooned, and extending
10/23 of total body length, the 2 lateral being single and simpler, and ex-
tending the whole body length; spicules 280 and 250 long.
Acuaria ornata, p. 223.
Body with 4 cordons, submedian; spicule lengths, where known. differ
Trom ‘Above tise toes! acl ets ee pie De, sickest AE eve evel 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 A. tenuis)___.--_ 3.
3. Male 7 mm. long, not otherwise described ; anus of female 1604 from posterior
end ; eggs said to be only 154 long; from Hirundo, species.
Acuaria attenuata, p. 220.
If only male known (A. papillifera), is 4.7 mm. long; anus of female 225
or more from posterior end, except in A. gracilis; eggs 32 to 48u long;
HOG. Ji Berwniio,-species: 1 AA SO) Ve LOR besip dein Papers 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 234y Jong.
Acuaria anthuris, p. 218.
Cordons, if described, extend for length of not more than 600; if length
not described (A. tenwis), spicules only 183 and 95y long_--__-_________ 5.
5. Spicules 133 and 95u long; cloacal aperture 1354 from posterior extremity ;
from’ Saavicola rubétra 22222 tele ee eae 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 400u from posterror extremity; not from
SOLICOLURTALDC Ti a Es San ee ae 6.
6 Male with, 10: pairstof caudal papillae®== 25 2s se eee ee eee Ge
Male-“with' 12 pairs*of caudal’ papilla#e_ 2222220) ee TOreess ae aes 8.
=]
. Vulva at middle or slightly in front of middle of body; eggs 32y long.
Acuaria gracilis, p. 222.
Vulva posterior to middle of body; eggs 37 to 39u long.
Acuaria subula, p. 224.
§ Only male known; 4.7 mm. long; cordons 190u long; cloacal aperture 168u
from posterior extremity 202. toe ieser ft Acuaria papillifera, p. 224.
Both sexes known; male 10 to 11 mm. long; cordons of male 280 to 320
long; cloacal aperture 800 to 4004 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; Filaria anthuris (Rudolphi, 1819)
Schneider, 1866.
Hosts —Primary: Corvus glandarius, C. cornix, 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 (1878); later Linstow (1901), in
describing his Dispharagus invaginatus, writes that the cordons
anastomose and are recurrent as in A. anthuris; Railliet, Henry and
Sisoff define A cuaria 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, 18735 he says 22 and 18 mm.,
NEMATODE PARASITES OF BIRDS 219
but his illustration and magnification show this an error) to 237
and 234n long (Linstow, 1895), the right 34 and the left 31 wide,
according to Linstow, or both 34 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, 92.64 by 26.24 (Linstow, 1873), or 39» by
26 (Linstow, 1895).
Fics. 281—-282.—AcCUARIA ANTHURIS. 281, MALE TAIL. @, CROSS SECTION; 0, VENTRAL
VIEW. AFTER SCHNEIDER, 1866. 282, a, MALE TAIL X ABOUT 50; Db, FEMALE TAIL
END. AFTER LINSTOW, 1873
Life history—Unknown; Linstow (1895) found an encapsulated
larva in Gammarus 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 Dispharagus; a well described and commonly known
type species aids in the definition of a genus and in the efforts to
Fic. 283.—ACUARIA ANTHURIS. OVEJECTOR. AFTER SpuURAT, 1915
promptly assign the species to a valid genus and retain it there.
Even yet, A. anthuris 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, Coracias 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 1s a synonym in part of Owyspirura sygmoidea
Diesing, 1851.
ACUARIA ATTENUATA (Rudolphi, 1819) Railliet, Henry, and Sisoff, 1912
Synonyms.—S piroptera attenuata Rudolphi, 1819; Dispharaqus
attenuatus (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 wide. Mouth with 2 lateral papilliform lips. Cordons
thick and sinuous. Buccal cavity narrow, 170» long. Anterior esoph-
agus 540u 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 160u from tail end. Eggs only 15y long,
according to Dujardin, who questions whether they are fully
developed.
Life history—Unknown; probably involves intermediate stages
in other hosts.
istribution.—Europe (Austria (Vienna Museum) ).
ACUARIA CORDATA (Mueller, 1897) Gendre, 1920
Synonyms.—Spiroptera anthuris Rudolphi, 1819 of Dujardin,
1845, and of Molin, 1860; Dispharagus cordatus Mueller, 1897.
Hosts —Primary: Lanius rufus, L. curullio; secondary : Unknown.
Mueller regards his 2). cordatus as identical with S. anthuris of
Dujardin; the hosts for the latter are Corvus glandarius, C. pica, C.
frugilegus, C. corax, C. corone, C. cornix, Coracias garrula, Oriolus
galbula, Pyrrhocorax alpinus. Other hosts listed are Corvus caryo-
catactes, C. corax tingitanus, C. 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 Ze
species is established on the material from Lanius rufus and L.
curullio.
Location.—In stomach wall (proventriculus?).
Morphology—Acuaria sensu stricto (p. 216): Head (fig. 284a)
with 2 lips, cordons joined for a distance of 20u 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 400p. 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, 270u long by 50p wide,
Fic. 284.—ACUARIA CORDATA. @, HEAD END; b, LATERAL AND C, VENTRAL VIEW
OF MALE TAIL. AFTER MUELLER, 1897
the other more curved, 2104 long, and slenderer; according to
Mueller, the right spicule is 150 to 170u long and the left 170 to
183 long by 17p wide.
Female 22.5 to 40 mm. long by 200 to 320% wide. Cordons 480 to
600pn long. Anus 270 to 400u 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, 1866; Dispharagus de-
pressus (Schneider, 1866) Gendre, 1912.
Hosts—Primary: Corvus cornix; secondary: Unknown.
222 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Location —Not given; presumably in proventriculus, gizzard or
esophagus.
Morphology.—Acuaria sensu stricto (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).
ies. 285-286.—285, ACUARIA DEPRESSA. Mate TAIL. AFTER SCHNEIDHR, 1866.
256, ACUARIA GRACILIS. @ AND b, ANTERIOR END; C, LEFT AND d, RIGHT SPICULE;
€, MALE TAIL; f, FEMALE TAIL. AFTHR GENDRE, 1912
ACUARIA GRACILIS (Gendre, 1912) Railliet, Henry, and Sisoff, 1912
Synonyms.—Dispharagus gracilis Gendre, 1912; Acuaria similis
Gendre, 1913c.
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 6) 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 380, 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 180% wide. Tail 1/39 to 1/42
of total body length. Buccal cavity or pharynx 150 to 170» long;
anterior esophagus 350 to 420» long; posterior esophagus T70p 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 170u wide. Tail 1/90 to
1/103 of total body length (fig. 2867). Buccal cavity or pharynx 160
to 200. long; anterior esophagus 530 to 660, long; posterior esopha-
:
£
Fic. 287.—ACUARIA ORNATA. d, MALE TAIL; 0b, LEFT SPICULE; ¢C, HEAD END; d,
RIGHT SPICULE; C, CORDONS OF THE LATERAL AND f, OF THE SUBMEDIAN FIELDS.
AFTER GENDRE, 1912
oy
gus 1 to 1.32 mm. long. Vulva near middle of body, about 10/21 of
body length from head. Eggs 32 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) Railliet, Henry, and Sisoff, 1912
Synonyms.—Dis pharagus ornatus Gendre, 1912.
Hosts——Primary: @orvus scapulatus; secondary: Unknown.
Location.—Between the tunics of the gizzard.
Morphology—Acuaria 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. 2877) 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 360p wide. Tail 10/204 to
10/228 of total body length. Buccal cavity or pharynx 220 to 260p
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 ).
<E——_ nn
ka SS nh
Pa
SSS
Sete
Fig. 288.—ACUARIA PAPILLIFERA. HEAD AND TAIL ENDS OF MALB. AFYER 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.
Morphology.—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 70p 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.
HTosts—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 6) 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 ¢) 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
Fic. 289.—ACUARIA SUBULA. @ AND Bb, HEAD END; C, MALE TAIL; d, RIGHT AND @€,
LEFT SPICULE. AFTER GENDRE, 1913
postanal papillae. Spicules (fig. 289 d and e) dissimilar, 190» and
120 long, respectively.
Female 18 mm. long by 290 wide. Head 51p wide. Buccal cavity
or pharynx 170n long. Tail straight, conical, 2254 long. Vulva
10 mm. from tail end and 8 mm. from head end. Eggs 37 to 39p
long.
Life history—Unknown; probably involves intermediate stages in
other hosts.
LDistribution.—Europe (France).
ACUARIA TENUIS (Dujardin, 1845) Railliet, Henry, and Sisoff, 1912
Synonyms.—Dispharagus 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 944 wide. Head 20u wide. Buccal cavity
or pharynx 140p long; anterior esophagus 250p long by 20 wide;
posterior esophagus 750y long by 44 wide. Spicules 133 and 95p
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 160n wide. Tail 280 long. Eggs 34p
long.
We history—Unknown; probably involves intermediate stages in
other hosts.
Distribution—KEurope (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. Jfa/e 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.
Ty pe-species—Chetlospirura hamulosa (Diesing, 1851) Diesing,
1861.
KEY TO SPECIES OF CHEILOSPIBURA
1. Male 7.9 mm. long, female 8.5 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
middlesof bod ys ek Meee te 2 TN EP) Re NST US eee A De
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 1924 and 700 to 720n long; eggs 39 to 43 by 25 to 27p.
Cheilospirura spinosa, p. 229.
Cordons not spiny, extending to posterior region of first part of esophagus;
spicules 170u and 560u long; eggs 33u by 22u__ 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» long, the
other 125 to 148, long.
CHEILOSPIRURA HAMULOSA (Diesing, 1851) Diesing, 1861
Synonyms.—Spiroptera hamulosa (Diesing, 1851); Dispharagus
hamulosus (Diesing, 1851) Stossich, 1890; Spiroptera perforans Cen-
toscudi, 1911; Acwaria hamulosa (Diesing, 1851) Railliet, 1911.
Hosts—Primary: Gallus gallus, Meleagris gallopavo, Phasianus
gallus; secondary: Unknown, probably arthropods, perhaps insects.
The turkey is reported as a host by Ransom and Hall.
NEMATODE PARASITES OF BIRDS 277
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 14mm. 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
Fic. 290.—CHEILOSPIRURA HAMULOSA, a, CORDON; b, HEAD, LATERAL VIEW; ¢C, MALB
TAIL. AFTER DRASCHE, 1883
papillae (fig. 290¢), 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 25mm. long. Posterior end pointed. Vulva slightly
posterior to middle of body. Eggs 30u long by 20 wide, according
to Centoscudi.
Life history—Unknown; probably somewhat similar to that of
Dispharynx 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, 1928.
Hosts—Primary: Caccabis rufa and Francolinus bicalcaratus;
secondary: Unknown.
Location.—Between tunics of gizzard.
228 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Morphology.—C heilospirura (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. 291g) 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
Hin
| os
MOUTHS
ia
Ch
ann
Ti
WIN)
Lf
_———
€.
Fic. 291.—CHEILOSPIRURA GRUVELI. d@, ANTERIOR END; 0, HEAD; C, CORDON ;
d, FEMALE TAIL, AFTER GENDRE, 1912. e, RIGHT AND f, LEFL SPICULE; g,
MALE TAIL. AFTER GENDRE, 1913
and 7) very dissimilar and unequal, the left 560, and the right
170» long.
Female 28.9 to 42.74 mm. long by 220 to 270u wide. Cordons
1.15 mm. long. Tail (fig. 291d) 1/128 to 1/144 of total body length.
Pharynx or buccal cavity 180 to 280n long; anterior esophagus
990n 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, 334 long by 224 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
Synonyms.—Dispharaqus 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 320.2 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 3202 wide. Esophagus 10/27 of body
length. Tail (fig. 292) 1/25 of body length. Four pairs of preanal
————
OOSTTV7
Figs. 292—293.—292, CHWILOSPIRURA ROTUNDATA. MALE TAIL. AFTER LINSTOW, 1907.
293, CHEILOSPIRURA SPINOSA. @, ANTERIOR END; Db, PART OF CORDON; C, VULVA;
d, FEMALE TAIL; €, SPICULE. ORIGINAL
papillae and 5 pairs of postanal papillae. Spicules dissimilar, one
750» 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, 34p
by 21p.
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution —Asia (Turkestan).
CHEILOSPIRURA SPINOSA, new species
Synonym.—A cuaria (Cheilospirura) species Stafseth and Kotlan,
1925.
Hosts—Primary: Bonasa wmbellus; secondary: Unknown.
Location.—Gizzard.
Morphology—C heilospirura (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
Kotlin) by 183, wide at the level of the posterior end of the esopha-
gus and 2324 wide in the middle of the body. Cordons 495 long.
Pharynx or buccal cavity 232» long; anterior esophagus 1.1 mm.
long; posterior esophagus 2.6 mm. long. Caudal extremity tightly
Fic. 294.—CHEILOSPIRURA SPINOSA. 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 7202 long (660 according to Stafseth and
Kotian), the other thick, 192 long (fig. 293e).
Female 34 to 38 mm. long (37 to 40 mm. in specimens of Stafseth
and Kotlan) by 315» wide at posterior end of esophagus and 348y
wide in middle part of body. Cordons 797 to 813» long. Pharynx
or buccal cavity 232 long; anterior esophagus 1.4 mm. long; pos-
terior esophagus 3.4 mm. long. Vulva (fig. 293¢) anterior to middle
of body, dividing body length in ratio of 17:23. Anus 250 to 300u
from posterior end. Tail (fig. 293d) slender and digitate. Eggs 39
to 42u by 25 to 27p.
NEMATODE PARASITES OF BIRDS 231
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution.—North America (United States (Michigan) ).
Type material.—N 0. 25538, U.S.N.M. (Bureau of Animal Industry
helminthological collection). Collected from a gizzard sent to the
laboratory by Dr. George R. La Rue, the gizzard having been sent
to him by Dr. Jan Metzeloas of the University of Michigan, Ann
Arbor, Michigan.
The description by Stafseth and Kotlin of a new species of
Cheilospirura 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 (.
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. gruwveli they
extend almost to the posterior end of the first part of the esophagus,
in (. 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.—sS piroptera Rudolphi, 1819, 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; they 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 part, 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.
Ty pe-species.—Chevreuxia revoluta (Rudolphi, 1819) Seurat,
1918c.
CHEVREUXIA REVOLUTA (Rudolphi, 1819) Seurat, 1918e
Synonyms.—Spiroptera revoluta Rudolphi, 1819; Dispharagus
revolutus (Rudolphi, 1819) Molin, 1860.
Hosts—Primary: Charadrius himantopus, Himantopus himan-
topus, H. melanopterus; secondary: Unknown.
Location. Between tunics of the gizzard.
Morphology.—C hevreuwia (p. 231) : Characters of the genus.
‘SJ
=e Ts
Co
: AOE Sa enn i
i
MIM Wm,
“SSE AZ
SERVLET 77
aD ; —— ane ata Sy
LT Ce SS
Fig. 295.—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,
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, long, wide and with
obtuse end; left spicule 750 long, slender, filiform and pointed.
Female 13.2 to 18.3 mm. long by 265) 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 Zao
border of collar 765. 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 ispharynw and Synhimantus; sphincter rectilinear ;
unpaired varnish gland or trompe 350» long. Eggs 32u by 18n,
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; Spzroptera Rudolphi,
1819, part; Dispharagus Dujardin, 1845, part; Hzstiocephalus
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.
Ty pe-species.—Cosmocephalus diesingi Molin, 1858, emend.
Cram, 1927.
KBY TO SPECIBS 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 button-like or
SUCKEr-IIKeY APDCHOAS Ott sere Alek dre Ge FN ta ee Zs
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% and 420 long.
Cosmocephalus obvelatus, p. 285.
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 1804 and 700 long.
Cosmocephalus aduncus, p. 254.
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 6) 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.
Ksephagus 1/3 as long as body and half as wide as body.
Male unknown.
Female 15 mm. long by 300y 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) ).
Fic. 296.—COSMOCEPHALUS DIESINGI. @, ANTERIOR AND MIDDLE PARTS OF WORM,
SHOWING POSITION OF VULVA; JD, ANTERIOR PND} c, FEMALE TAIL. AFTER MOLIN,
1861
COSMOCEPHALUS ADUNCUS (Creplin, 1846) Yorke and Maplestone, 1926
Synonyms.—s piroptera adunca Creplin, 1846; Dispharagus adun-
cus (Creplin, 1846) Molin, 1860c.
Hosts——Primary: Colymbus septentrionalis, Larus argentatus, L.
canus, L. glaucus, L. marinus, L. medius, L. ridibundus, Podiceps
auritus; secondary : Unknown.
Location.—Esophagus.
Morphology.—C osmocephalus. (p. 233): Mouth (fig. 297a) with
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 3800p wide. Caudal extremity
(fig. 297 6) obtuse. Caudal alae wide. Nine pairs of long, pedun-
culated papillae, of which 4 are preanal and 5 postanal. Spicules
180p. 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 34:39.
Eggs 39u by 23n.
Life history—-Unknown; probably involves intermediate stages in
other hosts.
Distribution.—EKurope.
Linstow reported this from Larus canus 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.—s piroptera obvelata Creplin, 1825; Filaria obvelata
(Creplin, 1825) Linstow, 1877); Dispharagus obvelatus (Creplin.
1825) Linstow, 1909a,; Histiocephalus spiralis Diesing, 1851; Cosmo-
cephalus papillosus Molin, 1859; Dispharagus papillosus (Molin,
1859) Stossich, 1898; Cosmocephalus alatus Molin, 1860d.
tii,
2b.
Fic. 297.—CoOSMOCEPHALUS ADUNCUS. 4d, HEAD PND; 0b, MALE TAIL. AFTER
STOSSICH, 1892
Hosts—Primary: Actitis hypoleucus, A. macularia, Alca torda,
Catorrhactes pachyrhynchus (new record), Larus argentatus, L.
argentoides, L. canus, L. fuscus, L. marinus, L. maximus, L. medius,
L. ridibundus, Mergus serrator, Puffinus kuhli, Sterna arctica, 8.
risoria, Totanus hypoleucus, T. maculatus, T. fuscus, Tringoides
hy poleucus, Uria grylle; secondary: Unknown.
Location.—Esophagus.
Morphology.—C osmocephalus (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; the 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 2554 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 260n 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,
igs. 298—299.—CoSMOCEPHALUS OBVELATUS. 298, a, DORSO-VENTRAL VIEW
AND b, LATERAL VIBW OF AN ANTERIOR END; C, FEMALE TAIL, VENTRAL VIEW.
AFTER SHuURAT, 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 380u% 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 364 by 202, embryonated when oviposited.
According to Linstow, in a specimen 9.77 mm. long the cordons were
NEMATODE PARASITES OF BIRDS 237
440 long, their anastomosis 110. from the head end, and the eggs
26 by 21u. 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 —Dispharynxe nasuta (Rudolphi, 1819) Stiles and
Hassall, 1920.
KEY TO SPECIES OF DISPHARYNX
1, Inadequately described ; from Falco minutus or Rhamphastos vitellinus__ 2.
Adequately described; from hosts other than above__-_---_------_----~-- 3.
“. Lips very small; cordons short; from Falco minutus.
Dispharynx capitata, p. 240.
Lips large; cordons long; from Rhamphastos vitellinus.
Dispharynx crassissima, p. 243.
3. Male with 7 pairs of postanal papillae; female 13 to 16 mm. Jong.
Dispharynx magnilabiata, p. 241.
Males with less than 7 pairs of postanal papillae; female not over 11 mm.
VOWS; Fearon. ase tar sR AS ott ee Eee re Sd ea ee 4,
4. Male with 4 pairs of postanal papillae; longer spicule twice the length of the
SNOUTCTHONC 2.2 2 = he es Dispharynx rectovaginata, p. 248.
Male with 5 pairs of postanal papillae; longer spicule either considerably
more or considerably less than twice the length of the shorter one___~ 5.
5. Vulva anterior to middle of body___-_--__-_--__ Dispharynx nasuta, p. 237.
Vulvar posterior: tovmiddle,,of bodyi. 1b lt ce ee he et en ce 6.
G. Postcervical papillae situated posterior to cordons; vulva just posterior to
middle of body; longer spicule 2604 long______ Dispharynx noctuae, p. 242.
Postcervical papillae situated between the cordons; vulva in posterior third
of body; longer spicule 4004 long or longer____--________----__-----~-- %.
7. Postcervical papillae bicuspid; tail of male about 2754, of female about
120u long; spicules 150u and 400u long__-__-__ Dispharynx spiralis, p. 238.
Posteervical papillae tricuspid; tail of male 3704, of female 1704 long;
spicules 240, and 865, long--_------------- Dispharynx laplantei, p. 241.
See Addenda, p. 391, for a species of Dispharynx recently described
hy Smit and Notosoediro.
DISPHARYNX NASUTA (Rudolphi, 1819) Stiles and Hassall, 1920
Synonyms.—Spiroptera nasuta Rudolphi, 1819; Lispharagus
nasutus (Rudolphi, 1819) Dujardin, 1845; Acuaria (Dispharyna)
238 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
nasuta Railliet, Henry and Sisoff, 1912; Cheilospirura nasuta
(Rudolphi, 1819) Ransom, 1916.
Hosts—Primary: Gallus gallus; secondary: Unknown, but prob-
ably arthropods. Porcellio laevis reported as intermediate host by
Piana; apparently he was dealing with Dispharyna spiralis.
Location.—Proventriculus and gizzard and connective tissue
between these.
Morphology—Dispharynz (p. 237): Two small lateral lips (fig.
3002). 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. 3000) 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.
Fic. 300.—DISPHARYNX NASUTA. HEAD AND TAIL OF MALE. AFTER DUJARDIN, 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, Porcellio laevis, but later writers
believe he was dealing with DP. 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 yarious localities are believed by Railliet to be records of
D. spiralis (pp. 238). Both species need reconsideration before one
may undertake to give their distribution with any certainty.
DISPHARYNX SPIRALIS (Molin, 1858) Skrjabin, 1916b
Synonyms.—Dispharagus spiralis Molin, 1858; Dispharagus nasu-
tus of Piana, 1897; Dispharagus spiralis columbae Bridré, 1910;
Acuaria spiralis (Molin, 1858) Railliet, Henry, and Sisoff, 1912.
NEMATODE PARASITES OF BIRDS 239
Hosts.—Primary : Gallus gallus, Bonasa umbellus, Caccabis petrosa,
Columbia livia, Meleagris gallopavo, Phasianus gallus, Numida me-
leagris, Quiscalus quiscula,; secondary : sowbug (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—VDispharynx (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.
Fic. 301.—DISPHARYNX SPIRALIS. MALE. AFTER PIANA, 1897
Male 7 to 8.3 mm. long by 230 to 3154 wide (fig. 301). Cordons
415 to 515y long. Five pairs of postanal and 4 pairs of preanal
papillae. Cloacal aperture about 275, from tail end. Long spicule
400 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
900p 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. 83030) 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 (trompe) unpaired, 250p long. Eggs 36
to 40p by 21h, embryonated when oviposited.
Life history—According to Piana, the life history of what he
called Dispharagus nasutus, but which was apparently Dispharynx
3612—27T——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.
Fies. 802-203.—DISPHARYNX SPIRALIS. 302, FEMALE. AFTER PiANA, 1897. 303,
a, HmAD END. AFTER SeEuURAT, 1916. 6, OvesEcToR. AFTER SEURAT, 1919
DISPHARYNX CAPITATA (Molin, 1860) Cram, 1927
Synonyms.—Dispharagus capitatus Molin, 1860; Spiroptera alata *
in Molin, 1860; Acuaria capitata (Molin, 1860) Railliet, Henry, and
Sisoff, 1912.
Hosts.—Primary : Falco minutus; secondary : Unknown.
Location.—Proventriculus.
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.—Acuarta (Dispharynx) laplantet Seurat, 1919.
Hosts.—Primary: Garrulus glandarius cervicalis; secondary: Un-
known.
Location.—Proventriculus.
Morphology.—Dispharynxz (p. 237): Body large, attenuated at ex-
tremities. Cordons similar to those of )). noctuae and DP. spiralis.
Postcervical papillae tricuspid, subsymmetrical; Seurat says they are
situated as in ). spiralis, directly in front of the level of the excre-
tory pore, but according to his figures the excretory pore is 590 and
390 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 3002; much
slenderer than female. Cordons 4554 long. Caudal extremity
coiled in spiral. Cloacal aperture 3704 from tail end. Four pairs
of preanal and 5 pairs of postanal papillae, arranged as in J). noctuae
and PD. 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 5654 wide. Cordons 8054 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 25p, 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.—Dispharagus magnilabiatus Molin, 1860c; Spiroptera
plataleae-ajajae * 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 the Vienna Museum.
QAI BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 7.5 mm. long by 200n wide. ‘Tail (fig. 805) 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 3800p 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 Chetlospirura, to which it had been assigned by Railliet, Henry,
and Sisoff.
Fics. 304—306.—304, DISPHARYNX LAPLANTEI OVEJECTOR. AFTER SEURAT,
1919. 305, DISPHARYNX MAGNILABIATA. MALE TAIL. AFTER DRASCHE,
1884. 3806, DISPHARYNX NOCTUAE. a, VULVA; b, ANTERIOR END, DORSAL
VIEW; C, FHMALE; d, ANTERIOR END, LATERAL VIEW} €, 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, 19138; Acuaria (Dispharyne)
noctuae Seurat, 1913a.
Hosts—Primary: Carine noctua glaua; secondary: Unknown.
Location.— Esophagus.
Morphology —Dispharynx (p. 237) : Mouth (fig. 3060 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 200u long. Caudal
alae very large. Four pairs of preanal and 5 pairs of postanal
papillae. Spicules 260» and 115m long.
Female 10 to 11 mm. long by 350» wide at the level of the vulva.
Buccal cavity 280 to 300u long. Tail 170» long. Vulva (fig. 306a
and e) opening on an oval prominence directly behind the middle
of the body. Eggs 484 by 21p, 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.
fHosts—Primary: Falco ater; secondary: Unknown.
Location.—Proventriculus.
Fic. 307.—DISPHARYNX RECTOVAGINATA. H®PAD END AND TAIL OF MALE. AFTER
DRASCHE, 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. 807), sinuous and recurrent, not anastomosing.
Male 4 mm. long by 200n 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, 1860) 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: Rhamphastos vitellinus; secondary: Unknown.
Location.—Proventriculus.
Morphology.—Dispharynax (p. 237): Head continuous with body.
Mouth with 2 large papilliform lps. 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; Dispharyna Railliet,
Henry, and Sisoff, 1912, part; Zamannia 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 (#. decorata).
Vulva usually in posterior part of body. Parasitic usually in pro-
ventriculus and gizzard, sometimes in tumors in gizzard, in birds.
Type species —H'chinuria 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
teovanterior portion. of (hodyiG 2. 2 UO Ce ee ee 2.
. Cordons claborate, their inner part made up of transverse bands, their
outer edge with posteriorly directed scales or teeth; esophagus one-
fourth of. total: body leneth#or, longer. 2.8 0s.) ee ee 3.
Cordons not as described above; esophagus length, if given, much shorter
Bhan: Dover kas se Le 2a 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 PARASITES 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. Body with 2 double rows of spines extending almost entire length of
ITCRA eee are ee oe ee ne eR EE a Be 6.
Body 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 euticular
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
PAR ASCOMIOS ES coe ree te ea) 2 A os ath re ee 8 EE de 3 ae ot Se dc
7. Male 9 mm. long; spicules 706, and 208 long, respectively.
Echinuria uncinata, p. 246.
Male 11 to 12 mm. long; spicules 8394 and 140u 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-
SDICUOUS IIS 2— = = 5 ee ee Echinuria contorta, p. 249.
Only female described; 8 to 10 mm. long; mouth with 2 large conspicuous
PLOJECEING IPSet: * Spe a ea Echinuria calcarata, p. 249.
10. Description incomplete; from Ciconia maguari.
Echinuria longeornata, p. 255.
on
Hrom? hostsgother than above.22 4. et a Eli
11. Left 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 180u, left spicule
900, long; eggs’ 32u Dy. 20eo22 = Echinuria leptoptili, p. 254.
Male 11 to 11.5 mm. long; female 13 to 15 mm. long; right spicule 190 to
210u, left spicule 650 to 675yu, long; eggs 284 by 18x.
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. 2085) each 57u
long by 32» 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 6 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
163» long by 47» wide.
Male 11 to 12 mm. long by 600p wide. Cordons 769» long by 14y
wide. Tail (fig. 308¢) curved ventrally. Spicules unequal, one 839.
long by 21p wide, and curved, the other 140. long by 33» wide, or
wider.
Female 15 to 16 mm. long by 800n wide. Cordons 792» long by
23 wide. Tail pointed. Eggs 35.4p by 21.84, with shells 3p thick.
246 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history Unknown; probably involves intermediate stages in
other hosts (see 4’. uncinata, p. 246).
Distribution—Asia (Russian Turkestan).
There is nothing in the description, given up to this time, of F.
jugadornata and E. wncinata 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, Heap
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 /. wncinata before placing
it as he did in his new genus with /. jugadornata, the present writer
accepts £’. 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 /. wncinata.
Since writing the above, the present writer has found specimens
agreeing with the description of 2. uncinata and finds that the lengths
of the spicules of the male differ somewhat from those given for £.
jugadornata.
ECHINURIA UNCINATA (Rudolphi, 1819) Soloviev, 1912
Synonyms.—Spiroptera uncinata Rudolphi, 1819; Dispharagus
uncinatus (Rudolphi, 1819) Railliet, 1893; Aeuaria (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 domesticus, Cygnus olor domesticus, Nettion
carolinense; secondary: Small crustaceans, (Daphnia pulew).
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. 244): Mouth with 2 lips and 6
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
iongitudinal 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
EI
ES
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=]
f—
Fy
=
>
RS
=>
Ss
S
eS
Fics, 309-310.—EcHINURIA UNCINATA. 309, HEAD END; VENTRAL
View. AFTER SEURAT, 1919. 310, a, HEAD END, DORSAL VIEW; D,
CROSS SECTION, AND C, VENTRAL VIEW OF MALE TAIL. AFTER
SCHNDIDER, 1866.
lines where they are joined a short distance beyond the excretory
pore, the change of course being especially noticeable in the latero-
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 9to10mm.long. Tail (fig. 3106 and ¢) with straight caudal
alae, somewhat vesicular. Schneider states that there are 4 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 Vettion carolinense and Anas rubripes, and is able to add
the following particulars: Male 8 to 10 mm. long by 300p to 500p
wide. In a male 8 mm. long, cordons 5002 long by 20 wide;
pharynx 133, esophagus 830,, long; cloacal aperture 330p from pos-
terior end; spicules 706 and 208, long, respectively.
Female 12 to 18.5 mm. long. In a female 12 mm. long by 515z
wide, the pharynx is 150 long; cordons 748 long by 25~ wide; vulva
1.3 mm., anus 250u, from posterior end; eggs 37% by 20n. Vulva 1
to 1.4 mm. from tip of tail. Ovejector of type of Dispharyna
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 pulew. 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 America (United States).
See the discussion of this species in connection with £. 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.—E chinuria (p. 244): Body slightly attenuated an-
teriorly. Cuticula rather coarsely striated transversely. Mouth
(fig. 311 @ and 6) 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
apastomose with the corresponding cordons from the opposite side.
Male unknown.
Female 17 mm. long by 700% wide. Head 150x wide at base of
lips. Anterior esophagus 800 long by 50 to 904 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. 311¢). Eggs not found.
Life history—Unknown; probably involves intermediate stages
in other hosts (see #’. 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: [bis guarauna,; secondary: Unknown.
Location.—Proventriculus.
Morphology —Echinuria (p. 244): Characters of the genus.
Male unknown.
\ sr
i
cit Theil MAE |
Wel
wo lity
Fig. 311.—EcHINURIA ARDEAE. @, FRONT VIEW AND 0, LATERAL VIEW OF HEAD; C,
FEMALE TAIL, AFTER SMITH, FOX, AND WHITE, 1908
Female 8 to 10 mm. long by 2004 wide. Mouth with 2 large con-
spicuous projecting lips (fig. 312 @ and 6). Cuticula transversely
striated. Anterior extremity of body appreciably attenuated. Tour
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 /’, wncinata, p. 246).
Distribution.—South America (Brazil).
This species may be identical with /. 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; Hamannia contorta (Molin, 1858) Stiles and Hassall, 1920.
i]
|
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
250
Spiroptera falcinelli Rudolphi, 1819, is a nomen nudum and hence
not available.
Hosts —Primary: [bis faleinellus, Falcinellus tgneus; secondary:
Unknown.
Location —In muscular ventriculus (gizzard).
Morphology.—chinuria (p. 244): Mouth with 2 small incon-
Cuticula with dense transverse striations which are
spicuous lips.
twisted irregularly. Four cordons (fig. 313a), anastomosing.
Male 7 to 8 mm. long by 200n wide. Tail (fig. 3130) curved.
Caudal alae wide, semilunar, and transversely striated. Molin states
that there are 7 pairs of caudal papillae, but only figures 3 pairs
cf postanal papillae.
ing
i
i
:
f
(I)
i
él
NUH
ad, LATERAL AND Db, VENTRAL VIEW OF
313, ECHINURIA CONTORTA. 4,
AFTER
Figs. 312-313.—312, ECHINURIA CALCARATA.
FEMALE TAIL. AFTER DRASCHE, 1884.
ANTERIOR BEND; b, MALE TAIL; C, FEMALB TAIL, SHOWING VULVA AND ANUS.
HPAD; ¢,
Caudal extremity (fig. 313c¢)
MOo.Lin, 1861
Female 19 mm. long by 40u wide.
with short, obtusely conical lateral appendages. Anus said to be at
raudal apex. Vulva somewhat anterior to anus.
Life history—Unknown; probably involves intermediate stages
in other hosts (see ’. wncinata, p. 246).
Distribution—Kurope (Italy (Padua) and Austria (Vienna) ).
ECHINURIA DECGRATA, 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 6)
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 7142 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 207
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 8304 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. 314¢) 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. 314d) large and projecting; diameter of body decreases
suddenly below 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 (fig.
514e); a narrow sphincter 664» long by 41.54 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 posteriorly and soon joins the ovary which has a con-
voluted course. Eggs 36n by 21p.
252 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history—Unknown; probably involves intermediate stages
in other hosts (see 2. wncinata, 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 /’. uncinata as described by Seurat. The lat-
ter species has a short ovejector of the type of Dispharynx with a bi-
Ff S
i 2
FS % LA»
SS BA
— ZZZx
—— ZY
ane
Fic. 314.—EcCHINURIA DECORATA. @, ANTERIOR END; b, CORDON; €, FEMALB TAIL; d,
VULVA AND VESTIBULE; €, VULVA, OVEJECTOR, AND UTERUS; f, MALE TAIL; g AND h,
SPICULES. ORIGINAL
partite varnish gland (trompe) which connects with the 2 uteri. As
a transition form between the didelphic /. wncinata and the monodel-
phic /’. 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 #. 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 #. decorata to EF. hargilae,
whereas the length of the esophagus and the elaborate nature of the
cordons relate it to /. sguamata.
ECHINURIA HARGILAE (Baylis and Daubney, 1923) Cram, 1927
Synonyms.—Acuaria (Echinuria) leptoptili (Gedoelst, 1916) of
Baylis and Daubney, 1922; Acuaria (E'chinuria) hargilae Baylis and
Daubney, 1923.
Hosts—Primary: Leptoptilus dubius; secondary: Unkown.
Location.—Not given.
Morphology— chinuria (p. 244): Cuticula finely striated trans-
versely. Cordons 20n wide, extending posteriorly for 950» to 1.1
mm., with a shght 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
Ringe
<a,
Fig. 315.—ECHINURIA HARGILAP. d, ANTERIOR END; 0, RIGHT SPICULE. AFTER
BAYLIS AND DAUBNEY, 1922
OM
115p 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. 315d), 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 170u 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 50y from tailend. Eggs
28 by 18z.
Life history—Unknown; probably involves intermediate stages
in other hosts (see /’. wncinata, p. 246).
Distribution—Asia (India (Calcutta) ).
ECHINURIA HORRIDA (Rudolphi, 1809) Cram, 1927
Synonyms.—Strongylus horridus Rudolphi, 1809; Spzroptera gal-
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; élaria aculeata (Creplin, 1825) Linstow, 1876;
Dispharagus aculeatus (Creplin, 1825) Stossich, 1891; L'chinuria
spinifera (Schneider, 1866) Soloviev, 1912.
Hosts.—Primary : Scolopax gallinula, Charadrius hiaticula, Tringa
variabilis, T. alpina; secondary: Unknown.
Location.—Proventriculus.
Morphology —E chinuria (p. 244): Mouth with 2 lips, dorso-ven-
tral, according to Stossich, but probably lateral. From these there
originate on each side an are 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-
‘viorly, 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 42u by 23y elliptical and thick-shelled.
Life history—Unknown; probably involves intermediate stages in
other host (see 2’. wncinata, p. 246).
Distribution.—Kurope (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.—Acuaria (Echinuria) leptoptili (Gedoelst, 1916) Bay-
lis and Daubney, 1928, not of Baylis and Daubney, 1922 (see Z.
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 simp!e bands with
margins scalloped, or, according to Gendre, denticulate as in Z.
squamata, 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 2602 wide. Tail (fig. 318 6) about 160
long, about 1/40 of total body length. Cordons 610» 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 ¢ and d) unequa!
and dissimilar; the left 900n 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
970u long. Gedoelst describes a membranous expansion of the cuti-
a
A
%
9999 Gg VIF IS
D
a
2
©
*°
oO
~o
A
a
4
Figs. 316-318.—EcHINURIA HORRIDA. 316, HEAD END. ArreR SCHNEIDER, 1866. 317,
HEAD PND. AFTER STOSSICH, 1891. 318, ECHINURIA LEPTOPTILI. @, HEAD END;
b, MALE TAIL; Cc, LEFT SPICULE; d, RIGHT SPICULE (THE SMALL DRAWING SHOWING
ORIFICE AT FREE END DILATED ON PRESSURE) ; €, FEMALE TAIL. AFTER GENDRE, 1926
cula along the lateral lines of the caudal region, extending from a
point 50u 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 95n, according to Gedoelst, or 50u, according
to Gendre (fig. 318 e), from tail end. Vulva 80 to 145y, according
to Gedoelst, or 210p, according to Gendre, anterior to anus. Eggs
32p. by 20p.
Life history.—Unknown; probably involves intermediate stages in
other hosts; (see /’. uncinata, p. 246).
Distribution.—Africa (Belgian Congo and Dahomey).
ECHINURIA LONGEORNATA (Molin, 1860) Cram, 1927
Synonyms.—Dispharagus longeornatus Molin, 1860c; Spiroptera
ardeae-maguari® Molin, 1860; Acuaria longeornata (Molin, 1860)
® Catalogue of Vienna Museum.
256 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Railliet, Henry, and Sisoff, 1912; Hamannia longeornata (Molin,
1860) Stiles and Hassall, 1920.
Hosts——Primary: Ciconia maguari; secondary: Unknown.
Location.—Proventriculus.
Morphology.—Echinuria (p. 244) : Mouth with 2 small inconspicu-
ous lips. Cordons very long, briefly recurrent (brevi regredientes),
according to Molin; Gendre (1919) says this may be a retraction due
to fixation, a thing he has seen in /. 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 200u wide. Anus near caudal apex. Vulva
small, anterior to anus.
Life history—Unknown; probably involves intermediate stages in
other hosts; (see #’. wneinata, p. 246).
Distribution—South America (Brazil).
ECHINURIA LONGEVAGINATA (Molin, 1860) Cram, 1927
Synonyms.—Dispharagus longevaginatus Molin, 1860¢; Spirop-
tera ciconiae-maguari* 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. 319@). Cuticula transversely striated.
Anterior extremity markedly attenuated. Cordons long and
straight, anastomosing. According to Molin, the cordons are briefly
recurrent here as in F’. longeornata,; 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 200u 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 Z’. 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 (Hamannia) phoenicopteri Seurat, 1916;
Hamannia phoenicopteri (Seurat, 1916) Stiles and Hassall, 1920.
Hosts —Primary: Phoenicopterus roseus; secondary: Unknown.
Location—In mucosa of proventriculus.
Morphology—k chinuria (p. 244): Anterior extremity (fig. 320 a
and 6) 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, wide, as viewed through the
Fics. 319-320.—319, ECHINURIA LONGEVAGINATA, 4, HEAD; b, MALE TAIL. AFTER
DRASCHD, 1884. 3820, ECHINURIA PHOPNICOPTERI. IMMATURE FEMALP. @ AND
vb, HEAD END; c, TAIL; d, VULVA AND OVEJECTOR. AFTER SEURAT, 1916
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 ¢) 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; Dispharagus squa-
matus (Linstow, 1883) Stossich, 1891; Acuaria squamata (Linstow,
258 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
1883) Railliet, Henry, and Sisoff, 1912; Hamannia 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. 321a and 6) 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
Fic. 321.—EcCHINURIA SQUAMATA, d@, ANTERIOR END; 0, 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
ijateral lines.
Male unknown.
Female 24 mm. long by 720p wide. Eggs 39u by 24u, 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.
Generic 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 ure
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 ef total body length; lateral
alae-presentou/ = ee Rusguniella elongata, p. 259
RUSGUNIELLA ELONGATA (Rudolphi, 1819) Seurat, 1919a
Synonyms.—s piroptera elongata Rudolphi, 1819; Dispharagus
species Wed], 1856; Dispharaqgus elongatus (Rudolphi, 1819) Molin,
1860; Lilaria elongata (Rudolphi, 1819) Schneider, 1866; Acuaria
elongata (Rudolphi, 1819) Raillet, Henry, and Sisoff, 1912.
Hosts.—Primary: Hydrochelidon nigra, Sterna nigra, Podiceps
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 6) originate immediately
posterior to cordons (epaulets). Lateral or precervical papillae
asymmetrical; in a specimen 28.2 mm. long by 312, wide the left
papilla 182,, the right 1924, from head end; the origin of lateral alae
105p, excretory pore 456p, the vulva 13 mm. from head end. Buccal
cavity or pharynx 135, long, anterior esophagus 865, long, posterior
esophagus 3.1 mm. long, tail 2752 long. Vulva with slightly salient
lips. Cuticular ovejector tubular, 450. long, directed anteriorly.
Eggs oval, 38u by 24u, thick-shelled, embryonated when oviposited.
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution—Europe (Vienna Museum) and Africa (Algeria).
260 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
RUSGUNIELLA VANELLI (Rudolphi, 1819) Seurat, 1919a
Synonyms.—S piroptera vanelli Rudolphi, 1819; Acuaria vanelli
(Rudolphi, 1819) Poche, 1912.
Hosts.—Primary: Tringae 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 2004 wide. Esophagus 10/68, tail 1/34 of total
FIGs. 322-323.—322, RUSGUNIELLA ELONGATA. ANTERIOR END. @, LATERAL VIEW; Bb,
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.—S piroptera 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, 1860) Skrjabin,
19160.
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 Sctadiocara.
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
SUE OV ea aed a a oe Sciadiocara umbellifera, p. 261.
SCIADIOCARA UMBELLIFERA (Molin, 1860) Skrjabin, 1916b
Synonyms.—S piroptera umbellifera Molin, 1860); Spiroptera
tantali rubri® Molin, 1860; Spiroptera totani*® Molin, 1860; Schis-
torophus umbellifera (Molin, 1860) Railliet, 1916.
Hosts—Primary: lbis rubra, Totanus melanoleucus, Scolopax
major; secondary: Unknown.
Location.—Between tunics of the gizzard.
Morphology—sSciadiocara (p. 260): Mouth (fig. 824 @ and 6) 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% wide. Two caudal alae (fig.
325 a) with 6 pairs of preanal and 5 pairs of postanal papillae. The
most posterior pair of papillae are 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 8ly long with a maxi-
mum diameter of 224. The other spicule is 330» long by 7.5y. 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 1024 wide. Pharynx 37y. long.
The characteristic caudal extremity (fig. 825 c) consists of a caudal
appendage 85, long rounded in an obtuse manner and attached to
the body at an obtuse angle. Vulva (fig. 8255) slightly posterior
to the middle of body, 4.08 mm. from tail end. Eggs 44 to 48y long
by 30 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: Unknown.
Location.—Between tunics of gizzard.
Fics. 324-325.—ScraADIOCARA UMBELLIFERA. 324, Hap. a, DORSO-VENTRAL VIEW; 0,
FRONT VIEW. AFTER DRASCHE, 1884. 325, a, MALE TAIL; b, VULVA; C, FEMALE TAIL.
AFTER 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 Skrijabin, 1916d
Synonym.—fRictularia Froelich, 1802, part; Acuaria Bremser,
1811, part; Spzroptera Rudolphi, 1819, part; Gnathostoma Owen,
1836, part; Prionostemma Gendre, 19200.
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. Vulv:
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, 1916d.
KEY TO SPECIES OF SEURATIA
1. Female 13 to 14 mm. long; from Procellaria anglorum.
Seuratia procellariae, p. 263.
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, 1851
(based on Bellingham, 1844) ; Spiroptera procellariae Diesing, 1851;
Prionostemma procellariae (Diesing, 1851) Gendre, 1920d.
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 species, Acuaria pelagica (=S. shipley?),
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-
dowxa Linstow, 1903), Rictularia shipleyi (Stossich, 1900) Johnston,
1912; Acuaria pelagica Seurat, 1916a; Prionostemma pelagicum
(Seurat, 1916) Gendre, 19206; Prionostemma shipleyi (Stossich,
1900) Gendre, 19205.
Hosts.—Primary: Diomedea exulans, Daption capensis, Larus
canus, Puffinus kuhli; secondary: Unknown.
Location.—Proventriculus.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
264
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 @ and 0) swollen to form a collar
with 2 large disks, posteriorly bilobed, and the free margin with
Large tricuspid cervical papillae a little behind the
strong spines.
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 ¢) 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
ey
———__—_—-#
rn
oe
A ai |
8
(il
(|
t
Oth
COOTER
mo
Oh
=
a, DORSO-VPNTRAL AND b, LATERAL VIEW OF
; €, EGG. AFTER STOSSICH, 1900
Fig. 326.—SEuRATIA SHIPLEYI.
ANTERIOR END} C, MALE 'TAIL; @, FEMALE TAIL
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
500u wide. Tail 180u 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 Reailliet, Henry, and Sisoff, 1912
Generic diagnosis—Acuariinae (p. 211): Mouth with 2 conical
: projecting lips, originating from a depression, the margin of which
Esophagus short, swollen, muscular.
forms a denticulated collar.
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 6
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.
Ty pe-species.—Stre ptocara pectinifera (Neumann, 1900) Skrjabin,
19160,
Vic, 327.—SHURATIA SHIPLEYI. a AND b, PROXIMAL REGION OF OVEJECTOR } C, DISTAL
REGION OF OVEJECTOR. AFTER SEURAT, 1916
KEY TO SFECIES OF STREPTOCARA
2: GCerviealpapillae iwith: 3 points2222) 2s) fn ee 2:
Cervical papillae with 5 or more points______________________________--- 6.
2. Female not.over 10 mim. long. =--2.-— 255 ae
HemMale iG emm: OF WONSCr =~. = = ee 8 a es 5
3. Cervical papillae over 400u from anterior extremity; eggs 27 long.
Streptocara triaenucha, p. 271.
Cervical papillae not more than 1404 from anterior extremity; eggs 39u
NI reek AAs eke Oe ee VDE We has Pat ba 26d 8B te ih skh te) ot 4.
4. Vulva anterior to middle of body (1/2.6 of body length) ; 5 pairs of postanal
papillae; spicules 140u and 6004 long____-- Streptocara cirrohamata, p. 266,
Vulva in posterior region of body (500u from posterior end); 38 pairs of
postanal papillae; spicules 68u and 237u long__ Streptocara decora, p. 269.
5. Male 5.5 mm. long; 6 pairs of postanal papillae; spicules 1104 and 400u
long; from Colymbus arcticus and Larus ridibundus.
Streptocara tridentata, p. 271.
Insufficiently described. From Fulmaris glacialis.
Streptocara stellae-polaris, p. 270.
6. Male 11 mm. long; female said to be smaller than male; spicules 2504 and
AGO MON Geet nn ee ek TEA ea ae Streptocara recta, p. 269.
Male not over 5.5 mm. long; female 6 to 15 mm. long; spicules not over 88
PATET COs 8 10a 1G) Ses a La a eS bs
7. Five pairs of postanal papillae; long spicule measures 2654; in Gallus gallus.
Streptocara pectinifera, p. 266.
Six pairs of postanal papillae; long spicule measures 314 to 3964; in other
HOSts thancanovesen.. 2222. 2d de ste ee Se ree ee 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. 268.
Marked transverse striations and swelling absent. In Anseriformes.
Streptocara crassicauda, p. 267.
266 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
This key does not include Streptocara, species Baylis, 1919), a
single female specimen of which was found in Uria gryile in Yukan-
ski (Arctic Russia). Baylis states that this is probably Strepto-
cara pectinifera.
STREPTOCARA PECTINIFERA (Neumann, 1900) Skrjabin, 1916b
Synonym.—sS piroptera pectinifera Neumann, 1900.
Hosts—Primary: Gallus gallus and Numida meleagris ; secondary :
Unknown, probably arthropod or other small invertebrates.
Location—In the mucosa of the gizzard.
Morphology.—sStreptocara (p. 264): Whitish worms, attenuated
anteriorly. Cuticle transversely striated. Mouth (fig. 828 @) 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 55z
posterior to head end, or 42 and 36y according to Gedoelst and Liége-
ois, are 2 cervical papillae (fig. 3285) 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 Liégeois.
Male 4 to 5.2 mm. long by 150 to 176u wide. Esophagus 2 mm.
long. The tail (fig. 828c) 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 300u long, ending in recurved barb, the right 75 to 88»
long, thick and barbed. Caudal alae, 200u long, unite anteriorly.
Female 6.5 to 9.6 mm, long by 275 to 280u wide. Esophagus 2.4
mm. long. Vulva slightly posterior to equator of body, about 5.47
mm. from head end. Eggs 33 long by 20u wide, according to most
authors, or 37 to 39u long by 20 to 21p wide, according to Gedoelst
and Liégeois, 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—Kurope (France, Belgium, and perhaps Italy and
Arctic Russia).
STREPTOCARA CIRROHAMATA (Linstow, 1888) Skrjabin, 1916b
Synonyms.—Filaria (Spiroptera) cirrohamata 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. 264): Head (fig. 829a) with 2 coni-
cal lips and rudimentary neck frills. Cervical papillae 140, from
head end, tricuspid.
Male 7.58 mm. long by 250un wide. Esophagus somewhat less than
1/2 the body length; tail 1/20 of body length. Posterior extremity
(fig. 8296) rounded; caudal alae broad. Four pairs of preanal, 5
pairs of postanal pedunculated papillae. Spicules very unequal, the
longer 600u, bearing a small terminal process projecting at right
angles, the shorter spicule 140. long.
Female 9.72 mm. long by 350u wide. Esophagus 1/2.6, tail 1/81 of
body length. Vulva just anterior to end of esophagus. Eggs 39u
long by 19» wide, with very thick shell.
Fies. 328-329.—328, STREPTOCARA PECTINIFERA. @, HEAD END; }b,
CERVICAL PAPILLAE; C, MALE TAIL, AFTER NEUMANN, 1900. 329,
STREPTOCARA CIRROHAMATA. @, HEAD; b, MALE TAIL. AFTER LIN-
stow, 1888
Life history—Unknown; see S. pectinifera, p. 266.
Distribution —Not given. Collected on Challenger expedition.
STREPTOCARA CRASSICAUDA (Creplin, 1829) Skrjabin, 1916b
Synonyms—S piroptera crassicauda Creplin, 1829; Dispharagus
crassicauda (Creplin, 1829) Molin, 1860; Streptocara crassicauda
anseri Skrjabin, 1916.
Hosts—Primary: Anas clangula, A. glacialis, A.tadorna, A. fusca,
A. boschas, Alea torda, Bernicla sandwichensis, Bucephala clangula,
Colymbus arcticus, C. rufovulgaris, C. septentrionalis, Harelda gla-
cialis, Mergus serrator, M. merganser, Nyroca clangula, Oidemia
fusca, Tadorna tadorna; secondary: Unknown; see S. pectinifera,
-p. 266.
Location.—Gizzard.
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 334 from head end, with 5 to 9 points of
different sizes.
Male 3 to 5.5 mm. long by 2002 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 360n wide. Buccal cavity
29u, esophagus 2 mm. long. Tail end (fig. 330 6) rounded, the anus
25u from end. Vulva (fig. 330 ¢) just slightly posterior to middle
Figs. 3380—331.—330, STREPTOCARA CRASSICAUDA. @, HEAD; Db, TAIL OF FEMALE;
C, VULVA. AFTER SKRJABIN, 1916. 331, STREPTOCARA CRASSICAUDA CHARA-
DRI @, HEAD; b, CERVICAL PAPILLA. AFTER SKRJABIN, 1916
of body, according to Skrjabin. Eggs 26 to 37 long by 17 to 22n
wide.
Life history—Unknown; see S. pectinifera, p. 266.
Distribution —EKurope 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. 266.
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, 19200.
Host—Primary: Alcedo ispida; secondary: Unknown, see S.
pectinifera, p. 266.
Location.—Between the tunics of the gizzard.
Morphology.—sStreptocara (p. 264): Collar (fig. 332 @) 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. 3325). Four pairs of preanal,3 pairs of postanal papil-
Jae. Spicules very unequal, the longer measuring 237» long by 8.3u
wide, the shorter 68 long by 14y wide.
Female 8 mm. long by 200u wide. Anus 130» from tail end. Vulva
500u from tail end. Eggs peculiar; they are 39, long, flattened in
one direction where they are no wider than 234 whereas they are
enlarged in the other direction to 31.54 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. californica 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.—Filaria 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 6) 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 360n 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-
Fies. 332—333.—332, STREPTOCARA DECORA. 4d, ANTERIOR END; 0b, MALE
TAIL. AFTER DUJARDIN, 1845. 833, STREPTOCARA RECTA. a@, DORSO-
VENTRAL AND 0b, LATPRAL VIEW OF HEAD; C, MALE TAIL; d, SPICULES.
AFTER MUELLER, 1897
tween the papillae of the most posterior lateral pair. Spicules (fig.
333d) unequal and dissimilar, the one 490» 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
Linstov give the length). Tail end rounded. Vulva not salient,
dividing the body length in ratio of 7:4. Size of eggs not given,
Life history.—Unknown; see S. pectinifera, p. 266.
Distribution—Europe (Germany).
STREPTOCARA STELLAE-POLARIS (Parona, 1901) Skrjabin 1916b
Synonyms.—Histiocephalus stellae-pollaris Parona, 1901; Y seria
stellae-polaris (Parona, 1901) Gedoelst, 1919.
Host—Primary: Fulmarus atibiabes" secondary: Unknown; see
S. pectinifera, p. 266.
Location.—Not given.
NEMATODE PARASITES OF BIRDS Zit
Morphology.—sStreptocara (p. 264): 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.
Distribution.—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 Yserta 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, 1916b
Synonyms.—Fularia triaenucha Wright, 1879; Acuaria triaenucha
(Wright, 1879) Ward, 1918.
Host.—Primary: Botaurus minor; secondary: Unknown; see S.
pectinifera, p. 266.
Location.—Proventriculus.
Morphology.—sStreptocara (p. 264): Cuticle densely striated. A
cervical frill present, the tops of the lateral loops being 180» from
anterior end; it extends posteriorly on the neck for a distance of
4054. Cervical papillae (fig. 334) tricuspid, the root being 60.
posterior to the end of the frill; the papilla itself measures 604 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, 1916b
Synonyms.—Filaria tridentata Linstow, 1877 a and 6; Spiroptera
tridentata (Linstow, 1877) Neumann, 1900.
Hosts—Primary: Colymbus arcticus, Larus ridibundus; second-
ary: Unknown; see S. pectinifera, p. 266.
Location.—Esophagus and intestine.
3612—27T——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. 3350).
Male 5.5 mm. long by 1304 wide. Tricuspid papillae 200% from
head end. Anterior part of esophagus 140p long; total esophagus
620 or 1/9 of total body length; tail 1/32 of body length. Caudal alae
(fig. 835a) 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, long, the latter with a
bulbous swelling at its proximal end.
Female 16.4 mm. long by 220n wide. Cervical papillae 160. from
head end. Esophagus 2.4 mm., tail 3002 long. Vulva divides body
ee 356
N 33S > =
SSF
Fies. 334-336.—334, STREPTOCARA TRIAENUCHA. CERVICAL PAPILLA. AFTER
WRIGHT, 1897. 335, STREPTOCARA TRIDENTATA. a, MALE TAIL. AFTER LINSTOW,
1909. 06, 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; Cheilospirura 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 tridentatus___ Synhimantus sygmoidea, p. 282.
Male (if known) 6 mm. long or longer; from hosts other than above____ 2.
NEMATODE PARASITES OF BIRDS Lic
. Description incomplete; female 20 mm. long; from Falco subbuteo.
Synhimantus denticulata, p. 275.
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 156u and 372u long..__.__--_________ Synhimantus affinis, p. 273.
Vulva posterior to middle of body (except in S. laticeps where perhaps
just anterior to middle) ; spicules, if male known, differ in length from
above (except possibly in NS. sagittata; lengths not given) ~___-_______ 4,
4, Only female known, 18 to 15 mm. long; cordons extend posteriorly for 24
or more of the length of the muscular esophagus, a distance of 8404 from
the head end; vulva 510n, anus 170u from posterior end; from Pele-
GdniusSpecies..: 2s2: bast fee eae Synhimantus raillieti, p. 283.
Both male and female known; female not agreeing in all respects with
above; from hosts other than Pelecunus__________--_-_-_-_---»_-_-_--_-______ 5.
5. Male with 6 to 7 pairs of caudal papillae___-___-___ ee 6.
Male with 8 to 10 pairs of caudal papillae_______________ (2
6. Cloacal aperture of male only 63 from tail end; vulva of female not more
than 700u from tail end____________.__ Synhimantus brevicaudata, p. 274
Cloacal aperture of male much more than 68 from tail end; vulva described
as posterior to middle of body______-_____ Synhimantus elliptica, p. 277.
7. Vulva near posterior end of body__-__-_______________________________ 8.
Vulva not-tar posterior to, middle of. body... 2e2 ~_=-_5_§__. sees 10.
8. Male with 10 pairs of caudal papillae; 6 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
VOTING sae ee Synhimantus invaginata, p. 279.
Male 7 mm. long; female 9 mm. long; cordons narrow throughout entire
COUTSO Hs ees fea!) te ee i ae et 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 190u and 720u long;
CEOS SOW Veto es ee Synhimantus hamata, p. 278.
Male with 9 pairs of caudal papillae; spicule lengths different from above ;
eggs at least 37 long and 224 wide_____________-______--__-_--__--~ Li:
11. Male 6.4 to 6.8 mm. long; female 10.5 mm. long; spicules 200 to 210u and
950 to 960u long; in Asturinula monogrammica,
Synhimantus subrecta, p. 282.
Male 7.2 to 10 mm. long; female 11 to 21 mm. long; spicules 170u and 5902
oT
long; in hosts other than above_____---___- Synhimantus laticeps, p. 276.
SYNHIMANTUS AFFINIS (Scurat, 1916) Seurat, 1919
Synonyms.—Acuaria affinis Seurat, 1916; Filaria laticeps in part
of Mueller, 1897; Acuaria laticeps, female of Seurat, 1915.
Hosts.—Primary: Strix flammea (=Tyto alba); secondary: Un-
known.
Location.—Ksophagus.
Morphology.—Synhimantus (p. 272): Body robust, elongate.
Cuticula thick, transversely striated. Buccal lips bear truncate tooth
and a pair of projecting papillae. Cordons (fig. 337b), 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. wide. Cordons 395 long. Tri-
cuspid papillae 4454 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 156y long, the left 372» long.
Female 11.2 to 27.5 mm. long by 300 to 6124 wide. Cordons 660,
long in worm 19.3 mm. long; tricuspid papillae 410 and 696y, and
<a a
Fic. 337.—SYNHIMANTUS AFFINIS. a, FEMALE GENITALIA. AFTER SEURAT, 1915.
6, 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, 420u long, the cuticular ove-
jector 210n long. Eggs 38u by 28, thick-shelled, embryonated when
deposited.
Life history—Unknown; probably involves ieee ate 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.—sSynhimantus (p. 272): Body filiform. Head 25 to
354 wide, with 2 conical projecting lips. Cordons (fig. 338)
transversely striated, extending posteriorly for 230 to 300n, 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
30n wide; posterior esophagus (ventricule of Dujardin) 2.16 mm.
long.
Male 10 mm. long by 160u wide. Cordons extend 228, from head.
Posterior extremity of body rolled spirally in 8 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 2904 wide (Skrjabin). Cordons extend 310» from head end
(Dujardin) or 4254 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
560u from head end. Pharynx 210» long, slender; anterior esopha-
gus 850u long. Caudal extremity thick, in the form of a short obtuse
cone, the anus 68 (Skrjabin) from tail end (fig. 3386). Vuiva 640
or 700 from tail end. Eggs elliptical, 20 to 234 long (Dujardin) or
30u by 254 (Skrjabin).
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution—KEurope (France and Russia).
SYNHIMANTUS DENTICULATA (Molin, 1860) Skrjabin, 1924
Synonyms.—Dispharagus denticulatus Molin, 1860c; Dispharagus
falconis-subbuteonis Diesing, 1851; Spiroptera falconis-subbuteonis
(Diesing, 1851) Diesing, 1851.
Hosts—Primary: Falco subbuteo,; secondary : Unknown.
Location.—Esophagus.
Morphology.—Synhimantus (p. 272): Mouth with 2 papilliform
projecting lips (fig. 339). 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 wnknown.
Female 20 mm. long by 200n wide. Cordons extend posteriorly
520u from head end; anastomosis 270» from head end. Eggs elon-
gate oval, 36u by 16y.
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution —Kurope (France (Rennes) ).
276 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
SYNHIMANTUS LATICEPS (Rudolphi, 1819) Stiles and Hassall, 1920
Synonyms.—S piroptera laticeps Rudolphi, 1819; Dispharagus lati-
ceps (Rudolphi, 1819) Dujardin, 1845; daria 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; Acwaria laticeps (Rudolphi, 1819) Railliet, Henry, and
Sisoff, 1912; Acuaria (Synhimantus) laticeps (Rudolphi, 1819)
Raillhet, Henry, and Sisoff, 1912.
Hosts—Primary: Falco lagopus, Strix bubo, S. alba, S. brachyo-
tus, Falco nisus, F. tinnunculus, Gallus domesticus, Falco cyaneus,
Fies. 338-341.—338, SYNHIMANTUS BREVICAUDATA. @, HEAD END; D,
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, 1866
Strix flammea, Bubo maximus, Buteo lagopus, Circus cineraceus, C.
cyaneus, Otus brachyotus, Cerchneis tinnunculus, Otus vulgaris, Ac-
cipiter nisus, Aegiolus otus; secondary : Unknown.
Location—Esophagus and proventriculus.
Morphology—sSynhimantus (p. 272) : Cutaneous cordons (fig. 341)
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. 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 170p
long by 18» wide, the left 5904 long, slender, its free portion alate
and its tip slightly dilated and divided into 2 lateral lobes. Six to
NEMATODE PARASITES OF BIRDS Zid
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 by 25y, 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.
Fic. 342.—SYNHIMANTUS LATICEPS. FEMALE GENITALIA. AFTER 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.—Synhimantus (p. 272): Mouth with 2 projecting
papilliform lateral lips. Body strongly striated transversely. Cor-
dons thick.
Male 7 mm. long by 200» 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 40u by 19 to 27p.
Gendre (1921a) notes that according to Molin’s (18610) 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.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
278
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 Vepervier of Dujardin, 1845; Filaria
Dispharagus hamatus (Linstow, 1877) Stos-
hamata Linstow, 1877;
sich, 1891; Acuaria hamata (Linstow, 1877) Railliet, Henry, and
Sisoff, 1912.
!
E
-
iS
Mm
nue tare
SOMO
CC ll
DS
2
Fics. 343-344.—343, SYNHIMANTUS BLLIPTICA. MALE TAIL. AFTER MOLIN, 1861.
a, HEAD END. AFTER LINSTOW, 1879. 06, MALE
344, SYNHIMANTUS HAMATA,
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 300u wide. Buccal cavity 180 long; esoph-
agus 7204 long. Cordons extend posteriorly 280 from head end and
anastomose 1404 from head end. Right spicule 720u long, the free
end shaped like a fishhook; left spicule 1902 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 8S. laticeps.
NEMATODE PARASITES OF BIRDS 279
Female 7.4 mm. long (Linstow) or 10 to 11 mm. long (Dujardin)
by 4802 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 30u by 19 (Dujardin).
Life history—Unknown; probably involves intermediate stages in
other hosts.
Iistribution Europe (Prussia (Hanover) ).
SYNHIMANTUS INVAGINATA (Linstow, 1901) Skrjabin, 1924
Synonyms.—Dispharagus invaginatus Linstow, 1901; Acuaria
(Synhimantus) 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.
Fic. 345.—SyNHIMANTUS INVAGINATA. @ AND b, HEAD END; C, MALE TAIL; d,
RIGHT AND €, LEFT SPICULE; f, FEMALE TAIL. AFTER GPNDRE, 1913
Morphology.—sSynhimantus (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 5h
apart anteriorly and 10 to 14p apart posteriorly. Cordons (fig. 345a
and 6) 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 ¢). Pharynx 250 to 270» long; anterior esoph-
agus 960 to 970u long; posterior esophagus 3.2 mm. long. Cordons
extend 550, posteriorly from head and anastomose 300. from head
end. Tricuspid papillae 630.2 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. 345d)
large and robust, 900p long by 35» 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 410u wide. Pharynx 280 to 310» long; anterior
esophagus 1 mm. long; posterior esophagus 3.5 to 3.6 mm. long.
Cordons extend 610 from head end and anastomose 330» from head
end. Tricuspid papillae 770.2 from head end. Posterior extremity
in a hyaline cylindrical sheath formed from the cuticula (fig.
345 f). 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 by 18 to 19p.
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution —Europe (Germany (Langenburg) and Corsica)
and Africa (Algeria and French Guinea (Labé) ).
SYNHIMANTUS RECTA (Molin, 1860) Gendre, 1920
Synonyms.—Dispharagus rectus Molin, 1860c; Spiroptera fal-
conis® Molin, 1860; Acuaria recta (Molin, 1860) Railliet, Henry, and
Sisoff, 1912; Chetlospirura recta (Molin, 1860) Stiles and Hassall,
1920.
Hosts.——Primary: Falco femoralis, F. unicinctus; secondary: Un-
known.
Location.—Stomach (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 Chetlospirura 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 100p 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 3004 wide. Posterior extremity
straight, the anus not far from the tail end. Vulva in posterior
part of body.
®° Catalogue of Vienna Muscum.
NEMATODE PARASITES OF BIRDS 281
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution.—South America (Brazil).
SYNHIMANTUS SAGITTATA (Rudolph, 1809) Cram, 1927
Synonyms.—Ascaris sagittata Rudolphi, 1809; Spiroptera alata
Rudolphi, 1819; Dispharagus analis Molin, 1860; Filaria 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.
—
a, ItL.\ eS
@e
oe So
= 0
S96 ft
. 3 r= 7
A W- ~
33 SS =x
: = »\
Figs. 346—347.—346, SYNHIMANTUS RECTA. MALE TAIL. AFTER DRASCHE,
1884. 347, SYNHIMANTUS SAGITTATA. a, HEAD END; b, MALD TAIL. AFTER
SCIHINEIDER, 1866
Hosts—Primary: Ardea nigra, A. purpurea, Ciconia nigra, Buteo
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 7mm. long. Tail curled in 8 spiral turns. Caudal alae thick,
vesicular. Nine pairs of caudal papillae (fig. 347b), of which 5
pairs are postanal.
Female 9 mm. long. Vulva very near the anus. Eges 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: Asturinula 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 250n 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 ¢
and @) 950 to 960, long, the right (fig. 348 6) 200 to 210u long; these
distinguish this species from S. laticeps.
Female 10.4 to 10.5 mm. long by 330 to 370 wide. Cordons 420
to 430p long. Anus 200 to 210n from tail end, or 10/495 to 10/518 of
total body length from tail end (fig. 348 e). Posterior extremity coni-
cal, rounded atend. Vulvaa little posterior to middle of body, or 10/19
of distance from head. Ovejector directed posteriorly, composed of a
large vestibule 270u long, followed by a sphincter 200, long; the
varnish gland or trompe is very short and divides into 2 branches
which immediately diverge. Eggs 37h by 224, embryonated when
oviposited.
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution —Africa (Dahomey).
SYNHIMANTUS SYGMOIDEA (Molin, 1860) Skrjabin, 1924
Synonyms.—Dispharagus sygmoideus Molin 1860e; Acuaria syg-
moidea (Molin, 1860) Railliet, Henry, and Sisoff, 1912.
NEMATODE PARASITES OF BIRDS 283
Hosts—Primary: Falco tridentatus; secondary: Unknown.
Location.—Not 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 300p 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.
Fic. 348.—SYNHIMANTUS SUBRECTA. ad, MALE TAIL; 0, RIGHT SPICULE; C, PROFILE
VIEW AND d, VENTRAL VIEW OF LEFT SPICULE; €, FEMALD TAIL, AFTER GENDRD,
1921
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution.—South America (Brazil).
SYNHIMANTUS RAILLIETI (Skrjabin, 1924) Cram, 1927
Synonym.—A cuaria 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 700n wide. Cordons extending pos-
teriorly for a distance of 8402; anastomosis 280. from head end.
Pharynx 425» long by 30n wide; muscular esophagus 560u by 100,;
glandular esophagus 3.57 mm. by 70z. Vulva 510 from the tail
end, a distance of 340, anterior to the anus. Tail conical, with
rounded end; anus 1704 from extremity. Eggs 38u by 20pn, swith: 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 GENERA OF SCHISTOROPHINAR
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
. Mouth with 6 lips; cephalic ornamentation of multiple appendices of various
shapes and sizes, some directed anteriorly, some posteriorly ; caudal alae
tbo
asymmetrical; vulva in region of anusi_. —-_~______--_~ 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
GEL CUS Oe = yee eS 2 a Md ee Sas oe Yseria, p. 292.
Mouth with 4 lips; vulva in anterior part of body; lateral papillae not
described: AS Tricuspid ase ae aE | awa ED Histiocephalus, p. 290.
Genus SCHISTOROPHUS Railliet, 1916a
Synonyms.—T etracanthus 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. JM/ale 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. /Hemale 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.
KBY 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 Glareola austriaca.
Schistorophus spinulosus, p. 288.
Male 4.5 mm., female 9 mm. long. From Tringa helvetica (=Squatarola hel-
vetica=Squatarola squatarola) ___-__-_-__- Schistorophus bicuspis, p. 286.
PeOAVD VARIG MN OGIe. Ol DOU Yn ee ee Fe 4.
Vulva DOSLCEIONA part Ob J00GV= = - a ee a
4. Female 23.5 mm. long. Cuticular processes of head broad. From Haemato-
pus ostralegus; Russian Turkestan_____~_ Schistorophus aulieatina, p. 287.
Female 14 mm. long. Cuticular processes slender. From Rdallus cayennensis ;
eZ MS ge a ee at 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. 285.
Head not cup-shaped, cuticular processes with only 4 points, posteriorly
UNS CEC ce se ed Ne ee ee Eo Se 6.
6. 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
SDC CIOS ee re ia tn Free 8 8 os oe Schistorophus longicornis, p. 285.
SCHISTOROPHUS LONGICORNIS (Hemprich and Ehrenberg, 1866) 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 and 6)
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.—S piroptera bicuspis Rudolphi, 1819; Dispharagus bi-
cuspis (Rudolphi, 1819) Dujardin, 1845; Histiocephalus gracilis
Diesing, 1851; Histiocephalus bicuspis (Rudolphi, 1819) Linstow,
1878.
a.
oe
2600 0 099000 000 00800% %e
Fig. 349.—ScCHISTOROPHUS LONGICORNIS. a, LATERAL VIEW AND b, FRONT VIEW OF
HEAD; C, MALE TAIL. AFTER SCHNEIDER, 1866
Host.—Primary: Tringa helvetica (=Squatarola helvetica) ; also
reported from Grus cinerea and Vanellus melanogaster; secondary :
Unknown.
Location.—Between tunics of gizzard.
Morphology.—sSchistorophus (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—Kurope (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-
laris Molin, 1860; Cheilospirura capillaris (Molin, 1860) 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—sSchistorophus (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% wide. Anus very near to tail end,
the width of body at anus being only 34p. Vulva almost in middle
of body, slightly anterior to middle. Eggs oval, 40u long by 26,
* wide.
Life history —Unknown.
Distribution—Asia (Russian Turkestan).
288 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
SCHISTOROPHUS LACINIATUS (Molin, 1860) Railliet, 1916a |
Synonym.—H istiocephalus laciniatus Molin, 1860c.
Host——Primary: Rallus 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 100p wide; caudal extremity twisted in spiral,
with long equal alae which have 24 single papillae.
Female 14 mm. long by 200u 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.—S piroptera glareolae austriacae’; Filaria spinulosa
Molin, 1860d.
Host—Primary: Glareola austriaca; secondary: Unknown.
Location.—Between the tunics of gizzard.
Morphology.—sSchistorophus (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» wide. Caudal extremity conical,
obtuse.
Life history —Unknown.
Distribution—Kurope (Austria (Vienna Museum) ).
SCHISTOROPHUS BIDENS (Rudolphi, 1819) Railliet, 1916a
Synonyms.—S piroptera bidens Rudolphi, 1819; Dispharagus bidens
(Rudolphi, 1819) Dujardin, 1845; Spiroptera denticulata Molin,
1860 6; Ancyracanthus bidens (Rudolphi, 1819) Schneider, 1866.
Hosts—Primary: Accipiter nisus, Astur palumbarius, Falco pal-
umbarius, Merops apiaster; secondary : Unknown.
Location.—Between the tunics of gizzard.
Morphology.—sSchistorophus (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 @) 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 ctip-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
Fics. 850—-353.—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; 0, MALE TAIL. AFTER LEPRI, 1898
2 pairs of postanal pedunculated papillae (fig. 353 6) but Schneider
describes and figures 16 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 the middle of the body; accord-
ing to Schneider 6 mm. from the posterior end in a speciman 15 mm.
long. Eggs 43» 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. Jfale with large caudal alae with pedunculated papil-
Jae of which 4 pairs are preanal. Spiculesequal or unequal. Female
with vulva in anterior part of body.
Parasitic under the tunic of gizzard of birds.
Type species. = ae laticaudatus (Rudolphi, 1819) Dies-
ing, 1851.
This genus was placed in the Acuariinae by Skrjabin (19165) on
the basis of the caudal papillae and the ornaments of the head.
Gendre (19215) states that because of the lip structure, the genus
belongs in the Spiruridae rather than the Acuariidae. Travassos
(19200) 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 700u long, the right 1554 long____________ Histiocephalus tridens, p. 291.
Head with 20 to 24 processes, each ending in 2, 3, or 4 branches; spicules equal
(GiaramMlong) 2a SS eee 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 6)
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 66u. 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 200n wide, or to 340n, according
to Skrjabin (fig. 355). Posterior extremity of body spirally twisted,
according to Molin, slightly enlarged and provided with 2 canal
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 16% wide. The
cloacal aperture in a worm this size is 110% from the posterior
extremity.
Female 7 to 14 mm. long by 200 to 400» 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: T'rachelotis 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
Figs. 354-355.—HISTIOCEPHALUS LATICAUDATUS. 3854, @, FRONT VIEW AND Jb, LATERAL
VIPW OF HEAD; C, MALE TAIL. AFTER DRASCHE, 1884. 355, MALE. AFTER
SKRJABIN, 1916
difficult to study. Body filiform; cuticle transversely striated.
fivad (fig. 356 @ and 6) distinctly set off from body by a furrow,
following which is a cuticular collar, 16» high in the male, 20 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 38 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 200z wide. Cloacal aperture 96, from pos-
terior extremity (1/54 of total length). Caudal alae (fig. 356 ¢) 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. 856 d, e, and f)
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 z) 155 long, its appearance different in different.
views.
N
Fic. 356.—HISTIOCEPHALUS TRIDENS. ad, FRONT VIEW AND D, PROFILE VIEW OF
HEAD; C, MALE TAIL; d, ANTERIOR END; €, POSTERIOR END, LATERAL VIEW; f,
VENTRAL VIEW, LEFT SPICULE; g, RIGHT SPICULE, LATERAL VIEW; hh, RIGHT
SPICULE, DORSAL VIEW; i, SHOWING NOTCHED APPEARANCE WHEN SPICULE RE-
MOVED FROM BODY. AFTER GENDRE, 1921
Female 12.1 mm. long by 220n 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. Eggs:
not mature in Gendre’s specimens.
Life history—Unknown.
Distribution—Africa (French Guinea (Labé)).
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.—Y seria californica Gedoelst, 1919.
NEMATODE PARASITES OF BIRDS 293
KEY TO SPECIES OF YSERIA
Female 12 mm. long by 100u wide; male 8 mm. long. In Alcedo americana
and Rallus cayennensis_______-____-_-_._-.__.---.---- Yseria coronata, p. 293.
Female 25.5 mm. long by 4164 wide; male unknown. In Oidemia deglandi.
Yseria californica, p. 293.
YSERIA CALIFORNICA Gedoelst, 1919
Host.—Primary: Oidemia deglandi; secondary: Unknown.
Location—Under mucosa of gizzard.
Morphology.—yY seria (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 185p 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.—s piroptera coronata Molin, 1860; Histiocephalus
coronatus (Molin, 1860) Skrjabin, 19160.
Hosts—Primary: Alcedo americana and Rallus cayennensis; sec-
ondary: Unknown.
Location.—Under mucosa of gizzard.
Morphology.—yY seria (p. 292) : Head set off from body, armed with
crown of sharply pointed, posteriorly directed processes. Mouth
with 2 small tooth-like papillae. Body slender, increasing in width
pes‘eriorly, 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 tail; 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.
T ype-species.—Serticeps vulvoinflata (Molin, 1860) Railliet, 1916a.
294 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
SERTICEPS VULVOINFLATA (Molin, 1860) Railliet, 1916a
Synonym.—sS piroptera vulvoinflata Molin, 18606.
Host.—Primary: Trochilus ochropygus; secondary: Unknown.
Location.—Between the tunics of the gizzard.
Morphology.—wSerticeps (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
Figs. 357—358.—SERTICEPS VULVOINFLATA. 357, a, HmsaD; 0b, MALE TAIL. 358,
LABELED SPIROPTERA MEDIOSPIRALIS, BUT APPARENTLY INCORRECTLY SO. SEE TEXT
FOR DISCUSSION. AFTER DRASCHB, 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. 162): Mouth with 2 lips pro-
vided with teeth on their inner surface. Head without ornamenta-
tion. Jale 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.
Ty pe-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.
fale 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.
Ty pe-species.—P hysaloptera clausa Rudolphi, 1819.
Travassos (1920c) divided the genus Physaloptera into 5 genera:
Physaloptera, Clamidonema, 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
i)
Oo
On
10.
a
13.
KDHY TO SPECIES OF PHYSALOPTERA
. True Physaloptera, i. e., species agreeing with the generic diagnosis____ 2.
Species inquirendae, placed in this genus but not agreeing entirely with the
SEMEL IC CLA INO SIS es eee oe a ee EE Se ee 16.
. Normally in reptiles; found as pseudoparasite in Ciconia alba.
Physaloptera abbreviata, p. 307.
INormall ying bird sset as oe ee ee ee eee 3.
= Teeth of Lipsaundescribed= sos = sau. ae ee ee 4.
Teeth of lipsxdesenibed 22 er ey a ee ee ee ee 6.
. Description very incomplete; from Lanius minor.
Physaloptera bilabiata, p. 307.
Krom: other: hosts than ab0Ve= 22-2 23 =. oe ee ee ee ee eee 5.
. Male 18, female 30 mm. long; cuticle enlarged posterior to the lips; eggs
8lu long by 264 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.
. Two teeth on each lip, an outer and an inner (tripartite) -___-_-__-__--__ Me
One: tooth or more.thean 2teeth on each lips]! <2) a ee 15.
. Relative sizes of teeth not described; distinctive character consists in the
one species (P. truneata) 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 iecentral caudal: papillae kesh ea aan sess ee Eh to SE ARs ae ee 8.
Relative sizes of teeth described; outer tooth not as above; pedunculated
Papiblae smu MPe! ry We EO TG ee es ae a 9.
. Male 14 mm., female 28 mm. long; 5 pairs of lateral pedunculated caudal
papillae; 1 pair of ventral oe A le he bak Physaloptera crassa, p. 302.
Male 25, female 38 mm. long; 4 pairs of lateral pedunculated caudal papil-
lae; 43 ventralspapillacs ieee _ ar eae Physaloptera truncata, p. 306.
. Outer tooth large, inner tripartite tooth small__.__-_=--_____~_-_=_-_= 10.
Inner (\tripartite,.tooth large,,outer tooth ;smalliis)) ee ee eee 14.
Caudal alae with 4 pairs of lateral pedunculated papillae.
Physaloptera acuticauda, p. 297.
Caudal alae with 5 pairs of lateral pedunculated papillae_____________ et
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) 225 2 Se Oe ek aa 8. 12;
. Male 7 mm. long; cervical papillae situated 100u posterior to the end of
muUuscularsesopha gus... 12M ees Physaloptera alata chevreuxi, p. 301.
Male 17 mm. or longer; cervical papillae at level of terminal region of
muscular. ‘esophagus i eiil ei) ot EG ie Va Ee ee 18.
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.
Maile 17 to 20 mm., female 19 to 27 mm. long; cloacal aperture of male 6504
from posterior extremity ; vulva of female posterior to end of esophagus,
at the 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 880u long) ; vulva a little anterior to middle of
DO Gyees se yt eet a Physaloptera galinieri, p. 303.
Spicules unequal (400 and S40u long); vulva at anterior third of body
ISU Op Nae ee ee IP aS. Physaloptera subalata, p. 305.
25. 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 [bis 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___________________ UG
17. Description very incomplete; alae described as vesicular; in Cuculus, species,
FS 1 7 en eee eines ee ee Physaloptera strongylina, p. 309.
Alae not described as vesicular; in other hosts than above, in Africa or
BEST CTS 0) 6) Cs ee a a cee a ore ee 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.
49. 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 39u
lone Dy 26m wide. --. = oe Se Physaloptera brevicauda, p. 307.
No papillae or teeth described on lips; spicules unequal; tail of male 1/33
of body length; eggs 46u long by 294 wide__ Physaloptera malleus, p. 308.
20. Spicules equal (290u 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 (SSOu 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.—P hysaloptera alata Rudolphi of Diesing, 1851 in part.
Hosts—Primary: Elanus caeruleus, Falco atricapillus (=Spizi-
aster melanoleucus), F’. cachinans (=Herpetotheres cachinans), F.
cayennensis (=Leptodon cayennensis), F. coronatus (=Harpyha-
liaéitus coronatus), F. dispar (=Elanus leucurus), F. gracilis
(=Geranospizias caerulescens), F. minutus (=Accipiter tinus), F.
ornatus (=NSpizaetus mauduyti), F. palustris (=Cireus maculosus ),
F. species, F. swainsonii (=campsonyx swainsoni), F. rutilans
(=Heterospizias meridionalis), F. unicinctus (=Parabuteo unicine-
tus), F. urubutinga (Urubitinga urubitinga) ; 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. wide. Caudal alae (fig.
3596) 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 50u wide.
Female 20 to 43 mm. long by 800n 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, shghtly enlarged
posteriorly; 2 uteri arising from the outer sides (fig. 359a). This
is the only one of the Physaloptera from birds known to have this
a4
—<————d
OSM
|
Kies. 359-360.—359, PHYSALOPTERA ACUTICAUDA. a, TERMINAL PART OF FEMALE
GENITALIA; 6b, MALE TAIL. AFTER ORTLEPP, 1922. 860, PHYSALOPTERA ALATA. 4,
HbAD; b, LIP; c, MALY TAIL. AFTER LINSTOW, 1877
arrangement of uteri; it is similar to that of P. praeputialis, a par-
asite of mammals. Eggs 51y 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 imperialis, A. pennata,
Astur nisus, Buteo vulgaris, Circaétus gallicus, Circus aeruginosus,
CU. 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. pygargus, F. rufus, F. rutilans, F. species, F. subbuteus,
F. swainsonii, F. unicinctus, F. urubutinga, Tinnunculus 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. 3600), 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 650u from caudal extremity. Circumcloacal region
covered with small tubercules. Caudal alae (fig. 360¢) very elon-
gated. Five pairs of long pedunculated papillae, of which 2 are
preanal, 1 adanal 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
Fic. 361.—PHYSALOPTERA ALATA RUDOLPHI OF ORTLEPP, 1922. POSSIBLY
EQUIVALENT TO P. GALINIPRI SEURAT. 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 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; trompe dilated
as a reservoir, beyond which it narrows again for a course of 3800p to
the branching of the uteri. Eggs thick-shelled, embryonated, 46 to
55pu long by 25 to 27n wide.
300 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Life history—Unknown; probably involves intermediate stages
in insects.
Distribution—Kurope (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 /. gallicus
and 7. 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’. biarmicus. Ortlepp (1922) examined Rudolphi’s
specimens from 7’. 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-
galimiert Seurat, that part of Rudolphi’s material may be the latter
species.
The above description of P. alata is based on those of Schneider,
Linstow, and Seurat.
PHYSALOPTERA MEGALOSTOMA Creplin, 1829
Synonym.—S piroptera megalostoma (Creplin, 1829) Dujardin,
1845.
Hosts.—Primary: Falco nisus and F. palumbarius; secondary :
Unknown.
Location.—Proventriculus.
Morphology.—P hysaloptera (p. 295) : Color brownish; head larger
than body. 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 Rudolphi’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 nisus; 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 515y wide. Muscular esophagus 260, long;
entire esophagus 1/4 of body length; cervical papillae 370 and 360y
from anterior end.
Fic. 362.—PHYSALOPTERA MEGALOSTOMA. AFTER CREPLIN, 1829
Female (immature), 8 mm. long. Tail long (310) 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, 191lic
Hosts—Primary: Accipiter nisus and Aquila chrysaetos ; second-
ary: Unknown.
Location.—Ksophagus.
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 8 points.
Male 22 to 28.5 mm. long by 900% wide. Tail 1.165 mm. long,
slender and pointed; small cuticular tubercles in circumcloacal re-
gion. Caudal papillae as in P. alata. Right spicule robust and
large, 5502 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 500u long, dividing posteriorly into 2 uteri. Eggs
50 by 25p, 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, and 360, long.
Female 23 mm. long by 2mm. 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.—P hysaloptera (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 6604 wide. Caudal alae (fig. 363 0) well-
developed; circumcloacal region covered with tubercles. Cloacal
aperture 9002 from tail end. Caudal papillae arranged as in P.
galiniert 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, 300u 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 300n wide; trunk (trompe) 400u
long, giving rise to 2 uteri. Eggs 55y 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: Micropogon, 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
Fics. 365-364.—363, PHYSALOPTERA CROSI. @, HEAD END; b, MALE TAIL; C, FEMALE
TAIL. AFTER SEURAT, 1914. 364, PHYSALOPTERA FUSIFORMIS. MALE TAIL. AFTER
LInstTow, 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 by 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 81p long by 26 wide, with very thick shells.
Life eS Galctaea: probably involves intermediate stages in
insects.
Distribution.—Europe (Germany, Breslau Museum).
PHYSALOPTERA GALINIERI Seurat, 1914i
Synonym.—P. alata Rudolphi of Ortlepp, 1922 may be a synonym
of this (see p. 299).
3612—2i—21
304 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Hosts—Primary: Aquila rapax belisarius and Melierax gabar;
secondary: Unknown.
Location.—Esophagus and proventriculus.
Morphology—Physaloptera (p. 295): Body robust. Mouth (fig.
65a) 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
Fic. 365.—PHYSALOPTERA GALINIERI. @, 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 8 pairs
occurring more posteriorly. Spicules short, subequal, 3602 and
380p long.
Female 17 to 34 mm. long by 1.04 mm. wide. Tail short (350,).
Vulva (fig. 365 0) unusually remote from esophagus, situated a short
distance anterior to middle of body. Vagina 2 mm. long, directed
anteriorly; egg chamber (reservoir) 600 long; triink (trompe) 200
long. Two uteri. Eggs 65p 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 domesticus) ;
secondary: Unknown.
Location.—Stomach and intestine(?).
Morphology.—P hysdloptera (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» wide. Tail 1/35 of total body length,
conical, rounded at end. Eggs 52y 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.—sS piroptera inflata Molin, 18606.
Host— Primary: Falco unicinctus; secondary: Unknown.
Location.—Esophagus and proventriculus.
Morphology—P hysaloptera (p. 295) : Head continuous with body,
with inflated cuticle. According to Drasche, this species does not
have the 8 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 3004 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.—P hysaloptera alata Rudolphi, 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 700u% 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. 8670), 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 6604 wide. Tail 240» long, conical.
Vulva not prominent, in anterior third of body, 3 mm. posterior
Fics. 366—368.—3866, PHYSALOPTERA GEMINA. MALE TAIL. AFTER LINSTOW,
1899. 367, PHYSALOPTERA SUBALATA. a, LIP; b, MALE TAIL, AFTHR SCHNEI-
DER, 1866. 368, PHYSALOPTERA TRUNCATA. a, LIP; b, 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) ).
PHYSALOPTERA 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. 3682).
a haw
NEMATODE PARASITES OF BIRDS 307
Male 25 mm. long. Four pairs of lateral pedunculated papillae
(fig. 368b), 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 cloacal 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.—sS piroptera 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 ¢
cone-shaped tooth. Esophagus 1/9 of total body length: tail very
short in both sexes.
Male 27 mm. long by 830p wide; tail (fig. 369) rounded, its length
1/69 of total body length; caudal alae with chitinous tubercules in
308 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 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 39u long by 26, 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.
Fies. 371-373.—371, PHYSALOPTERA, SPECIES. HEAD END.
1909. 370, PHYSALOPTERA MALLEUS. @, HEAD; b, MALE TAIL. AFTER LINSTOW,
18838
PHYSALOPTERA MALLEUS Linstow, 1883
Host.—Primary: Corvus cornia; secondary: Unknown.
Location.—Not given.
Morphology.—P hysaloptera (p. 295) : Head (fig. 370a) with 2 very
large rounded lips, hollowed out on the inner surface. Esophagus
1/6.5 of total length.
Male 20 mm. long by 760p 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» long by 29» 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.—sS piroptera affinis Dujardin, 1845.
Hosts—Primary: Cuculus melacoryphus and C. seniculus; second-
ary: Unknown.
Location.—Stomach and intestine.
Morphology.—P hysaloptera (p. 295) : Head continuous wtih body.
Mouth with lips. Body much attenuated anteriorly.
Male 6to8mm. 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 550» long spindle-shaped swelling over the
head end. ;
Male 13.8 mm. long by 660p wide. Esophagus 1/4.4 of total body
leneth. Caudal alae (fig. 372) oval, smaller in posterior part; large
310 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, 290u 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% wide. Vulva anterior to middle of body, ae the badly
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.
e
pire r
Fics. 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» from head end.
Male 18.4 mm. long by 4004 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 5102 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 long by 26p wide.
NEMATODE PARASITES OF BIRDS 31}
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 PAysaloptera 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, 19165. 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. J/ale 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.—T helazia Bosc, 1819.
Railliet included in this family the genera Schistorophus 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 GENERA OF THELAZIIDAE
1. Male with large caudal.alae_._._-.____.-/_-____________ Ceratospira, p. 319.
MSM WD NOML CHUdeWalaes = 25 22 e822 2 Se eo ee,
2..Vulya,in anterior part, of body,....-—— + — <2 48~—445---- = Thelazia, p. 311.
Vulva. in posterior part: of body .__.--...:-_._____... Oxyspirura, p. 321.
Genus THELAZIA Bosc, 1819
Synonym.—T helazius Bose, 1819.
Generic diagnosis —Thelazidae (p. 311): Mouth without lps,
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.
3612—27
99)
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 V’helazia lutzi
and 7. 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 Craz fasciolata.
Thelazia anolabiata, p. 317.
Mouth and lobes of buccal cavity not described as above; from hosts other
CHAN BD OVC 8 a ee a a Dn
2. Male 22 to 24 mm. long; female 25 to 27 mm. long; 10 pairs of preanal papil-
lae; cloacal aperture 854 from posterior end__-_-__~- Thelazia lutzi, p. 315.
Male smaller than above (up to 20 mm. in ZT. papillosa) ; female smaller
than above except in 7. papillosa (up to 26 mm.); in J. papillosa, as
in all other species where number is given, less than 10 pairs of preanal
papillae; cloacal aperture, if described, less than 854 from _ posterior
ONG ig = a ee ae ieee eer age sik Leu sae ee ie a ee 3.
8. 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 spicules ee eee sient eee See 5.
5. Esophagus not over 6004 long; anus of female 300u or farther from pos-
terior senso. 222 ee De ie SN pana So 6.
Esophagus 750 long or longer; anus of female not over 225u from posterior
Oe oe Se a a ee ae LOR a erence a RE ie
6. Maximum length of esophagus 340u; vulva 680u from anterior extremity.
Thelazia stereura, p. 316.
Length of esophagus 460 to 6004; vulva not over 500.4 from anterior extremity.
Thelazia campanulata, p. 313.
. Female with buceal cavity 334 deep; esophagus 9254 long; anus 85 from
=]
OOS CTO OTN Ca a Bee ee eg Thelazia cholodkowskii, p. 313.
Female with buccal cavity not over 284 deep; esophagus not over 850u
long-"anus' 120 orjferther fron’ posterior ends. ss. ee eee 8.
8. Female 18 mm. long; anus 120u from tail end_____ Thelazia digitata, p. 319.
Female not over 14 mm. long; anus 255 from tail end.
Thelazia dacelonis, p. 514.
NEMATODE PARASITES OF BIRDS 313
THELAZIA CAMPANULATA (Molin, 1858) Railliet and Henry, 1910
Synonyms.—Filaria campanulata Molin, 1858; Filaria falconis
magnirostris Molin, 1858.
Hosts——Primary: Falco magnirostris (Rupornis magnirostris) ;
secondary: Unknown.
Location.—Under nictitating membrane.
Morphology.—T helazia (p 311): Anterior extremity of body atten-
uated, obtuse. Cuticular surface annulated, the annulations compan-
uliform and imbricated. Buccal cavity (fig. 3742) 17 to 11p deep,
according to Travassos, by 28 to 35 wide. Esophagus subcylindri-
cal, 460 to 600n long.
Male 17 mm. long by 400% wide. Caudal extremity (fig. 374)
curved toward the ventral surface; cloacal aperture 140 from end.
rT]
a
re
TEL rH
f et tH A
<
cia an
05/1
ee
OSMM
lig. 874.—THELAZIA CAMPANULATA. @, HEAD END; 0, MALE TAIL} C, FEMALB
TAIL. AFTER TRAVASSOS, 1918
Seven pairs of preanal papillae and 1 unpaired median papilla
anterior to cloacal aperture; posterior to the cloacal aperture 1 large
median papilla and 38 pairs of papillae. Spicules very dissimilar
and unequal, one 190p long by 28» wide, alate laterally at its distal
extremity, the other 2.67 mm. long by 7p wide, with a slight swell-
ing 6402 from the proximal end.
Female 15 to 23 mm. long by 500 to 600% wide. Tail (fig. 374c)
nearly straight, pointed. Vulva about 350u 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.
Location.—Kye.
Morphology.—tT helazia (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, 470u 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 2204 from anterior end of body. Vulva
500u from anterior end of body. Oviduct 90n wide. Anus 854 from
tail end (fig. 875¢c). Eggs 58u by 30z.
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 leach; secondary: Unknown.
Location.—Conjunctival sac.
NAN
eet
Pm eS
iD i Ae
Itc. 375.—THELAZIA CHOLODKOWSKII. FEMALE. ad, HEAD END; JD,
CUTICULAR ANNULATIONS ; C, TAIL. AFTER SKRJABIN, 1922
Morphology.—tT helazia (p. 311) : Body tapering toward both ends;
anterior 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 long, cup-shaped. Esophagus 750
to 825u long, club-shaped.
Male 10 to 11 mm. long by 390 to 400u wide. Tail (fig. 376¢)
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 21p wide at
proximal end and 10» wide at distal end, the other 180p long by 30.
wide with its proximal end funnel-shaped and distal end rounded.
Female 11 to 14 mm. long by 500p wide. Vulva (fig. 376a) 525
to 555y from anterior end of body. Vagina 1mm. long. Anus 255
from tail end (fig. 367b), 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.—Kye?
WT
PET rrr cre Sl “
K >? TE ptr) 77s" aes ie > :
esi aN Soa uk as ais es wy BieMe tf fei hss
Fic. 376.—THELAZIA DACELONIS. a, HEAD END, SHOWING VULVA; 0, TAIL OF
FEMALE; C, TAIL OF MALE. AFTER BREINL, 1913
Morphology.—T helazia (p. 311): Transverse striations of cuticula
strongly developed (877a). Buccal cavity 35 to 494 wide by 28.
deep. Esophagus slightly claviform, about 670 to 740» long by 42
to 49u wide.
Male 22 mm. long by 400 to 500” wide. Caudal extremity (fig.
378) curved ventrally. Ten pairs of preanal papillae, 1 unpaired
Cl ‘ wt ann
Gi: i HH | it
AMTTTTTTATT
LT Ll ee ae
Fic. 377.—THELAZIA LUTZI. FEMALE. @, HEAD END, SHOWING VULVA; 0, TAIL.
AFTER TRAVASSOS, 1918
papilla anterior to cloacal aperture; 2 pairs of postanal papillae
and 1 unpaired papilla. Spicules unequal and dissimilar; one 740.
long by 7p wide, the other 190 long by 35h wide. Cloacal aperture
85 from the tail end, the latter obtuse and rounded.
Female 25 to 27 mm. long by 500 to 800n wide. Vulva (fig. 377a)
530 from head end, salient. Anus 100y from tail end (fig. 8770),
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.
OF 17177
Fic. 378.—THELAZIA LUTZI. MALE TAIL. AFTER TRAVASSOS, 1918
Hosts—Primary: Buteo vulgaris, B. vulpinus, Falco naevius
(=Aquila 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
Wie. 379.—THELAZIA STEREURA. FEMALE. @, HEAD END, SHOWING
VULVA; 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 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 51h deep by 68» wide in large specimens.
Esophagus 340, long. Tail (fig. 3795) straight, acutely conical,
prolonged into a styloid appendage with a rounded knoblike end;
NEMATODE PARASITES OF BIRDS 317
anus at base of the appendage, 3704 from tip of tail. Vulva (fig.
379 a) in anterior part of body, 6804 from anterior end in large speci-
mens, at a level just posterior to the middle of the esophagus. Eggs
oly by 25p.
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.—sS piroptera papillosa Molin, 1860b; Spiroptera fal-
conis leptopodis Molin, 1860; Spiroptera falconis gavial realis Molin,
1860; Spiroptera 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.—T helazia (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 300n 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 5004 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 history—Unknown; probably involves intermediate stages in
other hosts.
Distribution.—South America (Brazil).
THELAZIA ANOLABIATA (Molin, 1860) Railliet and Henry, 1910
Synonyms.—S piroptera anolabiata Molin, 1860); Spiroptera
cracis alectoris Molin, 1860; Filaria anolabiata (Molin, 1860) Stos-
sich, 1897; (? Oxyspirura) anolabiata (Molin, 1860) 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.—T helazia (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 4004 wide. ‘Tail straight, abruptly
conical, with blunt curved tip.
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution —South America (Brazil).
380
Fies. 380—382.—380, THELAZIA PAPILLOSA. HEAD; FRONT VIEW. AFTER
DRASCHE, 1884. 381, THELAZIA ANOLABIATA. HEAD, AFTER DRASCHE,
1884. 3882, THELAZIA (7?) CIRRURA. HEAD END, SHOWING VULVA.
ORIGINAL
THELAZIA (7?) CIRRURA (Leidy, 1886) Railliet, 1916
Synonym.—F daria cirrura Leidy, 1886.
Hosts—Primary: Quiscalus major (=Megaquiscualus major) ;
secondary: Unknown.
Location.—Orbit of eye.
Morphology.—T helazia (¢) (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 500n wide. Esophagus 863» long. Cau-
dal extremity slightly curved or nearly straight, conical, obtusely
rounded. Vulva (fig. 382) about 400, from anterior end of body.
Life history—Unknown; probably involves intermediate stages
in other hosts. s
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 (7?) DIGITATA Travassos, 1918a
Hosts.—Primary: Ramphastus, species; secondary: Unknown.
Location— Kye.
Morphology.—T helazia (p. 311): Anterior extremity digitiform.
Cuticula transversely annulated. Buccal cavity about 21h deep by
28u wide. Esophagus subcylindrical, about 850 long by 87p wide.
Male unknown.
é ii i el
———
m
05MM
LUTE]
‘b.
fil
Fic. 388.—THELAZIA (?) DIGITATA. FEMALE. a, HEAD END, SHOWING VULVA;
b, TAIL. AFTER TRAVASSOS, 1918
Female 18 mm. long by 500n wide. Vulva (fig. 383 @) 600» from
anterior extremity, its lips salient. Caudal extremity (fig. 383 6)
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. J/ale 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. Memale with very short blunt tail. Vulva in anterior
part of body. Sometimes viviparous. Parasitic in orbital cavity
of birds.
Type species —Ceratospira vesiculosa Schneider, 1866.
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' 9684 in\Jength?! 2ie2s2s2-2 5" spss 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 (lectus 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.—Ancyracanthus ophthalmicus Linstow, 1898.
Hosts—Primary: Carpophaga brenchleyi (=Zonoenas brench-
leyz) ; secondary: Unknown.
Location.—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 13u
wide, with swollen posterior edges.
Male 14.6 mm. long by 3902 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. 3850).
Spicules unequal, the right one short (264) and broad, the left long
(968) and slender.
Female 18 mm. long by 470n wide. Esophagus 1/23, tail 1/54, of
body length. Vulva near anterior end of body, about at the middle
NEMATODE PARASITES OF BIRDS Son
of the esophagus, dividing the body length in the ratio of 1:45.
Eggs 23 by 16, very numerous.
Life history—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
iS
RSS
S— -#
=
as
hs
LSS
SS wala
WS ZI
max Ah?
RSs ZA
SSS ZF
row
rh ne
Pe ZA
MS
Ss s
aS Ad
r
as TE
cS [Au
ens AK
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p59)
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Yl
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Fics. 384-385.—384, CERATOSPIRA VESICULOSA. MALE TAIL. AFTER SCHNEIDER,
1866. 385, CrRATOSPIRA OPHTHALMICA. @, HEAD END; b, MALE TAIL, AFTER
LINSTOW, 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. /ema/e with straight tail. Vulva
in posterior part of body, a short distance anterior to the anus.
Parasitic under nictitating membrane in birds.
Type species—Oxyspirura cephaloptera (Molin, 1860) Stossich,
1897.
KBPY TO SPRCIES OF OXYSPIRURA
1. Head with cuticular expansions, at least in the female_--_-~------~---- 2A
Head without cuticular expansions, or none described, or inconstant (in
<4
eo.
O. parvovum sometimes present in male) ~---~----------------------
322 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
2. Cuticular expansions absent in male, 4 in number in female in lateral,
1
10.
ek
Synonyms.
dorsal, and ventral fields; 2 pairs of postanal papillae in male.
Oxyspirura heteroclita, p. 331.
Cuticular expansions 2 in number in both sexes, situated laterally; more
than 2pairs“of“postanals papillae a= se ee ere oe
. 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.
. 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
preanall papillae cs se sa a Ses ee ee Oxyspirura cephaloptera, p. 322.
. 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. par-
vovum) ; female more than 9 mm. long (may be as little as 9.8 mm. in
O. parvovum) ; from hosts other than above_____----_---+--_----_---- 6.
. Spicules nearly equal in length and of same shape; position in genus ques-
ERGpT EAI) | @ ecetermsnae ee ea el Sw a Oxyspirura brevipenis, p. 332.
Spicules of unequal length and dissimilar shape_~_-4_~-----__--_-_-+- this
. Female 9.8 mm. long; male with about 28 pairs of preanal papillae; spicules
470u and 250u fone cl Sees SS AA 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
. Female 21 mm. long; male with about 26 pairs of preanal and 1 pair ot
postanal papillae; spicules 180% and 290. 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 lengthsidifferent: fromaboyes. 28 sna SE o}
. Preanal papillae apparently absent; 4 pairs of postanal papillae; left
spicule 2% 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 21% times the length of right spicule-____-______________- 10.
Mouth surrounded by a 6-lobed chitinous ring; 4 pairs of preanal papillae;
eggs 50 to 65u by 40 to 454_-----__------_--- Oxyspirura mansoni, p. 325.
Mouth not surrounded by a 6-lobed chitinous ring; not more than 3 pairs
of preanal papillae; eggs not over 46h by 294_-----_-------------_----- el
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 440u 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 600u long; anus of female 5104 from
posterior end s-e3 eae ee a ae Oxyspirura tanasijtchuki, p. 330.
OXYSPIRURA CEPHALOPTERA (Molin, 1860) Stossich, 1897
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 (—=Momotus momota)
and Jcterus croconotus; secondary: Unknown.
Location.—Under nictitating membrane.
Morphology.—Oxyspirura (p. 821): Body gradually attenuated
anteriorly, with rounded apex. ‘Tail awl-shaped, drawn out into a
slender acute point. Cuticula with dense transverse striations. Head
(fig. 386a) 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 200u wide. Tail (fig. 386 6) 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 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, 1860) Stossich, 1897
Synonyms.—Spiroptera anacanthura Molin, 1860; Spiroptera
crotophagae ani Molin, 1860; Spiroptera crotophagae majoris Molin,
1860.
Hosts—Primary : Crotophaga ani, C. major; secondary : Unknown.
Location —Under nictitating membrane.
Morphology.—Oxyspirura (p. 821): 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 Jarge, 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.
Male 8 to 10 mm. long by 100p wide. Caudal end (fig. 8872) coiled
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. 8876) 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 2002 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: Anthochaera carunculata; secondary: Un-
known.
3868.
\Wa
x
|
|
——
WOMMA
Fies. 386—388.—386, OxYSPIRURA CEPHALOPTERA. 4, MALE TAIL; 0, HEAD, FRONT
VIEW. AFTER DRASCHE, 1884. 387. OXYSPIRURA ANACANTHURA. @, RIGHT SPICULE ;
b, MALP TAIL. AFTER DRASCHE, 1884. 388, OxYSPIRURA ANTHOCHAERAE. ad, HBAD
END; b, TAIL END OF FEMALE. AFTER JOHNSTON, 1912
Location.—Kye.
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. Cloacal aperture 80» from
posterior extremity, which is spirally curled. Caudal papillae not
discernible.
Female 8 to 9 mm. long by 110% wide. Posterior extremity sharply
pointed. Anus 194y, vulva 320, from posterior end. (Fig. 388 0.)
Life history—Unknown; probably involves intermediate stages
in other hosts.
Distribution. —A ustralia.
NEMATODE PARASITES OF BIRDS 325
OXYSPIRURA BREVISUBULATA (Molin, 1860) Stossich, 1897
Synonyms.—sS piroptera brevisubulata Molin, 1860; Spiroptera
strigis Molin, 1860.
Hosts—Primary: Strix atricapilla (=Otus choliba) ; secondary:
Unknown.
Location.—Under nictitating membrane.
Morphology.—Cxyspirura (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 300, 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 500n 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-
ezu Emmerez and Mégnin, 1901; Spiroptera mansoni (Cobbold,
1879) Gedoelst, 1903.
Hosts —Primary: Gallus gallus, Meleagris gallopavo, 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 (fig. 389 a and 6) 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 @ 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 10u 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 4302 wide. Anus 400 to 530, from
tip of tail (fig. 390). Vulva 1 to 1.4 mm. from tip of tail. Vagina
FOX
Fic. 389.—OxXYSPIRURA MANSONI. @, DORSAL VIEW AND Bb, FRONT VIEW OF HBAD; C,
EGG; d, EMBRYO ESCAPING FROM THE EGG. (€.0.r., CIRCUMORAL CUTICULAR RING;
€s., ESOPHAGUS; l.p., AMPHIDS OR SO-CALLED “ LATERAL PAPILLAE”; m, MOUTH;
ph., PHARYNX; 8.l.p., SUBLATERAL PAPILLAE; 8.m.p., SUBMEDIAN PAPILLAE.) AFTER
RANSOM, 1904
1.5 to 2 mm. long. Eggs (fig. 389 ¢ and d) oval, 50 to 65 by 45
(Smit says 40u long by half as wide; see 0. parvovum, p. 328).
Larva, first stage, 225 to 250u long by 124 wide. Esophagus 50p
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 Owy-
4 7777
Fics. 390—391.—OxXYSPIRURA MANSONI. 390, POSTERIOR END OF FEMALE. (int., INTESTINE ;
0v., OVARY; 1rct., RECTUM; vg., VAGINA; vul., VULVA; @., CELLS SURROUNDING RECTUM.)
391, POSTERIOR END OF MALE. @, LATERAL view. (dct. ej., EJACULATORY DUCT; int.,
INTESTIND; 8p., SPICULES ; U.8., SEMINAL VESICLE.) 0b, 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 host 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, 1908a.
Hosts—Primary: Turnix taigoor; secondary : Unknown.
Location.—Kye.
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 42u wide; that is, about 1/24 of circumference of body.
Male 14.4 mm. long by 310n wide. Esophagus 1/16 of body length.
Tail 1/52 of body length finely pointed (fig. 392 6). 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 394 by
26n, 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 O. 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 300n wide. Tail (fig. 3936)
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 134 wide along most
NEMATODE PARASITES OF BIRDS 329
of its length and 24 to 30u wide at the base; short spicule 180 to
240u 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 780u 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 45h by 25 to 30u, rather square-ended, con-
taining embryos when oviposited.
Life history—Unknown. See remarks on O. mansoni, p. 326.
Figs. 392-393.—392, OXYSPIRURA OPHTHALMICA. ad, HHAD; b, MALE TAIL. AFTER
Linstow, 1903. 393, OXYSPIRURA PARVOVUM. a, HEAD; JD, MALE TAIL, AFTER
SWEET, 1910
Distribution.—Australia. The worms reported by Smit from the
Dutch Indies have egg measurements in agreement with this species,
40p by 202; he calls them O. mansoni, but they differ in some re-
spects from both O. mansoni and O. parvovum.
OXYSPIRURA SIAMENSIS (Linstow, 1903) Ransom, 1904
Synonym.—C heilospirura stamensis Linstow, 1903a.
Hosts —Primary: Centropus siamensis (=Centropus sinensis) ;
secondary: Unknown.
Location.—Probably the eye, according to Linstow.
Morphology—Oxyspirura (p. 321): Cuticula with fine transverse
striations. Buccal cavity small.
Male 8.9 mm. long by 260p 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 250p 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, 260, 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.—s piroptera anthuris Rudolphi, 1819, of Diesing, 1851,
part; Spzroptera sygmoidea Molin, 1860); 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.
Fic. 394.—OXYSPIRURA PARVOVUM. POSTERIOR END OF FEMALE. AFTER SWEET, 1910
Male 11 mm. long by 400u 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 21% times as long as the right.
Female 11 to 15 mm. long by 500n 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. Ouxyspirura (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 220n wide. Esophagus 518» long. Caudal
extremity (fig. 397c) tapering to a fine point. Right spicule (fig.
3976) 250p 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 320n wide. Esophagus 765p long. Anus
510u from end of tail. Vulva in posterior part of body. Eggs 46u
by 29.
Life history—Unknown; probably involves intermediate stages in
other hosts.
Distribution.—South America (Paraguay).
(?0XYSPIRURA) 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.—Oxyspirura (p. 321): Body attenuated anteriorly,
apex truncate. Cuticula with fine transverse striations. Mouth or-
bicular, large, without lips.
Male 12 mm. long by 300z 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% 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).
302 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
(?70XYSPIRURA) BREVIPENIS (Molin, 1860) Stossich, 1897
Synonyms.—Spiroptera brevipenis Molin, 18606; Spiroptera mi-
crodactyli margravii Molin, 18606.
Hosts—Primary: Dicholophus margravi (=Cariama cristata) ;
secondary: Unknown.
Location —Under nictitating membrane.
Morphology —?Oxyspirura (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, OXYSPIRURA SIAMENSIS. MALp TAIL. AFTER LINSTOW, 1903.
396, OXYSPIRURA SYGMOIDEA. MALE TAIL. AFTER SKRJABIN, 1916. 397, OxYSPI-
RURA TANASIJTCHUKI. a, HEAD; b, RIGHT SPICULP; C, MALE TAIL. AFTER SKRJABIN,
1916. 398, OXYSPIRURA BREVIPENIS. MALE TAIL. AFTER DRASCHE, 1884
Male 16 to 22 mm. long by 400u wide. Tail (fig. 898) 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» 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.
rs
NEMATODE PARASITES OF BIRDS gaa
Family TETRAMERIDAE Travassos, 1914
Family diagnosis —Spiruroidea (p. 162): Characterized by the
sexual dimorphism of the species. J/ale 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. /'emale red; body greatly
enlarged in comparison with that of male, its shape either globular,
with the two pointed extremities projecting (Z'etrameres), or the
long axis of the body coiled in a more or less complex manner (J/7-
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.—T etrameres Creplin, 1846.
KEY TO GENERA OF TETRAMERIDAB
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, 19156 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 7Z'etrameres (sensu lato), at least in the
present state of knowledge. 7’. 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
24 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 Z'etra-
meres (sensu lato) to one of the two genera, Z'etrameres or Microte-
trameres.
Genus TETRAMERES Creplin, 1846
Synonyms.—Tropisurus Diesing, 1835; Tropidurus Wiegmann,
1835; T'ropidocerca Diesing, 1851; Astomum 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.—Tetrameres paradoxa (Diesing, 1835) Travassos,
1914.
KEY TO SPECIES OF TETRAMERES
1. From Fulica atra; anus 2204 from posterior extremity; eggs 394 long by
DOL WAG Gee nena Rina skew vt ME Meares ere Senet Tetrameres globosa, p. 346.
From other hosts than Fulica atra, or if from that host, anus T1ly from
posterior extremity and eggs 48 to 56u long by 26 to 30u wide (Tetra-
MORES ATESSUS DING) oF a st a ee et Ee ee ee De
. Mouth of female surrounded by 8 projections, each of which bears a thorn
on its outer and on its inner surface; male unknown, female poorly
described; in Corvus cornig_.-.__--.-----_- 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
bo
CHAN OOTVUS CONE eee ae ane Ac IO Ease a Oe Ee 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_.-i-._=/-=:---+-++-=- 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 ald, mim, longs 3rd ht ey ete Hee he SEE Des pce eee 5
Female not over 4.5 mm. long (except in 7. confusa and T. fissispina
which may reach 5 and 6 mm. long respectively) ; male, when known,
not:-over “63mm, Tonge Ss ee ee Seer ete CREE eS eee es VEC eens eee ee (
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 468u 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 7404 and 16u long respectively.
Tetrameres gigas, p. 345.
Female 8 mm. long by 7 mm. wide, or larger; male without spines; spicules
saidito béeypabsen tesa 2 en oes ee Tetrameres gynaecophila, p. 347.
7. Anus of female 332u, cloacal aperture of male 232 to 290u from posterior
TN ee See ea gino Raleatend Ras De ae Tetrameres americana, p. 337.
Anus of female and cloacal aperture of male both nearer to the posterior
end) than a@bove.22.22. 2 ee 8.
«all
NEMATODE PARASITES OF BIRDS 385
8. Female with pharynx 400”, esophagus 2 mm. long; anus 2504 from posterior
end; cloacal aperture of male 70yv from posterior end.
Tetrameres confusa, p. 341.
Both pharynx and esophagus of female shorter than above, if described ;
anus, if described, not more than 175u from posterior end ; cloacal aperture
of male, if described, 100u or farther from posterior end__________ 9.
OOnly temale knOwWn2222 2525s ee ee 10.
oth male and. temale; KWOwmns 2-22 es ee ee i
10. Female 2.2 mm. long by 2.5 mm. wide; pharynx 350u, esophagus 1.4 mm.
long ; eggs 28 to 30u long by 15 to 184 wide___ Tetrameres coccinea, p. 339.
Female 3 to 4 mm. long by 1.5 to 2 mm. wide; pharynx 190u, esophagus 990u
long; eggs 42 to 49u long by 21u wide__-_- Tetrameres cochleariae, p. 340.
11. Male 2.2 to 2.6 mm. long; female not over 1.7 mm. long________________ 12,
Male larger than above or (7. dubia) smaller than above; female 2 mm.
COU TTL ETD LT) Oe a ee ee ae
12. Eggs 54 by 28 wide, with long filaments at each pole; one spicule 480” in
length, the other rudimentary________________ Tetrameres nouveli, p. 348.
Eggs 75 to 7T8u long by 214 wide, with no filaments described ; spicules 200z
and 22» long respectively__ _-______-_-______-__- Tetrameres tetrica, p. 350.
13. Female not over 2.5 mm. long; male 1.6 mm. long, its esophagus being 300u
long; long spicule 720u in length_____________- Tetrameres dubia, p. 342.
Female 2.5 to 6 mm. long; male 3 to 6 mm. long, its esophagus 78SOu or more
in length; long spicule not over 490 in length______-_____--__-_-_--- 14.
14. Ovejector with diverticulum or copulatory receptaculum 4004 long; eggs
not over 56u long; short spicule at least 82u long.
Tetrameres fissispina, p. 343.
Ovejector simple, without diverticulum; eggs 59 to G8 long; short spicule
Eyebeam 82S ee ee Tetrameres micropenis, p. 348.
This key does not include Tetrameres zakharowi Petrow, 1926. See Addenda,
p. 385.
TETRAMERES PARADOXA (Diesing, 1835) Travassos, 1914d
Synonyms.—T ropisurus paradowus Diesing, 1835; T'ropidocerca
paradoxa (Diesing, 1835) Diesing, 1851.
Diesing’s description of 1851 is evidently a composite description
as in 1861 he makes 7’ropidocerca paradoxa Diesing, 1851 in part a
synonym of his new species 7'ropidocerca inflata, and also transfers
the synonomy given originally under 7'ropisurus paradoxus (that is,
Spiroptera inflata and Tetrameres haemochrous) to Tropidocerca
inflata. 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, Cathartes 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 the 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 7’. fisstspina (p. 348).
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 9994 wide; see discussion under
size of female); according to Linstow (1879) 11 to 13.5 mm. long
by 750p wide. Body subcylindrical, threadlike, curved to crescent
shape, white, pointed at both ends. Drasche’s figure of the posterior
extremity (fig. 4006) 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
Fic. 399.—TmTRAMERDS PARADOXA. @, FEMALE, NATURAL SIZE; 6b, 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 187.)
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
(480n), the left one 8 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 214 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, Jongi-
NEMATODE PARASITES OF BIRDS 337
tudinal furrows (fig. 3996 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 7.
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 7. fissispina (p. 348).
Fic. 400.—TETRAMERES PARADOXA. a, HEAD (WHETHER OF MALE OR FEMALE NOT
STATED) ; J, MALE TAIL. AFTER DRASCHE, 1884
Location —Proventriculus.
Morphology—tTetrameres (p. 834): Mouth with 3 small lips; bue-
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 116 to 133” wide. Two double rows of
posteriorly directed spines extend throughout whole body length, in
the submedian lines. Buccal capsule (fig. 402a@) 27 deep by 4.5
Fic. 401.—TETRAMERES AMERICANA. POSTERIOR END OF MALE. ORIGINAL
wide; pharynx 365p long; esophagus 996u 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, cloacal aperture 2382 to 290. from
posterior end. Two unequal spicules, 100% and 290 to 312 long
respectively, not heavily chitinized.
Female 3.5 to 4.5 mm. long by 3 mm. wide; body globular (fig.
402f), blood-red in color (when alive), with 4 longitudinal furrows
corresponding to the lateral and median lines. ‘The anterior ex-
tremity (fig. 402¢) protrudes from the globular body for a length
338 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
of 913; the protruding posterior part of body is 860y long. Buccal
capsule 35 to 38~ long by 10p wide; pharynx 300 to 315p long by 27
wide; esophagus 1.4 mm. long by 50% minimum and 1254 maximum
width. Nerve ring 183, from cephalic extremity. Intestine saccular,
filled with black detritus. Anus 382, vulva 631 to 664» from caudal
extremity. Vestibule of ovejector with diverticulum or copulatory
receptaculum (fig. 402¢) 274, long. Uteri and ovaries very long,
their numerous coils filling the body cavity. Eggs 42 to 50u by 25p.
The American form of Z'etrameres from the chicken, as described
above, differs from the two forms reported from other countries from
—
O1mM
Fic. 402.—T@rrRraAMERES AMERICANA. dd, HAD OF MALE; D, JUNCTION OF TESTIS
AND VAS DEFERENS; ¢C, HEAD END OF FEMALE; d, JUNCTION OF UTERUS AND
OVARY ; €, COPULATORY RECEPTACULUM OF FEMALE; f, CROSS-SECTION OF FEMALE.
ORIGINAL
that host, 7’. fissispina and 7. 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—Probably similar in a general way to that of 7’.
fisstspina (p. 348).
Distribution —(North America (United States) ).
Type material —No. 26382, U.S.N.M. (Bureau of Animal Industry
Helminthological Collection).
TETRAMERES CERTA (Leidy, 1886) Travassos, 1914 .
Synonyms.—Filaria dubia Leidy, 1856, not Filaria dubia Crephn,
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 7'etra-
meres certa would have to fall if the older specific name dubia of
Leidy is available and in consequence 7'etrameres dubia Travassos,
1917, would fall as a homonym. ‘To avoid this needless rearrange-
ment I am interpreting /ilaria dubia Creplin as a good combina-
tion and aria dubia Leidy as therefore a homonym.
Hosts.——Primary: Albatross (iomedia exulans) ; secondary : Un-
known, probably similar in a general way to that of 7’. fissispina
(p. 343).
Location —Glandular stomach or proventriculus.
Male 12 to 18 mm. long by 375 to 5004 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 Zetrameres 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 7.
Jissispina (p. 343).
Distribution —South Atlantic.
TETRAMERES COCCINEA (Seurat, 1914) Travassos, 1914d
Synonym.—tTropidocerca coccinea Seurat, 1914m.
Hosts—Primary: Phoenicopterus roseus, Bubulcus lucidus, and
Platalea leucorodia; secondary : Unknown, probably similar in a gen-
eral way to that of 7’. fissispina (p. 348).
Location.—Glandular stomach or proventriculus.
Morphology.—tTetrameres (p. 334).
Male unknown.
Female 2.2 mm. long by 2.5 mm. wide. Body globular (fig. 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. 4036) circular, 20» 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, 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, filled with brown-black detritus. Anus
at a short distance from tip of tail. Vulva 480, anterior to anus.
Ovejector (fig. 403c) remarkable for the shortness of vestibule and
sphincter; they form a turnip-shaped organ 450» long. Sphincter
very straight. Trompe, Y-shaped, almost 1 mm. total length, the
340 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
unpaired branch being 350p 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. 403d), measuring 660% long by
215. wide. Diameter of oviduct only 35y; 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 7’.
fissispina (p. 348).
Distribution.—Africa (Algeria) and Asia (Russian Turkestan
(Lac Kulkainar) ).
Fies. 403—404.—403, TETRAMERES COCCINEA. FEMALE. a, TOTAL; Db, ANTERIOR EX-
TREMITY; ©, OVEJECTOR; d, SEMINAL RECEPTACLE AT JUNCTURE OF UTERUS AND
ovipucT. Arrer SeEuRAT, 1914. 404, TeTRAMERES COCHLEARIAE. OVEJECTOR.
AFTER TRAVASSOS, 1919
TETRAMERES COCHLEARIAE Travassos, 1917a
Synonym.—T etrameres micropenis Travassos, 19155, part.
Hosts—Cancroma cochlearia; secondary : Unknown, probably sim-
ilar in a general way to that of 7. fissispina (p. 348).
Location.—Proventriculus.
Morphology.—Tetrameres (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 28u
in height, 14 to 16 in maximum width; pharynx 190» long; esopha-
gus slightly claviform, 990pu long; nerve ring 140u 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, 1202 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 49u
long by 21p wide.
NEMATODE PARASITES OF BIRDS 341
Life history.—Probably similar in a general way to that of 7.
fissispina (p. 843).
Distribution.—South America (Brazil).
TETRAMERES CONFUSA Travassos, 1919b
Synonyms.—Tetrameres fissispina Diesing, 1861 of Travassos,
1914a; Tetrameres travassosi Skrjabin, 1920.
Hosts——Primary: Columbia livia domestica, Gallus gallus and
Meleagris gallopavo; secondary: Unknown, probably similar in a
general way to that of 7’. fissispina (p. 343).
Location —Glandular stomach or proventriculus.
Morphology.—tT etrameres (p. 334).
Male 4 to 5 mm. long by 160u wide. Body filiform, with 4 rows
of spines corresponding to the median and lateral lines; spines di-
Fic. 405.—TETRAMERES CONFUSA, a, FEMALE; Db, MALE TAIL. AFTER TRAVAS-
sos, 1919
rected posteriorly and about 100 apart in middle portion of body
and 20u in posterior portion. Tail ends in a hook about 4y long.
Buccal capsule cylindrical, 24y long by 802 wide; pharynx 250, long
by 124 wide; esophagus 740» long by 55 wide. Cloacal aperture
70» from end of tail (fig. 405 6). Short spicule 68. long and long
spicule 291 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 long. Buccal capsule 20u long by 14» wide;
pharynx 400, long by 30p wide; esophagus sinuous, 2 mm. long by
180 wide; intestine dilated to form a large sac, filled with black
detritus. Anus 250, 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» long by 24» wide.
Life history—Probably similar in a general way to that of 7.
jissispina (p. 348).
Distribution—South America (Brazil). —
TETRAMERES DUBIA Travassos, 1917a
Hosts—Primary: Gallinago paraquaiae, secondary: Unknown;
probably similar in a general way to that of 7. fissispina (p. 348).
Location.—Proventriculus.
Morphology.—T etrameres (p. 334).
Male 1.6 mm. long by 85 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
Fic. 406.—TETRAMERES DUBIA. 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 240u
long; esophagus 300u long. Anus 100n from posterior extremity.
Tail sharply pointed with 4 pair of very small spines on the ventral
surface. Spicules of very unequal dimension, measuring respectively
720u long by 6 wide and 64, long by 6u 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» wide. Anus 75 from pos-
terior extremity. Ovejector 420 long, of same type as that of 7’.
micropenis, the dilated part full of eggs. Eggs 494 long by 35y
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 7’.
fissispina (p. 343).
Distribution—South America (Brazil).
TETRAMERES FISSISPINA (Diesing, 1861) Travassos, 1914
Synonyms.—T ropidocerca fissispina Diesing, 1861; Tropisurus fis-
sispinus (Diesing, 1861) Neumann, 1888; Acanthophorus tenuis
Linstow, 1876; Acanthophorus horridus Linstow, 1876; 7'ropidocerca
tenuis Linstow, 1899; Filaria pulicis Linstow, 1894; Spiroptera puli-
cis (Linstow, 1894) Linstow, 1909. The nematode described by
Travassos in 1914 as 7. fiss¢tspina is 7. confusa.
Hosts—Primary: Anas boschas, Anas boschas domestica, Anas
boschas fera, Columba livia domestica, Cygnus melanocoryphus,
Fulica atra, Meleagris gallopavo, Merqus 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 7’. fisstspina.
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.—tTetrameres (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 7’. jfissispina; 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 (1918) 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. 407¢ and 6.) He
states that it is to the existence of these bifid cephalic spines that
the name jisstspina 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 84 deep by 3» wide, esophagus 780,
long by 52u wide, cloacal aperture 130 (250u according to Wharton)
from posterior end. The tail ends in a spine 44 long. The spicule
Fies. 407-409.—TETrRAMERES FISSISPINA. 407, a@ AND b, HEAD END
OF MALE, LATERAL VIEW (@), DORSAL VIEW (Db), SHOWING THE TWO
BIFID CEPHALIC SPINES AND THE SIMPLE BODY SPINES ; Cc, HEAD END
OF FEMALE, DORSAL VIEW. AFTER SEURAT, 1920. 408, HEAD AND
TAIL OF MALE. AFTER TRAVASSOS, 1919. 409, a, OVETECTOR; Db,
COPULATORY RECEPTACULUM ; C, FEMALE GENITAL SYSTEM. AFTER
SEuURAT, 1914. d, TAIL OF LARVA, VENTRAL VIEW. AFTER SEURAT,
1919
lengths as given in various descriptions are 150 and 320n (Seurat),
115 and 820% (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. 407¢c) 21 long by 10» wide; pharynx
2304 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» from the end of the tail.
The vulva is 310p from the end of the tail. It connects with a very
short vestibule (fig. 409 a, , 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 réle 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
long by 26 to 80 wide and containing well developed embryos when
deposited.
Larva, fourth stage, 1.65 mm. long, straight and slender (female) ;
tail (fig. 4092) long with rounded tip bearing 8 (?5) long spines
(Seurat says 8 and figures 5) and with 2 latero-ventral spines 70p
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-
marus 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 7’. americana (see p. 337).
TETRAMERES GIGAS Travassos, 1919b
Synonym.—T etrameres 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 7’. fissispina (p. 343).
Location — Glandular stomach or proventriculus.
Morphology.—tT etrameres (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 144 wide. Pharynx 370p 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 504 long by 21p wide,
containing embryos when deposited.
Life history—Probably similar in a general way to that of 7.
jissispina (p. 343).
Distribution —South America (Brazil).
SE sects ren
Sern
Pepe ae
=
Itc. 410.—TR?TRAMERES GIGAS. ANTERIOR AND POSTERIOR ENDS OF MALE. AFTER
TRAVASSOS, 1919
TETRAMERES GLOBOSA (Linstow, 1879) Travassos, 1914d
Synonym.—T ro pidocerca globosa Linstow, 1879a.
Hosts —Primary: Fulica atra; secondary: Unknown; probably
similar in a general way to that of 7’. fissispina (p. 343).
Location.—Proventriculus.
Morphology.—T etrameres (p. 334).
Male unknown.
Female globular, its two extremities slender projections. Cuticle
transversely striated. Buccal capsule circular with chitinous walls.
Pharynx 280; esophagus 780p long. Anus 220, from posterior ex-
tremity. Eggs thick-shelled, 394 long by 29” wide.
Linstow states that this species differs from 7'etrameres 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 7’.
fissispina (p. 348).
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.—T ro pidocerca 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. 3438).
Location.—Proventriculus.
Morphology.—Tetrameres (p. 334).
Male 6.4 mm. (Seurat) or 10 mm. long (Molin) by 600u 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 21p 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-
no
tu
Fic. 411.—TETRAMERES GYNAECOPHILA. @, HPAD OF FEMALBP; 6b, TAIL OF MALE,
AFTER SEURAT, 1915
tum 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 (1802). Anus 140» from posterior extremity. Tail sharply
narrowed behind anus, digitiform, ending in a blunt point. Vulva
in neighborhood of anus. Ovejector Y-shaped; vestibule and sphinc-
ter joined to form cylindrical tube 1.26 mm. long, with thick walls;
348 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
\
\
unpaired trompe 6002 long. Eggs 47p 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 7.
jissispina (p. 343).
Distribution.—Africa (Algeria) and Europe (Italy (Padua) ).
TETRAMERES MICROPENIS Travassos, 1915b
Hosts—Primary: Nicticorax violacens and Cancroma cochlearia,;
secondary: Unknown; probably similar in a general way to that
of 7’. fisstspina (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. 384).
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, 28p
long; pharynx 355 long; esophagus more or less cylindrical, 1.3
ee ee
SEES > ——
Zs meet 2
Fic. 412.—T®TRAMERES MICROPENIS. a, TAIL OF MALE; Db, OVEJECTOR, SHOWING
STRIATED APPEARANCE OF 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 14y wide, oval in form; pharynx 250,» long;
esophagus 1.5 mm. long, cylindrical. Vulva slightly anterior to
anus. Ovejector (fig. 4120) simple, about 710» long, claviform and
longitudinally striated. Eggs 59 to 63 long by 39u wide.
Life history.—Probably similar in a general way to that of 7.
Jissispina (p. 348).
Distribution—South America (Brazil).
TETRAMERES NOUVEL! (Seurat, 1914) Travassos, 1914d
Synonyms.—T'ropidocerca nouveli Seurat, 19142.
Hosts —Primary: Himantopus himantopus; secondary: Un-
known; probably similar in a general way to that of 7’. fissispina
(p. 343).
Location.—Proventriculus.
Morphology.—tT etrameres (p. 334).
NEMATODE PARASITES OF BIRDS 349
Male 2.16 mm. long by 75 wide. Body slender (fig. 413e and
413f) 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
(102); pharynx 280, long, encircled in its posterior third by the
nerve ring. Combined length of pharynx and esophagus 1/3 that
of body. Cervical papillae 60 and 90p, respectively, from an-
terior end. Cloaca 110. 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
JOO
Fic. 413.—TETRAMERES NOUVELI. a, FEMALE; 0, OVEJECTOR; C, SEMINAL RECEPTACLE
AND OVIDUCT; d, EGG WITH FILAMENTS ; €, HEAD END OF MALE; f, TAIL END OF MALE.
(5004 SCALE FOR a@ TO c; 1004 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. 418a). The
strongly distended median region of body measures 1 mm. long by
8502 maximum width. Buccal capsule 154 long; pharynx 210,
long, encircled in its posterior third by the nerve ring. Combined
length of pharynx and esophagus 1 mm. Cervical papillae 77» and
87, respectively, from anterior end. Excretory pore ventral, 50,
posterior to nerve ring. Anus 175» from posterior extremity. Vulva
120u anterior to anus. Ovejector (fig. 4136) 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 with 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. 413¢), its trans-
verse diameter being 125». Oviducts and ovaries 4mm. long. Eggs
350 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
54y long by 28» 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.
413 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 7.
fissispina (p. 348).
Distribution —Africa (Algeria).
Su
: KB
SES
Ee Be
ge ia a egg
s PE tS aS Re E
i
Fig. 414.—TETRAMBERES TETRICA. MALE, AFTER TRAVASSOS, 1919
TETRAMERES TETRICA Travassos, 1917
Host.—Primary: Aramides cajanea; secondary: Unknown; prob-
ably similar in a general way to that of 7’. fissispina (p. 343).
Location.—Proventriculus.
Morphology —Tetrameres (p. 334).
Male 2.6 mm. long by 130 to 1404 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 (20u long by 3» 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
long by 6p wide, irregular in shape. Pharynx 230» long by 42 wide.
Nerve ring 1404 from anterior extremity of body. Cloacal aperture
200% from posterior extremity which is long and sharply pointed.
Two spicules of very different lengths, the larger 2004 long by 6yu
NEMATODE PARASITES OF BIRDS 351
wide and the shorter which is only slightly chitinized, 22” long by
4u 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, 162 deep by 12” wide. ‘Tail slender and
very long. Ovaries 21p wide. Uterus ending posteriorly in a large
rounded seminal vesicle, about 10 in diameter. Ovejector similar to
that of 7’. micropenis. Eggs 75 to 78u long by 21 wide; ellipsoidal.
Life history.—Probably similar in a general way to that of 7.
fissispina (p. 348).
Distribution—South America (Brazil).
TETRAMERES UNISPINA (Diesing, 1861) Travassos, 1914d
Synonym.—tT ropidocerca unispina Diesing, 1861.
Host.—Primary: Corvus cornix; secondary: Unknown; probably
similar in a general way to that of 7. fissispina (p. 3438).
Location.—Proventriculus.
Morphology.—T etrameres (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 7.
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 Z'etrameres; if
present, these furrows also spirally coiled.
Ty pe-species.—Microtetrameres cruzi (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
Microtetrameres.
KEY TO SPECIES OF MICROTETRAMERES
1. Species wrongly placed in this genus, apparently identical with Hustrongy-
Hides mero orwmn 22-2 Microtetrameres inflata, p. 357.
2. Anus of female 450” from posterior end; male 6 mm. long; longer spicule
O-4mMmns in length +. see So nee Microtetrameres contorta, p. 353.
Anus of female not over 2254 from posterior end; male not over 5 mm.
long; longer spicule not over 3.6 mm. in length-_-__-________-_-___---- 3.
ga2 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
3. Female 780u long by 6404 wide, its pharynx 73u long; spicules 990% and 100u
long, respectively 222-22. 5" = ees Sea Microtetrameres minima, p. 358.
Female 1.2 to 2.56 mm. long by 1 mm. or more in width; pharynx 160y or
longer’: spicute “lengths ‘ditferent: strom ab0ve_2-- =eae ee 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
2S mms imy Lem ee is se ita eg ee ee 5.
. Anus of female 2254 from posterior end; size of male 4.75 mm. by 130z;
spicules 2.3 mm. and 145y long, respectively.
Microtetrameres spiralis, p. 360.
Anus of female not over 140u from posterior end; size of male not over 4
mm. by 1204; long spicule not over 1.32 mm., short spicule not over
ANC) ee 2 Ta a Ta a a 6.
6. Anus of female 74 to 100u from posterior end; long spicule 651 to 787u long.
Microtetrameres cruzi, p. 352.
On
Anus of female over 100” from posterior end; long spicule over 1 mm__-_~ 7.
7’ Mate’$.5 to’4) mm: longs 2 ee eee Microtetrameres pusilla, p. 359
Male 2:13: mm.clong#2! see ee eee Microtetrameres inermis, p. 355.
MICROTETRAMERES CRUZI (Travassos, 1914) Travassos, 1915b
Synonym.—Tetrameres cruzi Travassos, 1914d.
Hosts—Primary: Bucco swainsoni and Melanerpes flavifrons;
secondary: Unknown; probably similar in a general way to that of
T. fissispina (p. 348).
Location.—Proventriculus.
Morphology.—Microtetrameres (p. 351).
Male 1.17 to 1.4 mm. long by 85p wide. Body threadlike, white,
strongly striated transversely but having no spines (fig. 4150).
Buccal capsule 12” long by 4 to 54 wide; pharynx 93p long by 9p
wide; esophagus 290 long by 244 wide. Anus 132» from posterior
end. Spicules of very unequal dimension, the small one 82y, the
larger 651 to 787p 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 cor responding
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 620y long. Intestine saclike, filled
with black detritus, narrows toward posterior end and terminates
in a fine canal. Anus 74 to 100u from posterior end. Eggs 50 to
60u long by 24 to 28 wide, many of them containing well-developed
embryos. Vulva 300u from posterior end.
Life history—Probably similar in a general way to that of 7.
jissispina (p. 348).
Distribution —South America (Brazil).
NEMATODE PARASITES OF BIRDS 353
MICROTETRAMERES CONTORTA (Weidman, 1913) Travassos, 1915b
Synonyms.—T ropidocerca contorta Weidman, 1913; Tetrameres
contorta (Weidman, 1913) Travassos, 1914.
Hosts——Primary: Dichocercus bicornis; secondary: Unknown;
probably similar in a general way to that of 7’. fissispina (p. 348).
Location. —Proventriculus.
Morphology.—M icrotetrameres (p. 351).
Fig. 415.—MICROTETRAMERES CRUZI. a, FEMALE; b, MALB. ABTER TRAVASSOS, 1914
Male 6 mm. long by 125 wide. Body subcylindrical, filiform
(fig. 416 @ and }); 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 3004 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 7); no longitudinal furrows as
354 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
in species of Zetrameres. 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 f) sharply pointed; anus 450p, vulva
$00 from the posterior end, cuticle thickened into a rounded swell-
ing between the two openings. Eggs 40 to 45 long by 20 to 25p
w-de, containing coiled embryos when mature; some eggs show
peculiar unilateral bib attached to outside of shell (fig. 416 ¢).
Fic. 416.—MICROTETRAMERES CONTORTA. @ AND Bb, 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 FHMALE, SHOWING COURSE OF BSOPHAGUS AND INTESTINE;
i, HALF GROWN FEMALE, COILED IN ONE PLANE ONLY. AFTER WRIDMAN,
1913. j, WAX RECONSTRUCTION OF FEMALE, AFTER WBHIDMAN, 1923
Weidman made a very careful study of females of varying ages
(fig. 416 ¢ to e and h/ to 7); one of the wax reconstructions made by
him is shown in figure 416 7. 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 7.
fissispina (see p. 343).
Distribution —North America (United States (Pennsylvania
(Zoological Garden, Philadelphia) ).
NEMATODE PARASITES OF BIRDS 350
MICROTETRAMERES HELIX, new species
Host.—Primary : Corvus americanus; secondary : Unknown; prob-
ably similar in a general way to that of 7’. fissispina (p. 348).
Location.—Proventriculus.
Morphology—M icrotetrameres (p. 351).
Male 4.9 mm. long by 100n wide. There are no longitudinal rows
of spines on the body. Buccal capsule 21 deep; pharynx 274
long; esophagus 531p long. Nerve ring 191, from anterior ex-
tremity. Tail (fig. 417@) 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, 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 1.8 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 6 and c).
_ No longitudinal furrows as in 7etrameres. Cuticle very loose, pro-
jecting from the body in transparent folds. Head end blunt; buccal
capsule 22.54 deep; pharynx 225 to 250u long; esophagus very
thick, soon becoming obscured by the uterine coils so that its distal
end is not observable. Anus 141p, vulva 216 from posterior end,
which is finely pointed and projects out from a transparent cuticular
collar (fig. 417 d). Eggs 42u long by 33» wide; embryonated.
Life history—Probably similar in a general way to that of 7’.
fissispina (p. 348).
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, 1879a; Tetrameres
inermis (Linstow, 1879) 'Travassos, 1914d.
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 7’. fissispina
(p. 348).
Location.—Proventriculus.
Morphology.—Microtetrameres (p. 351).
356 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Male 2.125 mm. long by 90n wide (fig. 418 a tod). Buecal-capsule
19. long. Pharynx 225h long. Esophagus 395p 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 75u long. Tail tapering, ends in small button.
Female forming cysts 2 mm. large. Body coiled in complex man-
ner (fig. 419 a@ to ¢). No longitudinal furrows as are found in spe-
cies of Tetrameres. Cuticle of anterior extremity folded to give
“C/TVA
Figs. 417—-418.—417, MICROTETRAMERES HELIX. @, MALE TAIL; 0 AND C, FEMALES;
d, FEMALE TAIL. ORIGINAL, 418, MICROTHTRAMERES INERMIS. a, MAL; b, VEN-
TRAL VIEW AND C, LATERAL VIEW OF TAIL OF SAME; d, HEAD END OF SAME. (SCALE
APPLIES TO Db, c, AND d.) AFTER SEURAT, 1913
appearance of spy-glass (fig. 419d). Buccal capsule 20» long.
Pharynx 300p long (Linstow) or 2604 long (Seurat) by 40n wide.
Esophagus 1.125 mm. long (Seurat) or 2 mm. long (Linstow) by
244 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 young 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 1352 from
posterior extremity. Vulva (fig. 420d) 95p anterior to anus. Ves-
tibule of ovejector (fig. 420 a and 6) 875 long. Trompe in the form
of a large reservoir 375y long by 110y 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.
420e and 7) 36 long by 20u wide (Linstow) or 52» long by 37p
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, long, cuticle
finely striated.
500 u &.
Fic. 419.—MICROTETRAMERES INDRMIS. FEMALE. da, ADULT; b, ADULT EXAMINED
PERPENDICULARLY TO DETERMIND MANNER OF COILING; C, IMMATURE; d, ANTERIOR
END. AFTER SEURAT, 1913
Life history.—Probably similar in a general way to that of 7’.
fissispina. 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.
MNstribution—Asia (Russian Turkestan) and Africa (Algeria
(Bou-Saada) ).
MICROTETRAMERES INFLATA (Mehlis, 1846) Travassos, 1915
Synonyms.—sS piroptera inflata Mehlis, 1846; Z’ropidocerca para-
dowa Diesing, 1851, part; Z’vropidocerca inflata (Mehlis, 1846) Dies-
ing, 1861; T’ropidocerca paradoxa Linstow, 1877 (not T'ropidocerca
paradoxa Diesing, 1835); Tropisurus inflatus (Mehlis, 1846) Neu-
mann, 1892; Yetrameres 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 description
358 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
under 7 ropidocerca 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 paradowa 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 7’. paradowa is
a synonym of Hystrichis papillosus (Eustrongylides papillosus)
while Jaegerskiold (1909) has listed both 7’. paradowxa Linstow, 1877,
and Hystrichis papillosus Linstow, 1899 in part as synonyms of
Fig. 420.—MICROTETRAMERES INERMIS. a, OvesEecror. AFTER SHURAT, 1914.
b, PART OF SAME; C, OVARY, OVIDUCT, AND INITIAL REGION OF UTERUS; d,
VULVA; e€ AND f, HATCHING OF EGGS. AFTER SEURAT, 1913
Eustrongylides elegans. Jinstow’s description and figures are un-
questionably of a species of Hustrongylides and agree with L’. elegans
in all particulars except the length of the spicule. It follows there-
fore, from all the available evidence, that Aficrotetrameres inflata is
in reality a species of Hustrongylides and the present writer will not
include the description of it in the Tetrameridae but under /. ele-
gans, the latter, however, apparently also being synonymous with
EH. mergorum (p. 372).
MICROTETRAMERES MINIMA (Travassos, 1914) Travassos, 1915b
Synonym.—Tetrameres minima Travassos, 1914.
Hosts —Primary: Tachyphonus cristatus brunneus; secondary:
Unknown; probably similar in a general way to that of 7. fissispina
(p. 348).
Location.—Proventriculus.
Morphology.—M icrotetrameres (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% and 100u
long, respectively.
Female 780p long by 6404 wide. Body red, strongly striated trans-
versely; long axis coiled (this is assumed to be true as Travassos in-
cluded this species in Mtcrotetrameres). Buccal capsule 12u long;
pharynx 73» long by 10u wide; esophagus 490» long by 50u wide.
Intestine saclike, becomes thinner posteriorly. Anus 684 from pos-
terior extremity. Vulva slightly anterior to anus. Eggs 45y long by
24 wide, many of them embryonated.
Life history.—Probably similar in a general way to that of 7.
fissispina (p. 348).
Distribution.—South America (Brazil).
MICROTETRAMERES PUSILLA Travassos, 19156
Synonym.—T etrameres pusilla Travassos, 19150.
Hosts—Primary: Turdus rufiventris and Platycichla flavipes;
secondary: Unknown; probably similar in a general way to that of
T’. fissispina (p. 348).
Location.—Proventriculus.
Morphology.—Microtetrameres (p. 351).
Male 3.5 to 4 mm. long by 100 to 1204 wide. Body (fig. 422) fili-
form, white, cuticle transversely striated and without spines. Buccal
capsule cylindrical, 171 deep by 7n wide. Pharynx 300u long. Eso-
phagus yellow-colored, slightly claviform. Travassos says it is 42p
long but he evidently means width; judging its length as compared
with that of the pharynx, in his figure, the esophagus measures about
600u long. ‘Two slender spicules of very different lengths, the smaller
about 85u long by 5» wide, the larger approximately 1.32 mm. long
by 7» wide, having the basal part dilated while the distal end has a
rounded point. Cloacal 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, 94 long and 10u in maxi-
mum width. Pharynx 273p long. Esophagus cylindrical, 974, long
(in his description in 1919 Travassos says 530 long by 90 wide).
Anus 140, from posterior extremity. Eggs ellipsoidal, 42 to 49
long by 28» wide.
Life history.—Probably similar in a general way to that of 7.
fissispina (p. 343).
Distribution.—South America (Brazil).
360 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
MICROTETRAMERES SPIRALIS (Seurat, 1915) Travassos, 1915b
Synonyms.—T ropidocerca spiralis Seurat, 1915a; Tetrameres
spiralis (Seurat, 1915) Travassos, 19150.
Hosts—Primary: Bubulcus lucidus; secondary : Unknown; proba-
bly similar in a general way to that of 7’. fissispina (p. 343).
Location.—Proventriculus.
Morphology—Microtetrameres (p. 351).
Male 4.75 mm. long by 130n wide. Body filiform, white. Buccal
capsule (fig. 423a) 30» long. Pharynx narrow, its length 1/4 that
of the esophagus. Combined length of pharynx and esophagus 1/3
Fies. 421-423-421, MIcROTETRAMERES MINIMA. MALH. AFTER TRAVASSOS, 1914. 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, 25u in front of
the level of these papillae; it is connected with a cuticular canal 50y
long. Cloacal aperture (fig. 423 6) 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,) and curved, the left
2.3 mm. long, slender and filiform.
Female 2.5 mm. long by2mm. wide. Body red color, rolled spirally
(314 times). No longitudinal furrows as are present in species of
Tetrameres. Cuticle transversely striated. Buccal capsule bottle-
shaped, 302 long. Pharynx 175y 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 180p. Anus 225p 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 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 50u long by 30n wide, embryonated at
maturity, thick-shelled, oval, flattened on one face.
Larva 2.3 mm. long by 70u wide. Buccal capsule 20 long;
pharynx 320, long, surrounded in the middle by nerve ring; esopha-
gus 600 long. Cervical papillae 210, from the anterior extremity.
Tail pointed, 170» long.
Life history—Probably similar in a general way to that of 7. fis-
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. /emale 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.
Ty pe-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
KBY TO SPECIES OF ANCYRACANTHOPSIS
Male 3.5 to 4 mm., female 5 to 6 mm. long; vulva in posterior part of body;
INISCCING TASOTAG = 3s eee eee ee Se eet 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 Hurypyga helias.
Ancyracanthopsis bilabiata, p. 362.
ANCYRACANTHOPSIS BILABIATA (Molin, 1860) Diesing, 1861
Synonym.—A ncyracanthus bilabiatus Molin, 1860d.
Host—Primary: EKurypyga 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 long, similar to those of A. bilabiata. In addi-
tion, at the base 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 1804 wide. Caudal extremity
(fig. 424 ¢ 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.
494 ¢) 100» long, ending in 3 fine points, the left 34u long and, ac-
NEMATODE PARASITES OF BIRDS 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 200u wide. Vulva (fig. 424 g) slightly
prominent, situated shghtly 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 f).
Eggs 40n long by 83 wide, embryonated, bluntly oval, the shell
10p thick.
Life history—Unknown.
Distribution —Not given (Germany ?).
This species should certainly be in the same genus with A. bélabiata.
Mueller evidently did not know that Diesing had made a new genus
for the species bzlabiatus as he calls it Ancyracanthus bilabiatus and
—
Fic. 424.—ANCYRACANTHOPSIS BIHAMATA. @ AND b, HEAD END; ¢€ AND d, MALE
TAIL; €, SPICULH; 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. MMJale with caudal alae
more or less well-developed; two spicules. /emale 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.
Ty pe-genus—Gnathostoma Owen, 1836.
Subfamily GNATHOSTOMINAE Baylis and Lane, 1920
Subfamily 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.
Ty pe-genus.—Gnathostoma Owen, 1836.
Genus GNATHOSTOMA Owen, 1836
Synonym.—C hetracanthus Diesing, 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.
Ty pe-species.—G nathostoma 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
Pelecaius. Ongernotais20 2 Se fe ree 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 imperialis.
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 510n. Head (fig.
425a) set off from body by depression and encircled by 4 parallel
transverse rows of chitinous hooks or spines (fig. 425 6), 44 spines
in each circle. Head 150 long by 340u 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, 1874) 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.
mT i
ay Li uy fi | I ih
HI a WA Z, Z 5
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Pies. 425-426.—425, GNATHOSTOMA ACCIPITRI, 4@, GENERAL APPEARANCE: b, HOOKS
OF CEPHALIC REGION. AFTER SKRJABIN, 1916. 426, GNATHOSTOMA PELECANI. 4a,
GENERAL APPEARANCE; 0, 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—G@nathostoma (p. 364): Length 3 to 4 mm. long.
Denticulations very pronounced in the anterior region of the body.
Head (fig. 426a) rounded, flattened, with 4 concentric and super-
posed series of chitinous hooks. Glandular apparatus (fig. 4266)
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 70» long. The immature adults showed a
characteristic vulva (fig. 426 ¢) in the median region of the body.
Life history —Unknown.
Distri Surope (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 pelecant. '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. A/ale 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 eh surface of shell pitted or, rarely, non-
pitted but with ridges ( Hejstrtonis acanthocephalicus) .
Parasitic as adults in digestive tract of birds or kidneys and body
cavity of mammals.
Ty pe-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 DIOCTOPHY MIDAE Railliet, 1915
Synonym.—Eustrongylidae Leiper, 1908.
Family diagnosis —Dioctophymoidea (p. 366): Characters of the
superfamily.
L'ype-genus.—Dioctophyme Collet-Meygret, 1802.
KEY TO GENERA OF DIOCTOPHYMIDAR
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 Jagerskidld, 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).
Ty pe-species—Eustrongylides tubifex (Nitzsch, 1819) Jaegers-
kidld, 1909.
KEY TO SPECIES OF EUSTRONGYLIDES
1. Unrecognizable species________----------- Eustrongylides papillosus, p. 373.
Recognizables species. 2222. eg >
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. 2 = Be ee eee 2 eae Eustrongylides tubifex, p. 367.
Papillae of outer circle long (36 to 444), 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 14 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 Jaegerskidld.
EUSTRONGYLIDES TUBIFEX (Nitzsch in Rudolphi, 1819) Jaegerskidld, 1909
Synonyms.—Strongylus tubifex Nitasch, 1819, part; Hustrongylus
tubifex (Nitzsch, 1819) Diesing, 1851, part; Hystrichis tubifex
(Nitzsch, 1819) Molin, 1861, part.
Hosts —Primary: Anas boschas domestica, Colymbus arcticus, C.
septentrionalis; secondary: Unknown, but fish probably (see below
3612—27
368 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
(life history) and see also /’. ignotus, p. 371), according to Jaeger-
skidld 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. 4275) 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 160u long.
Male 34 mm. long by 2 mm. wide. Bursal cup (fig. 427@) trumpet-
shaped, its margin with 2 incisions on the ventral surface.
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Fic. 427.—EUSTRONGYLIDES TUBIFEX. @, MALY TAIL; b, HEAD
PND; C, FEMALE TAIL. AFTER JABGERSKIOLD, 1909
Female 35 to 44 mm. long by 2.5 to 3 mm. wide. Vulva near
anus (fig. 427.¢). Vagina 11mm. long. Eggs 65 to 75h long by 44
wide, oval with blunt ends and thick shells, the latter pitted.
Life history—Unknown; probably involves intermediate stages
in fish, according to Jaegerskidld 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 Jaegerskidld as the larva of a
species of Lustrongylides; see EL. ignotus, p. 371. Recently Ciurea
has found similar larvae in certain fish (Barbus fluviatilis, Lota
lota, F'sox luctus, and Perca fluviatilis) in the Danube and regards
them as larval forms of some species of this genus. As /. 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 Z'us-
trongylides, the specific identity of the larvae not being ascer-
NEMATODE PARASITES OF BIRDS 36S
tained and the life history still lacking experimental confirmation.
The larvae are relatively large, 28 to 70 mm. long by 264 to 539u
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 mouth
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 /’. ignotus,
p. 371.
Distribution.—Europe.
Fic. 428.—EUSTRONGYLIDES LARVAE. SPECIES NOT DETERMINED. @, FRONT VIEW OF
HEAD; 0, ANTERIOR END; C, OUTLINE OF MALE TAIL; d, OUTLINE OF FEMALE TAIL.
APTER CIUREA, 1924
EUSTRONGYLIDES AFRICANUS Jaegerskiold, 1909
Hosts —Primary: Anhinga rufa, Ardea goliath, Leptoptilus cru-
menifer, Pelecanus rufescens, Platalea leucorodia; secondary: Un-
known, probably fish; see 2. tubifew, p. 867, and FE. ignotus, p. 371.
Location.—Proventriculus.
Morphology.—Eustrongylides (p. 367) : Body almost equally thick
throughout, without noticeable swelling in the middle region. Head
(fig. 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 wide; those of the outer circle 48. long and 72 to 96 wide.
Cuticle with coarse transverse striations. Mouth cavity distinct
though not large, of triangular cross-section; its length is 80 to 100,.
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 575p, at tail end (fig. 429 6). Esopha-
gus about 16.5 to 18 mm. long. Nerve ring 160, from anterior end.
Eges (fig. 429 ¢ to e) 70 to 76u long by 36 to 42n wide.
Life history —Unknown; see /’. tubifew, p. 367.
Distribution.—Africa (Sudan) and Asia (Russian Turkestan (lac
Kul-Kainar) ).
370 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
EUSTRONGYLIDES EXCISUS Jaegerskiold, 1909
Synonyms.—Strongylus tubifex Rudolphi, 1819, part; Hustrongy-
lus papillosus Diesing, 1851, part; Hystrichis papillosus Molin,
1861, part; Zystrichis elegans Stossich, 1899, part.
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Fic, 429.—EuSTRONGYLIDES AFRICANUS. @, HEAD END; 0b, FEMALE TAIL; Cc, EX-
TERIOR OF EGG; d, PITTING OF SHELL; €, INTERIOR OF EGG. AFTER JAEGERSKIOLD,
1909
Hosts—Primary: Phalacrocorax carbo and P. pygmaeus; second-
ary: Unknown, probably fish; see H’. tubifex, p. 367, and EF’. zgnotus,
p. di.
Location.—Gizzard.
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Fic. 430.—EUSTRONGYLIDES EXCISUS. a, HBAD; b AND C, TAIL OF MALE. AFTER
JAEGERSKIOLD, 1909
Morphology—Eustrongylides (p. 867): Head (fig. 480a) 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 800n, width at an-
terior end of body 168 to 190,, at posterior end (just anterior to
bursal cup) 370 to 4802. Esophagus 7.3 to about 12 mm. long.
Bursal cup 480 to 560p long by 560 to 700 wide, trumpet-shaped
(fig. 430 6 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 F’. tubifex, p. 367.
Distribution.—EKurope (Austria (Museum, Vienna)) and Asia
(Russian Turkestan).
HoH
cen
Fia. 431.—EUSTRONGYLIDES IGNOTUS. a, HEAD; b, FEMALE TAIL; GC AND d, MALE
TAIL; € AND f, EGG. AFTER JAEGERSKIOLD, 1909
EUSTRONGYLIDES IGNOTUS Jaegerskiold, 1909
Synonyms.—F ilaria cystica Rudolphi, 1819; Agamonema eysticum
(Rudolphi, 1819) Diesing, 1851; Hustrongylus papillosus Diesing,
1851, part; Hystrichis papillosus Molin, 1861, part; Hustrongylus
tubifea Schneider, 1866, part; Spiroptera bicolor Linstow, 1899.
Hosts—Primary: ?Anhinga anhinga, Ardea cocoi, A. herodias,
Botaurus pinnatus; secondary: Fish; larvae in Symbranchus laticau-
datus and Galaxias scriba resemble this species, according to Jae-
gerskiéld, 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. 431a) with 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, long.
Male 36 mm. long; maximum width 1 to 1.4 mm., width at head
end 250p, at tail end (just anterior to bursal cup) 350 to 460z.
372 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Esophagus 8 mm. long. Bursal cup (fig. 481 ¢ and d) 290 to 350p
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 400un. Esophagus 11.6 to 16.4 mm. long. Eggs 58
to 664 long by 35 to 44» wide; shell 4.51 thick, with pittings rela
tively sparse (fig. 481 e and /).
Life history—Unknown, involving intermediate stages in fish.
Jaegerskidld 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
diaphanus 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 £.
tubifex, p. 367.
Distribution.—Kurope (Germany (Berlin) and Austria (Museum,
Vienna) ).
EUSTRONGYLIDES MERGORUM (Rudolphi, 1809) Cram, 1927
Synonyms.—Strongylus mergorum Rudolphi, 1809; Strongylus
papillosus Rudolphi, 1809, part; Strongylus elegans Olfers, 1816;
Strongylus tubifex Rudolphi, 1819, part; Zropidocerca paradoxa
Linstow, 1877 not 7. paradowa Diesing, 1851; Hystrichis elegans
(Olfers, 1816) Railliet, 1893, part; Hustrongylides elegans (Olfers,
1816) Jaegerskidld, 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 Red.
Hosts—Primary : Alca torda, Anas boschas 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 molisstma, Uria trotle;
secondary: Unknown; probably fish, according to Jaegerskiéld and
to Ciurea. See /. tubifex, p. 367, and /. ignotus, p. 371.
Location.—In tubercles in the esophagus and proventriculus.
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 Sha
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, to 175u 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. 484 ¢ and d@) almost trumpet-shaped,
275 to 320n long by 260 to 360n 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).
I'ies, 482—433.— EUSTRONGYLIDES MERGORUM. 432, a, FEMALE; b, MALE; NATURAL
SIZE; C, HEAD. AFTER LINSTOW, 1877. 433, a, FRONT VIEW AND D, LATERAL
VIEW OF HEAD; ¢, 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. 4846). Vulva close to anus.
Eggs 60 to 70 long by 33 to 38 wide, with blunt ends and pitted
shells.
Life history.—Unknown; see 2’. tubifex, p. 367.
Distribution.—Europe.
EUSTRONGYLIDES PAPILLOSUS (Rudolphi, 1802) Jaegerskiéld, 1909
Synonyms.—Strongylus papillosus Rudolphi, 1802, part; H'ustron-
gylus papillosus (Rudolphi, 1802) Diesing, 1851, part; ystrichis
papilosus (Rudolphi, 1802) Molin, 1861, part.
Hosts.—Primary: Anas boschas domestica, Anser cinereus domes-
ticus, and Nuecifraga caryocatactes; secondary: Probably fish; see
EL. tubifex, p. 367, and L. ignotus, p. 371.
Location.—In tubercles in the esophagus.
Morphology.—E ustrongylides (p. 367): Body not enlarged in the
middle portion, according to Jaegerskidld; 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» 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 Jaegerskiéld. The
above description is from Linstow.
Life history—Unknown; probably involves intermediate stages
in fish. See #. tubifex, p. 367.
Distribution.—Europe.
Fic. 434.—EUSTRONGYLIDES MERGORUM. a, HEAD END; Db, FEMALE TAIL; @ AND 4,
MALE TAIL, AIFTER 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 /. tubifex, p. 367.
Location.—Not given.
Morphology.—Eustrongylides (p. 367): A total of 18 cephalic
papillae (fig. 486 @ and }) 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, 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
375
NEMATODE PARASITES OF BIRDS
bursal cup) 270 to 290u. Esophagus 10.4 mm. long. Bursal cup
(fig. 436¢) 290 to 340u long by 320 to 350u 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.
o: maximum width 1.2 to 1.5 mm., width at
Female 17 mm. long;
head end 300y, at tail end (fig. 436 d) 250u. Ksophagus about 13.6
mm. long. Eggs 53 to 61p long by 31 to 33 wide, the poles very
wide, the shells thin, with closely-set pittings (fig. 436 e and /).
Life history—Unknown; see EF. tubcfex, p. 367.
Distribution—South America (Brazil).
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EUSTRONGYLIDES PAPILLOSUS.
436, EUSTRONGYLIDES PERPAPILLATUS.
VIEW AND 0b, LATERAL VIEW OF HEAD; C, MALE TAIL; d, FEMALE TAIL;
AFTER JAEGERSKIOLD, 1909
Genus HYSTRICHIS Dujardin, 1845
(p. 366): Body usually
Fics. 485-436.—435,
JAEGERSKIOLD, 1909.
€ AND f, EGGS.
Generic diagnosis.—Dioctophymoidea
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
Esophagus long, without bulb. dale with a bursal cup more
parts.
Female with anus in
or less bell-shaped. One very long spicule.
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.
Ty pe-species.—H ystrichis tricolor Dujardin, 1845.
3612—27——26
376 BULLETIN 140, UNITED ST/ ES NATIONAL MUSEUM
KEY TO SPECIES 0.’ HYSTRICHIS
1. Only one circle of spines present; it is directly posterior to mouth, the spines
Wey larg ees seen i ets ee iy ope Nope Hystrichis coronatus, p. 378.
Numerous circles of spines, on head and cervical region if not on entire
MD LY 5 2 ear bn sisal a a em ae ct 2
Z. Spines of head and of cervical region of approximately the same size, or
the 'Tattemsmailer than the rorm cre eee sees kee AREER ee 3.
Spines of head and those of anterior part of body of different size, the latter
being longer and wangen: 24632 sek ae spesh ue of Se ee Ee reg pee 6.
5. 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 1754 anterior to it; eggs not pitted but with network
Of LId PCR! =k Sn es eS Hystrichis acanthocephalicus, p. 377.
Spines of head thickly set, about 50 to 57 in each circle; vulva situated di-
rectiyaatvwanus'; esesspitted . spaeoms 8 ai EE se eee 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
1
region;-bentiior crooked. 2eesersn ds Hystrichis varispinosus, p. 387.
Spines, as far as can be told from incomplete descriptions, straight______ io
7. Female 100 mm. long by 3 mm. wide; 17 spines in each circle on head;
from Wuiica atrazs22es. oes ee Hystrichis wedli, p. 381.
Female 25 to 44 mm. long by 500u to 1 mm. wide; 23 spines in each circle
on head; from Cygnus olor and Ibis falcinellus.
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 bellonti, T. tadorna; secondary: Probably fish, according to
Jaegerskidld.
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 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 vo 40 mm. long by 350 to 500u wide. The vulva is at
the posterior end, just anterior to the anus (fig. 437). Eggs 85 to
88u long by 36 to 40n 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 Jaegerskidld.
Distribution—Europe (France, Germany, Italy, Hungary and
Treland).
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Fig. 437.—HyYstTrRICHIS TRICOLOR. HEAD AND TAIL ENDS OF FEMALE. AFTER
JAEGERSKIOLD, 1909
HYSTRICHIS ACANTHOCEPHALICUS Molin, 1861la
Synonym.—sStrongylus tubifex tantali™
Hosts.—Primary: /bis nudifrons, Phimosus infuscatus; second-
ary: Probably fish, according to Jaegerskiéld. Jbis tubifea, re-
ported as a host in the index-catalogue of Stiles and Hassall, is a
lapsus, the generic name of the host (/bés nudifrons) having been
combined with the specific name from the synonym.
Location —Glands of proventriculus.
Morphology.—Hystrichis (p. 875): Swelling of head end (fig.
435 a@and 6) slight or lacking. Head with 7 to 9 circles of spines, the
spines relatively sparce, not closely set, the largest 25 to 30n long by
15 to 17» wide. Mouth opening triangular; mouth cavity 130, long.
Male 25 to 45 mm. long; maximum width 1.5 to 2 mm., width at
head end 270 to 290p, at tail end (just anterior to bursal cup) 160p.
Spines on anterior part of body to a distance of 800u from head end.
Esophagus about 8 mm. long. Bursal cup 160u long by 240u wide,
a round bell-shape, its walls thick, muscular (fig. 438 ¢ and d).
Female 35 to 47 mm. long; maximum width 2 to 3 mm., width at
head end 300 to 8360p, at tail end 320 to 3504. Spines on anterior part
4 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. from posterior end (fig. 438e). Eggs 75
to 791 long by 40 to 44 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-
seris Diesing, 1861.
Host—Primary: Mergus merganser; secondary: Unknown, prob-
ably fish, according to Jaegerskidld.
Location —Glands of proventriculus.
Fic. 438.—HyYSTRICHIS ACANTHOCEPHALICUS. @, SIDE VIEW; b, FRONT VIEW OF
HEAD; c AND d, MALE TAIL; €, FEMALE TAIL; f, EGG. AFTER JAEGERSKIOLD,
1909
Morphology—Hystrichis (p. 875): 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. 4395), 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.—E chinocephalus cygni Molin, 1858; Hystrichis pachi-
cephalus Molin, 1861a.
NEMATODE PARASITES OF BIRDS 379
Hosts—Primary: Cygnus olor; secondary: Probably fish, ac-
cording to Jaegerskidld.
Location.—In vesicles between the external coats of the proven-
triculus.
Morphology.—H ystrichis (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.
Fic. 439.—HYSTRICHIS CORONATUS. «a, IPAD PND; b, FEMALE, NATURAL SIZE;
C, FEMALB TAIL. AFTER MOULIN, 1860
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 NEGLKECTUS Jaegerskidld, 1909
Synonyms.—Eustrongylus papillosus (Rudolphi, 1802) Diesing,
1851, part; Hystrichis papillosus (Rudolphi, 1802) Molin, 1861la,
part.
In addition to the above, Jagerskiéld lists Zystrichis, species Wedl
as a possible synonym and states that it is very close and possibly
identical with his new species. Hystrichis, species Wedl, however,
was named H. wedli by Linstow, emended by Jaegerskidld to /7Z.
wedli and if the two species, 7. neglectus and H., species Wed] are
identical, Jaegerskidld’s species will fall as a synonym of /7. wedi.
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 1. 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 Jaegerskidld.
Location.—Esophagus.
Morphology.—H ystrichis (p. 375) : Body long and slender. Mouth
opening small, triangular, surrounded by 6 small papillae, wart-
shaped with small points. Head (fig. 441@) with closely-set spines,
larger than those found on anterior cervical region. Largest spines
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Fias. 440-441.—440, HYSTRICHIS CyGNI, HAD END. AFTER MOULIN, 1861. 441,
HYSTRICHIS NEGLECTUS. @, HBAD END; 0, MALE TAIL; C, FEMALE TAIL; d, BGG.
AFTER JABGERSKIOLD, 1909
40 to 504 long by 20% wide. Spines-extend along body for distance
of 1.5 mm.
Male specimens imperfect; length more than 33 mm. Bursal cup
(fig. 4416) 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 500p, at tail end (fig. 441¢) 350u. Each circle of
the head contains 55 to 57 spines. Esophagus 8.25 mm. long. Kegs
79 to 84 long by 42 to 44» wide, with large but not closely-set pit-
tings (fig. 441d). 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, [bis falcinellus; secondary: Un-
known, probably fish, according to Jaegerskidld.
Location.—Glands of proventriculus.
Morphology —H ystrichis (p. 375): Head (fig. 442) distinetly
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, 500u 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 ZZ. tricolor,
p. 376.
Distribution —Europe (Italy (Padua) ).
HYSTRICHIS VARISPINOSUS Jaegerskiold, 1909
Host—Primary: Vergus serrator; secondary: Probably fish, ac-
cording to Jaegerskidld.
Location.—Not given; probably proventriculus.
Morphology.—H ystrichis (p. 375): Head (fig. 448@) only slightly
swollen, covered with 15 circles of small, closely-set strongly-bent
spines, the smallest of which are 27» long, the largest 48, long;
the highest number of spines in a circle is 52. Spines (fig. 4430)
of cervical region 88 to 90u 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,, the width of head 400z.
Esophagus 6 mm. long.
Life history—Unknown; possibly similar to that of HZ. tricolor,
p. 376.
Distribution.—N ot given.
HYSTRICHIS WEDLI Linstow, 1879 emended Jaegerskidld, 1909
Synonyms.—H ystrichis, species Wedl, 1856; H. wedlii Linstow,
1879a. H. 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 Jaegerskidld.
Location.—Esophagus.
Morphology.—H ystrichis (p. 375): Head a knob-hke 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, the spines disappear 3 mm. from the head end.
Male unknown.
Female 100 mm. long by 3 mm. wide.
fh EAN a
HUN if
f \ f.
\\ Na
H
d
iM
Fies 442-444.—442, HystricHIS ORISPINUS. HEAD BND. FROM JARGER-
SKIOLD, 1909, AFTER MOLIN. 443, HYSTRICHIS VARISPINOSUS. 4, H®AD
END; 0b, SPINE OF ANTERIOR CERVICAL REGION. AFTER JAEGERSKIOLD,
1909. 444, HYSTRICHIS WEDLI (LINSTOW’S MATERIAL). da, ANTERIOR
PORTION OF BODY; b, ONE OF THER MOST ANTRPRIOR SPINES, LATERAL VIEW ;
Cc, SAME, FRONT VIEW; d, ONE OF THE MOST POSTERIOR SPINES; €, POSTERIOR
END OF BODY (%, PROBABLE PRIMORDIUM OF VAGINA). AFTER JAEGERSKIOLD,
1909
Larva (found by Linstow) 24 mm. long by 600s 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 692 long by 29% wide at the base,
very thick at head end (fig. 444 @ 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 Wed] for adult.
Life history—Unknown; possibly similar to that of H. tricolor,
p. 376.
Distribution.—Not given.
NEMATODE PARASITES OF BIRDS 383
Jaegerskidld 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. varispinosus
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 Boulenger, 1926
Boulenger ** 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 ** has made a new genus based on three female speci-
mens of nematodes found along with specimens of Améidostomum
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.—P seudamidostomum loosi Boulenger, 1926.
PSEUDAMIDOSTOMUM LOOSI Boulenger, 1926
Boulenger** describes this species from stomach wall of /u/lica
atra; Egypt.
Females only, 7.3 to 7.8 mm. long by 160 to 170p wide. Cuticle
of head slightly expanded; four conspicuous submedian papillae;
1221926: Parasitology, Cambridge, Eng., vol. 18, pp. 94-96, figs. 18-22.
13JTdem, pp. 95-97.
%*Tdem, p. 96, figs. 23-25.
384 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
mouth opening small, circular. Buccal capsule broad (124) and very
short, without teeth. Esophagus 800 to 950% long. Anus 70p 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 325y; uteri
divergent. Eggs 57 to 69u long by 37p to 42u wide.
EPOMIDIOSTOMUM QUERQUETULAE Boulenger, 1926
Boulenger '° also describes this species from stomach wall of Quer-
quedula crecca (=Nettion crecca) ; Egypt.
Morphology. pomidiostomum (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 (2. uncinatum). 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 180n to 200n long, the distal portion divided into
two unequal branches. Gubernaculum absent. Memale 13 to 14.5
mm. long by 2202 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, 460n long. Eggs 80p to 90 long by 50n wide.
This species, as described by Bbileiber? is very oe to E'po-
midiostomum orispinum from the same host, te 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 /. orispinum to compare with that of /’. quer-
quetulae, the latter must be considered a distinct species.
EPOMIDIOSTOMUM SKRJABINI Petrow, 1926
Petrow *° reports this species from Anser anser Page and
Anser albifrons; Don district, Russia.
MGA Gy Wg litiast onan (p. 26): Male 9.5 to 11.5 mm.
long by 210u to 2404 wide. Esophagus 1.02 mm. long. Spicules
151926: Parasitology, vol. 18, pp. 91, 92, figs. 8-10.
161926: Trudy gossudarstvennogo instituta eksperimentalnoi veterinaril ; Moskow, vol.
3 (1), pp. 103-105; 4 figs.
NEMATODE PARASITES OF BIRDS 385
206n to 210% long, dividing into three branches distally. General
shape of bursa similar to that of #. wneinatum, 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 2. ori-
spinum also. Female 15 to 17 mm. long by 2704 to 290n wide.
Esophagus 1.29 mm. long. Anus 156 to 175y, vulva 3.32 mm., from
posterior end of body. Eggs 101p to 105 long by 58, to 62u wide.
The measurements given above for this species correspond very
closely to those of H’. orispinum (p. 28), of which species Petrow
makes no mention; the hosts of the two species are identical also.
Uowever, 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) ZAKHAROWI Petrow, 1926
This species Petrow*’ 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
only, 9.47 mm. long by 1702 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 ** has described four new species of Heterakis, two new
species of Pseudaspidodera, and a new subulurid.
HETERAKIS HASTATA Chandler, 1926
Chandler ¥° reports this species from cecum and large intestine of
Lophura rufa; Zoological Gardens, Calcutta, India.
Morphology —Heterakis (p. 50): Large stout worms. Esophagus
1/7, or a little less, of body length. Lateral alae well developed, in
female 80un wide, in male a little less; they extend about 2/3 the
Jength of the body. Lips prominent, papillae smaller than in Z.
lanei. Male 10.5 to 11.5 mm. long by 375» wide. Esophagus 1.5
mm. long by 80% wide (narrow part) ; bulb 8385p long by 240, wide.
Tail 550p 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,
171926: Trudy gossudarstvennogo instituta experimentalnoi veterinarii; Moskow, vol.
3 (1), p. 108, 2 figs.
18 Indian J. M. Res., Calcutta, vol. 13 (&), pp. 617-624.
191926: Idem, p. 619, fig. 3.
386 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
its antero-postero diameter 145y. Spicules slightly unequal, the
left 1.75 mm. long, with broad alae, which are slightly asymmetrical
near tip. Right spicule, 1.65 mm. long, with narrow alae. /emale
12 to 13 mm. long by 4204 to 460» wide. Esophagus 1.6 mm. long;
bulb 4202 long by 270” wide. Vulva approximately at middle of
body. Anus 1.2 to 1.3 mm. from posterior end. Eggs 70p by 40p.
HETERAKIS LANEI Chandler, 1926
Chandler 2° 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 75p wide, in
front of bulb, which is 300g long by 190” wide. Tail 400u to 4504
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 300u 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 65p to 68u by 40p.
HETERAKIS VARIABILIS Chandler, 1926
Chandler *! reports the discovery of this species in the cecum and
large intestine of Polyplectrum 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 404; gradually narrow
posteriorly, extending 2/3 of body length in female, but disappearing
anterior to middle of body in male. Male 9 to 10 mm. long by 310,
wide. Esophagus 1.5 mm. long; bulb 320p long by 230, wide. Alate
tail end narrow and long, 480 long by 220u wide. Caudal papillae
said to be normal in number; the first lateral paracloacals (adanals)
201926: Indian J. M. Res. Calcutta, vol. 13 (3), p. 618, figs. 1 and 2.
211926: Idem., pp. 619, 620, fig. 4.
NEMATODE PARASITES 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 180u
anterior to anus. Tail about 400u long. Spicules unequal, the right
720p to 880u, the left 1.04 to 1.18 mm. long, similar in form and
narrowly alate. Hemale 11.5 to 12.5 mm. long by 385 wide. Esoph-
agus 1.6 to 1.65 mm. long; bulb 360, long by 2604 wide. Vulva pos-
terior to middle of body, dividing body length in ratio of 6:5 or 5.5.
Posterior third of tail tapers rapidly. Eggs 60u to 65h by 38, to 40p.
HETERAKIS VULVOLABITA Chandler, 1926
Chandler ** describes this species from cecum and large intestine of
Arboricola torqueola; Zoological Gardens, Calcutta, India.
Morphology —Heterakis (p. 50): Small, stout worms. Esophagus
1/8 to 1/7 of body length. Lateral alae narrow. Male 6 to 7 mm.
long by 250u wide. Esophagus about 870» long; bulb 210» long by
165, wide. ‘Tail 230 long, the slender part posterior to papillae
only 70uz long. Sucker small, 50u in diameter, situated 90» anterior
to cloacal aperture. Alate tail end short, 3304 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 5354 long and very slender (3p to 9p
wide), the right 290u to 300u long by 184 wide. Female 7 to 8 mm.
long by 265 wide. Esophagus 875 long; bulb 235p long by 165
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 540y long. Eggs small,
55p to 6Op by 35p.
PSEUDASPIDODERA VOLUPTUOSUS Chandler, 1926
Chandler ** describes this species from the cecum and large intes-
tine of Argusianus argus; Zoological Gardens, Calcutta, India.
Morphology—Pseudaspidodera (p. 102): Small, stout worms.
Lateral alae narrow, 252 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 1/6 of body length. Male 6.25 to 9.25 mm. long by 325, to
380, wide; head about 100% broad. Esophagus 1.38 mm. long; bulb
300 long by 250% wide. Alae of tail well developed, their length
520pn, total width 270u. Tail 360n to 390» long. Sucker, 100 in
diameter, is 125u to 160 anterior to the cloacal aperture. Spicules
221926: Indian J. M. Res. Calcutta, vol. 3 (3), pp. 620, 621, figs. 5-8.
231926: 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 200p long. Female 9.35 to 10.15 mm. long by
460. wide. Esophagus 1.6 to 1.7 mm. long; bulb 320p long by 250.
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 35u
to 38p.
PSEUDASPIDODERA VOLUPTUOSUS MINOR Chandler, 1926
This species is reported by Chandler *4 as having been found in
large intestine and cecum(?) of Rollulus rowlroul; 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 225y 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, 360p to 400p. Female 6.75 to 7.85 mm. long by 310u 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 multipapillata Chandler, 1926.22 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 250n
wide, in female. Male 6 to 7 mm. long by 2604 to 280 wide.
Esophagus 1.12 mm. long. Tail 230% long. Sucker 480, 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 590u 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, 780u long and 224 wide. Gubernaculum in two parts, a nar-
row stout piece 1454 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’ 1i"12:
7% Tdem., pp. 622, 623, figs. 13-15.
NEMATODE PARASITES OF BIRDS 389
350 wide. Vulva inconspicuous, anterior to middle of body, divid-
ing latter in ratio of about 3:5. Ovejector directed forward. Tail
about 2 mm. long. Eggs 56u to 63u by 34p to 42u.
CHEILOSPIRURA SKRJABINI (Ozerska, 1926) Cram, 1927
Synonym.—Acuaria (Cheilospirura) skrjabini Ozerska, 1926.*°
From the house sparrow; Don district, Russia.
Morphology.—C heilospirura (p. 226): Male 8.7 to 9.2 mm. long
by 1402 to 1904 wide. Pharynx 160p to 194» long; first part of
esophagus 399n to 4564 long; second part of esophagus 930, to 1.08
mm. long. Cordons 250 to 285 long. Spicules unequal, the one
205 to 239n long, the other 125 to 148 long. Cloacal aperture
2004 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 shghtly greater than between the other pairs,
the spacing between the others being almost equal. Female 23 to
27 mm. long by 130n to 200% wide. Pharynx 182» to 228, long;
first part of esophagus 593p to 707» long; second part of esophagus
1.25 to 1.7 mm. long. Cordons 2854 to 387, long. Vulva 11.11
to 18.11 mm. from head end of worm; anus 170 to 2504 from
posterior end. Eggs 43 to 484 by 28p to 29n.
CHEILOSPIRURA PAVONIS (Ortlepp, 1925) Cram, 1927
Synonym.—Acuaria (Cheilospirura) pavonis Ortlepp, 1925.°*7
From under lning of gizzard of Pavo muticus, locality not given.
Morphology —C heilospirura (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 304 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 16m wide, with chisel-like tip; the right
2154 long and very broad. Gubernaculum absent. Female 14.7
mm. long by 3584 wide. Pharynx 305» long by 60n wide; muscular
part of esophagus 1.14 mm. long; glandular part 3.25 mm. long.
Dorso-lateral cordons extend slightly posterior to vulva; ventro-
2°Trudy Gossud. Inst. Eksper, Vet., Moskow, vol. 2 (2), pp. 103, 104, figs. 1, 2.
27 Journ. Helminth., Lendon, 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, 1906) Ortlepp, 1925
Ortlepp ** 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 325n 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. Female 24 mm. long by 455z 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 + of Smit and Notosoediro,
1926.22 From the crop of the chicken; Java (7).
Morphology.—Gongylonema (p. 203) : Cuticular bosses extend for a
distance of about 500y from the head end. Mouth cavity long (34,)
and slender (7u). Male 18 to 19 mm. long by 2154 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 580p. Female 40 to 56 mm. long by 330 wide. Anus 332
from tail end. Vulva 2.5 mm. anterior to anus. Eggs 55 by 35p.
The spicule lengths of Gongylonema ingluvicola Ransom being
very different from the above, Smit and Notosoediro are apparently
dealing with a new species.
231925: Journ. Helminth., London, vol. 3, p. 180, 4 figs.
» Nedrl.-Ind. Blad. y. Diergeneesk., Buitenzorg, vol. 38, pp. 92-94, 7 figs.
NEMATODE PARASITES OF BIRDS 391
DISPHARYNX, new species?
Synonym.—Dispharagus new species ? of Smit and Notosoediro,
1926.°° From the gizzard of the Java cock.
Morphology—Dispharyna (p. 237): Mouth with two lips. Mouth
cavity becomes narrower at base. Mouth cavity 100 to 116 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 500u, 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 335u long and thin, the right short (120, long) and thick.
Female 6 mm. long. Anus 166, from posterior end of body; vulva
1.22 mm. anterior to anus. Eggs 37p by 20p.
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 )). 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 *! 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 Streptocara, and this genus put in the Physa-
lopterinae; I have put Yseria in Schistorophinae, Gedoelst having
indicated that it is closely related to Histiocephalus, 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 Ysera californica, the second species which I included
* Nedrl.-Ind. Blad. v. Diergeneesk., Buitenzorg, vol. 38, pp. 95-97, 4 figs.
311926: The Nematode Parasites of Vertebrates. 536 pp., 307 figs. London.
~
392 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
in that genus, Y’seria coronata, having been left by them in /istio-
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. bélabiata, and the
only species in Ancyracanthus being the type species, A. pinnatifidus.
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: Rhabdiasata, 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
jatter 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 ail groups
from species to orders, inclusive. The descriptive material of each
species includes synonyms, 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, Cheilospirura, Hartertia, E’chinuria, Tetra-
meres, and Microtetrameres; 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 THEIR PARASITIC NEMATODES ©
The writer is greatly indebted to Dr. H. C. Oberholser of the Bio-
logical Survey, United States Department of Agriculture, for deter-
22 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 depressum.
Accipiter melanoleucus:
Physaloptera ovata.
Accipiter nisus:
Porrocaecum depressum.
Habronema leptoptera.
Schistorophus bidens.
Synhimantus elliptica.
Synhimantus hamata.
Synhimantus laticeps.
Physaloptera alata.
Physaloptera alata chevreuci.
Physaloptera alata nouwveli.
Physaloptera crosi.
Physaloptera megalostoma.
Microtetrameres inermis.
Accipiter superciliosus:
Dispharynz capitata.
Physaloptera acuticauda.
Accipiter tinus:
Physaloptera acuticauda.
Acrocephalus arundinaceus:
Porrocaecum ensicaudatum.
Acrocephalus palustris:
Acuaria papillifera.
Actitis hypoleuca:
Cosmocephalus obvelatus.
Actitis macularia:
Cosmocephalus obvelatus.
Aegialitis dubia; see Charadrius
dubius.
Aegialitis hiaticola; see Charadrius
hiaticula.
Aegolius brachyotus; see Asio flam-
meus.
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 fissispina.
Air sponsa:
Heterakis caudata.
Ajaia ajaja:
Dispharynz magnilabiata.
Ajaja ajaja; see Ajaia ajaja.
Alauda arvensis:
Physaloptera crassa.
Alauda, species :
Porrocaecum ensicaudatum.
Alca torda:
Contracaecum spiculigerum.
Cosmocephalus obvelatus.
Streptocara crassicauda.
Hustrongylides mergorum.
Alcedo americana; see Chloroceryle
americana.
Alcedo ispida:
Streptocara decora.
Alectoris barbara:
Heterakis tenuicauda.
Subulura seurati.
Cyrnea eurycerca.
Hadjelia thuillieri.
Dispharynex spiralis.
Alectoris barbara spatei:
Hartertia obesa.
Alectoris graeca:
Heterakis tenuwicauda.
Subulura curvata.
Alectoris graeca chukar:
Ascaridia compar.
Subulura curvata.
Alectoris graeca sacxatilis:
Heterakis gallinae.
Heterakis tenuicauda.
Asearidia compar.
Alectoris rufa:
Subulura allodapa.
NEMATODE PARASITES OF BIRDS
Alectoris rufa—Continued.
Subulura seurati.
Cheilospirura gruveli.
Cyrnea eurycerca,
Cyrnea seurati.
Alectura lathami:
Heterakis bancrofti.
Ascaridia catheturina.
Amazona aestiva:
Ascaridia hermaphrodita.
Amazona auropalliata:
Habronema incerta.
Amazona barbadensis:
Habronema incerta.
Amazona farinosa:
Ascaridia hermaphrodita.
Amazona farinosa guatemalae:
Habronema incerta.
Amazona festiva:
Ascaridia hermaphrodita.
Amazona leucocephala:
Ascaridia hermaphrodita.
Habronema incerta.
Amazona ochrecephala:
Ascaridia hermaphrodita.
Amazona vinacea:
Ascaridia hermaphrodita.
Amazona vittata:
Ascaridia hermaphrodita.
Anas acuta; see Dafila acuta.
Anas albifrons; see Anser albifrons.
Anas anser domesticus; see Anser an-
ser domesticus.
Anas anser fera; see Anser anser.
Anas boschas; see Anas platyrhyncha.
Anas boschas domestica; see Anas
platyrhyncha domestica.
Anas boschas 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 Mareca penelope.
395
Anas platyrhyncha:
Trichostrongylus tenuis.
Syngamus trachea.
Heterakis dispar.
Ascaridia galli.
Porrocaecum crassum.
Porrocaecum ensicaudatum,
Contracaecum microcephalum,
Streptocara crassicauda,
Tetrameres fissispina.,
Hystrichis tricolor.
Anas platyrhyncha domestica:
Trichostrongylus tenuis.
Epomidiostomum uncinatum.
Cyathostoma brinchialis.
Heterakis dispar.
Heterakis gallinae.
Ascaridia galli.
Ascaridia lineata.
Porrocaccum crassum,
Contracaecum microcephalum,
Ichinuria jugadornata.
Echinuria uncinata.
Tetrameres fissispina.
Tetrameres gigas.
Hustrongylides mergorum.
Eustrongylides papillosus.
Hustrongylides tubifex.
Hystrichis tricolor.
Anas querquedula;
querquedula.
Anas rubripes:
Echinuria wneinata.
Anas segetum; see Anser fabalis.
Anas sponsa; see Aix sponsa.
Anas tadorna; see Tadorna tadorna.
Anhinga anhinga:
Contracaecum Sspiculigerum.
Eustrongylides ignotus.
Anhinga melanogastris:
Contracaecum spiculigerum.
Contracaecum tricuspe.
Acuaria macrolaima.
Anhinga novaehollandiae:
Contracaecum spiculigerum.
see Querquedula
Anhinga rufa:
Contracaecum rodhaini.
Contracaecum spiculigerum.
Subulura plotina,
EBustrongylides africanus.
Anser acuta; see Dafila acuta.
Anser albifrons:
Trichostrongylus tenuis.
Amidostomum anseris.
396
BULLETIN 140, UNITED
Anser albifrons—Continued.
Epomidiostomum orispinum.
Epomidiostomum uncinatum.
Anser anser:
Trichostrongylus tenwis.
Amidostomum anseris.
Epomidiostomum orispinum.
Syngamus trachea.
Heterakis dispar.
Heterakis gallinae.
Anser anser domesticus:
Trichostrongylus tenuis.
Amidostomum anseris.
Epomidiostomum orispinvum.
Cyathostoma bronchialis.
Heterakis dispar.
Heterakis gallinae.
Ascaridia anseris.
Probably Ascaridia
“ goose.”
Echinuria uncinata.
Hustrongylides 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.
Heterakis 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-
lissima.
Anser nigra; see Oidemia nigra.
Anser penelope; see Mareca penelope.
Anser segetum; see Anser fabalis.
Anthochaera carunculata; see Crea-
dion carunculatus.
Anthropoides virgo:
Porrocaecum. serpentulus.
Antrostomus vociferus; see Setochal-
cis vocifera.
lineata; see
STATES NATIONAL MUSEUM
Aquila albicilla; see Haliaeetus albi-
cilla.
Aquila chrysaetos:
Porrocaecum depressum.
Physaloptera alata nouveli.
Aquila fasciata; see Hieraaetus fas-
ciatus.
Aquila heliaca:
Porrocaecum depressum.
Porrocaecum. kirghisensis.
Physaloptera alata.
Tetrameres, species.
Gnathostoma, accipitri.
Aquila imperialis ; see Aquila helica.
Aquila maculata:
Porrocaecum depressum.
Thelazia stereura.
Aquila naevia; see Aquila pomarina.
Aquila pennata; see Hieraaetus pen-
natus.
Aquila pomarina:
Porrocaecum depressum.
Thelazia stereura.
Aquila rapaxz belisarius:
Physaloptera galinieri.
Ara ararauna:
Ascaridia hermaphrodita.
Ara macao:
Ascaridia hermaphrodita.
Aramides cajanea:
Tetrameres tetrica.
Aratinga leucophthalma:
Ascaridia hermaphrodita.
Aratinga pertinaxr:
Ascaridia hermaphrodita.
Arceuthornis musicus
Porrocaecum ensicaudatum.
Spiroptera turdi.
Arceuthornis philomelos
Porrocaecum ensicaudatum.
Porrocaecum heteroura.
Spiroptera turdi.
Arceuthornis pilaris ;
Porrocaecum ensicaudatum.
Spiroptera turdi.
Arceuthornis torquatus:
Porrocaceum ensicaudatum.
Arceuthornis viscivorus ;
Porrocaecum ensicaudatum.
Archibuteo (Buteo) lagopus;
Buteo lagopus.
Archibuteo vulgaris; see Buteo buteo.
see
NEMATODE PARASITES OF BIRDS
Ardea agami; see Agamia agami.
Ardea caerulea; see Florida caerulea.
Ardea cinerea;
Porrocaecum reticulatum.
Porrocaecum serpentulus.
Contracaecum microcephalum.
Desmidocerca aerophila.
Desmidocerca nunidica.
Ardea cocoi;
Porrocaecum reticulatum.
Eustrongylides ignotus.
Ardea comata; see Comatibis eremita.
Ardea garzetta; see Garzetta garzetta.
Ardea goliath;
Eustrongylides africanus.
Ardea grus; see Megalornis grus.
Ardea herodias;
Contracaecum microcephalum.
Echinuria ardeae.
Eustrongylides ignotus.
Ardea leuce; see Casmerodius egretta.
Ardea major; see Ardea cinerea.
Ardea manillensis; see Phoyxr pur-
purea.
Ardea melanocephala:
Porrocaecum serpentulus.
Ardea minor; see Botaurus lentigi-
NOSUS.
Ardea nigra; see Ciconia nigra.
Ardea nycticorar; see Nycticorar
nycticorar.
Ardea pileata; see Pilherodius pilea-
tus.
Ardea purpurea; see Phoyx purpurea.
Ardea scapularis; see Butorides
striatus.
Ardea, species:
Contracaecum microcephalum.
Contracaccum multipapillatum,
Contracaecum tricuspe.
Ardea stellaris; see Botaurus stellaris.
Ardea violacea; see Nyctanassa vio-
lacea.
Ardeola grayii;:
Contracaecum microcephalum.
Ardeola ibis; see Bubulcus ibis.
Ardeola ralloides:
Contracaecum microcephalum.
Ardetta minuta; see Ixobrychus
minutus.
Arenaria interpres morinella ;
Porrocaecum ensicaudatum,
Porrocaecum heteroura.
397
Asio flammeus:
Porrocaecum depressum.
Porrocaecum spirale.
Synhimantus laticeps.
Asio otus:
Porrocaecum depressum.
Porrocaecum spirale.
Habronema leptoptera.
Synhimantus laticeps.
Astur gentilis:
Porrocaecum depressum.
Habronema leptoptera.
Schistorophus bidens.
Physaloptera megalostoma.
Microtetrameres inermis.
Astur melanoleucus; see
melanoleucus.
Astur nisus; see Accipiter nisus.
Astur palumbarius; see Astur gen-
tilis.
Astur trivirgatus:
Habronema leptoptera.
Synhimantus sygmoidea.
Accipiter
Asturinule monogrammica; sce Kau-
pifalco monogrammicus.
Athene noctua; see Carine noctua.
Athene noctua glauxr; see Carine noc-
tua glaun.
Babulcus lucidus; see Bubuleug ibis.
Balearica pavonina:
Ascaridia cristata.
Porrocaecum serpentulus.
Balearica regulorum:
Ascaridia cristata.
Balearica regulosum; see Balearica
regulorum.
Barnardius barnardi:
Habronema incerta.
Bernicla sandwichensis; see Nesochen
sandvicensis.
Bolborhynchus lineola:
Habronema incerta.
Bolborhynchus monachus; see Myiop-
sitta monachus.
Bonasa sylvestris; see Tetrastes bon-
asia.
Bonasa umbellus:
Heterakis bonasae.
Ascaridia lineata.
Contracaecum, species.
Cheilospirura spinosa,
Dispharyne spiralis.
398
Botaurus lentiginosus:
Contracaecum microcephalum.
Streptocara triaenucha.
Botaurus minor; see Botaurus lentigi-
NOSUS.
Botaurus mugitans; see
lentiginosus.
Botaurus pinnatus:
Eustrongylides ignotus.
Botaurus stellaris:
Contracaecum microcephalum.
Synhimantus brevicaudata.
Branta canadensis:
Heterakis dispar.
Branta leucopsis:
Amidostomum anseris.
Epomidiostomum orispinum.
Heterakis dispar.
Brotogeris virescens:
Habronema incerta.
Bubo bubo:
Porrocaecum depressum.
Porrocaecum spirale,
Synhimantus laticeps.
Bubo mazimus; see Bubo bubo.
Bubo virginianus:
Porrocaecum depressum.
Porrocaecum spirale.
Bubuleus ibis:
Habronema ficheuri.
Synhimantus invaginata.
Tetrameres coccinea.
Microtetrameres spiralis.
Botaurus
Bubulcus lucidus; see Bubuleus ibis.
Bucco capensis:
Subulura forcipata.
Subulura strongylina.
Subulura travassosi.
Bucco chacuru:
Subulura travassosi.
Bucco coilaris; see Bucco capensis.
Bucco macrorhynchus:
Subulura travassosi.
Subulura forcipata.
Bucco melanoleucos; see Bucco tectus.
Bucco rufiventris ; see Bucco swainsoni,
Bucco, species :
Subulura forcipata,
Bucco striolatus:
Subulura forcipata,.
Subulura strongylina.
Subulura travassosi.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Bucco swainsoni:
Subulura forcipata.
Subulura strongylina.
Subulura travassost.
Microtetrameres cruzi.
Buecco tamatia:
Subulura forcipata.
Subulura strongylina.
Subulura travassost.
Pucco tamatina; see Bucco tamatia.
Bucco tectus:
Subulura forcipata.
Subulura strongylina.
Subulura travassosi.
Bucephala clangula; see Glaucionetta
clangula.
Buchanga atra assimilis; see Dicrurus
adsimilis.
* Buse” ;=Buteo buteo.
Buteo borealis:
Synhimantus sagittata.
Cyathostoma americana.
Buteo buteo:
Porrocaecum angusticolle.
Porrocaecum depressum.
Habronema leptoptera.
Habronema mansioni.
Spirura talpae (probably
dental).
Synhimantus hamata.
Thelazia stereura.
Physaloptera alata.
Buteo buteo vulpinus:
Thelazia stereura.
Buteo lagopus:
Syngamus coelebs (nomen nudum).
Porrocaecum angusticolle.
Porrocaecum depressum.
Synhimantus laticeps.
Buteo vulgaris; see Buteo buteo.
Buteo vulpinus; see Buteo buteo vul-
pinus.
Butorides striatus:
Porrocaecum serpentulus.
Butorides virescens virescens:
Contracaecum microcephalum.
“ Buzzard ”:
Physaloptera subalata.
Caccabis chucar; see Alectoris graeca
chukar.
Caccabis petrosa;
bara.
acci-
see Alectoris bar-
NEMATODE PARASITES OF BIRDS
Caccabis petrosa spatzi; see Alectoris
barbara spatzi.
Caccabis rufa; see Alectoris rufa.
Caccabis sazratilis; see Alectoris
graeca saxatilis.
Cairina moschata:
Heterakis dispar.
Ascaridia galli.
Porrocaecum crassum.
Callipepla squamata:
Ascaridia cordata.
Calonectris kuhlii:
Cosmocephalus obvelatus.
Seuratia shipleyi.
Cancroma cochlearia; see Cochlearius
cochlearius.
Capella media:
Sciadiocara uwmbellifera.
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 derbyanus.
Caprimulgus campestris; see Podager
nacunda,.
Caprimulgus candicans; see Thermo-
chalcis candicans.
Caprimulgus cortopan; see Setochalcis
rufa.
Caprimulgus
nacunda.
Caprimulgus europaeus:
Subulura subulata,
Subulura suctoria.
Thelazia cholodkowskii.
Caprimulgus fossti:
Subulura leprincei.
Caprimulgus guianensis ;
dromus albicollis.
Caprimulgus leucopygeus; see Nycti-
progne leucopyga.
3612—27 27
diurnis; see Podager
see Nycti-
399
Caprimulgus mercurius; see Eleo-
threptus anomalus.
Caprimulgus nacandua; see Podager
nacunda,
Caprimulgus nattereri; see Lurocalis
semitorquatus.
Caprimulgus nigrescens;
polus nigrescens.
Caprimulgus ruficollis:
Subulura forcipata.
Subulura strongylina.
Subulura subulata.
Caprimulgus rufus;
rufa.
Caprimulgus scaphiuris ; see Lurocalis
semitorquatus.
Caprimulgus semitorquatus; see Luro-
calis semitorquatus.
Caprimulgus, species:
Subulura suctoria.
Caprimulgus trifurcus;
salis climacocerca.
Caprimulgus urutas (or wrutau) ; see
Nyctibius jamaicensis.
Caprimulgus vociferus; see Setochal-
cis vocifera.
Carbo brasiliensis; see Phalacrocorar
vigua.
Carbo cormoranus; see Phalacrocorar
carbo.
Carbo cristatus;
aristotelis.
see Nycti-
see Setochalcis
see Hydrop-
see Phalacrocorar
Carbo dilophus; see Phalacrocoraz
auritus.
Carbo graculus; see Phalacrocorazr
aristotelis.
Carbo pygmaeus; see Phalacrocorar
pygmaeus.
Cariama cristata:
Ascaridia pterophora.
Subulura allodapa.
Subulura forcipata.
Subulura strongylina.
Subulura suctoria.
Oxryspirura brevipenis.
“Cariama huppé” ;=Cariama
tata.
Carine noctua:
Syngamus trachea.
Heterakis dispar.
Subulura suctoria.
Porrocaecum spirale.
cris-
400
Carine noctua glaur:
Subulura noctuae.
Spirocerca sanguinolenta (larva).
Dispharynz 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.
Casuarius galeatus; see Casuarius
casuarius.
Catarractes pachyrhynchus:
Cosmocephalus obvelatus.
Cathartes urubu; see Coragyps urubu.
Catheturus lathami; Alectura
lathami.
Catorrhactes pachyrhynchus; see Ca-
tarractes pachyrhynchus.
Celeus grammicus:
Habronema mansioni.
Celeus jumana:
Habronema longistriata.
Centropus monachus:
Subulura similis.
Centropus siamensis;
sinensis.
see
see Centropus
Centropus sinensis:
Ascaridia circularis.
Ascaridia trilabium.
Subulura rimula.
Oxryspirura siamensis.
Centurus flavifrons:
Microtetrameres cruzi.
Cephalopterus ornatus:
Habronema unilateralis.
Cepphus grylle:
Contracaecum spiculigerum.
Cosmocephalus obvelatus.
Streptocara, species (? S. pectini-
fera).
Cerchneis naumanni:
Habronema seurati.
Tetrameres, species.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
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. r
Charadrius himantopus; see Himan-
topus himantopus.
Charadrius morinellus; see Arenaria
interpres morinella.
Charadrius oedicnemus; see Oedicne-
mus oedicnemus.
Charadrius 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 americana:
Yseria coronata.
Chordeiles semitorquatus;
calis semitorquatus.
Chroicocephalus ridibundus:
Contracaecum spiculigerum.
Chroocephalus ridibundus; see Chroi-
cocephalus ridibundus.
Chrysolophus amherstiae:
Heterakis isolonche.
see Pluvialis
see Luro-
NEMATODE PARASITES OF BIRDS
Chrysolophus pictus:
Syngamus trachea.
Heterakis gallinae.
Heterakis isolonche.
Heterakis neoplastica.
Chrysolophus pictus obscurus:
Heterakis neoplastica.
Chrysotis auripalliata; see Amazona
auropalliata.
Chrysotis festiva; see Amazona fes-
tiva.
Chrysotis guatemalae ;
farinosa guatemalae.
Chrysotis leucocephalus; see Amazona
leucocephala.
Chrysotis ochroptera;
barbadensis.
Ciccaba huhula:
Spiroptera penithamata.
Ciconia alba; see Ciconia ciconia.
Ciconia alba asiatica; see Ciconia
ciconia.
Ciconia ciconia:
Syngamus trachea.
Cyathostoma variegatum.,
Contracaecum microcephalum.
Cyrnea exrcisa.
Physaloptera abbreviata (as pseudo-
parasite).
Ciconia maguari; see Huxrenura ma-
guari.
Ciconia nigra:
Syngamus trachea,
Cyathostoma variegatum.
Contracaecum engonium.
Contracaecum microcephalum.
Synhimantus sagittata.
Eustrongylides mergorum.
Circaetus gallicus:
Porrocaecum depressum.
Physaloptera alata.
Circactus pectoralis:
Porrocaecum depressum.
Circestus pectoralis; see Circaetus pec-
toralis.
Circus acruginosus:
Porrocaecum angusticolle.
Porrocaecum depressum,
Habronema leptoptera.
Physaloptera alata.
Circus buffoni:
Physaloptera acuticauda.
Physaloptera alata.
see Amazona
see Amazona
401
Circus cineraceus; see Circus pygar-
gus.
Circus cyaneus:
Porrocaecum depressum,
Habronema leptoptera.
Synhimantus laticeps.
Physaloptera alata.
Circus maculosus; see Circus buffoni.
Circus pygargus:
Porrocaccum depressum.
Porrocaecum. spirale.
Habronema leptoptera.
Synhimantus elliptica.
Synhimantus laticeps.
Physaloptera alata.
Circus rufus; see Circus aeruginosus.
Circus spilonotus: :
Ascaridia dolichocerca.
Circus spilothoraz; see Circus spilo-
notus.
Clangula hyemalis:
Streptocara crassicauda.
EHustrongylides mergorum.
Coccyzus melacoryphus:
Subulura acutissima.
Subulura forcipata.
Subulura strongylina,
Subulura subulata.
Physaloptera strongylina.
Coccyzus melanocoryphus ;
cyzus melacoryphus.
Coccyzus minor:
Subulura acutissima.
Subulura forcipata.
Physaloptera strongylina.
Cochlearius cochlearius:
Tetrameres cochleariae.
Tetrameres micropenis.
see Coc-
Colaptes auratus luteus:
Habronema colaptes.
Colaptes campestris:
Habronema. longistriata.
Colinus virginianis:
Trichostrongylus pergracilis.
Heterakis gallinae.
Ascaridia compar.
Cyrnea colini.
Coloeus monedula:
Syngamus trachea.
Sciadiocara secunda.
Columba arquatriz:
Ascaridia columbae.
402
Columba domestica; see Columba livia
domestica.
Columba domestica laticauda,; see Co-
lumba livia domestica.
Columba gutturosa; see Columba livia
domestica.
Columba livia:
Ascaridia columbae.
Dispharynx spiralis,
Columba livia domestica:
Ornithostrongylus quadriradiatus.
Ascaridia columbae.
Tetrameres confusa.
Tetrameres fissispina.
Columba picui; see Columbina picui.
Columba risoria; see Streptopelia ri-
soria,
Columba speciosa; see Lepidoenas
speciosa.
Columba talpacoti; see Chaemepelia
talpacott.
Columbina picui:
Ascaridia columbae.
Colymbus arcticus; see Gavia arctica.
Colymbus atrigularis; see Gavia arc-
tica.
Colymbus auritus:
Porrocaecum praelongum.
Contracaecum spiculigerum.
Cosmocephalus aduwunceus.
Hehinuria decorata.
Colymbus cristatus:
Contracaeum ovale.
Contracaewm spiculigerum.
Streptocara recta.
Eustrongylides mergorum.
Colymbus dominicus:
Contracaecum spiculigerum.
Colymbus nigricans; see Colymbus
auritus.
Colymbus nigricollis:
Contracaecum nigricollis.
Rusguniella elongata.
Colymbus ruficollis:
Contracaecum spiculigerum.
Tetrameres fissispina.
Eustrongylides mergorum.
Colymbus ruficollis capensis:
Contracaecum praestriatum.
Colymbus rufovulgaris; see Gavia
stellata.
Colymbus septentrionalis; see Gavia
stellata.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Comatibis eremita:
Contracaecum microcephalum.
Conurus leucotis; see Pyrrhura leu-
cotis.
Conurus pavua;
cophthalna.
Conurus pertinaz; see Eupsittula per-
tinaz.
Conurus solstitialis ;
solstitialis.
Coprotheres pomarinus:
Contracaecum spiculigerum.
Coracias abyssinicus:
Subulura similis.
Coracias garrulus:
Hadjelia truncata.
Acuaria anthuris.
Acuaria cordata.
Coragyps urubu:
Tetrameres paradoza.
Corone cornix; see Corvus corniz.
Corone corone; see Corvus corone.
Corvus americanus; see Corvus bra-
chyrhynchos.
Corvus brachyrhynchos:
Syngamus gracilis,
Acuaria anthuris.
Microtetrameres heliz.
Corvus cajanus; see Cyanocorax cay-
anus.
Corvus caryocatactes; see Nucifraga
caryocatactes.
Corvus corar:
Acuaria anthuris.
Acuaria cordata.
Corvus corax tingitanus:
Spirocerca sanguinolenta (larva).
Acuaria anthuris.
Microtetrameres inermis.
Corvus cornizv:
Porrocaecum semiteres.
Acuaria anthuris.
Acuaria cordata.
Acuaria depressa.
Tetrameres unispina.
Physaloptera malleus.
Corvus corone:
Syngamus trachea.
Acuaria anthuris.
Acuaria cordata.
Microtetrameres inermis.
Oxyspirura sygmoided.
see Aratinga leu-
see Hupsittula
NEMATODE PARASITES OF BIRDS
Corvus frugilegus; see Trypanocoraxr
frugilegus.
Corvus glandarius; see Garrulus glan-
darius.
Corvus monedula; see Coloeus mone-
dula.
Corvus pica; see Pica pica.
Corvus pyrrhocoraz ; see Pyrrhocorar
pyrrhocorac.
Corvus scapulatus:
Acuaria ornata.
Coscoroeba coscoroba:
Cyathostoma coscorobae.
Cotinus virginianus; see Colinus vir-
ginianus.
Coturnix communis; see Coturniz co-
turniz.
Coturniz coturniz:
Heterakis gallinae.
Ascaridia compar.
Cyrnea eurycerca.
Coturnix dactylisonans; see Coturniar
coturniz.
Coturnix« delagorguet;
delagorguet.
Coturnix delagorguei:
Subulura suctoria.
see Coturnir
Cranorrhinus corrugatus:
Hadjelia inermis.
Craxr blumenbachii:
Heterakis nattereri.
Craw fasciolata; see Crazv sclateri.
Crac sclateri:
Thelazia anolabiata.
Crar urumutum; see Nothocrax uru-
mutum.
Creadion carunculatus:
Oxzyspirura anthochaerae.
Creciscus viridis:
Schistorophus laciniatus.
Yseria coronata.
Crocopus phoenicopterus:
Ascaridia columbae.
Crossoptilon manchurianum ;
Crossoptilon mantchuricum.
Crossoptilon mantchuricum:
Heterakis gallinae.
Heterakis isolonche.
Crotophaga ani:
Subulura reclinata.
Spiroptera saginata.
Oryspirura anacanthura,
see
403
Crotophaga major:
Subulura reclinata.
Cyrnea semilunaris.
Oxryspirura anacanthura.
Cryptoglaux funerea:
Porrocaecum depressum.
Porrocaecum spirale.
Cryptoglaur tengmali;
glaux funerea.
Crypturornis noctivagus:
Heterakis valwata.
Subulura strongylina.
see Orypto-
Crypturornis, species :
Heterakis alata.
Subulura strongylina.
Crypturornis tataupa:
Subulura strongylina,
Crypturus cupreus (this host uniden-
tifiable) :
Heterakis arquata.
Heterakis valvata.
Orypturus noctivagus; see Crypturor-
nis noctivagus.
Orypturus parvirostris ;
crypturus parvirostris.
Crypturus, species; see Crypturornis,
species.
Crypturus tatuapa;
turus tataupa.
Cuculus cayanus; see Piaya cayana.
Cuculus melanorhynchus; see Coccy-
cus melacoryphus.
Cuculus naevius; see Tapera naevia.
Ouculus seniculus; see Coccyzus
minor.
Cuculus tingazu; see Piaya cayand.
Cuculus tinguacu; see Piaya cayanda.
Cunewma leucogastris:
Contracaecum haliacti.
Cupidonia cupido; see Tympanuchus
cupido.
Curucujus melanurus:
Cyrnea semilunaris.
see Micro-
see Microcryp-
Cyanocorax cayanus:
Heterakis gallinae.
Subulura papillosa,
Spiroptera saginata.
Acuaria mamillaris.
Cygnus atratus ; see Chenopis atrata.
Cygnus melanocoryphus; see Sthene-
lides melancoriphus.
404
Cygnus olor; see Sthenelides olor.
Cygnus olor domesticus; see Stheneli-
des olor domesticus.
Cypselus apus; see Micropus apus.
Dacelo leachii:
Thelazia dacelonis.
Dajila acuta:
Amidostomum anseris.
Epomidiostomum uncinatum.
Ascaridia galli.
Dandalus rubecula; see Erithacus ru-
becula.
Daption capensis:
Seuratia shipleyi.
Dichocercus bicornis; see Dichoceros
bicornis.
Dichoceros bicornis:
Microtetrameres contorta.
Dicholophus cristatus; see Cariama
cristata.
Dicholophus margravi; see Cariama
cristata.
Dicrurus adsimilis:
Hadjelia inermis.
Acuaria gracilis.
Diomedea exulans:
Seuratia shipleyi.
Tetrameres certa.
Diomedea melanophrys; see Thalas-
sarche melanophrys.
Diomedia exulans; see Diomedea ex-
ulans.
Diplopterus
naevia.
Dryobates major:
Syngamus mucronatus (nomen nu-
dum).
Dryocopus martius:
Acuaria quadriloba.
Eclectus pectoralis:
Ceratospira vesiculosa.
Eclectus roratus:
Habronema incerta.
“Weret’’ (see also Casmerodius and
Garzetia):
Porrocaecum reticulatum.
Elanus caeruleus:
Physaloptera acuticauda.
Hilanus leucurus:
Physaloptera acuticauda.
Electus pectoralis; see Hclectus pec-
toralis.
naevius; see Tapera
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Eleothreptus anomalus:
Subulura subulata.
Subulura suctoria.
Emberiza pecoris; see Molothrus ater.
“Engoulevents”’ (goatsuckers) =Cap-
rimulgidae:
Subulura leprincei.
Subulura subulata.
“ Hpervier ” ;=Accipiter nisus.
Hrithacus rubecula:
Acuaria subula.
Iiupodotis senegalensis:
Hadjelia parva.
Histiocephalus tridens.
EHupsittula pertinaz:
Habronema incerta.
EKupsittula solstitialis:
Ascaridia hermaphrodita.
Hurypyga helias:
Ancyracanthopsis bilabiata.
ELurystomus afer:
Subulura recurvata.
Subulura similis.
Euvenura maguari:
Heterakis valdemucronata.
Cyrnea excisa.
Lechinuria longeornata.
Echinuria longevaginata.
Falcinellus igneus; see Plegadis fal-
cinellus.
Falco aesalon; see Tinnunculus aesa-
lon.
Falco albicilla; 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 Cireaetus
gallicus.
Falco brasiliensis; see Polyborus
plancus.
Falco buteo; see Buteo buteo.
Falco cachinans; see Herpetotheres
cachinnans.
Falco cayennensis; see Leptodon pal-
liatus. »
NEMATODE PARASITES OF BIRDS
Falco cenchris; see Cerchneis nau-
manni.
Falco cherrug:
Porrocaecum depressum.
Habronema leptoptera.
Falco chrysaetos; see Aquila chry-
saetos.
Falco cineraceus; see Circus pygar-
gus.
Falco coronatus; see Harpyhaliaetus
coronatus.
Faleo cyaneus;
Falco degener;
chima.
Falco deiroleucus:
Habronema leptoptera.
Falco destructor; see Harpia harpyja.
Faico dispar; see Elanus leucurus.
Falco femoralis; see Rhynchofalco
coerulescens.
Falco furcatus;
catus,
Falco gallicus; see Circaetus gallicus.
Falco gracilis; see Geranospiza caeru-
lescens.
Falco haliaetus; see Pandion. haliae-
tus.
Falco imperialis; see Aquila heliaca.
Falco lagopus; see Buteo lagopus.
Falco lanarius; see Falco cherrug.
Falco lithofalco; see Tinnunculus reg-
ulus.
Falco longipennis:
Physaloptera alata.
Falco magnirostris; see Rupornis mag-
nirostris.
Falco milvus; see Milvus milwus.
Falco minutus; see Accipiter super-
ciliosus.
Falco naevius; see Aquila maculata.
Falco nisus; see Accipiter nisus.
see Circus cyaneus.
see Milvago chima-
see EHlanoides forfi-
Falco ornatus; see Spizaetus ornatus.
Falco palumbarius; see Astur gentilis.
Falco palustris; see Circus buffoni.
Falco pennatus; see Hieraaetus pen-
natus.
Falco peregrinus ; see Rhynchodon per-
egrinus,
Falco pygargus; see Circus pygargus.
Falco rufus; see Circus aeruginosus.
Falco rutilans; see Heterospizias mer-
idionalis.
405
Falco, species:
Physaloptera acuticauda.
Physaloptera alata.
Physaloptera subalata.
Falco subbuteo:
Habronema leptoptera.
Synhimantus denticulata.
Physaloptera alata.
Falco subbuteus; see Falco subbuteo.
Falco swainsonii; see Gampsonyr
swainsoni.
Falco tinnunculus; see Cerchneis tin-
nunculus.
Falco tridentatus; see Astur trivir-
gatus.
Falco unicinetus; see Parabuteo uni-
cinctus.
Falco urubutinga;
urubutinga.
Faleo xanthothoraz ; see Micastur rufi-
collis.
Ficedula ficedula:
Acuaria muscicapae.
Flammea flammea; see Tyto alba.
Florida caerulea:
Porrocaecum serpentulus.
Contracaecum andersoni.
Francolinus adspersus:
Subulura poculum.
Physaloptera brevicauda.
Francolinus bicalearatus:
Heterakis brevispiculum.
Ascaridia francolina.
Subulura differens.
Subulura suctoria.
Cheilospirura gruveli.
Francolinus gularis:
Heterakis longecaudata.
Francolinus sephaena:
Subulura suctoria.
see Urubutinga
Framcolinus, species :
Subulura gracilis.
Fregata aquila:
Contracaecum spiculigerum.
Fulica atra:
Amidostomum anseris.
Amidostomum fulicae.
Amidostomum raillieti.
Epomidiostomum orispinum.
Tetrameres fissispina.
Tetrameres globosa.
Hystrichis wedli.
406
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 Fulia marila.
Fuligula mollissima; see Somateria
mollissima.
Fuligula nigra; see Oidemia nigra.
Fuligula nyroca; see Nyroca nyroca.
Fuliex marila:
Amidostomum anseris.
Fulmaris glacialis:
Streptocara stellae-polaris.
Gallinago paraguaiae:
Tetrameres dubia.
Gallinula chloropus:
Amidostomum anseris.
Porrocaecum ensicaudatum.
Galloperdiz spadicea:
Heterakis longecaudata.
Subulura galloperdicis.
Gallus gallus:
Trichostrongylus tenuis.
Syngamus trachea.
Heterakis beramporia.
Heterakis brevispiculum.
Heterakis gallinae.
Heterakis pusilla.
Heterakis putaustralis.
Ascaridia brasiliensis.
Ascaridia compar.
Ascaridia compressa.
Ascaridia galli.
Ascaridia granulosa.
Ascaridia lineata.
Ascaridia styphlocerca,
Subulura brumpti.
Subulura differens.
Subulura strongylina.
Subulura suctoria.
Hartertia gallinarum.
Spirocerca sanguinolenta (larva).
Gongylonema ingluvicola.
Cheilospirura hamulosa.
Dispharyne nasuta,
Dispharyne« spiralis.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Gallus gallus—Continued.
Streptocara pectinifera.
Synhimantus laticeps.
Histiocephalus laticaudatus.
Physaloptera gemina.
Physaloptera truncata,
Oxyspirura mansoni.
Oxryspirura parvovum.
Tetrameres americana.
Tetrameres confusa.
Tetrameres fissispina.
Gallus lafayettii:
Heterakis pusilla.
Gampsonyt swainsoni:
Physaloptera acuticauda.
Physaloptera alata.
Garrulus glandarius:
Acuaria anthuris.
Garrulus glandarius cervicalis:
Dispharynz laplantei.
Garzetta garzetta:
Ascaridia aegyptiaca.
Gavia arctica:
Contracaecum spiculigerum.
Streptocara crassicauda.
Streptocara tridentata.
EHustrongylides tubifex.
Gavia stellata:
Contracaecum spiculigerum.
Cosmocephaius aduncus.
Streptocara crassicauda.
Eustrongylides mergorum.
Eustrongylides tubifez.
Gecinus viridis; see Picus viridis.
Gelochelidon nilotica:
Cosmocephalus obvelatus.
Ancyracanthopsis bihamata.
Gennaeus nycthemerus:
Heterakis gallinae.
Geopelia, species:
Ascaridia longecirrata.
Geotrygon montana; see Oreopelia
montana.
Geranospiza caerulescens:
Physaloptera acuticauda.
Physaloptera alata.
Thelazia papillosa.
Geranospizias caerulescens; see Ger-
anospiza caerulescens. —
Glareola austriaca; see Glareola prat-
incola.
Glareola pratincola:
Schistorophus spinulosus.
NEMATODE PARASITES OF BIRDS
Glaucidium passerinum:
Heterakis dispar.
Glaucionetta clangula:
Amidostomum anseris.
Epomidiostomum orispinum.
Contracaecum micropapillatum,
Contracaecum spiculigerum.
Streptocara crassicauda.
“ Goose”:
Ascaridia lineata.
Grossiptodon manschuricum ;
Crossoptilon mantchuricum.
“ Grouse”:
Trichostrongylus pergracilis.
Grus antigone; see Mathewsena anti-
gone.
Grus australiasiana; see Mathewsena
rubicunda.
Grus cinerea; see Megalornis grus.
Grus communis; see Megalornis grus.
Grus grus; see Megalornis grus.
Grus paradisea; see Tetrapteryxr
paradisea.
Grus pavonina; see Balearica pavo-
nina,
Grus_ viridirostris;
japonensis,
Guara rubra:
Sciadiocara wmbellifera.
Guira guira:
Subulura forcipata.
Gypaetus barbatus:
Porrocaecum depressum.
Gyps fulwus:
Porrocaecum depressum.
Haematopus ostralegus:
Contracaecum spiculigerum.
Schistorophus aulieatina.
Halcyon chelicutensis; see Chelicutia
chelicuti.
Haliaeetus albicilla:
Porrocaecum angusticolle.
Porrocaecum depressum.
Hatliaeetus leucocephalus:
Contracaecum haliaéti.
Haliactus leucogaster; see Cuncuma
leucogastris.
Halieus brasiliensis; see Phalacrocor-
ar vigua.
Harelda glacialis; see Clangula hye-
malis.
Harpagus bidentatus:
Habronema leptoptera.
3612—27 28
see
see Megalornis
407
Harpia harpyja:
Thelazia papillosa.
Harpyhaliaétus coronatus:
Physaloptera acuticauda,
Physaloptena alata.
“ Hawk”:
Physaloptera alata chevreuvi.
Heliotreptus anomalus; see Eleothrep-
tus anomalus.
Helotarsus albicilla;
albicilla.
Helotarsus ecaudatus ; see Terathopius
ecaudatus.
Yerodias egretta: see Casmerodius alba
egretta.
Herodias tricholor;
tricolor.
“ Heron cendre’’—Ardea cinerea.
“Héron pourpré’’ (purple heron) =
Phoyx purpurea.
Herpetotheres cachinnans:
Habronema leptoptera.
Physaloptera acuticauda.
Physaloptera alata.
Heterospizias meridionalis:
Physaloptera acuticauda.
Porrocaecum depressum.
Hieraaetus fasciatus:
Porrocaecum depressum.
Hieraaetus pennatus:
Porrocaecum depressum.
Physaloptera alata.
Himantopus himantopus:
Amidostomun chevreusi.
Porrocaecum ensicaudatum.
Porrocaecum heteroura.
Chevreuria revoluta.
Tetrameres nouveli.
Himantopus melanopterus ; see Himan-
topus himantopus.
ITirundo riparia; see Riparia riparia.
Hirundo rustica:
Acuaria attenuata. .
Hirundo urbica; see Chelidonaria
urbica.
Houbara macqueent:
Subulura suctoria.
Hartertia rotundata.
Houbara undulata:
Subulura rima.
Subulura suctoria.
Hartertia confusa.
Hartertia rotundata,
see Haliaeetus
see Hydranassa
408
Hydranassa tricolor:
Contracaecum microcephalum.
Hydrochelidon nigra; see Chlidonias
nigra.
Hydroprogne caspia:
Schistorophus acanthocephalicus.
Hydropsalis climacocerca:
Subulura subulata.
Subulura suctoria.
Hylotomus pileatus; see Phloeotomus
pileatus.
Hypotriorchis subbuteo; 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:
Oxryspirura tanasijtchuki.
Icterus cristatus; see Ostinops decu-
manus.
Icterus croconotus:
Ozyspirura cephaloptera.
Irrisor erythrorhynchus; see Phoeni-
culus purpureus.
Ithagenes cruentus;
eruentus.
Ithaginis cruentus:
Heterakis isolonche.
Ascaridia galli.
Izobrychus 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 pergracilis.
Syngamus trachea.
Heterakis gallinae.
Lampronessa sponsa; see Aiz sponsa.
see Ithaginis
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Lanius collurio:
Viguiera euryoptera.
Acuaria cordata.
Lanius excubitor:
Viguiera euryoptera.
Lanius excubitor dodsoni:
Spirocerca sanguinolenta (larva).
Lanius minor:
Hartertia zakharowi.
Viguiera euryoptera.
Cheilospirura rotundata.
Physaloptera bilabiata.
Lanius rufus; see Lanius senator.
Lanius senator:
Viguiera euryoptera.
Acuaria cordata.
Lanius, species:
Microtetrameres inermis.
Larus argentatus ; 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 mazrimus; see Larus marinus.
Larus medius; see Larus hyperboreus.
Larus ridibundus:
Cyathostoma. lari.
Contracaecum spiculigerum.
Cosmocephalus aduncus.
Cosmocephalus diesingi.
Cosmocephalus obvelatus.
Streptocara tridentata. .
Larus, species:
Cyathostoma lari.
Rusguniella elongata.
Larus tridactylus; see Rissa tridac-
tyla.
NEMATODE PARASITES OF BIRDS
Larus varius:
Contracaecum spiculigerum.
Cosmocephalus aduncus.
Cosmocephalus obvelatus.
Lepidoenas speciosa:
Ascaridia columbae.
Leptodon cayennensis; see Leptodon
palliatus.
Leptodon palliatus:
Physaloptera acuticauda.
Physaloptera alata.
Leptoptila rufavilla;
rufavilla.
Leptoptilos crumeniferus:
Echinuria leptoptili.
Eustrongylides africanus.
Leptoptilos dubius-.
Echinuria hargilae.
Leptoptilus crumenifer ;
tilos crumeniferus.
Leptoptilus dubius;
dubius.
Leptotila rufazilla:
Ornithostrongylus fariai.
Lestris parasitica; see Stercorarius
parasiticus.
Lestris pomarinus;
pomarinus.
Leucopternis albicollis:
Habronema. leptoptera.
Limnocryptes gallinula:
Eechinuria horrida.
Lophoceros semifasciatus:
Hadjelia inermis.
Lophophorus impejanus:
Heterakis isolonche.
Heterakis longecaudata.
Lophophorus impeyanus; see Lopho-
phorus impejanus.
Lophortyz californicus:
Habronema incerta,
Lophostriz cristata:
Spiroptera penihamata.
Lophotis ruficrista:
Hartertia obesa.
Hartertia rotundata.
Lurocalis semitorquatus:
Subulura subulata.
Subulura suctoria,
Luscinia luscinia:
Porrocaecum ensicaudatum.
Luscinia philomela; see Luscinia lus-
cinta.
see Leptotila
see Leptop-
see Leptoptilos
see Coprotheres
409
Luscinia rubecula; see Hrithacus rube-
cula.
Lyrurus tetriaz:
Ornithostrongylus hastatus.
Ascaridia compar.
Ascaridia magnipapilla.
Mackenziaena leachii:
Spiroptera saginata.
Macrodipteryz longipennis:
Subulura leprincei.
Macrodipteryx macrodipterus;
Macrodipteryz longipennis.
Macropygia nigrirostris:
Ascaridia australis.
Malacoptila torquata:
Subulura forcipata.
Subulura strongylina,
Subulura travassosi.
Mareca penelope:
Amidostomum anseris.
Epomidiostomum orispinum.
Epomidiostomum uncinatum.
Echinuria uncinata,.
Mathewsena antigone:
Ascaridia cristata.
Ascaridia stroma.
Mathewsena rubicunda:
Porrocaecum serpentulus.
Megacephalon maleo:
Heterakis longecaudata.
Megaloperdiz himalayanus:
Heterakis macroura.
Megaloperdiz nigelii; see Megaloper-
diz himalayanus.
Megalopterus hawaiiensis; see Micra-
nous hawatiensis,
Megalornis grus:
Porrocaecum serpentulus.
Ascaridia stroma.
Schistorophus bicuspis.
Megalornis japonensis:
Cyathostoma variegatum.
Megaquiscalus major:
Thelazia cirrura.
Melanerpes flavifrons; see Centurus
flavifrons.
Melanitta deglandi:
Yseria californica.
Melanitta fusca:
Amidostomum acutum.
Amidostomum anseris.
Epomidiostomum orispinum.
Streptocara crassicauda.
see
410
Meleagris galiopavo:
Synganus trachea.
Heterakis gallinae.
Subulura brumpti.
Subulura suctoria.
Cheilospirura hamulosa.
Dispharynaz spiralis.
Oryspirura mansoni.
Tetrameres confusa.
Tetrameres fissispina.
Melierax gabar:
Physaloptera galinieri.
Merganser castor; see Mergus mer-
ganser.
Merganser serratus; see Mergus ser-
rator.
Mergellus albellus:
Eustrongylides mergorum.
Mergus albellus; see Mergellus albel-
lus.
Mergus merganser:
Contracaecum spiculigerum.
Streptocara crassicauda.
Tetrameres fissispina.
Eustrongylides mergorum.
Hystrichis coronatus.
Mergus serrator:
Cosmocephalus obvelatus.
Streptocara crassicauda.
EHustrongylides mergorum.
Hystrichis varispinosus.
Merops apiaster:
Schistorophus bidens.
Merula merula; see Turdus merula.
Merula nigra; see Turdus merula.
Micranous 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
cristata.
Micropogon, species ; see Capito, species.
Micropus apus:
Syngamus trachea.
Milwago chimachima:
Porrocaecum depressum.
Milvus ater; see Milwus migrans.
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Milvus govinda; see Milvus migrans
govinda.
Milwus ictinus; see Milwous milvus.
Milvus korschum; see Milvus migrans.
Milvus migrans:
Porrocaecum depressum.
Habronema leptoptera.
Habronema mansioni.
Dispharyne rectovaginata.
Milous migrans govinda:
Porrocaecum angusticolle.
Milvus milvus:
Porrocaecum depressum.
Habronema leptoptera.
Milvus regalis; see Milvus milvus.
Mimus polygiottos:
Porrocaecum ensicaudatum.
Mimus polyglottus; see Mimus poly-
glottos.
Molothrus ater:
Habronema leptoptera.
Momotus brasiliensis; see Momotus
momota.
Momotus momota:
Ozyspirura cephaloptera.
Monacha morpheus; see Monasa mor-
phoeus.
Monacha nigra; see Monasa nigra.
Monasa leucops; see Monasa mor-
phoeus.
Monasa morphoeus:
Subulura foreipata.
Subulura strongylina.
Subulura travassosi.
Monasa nigra:
Subulura forcipata.
Subulura strongylina.
Subulura travassosi.
Monasa nigrifrons:
Subulura travassosi.
Monasa tenebroso; see Chelidoptera
tenebrosa.
Monasa torquata; see Malacoptila tor-
quata.
Monasa tranquilla; see Monasa nigra.
Monticola sazatilis:
Porrocaecum ensicaudatum.
Motacilla alba:
Porrocaecum ensicaudatum.
“Mouette’”’ (European gull) ;=Larus,
species.
Muscicapa
ficedula.
atricapilla; see Ficedula
NEMATODE PARASITES OF BIRDS
Mycteria americana:
Contracaecum multipapillatum.
Myiopsitta monachus:
Habronema incerta.
“Nandou "=—Rhea rothschildi.
Necrosyrtes monachus:
Porrocaecum depressum.
Neophron monachus; see Necrosyrtes
monachus.
Neophron perenopterus; see Neophron
perenopterus.
Neophron perenopterus:
Habronema tulostoma.
Nesochen sandvicensis:
Heterakis dispar.
Streptocara crassicauda.
Nettion carolinense:
Echinuria uncinata.
Nettion erecca:
Amidostomum acutum.
Amidostomum anseris.
Epomidiostomum orispinum.
Nicticorax violacens; see Nyctanassa
violaceda.
Nisaetus fasiatus; see Hieraaetus fas-
ciatus.
Nisus communis; see Accipiter nisus
Nonnula rubecula:
Subulura strongylina.
Subulura travassosi.
Nothocraz urumutum:
Oxryspirura heteroclita.
Nothura maculosa:
Subulura olympioi.
Nucifraga caryocatactes:
Synganwus parvus,
Acuaria anthuris.
Eustrongylides papillosus.
Numenius arquatus:
Schistorophus longicornis.
Eustrongylides mergorum.
Hystrichis neglectus.
Numida meleagris:
Heterakis brevispiculum.
Heterakis gallinae.
Ascaridia compar.
Ascaridia galli.
Ascaridia numidae.
Subulura differens.
Dispharyne spiralis.
Streptocara pectinifera.
Numida mitrata rikwae:
Subulura acuticauda.
411
Numida papillosa transvaalensis:
Ascaridia numidae.
Subulura suctoria.
Numida ptilorhyncha; see Numida me-
leagris.
Numida rikwae; see Numida mitrata
rikwae.
Nyctale tengmalmi; see Cryptoglaur
funerea.
Nyctanassa violacea:
Porrocaecum serpentulus.
Tetrameres micropenis.
Nyctea nyctea:
Porrocaccum depressum.
Porrocaccum spirale.
Nyctiardea grisea; see
nycticorar.
Nyctibius aethereus:
Subulura subulata.
Subulura suctoria.
Nyctibius grandis:
Subulura subulata.
Subulura suctoria.
Nycticorar
Nyctibius griseus jamaicensis:
Subulura forcipata.
Subulura strongylina.
Subulura suctoria,
Nyctibius jamaicensis; see Nyctibius
griseus jamaicensis.
Nycticorax europacus; see Nycticorar
nycticoraz.
Nycticoraz griseus;
nycticorax.
see Nycticorar
Nycticoraxr nycticorar:
Porrocaecum reticulatum.
Porrocaecum serpentulus.
Contracaecum microcephalum.
Contracaecum rosarium.
Synhimantus sagittata.
Tetrameres gynaecophila.
Nycticorazr, species :
Contracaecum rosarium,
Nyctidromus albicollis:
Subulura subulata,
Subulura suctoria.
Spiroptera saginata.
Nyctidromus guianensis derbyanus:
Subulura forcipata.
Nyctipolus nigrescens:
Subulura suctoria.
Nyctiprogne leucopyga:
Spiroptera saginata,
412
Nyroca clangula; see Glaucionetta
clangula.
Nyroca ferina; see Aithyia ferina.
Nyroca fuligula; see Fuligula fuli-
gula.
Nyroca marila; see Fuliv marila.
Nyroca nyroca:
Tetrameres, species.
Odontophorus capueira:
Heterakis fariai.
Subulura strongylina.
Odontophorus guianensis:
Subulura strongylina.
Oedicnemus capensis:
Hartertia obesa.
Oedicnemus crepitans ; see Oedicnemus
oedicnemus.
Oedicnemus oedicnemus:
Porrocaecum ensicaudatum.
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 anseris.
Amidostomum monodon.
Epomidiostomum orispinwn.
Epomidiostomum uncinatum.
Oreopelia montana:
Ascaridia magathaesi.
Oriolus auratus:
Hadjelia inermis.
Acuaria gracilis.
Oriolus galbula; see Oriolus oriolus.
Oriolus oriolus:
Acuaria cordata.
Ortyx virginianus; see Cotinus vir-
ginianus,
Ostinops decumanus:
Spiroptera saginata.
Otis afroides; see Afrotis afra afra-
oides,
Otis houbara; see Houbara undulata.
Otis macqueeni; see Houbara mac-
queeni.
Otis ruficresta; see Lophotis rufi-
crista,
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Otis tarda:
Trichostrongylus tenuis.
Ornithostrongylus papillatus.
Syngamus trachea.
Heterakis gallinae.
Heterakis papillosa.
Otis tetra:
Heterakis gallinae.
Heterakis papillosa.
Subulura forcipata.
Histiocephalus laticaudatus.
Otus brachyotus; see Otus flammeus.
Otus choliba:
Subulura acutissima.
Spiroptera penihamata.
Spiroeptera 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.
Oxryspirura mansoni.
Pavo muticus:
Heterakis hamulus.
Pseudaspidodera pavonis.
Cyrnea bulbosa.
Cheilospirura pavonis.
Pavo spicifer; see Pavo muticus.
Pelecanus americanus; see Pelecanus
erythrorhynchos.
:
NEMATODE PARASITES OF BIRDS
Pelecanus carbo; see Phalacrocorar
carbo.
Pelecanus conspicillatus:
Contracaecum spiculigerum.
Pelecanus crispus:
Contracaecum micropapillatum.
Pelecanus erythrorhynchos:
Contracaecum spiculigerum,
Pelecanus fuscus; see Pelecanus oc-
cidentalis.
Pelecanus occidentalis:
Contracaecum spiculigerum.
Pelecanus onocrotalus:
Syngamus trachea.
Contracaecum spiculigerum.
Gnathostoma 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.
Perdiz cinerea; see Perdixz perdiz.
Perdiz coturniz; see Coturniz cotur-
ni.
Perdix dentata;
guianensis.
Perdixz graeca; see Alectoris graeca.
“Perdix grecque”; see Alectoris
graeca.
Perdix perdiz:
Trichostrongylus tenuis.
Syngamus trachea.
Heterakis gallinae.
Ascaridia compar.
“Perdix rouge ’’=Alectoris rufa.
Perdiz sazwatilis; see Alectoris graeca
saxratilis.
Perdiz, species:
Heterakis brasiliana.
“ Perdrix de roche ””=Alectoris graeca
saratilis.
see Odontophorus
413
Pernis apivorus:
Porrocaecum depressum,
Physaloptera alata.
Pernis, species:
Porrocaecum angusticolle.
Phalacrocoraz aristotelis:
Contracaecum spiculigerum.
Phalacrocoraz ater:
Contracaecum spiculigerum.
Phalacrocoraxz auritus:
Contracaecum spiculigerum.
Phalacrocoraxr carbo:
Syngamus microspiculum,
Contracaecum spiculigerum.
Desmidocerca aerophila.
Echinuria squamata.
Eustrongylides mergorum.
Eustrongylides excisus.
Phalacrocorax fuscicollis:
Contracaecum spiculigerum.
Phalacrocoraz graculus; see Phalacro-
corax aristotelis.
Phalacrocorar javaniscus:
Contracaecum spiculigerum.
Phalacrocoraz pelagicus:
Contracaecum spiculigerum.
Phalacrocorar pygmaeus:
Contracaecum spiculigerum.
Bustrongylides excisus.
Phalacrocorar sulcirostris ; see Phala-
crocoraz ater.
Phalacrocorar urile:
Contracaecum spiculigerum.
Phalacrocorar verrucosus:
Contracaecum spiculigerum.
Streptocara cirrohamata.
Phalacrocorax vigua:
Contracaecum spiculigerum.
Phasianus chrysomelas ; see Phasianus
colchicus chrysomelas.
Phasianus colchicus:
Trichostrongylus tenuis.
Syngamus trachea.
Heterakis gallinae.
Heterakis isolonche.
Heterakis neoplastica.
Phasianus colchicus chrysomelas:
Heterakis isolonche.
Phasianus colchicus satscheuensis:
Heterakis neoplastica.
Phasianus gallus; see Gallus gallus.
Phasianus nycthemerus ; see Gennaeus
nycthemerus.
414
Phasianus pictus; see Chrysolophus
pictus.
Phasianus
reevesii.
Phasianus satscheunensis; see Phasi-
anus colchicus satscheuensis.
Phasianus veneratus; see Syrmaticus
reevesit.
Phasianus versicolor:
Heterakis gallinae.
Phimosus infuscatus ;
nudifrons.
Phimosus nudifrons:
Hystrichis acanthocephalicus.
Phlegoenas luzonica:
Ascaridia columbae.
Phloectomus pileatus:
Acuaria quadriloba.
Phlogoenas luzonica; see Phlegoenas
luzonica.
Phoenicopterus roseus:
Porrocaecum serpentulus.
Echinuria phoenicopteri.
Tetrameres coccined.
Phoethornis pretrei:
Serticeps vulvoinflatus.
Phoyxs purpurea:
Porrocaecum. reticulatum.
Porrocaecum serpentulus.
Contracaecum microcephalum.
Synhimantus invaginata.
Synhimantus sagittata.
Piaya cajanea; see Piaya cayana,
Piaya cayana:
Subulura carlosi.
Subulura forcipata.
Subulura reclinata.
Subulura strongylina.
Subulura subulata.
Spiroptera saginata.
Pica caudata; see Pica pica.
Pica pica:
Syngamus pugionatus
dum).
Syngamus trachea.
Porrocaecum ensicaudatum.
Acuaria anthuris.
Acuaria cordata.
Picus campestris; see Colaptes campe-
stris.
Picus canus:
Syngamus mucronatus (nomen nu-
dum).
Syngamus trachea,
reevesi; see Syrmaticus
see Phimosus
(nomen nu-
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Picus grammicus; see Celeus gram-
micus.
Picus jumana; see Celeus jumana.
Picus major; see Dryobates major.
Picus martius; see Dryocopus mar-
tius.
Picus viridis:
Syngamus trachea,
Acuaria quadriloba.
“Pie griéche 4 Téte rouge ”’=Lanius
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 festiva.
Pionus (Psittacus) leucocephala; see
Amazona leucocephala.
Pionus (Psittacus) leucotis; see Pyr-
rhura leucotis.
Pionus (Psittacus) menstruus;
Pionus menstruus.
Pionus (Psittacus) pertinaz; see Ara-
tinga pertinaz.
Pionus (Psittacus) phoenicurus; see
Pyrrhura molinae.
Pionus (Psittacus) pulverulentus; see
Amazona farinosa.
Pionus (Psittacus)
Pionus fuscus.
Pionus (Psittacus), species; see Psit-
tacus, species.
Pionus (Psittacus) sulfureus; see Ka-
katoe sulphurea.
Pionus (Psittacus) vinaceus ; see Ama-
zona vinacea.
Pisorhina atricapilla; see.Otus choliba.
Platalea ajaja; see Ajaia ajaja.
Platalea leucorodia:
Tetrameres coccinea.
Eustrongylides africanus.
Platycercus adscitus:
Habronena incerta.
see
purpureus; see
NEMATODE PARASITES OF BIRDS
Piatycercus barnardi; see Barnardius
barnardi.
Platycercus eximius:
Habronema incerta.
Platycercus palladiceps; see Platycer-
cus adscitus.
Platycichla flavipes:
Microtetrameres pusilla.
Plegadis falcinellus:
Echinuria contorta.
Hystrichis orispinus.
Plegadis guarauna:
Echinuria calearata.
Plocepasser mahali:
Hartertia obesa.
Hlartertia rotundata.
Plotus anhinga; see Anhinga anhinga.
Ploius lavaillanti; see Anhinga rufa.
Plotus melanogaster; see Anhinga me-
lanogastris.
Plotus novae-hollandiae; see Anhinga
novaehollandiae.
Plotus rufus; see Anhinga rufa.
Pluvialis apricaria:
Porrocaccum ensicaudatum.
Porrocaecum heteroura.
Porrocaecum semiteres.
Eustrongylides mergorum,
Podager nacunda:
Ascardia amblymoria,
Subulura forcipata.
Subulura strongylina,
Subulura subulata.
Subulura suctoria.
Podiceps auritus; see Colymbus auri-
tus.
Podiceps capensis; see Colymbus rufi-
collis capensis.
Podiceps cristatus; see Colymbus cris-
tatus.
Podiceps dominicensis; see Colymbus
dominicus.
Podiceps fluviatilis ; see Colymbus rufi-
collis.
Podiceps minor; see Colymbus rufi-
collis.
Podiceps nigricollis ;
nigricollis.
Poeocephalus senegalus; see Poicepha-
lus senegalus.
Poicephalus senegalus:
Habronema incerta.
see Colymbus
415
Polyborus plancus:
Porrocaecum depressum.
Procellaria anglorum; see Puffinus
puffinus.
Prodiceps
cristatus,
Prodiceps minor; see Colymbus rufi-
collis.
Protogerys virescens; see Brotogeris
virescens,
Pseudotantalus ibis; see Ibis ibis.
Psittacula fasciata:
Habronema incerta.
Psittacus sinensis; see Eclectus pec-
toralis.
cristatus; see Colymbus
Psittacus, species:
Ascauridia hermaphrodita.
Psophia viridis:
Heterakis arquata.
Heterakis psophiae.
Pternistes, species; see Pternistis,
species,
Pternistes swainsoni;
swainsoni.
Pternistis, species:
Hartertia annulata.
Pternistis swainsoni:
Subulura suctoria.
Ptilopachus petrosus:
Acuaria ptilopachydis.
Ptilopachys fuscus; see Ptilopachus
petrosus.
Puffinus kuhli; see Calonectris kuhlii.
Puffinus pujfinus:
Seuratia procellariae.
Pulsatriz perspicillata:
Spiroptera saginata,
Tetrameres paradozra.
Pyrrhocoras alpinus; see Pyrrhocorar
graculus.
Pyrrhocorax graculus:
Syngamus trachea.
Acuaria cordata.
Pyrrhocorax pyrrhocoraz:
Acuaria anthuris.
Pyrrhura leucotis:
Ascaridia hermaphrodita.
Habronema incerta.
Pyrrhura molinae:
Ascaridia hermaphrodita,
Querquedula circia; see Querquedula
querquedula.
see Pternistis
416
Querquedula Nettion
erecca.
Querquedula querquedula:
Amidostomum anseris.
Hystrichis neglectus.
Quiscalus major; see Megaquiscalus
major.
Quiscalus quiscula:
Dispharyna spiralis.
Rallus cayennensis ; see Creciscus viri-
dis.
Ramphastos monilis:
Habronema unilateralis.
Ramphastos, species:
Thelazia digitata.
Ramphastos vitellinus:
Habronema unilateralis.
Dispharyne crassissima.
“ Rapace nocturne” (owl):
Habronema monoptera.
Ramphastus, species; see Ramphastos,
species.
Rhamphastos erythrorhynchus ;
Ramphastos monilis.
Rhamphastos vitellinus ; see Ramphas-
tos vitellinus.
Rhea americana; see Rhea rothschildi.
Rhea rothschildi:
Deletrocephalus dimidiatus.
Heterakis parisi.
Ascaridia orthocerca.
Spirura uncinipenis.
Spirura zeschokket.
Rhizothera longirostris:
Heterakis interlabiata.
Rhynchodon peregrinus:
Porrocaecum depressum.
Rhynchofalco coerulescens:
Synhimantus recta.
Rhynchotus rufescens:
Heterakis brasiliana.
Subulura olympioi.
Riparia riparia:
Acuaria attenuata.
Rissa tridactyla:
Contracaecum spiculigerum.
Rupornis magnirostris:
Habronema leptoptera.
Thelazia campanulata.
Salicaria turdoides; see Acrocephalus
arundinaceus.
Sazicole rubetra:
Acuaria tenuis.
Scolopaxr gallinula; see Limnocryptes
gallinula.
crecca; see
see
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Scolopax major; see Capella media.
Scops leucotis; see Otus leucotis.
Sephina francolinus; see Francolinus
sephaena.
Setochalcis rufa:
Subulura subulata.
Subulura suctoria.
Setochalecis vocifera:
Subulura forcipata.
Subulura suctoria.
Somateria dresseri:
Amidostomum anseris.
Somateria mollissima:
Amidostomum acutum.
Amidostomum mollissima.
Epomidiostomum orispinum.
Eustrongylides mergorum.
Spizaetus mauduyti; see Spizaetus or-
natus.
Spizaeius ornatus:
Physaloptera acuticauda.
Physaloptera alata.
Spizastur melanoleucus:
Physaloptera acuticauda.
Physaloptera alata.
Spiziaster melanoleucus; see Spizas-
tur melanoleucus.
Squatarola helvetica; see Squatarola
squatarola.
Squatarola squatarola:
Porrocaecum ensicaudatum.
Porrocaecum heteroura.
Porrocaecum semiteres.
Schistorophus bicuspis.
Stenopsis candicans; see Thermochal-
cis candicans.
Stercorarius 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.
Sthenelides melancoriphus:
Tetrameres fissispina.
Sthenelides olor:
Heterakis circumvallata.
Hystrichis cygni.
H ystrichis orispinus.
NEMATODE PARASITES OF BIRDS
Sthenelides olor domesticus:
Echinuria uncinata,.
Stictoenas arquatriz;
arquatriz.
“Stork, black ’=Ciconia nigra.
Streptopelia risoria:
Ascaridia columbdae.
Streptopelia turtur:
Ascaridia columbae.
Striz alba; see Tyto alba.
Strie albomarginata; see Ciccaba hu-
hula.
Strix aluco:
Porrocaecum depressum.
Porrocaecum spirale.
Strix atricapilla; see Otus choliba.
Striz brachyotus; see Asio flammeus.
Striz bubo; see Bubo bubo.
Strix dasypus; see Cryptoglaur fu-
nered.
Strix flammea; see Tyto alba.
Strix griseata; see Lophostrir cris-
tata.
Strix nivea; see Nyctea nyctea.
Striz noctua; see Carine noctua.
Strix nyctea; see Nyctea nyctea.
Strix otus; see Asio otus.
Striv passerina; see Carine noctua.
Strix, species:
Subulura lutzi.
Spiroptera penihamata.
Strix stridula; see Strix aluco.
Striz tangmalmi; see Cryptoglauc fu-
nered.
Striz torquata; see Pulsatriz perspicil-
lata.
Struthio australis;
melius.
Struthio camelus:
Ornithostrongylus douglasi.
Codiostomum struthionis.
Struthio camelus molybdophanes:
Codiostomum struthionis.
Struthio molybdophanes; see Struthio
camelus molybdophanes.
Sturnus vulgaris:
Syngamus pugionatus (nomen nu-
dum).
Syngamus trachea.
Porrocaecum ensicaudatum.
Porrocaecum heteroura.
Spiroptera turdi.
Surnia passerina; see Glaucidium pas-
serinum.
see Columba
see Struthio ca-
417
Sylvia palustris;
palustris.
Sylvia rubecula; see Hrithacus rube-
cula,
Sylvia turdoides;
arundinaceus.
see Acrocephalus
see Acrocephalus
Syrmaticus reevesti:
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 Mycteria
americana.
Tapera naevia:
Subulura forcipata.
Subulura subulata.
Terathopius ecaudatus:
Porrocaecum angusticolle.
Terpsiphone, species:
Hadjelia inermis.
| Tetrao bonasia; see Tetrastes bonasia.
Tetrao lagopus; see Lagopus lagopus.
Tetrao tetriz; see Lyrurus tetriz.
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
Thamnophilus funebris; see Macken-
ziaena leachii.
Thaumalea ambherstiae;
lophus amherstiae,
see Chryso-
Thaumalea obscurus; see Chryso-
lophus pictus obscurus.
Thaumalea picta; see Chrysolophus
pictus.
Thermochalcis candicans:
Subulura subulata.
Subulura suctoria.
Threskiornis aethiopicus:
Physaloptera, species Parona.
Tinamus, species :
Heterakis alata.
Heterakis skrjabini.
Ascaridia strelnikowi.
Subulura strongylina.
Tinamus tataupa; see Microcrypturus
tataupa.
Tinnunculus aesalon:
Porrocaecum depressum.
Tinnunculus alaudarius ; see Cerchneis
tinnunculus.
Tinnunculus regulus:
Porrocaecum depressum.
Totanus fuscus; see Totanus macu-
latus.
Totanus glottis;
larius.
Totanus hypoleucus; see Actitis hypo-
leuca.
Totanus maculatus:
Cosmocephalus obvelatus.
Totanus melanoleucus:
Sciadiocara umbellifera.
Totanus nebularius:
Schistorophus longicornis.
Trachelotis senegalensis ;
dotis senegalensis.
Tragopan satyra:
Heterakis bosia.
Heterakis gallinae.
Heterakis isolonche.
Heterakis longecaudata.
Tringa alpina; see Pelidna alpina.
Tringa helvetica; see Squatarola
squatarola.,
Tringa vanellus; see Vanellus vanellus.
Tringa variabilis; see Pelidna alpina.
Tringae vanelli; see Vanellus vanellus.
Tringoides hypoleucus; see Actitis
hypoleuca.
see Eupo-
see Totanus nebu- .
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Trochilus ochropygus; see Phoethornis
pretrei.
Trogon collaris; see Trogonurus col-
laris.
Trogon mellanurus ;
melanurus,
see Curucujus
Trogon, species:
Subulura bentocruzi.
Trogon variegatus:
Subulura bentocruzi.
Trogon viridis:
Subulura trogoni.
Trogonurus collaris:
Cyurnea semilunaris,
T'rypanocorax frugilegus:
Syngamus trachea.
Acuaria anthuris.
Acuaria cordata.
Oxryspirura sygmoided.
Microtetrameres inermis.
Turdus iliacus; see Arceuthornis
musicus.
Turdus merula:
Porrocaecum ensicaudatum.
Spiroptera turdi.
Turdus musicus;
philomelos.
Turdus pilaris; see Arceuthornis pi-
laris.
Turdus rufiventris:
Microtetrameres pusilla.
Turdus saxatilis; see Monticola sara-
tilis.
Turdus torquatus;
torquatus.
Turdus viscivorus;
viscivorus.
Turniz javanica taijoor:
Oxvyspirura ophthalinica.
Turnixz, species :
Subulura, species Baylis and Daub-
see Arceuthornis
see Arceuthornis
see Arceuthornis
ney.
Turnie taigoor; see Turniz javanica
taijoor.
Turtur sylvaticus; see Streptopelia
turtur.
Tympanuchus cupido:
Heterakis gallinae.
Tyto alba:
Porrocaecum depressum.
Porrocaecum spirale.
NEMATODE PARASITES OF BIRDS
Tyto alba—Continued.
Spiroptera penihamaia.
Synhimantus affinis,
Synhimantus laticeps.
Ulula aluco; see Strix aluce.
Upupa epops:
Hadjelia truncata.
Spirocerca sanguinolenta.
Uria aalge:
Contracaecum spiculigerum,
Eustrongylides mergorum.
Uria grylle; see Cepphus grylle.
Uria troile; see Uria aalge.
Urocissa erythrorhyncha occipitalis:
Acuaria anthuris.
Urocissa occipitalis; see Urocissa ery-
throrhyncha occipitalis.
Urubitinga wrubitinga:
Physaloptera acuticauda.
Physaloptera alata.
Utamania torda; see Alca torda.
419
Vanellus cristatus; see Vanellus va-
nellus.
Vanellus melanogaster; see Squatarola
squatarola.
Vanellus vanellus:
Amidostomum henryi.
Porrocaecum ensicaudatum.
Porrocaecum semiteres.
Rusguniella vanelli.
Streptocara crassicauda charadrii.
Vinago delalandii:
Ascaridia fasciata,
Vultur cinereus; see Aegypius mona-
chus.
Vultur fulvus; see Gyps fulvus.
Vultur monachus; see Aegypius mona-
chus.
Vultur percnopterus;
perenopterus.
Zonoenas brenchleyi:
Ceratospira ophthalmica.,
see Neophron
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1839—Hingeweidewiirmer, Binnenwiirmer, Thierwiirmer. Allg. Encyd. d.
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1846—Nachtriige zu Gurlt’s Verzeichniss der Thiere, bei welchen Entozoen
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1853—Eingeweidewiirmer des Dicholophus cristatus. Abhandl. d. naturf.
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1851—Systema helminthum. vol. 2, vit+-588 pp., Vindobonae.
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1861—Revision der Nematoden. Sitzungsb. d. k. Akad. d. Wissensch.,
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1883a—Helminthologische Notizen. Verhandl. d. k. k. zool.-bot. Gessellsch.
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(1883), vol. 33, pp. 107-118, pls. 3-5; pp. 193-218, pls. 11-14.
422 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
DUJARDIN, FELIX.
1843—-Mémoire sur les helminthes des musaraignes, et en particulier sur
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1863—Untersuchungen tiber Nematoden., 77 pp., 9 pls. Leipzig.
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1885—Observation sur un helminthe parasite de VOssifraga gigantea.
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1923—Disease in captive wild mammals and birds, incidence, description,
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1923—Nicotine as a poultry vermifuge. Science, N. Y., new ser., vol. 57,
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1789—Beschreibungen einiger neuen Hingeweidewtirmer. Naturforscher,
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NEMATODE PARASITES OF BIRDS 423
GENDRE, E.—Continued.
1920a—Sur une espéce nouvelle d’ “ Acuaria,’ parasite de “ Ptilopachys
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19206—Un genre nouveau d’ “Acuariinae.”’ Actes Soe. Linn. de Bor-
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1921a—Sur deux espéces de nématodes africains. Actes Soc. Linn. de
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1H.
1922.—Sur quelques espéce d’ “ Habronema,’ parasites des oiseaux. Actes
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[1790}—Caroli 4 Linné ... Systema naturae per regna trianaturae, secun-
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1885—A new entozoon from Struthio molybdophanes, Rehw. Notes Ley-
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1920—Nematode parasites of the domestic pigeon (Columbia livia domes-
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1912—Notes on some Entozoa. Proc. Roy. Soc. Queensland, vol. 24, pp.
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424 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
KRAEMER, A.
1853—Fragmentarische Notizen und Abbildungen zur Helminthologie and
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1856—A synopsis of Entozoa and some of their ectocongeners observed
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1855—Beitriige zur Anatomie der Nematoden. Arch. f. Anat., Physiol. u.
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1873—Hinige neue Nematoden, nebst Bemerkungen tiber bekannte Arten.
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1878a—Compendium der Helminthologie. Ein Verzeichniss der bekann-
ten Helminthen, die frei oder in thierischen Koérpern leben, geord-
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NEMATODE PARASITES OF BIRDS 425
yon LiNstow, O.—Continued.
1879b—Helminthologische Untersuchungen. Jahresb. d. Ver. f. vaterl.
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18883—Nematoden, Trematoden und Acanthocephalen, gesammelt von Prof.
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1888—Report on the Entozoa collected by H. M. 8S. Challenger during
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1889—Compendium der Helminthologie. Nachtrag. Die Litteratur der
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19034
426 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
von Linstow, O.—Continued.
1907—Nematoden aus dem Ko6niglichen Zoologischen Museum in Berlin.
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NEMATODE PARASITES OF BIRDS 427
Sitzungsb. d. k. Akad. d.
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Contribuzione allo studio dei vermi parassiti in
di storia nat. di Genova
428 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
PrRRoNcITO, Epoarpo; and ToMIOLO, ARISTIDE.
1901—Sulla elmintiasi mortale dei fagiani. Studi ed osservazioni. Ann.
r. Acecad. d’agrie. di Torino (1900), vol. 48, mem., pp. 63-72.
Prana, G. P.
1879a—Osservazioni sul Dispharagus nasutus Rud., dei polli e sulle larve
nematoelmintiche deel mosche e dei porcellioni. Atti Soc. ital.
di se. nat. (ete.), Milano, vol. 36, pp. 2389-262, figs. 1—21.
1879b—Idem. Mod. zooiatro, Torino, vol. 8, pp. 152-154, fig. 1; pp. 170-174,
figs. 2-14; pp. 189-198, figs. 15-21.
PoLonio, ANTONIO F.
1860—Novae helminthum species. Lotos, Prag, vol. 10, pp. 21-23.
RAILuiet, A.
1893—Traité de zoologie médicale et agricole. éd. 2. [fase. 1]. 736 pp.,
494 figs., Paris.
1895—Idem. [fasec. 2]. xv-+737-1303 pp., figs. 495-892. Paris.
1915—L’emploi des médicaments dans le traitement des maladies causées
par des nématodes. Rep. 10. Internat. Vet. Cong., London (Aug.,
1914), vol. 3, pp. 733-749.
1916a—La famille des Thelaziidae. J. Parasitol., Urbana, Ill, vol. 2, pp.
99-105.
191G6b—Nématodes parasites des rongeurs. Rec. de méd. vét., Paris, vol.
92, pp. 517-521.
RaILuier, A.; and Henry, A.
1909—Sur la classification des Strongylidae: 1. Metastrongylinae. 2.
Ankylostominae. Compt. rend. Soe. de biol., Paris, vol. 66, pp.
85-88; pp. 168-171.
1911a—Les helminthes du nandou. Bull. Soe. nat. d’acclimat. de France,
Paris, vol. 58, pp. 5388-541; pp. 573-582, figs. 1-6.
1911b—Helminthes du pore recueillis par M. Bauche en Annam. Bull. Soc.
de path. exot., Paris, vol. 4, pp. 693-699.
1912—Quelques nématodes parasites des reptiles. Bull. Soc. de path. exot.,
Paris, vol. 5, pp. 251-259, figs. 1-3.
1913—Un Haemostrongylus des bronches du léopard. Bull. Soc. de path.
exot., Paris, vol. 6, pp. 451-454, 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 nématodes du genre Goezia Zeder. Bull. Soc. de path. exot.,
Paris, vol. 8, pp. 270-275.
RaILuietT, A.; Henry, A.; and Sizorr, P.
1912—Sur les affinités des dispharages (Acuaria Bremser), nématodes
parasites des oiseaux. Compt. rend. Soe. de biol., Paris., vol. 73,
pp. 622-624.
Ransom, Brayton H.
1904a—Manson’s eye worm of chickens (Ovyspirura 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, pl. 1, 3 figs.
1904b—A new nematode (Gongylonema ingluvicola) parasitic in the crop
of chickens. Cire. 64, Bureau Animal Indust., U. S. Dept. Agri.,
Washington, 3 pp., figs. 1-2.
REIPISCH, JOHANNES.
1893—Trichosomum strumosum n. sp., ein Parasit aus dem Epithel des
Oesophagus von Phasianus ¢olchicus. Arch. f. Naturg., Berl., vol.
59, pp. 331-340, pl. 18, 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. .
RUbOLPHI, CARL A.
1803—Neue Beobachtungen tiber die HKingeweidewiirmer. Arch. f. Zool. u.
Zoot., Braunschweig, vol. 3, pp. 1-82.
1809—Entozoorum sive vermium intestinalium historia naturalis, vol. 2,
457 pp, pls. 7-12. Amstelaedami.
1814—Erster Nachtrag zu meiner Naturgeschichte der Hingeweidewtirmer.
Mag. f. d. n. Entdeck. in d. ges. Naturk., Berlin, vol. 6, pp. 83-113.
1819—Entozoorum synopsis cui accedung mantissa duplex et indices locu-
pletissimi. x -+ 811 pp., 3 pls. 8°. Berolini.
Rwust.
1908—Entenerkrankung durch Tropidocerca jissispina. [Abstract]. Ver-
Offentl. a. d. J. Vet. Ber. d. beamt. Tieriirzte Preuss., Berlin (1905),
vol. 6, p. 30.
SCHLOTTHAUBER.
1860—Beitriige zur Helminthologie. Amtl. Ber. ti. d. 31. Versamml. deutsch.
Naturf. u. Aerzte, Géttingen (1854), pp. 121-133.
SCHNEIDER, ANTON.
1866—Monographie der Nematoden. viii + 357 pp., 122 figs., 28 pls., 343
figs. Berlin.
SCHRANK, F.
1788—Verzeichniss der bisher hiniinglich bekannten Hingeweidewtirmer,
nebst einer Abhandlung itiber ihre Anverwandtschaften. 5 p. 1, 116
pp., 1 table. Mutinchen.
1790—Fortekning, pa nagra hittils obeskrifne intestinal-krak. K. Vetensk.
Acad. n. Handl., Stockholm [ners. ser.], vol. ‘141, pp. 118-126.
ScHWARTZ, BENJAMIN.
1925—A scaridia lineata, a parasite of chickens in the United States. Jour.
Agri. Research, vol. 30, pp. 763-772, figs. 1-18.
SruratT, L. G.
1913a—Sur un Dispharage de la chevéche et les affinités du genre Acuaria
Bremser. Compt. rend. Soe. de biol., Paris, vol. 74, pp. 103-106,
figs. 1-6.
19130—Sur le spiroptére des pies-griéches. Bull. Soc. d’hist. nat. de
V’Afrique du nord, Alger, vol 5, pp. 223-225, figs. 1-2.
1913c—Observations sur le Tropidocerea inermis Linst. Bull Soe. d’hist.
nat. de l’Afrique du nord, Alger, vol. 5, pp. 191-199, figs. 1-11.
1914a—Sur l’Habronema (Spiroptera) leptoptera (Rud.). Compt. rend.
Soe. de biol., Paris, vol. 76, pp. 21-24, figs. 1—5.
1914b—Sul la morphologie de l'appareil génital femelle des Spiruridae.
Compt. rend. Acad. d. sci., 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.
1914d—Sur un nouvel habitat et sur la morphologie du Subulura allodapa
(Creplin). Compt. rend. Soc. de biol., Paris, vol. 77, pp. 154-157,
figs. 1-4.
1914e—Sur quelques Hétérakis d’oiseaux. Bull. Soc. d’hist. nat. de
VAfrique du nord, Alger, vol. 6, pp. 195-202, figs. 1-65.
1914f—Sur un nouveau parasite du percnoptére. Bull. Soc. d’hist. nat. de
V’Afrique du nord, Alger, vol. 6, pp. 149-153, figs. 1-6.
1914g—Sur un nouveau parasite de loutarde houbara. Bull. Soe. @hist.
nat. de l'Afrique du nord, Alger, vol. 6, pp. 117-119, fig. 1.
430
BULLETIN 140, UNITED STATES NATIONAL MUSEUM
Srurat, L. G.—Continued.
1914h—Sur deux nouveaux Hétérakis du Sud-algérien. Bull. Soe. d’hist.
nat. de ’Afrique du nord, Alger, vol. 6, pp. 222-225, figs. 1-4.
1914i—Sur les physaloptéres des rapaces. Bull. Soe. d’hist. nat. de
Afrique du nord, Alger, vol. 6, pp. 244-253, figs. 1-3.
1914j—Sur la morphologie de l’ovéjecteur des Tropidocereca. Compt. rend.
Soe. de biol., Paris, vol. 76, pp. 173-176, figs. 1-3.
1914/—-Sur un nouveau spiroptere des rapaces. Compt. rend. Soc. de biol.,
Paris, vol. 76, pp. 427-429, figs. 1-3.
19147/—Sur un Tropidocerca parasite d’un echassier. Compt. rend. Soc. de
biol., Paris, vol. 76, pp. 778-781, figs. 1-8.
1914m—Sur un nématode parasite du flammant rose. Compt. rend. Soc. de
biol., Paris, vol. 76, pp. 814-817, figs. 1-4.
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. Compt. rend. Soc. de biol.,
Paris, vol. 78, pp. 279-282, figs. 1-4.
19150—Expédition de MM. Walter Rothschild, EH. Hartert et C. Hilgert dans
le Sud Algérien (mars—mai 1914). Nématodes parasites. Novi-
tates zool., vol. 22, pp. 1-25, figs. A—C, 1—27.
1915c—Un nouveau physaloptére des rapaces. Bull. Soe. d’hist. nat. de
lVAfrique du nord, Alger, vol. 6, pp. 157-159.
1916a—Sur un nouveau dispharage des palmipédes. Compt. rend. Soc. de
biol., Paris, vol. 79, pp. 785-788, figs. 1-5.
1916b—Contribution a l’étude des formes larvaires des nématodes parasites
hetéroxénes. Bull. sci. de la France et de la Belg., Paris and
London, ser. 7, vol. 49, pp. 297-377, figs. 1-14.
1916ce—Sur un nouveau type de Spiruridae. Compt. rend. Soc. de biol.,
Paris, vol. 79, pp. 517-519, figs. 1-38.
1916d—Sur un nouvel Habronema du Bubulcus lucidus Raf. Compt. rend.
Soe. de biol., Paris, vol. 79, pp. 295-297, figs. 1-3.
1916e—Sur la quatriéme mue d’un dispharage du flammant. Compt. rend.
Soe. de biol., Paris, vol. 79, pp. 439-441, figs. 1-4.
1916f/—Dispharages d’Algérie. Compt. rend. Soc. de biol., Paris, vol. 79,
pp. 984-9388, figs. 1-4.
1917—Nématodes de la perdrix de roche. Bull. Soe. dhist. nat. de VAf-
rique du nord, Alger, vol. 8, pp. 208-215, figs. 1-4.
1918a—Nématodes du Caccabis petrosa Gmel. (2e note). Bull. Soc. d’hist.
nat. de Afrique du nord, Alger, vol. 9, pp. 50-60, fig. 1.
1918b—Contribution a étude de la faune parasitaire de la Tunisie. Né-
matodes. Arch. de l’Inst. Pasteur de Tunis, vol. 10, pp. 248-275,
figs. 1-4.
1918c—Sur le dispharage de l’echasse. Bull. Soe. d@hist. nat. de Afrique
du nord, Alger, vol. 9, pp. 106-109, figs. 1-2.
1918d—Sur un nouveau strongyle (Trichostrongylide) de l’échasse. Bull.
du mus. d’hist. nat., no. 2, pp. 113-115, figs. 1-4.
1918ce—Sur les strongles du gésier des palmipédes. Bull. du mus. d’hist.
nat. no. 5, pp. 345-351.
1919a—Dispharages (Nématodes) de l'Afrique mineure. Novitat. zool.,
London & Aylesbury, vol. 26, pp. 179-189, figs. A—D.
1919b—Contributions nouvelles a l’étude des formes larvaires des néma-
todes parasites hétéroxénes. Bull. biol. de la France et de la
Belg., Paris and London (1918), vol. 52, pp. 344-878, figs. I-x1I.
NEMATODE PARASITES OF BIRDS 431
Seurat, L. G.—Continued.
1920—Sur une filaire du héron cendré. Bull. Soc. d’hist. nat. de l’Afrique
du nord, Alger, vol. 11, p. 142.
SHIPLEY, A. E.
1911—“Grouse disease’’—continued—strongylosis. Part 1. The thread-
worms (Nematoda). (Jn The grouse in health and in disease.
London. vol. 1, pp. 207-218, pl. 33, figs. 1-5).
VON SIEBOLD, C.
1836—Helminthologische Beitriige. Zweiter Beitrag Syngamus trachealis.
Hin doppelleibiger Eingeweidewurm. Arch. f. Naturg., Berlin, vol.
2, pp. 105-116, pl. 3, figs. 1-2.
1837—(Zusatz to Nathusius, H.: Helminthologische Beitrage, pp. 52-65.)
Arch. f. Naturg., Berl., vol. 3, pp. 66-68.
SKRJABIN, K. I.
1915a—Syngamus’y turkestanskikl ptits (Syngamus in birds of Turke-
stan). (Russian text.) Vestnik Obsh. vet., Petrograd, vol. 27,
pp. 645-658, figs. 1-5.
1915b—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.
1916b—Nématodes des oiseaux du Turkestan russe. Ann. Mus. zool.
Acad. imp. d. sci. de Petrograd (1915), vol. 20, mém., pp. 457-—
557, figs. 1-58, pl. 8, figs. 40, 41, 54, 59.
1916c—Contribution 4 l'étude de la faune helminthologique du Paraguay.
J. russe de zool., Petrograd, vol. 1, pt. 4, pp. 736-757, pls. 24-26,
figs. 1-27.
1916d—Seuratia n. g., nouveau genre de nématodes d’oiseaux. Compt.
rend. Soc. de biol., Paris, vol. 79, pp. 971-973.
1917—Sur quelques nématodes des oiseaux de la Russie. Parasitology.
Cambridge [Eng.], vol. 9, pp. 460-481, fig. 1, pls. 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 TVhelazia Bose
1819. [Matériaux pour servir 4 une monographie des Nématodes
d’oiseaux. II. Le genre Thelazia Bose 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. 144,
pls. 1-2, figs. 1-5.
1923—Nematody domaschnich ptiz. Opyt monografitscheskoi rasrabotki.
1. Kruglyje tscherwi kurizy, indjeki, zjesarki, pawlina i golubja.
[Abstract] Berl. tieriirztl. Wehnschr., vol. 39, pp. 297-299. From:
Iswestija Donskogo Wet. Inst., vol. 1 and 2, 1920.
1924—K faune paraziticheskikh chervei turkestanskikh pelikanoy (Pele-
canus onocrotalus L.) Trudy Gossud, Inst. Eksper. Vet., vol. 2,
pp. 149-154.
3612—2T——-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. 288-294, pls. 2-10.
Sonoviev, P. F.
1912—-Paraziticheskie chervi ptits Turkestana. [Parasitic worms of the
birds of Turkestan] [Russian text.] Ann. Mus. zool. Acad. imp. d.
sci. de St.-Pétersb., vol. 17, pp. 86-115, figs. 1-15.
Sonsino, P.
[1890]—Un nuovo Heterakis del Gallus domesticus, Heterakis differens
mihi. Atti Soe. tose. di se. nat., Pisa, proc. verb., vol. 7, pp.
136-137.
STAFSETH, H. J. and KorTnan, A.
1925—Report of Investigations on an alleged Hpizootic 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.) Cire. 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. scututus
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.
StimsEs, C. W.; and HAssAtt, 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.
STOSSICH, MICHELE.
1887—Il genere Heterakis Dujardin. Glasnik hry. nar. druztva, Zagreb,
vol. 2, pp. 277-801, pls. 3-9, figs. 145.
1889—Il genere Physaloptera Rudolphi. Lavoro monografico. Boll. Soc.
adriat. di sci. nat. in Tieste, vol. 11, pp. 36-59, pls. 1-8, figs. 1-42.
1890—Elminti della Croazia. Glasnik hry. nar. druztva, Zagreb, vol. 5,
pp. 129-136, pls. 4-5, figs. 1-15.
1891—I1 genere Dispharagus Dujardin. Lavoro monografico. Boll. Soc.
adviat. di sci. nat. in Trieste, vol. 13, pp. 81-108, 3 pls., figs. 1-27.
1892—Osservazioni elmintologiche. Glasnik hry. nar. drustva, Zagreb,
vol. 7, pp. 64-73, pls. 1-2, figs. 1-10.
1898—Filarie e spiroptere. Lavoro monografico. Boll. Soe. adriat. di
sci. nat. in Trieste, vol. 18, pp. 18-162.
(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—Contributo allo studio degli elminti. Boll. Soc. adriat. di sci. nat.
in Trieste, vol. 20, pp. 1+8-+1, pls. 1-2, figs. 1-18.
1902a—Sopra alcuni nematodi della collezione elmintologica del Prof. Dott.
Corrado Parona. Boll. mus. di zool. [ete], Genova (116), 16
pp., pls. 3-5, figs. 1-38.
1902b—Idem. Atti Soc. Ligust. di sci. nat. e geogr., Genova, vol. 13 guigno,
pp. 61-76, pls. 3-5, figs. 1-38.
1904—Sopra aleuni nematodi. Ann. Mus. zool. d. r. Univ. di Napoli, n. s.,
vol. 1, pp. 14, pl. 1, figs. 1-11.
SWEET, GEORGINA.
1910—Some new and unrecorded endoparasites from Australian chickens.
Proce. Roy. Soc. Victoria, Melbourne, new ser., vol. 23, pp. 242-
256, pls. 31-35, figs. 1-21.
THEILER, 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 pl., fig. 1.
THEILER, A.; and ROBERTSON, W.
1915—Investigations into the life history of the wireworm in ostrichs.
38 and 4. Rep. Dir. Vet. Res. Dept. U. 8S. Africa, Pretoria, pp.
291-345, 9 pls.
TRAVASSOS, LAURO.
1918—Sobre as especies brazileiras da subfamilia Heterakinae Railliet &
Henry. Mem. Inst. Oswaldo Cruz, Rio de Janeiro—Manguinhos,
vol. 5, pp. 271-818, pls. 27-31, figs. 1-38.
1914a—Contribuicao para o conhecimento da fauna helmintolojica brazi-
leira. 3. Novo genero da familia Heterakidae Railliet & Henry,
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. 1387—
142, pl. 15, figs. 1-3.
1914b—Tricostrongylinae brazileiras. (Nota prévia). Brazil-med., Rio de
Janeiro, vol. 28, p. 163.
1914c—Tricostrongylideos brazileiros. (3. Nota previa). Brazil-med., Rio
de Janeiro, vol. 28, pp. 325-327.
1914d—Contribuicoes para o conhecimento da fauna helmitolojica brazi-
leira. 3. Sobre as especies brazileiras do genero Tetrameres
Creplin, 1846. Mem. Inst. Oswaldo Cruz, Rio de Janeiro—
Manguinhos, vol. 6, pp. 150-162, pls. 16-238, 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, pls. 23-26, figs. 1-14.
1915b—Sobre as especies brazileiras de genero “J'etrameres Cliplin,”
[sic.] 1846. (Nota prévia). Brazil-med., Rio de Janeiro, vol.
29, pp. 297-298.
1917a—Tetrameridae brazileiras. (2. Nota prévia). Brazil-med., Rio de
Janeiro, vol. 31, pp. 65-66.
1917b—Contribuicao para o conhecimento da fauna helminthologica Sul—
Fluminense. Brazil-med., Rio de Janeiro, vol. 31, p. 149.
1918a—Contribuicao para o conhecimento da fauna helminthologica bra-
zileira. 7. Especies brazileiras do genero Thelazia Bose, 1819.
16 pp., 3 pls., figs. 1-13. 8. Sao Paulo.
434 BULLETIN 140, UNITED STATES NATIONAL MUSEUM
TrRavAssos, LAuro—Continued.
1918b—Observacoes sobre os Heterakidae. Rey. Soc. brazil. de sci., Rio
de Janeiro, No. 2, pp. 93-97, 2 pls., figs. 1-2.
1919a—Gastro helmintose das aves domesticas. Rev. de vet. e zootech.,
Rio de Janeiro, vol. 9, pp. 79-89, figs. 1-9.
1919b—Contribuicoes para o conhecimento da fauna helmintolojica bra-
zileira. 8. Sobre as especies brazileiras do genero Tetrameres
Creplin, 1846. Mem. Inst. Oswaldo Cruz, Rio de Janeiro—Man-
guinhos, vol. 11, pp. 71-79, pls. 25-28, figs. 1-14; French transl.,
pp. 63-71.
1920a—Trichostrongylidae brazileiros. Rev. Soc. brazil. de sci, Rio de
Janeiro, pp. 191-205.
1920b—Esboco de uma chave geral dos nematodes parasitos. Rev. de vet.
et zootech., Rio de Janeiro, vol. 10, pp. 59-70 bis, 1 chart.
1920c—Contribuicao a sistematica dos Physalopterinae. Rev. de sci., Rio
de Janeiro, vol. 4, pp. 57-58.
[1921]—Contribuicoes para o conhecimento da fauna helmintolojica bra-
sileira. XIII. Ensaio monografico da familia Trichostrongy-
lidae Leiper, 1909. Mem. Inst. Oswaldo Cruz, Rio de Janeiro—
Manguinhos, vol. 13, pp. 1-135, pls. 1—56, figs. 1-250.
1923.—Infromacoes sobre a fauna helminthologica de Matto Grosso. Folha
med., vol. 4, p. 38.
VEVERS, G. M.
1923—-Some new and little known helminths from British Guiana. J. Hel-
minth., London, vol. 1, pp. 35-45, figs. 1-3.
Warp, H. B.
1917—On the structure and classification of North American parasitic
worms. J. Parasitol., Urbana, Ill, vol. 4, pp. 1-12, 1 pl., figs.
1-14.
WALTER, H.
1866—Helminthologische Studien. 7 Ber. ... d. Offenbacher Ver. f.
Naturk., (1865-66), pp. 51-79, pl. 1, figs. 1-6.
Watton, A. C.
1923—Some new and little known nematodes. J. Parasitol., Urbana, II1.,
vol. 10, pp. 59-70, pls. 6-7, figs. 1-25.
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BIRD PARASITES OF NEMATODES 435
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INDEX
(Nematode synonyms and principal page references are printed in italics]
Page
abbreviata, Physaloptera.____....-----__-- 296, 307
PAT EULOL Caen en eee erie enn ee oe 295
abyssinicus, Coracias__..........------------ 127
acanthocephalica, Spiroptera_________--------- 287
acanthocephalicus, Hystrichis_-_.______-___- XVI,
366, 376, 877-878
Schistorophus---__- XIV, 285, 287
EA PONINODNOTUS! 228 a a en 334
Rorriduss. =. 5---.------ 2 343
VON AS ee ee ae ee 343
Meciplver WICOlOL a. -4==< seoasssce~s aon 2 138
TUES UIS aes Seer a Re EA a et A cs 138,
176, 276, 277, 288, 298, 299, 301, 302
LTRS 2 eee oe Rs Ah ee 297
accipitri, Gnathostoma_ _____________- XVI, 864-365
Acrocephalus arundinaceus_____________----- 139
IACLILIS NY POlCUCS «22062 ooo cone a we 235
WNACWIATIO = oe ee ee 235
Ja C0 CN (CASS ne cel an oe 210,
211, 212, 213, 214, 215, 216-217, 218, 219, 220, 221
222, 223, 224, 225, 226, 237, 244, 255, 258, 262, 272
OPIN oe ae eee ne 273
LOU a eee 281
ancnurise=--=--..--o-= XII, 212, 217, 218-220
NCE er ee as 248
ALUCHU ALE saa ee ee ee 217, 220
COLCOTOM ne os ee eee ee 249
CDI ee een 240
CONTE eee ara see an oe eau eee ances 249
CORGStA= aso 225) J) -~ 3238 XM, 218, 220-221
CLOSSISEIINIG te ee eee ce 243
Gepresstesss= ow ee a XU, 217, 220-221
CUIDUC Doe ae eee Meee 2 277
CLOT Er renee ee ee 259
GICINCU a= kn ee ane en ee ees 249
PEACIIS saat ees XI, 217, 218, 222-223
OEUT ES eee ea neta 227
FCT RED acne te re ya ed 278
PULTE LOS a ean a tata 226
MOERCE DG oe ieee es 273, 276
LOTIGCOT TOI nn en 255
longevaginnta..- 3 cece es 256
MACLOMIMA: 2 ee caccse 212, 213
MTTSTRU YAS Se eo een ee oe 212, 213
MpPSCICANHO =. a eee 212, 214
NOCLILOE = we eee ee nee ab ot 242
OLnata ae eee rece XII, 217, 223-224
papilufera.2---2.2----. XM, 217, 218-224, 394
DENGAN naan nen cee cate oe we 263
ptilopachydis_-:_._-.-___- XM, 212, 214-216
Gusdrilobae.-_------ >= = XM, 212, 215-216
POOL rn ae ee eae 283
FECLD ea 2 aren accgetenacada 280
TECLOOUGUTUI = ne oe ne ee 243
WOLUNGOIN. Dee ee sae amen ao 229
Page
A CUATIN StNiis_ = 2 Se 222
SDC Seo ee ee 238
SQUGMOQUG oe eee ee es 257
SU DU ate seeee eee. eaeeeee XII, 218, 224-226
SYOMOMC seca a oa ee ee 282
tarentolae__..._._..__________- XU, 212, 216
LORS ee 217, 218, 225-226
triaenucha__.._................-._--. 271
wpupae-epopis__..._____.___.-.-_---- 190
OTE ene ee os oe 260
Acuaria (Cheilospirura) gruveli_____..-.------ 227
magnilabiata________- 241
pavonis___...._____-- 389
Skrjabint=s ee 389
SPECS ss. ae eee 229
Acuaria (Dispharynz) laplantei_..._._______-- 241
MOSUtG: | eee 2 sens 237
MOCULGE. - 22a ee 242
Acuaria (Echinurta) hargilae____.._.____-__-- 253
leptoptilt__. .. 2 =e 253
Acuaria (Hamannia) phoenicopteri___________ 257
UNnCNOG =.= ees 246
Acuaria (Synhimantus) invaginata_________.-- 279
laticensa2- =~ sas 276
SUDTECH = eee gee 282
Acuariidae.--_-...-.... 163, 210, 211, 284, 290, 311, 392
A CUSTUNEO Sooo. han ee 211, 212, 216, 226, 231,
233, 237, 244, 258, 261, 262, 264, 272, 290, 391, 392
ICU LCOLO UA Ge 2 ee ee ee 254
acileata, Spiroptera...--...2.-22.-.-22- ene 253
aculeatus, Dispharagus__.......-------------- 254
UCT a ATA S Soe ore re ec ee re 27, 81
ANISOD os oe con oa ok eee oe 20
acuticauda, Heterakis_............---- vil, 52, 76,77
ODYS80MGs 2 ee ewe 133
Physaloptera_---- XIV, 296, 297-298, 305
Subulurs.-oocs]sce.-- IX, 106, 183-134
acuticaudatus, Strongylus___.-.....----------- 77
acutissima, Heterakis_......-.---.------------ 107
Subulura: <2. 52... vin, 106, 107-108, 109
acutum, Amidostomum__---_-------.------ 20, 24-25
Actas, SHY ONGUINWS 2 nan nese caseen eee tees 24
adspersus, He rancolinus= 2. ..-sasc<2----.---= 123, 307
QAUNCE I SDITODUCE.. cnacnacweeraaseassesSec> 234
aduncus, Cosmocephalus_--_..--.-.---- XU, 234-235
DiSPNOTOGUus. 5. sacecceanan-sccen ss 234
Aegolius brachyotus!-- 2-222... --=--.0-2-- 144
OUUS. = wave sece ancl So weenen newness 144, 276
aegyptiaca, Ascaridia-_-......-...-------- vu, 78, 83
HeleraKkisn oo a cncactase= 83
aegyptius saharae, Caprimulgus-_--.------ 118, 129
aerophila, Desmidocerca--_-.-.---.---- xu, 209, 210, 264
geruginosus, Circus.........------- 137, 138, 176, 298
BESAONs WAlGO Lo saceeweseaese<anosceesaaeeen 138
aestivus, Pionus (Psittacus)......-...-------- 80
438 INDEX
Page Page
aetherous, Nyctibius. 22.2228 eo 129,130 | americana, Tetrameres__--...---.-- XV, 334, 37-388
aothiopica; This 22 ee ee 297,309 | americanum, Cyathostoma----_---.--- VI, 42, 45-47
ater, Hurystomus: 2 oc see cee cee a ee 1245127] ‘americanus, Corvus. 2220222522 ae 220, 355
Oiiniss Actaria 2. 22 ee ee 273 Pelecanus: 2-22 s2222 eee 148
Spiropleras ies See ae ae 309 | amherstiae, Thaumalea_--_------------------ 64
Synhimantus 2:22 2222202222328 NIV, £79-274. |) Amidostomidae=. = 2 .--- a= se See agile’
africanus, Eustrongylides_-_-_------ XV, 367, $69"570" | Amidostominae. .----.--- 2-2-2 aS ecco se 18, 19, 26
afroides,OUIs oes ok rcs 2s ea ee 192,198 | Amidostomum.- ---- 17, 18, 19, 20, 22, 23, 24, 25, 26, 383
Aga VATGeR intense ees ne haere ae nee eons 142 BCU os see eee 20, 24-25
Agamonema cysticum .....:.-.=---=+---<-<-s< 371 alseris/2so22 25. eeee Vv, 19, 20-21, 29
Aaya alalece we oes aoe one eee eee 241 chevreuxi___-_- V, 19, 20, 22-24, 25, 383
EEN LEY Fret NG Fs Wap NS aces Ct a 241 Tiliches eee ose l eee eee 20, 25
PIR AiO a ssn ele ce nein oem era 241 Henry eee eee V, 20, 22
LOL, CA CHON Swen a ee ee ne ees eae 281 MmONnOGOM= a- on eee V, 20, 25, 26
LAVA oe eee a ee care eho 281 NOGULOSUIN. oa eee 20
Feterakise esse oe eons oe ceee ease VI, 52, 56, 56 faillieti 2.2. 2222-5 V, 19, 20, 21, 24, 383
Physalopterass=--.0-- se recesses XIV, skriqbint. io. oo ee eoeeee 22, 3838
296, 297, £98-$00, 301, 305, 394 | anacanthura, Oxyspirura_..-....--- XV, 322, $23-824
SPTOpEGa-+ cco scceceeseees 165, 240, 281, 282 Spiropter a. eee eee 323
STEALS nn ns eee ree ree 2814 anals, Dispnaragus: os. one asco eon eee e sees 281
alata chevreuxi, Physaloptera_-__--..--. 206;:$01,:394 1) (ANAS ACUtR= = Janos ones es a thee eee 27, 81
alata nouveli, Physaloptera-_-_.-.- 296, $01-302, 394 Blbrone ees trae eee ets 28
Glatus. \COSMOcePNQe =o. oo oes See wee 235 anser domesticus.) neck ance cas 28, 62
PSD GROG US set ue on ae 281 QnSer leran ete oS ke ee as 28
FRIRU GS SEVEDSIS: sooo. Me oo le Ue 302 boschas. .... 10, 34, 62, 136, 139, 152, 245, 267, 343
BDOCIOS cron ee OLE Ue UY eT 139 boschas domestica.....--.--. 10, 27, 42, 53, 62,
alaudarius, "Tinntineulus 22.2222 Sos Sees 299 81, 93, 136, 152, 246, 343, 345, 367, 372, 373, 376
alba; Ciconias coe eee 34, 44, 152, 168, 296, 307 bosehias férg 32 tec. bees 81, 343, 376
WP ot acta eee Sayaka ei econ eae 139 Canadensis: oo cose eco 62
BS rere ye Rh cca eta eee rae aI 276 clangula. toe oor ee eee eee 28, 147, 267
HY Bie po ee ee a Oy ld Wa a eR A ee 273 Cee eee ee eee ee ear 24, 28
a@lbajasiatica, Ciconiao: 22.2 ie oer eee oe oe 168 fuligulac. cc oo cet ecees cccnccssesemsece= 24, 28
Sipellas: Werpiisns = sen Ses ot ah ee 372 fusca soso cescceacesae= 24, 28, 267
SIDicii a PACUINAS oreo een ae eee 138 glacialig: 2 0s see see ese wenas 267, 372
Waleed eee cee see ce a a a eee 138 1OUCODS eas bccscoscs sowteeseeepenenees 28, 62
Piglideetiis snes ee ee eee 138 miollissima@. = 2222 oe eee 24, 28, 372
EISIQERISUS Tee. = ee ee ee ee eee 137 MOSCHAtG..-- <n 2 slo caneoe seeeee eee 62, 81, 136
albicollis, Nyctidromus.__.................-- 130 Nigras econ else eee see eee eee 24, 27, 28
aibiirons,;Anas© 2-28 2ces san we ere ee 28 penelope sone cece ees 27, 28, 246
JANSCP=\- eo eac ose 10, 20, 22, 27, 383, 384, 385 querquedulat i 2: 23-6 so cee coe ee 20
mibomarpinatatSerix ce tes eee ee eee 205 TUDTIPES see eee eee 246, 248
WMICAILOLOR essa ceccnsae eee ccaccecs 147, 235, 267, 372 Segetum) = cnc sc oso e ee aces ae enee noes 28
mAucedO AMer Cana cco eee sce e cones coeeeee 293 SPONSG! cas cescacet soe esee ee eae 59
ISDIG A= soc ee ee aed Seema 269 tadorna.c2 = ono aan cc oe se eee 53, 62, 267, 376
LEI G {1 Se ROSES IS ot ORB EEG RR I 104 | anatinum, Epomidiosttomum..--_.------------- 27
MUIDGUD Ose conn nee ee ae 108; 1261 Anavlostoma-- 2.2. -.sa-cee= eee nae one 29
DIAL =o ete ee ne re eee 1121| -Ancyracanthidae-_.-o-=------ceeeese 163, $61, 392
leprincei seo ec owe date Senta cas 118}'| Ancyracanthinae.-.--.--22-..----<se-ss—sece 392
MUI DODO 22a eee 338 | Ancyracanthopsis..-.-------------- $61, 362, 363, 392
TOCITUEE a aoc ae mn es ee 119 bihamata...=-—<-.. XVI, 362-868
SUCLOTID Soe ce nro teen a ec 112, 130 bilabiata._...... 361, 862, 363, 392
Cy DiC ae ee rade ieee ee ees 108 | Ancyracanthus_.........----..- 284, 319, 361, 363, 392
Alodape’ "Aloda pac See = eee 108, 126 Uidene. oe ae 288
FICC ORE =n coe oo ose c anaes 108 bikamata.: oon ee eee 362
OLY ES ee Ces Cah oe ee 108 bilabiatue ee ene 362, 363
Subulura-_._-_.-- vin, 105, 108-110, 126, 394 longicornis.._ =... Men sa-e 285
Biping. Lringae. oC osee seek een en ume 254 ophtialmicus. 2.022 - paste 320
SIpinus, SP VITOCOrax. ac ceo cs ccc esses 35, 220 pinnatinidus- =) 2. - 72222 cee 392
BIUCON PiTikin eons toon ace ee ate a 138, 144 | sndersoni, Contracaecum--.--.--.--- X, 147, 149-150
BYCMAsS ccceesceete cee ee eee eke es 144})| ‘anglorum, Procellarias: 2 -- 2-7 oo oe 263
MO ae hee ca eae. 144))| (Anguilla vipers: co 2cce eo cates ee ee - 146
Amazona ochrocephala. ....-.--------------- 80; | .anguttlae, Ascaris. 80.2... - 2-2-2 oe one 146
GMDGUUS, (SU ONPYIUS oe ene 287 Porrocaect0m 2.2) 225252 IX, 136, 146
amblymoria, Ascaridia._.-.----..-.-- vu, 78, 88-84 | angusticolle, Porrocaecum-.-.----..-- IX, 136, 187, 188
amblymoria, Heterakis............----------- 83 | angusticoltis, Ascaris... --..-.2----------- 137
americana, Alced0.....25..2.--c--cssncacoecs 293)/|" Anhinga anhinga.: - ooo 0 so senck cco coece seen 371
MURR eo ccs Ce eee ee 32, 69, 97, 165 Tufa_..----------------------------- 369
INDEX 439
Page Page
anhings, Anhinga. <.<.cc.~ occu ee. 371‘ | Aramides.cajanea........-.--..-2.-23.i22 S25 350
lotus si cower cece SN. 148 | ararauna, Pionus (Psittacus)_.....-.-.--_..- 80
ant. Crotophaga...-<.=...-....:..sa42: 124, 207,323 | Arboricola torqueola_._....-...-.-.---------- 387
IMBisakINae <= ..32ilsecc0csccescu cs teye Re 135,146 | Archibuteo (Buteo) lagopus-__-...-.-------- 137, 138
IRBISAKIS ces hoc co kocenecccwce Sen ao I 135 | Archibuteo vulgaris.............-.-.--..-.-- 138
annulata,. Hartertia.........-....... Masti 798-1956 | arctica, Sterna..-.........~... 223.2920 be 235
anolabiata, Mileria.:...:...--.<<2<-.255F 22334 317 | arcticus, Colymbus............ 148, 265, 267, 271, 367
(ROzyspirura)—..<...2<...8 Se 317 | arcuate, Kathleena.......2...-.s 2202220. 152, 153
WpiTOplerd =. -35ss5 25 Sif Ardea agaml: -. - 25.5<5.5224--ecesn--teenae se 142
"hhelaziaen coc ssccocek ees XV, 312, 817-318 caerulea .2.52.0072 hot Re ce teas 142
anomalus, Heliotreptus.............-----.---- 130 Cinereds_ 22 3h fe L. 141, 142, 152, 209
ANSON ACUUAS ox cosas eSdocesssesessenc ThE 20 COCOL <b see Eh ae 141, 371
albifrons: s2:2.2.<s--< 10, 20, 22, 27, 383, 384, 385 Comatas ces Loe ee ee 152
ONSGF cn. asses sassenssseens 10, 20, 28, 34, 53, 62 garzettan! 222.225.2125 $25 es eee nee 83
anser domesticus:...s<<.2s-2es222$ 20, 384 Olav. 6 os ee eeee eee 369
CinereuS: 2225255522065. ae 10, 20, 62 PTUS Honoka cae ee noe eae 142
cinereus domesticus-.-_--_- 10, 11, 42, 53, 246, 373 herodias”. 22.2. tL 141, 152, 248, 372
Clavigula secs as sesccsaccccccss 2 20 Wia] Obssone ao sase cee sa ew es eee 142
CTOCCAS 2 sctcscc nu acessustscenle Jb lie 20 manillensis’. 5.2.2.4 --.-.o5 eee 141
domesticus:2-<.<--<=--<ncncesesseceu? 10, 101 melanocephala......-..-/=422225 =. 142
fabalis. 5. <12.2 assed. te tttiees 20, 62 MINOM ooo. are nt ee eS 152
HERS: § 286 es 6 oon Sc RUIN 10 Vigra foe ot taco chee Be 281
ferusidomesticus....... .--J82280 22208 10 NY CviCcOrax .. <= >- 55.2 - 255 142, 152, 347
foligula 2252.62 see Sea see ea 20 piléatea-25222- cee -2-- se 142
PUISCHREs SEU aos onsen scaes sels 20 purpurea... -22225-.2-2.82223225228.-2 142
IBHCODS! 2n2- so sso. -sotecetesascc ce 20 scapularis: -.. 2-2 s2..5-2.8204 2 142
marilas. i225. 52. Boe ee eens BEE 20 Species: =~. 2-2 secs wens eee os 152, 154, 160
MOMISSIM Ae as 3. oon Seat. Se 20 Stellarisz:25<2+.-..2-. .< 1422 ees 152, 274
migra: 02 226 oo oe 2 Rk RLY 20 Wi0ldC@8: 2. ~ 2... 5.2 Ss2 sees ses seesee 142
penelope...=.<.--...- ie eee 201! | a@rdeae; Actarig......2--222222022bt3s2255..- 248
SORCtUMN | 222-58 ee 20, 62 ASCOT iS oo2 = a5 5 pasos sss Se 141, 142
inser; ANSer:22225-s<2-~525225-5 10, 20, 28, 34, 53, 62 Dispharagus.....-..=..----+25s82.5--- 248
anser domesticus, Anas....-...222252/2..322 28, 62 HWehinurig-*) 2222-5 ec XIM, 244, 248-249
AnS6r.2=-2.- 8b setes Sa 20,384 | ardeae-stellaris, Strongylus_.....-.------------ 274
griserifera WANA. 22. s22 25 onc aoac2tesecesc 28 | ardea-maguari, Spiroptera_.....-------------- 255
AMISOTMONMOS |. 5220225250555 oS 965)268u), Ardoola grayil_... 2-2. s2<-2-2s25- bese 152
ATISOVISs 2 on oe oo ssa EE 268 {DiSo2 cesses seseseeeaes ese eee 279
Amidostomum..-_.....-------- Vv, 19, 20-21, 29 ralloides=2 2s... 5.5 25-22-55 eee 152
Ascaridia..2..2-2-2.5.5. 228823 vi; 79,101-102 |, Ardetta: minuta_—__.......=--=----_.2 ee 274
Strongylus.....--.-----------1----- 191 20oR Arduerna- 22.225. 2s2c5 2-22-2222 cat eles 4
Anthochaera carunculata_--........-....-.-- 9007 argentatus; uarus..2.--.-.----22--- 25 148, 234, 235
anthochaerae, Ceratospira_.__.....-.------.--. $241] -argentoides; Larus. =.-2--.--.--.=-..s2232. 235
Oxyspirura...-.-..... 203 KV, $22; See) :argas, Argusianus_.....=..-==--=22552--23--— 387
Anthropoides virgo........-22% 1542. -uuehs2 #49/-|\, Argusianus argus_.......-.=-----..#s2s-s¥2e2. 387
ARMUTIE Qo. 5.28 eo ca xcs cllesy 212, 216, 217,:219)\| \arquata, Ascaridia__-_._-.....--.-~-22+--ssen= 55
anthuris, Acuaria......-.-...-- XU, 212, 217, 218-220 Heterakcis:2- 2... ses vI, 51, 55, 56, 71, 72
Dispharagus......:.---.. 286 b2Li. 18) |e arduatrix, Columba....--2...2-.--.-.. 2 87
PRlaritisos owns s225.-tathres.3 218, 220, 330 Stictoenas ..._...------.-.322 092. 87
Spiroptera_..-.---.-- 217, 218, 219, 220,330 | arquatus, Numenius-__--.---.---------- 285, 372, 380
enticone; Grus* <-.2222--252222-s0 3222... 89,99 | arundinaceus, Acrocephalus --_----.--------- 139
Antrostomus vociferus...............------ =. 129: |. abwensis,, Alandsa..—-.....- =... Sv oss2Ssse see 302
apidster, Merops 3). 2 = os ..s65 eels 32 988in}) \-ASCATIORS 2c. o.-wae sewn ssp cneen deel 48, 49, 135
apivorus, Maleo-220 222222. .cce sence. 198208 | ;Aiscaridata_____...-.-=-------=---=--222 1, 5, 48, 392
Pornis)= 22 --..2.292elentiwvel 2 ish 197i | 4 Ag@cOrideG-< 222-2 25225scceesss5oboutsteeseoe 48
apricaria, Pluvialis. 2>...2.2-.sidsasie+i- 139,141 | Ascaridia----- 49, 50, 55, 77, 78, 80, 81, 83, 84, 85, 86, 87,
apus, Cypselusi:22.4.~25----.2.ss2eeers25-36 34 88, 89, 90, 91, 92, 93, 95, 96, 97, 98, 99, 100, 101
Menila’al bicills. 25. 22h 5222 =.=. ete 138 aegyptiaca=.......-.---- 22828 Vu, 78, 83
chrysasetos -2225262 222-452-220 12222 138, 301 amblymoria......------.2- Vu, 78, 83-84
fasciatass soe... o--setene eres 138 anseris:--22- 2) 5-2 -secena vu, 79, 101-102
imperialisi 2220 .-ossescu2e 138, 145, 298, 364 GrQUAG. on 2 ans a teres 55
MACH iatals ee os ee 316 australis... .<.<--+2=35% Vu, 78, 84, 85
NEGSVidse oo Sas o ne os aalen he s--s 138, 316 Borealis... 5. .-5.- uss vit, 79, 84, 86
pennata: <2 25552256 .-ccebonabelt- sah 138, 298 brasiliang-_.--.-.---233-03-5s222e- 58
rapak: belisaritis 6-0-4 ed aot 322) 304 brasiliensis.__.........---=== Vil, 79, 84-80
ANHUAT BONY PGOtOS 2255-5254 267 oscars atu ade 138 catheturina__......--------- vu, 78, 86-86
Reauillse VA RCaTis= 5-9-4. Sy sccttoeh sor, 150 circnlaris.-.---<2--nqn------8 Vu, 79, 86
aracanga, Pionus (Psittacus).........---.-.-- 80 columbae...-....----- Vil, 79, 86-88, 89, 96
3612—27 30
440 INDEX
Page Page
Ascearidia compar_...-......---..-#222 VU, 78, 86,88 | Ascaris perspicillum-.___--.--------..-------- 81
COMIPressa_..- Jase seeds ee NE 78, 88 praclonge. =o. 22 ooo 140
cordata =) = 2.20 2. ba eines vu, 79, 88-89 MteroMhOr@ =. 3. 5-5-2. 2 ee 97, 98
cristata. =... S2sree! siee vu, 79, 89-90 Meclinata so ts a ee eee 124
cylindrica... 2-2 VU, Vil, 78, 90, 91 reticulata = sce Sar ee reat ented 141
dolichocercan 222252 ene viul, 78, 90-91 WOSOTiUS 22 8 eee ee 158
fascintawes oc os debe vin, 80, 91-92 Sagitiate. ooo 281
francoling: 2:05 0200 uaa VIU, 78, 92, 94 semiteres— =. 2322 Goa ns De 141
POLE ee TE eke a vil, serpentulus—.....___.. _gualepsiee! 22 142
49, 54, 79, 80, 81-82, 83, 84, 85, 86, 87, 88, BPRCLOS onc oan kee 324
89, 90, 91, 92, 93, 94, 95, 96, 97, 98, 99, 100, spiculigertm. 2... 2 tee a 147
101, 102, 108, 110, 111, 112, 113, 114, 115, SPP Gliss eo ue eee 143
116, 117, 118, 119, 121, 122, 123, 124, 125, strongyling 208. _ 22 2 eae 77, 128
126, 128, 129, 131, 133, 134, 135. subulata 53... once La 128
eraninlosa eae ee Vin, 78, 92-93, 94 truncata - 222035 3 ee i ee 80
WRB eek EAs Pe EL aaa 93, 94 valdemucronata.= 2.202.) See 74
hermaphrodita_____.._... vl, 77, 79, 80, 81 wesicilarisic8:. Leelee pula 52
ACT Gee Nh a e ageaga 813 PAscaroi desc ieee ea et ae 48, 49, 135, 391
lineata sce. Sa hana VINT;79; 80382; 98-94.) ‘Aspidodera... 2 28 o2b sooo Sone ee al 102
longecirrata._.._. 2 si2see vill, 78, 94-95 | Astomum-___...---- aici Lo 2 ean bear ® 334
miagalhaesin: Sooke oe Se vil, 79, 95,96 | Astur melanoleucus-_--_.....-...-...-.._---- 309
magnipapillas 2) sass vin, 78, 95-96 MISUSE tte ea 176, 277, 278, 298, 355
mumidse sii 2 aa vit, 79, 96-97 palumbarius: <2. 2.2. : 236 138, 176, 288, 355
orthocerca 3 eno ee vil, 79, 97,98 | Asturinula monogrammica--_._________-_- 273, 282
perspicillum. 2... see 49°81, 87.c} tater, Maleoess o_o se os cane 138, 177, 243
plerophora=22 20 aes vill, 79, 97-98 Mil Wis 224252628 oe ears 138
RUT SLs Ree 2 Vi,79; 98-99...) | Athene: noctiias2 222002 28 ee ee 139
Strelnikowisn22.-220252--4- vin, 79, 99, 100 noctua glaux. 2 2522 22 ee i ee 203
SUOUTS =(- 2s ue re Ste ee Vil, 79, 99-100 | atra, Fulica_.._ 20, 24, 25, 28, 334, 343, 346, 376, 382, 383
sty phlocerca =. 2-2-2 vill, 78, 100-101 | atra assimilis, Buchanga___.._____________- 188, 222
trilabium = 222223 ak witt, 79; 101. .| jatratayChenopis==- 222 eee 53
Wuncata.- -.52 2255 ae. 80..| {atratusp@syemasie 22 Ao eee 52, 61
ASSGATIS? See cers. on Soe 49, 104, 135, 146, 272 | atricapilla, Muscicapa__._..-___--___.._-_-__- 214
anguillge-_. 2°... plstpy eh ne. etal 146 Pisorhing 2 =) 3 4oae sea 107
angusticojlis -.- - 22 see eee Aan 137 SULT eet hart eel ea ts ae 107, 205, 207, 325
aquillae. 2. 025. Sooke ia 150%| jatricapillus, Maleo.s..20.. 22-2 Le eee 297, 298
GrUeR es oes EE Ree Be Oa 1415 142).,) 7atrigularis,/Colymbus.2o 0. ae 148
columbae. 22.20... ae 86) | iattenuata,Acuaria_.___./___ esate 217, 220
COMPGN. once eek eee. eee 88 Spiroptera..- 2-2). Lees 220
CT O880 ee oe oe es 136 | attenuatus, Dispharagus_._.......-.-_-------- 220
Mepressa oo SE 137 | aulieatina, Schistorophus______---._-_____- XIV, 287
dispar. 520 2 oo ae eae Glin! saurantius, Paleo... Ley ae 177
ensicQudata — 2.2 Lo beaters 1302) iauratus: Oriolus_...._...._eutiteneeey 2 188, 222
Sissilabium.-.-...2 Ben ees 206 | auratus luteus, Colaptes__._._._._._.__--___- 173
jorciparig. 223. 22.2 eee 1169) lauripalliata, Cbrysotis=. 2-2. 25s. cee ee 175
forcipatars 2.5 22-2. abana 116,131 | jaukituss Colymbuse -....-. .-. 2. estes 139, 250
Gata Fa, oe aD, ee 81 Podiceps. =. ..- 2.22 Sauer aeen 148, 234
gallinge:. 2-20 2+ ee 52) (australiasiana, Gros) 22/020 222s eee 142
gallopavonis...._._.. Pane ae 2s 8l4| jaustralis,“Ascaridias _._.._..__. 27S VU, 78, 84, 85
gibbosa._-..- solace. pat 81 Heterakigs joes 245550). 84
her.maphrodita...<_-_.- 2 5 Sajal hos 80 Struthio.. s..-232-. 2 eee! 31
FROLONS TT Oe So ee eee aa 1440;| jaustriaca,) Glareola.=...2.2:..2..-- sae 285, 288
WOLOT OUR Gas =k Se ee 144%) Babulcus lucidus:<2.5- 02-20... eee 279
irghisensis2 oo se eee 145 | bacaurau, Caprimulgus-_..-_.......-.--.----- 115
dricisas# Siu De: Be Se ee Ae Baca 139. | bancrofti, Heterakis._.........-....------ v1, 50, 66
inflera 22 AO AG G2 00. bet fe See 8c $1),,| {batbatus,.Giypaetus- 24. 22-5222 5 ss eee 138
Rirgnisensis:. 35... 136; 145, 147 | (Barbus fluviatilis. 2-4 sce- eee 368
lagepodis._________... Bee | 88.|\“barnardi, Platycercus_..-s=-1----. essa 175
daticatida.- 28. 25 s22U 222. See 97 | Bélearica pavonina......-2.-+-.--..- 222 89
lum pricoides.=2-0 20-0 ee 82 regulosum-s<-=-2.22+2. wee 89
MNBCULOSG = = — re ron 86 | bellonii, Tadorna._.....------------- eee 376
ANICTOCEDROUG neon 152 | bentocruzi, Subulura_____-------- vill, 106, 110-111
micropapillata...-.... mee 2882 153 | beramporia, Heterakis___.....--------- VI, 52, 56-57
ATVUCTONGLG oon cot SU 20 | Bernicla sandwichensis_-_......-.--.-------- 62, 267
muttipapillata— _ ....-- SLs ee. 154 | biarmicus erlangeri, Falco--__---_--------- 298, 299
OOO ere es UE 155 | bicalcaratum, Polyplectrum_---------------- 386
POP MWOsEsy. oo soe ELS 67, 68,122 | bicalcaratus, Francolinus--_----- 59, 92, 111, 180, 227
INDEX 44]
Page Page
Dicolor, sACCIpiter =... <2 35422 cc8 ssn ie 138 |. brumpti, Allodapa..._........-c- secu s--- 112
SPITODLENG ..- sencn ane = ee 371 Subulura-.-__.......- Vil, 106, 112-118, 131
bicornis, Dichocercus___._-...--------------- 353. |} Bubovmaximus..-_._..------ ope eens 138, 144, 276
Gieusmis, Disphavagus._-- 22522222 a eco 286 Vil giNlanuUss< =: -S2- =.= ects 138, 144
Histiocephalus.......-..-062-.s%--- 2865 li Dubo; Striv.s- 2226. 2.2. .5<2c leeatoese 138, 144, 276
Schistorophus..__-. 2-25.24. ce 985.286 | Bubulcus lucidus..-..--s-s<<<<.-ccccck ees 174, 339
WPT ODEN sn ae Sn hoon ces 986: |} Bucco:capensis..-.. 222+ sis. <kses2<opecseck =p 128
bidens, Ancyracanthus__.......-.-.----------- 288 CBacUNU sci sso cok ecu Seu eee coe 131, 132
SDROT OWS. cee nee ek ess 288 Collaris 22-4 22 oc ccchass cute eS 131
Schistorophus_-_-__----- XIV, 285, 286, 288-289 macrorhynchus=--2222522<adsesee ees 131
DITO DEN) aioe ae a eee 288 mielanoleucosi=<2 =2¢ 2222 2224- ae Se sce 128
ibidentatus, Waleo:...-.- 2.222222 22ccn- use 177 Tufiventris. -- 2. - see ss Meee e se sece 128, 131
IH ArpaGuSi = S22 nk ee ss 177 species: 22. 2-8: --2leicsc 5 sesh os 115
bihamata, Ancyracanthopsis-_-_..---- XVI, 362-363 Striolatus: <2. ec toe eee 128, 131
ANCYTOCONUMIS. 22 <~ a5 on ese nen 362 Swainson sie eee 218, 131, 132, 352
bilabiata, Ancyracanthopsis--_-.--- 361, 362, 363, 392 TaMAtIA. 222s 5 ~ 2 Sede seccest esos 131
Physalopteraciecas-<.5-55-22--4- 296, 307 tamMatina «22S ve SeSees Se esa eee = 128
ISPHIOMEN G2 tee cease ae ence oS an ee 307 teCtUSe2Sssosc2ck esas ekeesuee eee =k Ss 131
bilabiatus, Ancyracanthus-_.-_...----------- 362,363 | Bucephala clangula_....-........--.-.------- 267
Dlamenbachil, Crax=-=.-.<-2.._.=2<i-ses.-55 75 | Buchanga atra assimilis_.....-----...----- 188, 222
Bolborhynchus lineola_...-......------.----- 175" |) Dulbos «@ymea =. 222222 sees sec eeeece = 390
MIGNACHUS se. .o = =e aoe 175 Physaloptera..------- XV, 167, 297, 310-811
PONASA SVLVESUTIS.-- =< ---ceeeanacccanccscee~ 53 |; DuUOsasopiroptera... 25. s222S-.ssen5----- a 205
AMMEN WSs — 8 eS 7D;,93; 220.289: |: Buteo borealis: s2-.-..-2s<-5-.-Ssss8-- 42, 45, 281
bonasae, Heterakis.-._........--.---- vul, 51, 75, 76 IbWitdow==< 5 32 8 eee eee 137
DONASIS. OWA! .cc eso. sheen oc ot ees 53, 81 JaPOpUS: 2-<-<2issSoseds acc weses=s- sss 276
borealis, Ascaridia__........-.-......- vil, 79, 84, 85 VUIPATIS=.--.<.5< 137, 164, 176, 177, 278, 298, 316
Biteole sa 2 Me boo ete eee 42, 45, 281 VILLI PINUS; s.utsSse estas eee oe ae 316
CLERC RIS ot eas oa ee ot ee een see te Ste buted, Bulteds: os: sosss25 sec ecseseee ec dace 137
boschas, Anas_-__ 10, 34, 62, 136, 139, 152, 245, 267, 343 Walco se: = s6sessSuesuteee eteecoeee= 138, 177
boschas domestica, Anas__-___--- 10, 27, 42, 53,62,81, | Butorides virescens virescens....-..-.------- 152
93, 136, 152, 246, 343, 345, 367, 372, 373,376 | ‘Buzzard’’__.........-.------------eee- ee eee 305
HOSUNAS HOTA. “ANGS ec oo ee 815343, 376.) Caccabis chucan.... 2-522 osscastecsee ese 88
MOSIA FEOUCYAKISS 5.2... occ enescene VI, 51, 57-58 Chukoar’:.22.2-c=s Soi eee 114
IBGIAUTSaNINOR = 2. a are eee oe 271 petrosa ..sss<se52 73, 126, 167, 171, 187, 239
TMNUIGIDANS one ee eres Seta ee, 152 petross:spatzl: 2. .222s2 See ceeee ee 192
WinnaAtSse2 oes ee ea eos 371 FUlascc- esses wssd- Sess ones 126, 172, 227
SLOMATISs 22st ane oe eee 152, 274 Saxatilish 2 sc2222 Soe cate aceesesese 88
boularti, Cyathostoma-----..-.------- vi, 41, 44-465 | cachinans, Falco.......-..-.----------- 177, 297, 298
Sclerostomae- = 2-2 -ce2 eee ses oe 44 | cachinnans, Herpetotheres_......-..-.--.--- 177, 297
SUNPOUNMUS oo - ananassae sas seoeace @4: sl), caerulea. Wloridasc< -o2.s-ss5s. 25sec -caceeece 149
brachydactylus, Falco_......--..-------.---- 138 | caerulescens, Geranospizias_-_-------------- 297, 317
pruchyotos.-Aegolius 2222.2 2-_ 7-5 -Ssceec 144°) | caeruleus, Plans: 2s22 o26se wasenst cn aesecus, 297
Obs te. sentece so cscesee 144. 276)4|* ©airina moschates..222s2<+--5--<-22—--s< sem 81
Berit cance eases 138;:276: || cajanea, Aramides.-<-2.-.-=2--s---=---22- 350
brachyrhynchos, Corvus---....-...---------. 39 |' cajanus; Corvus —..s.----+--=------. §3, 122, 207, 213
brasiliana,: Ascaridia: 2-20... = 2- cee ena 2 58 |i calearata, Actuaries. 2so5- 2525-5 8o span nee 249
Fleterakis: =o. eee es Vil, 52, 58-659 Echinuria-2=..22-2ecseceeee XIM, 245, 249
brasiliensis, Ascaridia_...-....-------- vil, 79, 84-85 Hamannide 2. osnceceasceencaae es 249
@arboresso ss ticles ec on en 147 Teter aks aaos oo ceweeweeosaeecee= 96
AI COs tere SAE eee esas 188: | calearatus, .Dispharagus.._<..--2+-5-..c---5<~ 249
IatOuS= = a408 22 Sooo een oe 148.1], californica; Y seria =....<2.-.2+---< 269, 292, 298, 391
eter Okie nears een ee eee 84 | californicus, Lophortyx.......--------------- 175
Momiotusete:: oe 222-8 $23\ |? Callipepla squamats.. =< +..2<-----<-<n<acter= 88
brenchleyi, Carpophaga--._.--..--.------.--- 320 |: camelus; Struthi0;..cs<~.<----c2ee5-6--- =~ <= 14, 31
TOD OCD IS Cae ene 320 | campanulate, Filaria.........-.-------------- 313
brevicauda, Physaloptera___..___- XIV, 297, 307-308 helaviacsc<--sce2snesee XV, 312, 313
brevicaudata, Synhimantus_-_-_ xv, 273, 274-276, 276 | campestris, Caprimulgus....-..-.------- 78, 83, 130
brevicaudatus, Dispharagus_-_---.------------ 274 PICNS! ccs sodeccceco cess eee eee 178
Histiocephalus sesso c2 = 222. 274- |ecanadensis, ANAS. sosccecuec-cesesceenacaces 62
brevipenis, Oxyspirura-_--.------------ XV, 322, $82 | Cancroma cochlearia.-.....-------.-------- 340, 348
SUITO DUET GS 2 Soe ease cee t oe 332 | candicans, Caprimulgus_-...-.-------------- 129
brevispiculum, Heterakis__...-----.-- vu, 52, 69, 60 Stenosis! soe ones epsceneasessase 130
brevisubulata, Oxyspirura_._.-.---...-..-- 322), $85. \canus, Larus... -----=2-s<------<-- 148, 234, 235, 263
Sniropteras sce cee nce eo a 325 PiCdS ose se ee a etee ee ceceaeeeee= 34
bronchialis, Cyathostoma---------- VI, 35; 41, {2,47 | capensis, Bucco....-..---..---------0.-=---- 128
SUNG GTUSLED 2s eee eee 42 Daption seess22ceceso es aeseee= 263
442 INDEX
Page Page
capensis, Oedicnemus. ._..........----....2- 192") || catusidomesticus) Melisua he ene anne 304
POMICEDS! fs assets yeh eects 156 | caudata, Heterakis_......_.___- Vu, 52, 59-60, 61, 394
capillaris, Cheilospirura_.-2-..22.2022022 2222. 287 PCa ee eee ee ere 34, 139, 220
SCHIStOVODRUS se OE SST CAYaNG, PISVa wet eco se teees ee ee 113, 115, 124
Spiro pleras aren sconiey has ein pee 287-)|) CAyanus; Cuculus sooo ee eee ee 129
capistranus, Larus) 22222 ae 233) 42-CAYONNONSIS, MalCO! 222222 eee eee 297, 298
capitate; -Acuaria=~o 3222 a ee ea 240 eeptodon eee eae 297
Dispharynx esos sores 237, 240, 394 Rallusr yo eee 285, 288, 293
capitatus, Dispharagus. 2222220522 2c et PAO: |b concnris; WalCOe se conte ee nee ee een 182
Capito collaris.......-..-.. ASI Ta eatin neat TT5 Ole COndrG, PAOrOR= oe ace soon ee ieee een 209
INA CTOR He hs BAe DI ee een 11571) Centropus monachus. eee eee 127
mielanoloucts yee peor eee 115 SLAM ENSIS ee eee eee 329
Tufl ven trises oo se oe heeen 115 SiNeuSISe eee 86, 101, 125, 329
Striolatus= 2425s 115 | cephaloptera, Cheilospirura.__.....-.-.------- 322
TAMATIAS ee le NA aE eo 115 Oxyspirura__.._-- XV, 321, 322-323, $24
Gaprimulgidesss eeeskne Ska inien keene ne 115 SDIPODT OL ee eee 322
Caprimulgus aegyptius saharae__.-__...-_- 118, 129. |; Cephalopterus ornatuss 22-22 2-<--- 22s eee 186
DACaUTAUS ess See eee ee 115.) })) Cephalostronpylus: ono aa ee eee 11
campestris eis eee 78, 83, 130 quadriradiatus._.....----.- 12
Candicanss = +i ae eran 1290's Ceratospirass eo eee eee enone ae 311, 819, 320
COTtO Pa A) a eer ete aan 129 GNhOCh@eT ae es ee een ee 324
GETTIN SH et ane eer ee 129 oplithalmicas-22:2.-=-- XV, 320-321
enropaeus2 22.2. ee es 129, 130, 313 yesicilosacs ee XV, 319, 320, 821
FOSSILS Ra eee er Me Aarne enn 118 | Cerchneis tinnunculus_._..._....-.-.-------- 276
guianonsiss 2a ee £20; 207; :|'Ceriomis Satyrae coos es eae eee eee 53, 57
LELICODY GeuUsv< eel has we mae een 207)! 1e i corta,,,Revramereso.cs 2. once. ee ane 334, 338-859
TTLOLCUPIS es Ole een ene re 129 TLOPIMOCETCE ee ee ee ee ea 338
ACATIC UG Seite Sees e ra 1155128. chacurd BuUccO.s se. co eee ee eee ee 131, 132
met terer io: <= see eM ne 12077 Charadriformess ne a. ee ee 265, 268
nigrescens] eace sn ese es 1302] Charadrus.Gubius./2-c- 22 2-h oan ee --- 139,394
TUPLCOLS FP SE? 115, 128, 129 niaticil aes eee eee 139, 254, 394
PUTS ss Se ee rere nt 130 Himantopus2-- 2 oe aaa a ee eee 232
scaphinrise:sekshneersuemen ete 129 WOTINOUUS: 2 ee 139, 144
SOMICOrQUabUSs eee ete 129 OOCICNOMUS oa | eae 144
SDECIESH: ween eRe Rs eee a 130 PlaVvialiss 2 celle he Lewes 139, 144, 372
trifurcus!! stent mee eee 12951} Gheilospiruraces cso) con see ass ao eee 211,
MIT CASE 2 Sew ee eee eee eae 128 213, 214, 215, 216, 226, 227, 228, 229,
TINUE Re ee te eee eee 115 231, 272, 280, 389, 393.
VOCIORUS ae ene ee eee eee 130 COPINGTIS 220 52a eee 287
capueira, Odontophorus- -_-__--_.---.-.-_-- 63, 128 COPRGLODLENG. ooo nee eee 322
Oarbo brasiliensis <2 2222702 eee eee eae 147 QTUVelio see XII, 226, 227-228, 231
CONIDOTANUS*Ae eu Sere nes none chara 147, 258 aMUlOSAl cee eee X11, 226-227
Crista tristene Gs tate eae een ae 147 longistrigt@. 2022 ss nee ee 178, 179
GIL Oph us ieee Se Ses ae eee ee 147 magnilavial- =. 02-4 ene 241
Praculnighatnaass see ecee ree tees ere aL aE, NOSED. 22 oe ne eee 238
PVEMAGUS=? Ceceoe Sat ee ee See eee 147 ODRINGUNICE. oo = ae ree 328
carbo, Phalacrocorar--.-.-__. 38, 148, 209, 258, 370, 372 PAVONIS 2.2 0 oc. ees 226, $89
Canlamatcristataenn 25 ee ee es 97, 108, 332 OCLs ee cg Me ae ee eae 280
‘TCariamaihuppe sete eee ee ea 108 FOCUNG AAs coo soos See XI, 226, 229
Carine noctuaielaax? 220 siete 119, 242 siamensis.. 22.0 ee 329
carlosi; Subulurat." =) -<t2e ee VIU, 105, 113-114 skrjabini::)-0 022 oc aeeieeae 226, 389
carolinense; NWettion=<<2222222:==82 "~~ 7 f= 246, 248 SDINIOSA-- ose eee XII, 226, 229-281
Carpophaga brenchleyi_-_-.--_._._---_-_-_-- 320 wncinipenis. . ...caca-<acse----- 165
carunculata, Anthochaera---.......__---._-- o22 || Chetracanthidea:----- 222-2 hee aes aa 363
Caryocatactes; (Corvus= 22. 2°50 '" os oe 220713) CRE acONIAUS: <2. 3.52 2-2 2. ceeen eee ness 364
INICIPAg Re eee ee ae 39, 220,373 | chelicutensis, Haleyon._.........-.--.------- 188
Gascuraiosscar as cSt et esteem a re 42 | Chelidoptera tenebrosa..........---------- 128, 131
CASCarE AC ascara®: )t oven iee rn. Vee ee eae 42°] | Ghenonetta jubata:--2-.2 222 ee oe 60
CASDICA Starrs t sane sae ear oe 237 | chenonettae, Heterakis_.........------ vil, 50, 60-61
CAStOL, SWOrzeNSeN sooo eter ee ea 1485) i @hanopisialrata.- = 2-22 o sess eee spe etre 53
Castiariusralentus srs 2 ees ee 41,44 | chevreuxi, Amidostomum.... V, 19, 20, 22-24, 25, 383
Cathartesturubu------- 0°. ee $30); | S@HOVTONEIS. cone one ceo. ee ee oe 211, 231-232
catheturina, Ascaridia___-._________.- Vit, 78, 85-86 TOVOlUtA: = - <<. uae ces = XU, 232-233
catheturinus, Helerakis__...-2..2._-...-..-_.. 85 | Chloephaga poliocephala_..........---------- 20, 77
Catheturaslathami. 272 hoor eee 66,85 | chloropus, Gallinula__........-..-...- zececk 20, 139
Catorrhactes pachyrhynchus-_-...______.----- 236 "| icholiba, (Otws. 2.52... ono ae eee eaoa' IABeO
INDEX 443
Page Page
cholodkowskii, Thelazia._..--_.-.-- XV, 312, $18-$14 | Colaptes auratus luteus._...-.---.----------- 173
Chordeiles semitorquatus_-._...........---.- 129 | colaptes, Habronema---.-------.------- X, 178-174
Chroocephalus ridibundus_.____...-.-------- 148 | colchicus, Phasianus_-.-...-.------- 10, 34, 53, 64, 65
chrysactos, Aquila... 2 05.222 o 22a cee eck 138,301. "}) colini,; Cyrnea.<.-.-2--.2.2-2- 25524 ee x, 168-171
MalcO esc. cass oe es 138 | Colinus virginianus__-.--...-..-.-.---- 9, 53, 88, 168
Chrysolophus pictus_...__.........-----.-.-- 53.) ‘collaris, Bueco._._-2.----..<- caseseteeete st 131
chrysomelas, Phasianus....__........--.----- 64 Capito ..2. <2 22... 22. eee 115
Chrysotis auripalliata................--.--.-- 175 *BrogOn..222.-552-22¢0-e Sese 171
festivas ic. -<-sco22 22-2. tes 80 ;|, collurio;, banius.-_.--.2.2.-.-22ssseet 201, 220, 221
guatemalse...-..2---=-...-4-5.2. 175: || Columba:arquatrix......_._.....cuzeeee A eo 87
leucocephalus. ....-.-.---2-s<--..- 175 domestica ............--ueees2tL._- 87
ochroptera 222 == 22-4 een So 8 175 dom. laticauda. ... 21228224 87
ehucar., CACCaDSicc. on cc oa se aso n stone sok ce 88, 114 putturosa= 2.22.52... eshte e 87
Ciconia alba.............---- 34, 44, 152, 168, 296, 307 livia. 2au- sr es et 2c 87, 239
alba asiatica.. 2.222222 -nseeeer-ct 168 livia domestica__-.....-__---- 12, 341, 343
(a7). ee ee eee ee 168 Pleuis 2S. = 5s ee 87
maguari----sss2-s-225cs= 50, 74, 168, 245, 256 TiSOFia: .- -2 3. Ssoe=55- 55S 87
TMGTS aos ccccacccese 34, 44, 150, 152, 281, 372 Speciosa /--..-si sod tet 87
elconia; Ciconla-2.2<-s-. 22 ss6-< 222. wesc cice 168 talpacoti..c2.-- se ees 87
ciconiae-maguari, Spiroptera.._.....-.-.------ 256 | columbae, Ascaridia-__-_---_--- Vil, 79, 86-88, 89, 96
cineraceus, Circus............- 138, 176, 276, 277, 298 Ascaris: 252555 - sees son ees eae 86
AAT CO ea a ca = 177,277 Heteranis... -..ccocidoesees os 86
cinerea, ATdeS..:.. 222.2... ccceee 141, 142, 152,209 | Colymbus arcticus____--------- 148, 265, 267, 271, 367
GYUS oes 2 eee ee ae 142, 286 atrigularis.... ... Ge sasee". zesce 148
Pend i eae ca 2 ae 10, 34, 53, 88 quritus :--.._ 2-5. sesesesecsee 139, 250
GiNereus, VAMSCP ==... so a. hoa k laces 10, 20, 62 cristatis....... . so seaseteeiinced 2 270
WilOUhes 222 2 oe oon a 2 138 Nigricans:: 2. oes ap eae 148
cinereus domesticus, Anser_-.-- 10, 11, 42, 53, 246, 373 rufovulgaris. ....--<2scteci ese 148, 267
Circaetus gallicus_..........--.-..<--en:e-- 138, 298 septentrionalis_____- 148, 234, 267, 367, 372
Circestus pectoralis....._..-...-.-2..:<.--<e0 138 | comata, Ardea... 02 ..--cecustec os sess. 152
circia, Querquedula__............-----.------ $80 | communis, Coturnix.:-........----sssss.-- 53, 88
circularis, Ascaridia..........-.......---- vu, 79, 86 GQTUS 3+ 2. ccc sce cows s eee etes< 99, 143
HOUT ORS 223 cas cee ee caaces 86 Nisus: 2... ---0<c. Seuediaeas 277, 278
circumvallata, Heterakis_._........--.--- Vit, 52,61 || compar, Ascaridia.-._..2..------s<te2 Vil, 78, 86, 88
Circus aeruginosus.......-------.-- 137, 138, 176, 298 Ascaris. 22-252. oa5ee tee sesarees 88
cimeraceuS......-...-.-- 138, 176, 276, 277, 298 Heterakis 2.5... scx 555225 s ee 88
CYaNeUssces-c<-50n2 55 -255-5 138, 176, 276,298 | compressa, Ascaridia.-.~........-.----------- 78, 88
MAC IOSUS= 22 oe sa saat tose tee 207 Helerakis...-_----.ssousseeaes— 88
WV RareUS eee - coe 22 oe eens 998} .confusa, Hartertia. ....22--2---22-- XI, 191, 195-197
WUUIS 22-4 sees kee eS 138, 177, 298 Tetrameres_.. XVI, 334, 335, 338, $41-342, 343
Spilovhorax.~.2 2-2 -- 2 = eso te 90 | conspicillatus, Pelecanus-.-_-.-...-.-.--------- 148
cirrohamata, Filaria (Spiroptera)_..__.-.-.--- 266. | contorta, AcuariG.......-.<vssebueSieu-4-s-5 =m 249
ISU ODIEN Gann one or ens 266 Mchinurla= .20. 22 o6ezecce XIU, 245, 249-250
Streptocara..._.....- XIII, 265, 266-267 Hamannia: . -..2 22 casts --o sete n 249
CU TUS HGS. 22-2 2 ee sade lk OTe 318 Microtetrameres-----.-.-- XVI, 351, 353-354
PRCA A ee ee XV, 312, 318-319 Tetrameres......-seusncvs--2-c5--=- 353
Clamidonemna =). 2 = 6oe oS oon ecb sens 295 TropidOGerca.. -. +. -sscesekeest~seceoe 353
clangula; ‘Anas--22 2-2... .< =<... ence 28, 147,267 | contortus, Dispharagus-..---.--------------- 249
ASCP es eos esses sates A lk 20 | Contracaecum__-_. 135, 145, 146, 147, 148, 149, 150, 151,
IBNCepuSlas. sccssesccicccecs coon 267 152,153, 155, 156,157,158, 159, 160, 161
Gisuclonetta:--- =. 22546 <s52525-52 153 andersoni....-------- v, 147, 149-150
INS TOCR Ss scccceesaeeeee anes eee 20, 267 engonium)...--<...isss¢ x, 146, 150
clausa, Physaloptera:. 223.2202 eee 295 Haliseti_....-.-tcs2e- X, 146, 150-151
climacocerca, Hydropsalis._.....---.-------- 130 magnipapillatum--... xX, 147, 161-162
Coccigylormeziissac oe8 hoo ee so ss45-s5aesssen 127 microcephalum.. xX, 152-153, 159, 160
coccinea, Tetrameres-_-_._.--------- XVI, 335, 339-340 micropapillatum_. x, 146, 147, 153-154
TTOPIGOCET COS ae tence tgshn’ 339 multipapillatum -_-_- X, 146, 154-155
Coccyzus melacoryphus- ----.----.---------- 107 multipapillosa_.__...--.------- 154
melanocoryphus.-......-.-..------ 115 Ovals oe eee X, 147, 156-156
MinOes da—- oss =2 222252 2sceeeeetws 115 praestriatum__-.....- X, 147, 156-157
cochlearia, Cancroma.....-.-.------------- 340, 348 punctatum:.sss22--¢4.-< x, 146, 167
cochleariae, Tetrameres----------- XVI, 335, 40-341 quadriclepe..s.ccccnse2-=--< 152, 153
COCO WATOGR. so see sce acne e ane ences sean 141, 371 rodbaini.._.<w<se-s X, 147, 157-158
Wodiostonmiuuini {os 28 =~ see eee s- 255222 80-81 FOsaTiUM=osses02—ssc< X, 147, 158-159
struthionis: 22.2.2 52222 V, 1-82 scotiin.--.2iG.2leuecs X, 147, 159-160
eoclebSx SYN Samus. .2es-s5--nacseeens a ---se=— 34 spiculigerum_---. X, 141, 146, 147-149
444
INDEX
Page Page
Contracaecum tricuspe__-.------_--- X, 147, 160-161 | crassissimus, Dispharagus.._._......------.-- 243
Contrusileucotis--.-.-- 2 eS 175 | crassum, Porrocaecum........._-.--------- 136-187
PAVUS. Wo see SE ees BO Crassus. sispharagusecc sa eee 215
pertinax Ss ee £75 Wo Crax'biumenbachii. ee ee ee 75
Solstitialis.c.22 2.) a 80 fasciolata css Succes Me need 312, 317
Coracias:abyssinicuss..2222522- 127 LEUNG URNA 2 en ne Sac ape eo 331
corax, \Corvuls:2.5.3cceese Goes 220.3) CRECCAs WA MASE Aes mean eny Meluna we ee ee, 24, 28
corax tingitanus, Corvus___-_-.-______- 203, 220, 355 ‘ASOT RHIAN od RR ER on eee ee 20
cordata, ‘Acuaria..222-.22-222-22222 XU, 218, 220-221 Nettion i.e beer en ered 384
Ascaridia. 22-5252 2=2 ees vul, 79, 88-89 Querauctialassies eee ee 384
Heterakiss2 32205 ae 88 | crepitans, Oedicnemus.-_-_........._.-_-_--- 139
cordatus, Dispharagus__..-..-...----.-------- 220:| ‘erispus; Pelecants’ 272 2e eee ee 153
cormoranus;'\Carb0=<..2< 252.5 eS 147, '258)) ‘cristata, VAscaridia = ees sao ee vil, 79, 89-90
cornix, Corvus. ---- 34, 141, 218, 220, 221, 308, 334, 351 CATIA TN els 207 2 Ae anes 97, 108, 332
COTONALA SpiTODler G2 oe IE 293 uri gurl pert A Ne ea eae ne 24
MV SETA sR asec eee 269, 293, 592 LET OUT ONES SIMRAN CARR Se ee ne 89
coronstus) Maleoiss 22 eee ae 297,208: Uneristatuse Warps eee eee ana 147
Harpyhaliaetus....-2 2252422 297 CGlyni bush ere Wet enn ie eee erauee 270
Histiocephalus= 22.2 aes 293 Dicholophus! tse 115
Eystrichis:..-.. 22.32 XV, 376, 378, 379 Fotertisx2 i tse eee A a ae ore 207
corone, Coryus__.--_--.----.--- 34, 218, 220, 330, 355 Microdactylus___....._.__._.- 97, 128, 130
corrugatus, Cranorrhinus_._-__...-.--------- 188 Pavosse asa: 34, 53, 63, 103, 310, 311, 325
cortopsan, Caprimulgus__.._....__.....--.--- 129 Podicepse st:3222 0 ak aerate 148, 155, 270
corvi-cajani, Spiroptera._...........-------- 122, 213 ‘Prodiceps ec as2e2 kits See ee ee han. 372
Corvus americanus2.2 2... 22222... AUP. = 220, 355 Wanellus22ic222e25222 22, 139, 141, 260, 268
brachyrhynchos: .-2-2 2022222... 39 | cristatus brunneus, Tachyphonus- -___-_--_- 358
cajanuse222 2k ee 53, 122, 207, 218 | Crocidura leucodon............-------------- 206
earyocatactes). 2.2. 2 ee 220: | crocomotus! Teterus: ce 2822 ee fee cee 323
COFSKee Sots tates aoe See e cone 220 | Crocopus phoenicopterus_.......------------ 87
corax tingitanus_-_..-_..-...--- 203, 220,355 | crosi, Physaloptera___...------ XIV, 296, 302-303, 394
cornix =... 34, 141, 218, 220, 221, 308, 334, 351 | Crossoptilon manchuricum___--.-.---------- 64
COTONG Ss oe ee. 34, 218, 220, 330,355 | Crotophaga ani_...........-.---------- 124, 207, 323
frugilegus=2..---<<-==--. 34, 218, 220, 330, 355 MAJOR oi 22 Seta eee 124, 171, 323
glandariusci 22.22: +2.-- Sessa. 218, 220 | crotophagae ani, Spiroptera_....-..-.--------- 323
monedulazeccestecee ects ee 34, 261, 262 majoris, Spiroptera._.....-------- 323
Please eee 34, 220'| cruentus, Ithagenes__..........-..-.--------- 64, 81
pyrrhocorax22222-- 2 lees a 220 | crumenifer, Leptoptilus__........--.------ 254, 369
scapulatusic..-22222-2- Sots 223 | cruzi, Microtetrameres_....-.----- XVI, 351, 852, 358
Species sce sekece ect 35 Tetramevesnct iene ee ese er 333, 352
Coscoroba coscoroba--.---------------------- A247; || \Crypturus CUpreUS #2 otter eee 55, 72, 74
coscoroba, Coscoroba. - 2.222 SS eee 42,47 NOctivagus: 222: [ioe eee 74, 128
coscorobae, Cyathostoma__.--..---._-- VI, 42, 47-48 DALVITOSiTIS 32 oa le eee 121
Cosmocephalus.......--------..--. 211, 233, 234, 235 SPOCIES =e ie eee ee eg ee 55, 128
aduncus=.2= 2s XH, 234-235 TALC ADA cre. Sacer ees 128
lates ees 8 295) |) \Cuculusicayanuse) ccc eee eee 129
diesingi. =... 2-22 uaae XII, 233-234 melacoryphus =< ~ 222.2252 22 2222.-.. 1291309
dresingtt 229 Oe 233 melanorhynchus........------------ 128
obvelatus_-__.-___- XU, 234, 235-237 NAOVIUS Se ae oer es ee ee 115, 129
paptilosud) 422. IPS 235 SOT CUTTIS MR eh eee eee ee 107, 115, 309
@oturnix'communis-2 224 Seer! aot 53, 88 SPOCIOS EE tee a ensue eae 297
CObuENix 22. - 2S. See Se 167 tingazraetee Stee ooo eee 115, 128, 129
dactylisonans..2t@2uoRne _- 53, 88 tinguAciise 57525828 Wee ese eee 207
Golagorgnel: 22. Aen = 130) || <cupido, Cupidoninesseensn ieee ene 53
coturnix,(Coturnixs see Se 167i}; Capidoniaeupido”*2=2s5 2 ea eens 53
Per dixtss; Me a ee hs 53 Meupreus Oryptunuses sss esas stk one 55, 72, 74
chacts;iSpiropter ge se Tg yON es ee Boll. lM curcata,Nielerakigsca ees eee 114
cracis alectoris, Spiroptera 2.22 222k 317 Subulurasi20= 2a vil, 105, 114-116, 116
Cranorrhinus corrugatus.---..-....-.......- 188:\\ (eyaneus Circus! s:=2!s= 5.520 138, 176, 276, 298
CHGS ABCURIS yo Oba ee 136 Falcons savexs Hee Ces eee renee 138, 177, 276
Physsloptera.c 222k ae 296, 802,394 | Cyathostoma_-_._-.----.--- 33, 34, 41, 42, 43, 44, 45, 47
crassicauda, Dispharagus__..........---.----- 267 americanum. ___._------- VI, 42, 45-47
Spiroptera.. 525 Cage 35k 267 Dolartics se eee VI, 41, 44-45
Streptocara-----.---- XIII, 265, 268-269 bpronchialis®*2<""222= VI, 35, 41, 42, 47
crassicauda anseri, Streptocara_......-.--- 267, 268 coscorobae 22-5 2s22 2822" VI, 42, 47-48
charadrii, Streptocara. xml, 265, 268-269 aries «o> xe eevee et VI, 41, 42, 43-44
Crassissima: cA CUaTiOs = ose eee «eee 243 tadormae 2222 222h ween VI, 41, 42-43
Dispharynx.22os-ee 5 eee 287, 243-244 prariogatum._). 2 2-- 22-2205" VI, 41, 44
INDEX 445
Page Page
cyathostomum, Sclerostoma__......-.--------- 43 | diesingit, Cosmocephalus_...--.------------.. 233
G@yclistomeney.-.....-..----ssseeeetisvecs.__- 30:}| sdifferens:, Heterakis 2 <~ 235-8 oS c ccs 111
cygni, Echinocephalus__.......-.-.....------- 378 Subulura.......--s-4; vil, 106, 108, 111-112
Hystrichis—.- 2-222... XVII, 376, 878-379, 380 | digitata, Thelazia (?)__._......-...___- XV, 312, 319
@ygnus'atratus.22. =~ S25 atts. 52,.61-}} tdilophus, Carbo...-<.-=<.-2<<-s2-<c20-- 2-2 147
melanocoryphus- - ..=--..-.=--u2---- 343 | dimidiatum, Sclerostoma_._......-...-------- 32
OlOR? 222 3 2 once os eS 61, 376, 379, 381 | dimidiatus, Deletrocephalus.____......__- V, 82-83
olor domesticus. ...2se:s-2e22t cc 246;;| 2Dioctophyme. _ = s2...2-s-2eseeeeee en soos =e 366
cylindrica, Ascaridia_.........___- Vil, Vul, 78, 90,91 | Dioctophymidae-....-..--....--.--.--_--. 866, 367
eter kis. =. se ee 90:'| Dioctophymoidea.-=-=.2.-.2.2-22s--_.- 162, 366, 375
@ypselus apus....-.---- espe - ssc2ee03. seek 34 | Diomedea exulans_____...-..------_---_-_- 263, 339
Cyrnea____ 164, 166, 167, 168, 171, 172, 310, 311, 390, 393 melanopnorys =< eon te ees oe 159
bulbosa: = 22-2... see sed 890 | Diplopterus naevius__....._......--------.-- 115
colini.- =. =< s<i2essesesedect. . MS 167,,168-171 || <dispar, Ascaris.........---=.42.s..-<-.------ 61
GEV COrent o-oo X, 167, 168 SICO see eee ses ean eee 297, 298
OXCISH aos eee coe eke X, 167-168, 169 Heterakis:.- 222-222. -2a32-- Vil, 52, 61, 62
parrotitt: 2... 2.252. 2Se es be, 167291 {| Disphoragus:.----...2<2 2s sose0--co-<-- 212, 216,
SCT GU 2 2 > Gaon tape 172 217, 219, 226, 231, 233, 237, 244, 258, 272
Semilumnaris.. 2226822555... .....-- 167, 170 acwleniise = a. ae ete eeeeeeasees 254
Seurati. .. sagt beset 8 e.. X, 167, 169, 172 OOUNCUGs..n2 = = ese ts notaar secs 234
ROUT OLE Ss c= 22 ae Saris Qe Se 172 Glatus 2 se525 sod ee 281
eystied, Wilaria - =... 2282. eee 368, 371 Gnas oe eens enaes 281
cysticum, Agamonema_.__......--------------- 371 OnthUtisn 2 cate eee e eee eee ee 218
Maccloleachti_ =— 22 ssa selon. cee dsds 314 OTCCGE: 2 5-8 Pee ed 248
aacelonis, Wilaria...-2222- %22852 - sosdeces ke 314 attenwatus: ©2222 22255- sone accncem 220
Thelaziaotz ssc. 2-53.54... XV, 312, 814-3815 UiCUSDIS se oe ae eee eee 286
dactylisonans, Coturnix-____- ee ent 53, 88 Bidens =o js cee ee 288
Paphnia pulex.—.--=<..-...<s.=--- 246, 248, 343, 345 brevicaudatus..-....--.<..-----<- 274
iDaption.capensis........=...----=--=st2 5522+ 263 COICGTOLUS == oa nase ete eee 249
Gasypus< Strix... ....--. .2usecos cue doco 138 Capitates 2a. ose ee ee oe 240
decora, Streptocara---_--.------ XII, 265, 269, 270, 394 CONLOTIUS 22> o-oo eee coe 249
RY SOTO) 2 = ooo oan ee SE Le 269 CONGQUIS == 358 eee ee 220
decorata, Echinuria_..-...---.---- XIII, 244, 250-253 ChOSSICOUNG a3 eo ee eae 267
decorum, Prionostemma__..------------------ 269 CHOSSISSUINUS omen ee eae 243
decors, Dispnaraguel....-.-2.<<.-sasect ees 269 ChOSSUS a eee cose eee eee 215
Histiocephalus.....2<-..< 2S2z secu 269 CCCOTUS Sd daseccen eo sacncseet = 269
degenery:Waleo_3s25s2 38s 2c _-_satiiesscs 138 denticulatus.........-..-.-..-~-- 275
delagorgnei, Coturnix.......------.---.------ 130 GepressS=. <2 2 oceans sess 221
degisndi; Oidemia.-_......22cast......=.--- 293 CUiptiCussaccsen~ scnasennsen-s-=— 277
délalandiit Vinago_.. =... ssise--.----5-6- 91 Clon gals? 22322 n eee noes 259
IClELV OCEDNOIUS 8 2— ee oo Sa eo nas oases 30 falconis-subbuteonis......------- 275
Deletrocephalese = <<< 2— sates ao sacooccan cece 30 ORACHiS hone ee eee 222
Poeletrocephalust=.2- 226 ...- 2.222522 -5<-.-- 30, $2 ONUve ose os nace wsene coe 227
dimidiatus:2:25:2- ...-.-- V, 82-88 ROMAUUG 2 5 oslo on asesaeonse= 278
dentata; *Perdix...---Sd2ie20sete-. .-=.------ 128 hamulosus: 5. ---ascnceecssoune= 226
denticulata, Spiroptera_...-.----------------- 288 INvOgiINGWwS == =2- 2 secczo=--= 218, 279
Synhimantus......... XIV, 273, 275, 276 INVOUIUE: on oo ce cant eneecee-oae= 276
denticulatus, Dispharagus -------------------- 275 laticaudatis:..2.-2.-52------=-= 290
depressa, Acuaria.....-.----------- XU, 217, 220-221 LACE DS) ee ease ease see ee 276
tASCETIS 2 co a coce set ie ewes casas 137 1ONQCOTNOANUS =< ac nee owe = 255
Wilavid..o<s22d2-25c0c tae eeotsesee 221 longevaginatus......------------ 256
WUGONI Oe e on2 s cawctns os 2250 137 macrolaimus._.-.--------------- 213
depressum, Porrocaecum.-.------------------- Ix, MOQNMAUIOIS. on noose enn eeeee= 241
136, 187-138, 189, 144, 145 MAMIUOTIE 2s cee amaasesessssos54 213
depressus, Dispharagus.-...-.-.-------------- 221 MiUsCiCOPGC... . .-0--asa-sa-5~=--- 214
Desmidocerca:...3- 2. so ast. tettssets 208-209 MOASULUS Son on ae nee ene nn a 237, 238
aerophila. . ......=s2- XU, 209, 210, 264 Obnelatue-< 2+ 2-825 cSanetesenn-o= 235
NuUMIdICAW 4. 2-~ Sethe set. 209-210 ONNMUUGSL ososc nee oceans o> aaasoe 223
Mesmidacercidaes-..22-2---.5----5 55552 — 163, 208 DOD US2 a cceea= ae sea = 224
POST COCELCINAG occ 2e4 cote conn a nsesena sa aoe 208 papillosus....2.-----<-c0----=-5 235
destructor, Paleo...--.--....~.ssseusue-~---- 317 GQUaAdTHOUUS 2 222.- cemacccsncese= 215
Diaphanocephalidae.._....-.-.-.------------ 6 rectovaginatus ....--------------- 243
diaphanus, Fundulus-....-.-----.---------- 371, 373 VOCUS ont Sao onan ennanoennaeans 280
Dichocercus bicornis......----...-.-.=-s-==> 353 FEVOVMUS co ceacclesacesacensenne 232
Dicholophus cristatus---.-.------------------ 115 TOUNNGGIUS <<—- cece naaosanense oa 229
MArgravinca-a.sesncc=5 97, 108, 130, 332 SPeCles.2 28 2-2-5 2 secee ene == 227, 259
diesingi, Cosmocephalus. -...---------- XI, 233-284 species NOW ?...--<ss<-<<--s=e--- 391
446 INDEX
Page Page
Dispharagus spiralis. ...-...----.---------- 238/276) || elliptica, Acucria..-=-=- 2200 ee, ee 277
spiralis columbae._.-..-.------- 288 Synhimantus.-_-. -.---_-- XIV, 273, 277-278
aquamatus === 22-3 257. | ‘ellipticus, Dispharagus--.-.-.25 22222222 277
sygmoideus:- 2252522 MORE 23 MG 282, 1) elongata; -Acuartal osc soos a 259
subulesirccoscsecses SEE 224 Htlarie 2 222552235 5552-0- SS 259
tentligs. 2. SP Se 225 Rusguniella. - 2. Sues eae xm, 259
truncatus 2 Cas 8 ea 190 Sproplergse<cas3sss3s-see eS 259
uncinatus:=25is2222s25- tees 246 | elongatus, Dispharagus__..-.--.-------------- 259
Dispharynxr. 2-22-2225. 52544 211, 126, 233, 287, 238, | Emberiza pecoris_----.-..--.---------------- 177
239, 240, 241, 242, 243, 244, 248, 252,391 | emmerezii, Spiroptera_.......-..-.----------- 325
capitatascsss222222 See 237,240 | engonium, Contracaecum.-_-...--_..---- X, 146, 150
crassissima .222 2 SSeS 237, 248-244 | ensicaudata, Ascaris..._....-.-.-.---.---.--- gov3e
laplantei..<<:-2:2-22525 XI, 237, 241, 242 Pusarignccssacccsscse see 139
magnilabiata-......... XU, 237, 241-242 | ensicaudatum, Porrocaecum-----..-.-------- Ix,
nasttassszsscevscsscesseo% XII, 237-238 136, 189, 140, 142, 145
HOCtUREs=s252522552 MIT 297/241, 242-048.) “!Eipervier 7© 2. ooo onan eee om er en eae 355
rectovaginata_-..--..- XI, 237, 248,391 | Epomidiostomum----.--.------ 18, 19, 26-27, 28, 384
SPACieS MOWiT esos fe Sacco nec ee lS 391 enatinum.....55-. eee ee 27
spiralisssss2 eee XU, 237, 288-240, 241 orispinum_-.-_.- V, 28-29, 384, 385
aiurnus; Caprimulgusess2so0 Mies oes. 129 querquetulae___...--..-.. 27, 884
dolichocerca, Ascaridia-_--.....------ vii, 78, 90-91 skrjabiniss-- 22.2223 27, 384-385
Feber akiten c= 22 sae eS 90 uncinatum_----. V, 27-28, 384, 385
domestica; Columba a222 2s) MiP 87 || epops, Upupa...2...-.25-.- eee ee 190, 203
domestica laticauda, Columba_--.._...------ 87 | erythrorhynchos, Pelecanus-----.----- 148, 153, 154
domesticts;-Amser=* 2.52 2c Cen ena 10,101 | erythrorhynchus, Irrisor_.-.------.---------- 188
Passer sc sscey so Sie to 355 Rhamphastos---.--------- 186
domesticus tingitanus, Passer___....-_.------ 203, i] WWsox lucius. 22-22 55.5255 SD See 368
dominicensis, Pionus (Psittacus) -__-..-.-_-- go || ‘Wunematedas. -..2-.022.252-222 eae 391
Podiceps 2222 ee ee ee 148 || ‘europaea, ‘Talpa.l_22220- 2.2.22 See 138
douglasi, Ornithostrongylus--_-..__.-.-- v, 11, 14-16 | europaeus, Caprimulgus------.-------- 129, 130, 313 .
Strong ylussre asses eee eR: ek 14 INyctieorax.. 2222-2 2easecc RS 152
Trichostrongylus_...-._------------- 14.;| eurycerea, Cyrnea_._-_2--2--s22--- S238 X, 167, 168
douglassii, Strongylus_._-..-..--.------------- 14 | ‘euryoptera, Spiroptera__..-_-.---2-2---=------ 201
dresseri: Somaterias=2> et eeieei 20 Viguiera_.-.. 2. -sieteseass XI, 201-202
Dyryocopusimartius*=s*"="+=~- steer 916 | ‘Hurypyga helias-----.-_-___. Seseaaseess = 362
GRU AUar tir | ee Senet Ane been ene WE es 338 || wurystomus afer. ---..- 2 ea leseeeeee see 124, 127
Tatranteres a7 ne tao ee XVI, 335, 342-343 | Eustrongylides--.- 358, 366, 367, 368, 369, 370, 371, 372
dubius,* Charadrits(*2=2#s<s6s or Mane oS 139, 394 africanus_. --_.--- XVII, 367, 369, 370
Eaptopellus22 2 sessee tee ates 253 elegans...---—- <2 ueeeee 358, 372
ecauidatus: Helotarsus 22222002 ee eee =e 137 excisusc--2--- 33 XV, 367, 870-871
EChINOCEDRAVUSACYU NI co enc ce ee ee tene ence 378 ignotus..- XVI, 367, 368, 871-372, 373
Mchinuria--ss2s 1, 211, 244, 245, 247, 248, mergorum..---= sssseace==se Xvi,
249, 250, 253, 254, 256, 257, 258, 393 335, 351, 358, 367, 72-373, 374
FEN (0 Uo Ce ae aged por apa XI, 244, 248-249 papillosus _ XV, 358, 367, 878-874, $75
cealcarata ses. nc soeec coe XII, 245, 249 perpapillatus-_--- XVU, 367, 874-375
contortal Sees ~ Xml, 245, 249-250 Species= 2: <= sssseaeare XVI, 369
decoratanw ees XII, 244, 250-253 tubifex.. 2d2uaieeee XVI, 867-369,
argine ee sen. XII, 245, 252, 258, 254 370, 371, 372, 373, 374, 375
Horrida seers he See XIU, 245, 253-254 | Hustrongylus papillosus_.....-- 370, 371, 373, 374, 379
jligadornata® aes XIU, 244, 245-246 tubifer._-_---_------ 225-55... 367, 371
leptoptili-_----_-- __ XI, 245, 254-256, 256 | Euxenura maguari_-.---...------------------ 50, 74
lonlgoormatarce ess eat eos 245, 256-256 | excisa, Cyrnea._.----------------+--- X, 167-168, 169
longevaginata___----_-. -. -- XIN, 245, 256 Spiroptera.. J... eae ee 167
phoenicopteri-___--...--- -- -- xu, 245, 257 | excisus, Eustrongylides----------- XVU, 367, 370-371
SDN era res ere res ee ed 954 4| ‘excubitor,. Lanius. __--_ >_> -eeesseeee ee 201
squamata___._. XII, 244, 253, 254, 257-268 | excubitor dodsoni, Lanius------------------- 203
NricinAtagee sce eee eee xm, 245, | eximius, Platycercus__.-...-.-..------------- 175
246-248, 249, 250, 252, 253, 254, 255, 256 | exulans, Diomedea------------------------ 263, 339
Hiclectts pectoraliss-2°2 2232 — wie aoe os 990} fabalus, Anser...=...-.- 25-2453. 2a 20, 62
TOLAUUS Soe ee ee ee eee 175 || *{eleinelli,, Acuartas. 252-23 249
wR TOG eae er ee a ee ome Ne 141 Sptropterg..----.- 2- == Jee 249, 250
Ggretia, Herodiase= a0 io ne ee eee 152)] ‘Falcinellus-igneus......--.._ _sSsedse ee eee 250
ulanuS CaCruleus sen nae eee seen Sake 997. | Mfaleinellus, Ibis.2..=2-2.=-. 29 ies 250, 376, 381
Weuicnrns aa ates see eee aS 997: 1) SP alo AeSGION 3552 sa es ee ee ee 138
elegans, Eustrongylides_......-.---.-------- 358, 372 Slbicillaccc tase Ee eee 138
15 CE tg tl (Ub jes eee teas ah i a 370, 372 AIDICOMIS eso SS es 177
Sivongyitist see awe ee coc noe eee 372 APVOrUs =. «21 c.~ SU 138, 298
INDEX 447
Page Page
T CEN COGS 22) ee a 138, 177, 243 | femoralis, Falco_.............-..---.-------- 280
SUrICapUlus. - S22 52.2 oo ee ee ek 207,208.) ferina, NYrOC8=. =~ - ceo cecen ca nececuce cca 343
BULANGUS: 225 eo ee ek L7G ACTUS, “ANSCD 2 2a oe ee ele 10
biarmicus erlangeri__.-.....-----_-_- 298, 299 | ferus domesticus, Amser__...____.---.-_-.-... 10
Didentatus. ——<------- = 2. eecacece 177 | festiva, Chrysotis...........-..----.2-.-- 2. -- 80
brachydactylus.-.......-....2-.-...... 138 | festivus, Pionus (Psittacus)_--........-._._- 80
prasilionsisis 22 oo se se cee eS 138 | ficheuri, Habronema._-_________.-.-- X, 172, 174-176
TUCO Se a a re ISR Litho GON a a2 ocec a eons oe 203, 233, 244, 258, 272
CAaCHiNANS 5 =. 2.2.28 eee 177, 297, 298 aculeata..........------------anececece 254
CAVOUNSIS. 222 Snr eee ia eee = 297, 298 LO Fae ee en 281
CONCITIS ee Soco8 ete ode eee 182 TOLLE are ce ee ee 317
CHE VSACLOS oak ee ee ee so case oe ial 138 ANB Se eee 218, 220, 330
CINGLACOUS << oc n essen dacsowaee oe WT lhe campanulata_._._.-.--_--- eee 313
COTOWAUIS! = 2 ee ee 297, 298 COMMCN OLE 22 he eee 220
CVANCUS 322627 oe ease 138, 177, 276 CUGSAGT Oeete ees eee eee 318
GOPONCI 8a oe es ae 138 CU SUC carta een ne et 368, 371
@eSiTUClOn. <5 a2 osc sca ceus can oetencwce 317 OQCClONIS 2 = oe es ee ae 314
GSD Oe er rae Na eeeaeec a anne see 297, 298 EDT CSS Wa Soe a are fee ee ee 221
TTA ESS ene ee ee es 280 UU oe oe a ey 338
ACCA G LIS nee ee Se 207 CLON DOLD = Baas ener ee 259
gallicus:....-.<-<22s5-2sce5: 138, 297, 299, 300 falconis magnirostris..............---.- 313
gracilis... .2 2 ss esas ence sectors ets ce 298, 317 QOUINATU Moo oe. anno meee oe 197, 198
hhalisetus. -...-gicctzzcSeccwedaccesce 137, 207 REIN OL Ge ot Bes ee 278
BT PORIANS hoes te at inl Saag 138 taticaudalas 225. eee n ee 290
RE ZOPUS en <n e oe et 34, 138, 276 LQliCODS S22 2 ee 273, 276
MANAUS. 8 = cote aa hector ees 138, 177 leploplend 6 22222. one 176
lithofaleo. ===. 25.25.6025. sonst Ack sss 138 MANSON 222 = 2 oe eee 325
loneipennis:..-. <2. e222 toe ats ce 298 MUS CLCODGE se ae ae eee eeesn 214
MACNITOStTIS: = noe ee ee 177, 313 MUGS CU ie rts gee ee 203
PRR RES oe sw ne a re ts 138 ODU ELLOS ees ee hoe ee ee 235
MINUUS= = ooo - Sa este eee wee 237, 240, 297 MOU er@. 2 230 se eee ne 224
MAOVIOS S22 one San s- oa a eee eS 138, 316 LACES ae ae ee eee 343
MISUS 22-5.2-5< 138, 177, 276, 277, 278, 299, 300 Quadrilovdes 2) 28 eee een oe 215
OINBUUS fa25 ceo coee- esos caneeeeacs ses 297, 298 ECU 8 te ee ne oe eee as ae ee 269
palumbarius-..._....---.--- 138, 177, 288, 300 rotundata.._.___..._------------ 195, 196, 198
PAUSGNIS! 22>. = hoe oes eee 297, 298 SDI ER Cs ras oe ae ee ere 254
Pennatuse 22. oes n2 a cccte een 5 299, 300 SDUNUL08 Wass a oe ee oe 288
CTOLTINUS! 2 3522 sc5 2 oo eee Sete Fee cce 138 SOUT a ee 257
WV Gargus <2 2552.2 eee else 144, 299 SUPU MOS ae eee ee eee 164
PULUS) Soe: 32 oo Se etese ss 138, 177, 299 tridenwcht 22 ot eee aeeata 271
PUUMANS 2s eos beeen St echt cts 138, 297, 299 tridentata. 26 5-o eee ese see ee 271
SPOCICS = 3552s oS cen cows 297, 299, 305 PULOSLOM Gar a a ee eee 184, 185
subbuteo: - s--<23e.-¢s54 cssccecs 177, 273, 275 VUE OT se eae ae ee ee ae oe 206
subbuteusSs. == 22 - on tcecst iw ete st 299 | Filaria (Spiroptera) cirrohamata___....------- 266
Swalnsoniic- ~~ 226e2c0 oso ee 207s 200 «OIL ae ance sea wean cen eee eee ences 162
tinnunculus:=-«....-.2.=-<<sss 138,177,183;.216 | Wilarioidea..- 22. -52.-.5...22---5-- 1, 2, 162, 311, 392
tridentatus_<..-.<--.--<ssccchec- Li Tp eler2ed 1) PASTQUILM,) ASCOPS = snnccoscs sn seeactese eee 206
unicinctus._....-.------- 177, 280, 297, 299, 305 | fissispina, Tetrameres... XVI, 334, 335, 336, 337, 338,
RINMOULING A= 22.2 <3 ace aenaweases 297, 299 339, 340, 341, 342, 348-345, 346, 347, 348,
xanthothorax. ....----s-s-sessec0+esee 177 350, 351, 352, 353, 354, 355, 357, 358, 359,
Naiconis, Spiroptera....- 0 Ssacaisecao-sse 280, 317 360.
falconis gavial realis, Spiroptera__._..--------- 317 TrODMOCENCH <2 2a- oem enee coe anews 343
leptopodis, Spiroptera_..---.--------- 317 | fissispinus, Tropisurus.......--...----------- 343
magnirostris, Filaria.......-...------ 313:\ Hlammeaiflammea..- -. .<.-.-s2nc-a05--5-0<- 144
Jalconis-subbuteonis, Dispharagus._.......---- 275 | flammea, Flammea..-_.-..-..---------------- 144
Spiroptera....---2<=-<-2s 275 SErikis = soc cusecos ss 138, 144, 205, 273, 276
falas? SpirOpera.. « <sccnantarkzwracsecnassseo 276 | flavifrons, Melanerpes_-_...--.--------------- 352
farial, Hoterakis. ... .....-c<ccndeses-nece vil, 61,68 || flavipes, Platycichla.........--...---------.- 359
Ornithostrongylus.___.__--- Vi 11; 12, 18=14;15:4|) lorida caerolea._.-.-.-<.....--..----------= 149
fasciata, Aquila...-...<2<- ssyenatetseoecsce-= 188.4 fluviatilis; Barbus:_<.-......222----<-<co5-56 368
Ascaridia. o..-ficceebee « vill, 80, 91-92 POT CA she ct ae ee noeee 368
fasciatus, Nisactus.........-.---- -cnqiesnaud= 138 Podicéps2 =. 2--. ace stanncaceteae 343
Palacormiss:2-- 222.2 2c22. eee toe 176 AN forciparia, (ASCOVits - <2 co- 20 -csnse-sceso--n0 115
Fasciola trachea...-.-.-------------<--ceeccee 34 WCET ORB ied asco eanasae asccone= 108, 131
fasciolata, Crax...--..--<-2-s0--sieeeesS> = 212417 || jorcipata, Ascayvis.....22~---2--2-na--=n=-= 115, 131
Felis catus domesticus... ....-.--..---.------ 304
Subulura..- v1, 106, 111, 115-116, 117, 126
448 INDEX
Page Page
fossils ;Caprimuleus 2s ese 113 )3| "Gallinulachloropus:222:2=-% “e222 scr t ace 20, 139
francolina, Ascaridia_-__....-_.------- Vil, 78, 99; 93:4) ‘gallinula; Scolopaxt=\221 22522253 toes 254
ET ECE T KES es eee eS aor eae G2 3) -gallinulae, Spiroprerd= 2522 faa e kt eee 253
Francolinus adspersus___.-_.-_-.-.---.---- 123,307 | gallopavo, Meleagris. (See Meleagris gallo-
bicalcaratus-__--_ _--_-- 59, 92, 111, 130, 227 pavo.)
Une ES 2) See ee ithe ee 66: |. Galloperdix'spadiceat:! 22852222 =s2ce cee ene 66, 116
SPOClGS: 2 2 eee yeas 134 | ‘gallopavonis; Ascaris: 2222222222 5220 ieee eee = 81
francolinus; Sephinas 22222. Sse 130 | galloperdicis, Subulura_-_-___-____- vit, 107, 116, 117
frugilegus, Corvus_-_------------ 34, 218, 220, 330,355 | Galloperdix spadicea__----_.__._---_-------- 66, 116
Fulica atra__._ 20, 24, 25, 28, 334, 343, 346, 376, 382,383 | Gallus gallus___-.-..-..-__-_----------------- 10,
leulcopterazeese eae 148 34, 53, 56, 59, 70, 71, 78, 79, 81, 84, 88, 92,
fulicae, “Amidostomum::2222o ae 20, 25 93, 100, 111, 112, 128, 130, 197, 203, 204, 226,
Spiroptera=s2 2S ey Nine tes ks 25 238, 239, 265, 266, 276, 290, 304, 306, 325,
Kulignla eristatars. toss ees eee 24 328, 337, 341.
fitipnl ats ee eee 20 latayettiitt SS ee eee 70
fusca eee See eee ee 24 | gallus, Gallus. (See Gallus gallus.)
TYVERD LL yee ee ene hn ete oD 20 IPHasianUs22 si aeaees 34, 53, 81, 226, 239, 306
mollissima S\ soso oe aS ee 24°) ‘Gammarus pulexe 222222 oss eee 343, 345
Nigra eS eee eae 24597 4. Gampsonyx' swainsoni= 2! 20s eee 297
fuligula, Anas ooo foe oie eee Pe eee 24298) Ganguteterakiss see ee eee 50
I TISOr teeter A antler Soran ce Maen eee 20 PIOPhidesss 5023 69
FF idlie ula ee nese Retee ola 29 | Garrulus glandarius---.._....-------------- 220
INyroca. ees peer erent 20, 24 glandarius cervicalis_--____-------- 241
Fulmarus glacialis______......-.---.------- 265,270 | Sarrulus, Coracias_--_...------.----------- 190, 220
Pulytis GV DS esa a Ter sia 13g | Barzetta, Ardea_____.------------.----------- 83
LLC t eee ah a er tetera BO el 138 | gastrophila, Spirura_---__---_------=--+-252 165, 166
Fundulus diaphanus_._.....__.-_-_-------- 371,372 | Gecinus viridis. .........-.------------------ 215
funebris, Thamnophilus....______.__--___--- 207 | Semina, Physaloptera-___----.---- XIV, 297, 304-305
TUTCAtUS: Walcos Neue te tree ea. 207 || “Geopelia species-+ 1 224+22222:2-==-2 72S 94
Fusaria depressac 2222.22.22 282222 137 | Geotrygon montana-.-_--_-------------------- 95
ENFICUUMELE a oe RAR En oa 139 | Geranospizias caerulescens_---------------- 297, 317
ATU LEL ON Wee PSS ARTE eeue Si | hotbesa, sAscarisis 2522252 ae ken 2s a eee 81
ETT LER EST aed Oa iene eee S 141 | gigas, Tetrameres----------------- XVI, 334, 335-336
STEN CLES owe as ee a ee Ci 143 | Gilsonia-. ----2---222---+----=2222------=--=- 187
ETN CUL Ee EIS Pe ee ee 80 INETMIS Hades 2 ee 188
HLISCAS TAMAS = oo ots tee SE ne 27, 28,267 | glacialis, Amas__--..----------------------- 267, 372
(Agi sen eee ae eae 20 Kulmarus{4-222 2-22 265, 270
eas rar ah Ear eee ee ee 24 Harelda.22.<-~-~ 22 so SE 267, 372
OL GET yee eee om eee 24,267 | glandarius, Corvus. ----------------------- 218, 220
TISCLIStLarUS eee hr aces 43, 148, 235 Garrulus: <= <s2s-ces5 eo 220
Pelecantishes. te ae eee 148 | glandarius cervicalis, Garrulus--_------------ 241
Ptilopachys222 62 Ss See 914.) ‘Glareola austriaca: 22-22 2=--- eee 285, 288
TT OE ATAUIS ee ee ot RR Sale ee 235 | glareolae austriacae, Spiroptera.__.----------- ares.
fusicollis, Phalacrocorax- --_.---------------- 148 | Glaucionetta clangula---_-------------------- 153
fusiformis, Physaloptera__------------ xiv, 296, 303 | glaucus, Larus------------------------------ 234
gabar, Nelieraxs 2 ee eas 304 | globosa, Tetrameres--.-------------------- 334, 346
Galaxias scribes wea ae eee cee 368, 371 Tropidocerca 232 ssaasee at anee 346
Palbula, OviOlUses oo 26 ote eee rn eee D0) || glottissTotanus: |. 2-25-22 eee 285
galeatus, @aswariuUss.- 90-8) 2oee neo meee 41,44 | Gnathostoma--.--.--------------------- 262, 364, 365
galinieri, Physaloptera___ xiv, 297, 300, 302, 303-304 accipitri-2 =... See XVI, 364-365
galli;/Asenri@ias. cl --. soos et oe eee eee VU, pelecani=4--2-:-.2 2 XVI, 365-366
49, 54,79, 80, 81-82, 83, 84, 85, 86, 87, 88, 89, shipleyi 2) covet. 2a Sales 263
90, 91, 92, 93, 94, 95, 96, 97, 98, 99, 100, 101, spinigertimmt) e. Ss. seen = 364
102, 108, 110, 111, 112, 113, 114, 115, 116, | Gnathostomidae-.--------------------- 163, 363-364
117, 118, 119, 121, 122, 123, 124, 125, 126, | goliath, Ardea__..--------------------------- 369
128, 129, 131, 133, 134, 135. Gongylonema- 2223 se --- ------ 208, 390
SA SCOTIS CED Re ee eee 81 ingluvicola.--------- xu, 204-205, 390
gallicus,'Circsetuss 222 ase e aa eee 138, 298 MINIMUM ~<252 ses seee ee ee 203
WATCH ieee ee ee eer 138, 297, 299, 300 musculi.---2t2 22S SS 203
GallifOrmies ee ene eee 35 scutatum..225-45-S- eee 205
Guilinde; AsCariSce eee sen oe en nee == 52 species new? .----------------- 390
eterakis!. = eae sone VI, VU, Gongyloneminae. -_.---------------------- 163, 208
50, 2-54, 55, 56, 57, 58, 59, 60, 61, 62, 68, “GQ 00S6 Vip Ree eee eee eee 93
64, 65, 66, 67, 69, 70, 71, 72, 74, 75,77, 104 | Gordiacea----------------------------------- 391
gallinago paraguaiae_--_-_--.-._--------------- 342 | govinda, Milvus-.-------------------------- 137
gallinarum, Hilaria 22-22 20h nee 197,198 | gracile, Orysoma_-_-.-------------------------- 134
Hartertia___ x1, 191, 193, 195, 196, 197-198
gracilis, Acuaria---_--.---------- XU, 217, 218, 222-223
INDEX 449
Page Page
Pracilis, Mispharagus.....-....=86 2 Se 222 | haliaeetus albicilla..............-_--._-_____- 138
Maleo: s.2scissseccisceteel aes. 298, 317 leucocephalus...........=-:--.-22. 150
Histiocephalus.« 22.2222. 520th 286 | haliaeti, Contracaecum____________- XI, 146, 150-161
Subulura..-<--3--..-. 2" Ix, 106, 134-135 | Haliaetus leucogaster___._.......___________- 150
SynPamus= ==222-222 2222204 VI, 34, 35, 39-41 | haliaetus, Falco__.....-----_--2---2- 2-2. 137, 207
graculus, Carbo. ..=-.-..-.. 24202 Uae. 147 Pandion ss 20.. =5. 29a 2 Soe 137
Phalacrocorax..52: 22220 S 232... 148 | Halieus brasiliensis__...._.._...__--__________ 148
#raecs; Perdix::++<s<sscscseeseec Rt Weccce io) alli, Subulura..... 22.225. ee viul, 107, 117
grammicus, Picts 2252222222252 522 sooo 179] sHamennia. 225. ...2222.2.22 502 Oe 244
grandis; Nyctibius..2--<225222. 5b 2%... 129, 130 Calcarate. 22 se ee 249
granulosa, Ascaridia__._........_- Vu, 78, 92-93, 94 CONT ~ 2 ss 55 se 249
Heterakiss>. 2222220. 92 longeornata..2 222222252520 ss 256
prayli, Ardeola...22s222522-.-.- 24 Rsk 152 phoenicoptert.....-- S38. 2... 257
prises; INyctiardes==.22=2-2s60555-2 Et. 152 squamata.. =... 222 Niece 258
prises ta, Stik. .2 5202-3204 25.-5 ee 205 WNC = oc ss Se 246
griseus, N ycticorax.<--2222-/2.2221L0 141, LO8i281 Wi hamata, Acuaria = 2-22 osesec2l sheen 278
Grossiptodon manschuricum_.--__.___..--_.-- 53 Pilantae.- as523 55222, See Se 278
Grus/antipone:=<222..cccosses5 PE 89, 99 Synhimantus--._........._ XIV, 273, 278-279
australasiana.2---=-..5-22e2 see: 142 | hamatus, Dispharagus_....--......._.------- 278
clnereds—<555--26-5ce5eso8 Hed. 142. 2863 |"hamia, Ascaridta 22222222224 2 ee 93, 94
COMMUNIS. 5 S225-2222 ees ee 99,143 | hamulosa, Acuaria__...........-..--..------- 226
PSTAGISOS 32-255 SPS a oo 99 Cheilospirura: ....-2..2222522 XII, 226-227
AVONING 222472 neon. soe Fo 143 Spiroptera. <220 522 ee 226
WiridirostriS:=2.<ssscosce te SS tee 44 | hamulosus, Dispharagus_-._-_._.______-____- 226
PTTIS WANOCA = Soe oan se EE 142 | hamulus, Heterakis___.........___- vu, 52, 63-64, 65
OruUveli: *ACUATIGs=. 22-2 5e soe eee 227 |) Harelda glacialis_.........22.-..2 220 .. 267, 372
Acuaria ( Cheilospirura) _........-.--- 227 | hargilae, Acuaria (Echinuria)....._________-- 253
gruveli, Cheilospirura--_.----.. XU, 226, 227-228, 231 Echinuria___.-...---- XIN, 245, 252, 253, 254
Dispharagus=-222.. Fe 227 | Harpagus bidentatus___._._.........__....- 177
erylle; Uria--22:-2sccccsccc eRe. 148, 235, 266 | Harpyhaliaetus coronatus__.._____.________- 297
guarauna, Tis. -..55. 22.2 249 ;| Harpyia, Thrasactus....2-. 2222222222... 317
guatemalae, Chrysotis._.......-.........---- 175 | Hartertia_. 164, 191, 192, 193, 195, 196, 198, 200, 206, 393
guianensis, Caprimulgus__.__......____-_- 129, 207 annulata. 22a XI, 191, 198-195
Guiry guiracc. ss22s5--222-5222- sibs 115 CONfUSHE <=. = 23 ee XI, 191, 195-197
gnira>Guirassess2.2c0<sc2Lbeie ose Secs 115 gallinanims 235258 2sceeeeee eee x,
gularis,"Francolinus.- -222222222522-0........ 66 191, 193, 195, 196, 197-198, 207
gutturosa, Columba:=.<.-<2.22222222 022.0522 87 obesa2=.-2= = XI, 191, 192-1938, 196, 206
gynaecophila, Tetrameres_-_-__-___- XVI, 334, 347-348 rotundata=..-....=... XI, 191, 198-199, 200
Fropidocerca__..2 2202-22222. 347 zakharowi._.....-.---.-.2-22-2'--- 191, 200
gynecophila, Tropidocerca__..........-----.-- 347 ZOTWIES - ee. oa Soe XI, 191, 200-201
Gypactus barbatus-==--..2s-s2<-t222b2 2... 138 | hastata, Heterakis_.....................-.4.. 385
G@yps fulvUs 25222222226 cssecccsessseee_ wk 138 | hastatus, Ornithostrongylus_____.__.______ V, 12, 17
Ma bronemss 2. 22s es oo 52 cess Eke 164, SU ONGULUS:. 222. - 2 ee ie eee 17
171, 172, 173, 174, 175, 177, 178, 179, 180° | hawaiiensis, Megalopterus_......._-.._-.---- 161
182, 183, 184, 186, 196. MAW ten soca k release Bae Ges Waters eee 301
colaptes'=. 22.22 2282 st 2s 78-174 3) helias: Hurypy ease: #222 2-20 os ese 362
fichouri..22 252222223 MR, 172,.174-176 ‘|| Heligmosomidae.s..-..<..2=.5 ili cece 6, 7,8
Mcertars2- 2. bss LS. XI, 173, 175-176 | Heligmosominae.-.............-------------- 8
leptoptera...-- 2222 22 Xi, 173, 176-178 || Heligmosomum=....... 22-2. 24 2S Sai scce 8
longistriata=--.-220=- XI, 173,.178, 179 || Heliotreptus'anomalus:- =. -.2.-- -282222_.-_- 130
706029 (0 fe feeder eh Ai be rrr 162 | helix, Microtetrameres__..__---.-- XVI, 352, 355, 356
mansionl 5202 6223es45:- Xu,173;.179-180 ‘|; “Helodrilus' parvus: = -22-2.--2t't2- 2222. 82
monoptera._.-----. XI, 173, 180-182, 183 | Helotarsus albicilla_...........--.----------- 137
TRUSCAG Soars aa ees 2 oe cn 172 OCA GAUS ss rest eens ee 137
FOCUNAALE 2 Fos BAe OPI SOE 195,198 | helvetica, Squatarola_-_-_-_..------ 139, 144, 285, 286
SULA tiers Aes Fen XI, 172, 173, 182-183 IE TIN Rian oases eee eee oe 285, 286
Sspinosas242 east XI, 173, 183-184 | henryi, Amidostomum..---.....---------- Vv, 20, 22
tulostoma_______._- XI, 173, 180, 184-186 | hepatica, Hepaticola_...--....-.------------ 377
unilateralis____ x1, 172, 184, 185, 186-187 | Hepaticola hepatica__-......-.-------------- 377
TIBU JCA >. ss sess sores cas 164, 187, 188, 189,190 | hermaphrodita, Ascaridia___-_-.-..-- Vu, 77, 79, 80, 81
INGLIS 2 ee ee XI, 187, 188-189 PACCOTIS= ooo tacos eer eases 80
Lhuillier? t+ sa<r2 5-5 Sees Ni fo7-188 |) “Hlerodiasegrettacsas22052 toot ei sceceea-s 152
DAL VAs ear et ss sane XI, 187, 189-190 THICDGON aces see naaenesceeseseeees 152
truncata asso eee XI, 187, 190-191 herodisswArdea. 25-22-2252 52 sc<-- 141, 152, 248, 372
Haematopus ostralegus________-____--- 148° 285s287, |! "Eléron’cendré...2.2_--.2...--22 222022222. 209
haemochrous, Tetrameres- ----.--.-------- 335, 357 MOULD Os seas. Ue we ccuk cea aoe 279
Halcyon chelicutensis_............-....-...- 188 | Herpetotheres cachinnans--.--.....------- 177, 297
450 INDEX
Page Page
bleteracidae ss. nas a a oe 49 | Heterakis papillosa_-.-.______--_.. VU, 52, 53, 67-69
WFTELET CIS. ooo oko S| el at a euce tS oo 50 parisiis= 5322 le oe ae vu, 50, 69
Meterakidae <2 oS. seas 49, 75, 104, 363, 391 DErspicilm 22" et eee 81
Meterakinae:. 220.2525. 0- <a 49-50, 75, 77, 102 psophiae:s- = 24: els ee. Vil, 52, 69-70
Heterakis 22. o2j5.2 ono 1, pusilla 22223 alee ae vu, 51, 70, 71
49, 50, 52, 55, 56, 57, 58, 59, 60, 62, 63, 64, putaustralis: _ 2. =. >= 22s ee vo, 51, 71
65, 67, 68, 69, 70, 71, 72, 73, 74, 75, 77, 104, FECUTUOIG... =e eteeeee a heey 124
385, 386, 387, 393. FUNG S hokin kL ke S 125
acuticandas 222s. tees VU, 52, 76, 77 rimula=—o-_ 2 =.) =) ee 125
aculissima.. << =< — 2 Ae S 107 Senrate. 2. 2 ee ee 98
aegyptiaca...\._.-_- eee 83 similig.22- ee ae 127
alata 2-4 2.222 eh roteet VI, 52, 55, 66 skrjabint:2 - 222 2 sys3-tucs vu, 52, 71-72, 73
allodapa....... - saetn2 eee sess on 108 Spiculatus 5-2 <3 ue ee ee 128
amblymoria......-- 2$5-5-22.-~-.- 83 StYOMMs= 7 2< 22 <= - ae e 99
arquata- 2° =) 2 2c vi, 51, 55, 56, 71, 72 Slglosa. 2) 228 oa i oe 67, 68
GUstt lie... 2-82 = steete - 84 styphlocercas.5=<.- 2+ see 100
bancrofti_ 2... --..2- 2. seaste VI, 50, 66 subtleta.-. 3.22) -doap se ee 128
beramporia. 2.0 - - 245-te VI, 52, 56-57 StLCLON Ee wae) ed oe 108, 112, 130
ousasse ==... 2S eee vu, 51, 75, 76 tenuicandas) 290 =e Vil, 51, 78-74
DOV eG seek g net os eee Bete 84 trilabium._ = 42 22-2 eee 101
Ibosia: -22 2222.2. eens VI, 51, 57-58 truncata? 2: 22. so eee 80
brasiliana.....< s2sege-2e5 52 Vu, 52, 58-59 valdemucronata__.-____------_- 50, 74-75
brasiliensis... =... - -wereen cee US 84 Wal vataets 542s ere. Saag vu, 50,74
brevispiculum_-...-..------ Vu, 52, 59, 60 Variabilis ec o22- 2k bees 886-387
ealcarate.. <2 2222s. eet Ot 96 pesicularige. yee ee 50, 52, 67
cathelurinus:. 222.2 paso 85 Vulvolabitac# 220-55. 0 2 ees oe 887
caudate. << (tyes vu, 52, 69-60, 61 | heteriira, Ascaris_.......--------------------- 144
chenonettae-._....--------- vu, 50, 60-61 | Heterocheilinae...-.....-.--------2+----2---- 135
circularis—_..-- 250 ao kee ee 86 | heteroclita, Oxyspirura............-.-_---- 322, $31
circumvallata....-.=-..+.----= vu, 52, 61 Sniroptéra. 8 oS eee 331
COULTUG Es ae acs meee ee 86 | Heterospizias meridionalis_________.._.._.--- 297
COMPON= = ans. se 25-288 Sis- 0 ote 88 | heteroura, Porrocaecum-.-----.---- 1X, 136, 144-145
COMPTESSA.-- 8 = =e tees <5 = 88 MAS COTIE =.=. nto ta. als 144
CONGGLG Son nn noe ene na a= 88 | hiaticula, Charadrius._.........-----.----- 139, 254
Cristatae: 322222. <2 ee team 89 | Himantopus himantopus-_--.-..-------- 22, 232, 348
CUmuGl@n ane tes oases see 114 melanopterus---_-------- 139, 144, 232
cylindiic@a< = 2. 2-=--5-2asacnss=<e 90 | himantopus, Charadrius___.....-....-------- 232
divierens...- = 2. -- - eh eee ee se 111 Himantopus-s-cons=28~ 254 22, 232, 348
dispare Win. oes i26 Bes vu, 52, 61,62 | hirghisensis, Ascaris__-_---.------------------ 145
dolichocerca.. 25 --. -= Sa cee 90)}1| SHirundo riparia’ 20-0... cseaok eee eee 220
pag ORIOL ste eS ee vil, 51, 63 rustica $2 oS i 220
Worcipanig: <2 ac ook ee 108, 131 SPOClESE S. 2.5 = 25 oho ose ae 217
WRONCOUNG = 2222s ae 92 urbicaht) . Asner te ee 220
gallinse:. 22-2 oe te Ni, Vil,;|: hirundo; Stemmas 244 (242. 285. 222s seas 287
50, 52-54, 55, 56, 57, 58, 59, 60, 61, 62, Histiocephalus tread 233, 244, 272, 284, 290, 391, 392
63, 64, 65, 66, 67, 69, 70, 71, 72, 74, 75, bicuspis: {-- 2. ade saeco 286
77, 104. brevicaudatus_._.--.---------- 274
GTanwlos@ oso ae ee 92 COFONALL Se. ess. Saat ee 293
hamulus:es2220 So eee Vu, 52, 638-64, 65 decoruse aoe eee 269
hastates--2-- eee ee ee eee 386 OTOCHIS steers eae 286
ANALG soo ooo eee a a 81 LOCINNAUS oo oe eee 288
interlabiatas: 02 2202. wk vi, 51, 64, 65 laticaudatus_-_-..----- XIV, 290-291
isolonche.2e2 202 seca vu, 51, 64-65, 66 BDU CUS epee noee 235
Janel 222-2 ode os a 385, 886 tridens 2202s eek XIV, 291-292
tatCGUad..- <5 2 ee eS 97,98 | Hodotermes pretoriensis__---.-.------------- 197
LOPriNnces. 22 es ee ee ee, Le 118 | horrida, Echinuria_____-._--.---.- XIN, 245, 253-254
Unegta. 3. .5- 5-2 oe ere 93 SD Open Sse asap eheeeaceses 254
longecaudata________- vu, 51, 66-67, 68,95 | horridus, Acanthophorus____------------------ 343
longecirrate...2.-.- S 94 StrongylUssses a3 oan soso ws Sane 253
MACTOUFA:. <2 el eS vu, 52, 67,68 | Houbara undulata.__._._..........-.-- 130, 196, 198
MOCULOSO: ooo Se ee ay eee 86: | choubarasOtiss.2-e 2 seates 130, 195, 196
ANOGNUDODIUG So oo ct de ot 95' || shumeralis;;Penelope---...=.--.2-e2ete eee 79, 98
monticelliana::. 5. ..5- 5552-8. 53,67, (68, || -huppéy Cariama—. 2_- 2 = ase eee 108
mattererc <2) cnc. Son so ee 51, 75-77 | Hydrochelidon nigra--......-...--..------<- 259
neoplasties =... soe aen vu, 51, 64,65-66 | Hydropsalis climacocerca--_----.------------ 130
MUMAGGE 2 oes ke 96: || -hydrus, ‘Tropidonotus._ 2. ee 168
Orthocerca ==. os oe see ad ais 97. ||: Eeylotomus:pileatus.._-..-- -=--- 355 325-2 215
INDEX 451
Page Page
hypolencs, A.ctitis+...2s2c-s2e 25220222 235 | javanicus, Phalacrocorax__.............----- 148
bypoleucus,-Totanus... 2-222) 22202-2001 235. |; jubata, Chenonetta.....-...........-.----..- 60
Tringoides... - Wii" aie fei 21 235 | jugadornata, Echinuria___--..__-_- XIN, 244, 245-246
Hypotriorchis subbuteo_________-_.--------- 172 ( Jamana, Pious. .2252<2s02sc<seseccs st ths 178
‘Heystrichis=-.52<>=-= 366, $75,376,377, 378, 379,381 | Kathlaniinee....<...<..<s- 2-2 -- seit 2ei ioe 104
acanthocephalicus_ xvu, 366, 376, 377-878 | Kathleena_.........------------------_------- 146
coronatuS---......-- XVII, 376, 878, 379 arcuate cu) 2c s5255 SEES 152, 153
Cygnl..2 =. s<s2<<5 XV, 376, 378-379, 380 DUNC. ook ose I 157
CLEDANS oo ee oe 370, 372 TOGNGING = 2sscsse ose a eset t a 157
mergi-merganseris_._...-...------- 378 Scotti... o-2 .- . ssecnsteoeeeere ie. 159
neglects... 22.22 XVU, 376, 379-381 UriCUSPIS. 2225-2 ssccuBe eaes ee ceae 160
OTSPINUS =s52555-4-<6 XVI, 376, 381, 882 | kirghisensis, Ascaris..............-...------ 136, 145
pachicephatusx20 2822220. Sees 2 378 Porrocaecum_.--------- X, 136, 145-146
papillosus ___._- 358, 370, 371, 373, 374, 379 | korschum, Milvus_...-.............-_-_--- 179, 180
pecies <5 55. Te 378,379, 981' | kuhli, Puffinus: ....-2222-...22. 222802. ..- 235, 263
tricolor.__ xvul, 375, 376-877, 378, 379, 381 | laciniatus, Histiocephalus__-........-------.- 283
Cub fee ee 367 Schistorophus__...._....._----- 285, 288
varispinosus___..._... XVII, 376, $81, 383 | laevis, Porcellio_....-.....-.-..----.------- 238, 239
wedli___..__.. KV} 376,'379)' 380; $81-S8s: | lafayettii, Gallus__...-........2.22.322ssc. 70
wedi 22-22. 2 SIS 379:38i" | tagopodis, Ascoris.....-.. 222025 fo eee 88
Ibis aethiopics.-- sess te ae. ce 297;'309: |; Lagopus mutus...:....--.-2224225...2.22ch<8 53, 84
falcinellus-..--.--.- 22. oe 250, 376, 381 SCOUICUS! <2 552252 c oe eee ne aae 9, 34, 53
Rasraunas =. 23.3 22.2 so ee 249 | lagopus, Archibuteo (Buteo)___.._.-______- 137, 138
MUGHLONS asec en Hee ese 377 Buteo. 2052-5 ee Se 276
TUBS. 222 css Seok SIL ee 261 Wal@0sscccsc ose cese sibs ees 34, 138, 276
PUL O Te Ne are cee ee 377 TOUra0.oc5- cas sec ace eho eeee ee 53, 84, 88
ibis; /Ardeola:. 2: 2225-523. eo AS. ot 279 | Lampronessa spomsa___---------------------- 59
Pseudotantalus._...... 2282-24. cee Pots); lanarius, Malco. <.22.-2-2-5.2seseceetaose 138, 177
Teteridae.. 25.20 Let ee A Leak oetee 5 330 | lanceolata, Spiroptera_...---...---------.---- 171
Feterusicristatus.-.--2-. Sse ei Fe... 207 |" lanei;Heterakis 22. -.22s.-s<252.82223.25 385, 386
croconotus_...... 22 2 eee. $23. |: Eanius'collurio_...:-..---~2-...-35.4 201, 220, 221
igneous; Falcinellus.....222002¢seen Sc tee sees 250 excubitor...<...) oS ee 201
ignotus, Eustrongylides__.. xvu, 367, 368, 371-372 excubitor dodsoni___.__.__..___-----_- 203
iHacus: -Turdus=s-. 23. <2. sc SSMS. 22 139, 206 THING? oes eee ee eee eee 200, 201, 229, 296, 307
imperialis, Aquila. ._........-.---- 138, 145, 298, 364 TUfUS. 20s 282054 2220555 SES 201, 220, 221
Malco. 525 Sascha ats 138 SDOCIOS. 222 5. 2 se onc soca Aces ese 355
impeyanus, Lophophorus__-__-._.-----.------ 64,66 | laplantei, Acuaria (Dispharynz)_--- XU, 237, 241, 242
incerta, Habronema--____--_-_-__- XI, 173, 175, 176 Dispharynx.--32..22028" XIU, 237, 241, 242
SpiroplerG ccc. 2255.2 2. SS 175: | lari, Cyathostoma-.--.-4..--+.-2-- VI, 41, 42, 43-44
INCIsAsVA SCAMS. 52.52 on oe SS 137 SclerostoOma _- 3-252 = seh sss SS 43
smevniss GilsOniG.. - oo een sks 188 SYNGAMUS._.2. cae FA esa lel wah seets 43
Hadjalig::.2225-2..22 5-8 XI, 187, 188-189 | Larus argentatus_-_-.--.-_.-----.------ 148, 234, 235
Microtetrameres ------ XVI, 355-357, 858, 361 argentoides!..2—.:.-ssti2s2ei22t See 235
ENGINE C8 5222 eee sos S asec 349, 355 CSNUS Ss eo cece tees 148, 234, 235, 263
Propidocerca-2s--<.- 3 en 355 CADIStranus!<2.5-cc2- 3-2 233
inflata, Microtetrameres-__-----.---- 335, 351, 367-358 fUSscuS: 2 Lacs. secs Beas 43, 148, 235
Physaloptera..<52.. 252. 2et8s22e se 297, 305 glaticus:2.02...£52.5-8¢ 22 RON W.- 234
Spiroplera’..cs..ss..52c8teh eee 305, 335, 357 marinus# 2. 22.2253. ss 148, 234, 235
Tetramenes 2 = -.sccest test 357 MAXIMUS. .__.-.5- 2255. S. Setzb.a Biases 235
TOM UMOCET CR 22255525282. 2aSi 5s 335, 357, 358 medins!. 2. o=-55 05. 3a Ss ae 234, 235
taflaties Tropisurus 2-22. 2222 kook w22 Ss 357 ridibundus_____-_- 42, 43, 148, 234, 235, 265, 271
infleza, Ascaridiag=—=--..22- Ses sls... 81 species... <...-- pI ARGID SL Sees 43
ASCOris. 82s ORT et 81 tridactylus.-2 -2ss-Re beet oe 148
userid. sooo. asses a sesec ce Si |) lathami: Catheturtis....-- fo. 520cu-cit4e...-- 56, 85
Heterakis:. 2. Seataeeos } Lu S285 81 Talegallus...22.-.-..-222:6-2 ost 85
infuscatus;‘Phimosis._.......ssosessuieltesus 377 | laticauda, Ascoris_--.-...--.-----------+----. 97
ingluvicola, Gongylonema__-_-__..-.-.--- XU, 204-205 Heterakis.....2.-.scs0veisii...-. 97, 98
interlabiata, Heterakis_...-.....-.-._- Vil, 51, 64,66 | laticaudeta, Filaria_.........---.--.---------- 290
invaginata, Acuaria (Synhimantus) ----------- 279 Spiroptera.._ssssureseaheeeelas 2 290
Synhimantus-_--------. XIV, 273, 279-280 | laticaudatus, Dispharagus_-_-.---.------------ 290
invaginatus, Dispharagus_....-.------------ 218, 279 Symbranchus-.--.-.-..-.------- 368, 371
involutus, Dispharaguss:_22.---2222-2-222.-. 276 Histiocephalus-_-....-----. XIV, 290-291
Irrisor ery throrhynchus....-:..-..-2-.22221:. FSS | |taticenes Acuaria......2-..--------.--=20% 273, 276
isolonche, Heterakis_........-.-.--- vul, 51, 64-665, 66 Acuaria (Synhimantus)-_...---------- 276
isda, Alcedo_2- 25/22 22-<222- 3 = ee ee 269 Dispharagtiss.:. 2 .cc22s525-2522ee8 276
htiayenes'crucntus.. . 2222+ .22..<2 299-1 2 64, 81 Pilavids.<cceccocndccoos- Stel eles 273, 276
jamaicensis, Nyctibius_.._.........---------- 130 Spiropleas-o6sc0.------5 eet. 276
452 INDEX
Page Page
laticeps, Synhimantus----.--------------- RIVA272;)|t Ineius; sox. 220 25 2s see ee es 368
273, 274, 276-277, 278, 282 | lumbricoides, Ascaris........._---=-------.-- 82
lavaillanti:;Plotus: = 2.2222 22ssses0 Sess 148 | Lurocalis semitorquatus -___._..-_-_-.------ 130
Jedehii Dacelo-=25.-25-..5455424520 ss oete 314) |) Tauscinia philomela. <2 2+ =: 2oee hee 139
leprincet, Allodapa.-=>-<=2=--2<5225 522 sestie 118 ru bectilan: sss: 540 nnn sue Eee 224
eter akie. &- 02s soso eee 118°"|' lutzi;Subuluras 232-62 Ses asee 1X, 105, 118-119, 120
Subulura.--=-=-2° 1X, 106, 111, 118, 119, 126 PB elazigeseee Sos. a bs oi ee XV, 312, 315-316
Leptodon cayennensis_--..-.---------------- 297 |) luzonica, Phlogoenas=- <=. = 2-222) sabato 8 87
leptoptera, Filaria 2 +=. => sot ieeees 176; thlybicayZorillaves <8 5 na koe oe eee a ie 200
Habronemess2 4-282 22228 x5173; 176-178 ||) Tayrurus tetrix_/ 2.2 = 23g eee eee 17, 95
Spirdpterde ce = 2 eee es 176 | macqueeni; ‘Houbara.-=- += -- 2 sa-hotee ss 2 ee 130
leptoptili, Acuaria (Echinuria) -_--_--._------ 253 Otitis seleee scaae 196, 198
Bchinuria-= 2 ase XM, 245, 254, 255, 256 | macrodipterus, Macrodipteryx-_----.-------- 118
Leptoptilus crumenifer--____.-..__-------- 254,369 | Macrodipteryx macrodipterus---___-_.------ 118
GU DIUS Sos Se = eee en EE 253; |) macrolaima; “A cuaria =<. 2-2 -- == Seema aes 212, 213
eptosoma. oo ee iene abet 2a 295 | macrolaimus, Dispharagus-_--.--..------------ 213
eptotila rufaxillas 22): 2 ~ ep inespitee tt? 13 | Macropygia nigrirostris_.__.......-2__...---- 78, 84
Lestris parasitica =-=ssee= eee Le gee et 148 ||) macror; Capitosss is. - =<. 4 92a eee 115
PoMarinus 292.5 5. ee Sees 148; || macrorhynchus;)Buccosss----- = Seese2=2= 131
leucoc., Pionus (Psittacus)__.._-..2-+-22+---- 80 | macroura, Heterakis............------- VI, 52, 67, 68
leucocephalus, Chrysotis_-....-..----------- 75y|) macularia, “Actitiss: {= .22 2-2 © hao eek eea 235
HYalineetus sas 2 ieee 2 150: |) maculata; Aquila: 30220 eke ts. = eet aes 316
leucodon, Crocidura.-c22tett es beeen 2067}, maculatus, Totanus: + -~.2) 2-0) eee 235
leucogaster, Haliaetus_ =... -.-.-- 282-2 150)'|)) maculosa; Ascaris: to-8- xo eee 86
1evCOpS; Amast 22. (ce Le aes Sails ts eee Se 28, 62 Teter Chis Sh ose e ee ee 86
PATS SC ik Sn NE SS Copies oe 20 Notre ete iictuue is, Ses ee 121
Moonasaei 22 22 cee aus pueteiedn See 115;028% |); maculosus: ‘Circus: 6220.20 022 250. 297
léucoptéra,CMmlica 25.) seus Saiki Vic eae erste 148 | magalhaesi, Ascaridia._.._--.------- Vill, 79, 96, 96
leucopygeus, Caprimulgus-_-__-__--_---------- 207 | magnilabiata, Acuaria ( Cheilospirura) --__---- 241
lemcorodia, Plataleazss nso eee ees 339, 369 Cheilospirura-.-->= 2) - Ae 241
legeotisy (Conpris = oe es are pene 175 Dispharynx_22-=2 = XU, 236, 241-242
Pionus) (Psittacus)_- =. 22: seni ees 80 | magnilabiatus, Dispharagus ----.------------- 241
Scops:. ooo 2 A tet ek pega 127 | magnipapilla, Ascaridia__________---- VII, 78, 95-96
leucumis #Blanus ieee soe en eee age 297 eterakis.-: 2-28. 2.» sapeegeetey 95
[hpi lieri phadielia.ce*- 24-2222 ses oe x1, 187-188 | magnipapillatum, Contracaecum_-_-__ X, 147, 161-162
lineata; Ascaridia. 222-2222). VII, 79,'80;82, 98-94 |. magnirostris, Falco -.-..-.------..2232%-- 177, 313
ieterakis 52 sce sends tyes Se 93 Rupornis::..s:2e2e42nie tees 313
linéolasBolborhynchus:-.~.-..22sset sess | 175 |) maguari, Ciconia-22222— -2 222-2 50, 74, 168, 245, 256
lithofaleo, aleo-- 2200-23-32 5 ee ees 138 Buxentra-- 2222-28262 -5 ee eee 50, 74
livia, Columba. 2-2-8 =. S-2- = eet 87, 239 | mahali. Plocepasser__....---.-.-------+-s2: 192, 198
livia domestica, Columba-------------- 12341, 343 |: major, Ardeas::=..24--. 22. = 2222s Saeeee 142
loculators@antalus- 22-268 se Sater 154 Crotophaga.:...----=-2--.-=288 124, 171, 323
longicaudata, Heterakis__-______- vu, 51, 66-67, 68, 95 Megaquiscalus.....-sseenses2 ate et 318
longecirrata, Ascaridia_..___._.._._._- Vill, 78, 94-95 Pics: Ue Ses od Cee Se 34
longeornata, Acwaria-=.--...-.2-. Apes 255 Quiscalus... 2-22-2222 2.2 318
Kichinynia ss Sse ees 245, 255-256 Scolopax®.< .22°>.---eesspeeeiasinly® 261
Hamannia= = 2222-3 eos 256 "| ‘majusabronema 2222222 == see eects 162
longeornatus, Dispharagus_._____------------- 255 | Malacoptila torquata..-..----...-2 2222223 128, 131
longevaginata, Acuaria_-___..-----.--------=: 256 | maleo, Megacephalon___.....-_-.--.--------- 66
Hchinuria! =". Secewew les xi, 245, 256 | malleus, Physaloptera_.-...-.----- XIV, 297, 308-309
Synhimantus.—2 2 eee 256 |) mamillaris,, Acusria= 22 --—— - see 212, 213
longevaginatus, Dispharagus______-.---------- 256 Dispharnagus = +. >>> seek 213
longicornis, Ancyracanthus..._____..--------- 285 | manchurianum, Crossoptilon.___...--------- 64
Schistorophus- -----.-. XIV, 285-286, 287 | manillensis, Ardea__.-.----.----------------- 141
lonigipennis; ‘Walcoz: = =<2- 225 peat 298 | manschuricum, Grossiptodon_._--.---------- 53
longistriata, Cheilospirura__-_...----------- 178,179 | mansioni, Habronema-------.----- XI, 173, 179-180
* “Habronema..-. 22-22 XI, 173, 178,179 | mansoni, Filaria_.......-...22.2222.22-2----- 325
Spiroptera. 22-22. sta ae 178, 179 Oxyspirura 2: =: eee XV, 322, 825-328
loosi, Pseudamidostomum_-_-_____---------- 883-384 Spiropter ase oei soe > nee ee 325
Lophoceros semifasciatus-_-_..._..--.-------- 188: | | Mareca' penelope = -_-..-.--.ewssnenesees pte 27
Lophophoros impeyanuS- ~__-_--------------- 64,66 | margravi, Dicholophus-_-_----------- 97, 108, 130, 332
Lophortyx californicus-.:2-20a@gesi 5-2: _----- 175 | margravii microdactyli, Spiroptera_._-__..---- 332
ephiire rofasteee ee sss of ees ies 385,386) )marila, Anser. o.oo. 22... po eee es 20
Tote lota si. 22 sheets Brest 2 368 Mulignla ei 2k oe ee ee 20
lot; Lota ete ee eee > 368 INGE OCS ROO es es ee 20
lucidws) Babuleus: = 22-3222 22sc222 2s See 279.)| trdarinus: Tards 2 8 eo 148, 234, 235
Bubulcus22522 2222-22522 eee 174,339 | martius ,Dryocopus---......----.--.-------- 215
INDEX 453
Page Page
WHATTIUS!, PiCUS. 22 ote 215 | Microtetrameresinermis-_.._._. XVI, 355, 357, $58, 361
mauduyti, Spizaetus_......._.....--..-____- 297 inflata__..___- 335, 351, 867-868, 372
mauritanica, Tarentola__...........-...._-- - 216 minima_._.__-.-- XVI, 352, 358-359
maximus, BUDO...-2----)-- sc. tee 138, 144, 276 pusilla...=-..-<..-.<. XVI, 352, 859
NAYUS 2S ee en Se Rs 235 spiralis_........_- XVI, 352, 860-861
mediospiralis, Spiroptera__......_.._------ XIV, 294 | Milvus ater..._...-..- 2-2-2 138
medins: Waris: -. =. 2.<<2.-2<0 sees ets 234, 235 POVINnd Ake) 137
Megaloperdix nigelii_....._._..-..-------_-- e 67 korschum. ....-<-<<cc2csecusce- 2. 179, 180
Megalopterus hawaiiensis_.._.____...._-___- 161 MAUVUS 2-3 we hoc cks ace 138
megalostoma, Physaloptera._. XIV, 296, 298, 300-301 MOS ee ee ee eee 138, 177
Spiroprerd... 2-322 -2- see see 300s |i milvus; Faleo-. 2.02.23. scescseceseneee luce 138
Megaquiscalus major____._....___.-----_---- 318 IMEI US: oe ooo ee 138
melacoryphus, Coccyzus.__..___-.-_--------- 107 | Mimus polyglottos....._._....--.---_----__- 139
Ouculusse2 26853-5222 245 129, 309 | minima, Microtetrameres_______-__ XVI, 352, 858-359
Melanerpes flavifrons_.............---.------ 352: |: minima, Tetrameret....<-.-<<-<<0.ssao2-225-- 358
melanocephala, Ardea____________.__--___--- 142 | minimum, Gongylonema-.-______.._______-_- 203
melanocoryphus, Coceyzus_______-_-_.___-_- 1iheMmMINOL,,-ATdeas2.-22255-5...2255-2--5-Seccce 152
@yPnusies4 sate 343 IB OtHUTUS #222" 25a ee 273
melanogaster, Plotus__..._..__.__.___- 148, 160, 213 @Wocty7s=--222 52 4:20. 25 ee ee 115
Vanellus: <.-.<. 342.50 139, 141, 286 Tans. 2222 pees 200, 201, 229, 296, 307
melanoleucos, Bucco__._..-___....----------- 128 POdIC6DS= = 2322222. i ee 148
melanoleucus, Astur_.................-.----- 309 IPrOGICEDS == 2 ce od coe ee ee 372
CAD IGG 22 es so ee 115;.) minute, Ardetta..-...-.-- 22-62-2502 34-3- 274
Spiziaster_.............-=...:- 297-| minutus, Falco.......=..-...--2s2-<-<- 237, 240, 297
TOtanus=-..2. 22252 cee 2612} mollissima, Anas. -.- -- 5 222 See eee 24, 28, 372
melanophrys, Diomedea__.._._.___._-----_-- 159 SANSCY Uo 35 cc cso eseass eee 20
melanopterus, Himantopus___________- 139, 144, 232 Bualiguila.< 232252 ee 24
melanorhynchus, Cuculus____.._--__-___----- 128 Bomateria= 2-2. eee 24, 372
melanurus;, Trogon. ---:=.-2-<.-=<s-..------ 171 | molybdophanes, Struth‘o________.___--_-_-_- 3h
Meleagris gallopavo-_-.____.____.-.---.---_-- 34, |i momota, Momotus-......--.-2==s<2-_-+<<-:5 323
53, 112, 130, 226, 239, 325, 341, 343 | momoti brasiliensis, Spiroptera.________-__---- 322
meleagris, Numida_-_____________.____--_.--- 53, | Momotus brasiliensis_____...._-.------------ 323
59, 81, 88, 96, 111, 130, 136, 239, 266 MOM O bas a 323
Melierax gabar...___..--- 22 ete ne 304]: Monaca morpheus. .........--2::-.------=-< 131
menstruus, Pionus (Psittacus)__...__________ 80 MPT A 2a ee ea en 131
mercurius, Caprimulgus____._.....__-_____-- 129 | monacus, Bolborhynchus_-________-___-__--- 175
Merganser castor.-....--.--. 222-2222. c03l2- 148 C@entropuS: 222225. -2262-5-- = 127
SOVTAGUSs oe ooo tha oe eee 372 Neophrons.i2s522 2221s se 184, 185
merganser, Mergus--.__.-.-__- 148, 267, 343, 372, 378 Wil tite oo nes ee 138
mergi-merganseris, Hystrichis_._......_-___-_- 378 oli, Mionasa leucops 22222223 25.2.cckt eee 115, 128
mergorum, Eustrongylides___....__.__-_-___ XVI MIPTULONSs. 2 se ack see ee eee 131
335, 351, 358, 367, 872-373 tenebrosas222282 0.22 22. Scene oan 115
SHOngy lus. 56s ops ose o 372 TONGA tatede ee 5A Sd aoe 115
IMerrus albellus..--...22--.5------eavecsec- 372 tranquillss 29-2.) js 22 eet 115, 128
merganser__..._..-__-- 148, 267, 343, 372,378 | monedula, Corvus_._-_...-....--------- 34, 261, 262
SORTALO 2 sooo aan ee 235, 267, 372, 381 | monodon, Amidostomum.._--..-------- V, 20, 23, 26
meridionalis, Heterospizias_____...--_-____-- 297 SCLEROSLOMG- ==... onacanaceeenes oes 26
Meropsiapiaster =. -=.-.-------=.2..0-%--5-—- 288 Strongylus...----.<s<--<-- eee 26
Miertlaimipra. 22. . 2.2222 esse 139 | monogrammica, Asturinula_.._...-.------- 273, 282
merle, MUrdus. == 252-22 2- ccnp noes 139,206 | monoptera, Habronema-.------- X1, 173, 180-182, 183
Metastrongylidae___._______..._-.-_-_-------- 7. |) montana, GeOuryeOn.-.2-cssssesseesnte anna = 95
Metastrongyloidea____......-.--------------- 6,7 | monticelliana, Heterakis._.....------------ 53, 67, 68
Microcarbo pygmaeus-.-....------------- 148 | morinellus, Charadrius_-___-..---.--------- 139, 144
microcephalus, Ascari$_.....<-+~---~=<---==<— 152 | morpheus, Monacha..-..--..-.-------------- 131
microcephalum, Contracaecum _-_---_------ %147., |; Moschata,-Amas._.< o52- 27 wccnceenassa 62, 81, 136
152-158, 159, 160 Cainina. 24. 26s oscko essen 81
Microdactylus cristatus._..-_----------- 07-128150), ||) JMotacilla albas. 2.2.2: ---- sessqonaseoecan 139
micropapillata, Ascaris.........-----------=-- TGS) MULCTONGL, -ASCOMS: 2. ~~ 2 o-oo cece aee naan 20
micropapillatum, Contracaecum ---- x, 146, 153-154 | mucronatus, Syngamus----...-..------------ 34
micropenis, Tetrameres_-- XVI, 335, 340, 342, 348,351 | mugitans, Botaurus-----.......------------- 152
Wilcrapogon SpecieS....-.2-2.2--se=-c2<c-5—5 296 | multipapillata, Allodapa_-_-_--.-------------- 388
microspiculum, Syngamus-_-___------------ VI, 34, 38 Ascaris: 2-..ccsces. eae 154
Mucrotetrameres.- <<... < --<.<202e—4 these wan 333, SUDUIES, 2.csn8 ¢>44.55058 888-889
351, 352, 353,355, 358, 359, 360, 393 | multipapillatum, Contracaecum----- X, 146, 164-166
contorta...------ XVI, 351, 353-354 | multipapillosa, Contracaecum-----.---------- 154
CTU oo XVI, 351, 352, $53 | muscae, Habronema.-.--..-------------------- 172
helix.-<25 26 XVI, 352, $55, $56 | Muscicapa atricapilla__.....----------------- 214
454 INDEX
Page Page
muscicapae,“A cuaria. <sseser eee ot ssee ee 212,214 | nodularis, Trichostrongylus._.....-.---------- 20
‘DPispnaraguese ori ee sae 214 | nodulosum, Amidostomum.--....------------- 20
ilar 7ae sea snin wr nts enor 214 | nodulosus, Strongylus..._.-.-.-----2-2--22- 2 19, 20
ANUSCULIT ME TGTIGococe NEES 203 | Nonnularubecula_.......----------------- 128, 131
GORGYLONE NG soe ieee 203) |) Nothocrax-urumutums::+)- 220. eee 231
WMUSICUSH DUT AUSe Pee ees tie 139/144; 206" |* Nothura maculosa:.222)2 26" eee eee 121
MUTICUS PE AV O0 2228 5 oe ee SO ees 103,389 | nouveli, Tetrameres_-_..----------- XVI, 335, 348-350
NIELS: Vago pus! ee es ees 2h le ee 53, 84 Tropidocerca-22222. 2-2 BOA Ae 348
NEvOosyringateat==} cles n mee 4, 48, 162, 391,392 | novae-hollandiae, Plotus___........---------- 148
My ZOmimussests: 28 eee eres ines ees 203 | Nucifraga caryocatactes_-__--.---------- 39, 220, 373
nacandua, Caprimulgus--.__.....--..----- 1155 128" MG LFONS;RDISA ates ae 33 > oP ae 377
nactinds;“Podager2 2U seen! escent eee eee ee 128,130 | Numenius arquatus.-..-_..------------ 285, 372, 380
Tas yia, Aq Uilasiees = bos ee ee eee ae 138,316 |} Numidasmeleagrisss22-sesecee eet eee 53, 59,
MAG VIUS AO UCHIUSHseee ene ene fan ee 115, 129 81, 88, 96, 111, 130, 136, 239, 266
Diplopterus= 22:2 esse es 115 papillosa transvaalensis_____------- 96, 130
ft COS AMM ee ene re ese ae 138, 316 ptilorhynchas=2-c2-. 69802 se 96
SEIN RTLCLOUL ee ene ee tH MINE ER iri rIE ce Vo 69 PULCWike Aes Se EEE hea 133
nasuta, Acuaria (Dispharynz) -_....-----.--- 237 | numidae, Ascaridia_—_._........-.---- vil, 79, 96-97
Chetlospinun dese te s ee 238 Teter akigesve ss ccutn seve eee 96
Dispharynxssocs Se XU, 237-238 | numidica, Desmidocerca_-__....------------ 209-210
Spiropter ase ee eters ee ee ee 237) Witty ChCA HS briK sae oe Ha ek ae nese eres 138
nasulus, Dispharaguss ie. 222-2 sect 237, 238, 239 SYTM GAY sh ie oui shee ee ees 144
mattereri,;Caprimulgusesscos2222 ee eee 129 | nycthemerus, Phasianus__......---....-.---- 53
Hoterakcistsusics 2 sete nea 51 76-77 ay chiard eal grisea: = 2426) Sel pie tare se ee 152
neglectus, Hystrichis........_...- XVII, 376, 379-881 | Nyctibius aetherius_--__-----.-----.------ 129, 130
NY GTA E OG Bie eee ale hres ahr ee aS ee S QEATIGISH 241 = 244 Monee ENA ae 129, 130
Neophron‘monachus2 22222) 22ers 184, 185 Famaicensis: .27E es ery ON eae 130
Percnoplerusse ao. e cee sees 184,185 | Nycticorax europaeus_-_.___-___------.-------- 152
SPOCIES SSL e none et ae eee 186 TISHUISAE seat eo NS 141, 158, 281
neoplastica, Heterakis_.......___-- vil, 51, 64, 65-66 my chicoraxeswaenEee ses 143, 152, 281, 347
Nettioncarolinense 2. - 2 ae wee ee 246, 248 BPOCESNe Mee Fane ae te 158
OP ECCA 22 Se Pema en: hn eee 384\\nycticorax ‘Ardea: ed etee sent os See e 142, 152, 347
Nicticoraxiviolacenss 22223 °2ssee eee 348 Niycticoraxs5--5— 143, 152, 281, 347
nigeliisMevaloperdix=2=222422s2 2 ee 67°'| UNyctidromus’albicollis#22¢ == 22-2220 eee 130
TNT A PATTAS So ees oa in ERR Ree eee eee 2427/28) || tNivrocaclangula:t228ect ese eee 20, 267
STIS OT ates he SSeS at armen ee ee 20 {Oring sss hte Asie e ee eee 343
CAT OS tee as ee emt Bale Ne eae 281 filiguilasse ses ee cee e een eee Sue 20, 24
CWiconiae settee ee 34, 44, 150, 152, 281, 372 TAT ee oss Neer ria ash RE en ee 20
Maligulasstestecst cei e ee 24:27. | sODeSA, ELArlertides +. = fo ee XI, 191, 192-198, 196
Hey drochelidon=<22te1s 352 Se eee 259") sobscurus; +l haumalease: = coowners eee 65
Monachas2 222s oe eee 131: obvelatay Mitlarian sees een cee eae 235
Werle me taast sre re rahe ae eee 139 SINTON Esa os Sa ee 235
Oidemiasst es ve 2 aS ee 24,26 | obvelatus, Cosmocephalus ---.-..-- XII, 234, 235-287
SCOR ri en tre a hie Neen 259 Pisphavaguesssscsce sees ee ee 235
nigrescens, Caprimulgus_.._.........-------- 139 || :oceipitalis; (Urocissa:c2o2+ sass eeneeee een 220
nigricans, ColymbDuUS #22222 see eae eee 148 | ochrocephala, Amazona- -.........---.------ 80
nigricollis;;Podicops2s snus eise esses essere 1485250) \l'tochroptera, (Chrysotisi2 os seeoe eae e eee 175
MIPTUTONS: Wionasae eshte s 2 Se eee eae 131, |Gochropyeus}Erochilus:{s2s2s22eeene cee eee 294
nigrirostris, Macropygia.......-..--_-------- 78,84 | Odontophorus capueira.._--...--..--------- 63, 128
INissetus fasciatus ese ee ere 188 | Oedicnemus capensis .............---.------- 192
NESS (COMMUN IS = hos we ee eee Wench 277, 278 erepitans cis cee See ee oe 139
nisnsAccipiter ee eee sheek sees 138, oedicnemuse2.2.24i222-22 8 196, 198
‘ 176, 276, 277, 288, 298, 299, 301, 302 Vermicularis(*cei sss es sone 198
Asturst22 etait: sos ce 176, 277, 278, 298,355 | oedicnemus, Charadrius-_-....-...--.-.------ 144
Palcon sites 22225 138, 177, 276, 277, 278, 299, 300 Oedicnemus2 seo eoeeeecee 196, 198
Nivea; Strix: {tots Sees patente een ee 144 | Oesophagostomum_ --.-_-_----. on eee 29
noctivagus; ;Crypturus.... 2 eS 74,128 |? Oidemia deglandie 222-22 eee 293
moctia Athenee st ects eee 130 TUSGaA Le ESs tees Hee 24, 267
Strizhisi sac eee eee 35, 144 TUT ey ee eres nate ere eee 24, 26
rioctua glaux, Athene.r2-2-< 2222 2 203? }2Ollulaninae tes. tetas seen cere eee 6
Carine? 2 eee 210,242 "| Folor Cygnus een lees seh ee 61, 376, 379, 381
noctuad, *Actaria = ee ek ee 242 | olor domesticus; Cygnus—- = --2-- 22-2222 22s 246
Acuaria (Dispharynz) --_------------ 242 | olympioi, Subulura--_.......__-.-- 1x, 107, 121-122
AUGTADR eta ee e eeeee 119 | onocrotalus, Pelecanus---__...-..--- 34, 148, 364, 365
Dispharynx-_____-.-.-- XI, 237, 241, 242-248 | ophthalmica, Ceratospira_.-...-.---.-- XV, 320-821
Subuluras:sf2is2esss eee Ix, 106, 119-121 Chetlospirurasc: cece eee 328
nodularis, Strongylus....2-.-.22 222 -s cece 20 sOxyspirura.. 22:2. 222 XV, 322, $28, 329
INDEX 455
Page Page
ophthalmicus, Ancyracanthus__....__._-______- 320 | papillatus, Ornithostrongylus__._______ V, 12, 16, 17
orioli; “Spiroptera:: =... -..-.Geceuliee.--n- 322 Strongylus_......-- = tbs. 16
Oriolus guratus22< 226 . 424... sete ou. 188, 222 | papillifera, Acuaria__......__-._____ XU, 217, 218, £24
galbula . «2... 21ers. Sethe coe 58. 220 Filaria....<.2-.2-..-- 2 td ae 224
orispinum, Epomidiostomum _____ V, 28-29, 384,385 | papilliferus, Dispharagus___.._______________. 224
orispinus, Hystrichis__....______- XVI, 376, $81,882 | papillosa, Ascaris........_..__._---_---_. 67, 68, 122
Wirongylus: ae bee ce 28 Heterakis......._...-.._- Vil, 52, 53, 67-69
ornata, Acuaria:::-..........2sc2 XU, 217, 223-224 Oxyspirura_...22- 22.2. Seusn is... - 317
ornatus, Cephalopterus__..........-_-______- 186 Spiroptera..-_.. .b22822 317
Dispharagus _....... <= Sas: 2.2. 223 Subultira22.-.......2...4 1X, 105, 122, 123
(alee. 222-3 eee 297, 298 Thelavig-- 2225.22.26 5.8 XV, 312, $17, 318
Ornithostrongylus___.._..._..- 8, 11, 12, 13, 14, 16, 17 transvaalensis, Numida-_-._____... 96, 130
douglasi..2222i2:.. Vv, 11, 14-16 | papillosus, Cosmocephalus__._..._...-------.. 235
fariai- . 22-32" V, 11, 12, 18-14, 15 Dispharagus_._._-_.----_.-------. 235
hastatus..<.2cs2842. 2s. Wp 12.17. Eustrongylides._................- XVH,
papillatus. 2c: - - V, 12, 16,27 358, 367, 8738-374, 875
quadriradiatus_-_-_-__---- v, 12-18 Eustrongylus_..___- 370, 371, 373, 374, 379
orthocerca, Ascaridia._........_.-.._- VII, 79, 97, 98 Hystrichis__._.. 358, 370, 371, 373, 374, 379
Heterakis:. . ..2vo: steers... 25. 97 Strongylus......--...-.-.-230 22 372, 373
Ortyxivirginianus...__._...2seuebeL-_.s-- 53,88 | Parabuteo unicinctus..............2cstzc2s.- 297
ostralegus, Haematopus------.-.____- 148, 285, 287 | paradises, Grus_..............-.--..eetnen. 99
Otis afroidess:.- + 2-224 sles ebsites 192, 198 Tetrapteryx......--222 2. B23 99
Hovbaraxt._.... 2.5. co eee 130, 195,196 | paradoza, Rictularia_................-_------. 263
macquoeni. 2.2225. = ee 196, 198 Tetrameres_.___._______- XV, 334, $86-337
Muficresta).. 225 ---..22 set vasea kn 192, 198 Tropidocerca___._-.----- 335, 357, 358, 371
tarda gece: Stee eek 10, 16, 34, 58,67 | paradozus, Tropisurus_.........-.-.-.-.----- 335
Netrax.t S$. cL¢-.. =. 2c eeuee 53, 67,117,290 | Paraguaiae, Gallinago_.............---..----- 342
Otus: brachyotus- —2_-2...steut 2482... cs 144,276: ‘| parasitica, Lestris........-2.<-s<5....222i8. 148
eholihas=..... sco... Js e. ce 325: | parist;Heterakis: .. set: ssasetiesst nese vu, 50, 69
WHIBATIS=<2 S222 25 apa ae 44°276.1 patroti; Cyrnea_--......<...s222c2ee.-2 2c 167, 171
METIS PA CLGUUS = = 2 =. 222 Saket tiveie 144,276 | parva, Hadjelia__......--..-.-.----- XI, 187, 189-190
SUN ns ec ae Sa 138,177. | parvirostris,.Crypturus....:22--252h.-<2--.. 121
ovale, Contracaecum...__..------.-- x, 147, 166-156 | parvovum, Oxyspirura___.---- XV, 322, 328-829, 880
prise ASCOrie® gn oe nn ete 155 | iparvus, Helodrilus.....2.c22s2oss.2e25 522 ...- 82
ovata, Physaloptera_--.....__----- XV, 297, 809-310 Syngamus.:3.........12. 24202 VI, 34, $9, 40
Oxysome acuticauda.......---.2---.-.susc8s 2 133: || ‘Passer domesticus... -.<si23s5.).22uc286 335
O8ACiUe So en nn 134 domesticus tingitanus-_-__--_-..---.-- 203
Oxyspirura_------- 311, $21, 323, 324, 325, 328, 329,330 | passerina, Strix_............------.-----.-.-- 62
anacanthura_.....----- XV, 322, 8328-324 Sumi. ..2.5...-Uieteu ees. 62
@nolatiatd «. «..n<-<aco28ussests oc 317 | Pavo cristatus............ 34, 53, 63, 103, 310, 311, 325
anthochaerae_-.....------- XV, 322, $24 mubletis.c 2222-65 a 3 eS senses 103, 389
brevipenis...._-.....-2u22s XV, 322, $82 spleifers= 5.2. s= esas 63, 310
brevisubulata_...........----- $22,525 | pavonina, Belearica_.....+.:-2:-.--.--------. 89
cephaloptera_.____- XV, 321, $£2-323, 824 GnUS 5 Q.65 322 bean eee 143
heteroclita.......-...-.--2si68 322,831 | pavonis, Acuaria ( Cheilospirura).....-.-..--- 389
mansoni_......--.- XV, 322, 826-328, 329 Cheilospirura.....-c2s292scib leet 226, $89
ophthalmica_-_..--.-.-- XV, 322, $28, $29 Pseudaspidodera. -.------ vim, 102, 108-104
POPUlose = «nap aacenananncdtien 317. | pavua, ConuruS. 22.-.....-----2-=204it 2... 80
parvovum.-..-. XV, 322, 326, 528-829, $80 | pecoris, Emberiza-................2---+---.-- 177
siamensis.....<.-.<- XV, 322, 829-S00, $82 | pectinifera, Spiroptera_.........-------------- 266
He EUIG. . <2. acc ce tctertestatie se 316 Streptocara....-22..- XII, 265, 266, 267
sygmoidea_......-- XV, 220, 322, $30, $82 | pectoralis, Circestus__.........-.-----..----- 138
tanasijtchuki._.... XV, 322, 330-331, 882 Eclectuig....--22.2. 2822282. ne 320
Oxvyurata:...<..-..c-2-cscersteen stpsetsipocke 53892: | Melagica, Acuaria.........----- 20-020... - 263
Oxvurids:<. 35-2 22 eee oes 49, 363 | pelagicum, Prionostemma-.----..------------- 263
Oxvuris 8). nos PRU S pcccceccn ence 108 | pelagicus, Phalacrocorax_....-.-.------------ 148
allodapa .z 22esascalisik. 2. --se-0e 108 | pelecani, Gnathostoma-.--.-.-.-------- XVI, 865-366
Oxyotoidea=:< =. 25.5. oe Be. 52222222 49, 391 Sclerostoma: tee.) Sete. Uae 365
pachicephalus, Hystrichis.........------------ S78") PelécanGss 5..-2--4 occ SUSU concn cs 148
pachyrhynchus, Catorrhactes-__....--------- 235 americanus.....2.0 0220246 ..--..-. 148
Walacornis fasciatus. Aes isUe.....--a-----~ 175 conspicillatus.....sU2203.. el 148
palladiceps, Platycercus........-.----------- 175 crispuss..2 5s JaN2Gcos.1 SMBS 153
palumbarius, Astur__...----------- 138, 176, 288, 355 erythrorhynchos._-------.--- 148, 153, 154
Faleos 22... . 138, 177, 288, 300 (IBCUS acess awssoessaescssSe se 148
palusizis, \Nal0o..ca2s sesso eo ote eeonaaceee 297, 298 onocrotalus..-...--------- 34, 148, 364, 365
Pandion: haliactus.-:-../f2222-<--.--------- 137 PYGMACUS. {52022 0e =) Aes eae wee 148
456 INDEX
Page | . Page
Pelecanus rufescens__2o2 sss se es 369 | Physaloptera abbreviata__...-.....-------- 296, 307
species= 22. =— <3 sahara 148, 153, 273 acuticauda._----- XIV, 296, 297-298, 305
trachyrhynchus.. ..=. 4s. 148 alates = XIV, 296, 297, 298-300, 301, 305
Penelope humeralis___...__.._.. Aves. 79, 98 alata chevreuxi__....----2.. 2 296, 301
species... =< saat s eae 315 alata nouveli___._._--___- 296, 301-302
penelopey:Anas-- 2220 tos hain 27, 28, 246 bilabiata..-- 2. eee 296, 307
*Amser. - 5 ee ee eo 20 brevicauda-_--__----- XIV, 297, 307-308
Marea = 2222 era eee ict 2 27 pulbesatessss525 XV, 167, 297, 310-811
penihamata, Spiroptera___._------__-_- xm, 205-206 Clansascssc REE eee 295
pennata tA quila-——--- = 22. andy seine 138, 298 CRASSA ses 5 296, 302
pennatius) ih alco.. <=... -. -_.__.. seen 299, 300 CroSloe.2 see soo sse eee XIV, 296, 302-303
Peres fluviatilis. 2i300:2%. ses ase 368 fusifermis__..-2.2--202e53 XIV, 296, 3038
percnopterus, Neophron__..-_------------- 184, 185 galinieri_____- XIV, 297, 300, 302, 303-304
Wilt 5. eis ae§ 184 gemina._..3=.22285ui2- XIV, 297, 8304-305
Perdix.cinerea==-- = sarees ayes 10, 34, 53, 88 inflata ..vsPHeewee_ Le es 297, 305
COULLER EK Sens re ie ae ae 53 malleus. sah He XIV, 297, 308-309
Gentata Ate 3... Mubespe eee 2 128 megalostoma____- XIV, 296, 298, 300-301
STARCH Pe EES 8 ay Se 73 ovata. 2-.-.c2 ee XV, 297, 309-310
DONGLE ee 53 pracputialis...... Sse p9sebe 298
Saxatiliss: 2 =<). .2. Seelam 53 rotundata-..2. 5... tee 195, 196
SDOCIOS 2 ae ee ee 58 saginata..... SRA ee 207
Pergix,; Perdixo----...-- 2. sea e is 53 saginata strigis brasiliensis__..__- 107
peregrinus;, Waleo.2 2-0. ea 138 speciesisss. 20s. 5 cess XIV, 297, 309-310
perforans; Spiroptera___..... .2ateeneree to 226 striata. 2. on 167
pergracilis, Strongylus._... 22 Soie ee 9 strongylina ...-- 222.2. 8seeee 297, 309
Trichostrongylus____..-.-2-----_ v, 9-10 strongylina cuculi-seniculi_____- 107
Pernis'aplvorus 22220-2235 SRE pes 137 subslata)- <2. -2- XIV, 297, 302, 305-306
SPOCIOS oot ce ape apa 137 tenuicollig-2 2-2... Serna eae 207
perpapillatus, Eustrongylides__.. xvu, 367, 374-375 truncata. ..2--- 32-2 XIV, 296, 806-307
perspicillum, Ascaridia__........-..------- 4981.87. | ‘Physalopteridae. 2-2-2...) 2. 2 163, 295
CA SCOT IS oo ele ea eee 81°}: Bhysslopterinaes}. 22s 2965, 391
eter akis.... oi eineat ots ee 81-| ‘pliysalura, Spiroptera--— 2s oS 298
pertinax,;Contrus_... 2... etn tsevalt agers a5 || deiayaicayanallss one 8 ages 113, 115, 124
Pionus,,(Psittacus)_..-28ccss2k: 2 8) 4: Picacaudatace seo eo ee 34, 139, 220
petrosa, Caccabis__.-.------ 73, 126, 167, 171, 187, 239 PCH eo Se os PRES 34, 37
‘petrosa.spatzi,.Caccabis......-. seesaw: 192 | pica, Corvus. ::...:.--3__..3 . eee 34, 220
Phalacrocorax carbo-------- 38, 148, 209, 258, 370, 372 Pica 64546 5 eo eae 34, 37
fusicollis -. 23422-4223 148.) pictazgDhaumalessos 28. J. 2 SS 64, 65
graculus.c.22¢222~ See. SS 148 | pittus; Chrysolophus..... <u 53
javanicus.....-..<.-. tas 148 Phasianus 22 =.=. Sooo eee se 34, 53
pelagicus.5..32.3225 Seo 148: | pituiiC olumba_.....--.. 2S 87
pygmaeus. 2s3.-c2- 52 Sess 370 | ‘Picuscampestris-—_..-. -.. -Saee 178
SUlcirostris....._- 2 22eee) Fae 148 CANUSivjcc eee PSR SS 34
UT ee eee 148 prammiicus % 32. Sa ee 179
VOLTUCOSUSLEZaRAk | Fast | 148, 266 Jumana sess oun cot 55 178
Phasianus chrysomelas--.-_..-.--------------- 64 maj OR ee ees ose SORE = oe 34
colchicus.. . ..- 2aesab ieee 10, 34, 53, 64, 65 mMartivs: 2.2 2S... see os 215
gallus.) 22 tt 34, 53, 81, 226, 239, 306 Wiridis{ 20ers Se 34, 215
nycthemerus..-.=-2--esieuiste 63 | pitariS}-Turdussat 2 a... See 139, 206
pictus: === eee eee 34753 | pileste, “Ardeai 65252-5255. 5S:2eeueet 142
reevesi.2-.-.-_- se¥asetas |. oS 34 | pileatus,. Hylotomus:...-..+=-.-52292=_--_-- 215
satscheunensis_.........---------- 65 | pinnatifidus, Ancyracanthus-------.-------- 392
Veneratusies 545.258 sven 5c: 63: |. pinnatus}"Botauftiss.2.-s4 ee es ee 371
wersicolor:..-.-.._.-. 22ebe sees 53 | Pionus (Psittacus) aestivus__.-....--..-.-.-- 80
philomela, Luscinia..___ss2cseebseseiet eee 139 Saracanzacen. we 80
Phimosusinfuscatus_....- sseseese ee pe 377 araraUnaccccc esse 80
Phlogoenas luzonica__......-..-------------- 87 dominicensis___..___----- 80
phoenicopteri, Acuaria (Hamannia) ___.------ 257 festivus. 25.32 ees 80
Eichinuria so. ts joke ee XI, 245, 257 leucoc. s432eat Jos 80
Hamannig-_-.. 22225 Seseee JS 257 leucotistewts®: 20ers 80
Phoenicopterus roseus-_-..-.-.--------- 143, 257, 339 menstruus. 22s25 22) Sees 80
phoenicopterus, Crocopus-_..-.-------------- 87 DPertinaxeeessos the - pe 80
phoenicurus, Pionus (Psittacus) -.-.----.---- 80 phoenicurus____...-.----- 80
Physalopteras.. 2-22 2 oe ce Saree oe 4, pulverulentus_____-__-_-- 80
207, 208, 295, 296, 297, 298, 299, 300, purpureus - - .-- -sef8_ 222 80
301, 302, 303, 304, 305, 306, 307, 308, species - . 22. — state at 80
309, 310, 390. . sulfareus.=.. 52. 0ee ees 80
INDEX 457
Page Page
Pionus (Psittacus) vinaceus.-__...--.-._____ 80 | Prodiceps cristatus...-2:-.. 25202. 22 2=t22.. 372
Pisorhina atricapilla._-__.-2 22-22. 222222¢ 222 107 MUN OF een ace ee a ee 372
Platales:aiaia..---~.-< 2-22 ce et Se SE 241 | Protogerys virescens.__-.....-..2...2.0.2-..- 175
leucorodia:<.--=<<. 22. 339, 369 | Pseudamidostomum_-____..-_--.--...-___--- 18, 8838
plataleae-ajajae, Spiroptera_......__.__-..--_-- 241 loosi= = 22252 see 883-884
Platycercus' barnardi-..-.=..2......-720 175 | Pseudaspidodera-.-_-......--.-- 50, 102, 103, 387, 388
Oximius.- 22222252 ~ FP eee 175 pavonis___-_-_-- vil, 102, 103-104
Palladiceps vests se 175 voluptuosus.-2 9222.28! 22 387
Platycichia flavipes:_... =... et se 359 voluptuosus minor_______~- 888
Plocepasser mahali-___-_...22.-.22-..-_--. 192,198 | Pseudotantalus ibis......----[¢ 222222222222 157
Plotina, Subtilura--=-:.::--...22.22 1X, 106; 1994798 | Fsittacus’sinensis-.<---:. 2/5 /2-22.222__-.-..- 320
Plorastamhingss]s~Ss0 ts fees ee dest eet 148 | Psophia viridis. ..-.-2..2 S82, BRU Es 55, 69
lavaillanti=:-= 222222222 22---- bt 148 | psophiae, Ganguleterakis___..._.....___------ 69
melanogaster __....--...-------- 148, 160, 213 Hieterakis <2 222-25. -e aes vu, 52, 69-70
novae-hollandiae____.____..______.___ 148 | Pternistes'species:_..__......--23.022222....- 193
PUfUSe sean De eee ee 122, 157 SWainsoni-..- --. =. 6-2 Eee ae 130
Pluvialis’apricaria.-<-2 -2-:.<22s220- 22 139°1417 | Pternistis species........-..---=-..-22-22!-.- 111
pluvialis, Charadrius_.............__-. 139, 144,372 | pterophora, Ascaridia_-.-..-.-------- vin, 79, 97-98
Poculum, Subulura__.____..----..-.-.-- Ix, 106, 123 ASCONNS sooo o == ofS 97, 98
odacer nacunda:=- =-=<- v=see ih! tele: __ 128,130 | ptilopachydis, Acuaria___._....-.-- XU, 212, 214-216
Podiceps'atirituss.--.- 62.202 148934. | Ptilopachys fuscus¥: 2200. 2.22-2 B22 bees 214
capensis: = +2222) FFP hk 156 | ptilorhyncha, Numida.=-~..-2.-222/.-.2 96
cristatus:<. ee". 2 - See: 148, 155,270 | Puffinus kubli___....-.2.2.-..sssisski.- 235, 263
Gominicensis:3 225 26325220 ve bs 148 | pugionatus, Syngamus. ...=.2 122122222 222t2 34
fiviatiliss-2 ee Jee Re 248) 1 “pulex, Daphnis..2-<<c.2ss6s-2e2- 246, 248, 343, 345
MINOP. 4.2 Sas SSS 2 SUSI NN oe 148 Gammarus... 25 eee eee 343, 345
MIPTiCOlUS#a2—2=2eAeces. cess tS 1487959 | Mulicts,, Wilaria__.-_--_..-- 3. ALP tee 343
SONOPAlUS Jans 3 Fe MEE ES ee 175 Spropler a... assess 2 sees SSS Sek. 343
Poliocephala, Chloephaga_____._____.____-.___- 20,77 | pulverulentus, Pionus (Psittacus) --_-------- 80
polyclottus; Mimus-:.2 S222 22h eer 199.| punctate, Kathleena....22-. 2 ee: 157
Polyplectrum bicalearatum______._-..-_-_-.- 386 | punctatum, Contracaecum_-___--.------- X, 146, 157
Pomerinus; Westris:22-.-22-2<. 52. P et eS 148 || purpures, Ardea_...........5--24 eS 142, 152, 281
Worcellio lasviS-= == 222. 2cesenese co 238, 239 | Purpureus, Pionus (Psittacus)__.------------ 80
Porrocaectims- eee et 135, 136, 137,138, | pusilla, Heterakis_--...-------------- vil, 51, 70, 71
139, 140, 141, 142, 143, 144, 145, 146, 147 Microtetrameres-.....-.------ XVI, 352, 359
anguillaes-.22_..<. 2a" IX, 136, 146 DCM UIMNEL CS ne saa ae ae eee 359
angusticolle__..__.___- IX, 136, 137, 138 | Putaustralis, Heterakis-_--_-.------------ vu, 51, 71
crassuim: 2422222... 156-187 | Pygargus; Circus. <_... -.---------2-SS 2 2s 298
depressum___________-__-..-_-- Ix, Falco-.--.---------------------- 144, 299
136; 187=198° 189) 144.445 | Dygmacus, Carbo.__-.-------<---224-55===2_- 147
ensicaudatum=”.2220 22222922! TX, Microcarbo. 225-222 s2L2 2 best 148
136, 189, 140, 142, 145 Pelecants. a. ssseee = eee eae ae 148
heteroura____________- IX, 136, 144-145 | Pyrrhocorax, Corvus-----..----------------- 220
kirghisensis___________- X, 136, 145-146 | Pyrrocorax alpinus------.------------------ 35, 220
praelongum______________ 136, 140-141 | Quadricuspe, Contracaecum_--..------------ 152, 153
reticulatum__________. IX, 136, 141, 142 | Guadriloba, Acuaria__-_--..-------- XI, 212, 215-216
semiteres_.__._____--- IX, 136, 141-142 Filaria...--.-------------------- 215
serpentulus___._.-_- 1X, 136, 142-143 Spiroptera___.--.---------------- 215
spirale:~:-. 7S IX, 136, 143-144 | (wadrilobus, Dispharagus__.-.---------------- 215
pourpré; Héron-..-~-< 5. Se 979 | quadriradiatus, Cephalostrongylus --_---------- 12
Praclonga Ascaris == <2 22a sexe dees. AE 140 Ornithostrongylus_ -_------ v, 12-183
praelongum, Porrocaecum__-_-._______- 136, 140-141 Trichostrongylus. -_.----------- 12
praeputialis, Physaloptera___..._.-....------ 298 Strongylus__.----------------- 12
praestriatum, Contracaecum_---_-_- x, 147, 166-157 | Querquedula circia-------------------------- 380
pretoriensis, Hodotermes_____.._____.._____- 197 | GQuerquedule; Anas._._.-..---20s22e20ie_.__. 20
primitious, Strongylus.._---2222222.22------ 34 | Querquetula crecea______.. 32322222-.-.-.- 384
SUNGGMUSzenccwsereeeet S45. Hs 34 | querquetulae, Epomidiostomum___------.---- 27, 384
iPrionostem MG 2. anes ese S ae 35 262,269 | Quiscalus major.....-.-..-...---.---25t52.=- 318
decOTwWM 29 ODI es 269 quiseula.<.----.6-e Be ee 239
pelagicwmere Sse a= =~ 263 | quiscula,.Quiscalus.._........--.s-222-----6-0 239
procellariae....--rtes259=..--- 263) | -FAtlsell, ACUATIO <connscowseeneas 2 EEE = 283
RUDY so SINS awe 263 Amidostomum___--..--- V, 19, 20, 2/, 24, 383
Procellaria anglorum=-__......-..suzalls> 22s 263 Synhimantus........---t223 273, 283-284
procellariae, Prionostemma____-.__---------- 263\ | eatloides, -Ardeola:_s-—-<-s--<-s--=---~ sees 152
Seuratias...-s.2...~.----¢- ent 268 | Rallus cayennensis---.---------------- 285, 288, 293
Spiroptey a... <202-5-.<2588 292 2! 263 | Ramphastos vitellinus.__-------------------- 237
procellariae anglorum, Spiroptera_..---------- 263 | Ramphastus, species. -------.-.------------- 319
458 INDEX
Page Page
rapax belisarius, Aquila../..._..-..-.....--- 304>|: nafay Lophura.2 6222s l deep ooets ee eee 385, 386
reclinata, “Ascorie o-oo ee 124° | vufaxilla, Leptotilas2ec2. >>) We ee 13
Subuluras- 22222... she 1X, 106,124.25 | rafescens, Pelecanus..-_-02.22-4_2. Sees 369
mecta,ACwariGi eo! ek oe oo ea a 280 Rhyzichotussss0s <2 sk eae 58, 121
Chetlospirurge 2 eke ae ee 280 | ruficollis, Caprimulgus_-_--_.----_----- 115, 128, 129
Fr alarieenytene fo at 269) | neficresta; Otiss ieee ro. ae ae ee 192, 198
Spiroprengs.. i: 88 pegs ee Ss 269i) rahiventris, Buceos 22224. 2.) c 128, 131
Streptocara. .- -seussesess2- XIN, 265, 269-270 Capitosse 3-5 eee 115
Synhimantugi siscns252742 - XIV, 273, 280-281 TURGUS:. <2 - oc e 359
mectovaginate, Acuaria.__...-- 5 eo 243 | rufovulgaris, Colymbus----..----.-------- 148, 267
Dispharynx. -~-.. ..=ce ME 2377248). |; rafus; \Caprimulgus..._-- ee 130
rectovaginatus, Dispharagus___-----.-.------- 243 Circus 222 ee 3 138, 177, 298
gectus, Dispharagus= = 22-8. pei es ae 280 Maleos 22 2teni. 2 are eae 138, 177, 299
mecurvata..Heterakige.2 -. 2 3s ee 124 anius:3423 yes os 2 = 201, 220, 221
Subulura-2- => 222253 IX, 106, 124, 126, 127 Plotus. 23sc42 20 os ese ee, 122, 157
meevesi, Phasiants: 2 82 os Seder ates k 34 | Rupornis magnirostris.-._:.-.--...- 2-2. .22-- 313
TORALIS MG WISE oe a ea 13877) | Rusguniella. 22432 eB ee 212, 258-259, 260
regulosum, Belearica__.._._....._-.-.-_-___-- 89 elongata; -- 2.053 Sse XII, 259
SELICILIGULS VASCONIG.- 5 eee 141 Vanelliiibtse. ues XII, 259, 260
reticulatum, Porrocaecum__-_-.----- 1X} 1360447/5148" |; rustica, Hirundos2/2-2- 2222. = = 22k ee ee 220
retortaeformis, Trichostrongylus____.___.._- of |i rutilans, Ralcooseew ees eee 138, 297, 299
revolute, Chevreuxia..- 2 isles XII, 282-283 | saginata, Physaloptera__.-__.-.---.--.=------- 207
Spiroptera) io. 2 oo . Seeae en 232 Spiropterat. 2:2. 2 ee eee 207
revolutus, Dispharagus__...-..--------------- 232 | saginata strigis brasiliensis, Physaloptera____.- 107
ithahdissatalei we) ua. NN ele ee A-302''|’ sagittata, Ascaris: clea. bien ae 281
Rhamphastos erythrorhynchus_-_________.___ 186 Synhimantus-________---- XIV, 273, 281-282
Witellinng.- oe ae 186244" |’ Salicaria turdoides: 2008 0. 1) 2. pues es © 139
Rhea americana...) -.--------...e=s8< 32, 69, 97,165 | sandwichensis, Bernicla_..._..........---.-- 62, 267
rhodes) Thelazity oust ger en ort ghees 312 | sanguinolenta, Spiroptera_.........-.-.------ 203
Rhynchotus rufescens_--.....-.-.-.-------- 58,121 | satscheunensis, Phasianus_-_-.-.-.-.--.-...- 65
Rictolaria... 2 bere _ pes ates Nagi 18262° |: satyrasiCeriormis.. | ge 53, 57
MOTMAOTEZ Ls 2 22 ie ee 263 Mragopan ecto.) 28 oo see 57, 64, 66
Shipleyi.. 25 ceo ae OE. sake, 263): ‘saxatilis“@accabis. 2 22 a ee eee 88
ridibundus, Chroocephalus____._____-_-_-___- 148 POrGiKe ee Le ee 53
uarus: 82 3 42, 43, 148, 234, 235, 265, 271 MurGuse= che $y 2 pep ae 139
rikwae; Numida-<- =~ 22.22 eee 133). |) Saxicolawmbetra 2280 a2 gery See 218, 225
"ima, sHeterakis. 22: _-_-_ Lippe ane ees 125 | scaphiuris, Caprimulgus_.....__...--..------ 129
Subulura*s- heres 2555 eee 1551052125) || seapularis;sArdeast 2-20-2252 2 poe nee 142
mama, eleterakis === — ! <8 ee S 125),.\ scapwlatuss Coryiisies = 2 a2 eee ee ee 223
Subulurasse ee et Ix, 106, 125,126 | Schistorophinae_._ 211, 284, 200, 292, 293, 311, 391, 392
riparia, Hirundos2) 2.5) -) acted te 220 | Schistorophus-_--_- 260, 284-285, 286, 287, 288, 289, 311
risoris. ‘Columba 22228 22: 2) een dnt SG. 87 acanthocephalicus----_-_- XIV, 285, 287
Stermas2 tao cere he beers 235, 362 aulieatina--- Sie. go XIV, 285, 287
rodhaini, Contracaecum____.._._._- X, 147, 157-168 IDICUSPIS. © haps 285, 286
Kathleena 22s ctu detest i. keys 157 bidens:=.22-ece XIV, 285, 286, 288-289
Rolinius roulroul 22-4... eas A 388 COP OTiIsi$ se ee 287
roratus, Ecloctus2 23.0205 20 jee eo 175 laciniatus. =“ ULe sae ea 285, 288
rosarium, Contracaecum.--___._-_--- X, 147, 188-159 longicornis.. 2.-.<-. XIV, 285-286, 287
MOserTius, ASCOTIS Kone. ewan Sunes - lett 158 Spinvlosus:: 220 a ree es eee 285, 288
roseus, Phoenicopterus_....-....-____- 143, 257, 339 ALU ELLALEr G2. =the ene wae 261
rotundata, Acuaria.......-..-.--------------- 229 | Sciadiocara_.__.....-.----- hae 212, 260-261, 392
Cheilospirura___........-_- XII, 226, 229 Secunda... -cebeee cages 261, 262
Hila rig tsos eine ee 195, 196, 198 umbellifera. cic taatase2s- XIU, 261-262
Hobronemsa ...- 3... wee ets 195,198 | Sclerostoma boularti__.._......-..-------+---- 44
HMartertia.. 28-253 XI, 191, 198-199, 200 cyathostomim=o22 cs Oe ee 43
Physaloptera....<...- 2 oda 195, 196 dimidiatum. -2-— <23=s20 3s. <2 32
rotundatus, Dispharagus__.....--.----------- 229 Lari. 3: . Sot Re toast ERE 43
molige; herdix. . 2222225. Sh os eaegers reels 167 MONOGON 22222 a nn 26
Rdplroul, Roliing= 2. <2 = stsenabpapp ee 2 388 Peleca Nie. oc. ENE 22 365
rubecula, Luscinia: 22... |. eet k aht 224 struthioniss~<seeebo nines et 31
Non a aes on) eee 128, 131 tadornae. a2e2 Ste one eee 42
SylWias. o<n > 22. stan Sees F _ . 224 trachealescoks WG 34
rubetrasiSaxicolas4-— 2855) eeteraenes 5 218, 225, |; Seblopax gallinula..223.-=- 2 ato ees 254
mubra, Lis abe fra 261 major. 232 eae ees 261
rubripes; Anos. 0.) 224 st ue gap 246;248: |. Seops leucotis....- cco Ae See 127
rufa, Anhinga: ..-<:) S23 eee ay aed 368. | scoticus,, Lagopus...c.c_22s_ Ae 9, 34, 53
Caccabis... 2.06.2: dak eaters 2 126, 172,227 | scotti, Contragaecum_-_....__..-._.-- X, 147, 159-160
INDEX 459
Page Page
Scotti Kathlieena--. <=... ES 159 | species, Dispharagus_.....-..---------------- 227
scriba, Galaxias.-~-.:--.=--:..s.20t22224.-. 368, 371 Eustrongylides_._........------.- XV, 369
RCUTO, CUTNEG.2 0. acetone 172 Baleos=- =: 2242222 ee 297, 299, 305
scutatum, Gongylonema__._______.---------- 205 Brancolinus'==:-==<===-. 5 suas soe 134
secunda, Sciadiocara___.__._.__._---------- 261, 262 Geopellasssssssssehssas=so¥inn ee 94
segetum, Anse c:-. 22+ = 29S s vis 28 EE GO eae ae ee eee 217
ANISOR 22 sees s- 2S 20, 62 HLYStricnis 5 sq oases sess cee oe 378, 379, 381
semifasciatus, Lophoceros.--.......---------- 188 Wanius': 5-4 s2i2222---32202" 2525 ---. 355
semilunaris, Cyrnea__..........._--------- 167, 170 MUSTUS sos 2! s 32 sees a5 ES ee 43
Spiropteras 2-222. 2 lak 171, 205 Micropogon==sa0s-2-- 8 eee 296
Wamiteres, AScayiss< 2 --- 2 2 ese 141 Neophron disse. <ses eee eee see 186
FOUSOTIGS ooo 141 INiveticorax 2: ="=25 Sues ee 158
Porrocaecum._____._--_-_- IX, 136, 141-142 Pelecanus= =: 2-222 ce 148, 153, 273
semitorquatus, Caprimulgus_--......-------- 129 Fenelopes: 22: s2iic5 ioe tee see 315
Chordeiles- 2222 et See th 129 POL hn 228.2 © Se eee 58
Luroéalisessc2.tssstee teh 130 IPOrnis ri 2ene oh <3 sos eee 137
senegalensis, Trachelotis__.....-..--------- 189, 291 Physaloptera -_--.-.------ XIV, 297, 809, 310
senegalus, Poeocephalus.__.___-.------------ 175 Pionus (Psittacus) -.-1:..2-._..-.... 80
seniculus, Cuculus..........-..-=------ 107, 115, 309 Pternistes= 2.525 < 2: =< sos-aee eae 193
Sephina francolinus:_...-....2 iv 2-224.-..- 130 Ptermistis 223 = aso ot ee ee 111
septentrionalis, Colymbus____- 148, 234, 267, 367, 372 Ram phastis 22 ee es 319
serpentulus, Porrocaecum-____----- IX, 136, 142-143 Sternaeec 22 Shoe 22 sce ee 285
serrata, Ascaridia...==...:22-=2=22252: VII, 79, 98-99 PSE I el meee ea 118
Hetevaktg sot nt cee el eee 98 UONGULUS = ax oe se oe ea 43
Serrator, Moergus:---- ===: .-.-.--2- 235, 267, 372, 381 Sibulura--2227222- 22 ee 105, 132
serratus, Merganser- ---- SSeS RE ees 372 Terpsiphone::-2 2522522. oe see 188
Strongyluss. 2222223 10 Tinsmus <2. 25 ae2ee nee 55, 71, 72, 99, 128
Serticeps 2) 22* ...2...----22.2. _--- 284, 293, 294, 311 ERrovones- a5 2 oe. ee 110
vulvoinflata__.......---- XIV, 293, 294-295 UIDIK ooo 2 8 tee eh 132
Sourati;. OyImea..s-2222222522 2220520 X, 169,172 | species new?, Dispharagus__._.-.___________. 391
Habronema..-.__-_----- XI, 172, 173, 182-183 Dispharynx. = s2-.2ss222-2 fee 391
Subulura......-...-.--- Ix, 105, 111, 126, 127 Gongylonema_.----.-____-.-__- 390
Bouratiaa--..! 22.22.22 2.- 212, 261, 262-263, 264, 269 | speciosa, Columba_.......-.--.--.------____. 87
LOCOMaAriae!: 2-2 ee — eae ae Ie woo.” | spiciior Pavo:.222-2--=seee- hee Se 63, 310
shipleyito vss .o2 426.22 XII, 263-264, 265 | spiculatus, Heterakis__.._...........----.-__- 128
aetratite Cyrned:. =<: 22a 172 Strongylus-. 2-2 ee 77, 128
shipleyi, Gnathostoma_.-__-_.----------------- 263 | spiculigeruwm, Ascaris___.._.......---.--____- 147
(Prionostemma.. 263 Contracaecum.-_-_. X, 141, 146, 147-149
Rictularia <1 tt aia ae 263 | spilothorax, Circus...............----.---_-- 90
Seuratia_____- ssbane ss ss XML, 263-264, 265 | spinifera, Echinuria__.................-.----- 254
Siamensis;Centropus=-2-- st ae Mets. 329 DEAN sooo eae tae ee 254
CRElOsPiTure soon ee ee ee 329 | spinigerum, Gnathostoma__________..-_____- 364
Oxyspirura ______-- XV, 322, 829-330, 332 | spinosa, Cheilospirura__......._.-- XU, 226, 229-231
RUE) ACUATIG 28 tenes kana neta eaceacee 222 Habronema.2222-- 2-3-2524 XI, 173, 183-184
SELeL en A iSite cee Cows abe REPOS te 127 | spinulosa, Filaria............-.-....--------- 288
Subulura.........-..--- Ix, 107, 127-128, 129 | spinulosus, Schistorophus_-_...-...-...__-- 285, 288
sinensis, Centropus...-------------- 86, 101, 125,329 | spirale, Porrocaecum_--.._-...----- IX, 136, 148-144
EPSIGEACUS Se 22s 2 eye A a SAU) i) spiralis, Acwaria. 3.5. 222-2 Soeceeeesse anes See 238
skrjabini, Acuaria ( Cheilospirura) __.-.------- 389 ASCH Sa ose. St ee eee 143
AMIGVSIOMUM ssa 2a ee eto 22, 383 Dispharagus: ..o2c22ses>-e2225-2-= 238, 276
@heilospirura= <2 ee 226, 389 Dispharynx-_-....--.-- XU, 237, 238-240, 241
Epomidiostomum.. -_-------- 27, 384-385 USUI coos osname eu cadee tet Sees 143
Heterakis -.- ses Vil, 52, 71-72, 73 Histiocephaluse.............2...-.=--- 235
solstitialis: Conurns=.<=24s2s22.2 220.222. 80 Microtetrameres-_-_------- XVI, 352, 360-361
Somaterta dresseri==<- +=2282*9e% => ses 20 CNG MCT ES ssn was aaen aoa een aac eeos 360
MOMSSN asec Ae eee ok 24, 372 TOP MOCEN CG a= ane fame na cn kane 360
Sorex: tetragonurus} 22-2252 -2=225.225-2-25--= 138 | spiralis columbae, Dispharagus__.........-.--- 238
spadicea, Galloperdix::*s:..22-.2..---.-.--- 66,116 || Spirocerta.2. 22-222 -cecen-eceoce 163, 164, 202-203
MOS DATTOW. nak oi Cts eae Se eet swans 389 sanguinolenta..........-....------ 203
species, Acuaria ( Cheilospirura).........----- 229| || Spiroptera .__-.........<- 203, 205, 206, 207, 212, 216,
ilaudars 202 tee Seah ete OE. 139 219, 231, 233, 237, 244, 258, 260, 262, 272
IATOeR seat ec = Sette eet 152, 154, 160 acanthocephalica._..........------- 287
a SCOTis: 8 ee Te ea ee oes 324 GQCUNEGE 22sec eeaseseneeaee ase 253
UCC lS See ets Sn eee 115 WHINE 2 dccsnccewc Sscesectesnnooe 234
Caprimulgns 220-2 eet oes 130 apinis......-. Wsaeesaeceast ee oeee 309
Cry piiris:(.525 ke a canes 55, 128 COM sscswecececece ees 165, 240, 281, 282
@rcaltiss = 5225 ose ee eccesese 297 GRACONIRUG: 2 cee -5 525. se eee 328
460 INDEX
Page Page
Spiroptera anolabiata____....-------------.-- 317 | Spiroptera semilunaris -........-- s.22ssss5 171, 205
Onthuyige 22 oo. eee 217, 218, 219, 220, 330 Stereura 22-2228 316
ardeage-Maguart. 2-2 ee 255 slernge 22 oo a ee 287
atientiata. 2 oo. Sean ae 220 sternae hirundinis Poses ee 287
DECOLOT eee ee ee eee 371 Strigiss. <0 5222225 Sareea ek 325
bicwspisi i. 22 eee 286 strigis-atricapillae____.__.-...---_. 205
Dideng ness ae Soe ee ee a ee 288 flammedeé: 22 228 ee es Ss 205
Dslabiata oe eet ee ee 307 OTISEOLEE sone eh en eas 205
breorpenig ss 2 Fo Ee ee 332 lineatae!.- asses ere ans 205
brevisuoulata. = ae ee 325 sygmoidea..- 2. ta eee a 330
DUllosa eee ae eee 205 tadorn@e: 2-22 ee ee 376
Capillarige o-oo. 25. acc eee 287 tantal? rubri_.. 2... 2. se 261
cephaloptera.. 2. eee 322 tenuicollis _ -... = sasocsenee SEZ 207-208
ciconiae-maguari_ _.--- 5-222 _- 256 totant.. ..- avd petise eT Ape 261
cirrohamata 2.2 = oo oe 266 tricolor... -=— nates ite 2c eee 376
COLGNGIGs conan oon coe oe == 293 tridentate... =. s-toeeee ae 271
COMECEIONIS 2 =o ee ae meee 122, 213 truncata. 23325 8.Fe ath. oe 190
CHOC oe een ee ee 331 tulostoma= - 2. 2 tee ee EE 184
ChOCIS QLECLOTIS. 2-2. ohare = 317 turdi. 2252 ea tae XII, 202, 206-207
ChaSssicauda.--- -- ke ees 267 ambellifera. 2-2 5_* ele es 261
crotophagae Gni__....<.--.=----2-- 323 auncinata 2 = 222 setae N_ 246
MOAIONIS: oo ae 323 UNCINIPENIS _ <a ae ee 165
GONEICULOG ee on oa a Ne 288 anialata 2 2522s 2a ga 186
CLONG OG eee aoe ee es 259 anilatenaus . 8 8 ee 2 186
CIMIMEN E218 22 wos ers Doe ae 325 UDUW DOE Oo oS Sante ee 190
eunvoplerd= <2. = 2s cen eee es 201 DONEUG 2 8 oe Cee a 260
CL CLS epee eee a ne ea ee pate 167 oulvoinflata = soo — 2. eee ere 294
TOUCUN CU es Se oe Se see hs 249, 250':'|, SDinpre Wie hye Ee ee 163, 164, 165, 392
UCONN AG em a tue tote ane ee eS 280, 317 gastrophila.. 20.2 51 = ste etre 165, 166
falconis gavial realis_._.....------- 317 talpag: eto a Le 164, 165, 392
lepltODOd1S = ae os eee 317 uncinipenmis. +. =. 228 X, 164, 165-166, 392
SUUDUULEON IS ape ote 275 gschokKeie 2 fos. eee 164, 165, 392
PULICRE re Coe a seta wetter e ah Canes 25 |: Spiruratas woe es: 3 pies te od 1, 5, 162, 208, 366, 392
GOLINUe eo en eee a 253) SPITUTGAO see ee eee oe 162, 163, 164, 290, 392
glareolae austriacae_....-.-_------- 288 | Spirurinae___------ 163, 164, 172, 187, 191, 201, 202, 392
NOMMULOSE > o2 eee re eee 226 >|) :Spirunoidea =: 20 === fos Sree 2, 162, 163,
Weleroclita cee eee eee 331 208, 252, 295, 311, 333, 361, 363, 392, 393
RORY ICG oe ec oe ote ee ar Ser ee 254: | Spizaetus mauduyti_...-2-—sseeet ee 297
AN CON aoa en oe wae we Ee 175 | Spiziaster melanoleucus--_-_----...----------- 297
ANLOLDaa ho ws een ee 305, 335, 357 '|- SPONDS, JAMAS.= 22 2-2 a ee 59
LEUYCCE OU G tmeerae tanr 171 Lampronesa. = -2) == === es 59
taticnveale= so esos ee een 290\ | squamata, Acuaria_.. 2 2-22 t pote oe 257
MUICEDS hae a wee aa ee 276 Callipepla- - == --seseseeee sso 88
LEDLODIEN Oe seen Soe eee a 176 Echinuria___.-- XIII, 244, 253, 254, 267-258
MONIUISUTEOL Os nes 2 cee ager 178, 179 Milani. <2. 22325 See 257
MLNS OT ooo aa ee ee 325 Fqmanni se 2222-2 See 258
TMeEGiOSpInAliss see ae ee XIV, 294 | sqguamatus, Dispharagus_-_-_----.------------- 257
MILE CLOSLO MU 3 mene he ee 300 | Squatarola helvetica__..-__..--__-- 139, 144, 285, 286
microdactyli margravii____.-_--_--- 332 squatarola) 2). Jo go 8S oe Me 285
MOMOL OTASIIENSIS oc ho 322 |) -squatarola, Squatarola.--<- ose 2 285
ASUS = St cee stn ne eee me a 237 | stelli-polaris, Streptocara___..-..-..--- 265, 270-271
ODUVELGLE ae er a ee anen,) 235. | Stellanis, wArdea. - 2220. No hacks Bie gs ae 152, 274
ONO sees Sete mal cee nn a eR 322 Botaurus 2-2 2 ee ere 152, 274
MUL DIELO SU me meena ee ee ees 317} Stenopsis candicans ~~ - "<= 2-2 Sawa 130
DCCLITUNEN Cc ene ene a ae 266. Tl iSiephanuruses 5-222 — 2. - oon ee ee 29
PONIOVAMALA ee eee eee XU, 2056-206 | stereura, Oryspirura___--------------2------- 316
DETIOTUNS Senne era ee ee 226 Spinoplera=.2=---- 225. ease 216
DY SOUL Dene an nen eee eee 298 DWHOLAZIG- 2 hon! See ee XV, 312, 316
DLOLUe0e-0j Oj Ol. eae 241>) (Sterma arctica. --25 = ees oe oe oo ees 235
WVOCEWATIOC te eee see 263 CaSDita = 225-2 ase a ee es 287
procellariae anglorum__.....--..--- 263 hirundo<.. =-2:<22_ +021 coe eee 287
TAC ES re ae ae tie ea ree ea 343 DIPTA So oo an I eee oe ee 259
NCCE NE? SOTA ns enn Ne ee 269 TISOLVIA. c= =< soos oc see 235, 362
TEDOUL ee ee ee ne ee en oe 232 SDECIOS2 = 2 5a5 2-82 eee ee eee 285
quadrilonr 2 oa eee 215. Wislepnae, SPINDLE E.- 222 =< eee 287
SAVITIGC A Pose ee ee ee eee en 207 | sternae hirundinis, Spiroptera__-------------- 287
SUIUUIMOLEN =e nee ana ee 203
Stictoenas'srquatrix= = ---- = eee 87
—
INDEX 461
Page Page
Stomoryst fo... eke Ree 162 | Strongylus orispinus_.........-2--.---------- 28
strelnikowi, Ascaridia_____.--------- vil, 79, 99, 100 papillatus._ 222.2... 16
Streptecara_________-_ -_-_- 2 <2. ad e221, 264-265, papillosus.. 22222-2202 SS. 372, 373
266, 267, 268, 269, 270, 271, 391 Pergracilis.. sas IAS Ce 9
cirrohamata_____----- XIN, 265, 266-267 Primilvus..csrcccocase Sth Ke 34
crassicauda____.--.--- XIU, 265, 267-268 quadriradiatus......<-s<...-22--5 12
crassicauda anseri_..----------- 267, 268 SEMNAWUS oc ccs scsceeson cess LB 10
charadrii____ x1, 265, 268-269 SPeOClOS 0 sedate 43
ecora == a wee XII, 265, 269, 270 SPICULOLUS ooo crds 77, 128
pectinifera______---... XIU, 265, 266, 267 tenvises 3h ee 10
FOCba «eae sa aes XIN, 265, 269-270 trachealis..... A232 Re Sas 34, 44
stellae-polaris______._----- 265, 270-271 tubifer..... 2 ites ee 367, 370, 372
triaenucha________-__ XII, 265, 271, 272 tubifertantali- a. 2 sscase fs 377
tridentata__--_-----_. XIII, 265, 271-272 UNCINGLUS . 2-2 scosss estes. 27
striata> Physaloptera.._ ..-2-22/2--22222d2e224 167 variegatus. ose 44
Sstridtila, Strix. _.<-=.2---<-<-.-.--22284e¢én 138,144 | strumose, Filaria__._....--.----------2----- 164
Serigiforme. = 20-2 22 ean eins Sehis 1270) (Struthio australis! 2. s-css2s22255 Eh tt 31
strigis, Spiroptera._—.-.--.-----~-eesasesie2e2 325 Camels: 552s 3s ssass oe SER 14, 31
strigis-atricapillae, Spiroptera___--.-.--------- 205 molybdophanes..........-.--..---- 3ibe
strigis-flammeae, Spiroptera___.-------------- 205 | struthionis, Codiostomum___.._.-.-----_-- Vv, 31-32
strigis-griseatae, Spiroptera____--------------- 205 Sclerostoma@-..2 2222922. ee 31
strigis-lineatae, Spiroptera__------------------ 205: | Sternus vulgaris......-...-22222- 34, 35, 139, 144, 206
simolatus, Bucco: .-.-.----=-2 2s sosaeeats 128,191 || stolosa,, Heterakis ....22-+-22402222-220 9222222 67, 6&
WSDItOn seksi see a 115 | styphlocerca, Ascaridia_-.........-- vu, 78, 100-101
Strimalbai-. -6:... 222062. Stee te 276 HMeterahkis ss -.c2.czssest oeae ence 100
albomarginats.<...-..:---2esasstf..22- 205 | subalata, Physaloptera_-_--.... XIV, 297, 302, 305-306
geucos 225.5255... s -eesseeth eke 138,144 | subbuteo, Falco.....-.-22-2-2bs4ie122 177, 273, 275
atricapilla...-..-..---ssntss's 107, 205, 207, 325 Heypotriorehis: --=<-22:2s222220-"22 177
brachyotus: —....2s:.2<. ese neestlee 138,376 | subbuteus; Paleo.2:.<.<:-----ss:s:s2:ss222¥ 299
DUO: -- 22-25-2522. 25-feea ene 138, 144,276 | subrecta, Acuaria (Synhimantus) -_...-------- 282
GASVDUS 2. oF 22 4 ieee sce cokine ek 138 Synhimantus__--.-__- XIV, 273, 280, 282, 283
Hammes 2220222555 -22.2= 138, 144,:205;.273, 276 | subula, Acuaria_................--- XII, 218, 224, 225
priseats:i2. = 22520222 =p wabeeeestse 205 Dispharaguescs 2. seas 2 224
MiVes..--2 22520 265 eesti k- V44, | eubulata,Ascarise22 sso cscesscccce tt eccce 128
NOCtWa 222 as see 35, 144 Heber AKG in: = seston eee 128
NYctes..=.---2----- 225 -.s ess 138 Subuluraties. 2222 eS 105, 126, 128, 129
OUUS ee eae ee 138; 177 | Subulura---.....---.:- 49, 104-105, 107, 108, 110, 111,
MASSING. = sca e oo eS 62 112, 113, 114, 115, 116, 117, 118, 120, 121, 122,
species..<22 2.025... -s.co ee ee 118 123,124, 125, 126, 127, 128, 129, 130, 131, 133,
Stridula-o2= 2-5-2282 ee 138, 144 134, 388.
fangmalmi _12- ts ee 82 tS...-5.-. 138 actiticatida=.~ 42.2.2 IX, 106, 133-134
torquata s2. 2 ht 2te G1 22i..2255 207, 335 acutissima---------- vill, 106, 107-108, 109
stroma, Ascaridia_-----_....--222.22% vull, 79, 99-100 allodapa_._.-.-.----- vitt, 105, 108-110, 126
ereteyvakis.. 2-22 ono SP eSeB si. 99 bentocrtzizss+----- 22 vill, 106, 110-111
Strongyiste.2.22--.225=.--2223-2.S3834 1, 5, 7, 29, 392 prumpti: =~ 5.2 28%: vul, 106, 112-113, 131
Strongylidaess......---.--...cesles22= 6, 18, 29, 30 Garlosinsi@ 2-2-2224 vill, 105, 113-114
strongylina; Ascaris._.....-. 22)28202222.-.-- 77, 128 Gurvata:-s---:--:-.2s vult, 105, 114-115, 116
Physaloptera::....2322t223..-- 297, 309 differens....-.._...-- vill, 106, 108, 111-112
Subulura____--.---- IX, 107, 128, 129, 131 forcipata_... vit, 106, 111, 175-116, 117, 126
strongylina cuculi-seniculi, Physaloptera_..... 107 galloperdicis.....------- vu, 107, 116, 117
Strongylinses ........---.-2--2fUL0.----- 30 PTaciliSe 2.22 25.08 eee oe 1X, 106, 184-185
Strongyloidea....---.---..-.-tieee3.. 6, 7, 29, 33 halif seccecesssrect vill, 107, 117
Strongylus 2222225. 2. PR CES 82t_-_- 29, 30, 272 leprincei_____----- Ix, 106, 111, 178, 119, 126
acuticaudatus__-_..---.---. eee 77 itz o<2 23 s2osestec8 Ix, 105, 118-119, 120
cubits PIN SSO SES See bee. ce 24 multipapillata:-........--------- 888-889
ORBIGUUS ce Ssacs ro Ss. ce 287 noctuae___.__._.---------- 1X, 106, 119-121
ONSETIS =. -scnecce cnn esos... 19, 20, 28 olymipidig.= 222225-2-- =. 1X, 107, 121-122
ardece-stellaris_oie se 222 £2. 274 papillosa........---------- IX, 105, 122, 128
douglasigg<<< 2 AUS 14 Miobinast 2. se ee eee Ix, 106, 122-123
QOuUuglasstt = foc. o52~ = es 14 poctium?...-_.....22--- 22... Ix, 106, 128
CeGGNS ae eent te 372 reclinatas::....2205-.2---* Ix, 106, 124, 125
astatiie 02-525 EMO 17 recurvata_...--------- 1X, 106, 124, 125, 127
horridtse: o222---2 ee sees 253 rirnatee ss 8528 2" se IX, 105, 125
METJOTWIN sss dors cesse 5 FSS nee 372 PA aa eee ae IX, 106, 125, 126
MON0OGON ono Ee none 26 seurati_.....-.------- IX, 105, 111, 126, 127
modularis._--2--22 55h. ne 20 similis-__......------- 1x, 107, 127-128, 129
Modulosus.-—aa2~ sees eens toons 19, 20 SDECIOS Ne sate se hoo nese ee 106, 182
462 INDEX
Page Page
Subulura strongylina.-.-.-..--.- 1X5107;. 128; 129513" |i stadorma, Anas. 2222-57222. oe 53, 62, 267, 376
subulata--.:- 23... 4. se 105, 126, 128-129 ‘adorna...-.----.- 4 41, 42, 53, 62, 267, 376
suctoria-_-. 1X, 107, 112, 125, 127, 128, 129, 180 | tadornae, Cyathostoma__..__..__-.__-. VI, 41, 42-43
trayvassosi_ 2 ees 1x, 107, 181, 132 OCTERO SLOT SPR aoa ee ae 42
trogoni. 2.22. 222 IX, 107, 181-132, 133 Spiroptera-3..5-. SARE 376
Suabuluridae: 2.2... ..--_ st eee 391 Syngamius2 2225... Bee 42
Subulurinaess. . 2... 1.52. eee 49, 104539% |}: tatgoor, Turnix® “22 22 arene 2 328
suctoria, Allodapa..... 2...) 5 Samages 2 1125130) ||) Palegallusilathamics- ios 85
HHeterakis'.5 2. 2. asec 108; 1125130: | Palpsieuropaca:: --255..- 2. ee 138
Subulura.-. Lx, 107, 112, 125, 127, 128, 129,180 | talpacoti, Columba__............._._-_.----- 87
sulcirostris, Phalacrocorax-....-...-...-----. 148° |) falpaesSpirtrat_.--.--- --<.2-.-. 164, 165, 392
sulfureus, Pionus (Psittacus) ......-.-...-.-- 80 |) tamatiatBuccoX222... esos eee 131
Surnia passering= o.. - | teenth 62 Capito)... 225 pee: 115
swainsoni, Buccos2--222.- ==. 4 128,131, 132; 352: |) tamatinaBueeo-..-----.-- esas 2 128
Gampsonyx...=<-) Steins es 297 | tanasijtchuki, Oxyspirura___-_- XV, 322, 830-331, $32
Pternistes. 5. ==... = eee as, 130: | tangmalmi) Strix_.-...<-.....-.... 222 138
swainsonil, Kalco)... ....-. 2. seeders 297,299 | tantali rwbri, Spiroptera_......-.------------- 261
sygmoided: “Acunria....- <=... .23- seeeteeeieees | 282 | Tantalusilocunlator...-<---...-.-_ see 154
Oxyspirura_...---- XiWI;2207822, $90; 682 | tarda, Otise 22... 2 ee ae 10, 16, 34, 53, 67
Spiroplera ... sees see aes 330 | Tarentola mauritanica.._.2-.--_..--...+----- 216
Synhimantus....<2.<:~-2!-2 272, 282-288 | tarentolae, Acuaria__......--..-------- XU, 212, 216
sygmoideus, Dispharagus_...-...------------- 222: | tataups, “Tinamus: 2.2.2. see ae 128
SulVaticus, Purturs..-..<-.----- Loe 86: |. tatuapa, Crypturus 22... -2...- Se Ae 128
sylvestris;“Bonasa ..2 500 cathe See 53: | tectus: Buccot! exo. 2 1. Joe eee ee 131
Sylvia rubeculai?:....... Basse ee 224 | tenebrosa, Chelidoptera_-__...-.-.--------- 128, 131
turdoides2ss. oo. et ae 139, 224 Monasaiji cs: 22 ae 115
Symbranchus laticaudatus...........------ 368, 371 | tenuicauda, Heterakis_-...-..-------- vu, 51, 73-74
Syngamidae! eee... oe ed 6, 29, 33, 34,41 | tenwicollis, Physaloptera.....-..-------------- 207
SyNgamuse sess ooo les ee 29, 33, 34, 35, 38, 39, 40 Spiroptera.2.2 uc. - 2s 207-208
boularth: 2oh2 ccc sdeeee iets 44 | tenuis, Acanthophorus_...-------------------- 343
OPONCHIGHS) 2. 2 ak Utes 42 PA Ctiariat meister ee 217, 218, 225-226
coplebse eke 22 ke ce ee 34 Dis pharagus.. 5.62 22e55 4. ee 225
gracilis. 2.223 es VI, 34, 35, 39-41 Strongylussco22cesoec eR ee 10
LOTR oe ee a epee 43 Drichostrongylus:.+222es-s524-e6 Vv, 9, 10-11
microspiculum:.-..--.b.=2224 VI, 34, 38 Tropidoceréai.222.2. 2 eS ee 343
mucronatus....:...=--=2s3 Bhs ee ih. 34 |) ‘Berpsiphone:species: ...-=-==.+2<.-2.--2 2293 188
PATVIUNS2. 553 oS. oc csecese es VI;34;:$9540' || Terranova... 5222255. Se. ee 135
primiligusss: cee bss same Y 34. [oO Wletracanthte=. .-...=. 6 -25-se5ctee aa 284
pugionatuse=..s2_264 Ses seb $4 |) tetragonurus, ‘Sorex::.........--2222225522225 138
tadOrNnade so 52 aos ERO 42 ‘| MIRStrAMOreS 22. oc se eeee e S 333,
trachea... VI, 34-38, 39, 40, 42, 43, 44, 45, 47, 48 334, 336, 337, 338, 339, 340, 341, 342, 343,
trachealis.....<-- os nssteteeee <= = 34, 39 345, 346, 347, 348, 350, 351, 354, 355, 356,
parniegatis. oa oe to ee ee 44 360, 393.
Synhimantusss) 2... csc ceca eet xi, 211, americans. —-22<- 222 XV, 334, 337-838
216, 233, 256, 272, 273, 274, 275, 276, Cette. oust s A ee 334, 338-339
277, 278, 279, 280, 281, 282, 283. cocciness.. +~=-222<-L = XVI, 335, 339-340
SMNIS 3. oo. ce Seekers XIV, 273-274 cochleariae__._-.------ XVI, 335, 340-341
Glateee 2 5 oe tees eee 281 confusa___ XVI, 334, 335, 338, 341-342, 343
brevicaudata__.. XIV, 273, 274-276, 276 Contoyia... 2 ee ee 353
denticulata_-....---- XIV, 273, 275, 276 cruziiniocy So. sbseeteet sau s es 333, 352
Glliptica-~322-2-2525-~ XIV, 273, 277-278 Guia. oe ae XVI, 335, 342-343
hamata.ccs--2s--sse XIV, 273, 278-279 Nssispina 5223-5 5-- eee SISO)
invaginata.....-.---- XIV, 273, 279-280 334, 335, 336, 337, 338, 339, 340, 341, 342,
laticeps:2 aos oo ae XIV, 343-845, 346, 347, 348, 350, 352, 352, 353,
272, 273, 274, 276-277, 278, 282 354, 355, 357, 358, 359, 360.
longevaginata.—.. -2 2 256 gigas. .< 00). 3 oo eee XVI, 334, 335-336
Taiiie tis c-c eet ees eee 273, 283-284 plobosac 2-2 2- oc). see 334, 346
Pectae see c eas XIV, 273, 280-281 gynaecophila__.-..---- XVI, 334, 347-348
Sagittata.........-..- XIV, 273, 281-282 haemochrous ------- eosaaeenn ae 335, 357
subrecta__......- XIV, 273, 280, 282, 283 inermis....-...-.-sdessssede--= 3407855
syguioidea!-2 = 22-2. 272, 282-288 inflata. ...--.------<-sdassue----5 397
SYENIs AlUCO een ce ee al wie ee 144 micropenis.... XVI, 335, 340, 342, 348, 351
MV ClGS to oe Soe oan aoe sues 144 minima..-.---.2..- shee 358
RPA YPAtesIAQ Mae. ose ose ee eee 148 nouvell: |=. 523. - 4.8 XVI, 335, 348-350
Tachyphonus cristatus brunneus-_-._.---.--- 358 Paradoxa. = --.-.-- 2585 XV, 334, 335-837
"Tadorns pelonil 222. cs ee eee 376 pusilla 2 22... aetetey--2<- 359
tadorna e222. 5 -25--2 41, 42, 53, 62, 267, 376 spiralis SecerssoronhRaeeeeencewe - 360
INDEX 463
Page | Page
Tetrameres tetrica__.--...-------. XVI, 335, $50-351 triaenophora, Spiroptera__.......-.-.--.----.- 274
UruvdSS0Gten aoe ons Sree ee 341 triaenucha, Acuaria.___.._....--.-------2 8 971
ONISDING 2 8 ese ee _-. 334, $51 IPO oe ome eee OT
gakharowi..=2..------2<s2a+--- 335, 385 Streptocara____-.---.- X11, 265, 271, 272
Tetrameridae____--__-____----- 162, 163, $33, 351, 358 tricholor, ‘Herodias... - 2.) 22-3 tects. coca 152
PPetrao DONGSIa ~~ aoe ete ee 53,81 | Trichostrongylidae.-_-.............---..-...- 7, 8,18
WAZODUS. 2 2 see ena2e aac sane eS §3, 84,88 | Trichostrongylinae...........-.....--.-..-- 80.11
LO Gite oe ee 17, 88,95 | Trichostrongyloidea_____-..-._..-._.-..-- 6, 7, 8,17
WTOZALUS. sce ees sees oe 35,53, 78, 81, 88,90 | Trichostrongylus............---....=..=.... 89,10
PANU eee oe ee oe aan a eee 115, 128 CGOWGUGS see eee 14
Tetrapteryx paradisea_._.......-..---------- 99 modularis._.......--.---.-- 20
Retrak. Obis- 2:5 seo ano ee or. Sk 53, 67, 117, 290 pergracillis............-... v, +10
tetrica, Tetrameres______-_-------- ¥VI, 335, 850-351 quadriradiatus_.......22.2. 12
RGUIee a VTUTUS= 2 22 eco ee ea Aes Fee 17, 95 retortaeformis__._..-....-- 9
PROULBOMS oe Soe e ee 17, 88, 95 NONUIS = 22 a eee Vv, 9, 10-11
Thamnophilus funebris__.-_----_-----_-_---- 207 | “Trichosyringata..... 222 -2.s-22s-2222-- 4, 391, 392
Thaumalea amherstiae___.-_-.-------------- G4: We inichuratge: 2° soe sas So on a ee 2, 5, 392, 393
OUSCULUSS oe re ce ee Sek 65 | tricolor, Hystrichis_. xvn, 375, 376-377, 378, 379, 381
IC LAME ae ee ee a een 64, 65 Spiroplera- 2. =. Sa ce 376
Thelazia____--- 311-312, 313, 314, 315, 316, 317, 318, 319 | tricuspe, Contracaecum _---.-.---- X, 147, 160-161
anolabiata__-__._.....__-- XV, 312, 317-818 | tricuspis, Kathleena__.............--.-------- 160
campanulata__.-.---_--.----- RV cl2 sio. |, tridactylus, Marus_.-.. 2222 2.22.2. sa nse 148
cholodkowskii-_-_____- _-.-. XV, 312, $/3-814 | tridens, Histiocephalus-_---.--..-...-- XIV, 291-292
CIETULS = eee seh ac ee XV, 312, 318-819 | tridentata, Filaria_...............----.--.--.- 271
dacelonis_...--.......2--_ XV, 312, 314-316 ISUITODLERG ste 2 ee ean ee 271
GIGILAtA = 2 enc s se. ~ ee comes XV, 312, 319 Streptocara_-.....-- a..< XU, 265, 271-272
MG 7 tae een eee oe XV, 312, 816-816 | tridentatus, Faleo_.-..--...--....--.--- 177, 272, 283
MapllOsae o- a2 ee ene eee KV, 312, $17, $18 | trifurcus, Caprimulgus.-.....-.-..-.-.--<...< 129
PDO GOS teres eee te ae es 312 | trilabium, Ascaridia.=_.............-- vin, 79, 101
BlOvOULB= ee ene Bee eee XV, 312, 316 TTCLEF OK iS 2 oe oe 101
nel agudee 2 = tee Se 1, 163, 871, 319, 321 DTT ee ADIN ee eee ne eee 254
RELA ZIUS a rn Se ssc So eae 311 Helvetica <2. 2 “sss 222 Seseeeecese 285, 286
Thrasactus harpyia........---_-_--.----..--- 317 Wanellits cote Se eee 141
Tinamus:species.:----...=.-.=.-- _. 55, 71, 72, 99, 128 VATIADIIS 35-25-22 ones hee 254, 285
CSC AU DA ee ete eee 128) |), Tringae vanelli_._-s- 222222 2e<2 6-2. S25. 260
tingazu, Cuculus.<.¢ -22-..-5-20 casas 115, 128,129 | Tringoides hypoleucus_-._..---..-.-.--------- 235
tinguscu, Cuculus_....;--..2--2--..--2s-2i-. 207 |}. 7Trochilus\ochropygus.__--.- 2. .---h 225-2 294
Tinnunculus alaudarius--.--.--------------- Door, Trogon Collaris. - 2.227 22 asec ase 171
tinnunculus, Cerchneis__.-.-.._------------- 276 MEIANUGUS: 22 sew eee 171
ALCO me eee ee 138, 177, 183, 276 SPOClOS Sos ee eS Sen oe ame 110
DIMUIS PA CCLOIGCR =.<a 9202 aaa ae oes eee 297 Wanlegatuses 2-2 ee none 110
(0) Ot: (Cc: eee ere ee a 147, 235, 267, 372 SAP YS a a es ee 131
Rta oee es eee eens 148 | trogoni, Subulura.--.--.-...--- IX, 107, 181-132, 138
torquata, Malacoptila_._.._--------------- TORTS Tey MUROUGH WLS = onan 148, 372
WWONSSA = ee ee ee 115 STO COCO CI eer ee eee ee 334
URLs eens eee 207, 335 ert. oe ae nek Soa oa 338
NOTQUaAbUS) UTOUS 8.2 U5 52s soos ones 258 189 COCO CW. Pane a sheen anaes 339
torqueola, Arboricola- ...-=-2<-2-<2---- 22-224 387 COMO ie oa 2 oe Oe ee 353
LOL SDITOPUEN OS. 2. oo asa a sews ae t- = 261 WISSISDING soo coe ieecrwcc—oses 343
MOLANUS USCUS..<2<2-2--sciacsmasccssc-—5--5 235 GlO0GS@..- >. Jee eee eae 346
PlOu Gea ae ee eee! 285 gynaecophila______-__-.._----.-- 347
INV DOIGUCUS ce == 32 eee eces oa ak 235 gynecophila___......-----------. 347
MACUIOUUS eee ea eee ok ae 235 INET IS <a seed eins aso 355
melanoleucus.-<c22s<--254.ese50.0% 261 PC 2. a ouee SOAs nan 335, 357, 358
trachea, Fasciola... .<.02--. se 5-5-2 a 34 MOUVEM o-oo co cccccccwcnccccnns 348
SVN PAMUS cee ee eco ese ae se eb ae VI, paradora..._..------- 335, 357, 358, 372
34-88, 39, 40, 42, 43, 44, 45, 47, 48 SDNGUES. cet. cusses des epson ee UU
tracheale, Sclerostoma.._...------------------ 34 LONNIE St Ses 2 ac ete ene anee 343
trachealis, Strongylus.....-=.\..2..------------ 34, 44 WNISDING oo sc caasudesntoceses . 361
Syngamus. oe eee oc eee 34,39 | Tropidonotus hydrus__--.-...----.---------- 168
Trachelotis senegalensis -___--------------- 189, 291 TTODIGUTUS! 5b 2c8 2a ce acdoas bkteeeaee = 2s OOe
trachyrhynchus, Pelecanus_------.---------- 148 TRODIGUPUS Oe oe oS Seco cena see tee 334
WTragopal Satyra-—.-2 04-52 5226 2 ote te oe 57, 64, 66 PSSISDINUGc 2 < os core ne ae eece ee 343
tranguilla; Mouasa-+-23._- 2 22225225.---.< 115, 128 infldtus. 9... 2. ceeste ees senle 357
travassosi, Subulura_.__---_- eee aee 1X, 107, 13/1, 182 MUTadorUs sc Ioosoehaeaet aces ewes 335
TelrQmen eS... a ote nen seco eas 341 | truncata, Ascaridia___...----------- say 80
3612—27——31
464 INDEX
Page Page
UMN CULO, NA SCATIO one se nes en ep eae ney 80)" | “Urocissa occeipitaliss) 2222 ea eee ye aE DO
USOT yee et ooo eee eins nes 80) | huogallus) Wetrao see 35, 53, 73, 31, 88, 90
Hadielinee aoe oe eres 1X, 187, 190-191 (DEFT ETA BX) 7 V0 op me MURR eo PAUSI RICE) Sr paren > PERE 113, 128
TELLER RASS wer cee cee eee 80) "| Urubitinga urubitinga_-- 225-22 = eee 297
Physaloptera---------- XIV, 296, 306-307 | urubitinga, Urubitinga_..__......_______.__- 297
DIODE co eee ee hae ok 190)"| SurabuyCathartes:..2 2222-22222) 2s ee 335
truncatus, Dispnaragus.--.- 2-2 ee 190)" urubitinga, Paleo. 222-2 > 82 eee 297, 299
tubifex, Hustrongylides_------ =- =. VS | UnumUtiwm, Craxc2 8 2 22 ee 231
367-369, 370, 371, 372, 373, 374 INOtnOCTaX! ooo ee ee 331
Bustrongylus se en es 367,371, | untitas, Caprimulens._-. >. =e eee 128
FLY Strichise Sosa oe. eee cote eee 367" |-uputan);Caprimulgus! 222-2 a eee 115
EDISE Sain eee apes are oie heme eee e 377, eutamaniatordas ses: ose ooo eee 148
Strongylus--—- 22222 Le ae 367,370,372 | raldemucronata, Ascaris__-------------------- 74
tubifex tantali, Strongylus............-.------- 377 Heterakis: 2222 te oes 50, 74-75
LULOSOM A KE ALAT ae ee ee ee 184,185" :| valvata; Heterakis:.--- -42-saseos soe vil, 50, 74
Habronema---_--_---- KI 173, 180, 184-185. | wanelli,, Acwarid. 22 - 5+ cos yee ee 260
SDINODLEr Ae ote ee ee ee mee 184 Rusguniellas-- == Se ees XIII, 259, 260
LTR AYE ST 7 as eA a Seer ee 206 | SDINODLEna scene ee recs epee ee 260
Spiropteras es ces set eee XII, 202, 206-207 TBringae. wre fee wee eok kes ea eM 260
turdoides: -Salicaniags.- ose c ce Ne eee 139 | Vanellus cristatus__-_.--...._._- 22, 139, 141, 260, 268
Sylyiaesss: save ec soe es 139, 224 melanogaster. _ 22-22 ==-beeeeet 139, 141, 286
sRurdtis iliacuguessscs. ccnses aa tue Nba ee 139,206 | vanelluswPringa::-<4o2- 0. + sen ae ener 141
MOP Was a cse ee eo eee 139, 206 | variabilis, Heterakis__-_---.-.---.-------- 386-387
IMUSICUS 2 eres ne eee yatecias 139, 144, 206 Mrin gays. ute PUSS eee eee 254, 285
Dilarises ose eee oer eee 139, 206 | variegatum, Cyathostoma_-.-_---__------- VI, 41, 44
FufVentris 22.328 o 22 Uohe soe es 359) |) vaniegatue: WSErongylus.. 9 stesoee e e 44
SAKALINIS oS o eee aeons ee 139 SYUNGMINUG oo) LeeLee ae ae ecto ne 44
CORQUS CUS! nets oe Oe ee eee ee 139 Trogone 22 one eae 110
WASCIVIOTUIS 2 oe ne eee ae eons eerie Rie 159 | varispinosus, Hystrichis___-_--_- XVI, 376, 381, 383
AT fF Ce ct i Se oe 4,295" | |‘ Veneratus; Phasianus. 20: -0.2 22-222 oe 53
ARUEDIZ;SHOCIOS so. asec ee ee ea ORL Aaa are 132 | vermicularis, Oedicnemus- -_---------------- 198
AIG OOLe ates Se eae emt See 328 | verrucosus, Phalacrocorax- ---------------- 148, 266
ATUrCuT SVIVatiCUSs 2 ae ee eae eee ee 87 | versicolor, Phasianus__....._.-----.--------- 53
typica; AlLOdapts = weet eee ena eee ae 108'"| -nestcularis,, Ascaris. .-2 52-22-22 eee 52
EE VCO AM Das pee ee eee ee ee 273 Hetenakis. 200 C2 Ue e eae 50, 52, 67
Wihtilavalucoste es ae neo nee eee ere en 144 | vesiculosa, Ceratospira_-___-------- XV, 319, 320, $21
umbellifera, Schistorophus...........-.------- 261°) Wignlera.2: 552. vee* sooo eee 164, 201, 207, 392
Scisdiocara-=-22--=-cceeee XI, 261-262 QULVODLOTa. 3-5 eee XI, 201-202
SDITODLCNE ssc a ee eee een 261 | vinaceus, Pionus (Psittacus) __.-------------- 80
em pellus VBOnasae ao = 2 eee ee 15;.93, 229, 239} Vinago delalangil\s-- 222222" Sone aes 91
uncinata, Acuaria (Hamannia)......-...---. 246 | violacea, Ardea___---- PU eee ee 142
Bchinurige sos 02s esos anes XU, | violacens, INicticorax. 2-22 2 2- --ee eee 348
245, 246-248, 249, 250, 252, 253, 254, 255,256 | virescens, Protogerys_----------------------- 175
Rl amanniaeeseess case tenon ees oe 246 | virescens virescens, Butorides___-__-----_--- 152
ISDINO DUET Ne hee eee ee ee eee 246...) ‘virginianus, /B UDO. s2c- oon <= ee oe see 138, 144
uncinatum, Epomidiostomum___. Vv, 27-28, 384, 385 Colinus ete ooo Saat 9, 53, 88, 168
UNCiNALULs Dis paanggus. 2-02 eee 246 OFty x25. 2 a eee 53, 88
Strongylus. o> 2220 ee ee 27 -| virgo; Anthropoides_ -°-_=-- 22. = 222 22bee 142
uncinipenis, Cheilospirura.........-.-------- 165\ ||| “yatidirostris; Grus=-2----- eee 44
ISDIO DET hse eats esa eee 165) yanidis; Gecinuss see oc see eee 215
OpirUra soo ese -seeeee X, 164, 165-166, 392 PicUseecesese cess soa eee 34, 215
undulata, HOuUparTa oe ee ea 130, 196, 198 Psophiae soe. oe ee ee 55, 69
ALTGatG. Spiro pent - 2 a, ee ee 186 TTO RODE ines cee es oe eee eee eo 131
lnicinetus,Waleoe sno shoes 17;7,,280%297, 299; 605) ||) WASCLVOFUS, ,LUTGUS"= 2-22 aoe eres ee 139
Pareputeoe see coe ae ee eee 297.5 ‘vitellinus; Ramphastos::=2 = 2 eee eee 237
unilateralis, Habronema_-_-- XI, 172, 184, 185, 186-187 IRnamphastoss.- 22-252. = ae 186, 244
SDITOD ET aa eee ee ae see es 186.) vociferus, Antrostamus----2"-- 2 —-" nee ee 129
wunispina, otrameres-2- 0 ee 334, 351 Caprimul gus? 2222 2. ooo. oo. een 130
RTO DIGOCEN CO arn oe eae 351 | voluptuosus, Pseudaspidodera__--.---------- 387
Uppal epopsssen Meee eee Onn ee ets 190,203 | voluptuosus minor, Pseudaspidodera__------ 388
UDUDAL SDNO DENT 2 ones ca ae eae en oon 190: |) wulgaris, Anguilla_-- 20-2022 ee eo ee ee 146
upupae-epopis, Acwaria_.....-..t.-.--------- 190 Archi DMG0 ee eee ec ae 138
WG DIC EVATUTYG Oe eee oe Mee pele ees aes 220 iButeosce-ee- 137, 164, 176, 177, 278, 298, 316
Upigserollo nse ee est eo en ea een tes 148, 235, 266 Opus 2 eet 144, 276
Car (Ne ie adele se cabal ing 148, 372 Stumusio-- 2s. oe ee 34, 35, 139, 144, 206
urile. “Phalacrocorare 22. uae ee nie 148) 4| avnipinws;BtleOesss ase en eee ee 316
INDEX
Page
Vultur cinereus: .....2 2... 22252522242 -- == 138. |) VSsenlascc--c 2s .-
POU VUS = See eee eee ee cee 138 californica. ____-
MONACHUS! 2-2 - Sas ons eee 138 coronata_.______
percnopterus.-.-- See eee a ease eee oes 184 decora_.___---_-
vulvoinflata, Serticeps_...--..---- XIV, 293, 294-295 | zakharowi, Hartertia__
SDPiIropteras2ascos28- cece eek 294 Tetrameres
vulvolabita, Heterakis.................-.-.-- 387 | Zonoenas brenchleyi_--
wedli, Hystrichis_._..___- XVUH, 376, 379, 380, 381-383 | Zorilla lybica._.-..-__-
wealit; Hystrichta = 522--2-<. 222 S-22<- 379, 381 | zorillae, Hartertia__.___
xanthothorax, Malco. ....-2..-.22--2.22 45. 177 | zschokkei, Spirura____-
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