A Key For The Identification of Genera of The Nematode Family Trichostrongylidae Leiper, 1912 PDF

Journal of Helminthology (1982) 56, 185-233
A key for the identification of genera of the nematode family

Trichostrongylidae Leiper, 1912
LYNDA M. GIBBONS and L. F. KHALIL
Commonwealth Institute of Parasitology, 395A Hatfield Road, St. Albans, Herts ALA OXU, England
ABSTRACT
The nematode family Trichostrongylidae is divided into six subfamilies Trichostrongylinae,
Libyostrongylinae, Haemonchinae, Cooperiinae, Ostertagiinae and Graphidiinae which are briefly reviewed. The key is based mainly on specimens examined by the authors and includes one genus in the
Trichostrongylinae, three genera in the Libyostrongylinae, six in the Haemonchinae, seven in the
Cooperiinae, 17 in the Ostertagiinae and two in the Graphidiinae. Pseudostertagia, Pararhabdonema,
Parostertagia, Obeliscoides and Travassosius are briefly described, figured and considered as genera of
uncertain affinities. Nine genera, namely, Moguranema, Neocooperia, Bergheia, Mazamastrongylus,
Paramecistodrrus, Ostertagiana, Ostertamia, Graphidiella and Trichostrongyella, specimens of which have
not been available to the authors for examination, are listed giving details of host, habitat and geographical
location, together with a summary of the diagnostic characters as described by the original authors.
INTRODUCTION
The family Trichostrongylidae is of both considerable scientific interest and great
economic importance. The comparative morphology, and phylogenetic and evolutionary relationships of the genera of this family to the other families of the superfamily
Trichostrongyloidea have been extensively studied by Chabaud, Durette-Desset and
co-workers. Their work is summarized in CHABAUD (1965), CHABAUD et al. (1970),
DURETTE-DESSET (1971), DURETTE-DESSET & CHABAUD (1977), (1981) and is of great
zoological interest and significance.
The aim of this key to the genera of the Trichostrongylidae is to provide a working
tool for those who have to examine and identify members of the family, especially those
nematodes which are the known or potential causes of disease. The key, which is based
on the authors' own observations and re-examinations of specimens, includes all
but nine of the genera. These nine (Moguranema, Neocooperia, Bergheia,
Mazamastrongylus, Paramecistodrrus, Ostertagiana, Ostertamia, Graphidiella and
Trichostrongyella), specimens of which were not available, are excluded from the main
key but are listed with a summary of the diagnostic characters described by the original
authors. Five genera which have been re-examined (Pararhabdonema, Parostertagia,
Pseudostertagia, Obeliscoides and Travassosius) are considered to have uncertain
affinities.
Earlier workers on the group did not recognize the. importance in generic
differentiation of such characters as the synlophe and the genital cone. The synlophe
was first studied in detail and applied to the classification of the heligmosomes by
DURETTE-DESSET (1971) and the genital cone by ANDREEVA (1957). In descriptions
published before this work appeared these characters were either ignored or poorly
described by other research workers.
Because no single character can be relied upon to distinguish between members of
the group and because of the inadequacy of some of the earlier descriptions we decided
that re-examination of specimens was essential. In preparing the key, we have used a
combination of characters: the structure of the cephalic region, the size of the cervical
186
L. M. GIBBONS and L. F. KHALIL
papillae and their position in relation to the body-wall, the synlophe, the shape and
structure of the spicule, the bursal lobes and arrangement of the rays, the structure of
the genital cone, the female reproductive system and the shape of the female tail.
OSCHE (1958) and CHABAUD <?f al. (1970) numbered the papillae and rays of the male
tail so that the bursal papillae of rhabditoids and strongylids could be directly
compared. The numerical system (after CHABAUD et al., 1970) and the terminology for
the papillae and rays of the male tail are listed below and illustrated (Fig. 1). Drawings,
unless otherwise stated, are original.
Ray or Papilla Number
Description
0
Papilla(e) which appear to be supplied with nerves by the
nervus terminalis surrounded by a simple membrane,
ventral to the cloaca
1
Prebursal papilla
2
Anteroventral ray
3
Posteroventral ray
4
Anterolateral ray
5
Mediolateral ray
6
Posterolateral ray
7
Rays which appear to be supplied by cloacal nerves, which
may be associated with an accessory bursal membrane,
dorsal to the cloaca
8
Externodorsal ray
Dorsal ray
Although the genital cone is a complex structure which is sometimes difficult to
interpret and describe, its value as a taxonomic character in the identification of
trichostrongylid nematodes has been discussed and its different components described
by GIBBONS & KHALIL (1982). The genital cone is particularly important in the
classification of the subfamily Ostertagiinae. The various components of the genital
cone are: the papillae of the genital cone ventral to the cloaca referred to as "ventral
rays" by ANDREEVA (1957), "papillae 0" by CHABAUD et al. (1970) and "ventral raylets"
by STRINGFELLOW (1970,1972). Although usually paired in the genera of the subfamily
Ostertagiinae, in the other subfamilies there is only a single papilla ventral to the
cloaca. The pair of rays of the genital cone dorsal to the cloaca and associated with
an accessory bursal membrane are referred to as "dorsal membrane and rays"
by ANDREEVA (1957), "ray 7" by CHABAUD et al. (1970) and "dorsal raylets" by
STRINGFELLOW (1971).
According to DR6D (1965) a modification of the dorsal raylets and accessory
bursal membrane was first recognized by Sjoberg in Ostertagia lyrata and described by
him in 1926 as the additional membrane. DR6D (1965) suggested that the structure
was formed by secondary joining of an extension of the dorsal wall of the genital cone
with the accessory bursal membrane and called the structure the "Sjoberg organ". It is
also referred to as a "modified dorsal membrane" by ANDREEVA (1957) and a
"sclerotized cloacal plate continuous on the dorsal surface of the accessory bursal
membrane" by STRINGFELLOW (1971).
The terms used in this key are "ventral raylets", "dorsal raylets" and Sjoberg organ.
The arrangement of the various papillae and rays in relation to the cloaca is shown in
Fig. 2.
A key for the genera of Trichostrongylidae
187
GU8ERNACULUM
PROCONUS
VENTRAL
RAYLETS
ACCESSORY
Of
PAPILLAE
BURSAL
MEMBRANE
DORSAL LOBE 1 RAY
LATERAL
BURSAL LOBE
DORSAL
FIGS. 1-2.1. Diagram of the male tail of the Strongylida. 2. Diagram of the lateral view of the male tail of an
ostertagid nematode.
Family Trichostrongylidae Leiper, 1912

LEIPER (1908) erected the subfamily Trichostrongylinae and later (1912) the family
Trichostrongylidae, for Trichostrongylus and related genera. Unaware of Leiper's
work, RAILLIET & HENRY (1909) placed this group of genera in the subfamily
Metastrongylinae. Since then several changes and additions to this group have been
made. The major systems of classification proposed are those of RAILUET & HENRY
(1909), TRAVASSOS (1920), BAYLIS & DAUBNEY (1926), YORKE & MAPLESTONE (1926),
NHVEU-LEMAIRE (1934), TRAVASSOS (1937), SKRJABIN, SHIKHOBALOVA, SCHULTS,
POPOVA, BOEV&DELYAMURE( 1952), YAM AGUTI (1961), CHABAUD(1965) and DURETTEDnssirr & CUABAUD (1977, 1981).
188
The genera of the family Trichostrongylidae are here allocated to six subfamilies
which are reviewed below.
Trichostrongylinae
The subfamily is considered to contain only one genus, Trichostrongylus. The
genera Buckleya Sarwar, 1956, Cobboldostrongylus Sarwar, 1956, Gilesia Sarwar, 1956
and Probostrongylus Sarwar, 1956 are confirmed as synonyms of Trichostrongylus.
Libyostrongylinae
The subfamily was erected for the genera Libyostrongylus Lane, 1923,
Cnizostrongylus Chabaud, Durette-Desset & Houin, 1967, Obeliscoides Graybill, 1924,
Paralibyostrongylus Ortlepp, 1939, Pararhabdonema Kreis, 1945 and Pseudostertagia
Orloflf, 1933. DURETTE-DESSET & CHABAUD (1981) added Moguranema Yamaguti, 1941
to this subfamily.
Libyostrongylus, Cnizostrongylus and Paralibyostrongylus share the following
characters: the absence of a synlophe except for a few discontinuous cuticular ridges in
the vulvar region; short pointed spicules; genital cone with single large papilla ventral
to the cloaca and a pair of modified papillae or rays dorsal to the cloaca and a long
dorsal ray with characteristic branching. For these reasons they are the only genera
retained in the subfamily. The genera are separated by the number of branches of the
dorsal ray and the presence or absence of a modification to the tip of the anterolateral
ray of the bursa. Moguranema Yamaguti, 1941 resembles these genera in the absence of
a synlophe but, as we have been unable to examine specimens, is not included in the
main key.
Haemonchinae
The subfamily was originally erected for genera with one or several teeth in the
buccal cavity, possessing cervical papillae and short massive spicules and lacking a
spine on the female tail. This is now modified to include Mecistocirrus which has long
spicules. All the genera have large cervical papillae projecting from the body surface
and numerous longitudinal cuticular ridges but these, in some genera, are restricted to
the anterior half of the body. Characters separating the genera are: the number of teeth
in the buccal cavity, the shape and symmetry of the dorsal ray and lobe, the presence or
absence of chitinization on the bursal rays, and the length of the female vagina.
Only female specimens of Leiperiatus Sandground, 1930 were available to us and
they have the characteristics of the subfamily. The genus is therefore included in the key
to the subfamily and the description of the spicules is taken from the literature.
Cooperiinae
& SHIKHOBALOVA in 1952 erected this subfamily for genera with a
cephalic vesicle. The genera also have in common poorly developed cervical papillae, a
synlophe with only a small number of longitudinal cuticular ridges and divergent
ventral and lateral bursal rays. Characters separating the genera are the number and
arrangement of the longitudinal cuticular ridges, the shape and branching of the dorsal
lobe and ray, the structure of the spicules, the genital cone, the single or double
reproductive organs of the female, and the shape of the female tail.
FOTEDAR & BAMDROO (1971) proposed a new genus Neocooperia for their species
N. magniovata from sheep in Kashmir. They placed this genus in the tribe Cooperiea
but the species is badly described with no illustration. As it is stated to have a poorly
developed head vesicle and well developed cervical papillae it is unlikely to belong to
either the tribe Cooperiea or the subfamily Cooperiinae. No further description of this
SKRJABIN
189
species appears to have been published and the original type specimens were not
available to us for examination. The determination of its taxonomic position must,
therefore, remain in doubt.
Ostertagiinae
(1947) erected this subfamily for genera with an accessory bursal
membrane. The subfamily name was given as Ostertaginae but this has been corrected
to Ostertagiinae in accordance with the International Code of Zoological
Nomenclature 1964, Article 29. The subfamily is generally attributed to SARWAR (1956)
who, unaware of Lopez-Neyra's earlier publication, also created a subfamily
Ostertagiinae.
Characters which are common to all the genera are: a slight head vesicle with a
small region of prominent transverse striations posterior to it; numerous longitudinal
cuticular ridges, more or less even in height, which arise posteriorly to the region of
transverse striations; small prominent cervical papillae which project from the surface
of the body; a genital cone with a single or double papilla(e) ventral to the cloaca and a
pair of rays or papillae, usually associated with a distinct accessory bursal membrane or
with a modification of this structure dorsal to the cloaca. Characters separating the
genera are: the size of the proconus in the genital cone; the structure and arrangement
of the rays or papillae dorsal to the cloaca and the modification of the accessory bursal
membrane associated with them; the size of the dorsal lobe; the branching of the dorsal
ray; the arrangement of the ventral and lateral bursal rays; and the structure of the
spicules.
The last comprehensive revision of this subfamily was that by DROZDZ (1956) and
this key follows his classification with some additions and modifications.
The genus Hyostrongylus Hall, 1921 was included in the tribe Ostertagiea of the
subfamily Trichostrongylinae by SKRJABIN & SCHULZ in 1937. Later, SKRJABIN &
SHIKHOBALOVA (1952) transferred the genus to their subfamily Cooperiinae because of
the presence of a cephalic vesicle. DURETTE-DESSET & CHABAUD (1977) transferred the
genus to the subfamily Graphidiinae. On the basis of size and shape of the head vesicle,
the small prominent cervical papillae projecting from the body surface, the synlophe
and the structure of the genital cone this genus is placed in the subfamily Ostertagiinae.
The genus Spiculopteroides Jansen, 1958 was considered a synonym of Apteragia
Jansen, 1958 by DROZDZ (1965). Although the two genera are closely related there are
some differences and the genus Spiculopteroides is maintained here as valid.
VAN DEN BURGHE (1937), in his description of Ostertagia okapiae, separated his
species from 0. rubida (now Hyostrongylus rubidus) and from 0. trifurcata. The
taxonomic position of this species is discussed by GIBBONS & KHALIL (1982) and is here
briefly reviewed. BAER (1950), in his review of the helminth parasites of the okapi, had
difficulty in placing the species with the genus Ostertagia as defined by TRAVASSOS
(1937). DR6ZDZ (1965) overcame this by creating the genus Bergheia for 0. okapiae. No
specimens of this species are available for examination and the description of it shows
many similarities to H. rubidus, particularly in the shape and branching of the dorsal
ray. Until specimens of this species can be obtained for study the taxonomic position of
this species and genus is in doubt.
The systematic position of Mazamastrongylus Cameron, 1935 is also uncertain.
Cameron was of the opinion that the type and only species, A/, trinitatis, had affinities
to both Ostertagia and Cooperia. Unfortunately the type specimens are no longer
available. HINAIDY & PROSL (1981) and JANSEN (1981) consider that the genus
LOPEZ-NEYRA
190
Spiculopteroides Jansen, 1958 is similar to Mazamastrougylus and that when specimens

of Mazamastrongylus are available for study the two genera may be found to be
synonymous.
The genus Paramecistocirrus Roetti, 1941 was originally placed in the subfamily
Nematodirinae. DURETTE-DESSET & CHADAUD (1977) moved the genus to the subfamily
Ostertagiinae. Unfortunately, no specimens were available for examination and
consequently the genus has not been included in the main key.
The type species of the genus Teladorsagia, T. davtiani Andreeva & Satubaldin,
1954 and T. trifurcata (Ransom, 1907) Drozdz, 1965 have been regarded as synonyms
by several authors although BECKLUND & WALKER (1971) considered them to be
separate species. More recently, DASKALOV (1974) has regarded the two species as part
of a species complex of T. circumcincta (Stadelman, 1894) Drozdz, 1965. The genus
Teladorsagia has been included in the main key to accommodate these and other
species for the time being.
Ostertagiana Dikov, 1963 and Ostertamia Dikov, 1963 were described from the
abomasum of sheep in Kazakhstan. The two genera were placed in the tribe
Ostertagiea by DIKOV (1963). Unfortunately, no specimens were available for study and
the genera are, therefore, not included in the main key.
The synonymy of the following is confirmed: Capreolagia Schulz, Andreeva &
Kadenatsii, 1954 and Gruhneria Sarwar, 1956 with Ostertagia Ransom, 1907;
Mujlonagia Schulz, Andreeva & Kadenatsii, 1954 and Sjobergia Sarwar, 1956 with
Skrjabinagia (Kassimov, 1942) Altaev, 1952; Altaevia Sarwar, 1957 with
Spicitlopteragia (OrlofT, 1933) Travassos, 1937; Ostertagiella Andreeva, 1956 with
Grosspiculagia (OrlofT, 1933) Sarwar, 1956; Stadelmannia with Teladorsagia Andreeva
& Satubaldin, 1954.
Graphidiinae
The classification of this subfamily follows that of SKRJABIN et al. (1952). Characters
common to all the genera are: the absence of a cephalic vesicle; a synlophe with a very
large number of longitudinal ridges; a stout dorsal ray only divided at its distal tip; and
long filiform spicules. Characters separating the genera are: the length of the
externodorsal ray; the arrangement of the ventral rays; the size of the spicules; and the
formation of the genital cone.
Graphidioides Cameron, 1923, previously placed in the Trichostrongylinae, is
considered to have more affinities with this subfamily.
Graphidiella was erected by OLSEN (1948) and for the species G. ochotonae from
Ochotona princeps figginsi in Nebraska. Unfortunately the type specimens are no
longer available and no other specimens could be obtained. The genus is consequently
not included in the main key.
Abbreviations used in figures:
0=ventral raylets
7=dorsal raylets
D = Dorsal
V = Ventral
Dorsal ray
S spicule(s)
R Right
mm = millimetres
PC=Proconus
TTclamon
191
KEY
Family Trichostrongylidae Leiper, 1912

Key to subfamilies
l-(4) Synlophe absent or reduced to a few discontinuous, poorly developed longitudinal
cuticular ridges in the female vulvar region (Figs. 3, 4).
2-(3) Excretory pore opening in a notch on the body surface (Fig. 6) Spicules short, generally
twisted, irregular in shape, often with barbs near their distal tips (Fig. 5).
Dorsal lobe generally reduced, dorsal ray only branching at its distal end (Fig. 8).
Genital cone simple in structure with a single ventral raylet and a pair of short dorsal
raylets (Fig. 7)
Trichostrongylinae
3-(2) Excretory pore not opening in a notch on body surface.
Spicules short and pointed (Fig. 9).
Dorsal lobe not reduced, dorsal ray long (Fig. 19).
Genital cone with single ventral raylet, dorsal raylets and accessory bursal membrane
modified (Figs. 12, 13)
Libyostrongylinae
4-(l) Synlophe present and well developed.
5-(6) One or more teeth in the buccal cavity (Figs. 20, 21).
Large cervical papillae projecting from the surface of the body (Fig. 14).
Longitudinal cuticular ridges sometimes restricted to the anterior part of the body.
Dorsal ray symmetrical or asymmetrical (Figs. 23, 24, 26)
Haemonchinae
6-(5) No teeth in the buccal cavity.
Small cervical papillae.
Longitudinal cuticular ridges always extend along the full length of the body.
Dorsal ray symmetrical.
7-(10) Cephalic vesicle present (Figs. 15, 17).
Parasites of artiodactylids.
8-(9) Cephalic vesicle large (Fig. 17).
Cervical papillae little developed and difficult to see.
Number of longitudinal cuticular ridges small, ridges sometimes enlarged in the lateral
field (Figs. 35, 49, 50, 52)
Cooperiinae
9-(8) Cephalic vesicle restricted and small (Fig. 15).
Cervical papillae small, project from the body surface (Fig. 16).
Number of longitudinal cuticular ridges large, not enlarged in the lateral field
(Fig. 18)
Ostertagiinae
10-(7) Cephalic vesicle absent (Fig. 113).
Parasites of rodents, lagomorphs and primates.
Number of longitudinal cuticular ridges very large, the height of the ridges sometimes
very low (Figs. Ill, 114).
Spicules long and filiform.
Dorsal ray stout divided only at its distal tip (Figs. 107, 112)
Graphidiinac
Subfamily Trichostrongylinae Leiper, 1908
One genus (Figs. 5, 6, 7, 8).
Trichostrongylus Looss, 1905
( = Buckleya Sarwar, 1956; = Cobbohlostrongylus Sarvvar, 1956;
= Gilesia Sarwar, 1956; = Probostrongylus Sarwar, 1956)
Subfamily Libyostrongylinae Durette-Desset & Chabaud, 1977
Key to genera
H2) Dorsal ray with two pairs of branches from the main trunk (Fig. 10).
Parasites of rodents, lagomorphs and primates
Paralibyostrongylus Ortlepp, 1939
{ = Baylisiella Skrjabin & Schulz, 1937)
2-0) Dorsal ray with one pair of branches from the main trunk (Fig. 11).
3-(4) Tip of antcrolatcral ray modified to form a chitinous pincer (Fig. 22).
Genital cone with a pair of dorsal raylets associated with an oval membrane (Fig. (13).
Parasites of Madagascan rodents {Hypogeomys antimena)
Cnizostrongylus Chabaud, Durette-Desset & Houin, 1967
192
0 2mm
0 1mm
4sa
0-1 mm
0-OSmm
FIGS. 3-8. 3. Libyostrongylus douglassii, cross section of male in the middle region of the body.
4. Paralibyostrongylus vondwei, cross section of female just posterior to the vulva. 5. Trichostrongylus
retortaeformis, spiculcs. 6. Trichostrongylus cotubriformis, anterior end. 7. Trichostrongylus colubriformis,
ventral view of genital cone. 8. Trichostrongylus colubriformis, male bursa.
193
10
Of
11
12
0-2inm
11
O'lmm
t...
\"
*.
O-1nnn
v0
004mm
0 03mm
13
Paralibyostrongylus cassonei, dorsal ray.

, * . of 8 c n i l a , cone.
13. Cnizostrongylus kleini, ventral view of genital cone
194
0-1mm
0 05 mm
FIGS. 14-18. 14. Haemonchus contortus, cervical papillae. 15. Ostertagia ostertagi, cephalic region.
16. Ostertagia ostertagi, female, cervical papillae. 17, Cooperia curticei, anterior end of female. IS.Ostertagia
ostertagi, cross section of male, midbody region.
4-(3) Tip of anterolateral ray not modified (Fig. 19).

Genital cone with single ventral raylet and a complex arrangement of sclerotized plates
with a reduced accessory bursal membrane dorsal to cloaca (Fig. 12).
Parasites of birds (Struthioniformes, Rheiformes) and hyracoids
Libyostrongylus Lane, 1923
Subfamily Haemonchinae Skrjabin & Schulz, 1952
Key to genera
l-(2) Buccal cavity with more than one tooth (Fig. 21).
Lateral rays and sometimes externodorsal rays of bursa chitinized (Fig. 23).
Parasites of rodents
Bochmiclla Gcbaucr, 1932
2-(l) Buccal cavity small with only one tooth (Fig. 20).
Lateral rays of bursa not chitinized (Figs. 24, 26).
3-(6) Well developed synlophc extends along full length of body.
195
19
O 3mm
.99
0 2mm
0 0 4mm
19.23
.90
FIGS 19-23 19 Libyostrongylus douglassii, one lateral lobe and dorsal lobe of bursa. 20. Haemonchus
contortus, cephalic region of male. 21. Boehmiella perichitinea, en face (after GEBAUER 1932).
22. Cnizostrongylus klcini, part of one lateral lobe of bursa showing the distal tips of the ventral and lateral
rays. 23. Bochniella perichitinea, one lateral lobe and dorsal lobe of bursa.
196
4-(5) Spicules long and thin (Fig. 25).

Parasites of lagomorphs
Biogastronema Rohrbacher & Ehrenford, 1954
5-{4) Spicules short.
Parasite in the stomach of Hippopotamus
Leiperiatus Sandground, 1930
6-(3) Well developed synlophe restricted to the anterior half of the body.
7-(8) Dorsal ray asymmetrical (Fig. 24).
Spicules with barbs at their distal tips (Fig. 30).
Primarily parasites of artiodactylids
Haemonchus Cobb, 1898
8-(7) Dorsal ray symmetrical (Fig. 26).
9-{10) Spicules long and thin slightly swollen near distal end (Fig. 29).
Vagina long, vulva opening near anus (Fig. 28).
Parasites of bovids
Mecistocirrus Railliet & Henry, 1912
10-(9) Spicules short with barbs and projections near distal end (Fig. 31).
Vagina short, vulva opening in posterior part of the body some distance anterior to anus
(Fig. 27).
Parasites of artiodactylids
Ashworthius LeRoux, 1930
Subfamily Cooperiinae Skrjabin & Shikhobalova, 1952

Key to genera
l-(2) Dorsal ray divides towards the middle into two branches to form a lyre shape
(Fig. 34i, ii).
Spicules short, thick and generally ornamented with "striations" (Fig. 32).
Cooperia Ransom, 1907
2-(l) Dorsal ray not forming lyre shape (Fig. 33).
3-(8) Longitudinal cuticular ridges number 16 in the middle region of the body with one small
ridge in each lateral field (Fig. 35).
4-(5) Spicules short without ornamentation and with a single branch (Fig. 39).
Genital cone conspicuous with a complex arrangement of sclerotized plates and bars
surrounded by a reduced accessory bursal membrane dorsal to the cloaca (Fig. 40).
Female didelphic (Fig. 36).
Female tail without papillae or terminal spike (Fig. 37).
Parasites of African tragulids {Hyemoschus aquoticus)
Chabaudstrongylus Durette-Desset & Denke, 1978
5-(4) Spicules long, thin, unbranched (Fig. 38).
Genital cone not conspicuous or complex (Fig. 41).
Female monodelphic, tail with terminal papillae or spike (Figs. 44, 46).
6-(7) Spicules pointed at distal tjps (Figs. 42).
Distinct gubernaculum present (Fig. 43).
Female tail with papillae (Fig. 46).
Impalaia Mdnnig, 1923
(=Anthostrongylus LeRoux, 1936;
= Mimitostrongylus Croveri, 1929)
7-(6) Spicules with a distinct foot at distal tips (Fig. 45).
Distinct gubernaculum absent.
Female tail with terminal spike (Fig. 44).
Parasites of bathyergid rodents (Bathyergus suillus)
Ortleppstrongylus
Durette-Desset, 1970
8-(3) Longitudinal cuticular ridges number 10 in the middle regional of the body,
occasionally one additional smaller ridge in each lateral field (Fig. 49).
9-(12) Longitudinal cuticular ridges perpendicular to the surface of the body.
10-( 11) Longitudinal cuticular ridges of more or less even height (Fig. 49).
Dorsal ray dividing in its distal half forming an inverted Y-shape (Fig. 47).
Female tail pointed (Fig. 48).
Parasites of African bovids
Cooperioides Daubney, 1933
1H10) Longitudinal cuticular ridges with two larger equal ridges in each lateral field (Fig. 50).
Dorsal ray dividing in its distal quarter each branch dividing again into two unequal
branches (Fig. 51).
Female tail with a swelling at its distal tip (Fig. 53).
197
FIGS. 24-26.24. Ilaemonchus contortus, male bursa. 25. Biogastronema leporis, spicules. 26. Mecistocirrus
digitatus, composite diagram of male bursa.
198
,L. M. GIBBONS and,L. F.-KHALIL
28
30
31
01mm
O'lmm
I 30
-129,31
FIGS. 27-31. 27. Ashworthius pattoni, vulvar region (not drawn to scale). 28. Mecistocirms digitatus
posterior end of female (not drawn to scale). 29. Mecistocirms digitatus, distal ends of spicules. 30
Haemonchus contortus, distal ends of spicules. 31. Ashworthius pattoni, distal ends of spicules.
Parasites of African bovids {Nesotragus moschotus)

Megacooperia Khalil & Gibbons, 197(
12-(9) Longitudinal cuticular ridges uneven in height and not perpendicular to the surface o
the body.
13-(14) Two larger longitudinal ridges in each lateral field which curve dorsally, dorsal pai
larger than ventral pair (Fig. 52).
Spicules with a main stem and two branches one with serrations (Fig. 54).
Parasites of bovids
Paracooperia Travassos, 193:
(=Schwartziella LcRoux, 1936
14-(13) One larger longitudinal ridge in each lateral field.
Spicules without serrations (Fig. 55).
Parasites of African bovids (Pelea caprcolus)
Paracooperioiilcs Boomkcr, Horak & dc Vos, 198
199
O 3mm
O-Itnm
33
0 0 5 mm
0-2mm
O'lmm
-I
36
0 05mm
32
J37
J34
_I3S
FIGS. 32-37 32 Cooperia pectinata, spicules. 33. Impalaia tuberculata, male bursa. 34. Cooperia curticei,
lateral (i) and ventral (ii) views of dorsal ray. 35. Clmbaudstrongylus dubosti, cross section of male, midbody
region. 36. Clmbaudstrongylus dubosti, vulvar region. 37. Chabaudstrongylus dubosti, female tail.
200
FIGS. 38-45. 38. Impalaia tubcrculata, spiculcs. 39. Chabaudstrongylus dubosti, spiculcs.
40. Chabaudstrongylus tlubosti, ventral view of genital cone. 41. Impalaia tuberculata, ventral view of genital
cone. 42. Impalaia tuberculata, distal ends of spiculcs. 43. Impalaia tuberculata, gubcrnaculum.
44. Ortleppstrongylus bathyergi, posterior end of female. 45. Ortleppstrongylus bathyergi, distal ends of
spiculcs.
201
50
0-15nim
0-irnm
0 0 5mm
46
-47,48,49
ISO
J I G S . 46-50. 46. Impalaia tuberculata, posterior end of female. 47. Cooperioides antidorca, dorsal ray.
48. Cooperioides hamihoni, female tail. 49. Cooperioides hamihoni, cross section of male, midbody region.
50. Megacooperia woodfordi, cross section of male, midbody region.
202
55
\
0*2 mm
0-imm
O'lmm
004mm
FIGS. 51-55.51. Megacooperia woodfordi, one lateral lobe and dorsal lobe of bursa. 52. Paracooperia serrata,
cross section of male, midbody region. 53. Megacooperia woodfordi, female tail. 54. Paracooperia nodulosa,
spicules. 55. Paracooperioides peleac, spicules.
A key for. the.genera of-.Trichostrongylidae
203
Subfamily Ostcrtagiinac Lopez-Neyra, 1947

Key to genera
l-(8) Bursa with dorsal lobe and ray not reduced (Fig. 60).
2-(5) Spicules with two almost equal branches, without barbs and a main stem.
3-(4) Spicules with two long, thin pointed branches and a thick main stem (Fig. 59).
Genital cone with a single ventral raylet and dorsal raylets associated with a thickened
accessory bursal membrane divided into two lobes, one for each raylet (Fig. 63).
Parasites of African bovids (Antilopinae)
Gazellostrongylus Yeh, 1956
4-(3) Spicules with two branches and a main stem all more or less the same width, branches
with bluntly rounded distal tips (Fig. 56).
Genital cone with a pair of ventral raylets joined at the base surrounded by a membrane
which forms two lobes, one for each ventral raylet, and a pair of long dorsal raylets
surrounded by a simple accessory bursal membrane.
Marshallagia (Orloff, 1933) Travassos, 1937
5-(2) Spicules with two equal or unequal branches one with a barb at its distal tip and a main
stem (Fig. 65).
6-(7) Genital cone with a pair of ventral raylets joined at the base and a pair of short dorsal
raylets surrounded by a simple accessory bursal membrane (Fig. 58).
Parasite of artiodactylids
Grosspiailagia (Orloff, 1933) Sarvvar, 1956
( = Ostertagiella Andreeva, 1956)
7-(6) Genital cone with paired ventral raylets joined at the base and Sjoberg's organ dorsal to
the cloaca (Fig. 62).
Skrjabinogia (Kassimov, 1942) Altaev, 1952
( = Muflonagia Schluz, Andreeva & Kadenatsii, 1954; =Sjobergia Sarwar, 1956)
8-(l) Bursa with dorsal ray and lobe short or reduced (Figs. 64, 66, 73, 74, 78, 89, 97, 104).
9-(14) Bursa with very reduced dorsal ray and lobe, length of main stem of dorsal ray very
reduced (Fig. 66).
10-(13) Spicules with two equal branches and a main stem (Figs. 61, 67).
Genital cone with a pair of dorsal raylets undivided at their distal tips and associated
with an accessory bursal membrane which may be lobed at its distal margin (Fig. 69).
H-{12) Spicules slender with two short sharply pointed branches (Fig. 61).
Dorsal ray stout, bifurcates distally for up to half its length (Fig. 66).
Longistrongylus LeRoux, 1931
Parasites of African artiodactylids
( = Bigalkea Monnig, 1931; =Bigalkenema Ortlepp, 1963;
= Kobusinema Ortlepp, 1963)
^-(l 1) Spicules thick, one branch sharply pointed, one with a barb or triangular projection on
its distal tip (Fig. 67).
Dorsal ray bifurcating near its base, each branch widely divergent and divided at its
distal tip (Fig. 68).
Sarwaria Drozdz, 1965
Parasites of bovids [Bubalus bubalis)
13
-(10) Spicules with a single, long branch (Fig. 70).
Genital cone with a pair of dorsal raylets which are close together and bifurcate at their
distal tips (Fig. 71).
Dorsal ray with a swollen base bifurcating only at its distal tip (Fig. 72).
Parasites of bovids (Ethiopian sheep and goats .... Pseudomarshallagia (Roetti, 1941)
Graber & Delavenay, 1978
*4-(9) Dorsal lobe and ray not very reduced, length of main stem of dorsal ray not reduced
(Figs. 64, 73, 74, 78, 89, 97, 104).
15
-(24) Anterolateral and mediolateral rays close together or parallel for most or all of their
length, posterolateral ray widely separate from the other laterals and curving towards
the externodorsal ray (Figs. 74, 78, 104).
16
-(17) Anterolateral and mediolateral rays slightly divergent at their extreme tips (Fig. 74).
Dorsal ray with three pairs of branches (Fig. 75).
Spicules tapering with a single branch and a main stem (Fig. 77).
Genital cone with two very reduced dorsal raylets (Fig. 76).
Parasites of suids
Hyostrongyhts Hall, 1921
17
-(16) Anterolateral and mediolateral rays parallel for their entire length, anterolateral ray
generally shorter than mediolateral ray (Fig. 78, 104).
204
56
0-15 mm
59,60
008mm
005mm
56,58
57
FIGS. 56-60.56. Marshallagia marslialli, spi'cules. 57. Marshallaqia marshalli, ventral view of genital cone.
58. Grosspiculagia occidentalis, ventral view of genital cone. 59. Gazellostrongylus lerouxi, spicules.
60. Grosspiculagia occidentalis, one lateral lobe and dorsal lobe of bursa.
Olmm
0 0 5 mm
205
163,64.65
I 62
61-65. 61. Longistrongylus meyeri, spicule. 62. Skrjabinagia kolchida, ventral view of genital cone.
->. Gazellostrongylus lerouxi, ventral view of genital cone. 64. Ostertagia ostertagi, bursa. 65. Grosspiculagia
Occidentalis, spicules.
6
206
"L. M. GIBBONS and'L.T. KHALIL
66
67
68
0-1mm
69
i 66
0-imni
005mm
J69
FIGS. 66-69. 66. Longistrongylus meyeri, one lateral lobe and dorsal lobe of bursa. 67. Sarwaria bubalis,
spicules. 68. Sarwaria bubalis, dorsal ray (reconstruction from specimens examined). 69. Longistrongylus
meyeri, ventral view of genital cone.
key for the genera of Trichostrongylidae
207
73
0 1mm
0 1mm
0 ' Imm
70
71.72
FIGS. 70-73. 70. Pseudomarshallagia elongata, spicules. 71. Pseudomarshallagia elongata, ventral view of
genital cone. 72. Pseudomarshallagia elongata, dorsal ray. 73. Camelostrongylus mentulatus, one lateral lobe
and dorsal lobe of bursa.
208
O'lmm
74
0-1 mm
\
005mm
75.76
FIGS. 74-77. 74. Hyostrongylus rubidus, bursa. 75. Hyostrongylus nibidus, dorsal lobe. 76. Hyostrongylus
rubidus, ventral view of genital cone. 77. Hyostrongylus rubidiis, spicules.
18-(19) Dorsal ray with branches that divide as soon as they arc formed (Fig. 78).
Spicules with two distinct branches, equal or unequal, and a main stem (Fig. 79).
Genital cone with two modified dorsal raylets (Fig. 80).
Parasites of lagomorphs and artiodactylids
Cervicaprastrongylus Gibbons & Khalil (1982)
19-(18) Dorsal ray with branches that divide only at their distal tips (Figs. 83, 84).
20-(21) Branches of dorsal ray stout (Fig. 83).
Tclamon strongly developed (Figs. 85, 86).
Spiculcs with one thick branch which has triangular distal tip and one pointed branch
(Fig. 87).
Rinadia Grigorian, 1951
209
78
79 t
jl !}' 78 ~ 82 - 78. Cervicaprastrongylus mahiyai, one lateral lobe and dorsal lobe of bursa
y. Cervicaprastrongylus makiyai, spicules. 80. Cervicaprastrongylus mahiyai, composite drawing of ventral
ie\v of genital cone. 81. Camelostrongylus mentulatus, section of spicule. 82. Camelostrongylus mentulatus
a|
stal tip of spicule.
'
210
21-(20) Branches of dorsal ray slender, usually curved (Fig. 84).

22-(23) Spicules with a single branch surrounded by a striated membrane and a main stem with
many barbs and projections (Fig. 105).
Spiculopteragia (OrlofT, 1933)
Travassos, 1937
(=AUaevia Sarwar, 1957)
23-(22) Spicules with two equal branches, one pointed and one with a barb or triangular
projection at its distal tip (Fig. 88).
Parasite of artiodactylids
Apteragia Jansen, 1958
24-(15) Anterolateral ray curving towards ventral rays, mediolateral and posterolateral rays
curving towards externodorsal rays (Figs. 64, 73, 89, 97).
25-(26) Lateral rays widely separated.
Ventral rays almost parallel.
Externodorsal rays sharply curved (Fig. 89).
Parasites of cervids
Spiculopteroides Jansen, 1958
26-(25) Lateral rays not widely separated.
Ventral rays not parallel.
Externodorsal rays only slightly curved.
27-(28) Spicules very ornamented (Figs. 81, 82).
Dorsal ray with one pair of short branches from main trunk each of which divides again
(Fig. 73).
Genital cone with a pair of dorsal raylets which are partly fused along their length
(Fig. 90).
Camelostrongylus OrlofT, 1933
28-(27) Spicules not ornamented (Fig. 91).
Dorsal ray with one pair of long branches from main trunk each of which divides again
(Fig. 92).
Genital cone with a pair of long dorsal raylets which are separate along their length
(Figs. 93, 94).
Orlojfia Drozdz, 1965
29-(30) Proconus enlarged (Figs. 95, 96).
Spicules with two short slender branches shorter than the main stem, one or both hooked
or barbed at distal tip (Fig. 98).
Genital cone with short dorsal raylets associated with a simple accessory bursal
membrane (Figs. 95, 96).
Ostertagia Ransom, 1907
( Capreolagia Schulz, Andreeva & Kadenatsii, 1954;
Gnihneria Sarwar, 1956)
30-(29) Pronconus not enlarged.
Spicules with two sharply pointed branches (Fig.' 106).
Genital cone with Sjoberg organ dorsal to cloaca or pair of dorsal raylets associated with
a simple accessory bursal membrane (Figs. 99, 100, 101, 102, 103).
Teladorsagia Andreeva & Satubaldin, 1954
(=Stadelmannia Sarwar, 1954)
Subfamily Graphidiinae Travassos, 1937

Key to genera
l-(2) Externodorsal ray longer than dorsal ray, ventral rays close and curved (Fig. 107).
Synlophe with numerous longitudinal cuticular ridges (Fig. 111).
Genital cone with single ventral raylet and modified dorsal raylets and accessory bursal
membrane (Figs. 108, 109).
Parasites of lagomorphs
Graphidium Railliet & Henry, 1909
2-(l) Externodorsal ray shorter than dorsal ray, ventral rays curved and widely separated
(Fig. 112).
Synlophe with a large number of small longitudinal cuticular ridges (Fig. 114).
Genital cone with a single ventral raylet and a pair of short, thick, dorsal raylets close
together (Fig. 110).
Parasites of rodents (Cavoidea, Chinchilloidca) and primates (Platyrrhina)
Graphidioides Cameron, 1923
211
0 1mm
86 ,88.89
O'tmm
87
005mm
83 .84.85
FIGS. 83-89 83 Rinadia mathevossiani, dorsal ray. 84. Spiculopteragiaaspnmetnca, dorsal ray. 85. Rinadia
nathevoJani vent al view of genital cone. 86. Rinadia mathevosmm, lateral view of gemtal cone. 87 Rinadia
mathevossS, lcScs. 88. Apteragia quadrispiculata, spicules. 89. Spiculopteroides odocoiki, one lateral lobe
and dorsal lobe of bursa.
212
91
92
93
0 1mm
.01 Qd
OR
0 1mm
0 05mm
92
DO.93
FIGS. 90-94. 90. Camelostrongylus mentulatus, ventral view of genital cone. 91. Orlojjia orlojfi, spicule.
92. Orloffiaorloffifdorsalray.93.Orlojffiaorloffi, ventral viewofgenital cone. 94.OrlofliaorloJfi, lateral viewof
male tail.
213
96
97
0'2mm
OlSmm
O-Imm
_i97
J95.96
J98
HGS. 95-98 95 Ostertaaia ostertagi, male tail, ventral view. 96. Ostertagia ostertagi, male tail, lateral view.
97- TelairsagilûSTe
lateraVlobe and dorsal lobe of bursa. 98. Ostertagu ostertag,, sp.cules.
214
99
101
102
103
O'tmm
O'OSmm
_f 101,103
J09.1OO.1O2
FIGS. 99-103.99. Teladorsagia davtlani, ventral view of genital cone. 100. Teladorsagia trifurcata, ventral
view of genital cone. 101. Teladorsagia davtlani, lateral view of genital cone. 102. Teladorsagia circumcincta,
ventral view of genital cone. 103. Teladorsagia trifurcata, lateral view of genital cone.
215
104
106
O'lmm
.1104.105.106
. 104-106. 104. Apteragia quadrispicuhta. bursa. 105. Sptculopteragta asymmetric* spicules.

'06. Teladorsagia davtiani, spicules.
216
107
FIGS. 107-111. 107. Graphidium strigostim, one lateral lobe and dorsal lobe of bursa. 108. Graphidium
strigosum, ventral view of genital cone. 109. Graphidium strigosum, lateral view of genital cone.
110. Graphidioides affinis, ventral view of genital cone. 111. Graphidioides ajjinis, cross section of males,
midbody region.
217
112,114
113
ceoh-jl
) * ' ' ^ - Graphidioules qffinis, one lateral lobe and dorsal lobe of bursa. 113. Graphidium strigosum,
H anc region of female. 114. Graphidioides qffinis, cross section of male, midbody region.
Genera of uncertain affinities

tne
A?,
family Trichostrongylidae there are five genera of uncertain affinities.
though representative specimens of these genera have been made available to us for
lamination, the characters have not enabled us to assign them with certainty to any of
e
existing subfamilies. Details of host and locality refer to the specimens examined.
* ârarhabdonema Kreis, 1945 (Figs. 115-122)

type and only species: Pararhabdonema longistriata Kreis, 1945
Host: Lepilemur sp.
Habitat: Intestine
Locality: Madagascar
Cephalic vesicle absent; cervical papillae small; numerous longitudinal ridges
^tending along the length of the body, closer together in the lateral field, short in
tgntl prebursal papillae large; bursa with two large lateral lobes and an unreduced
orsal lobe, ventral rays widely separated, lateral rays separated at distal ends, all
rv
'ng towards dorsal rays, externodorsal ray arising in the second quarter from'the
218
115
119
118
0-2mm
01mm
120
J115.117.12O
J116.119
r
0'1mm
J118
FIGS. 115-120. 115. Pararhabdonema longistriata, one lateral lobe and dorsal lobe of bursa.
116. Pararhabdonema longistriata,cepha\ic region, female. 117. Pararhabdonema longistriata,cross section of
female, mid body region. 118. Pararhabdonema longistriata, ventral view of genital cone.
119. Pararhabdonema longistriata, spicules, in situ. 120. Pararhabdonema longistriata, cervical papillae.
219
-""""
121
l G S . 121-122. 121. Pararhabdonema longistriata, female tail, lateral view. 122. Pararhabdonema longisWata, vulvar region.
220
proximal end of the main trunk of dorsal ray, dorsal ray long, dividing only at its distal
tip with each small branch dividing again at its tip; spicules short, pointed; genital cone
with single ventral raylet and two short, thick dorsal raylets surrounded by a small
accessory bursal membrane; female didelphic.
Comments
KREIS (1945) described the female specimens from a' Madagascar lemur as
Pararhabdonema longistriata. Later, CHABAUD & CHOQUET (1955) obtained and
described the male and considered this species to have many affinities with
Pseudostertagia bullosa (Ransom & Hall, 1921) Travassos, 1937. DURETTE-DESSET &
CHABAUD (1977) placed the genus with Pseudostertagia in the subfamily
Libyostrongylinae. The specimens examined had a synlophe, the longitudinal cuticular
ridges extending along the length of the body; the ventral rays are widely divergent, the
dorsal ray divides only at its distal tip into two main branches, each of which forms a
small division at the extreme distal tip, and the externodorsal rays arise part way along
the length of the dorsal ray. These characters differ from those of other genera'of the
Libyostrongylinae and, therefore, the taxonomic position of the genus remains
uncertain.
2. Parostertagia Schwartz & Alicata, 1933 (Figs. 123-129)
Type and only species: Parostertagia heterospiculum Schwartz & Alicata, 1933
(Paratypes USDA Collection No. 30166)
Host: Pecari angulatus angulatus
Habitat: Small intestine
Locality: near Raymondville, Texas, USA
Cephalic vesicle absent; cervical papillae not seen; longitudinal cuticular ridges 20
in the middle region of the body of the male, of more or less even height, extending
along the length of the body; prebursal papillae large; bursa with two large lateral lobes
and an unreduced dorsal lobe, ventral rays separated at their distal tips, curving
anteriorly, lateral rays separated at their distal tips curving towards the dorsal rays,
externodorsal ray arising at the base of the dorsal ray, dorsal ray long, dividing at its
distal tip forming two pairs of branches, the internal pair being long and having each of
its branches dividing again at their distal tips; spicules short and pointed; genital cone
structure not clear in specimens examined; female didelphic.
Comments
The genus Parostertagia was established by SCHWARTZ & ALICATA (1933) for
P. heterospiculum from North American peccary. TRAVASSOS (1937) considered
Parostertagia to be a synonym otHyostrongylus. SKRJABIN et al. (1952) disagreed with
this synonymy and separated the two genera maintaining Parostertagia in the tribe
Ostertagiea of the Trichostrongylinae. In the specimens we examined, no cephalic
vesicle or cervical papillae could be seen and the structure of the genital cone could not
be determined. Although this genus has some affinities to the genera of the subfamily
Ostertagiinae, its taxonomic position remains uncertain.
3. Pseudostertagia (OrlofT, 1933) Travassos, 1937 (Figs. 130-136)
Type and only species: Pseudostertagia bullosa (Ransom & Hall, 1912) Travassos,
1937 (USDA Collection No. 45552)
Host: Antelope
Habitat: Abomasum
Locality: Hammond, Montana, USA
221
123
125
124
126
O'tfflm
123
Olmm
_i!24,125
005mm
_|126
1GS. 123-126 123 Parostcrtagia hcterospiculum, reconstruction ofone lateral lobe and dorsal lobeofbursa
from specimens examined. 124. Parostertagia heterospiculum, spicules, in situ. 125. Parostertagia heterospicul, gijbernaculum, in situ. 126. Parostertagia heterospiculum, cross section of male, midbody region.
222
L. M. GIBBONS and L.F. KHALIL
129
127
FIGS. 127-129. 127. Parostertagia heterospiculum, female tail, lateral view. 128. Parostertagia heterospicuhtm, vulvar region. 129. Parostertagia heterospiculum, anterior end.
130
13t
bun
*~ 133 - 1 3 - Pseudostertagia bullosa, cross section of male, midbody region. 131. Pseudostertagia
malc
anterior end. 132. Pseudostertagia bullosa, spicules, in situ. 133. Pseudostertagia bullosa
emu '
suoernaculum, in situ.
223
224
134
135
136
V-b
0 2 mm
135
01mm
134.136
FIGS. 134-136.134. Pscudostertagia bullosa, one lateral lobe and dorsal lobe of bursa. 135. Pseudostertagia
bullosa, vulvar region. 136. Pseudostertagia bullosa, female tail, lateral view.
Cephalic vesicle absent; cervical papillae not seen; longitudinal cuticular ridges 28,
32 in the middle region of the body of the male and female respectively, more or less
even height, extend along length of body; prebursal papillae large; bursa with two large
lateral lobes and an unreduced dorsal lobe, ventral rays separated at their distal tips,
anterolateral and mediolateral rays close together for most of their length, diverging
slightly at their distal tips, posterolateral separate from the other laterals curving
towards the dorsal ray, dorsal ray long, bifurcating at its distal tip, with each branch
225
giving off a small external branch halfway along its length, and dividing at its extreme
distal tip; spicules short and pointed; genital cone structure not clear in the specimens
examined; cuticle of bursa heavily striated obscuring rays; female didelphic.
Comments
RANSOM & HALL (1912) described this species from the abomasum of Ovis aries in
Colorado and Montana, USA and originally placed the type and only species P. bullosa
in the genus Ostertagia. ORLOFF (1933) in his revision of the genus Ostertagia placed it
in the subgenus Pseudostertagia and later TRAVASSOS (1937) elevated the subgenus to
generic rank. The genus was placed in the Libyostrongylinae by DURETTE-DESSET &
CHABAUD (1977).
The specimens we examined did not have a cephalic vesicle, the cervical papillae
could not be seen and the structure of the genital cone could not be determined.
Although the specimens showed some affinities to genera of the subfamily
Ostertagiinae, the taxonomic position of the genus remains uncertain.
4. Obeliscoides Graybill, 1924 (Figs. 137-145)

Type species: Obeliscoides cuniculi (Graybill, 1923), Graybill, 1924 (USDA
Collection No. 30174)
Host: Rabbit
Habitat: Stomach
Locality: Washington, D.C., USA
Cephalic vesicle absent; cervical papillae small, projecting from body surface;
longitudinal cuticular ridges 26 in the middle region of the body of the female, of more
or less even height, extending along length of body; prebursal papillae not seen in
specimens examined; bursa with two large lateral lobes and a reduced separate dorsal
!obe, ventral rays diverging and coming together at their distal tips, anterolateral ray
tong, curving towards the ventral rays, mediolateral and posterolateral rays shorter,
c
'ose together and curving towards the dorsal rays, externodorsal ray arising from the
base of the dorsal ray, the short dorsal ray with one pair of branches halfway along its
length projecting ventrally and another pair of branches at its distal tip, each branch
dividing again; the ventral part of the genital cone appears to be lobcd and the dorsal
raylets are fused for most of their length, separating and branching at their distal ends;
surrounded by a lobed accessory bursal membrane; spicules long, branching only at
their extreme distal tip; female didelphic.
Comments
The species of this genus are parasites of lagomorphs. According to SKRJABIN et al
(1952), SKRJAIUN & SCHULZ in 1937 included the genus in the tribe Ostertagiea.
D
URniTE-DESsET & CHABAUD (1977) moved the genus to the Libyostrongylinae. Our
lamination of specimens of Obeliscoides cuniculi, the type species of the genus,
revealed that there is no head vesicle, that small prominent cervical papillae are present
a
nd that the synlophe has 26 small longitudinal cuticular ridges. These characters show
* e genus has some affinities to the subfamily Ostertagiinae but the taxonomic position
f the genus is uncertain.
5
- Travassosius Khalil, 1922 (Figs. 146-151)

Type species: Travassosius rufus Khalil, 1922
Host: Castor canadensis
Habitat: Stomach
Locality: Alaska, USA
226
138
J137
0-1mm
0-1mm
0 05mm
J141
j139.14O
FIGS. 137-141.137. Obeliscoides cuniculi, male, anterior end. 138. Obeliscoides cuniculi, one lateral lobe and
dorsal lobe of bursa. 139. Obeliscoides cuniculi, ventral view of dorsal lobe and ray. 140. Obeliscoides cuniculi,
ventral view of genital cone. 141. Obeliscoides cuniculi, male tail, lateral view.
for the genera of Trichostrongylidae
227
U2,
144
0 2 mm
O-Inrim
_L-|143
J142.144.145
FIGS. 142-145. 142. Obeliscoides cuniculi, spicules. 143. Obeliscoides cuniculi, vulvar region.
144. Obeliscoides cuniculi, female tail, lateral view. 145. Obeliscoides cuniculi, cross section of female, midbody
region.
Cephalic vesicle absent; cervical papillae small, projecting from body surface;
longitudinal cuticular ridges 32 in the middle region of the body of the male, extending
along the length of the body and of more or less even height; prebursal papillae large;
bursa with two large lateral lobes and an unreduced dorsal lobe, ventral rays widely
separated, lateral rays separated at their distal tips, anterolateral curving towards the
ventral rays, posterolateral curving towards the dorsal rays, externodorsal rays arising
in the posterior part of the proximal third of the main trunk of dorsal ray, dorsal ray
bifurcating in its distal half, each branch dividing again at its distal tip; spicules short
with thick main stem and a single thinner branch; genital cone with small ventral raylet
228
K7 /[mm
148
U9
150
O'lmm
0'1mm
0 0 5 mm
1140,147.148.151
i14S
_|1SO
FIGS. 146-151.146. Travassosius rufus, one lateral lobe and dorsal lobe of bursa. 147. Travassosius rufus,
cephalic region of male. 148. Travassosius rufus, male, cervical papillae. 149. Travassosius rufus, cross section
of male, midbody region. 150. Travassosius rufus, ventral view of genital cone. 151. Travassosius rufus,
spicule.
229
and a pair of short dorsal raylets surrounded by a reduced accessory bursal membrane;
females not seen.
Comments
The genus is parasitic in beavers. TRAVASSOS (1937) considered it to be near
Trichostrongylus and Ostertagia. According to SKRJABIN et al. (1952), SKRJABIN &
SCHULZ in 1937 included the genus in the tribe Ostertagiea. DURETTE-DESSET &
CHAIMUD (1977) placed the genus in the subfamily Trichostrongylinae.
The specimens we examined had longitudinal cuticular ridges of more or less even
height and extending along the length of the body, and prominent cervical papillae,
showing the affinities of this genus to the subfamily Ostertagiinae. However, the
cephalic vesicle found in all other genera of this subfamily is absent in Travassosius. The
simple structure of the genital cone shows the affinities of this genus to the subfamily
Trichostrongylinae but the presence of a synlophe and prominent cervical papillae
separate it from the subfamily. For these reasons the taxonomic position of this genus is
uncertain.
Genera not examined
As stated in the Introduction, these keys are based on the authors' own
observations on the nematode genera in question. Specimens of the nine genera listed
below were not available for examination. The type specimens have either been lost
(Mazamastrongylus and Graphidiella) or too few specimens exist to be lent and no
further material has been collected.
The diagnostic characters, as described by the original authors, are summarized
under each genus.
! Moguranema Yamaguti, 1941
Type species: Moguranema nipponicum Yamaguti, 1941
Host: Mogura wogura wogura
Locality: Yamasina, near Kyoto, Japan
Diagnostic characters: Cephalic vesicle present; cervical papillae absent; integument double-layered, thrown into conspicuous transverse folds; longitudinal ridges
absent; mouth with an internal circle of minute "comb-teeth-like stnaUons; oe oPhagus muscular, narrowed behind nerve ring; prebursal papillae absent bursa
symmetrical with well developed lateral lobes and poorly diflerentiated dorsal lobe
anteroventral ray smaller than posteroventral ray, separated at their disUd d j
anterolateral ray largest, fused at base to other lateral rays, externodorsal ray tok and
^rved, dorsal ray with stout trunk divides into two branches which may or may
not divide again; sîcules simple, equal, connected by a membrane,joined togethe a
^stal tips; gube naculum absent; female body tapering a b r u p t l y ^ J * - * n
opposed; vulva opens some distance anterior to anus; eggs thin-shelled, large,
n
umerous; tail more or less digitiform.
2
- Neocooperia Fotedar & Bambroo, 1971

Type species: Neocooperia magniovata Fotedar & Bambroo, 1971
Host: Sheep
Habitat: Duodenum
Locality: Kashmir, India
long, 0-21-0-23mm wide (male),
'"conspicuous, without teeth; cervical papillae well developed; prebursal papillae
230
present; bursa symmetrical with two large lateral lobes longer than broad and a small
dorsal lobe; ventral rays with common trunk, directed inwards, posteroventral longer
and thicker than anteroventral; anterolateral thick, isolated from other two laterals,
mediolateral and posterolateral rays with common trunk, longer than anterolateral,
externodorsal rays arise from base of dorsal, dorsal ray bifurcates at beginning of distal
quarter, each branch with lateral offshoot bifid at its distal tip; spicules short, complex
with 2-3 distal offshoots of varying length and thickness; gubernaculum and telamon
absent; genital cone complex; vulva in posterior quarter of body; cuticular flaplike
structure present immediately behind vulva, along its dorsal side; eggs O15CMM80
x 0-075-0-100 mm.
3. Bergheia Drozdz, 1965
Type species: Bergheia okapiae (van den Berghe, 1937) Drozdz, 1965
Host: Okapia johnstoni
Habitat: Intestine
Locality: Zaire (Belgian Congo)
Diagnostic characters: Bursa with dorsal ray of "mean" size, which has three pairs
of small branches one in the middle, one near the distal end, one at the extreme distal
tip; spicules short, slender, tapering distally with blunt bifurcated tips; gubernaculum
absent.
4. Mazamastrongylus Cameron, 1935
Type species: Mazamastrongylus trinitatis Cameron, 1935
Host: Mazama simplicicornis
Locality: Trinidad
Diagnostic characters: (generic diagnosis not separated from species description):
Cephalic vesicle absent anterior end dome-like; cervical papillae small; transverse
striations visible in cephalic region; longitudinal cuticular ridges approximately 20
tending to be discontinuous and variable in number, numerous fine longitudinal
striations occur between these; bursa with two large lateral lobes and a small dorsal
lobe, ventral rays parallel for entire length separated from lateral rays, anterolateral
and mediolateral rays lie close together, diverge at distal tips, posterolateral ray
separate from other laterals, externodorsal rays stout, curved and shorter than the
posterolateral rays, the dorsal ray is short, stout, and divides approximately halfway
along its length, each branch divides to form three smaller branches, two medial, long
and pointed, one short; spicules similar, short, irregular in outline; gubernaculum
absent; small prebursal papillae present; the genital cone has an accessory bursal
membrane surrounding two weakly curved rays.
5. Paramecistocirrus Roetti, 1941.
Type species: Paramecistocirrus dimorphus Roetti, 1941
Host: Goat
Habitat: Abomasum
Locality: Ethiopia
Diagnostic characters: Genus based on the dimorphism of the male spicules.
6. Ostertagiana Dikov, 1963.
Type species: Ostertagiana andreevi Dikov, 1963
Host: Sheep
Habitat: Abomasum
Locality: Kazakhstan
231
Diagnostic characters: Small oral opening; bursa with three lobes; dorsal ray
bifurcates halfway along its length, each branch divides two thirds along its length
forming a lateral branch and divides again at its distal tip; genital cone large, rounded
Projection, proconus absent, accessory bursal membrane, dorsal and ventral raylets
absent; presence or absence of telamon not established; gubernaculum present; spicules
with main stem and two short branches which arise in distal third; female unknown.
7
- Ostertamia Dikov, 1963

Type species: Ostertamia orientalis Dikov, 1963
Host: Sheep
Habitat: Abomasum
Diagnostic characters: Small oral opening; bursa with three lobes; dorsal ray
bifurcates in the proximal half of its length forming long widely separated branches;
genital cone semicircular in shape with broad base, telamon weakly developed
v
entrally, well developed dorsally particularly at the distal end; accessory bursal
Membrane, dorsal and ventral raylets absent; spicules typical for Ostertagia sens, lat.;
gubernaculum present, little developed; female unknown.
8. Graphidiella Olsen, 1948
Type species: Graphidiella ochotonae Olsen, 1948
Host: Ochotona princeps figginsi
Habitat: Stomach
Loca//fy.\Gunnison County, Colorado, USA
Diagnostic characters: 'Cephalic vesicle absent; cervical papillae present; fine
transverse striations present; longitudinal cuticular ridges numerous; bursa symmetrical with dorsal lobe slightly differentiated from lateral lobes, all bursal rays
relatively straight throughout their length, ventral rays with short common trunk,
e
qual in size, slightly divergent, lateral rays with short common trunk, approximately
e
qual in .size, slightly divergent, dorsal ray longer and wider than other rays, divides
n
ear its distal tip, each branch dividing again; spicules equal or subequal, long, filiform
with fasciculate points and transparent alate processes; gubernaculum absent; genital
c
one well developed; vulva opens in third quarter from anterior end; body not
narrowing abruptly in the post-vulvar region; ovejectors approximately equal, each
With three distinct bulbs; eggs with thin, unmarked shells, those near vagina
e
mbryonated at morula stage; tail with obtuse conical terminal point.
9- Trichostrongyella Dikov, 1961.
Type species: Trichostrongyella schulzi Dikov, 1961
Host: Sheep
Habitat: Abomasum
Diagnostic characters: Cervical papillae absent; prebursal papillae small; bursa
, symmetrical with little differentiation of dorsal lobe; dorsal ray long, divides into
branches each bifurcating halfway down their length; ventral rays parallel,
Un
equal; well developed telamon present; spicules short with poorly developed ventral
a
la and well developed dorsal ala; gubernaculum absent; female unknown.
ACKNOWLEDGEMENTS
w
are very grateful to Dr. Sheila Willmott, former Director of the Commonwealth Institute of
inthology for her tremendous encouragement and continued interest in our studies since the project
n. The authors arc also grateful to Dr. A. Verster of the Veterinary Research Institute, Onderstepoort
232
and Dr. J. Boomker of the Department of Parasitology, Faculty of Veterinary Science of the University of
Pretoria, Republic of South Africa, Dr. J. R. Lichtenfels of the National Parasite Collection, USDA,
Beltsville, Maryland, USA, Dr. K. A. Neiland of Wildlife Diseases Studies Section, Department of Fish and
Game, Alaska, USA, Dr. J. Jansen of the University of Utrecht and Dr. F. H. M. Borgsteede of the Central
Veterinary Institute, Lelystad, The Netherlands, Professor A. G. Chabaud and Dr. M. C. Durette-Desset of
the Museum National d'Histoire Naturelle, Paris, and Dr. H. K. Hinaidy of the Institute of Parasitology,
Vienna, Austria for making specimens available for this study.
We would also like to acknowledge Annales de Parasitologie Humaine et Comparee, Journal of
Helminthology, Systematic Parasitology and Zeitsclirift fur Parasitenkunde in which some of the drawings
have previously been published.
REFERENCES
ANDREEVA, N. K. (1957) (Atlas of helminths (Strongylata) of domesticated and wild ruminants in Kazakh
SS.R.) Tashkent: Institut Veterinarii Kazakhskogo Filiala Vashknil, 215pp. (In Russian.)
ANDREEVA, N. K. (1957) (Revision of Ostertagia (trichostrongylids) of ruminants.) Trudy Instituta
Veterinarii. Akademiya Nauk Kazakhskoi SSR, Alma-Ata, 8, 473-487. (In Russian.)
BAER, J. G. (1950) Etude critique des helminthes parasites de l'Okapi. Ada Tropica, 7, 164-186.
BAYLIS, H. A. & DAUBNEY, R. (1926) Synopsis of the families and genera ofNematoda. London: British
Museum, 277 pp.
BECKLUND, W. W. & WALKER, M. L. (1971) Nomenclature and morphology of Ostertagia trifurcata
Ransom, 1907, with data on spicule lengths of five stomach worms of ruminants. Journal of
Parasitology, 57, 508-516.
BERGHE, L. VAN DEN (1937) Contribution a l'etudedes parasites de l'Okapi (Troisieme partie). Revue de
Zoologie et de Botanique Africaines, 30, 117-139.
CHABAUD, A. G. (1965) In: Traite de Zoologie, Anatomie, Systematique, Biologic Tome IV, Fasc. III.
Grasse, P. P. (Editor). Paris: Masson et Cie.
CHABAUD, A. G. & CHOQUET, M.-T. (1955). Deux nematodes parasites de lemurien. Annales de
Parasitologie humaine et comparee, 30, 329-338.
CHABAUD, A. G., PUYLAERT, F., BAIN, O , PETTER, A. & DURETTE-DESSET, M. C. (1970)
(Presentee par M. Pierre-Paul Grasse.) Remarques sur 1'homologie entre des papilles cloacales des
Rhabditides et les cotes dorsales des Strongylida. Compte Rendu Hebdomadaire des Seances de
VAcademie des Sciences, Paris, 271, 1771-1774.
DASKALOV, P. (1974) (On the reproductive relationships between Ostertagia circumcincta, Teladorsagia
davtiani and 0. trifurcata (Nematoda: Trichostrongylidae).) Izvestiya na Tsentralnata
Khelmintologichna Laboratoriya. Sofia, 17, 59-72. (In Bulgarian.)
DIKOV, G. I. (1961) (Trichostrongyella schulzi n.g., n.sp. (Nematoda: Trichostrongylidae) from sheep.)
Helminthologia, 3, 81-84. (In Russian.)
DIKOV, G. I. (1963) (Two new genera and species of Trichostrongylidae of sheep.) In: (Helminths of man,
animals and plants and their control. Papers on helminthology presented to Academician K. I.
Skrjabin on his 85th Birthday.) Moscow: Izdatel'stvo Akad. Nauk SSSR, pp. 178-182.(In Russian.)
DR62D2, J. (1965) Studies on helminths and helminthiases in Cervidae. I. Revision of the subfamily
Ostertagiinae Sarwar, 1956 and an attempt to explain the phylogenesis of its representatives. Acta
Parasitologica Polonica, 13 (44), 445-481.
DURETTE-DESSET, M. C. (1970) Affinites de lTieligmosome Ortleppstrongylus bathyergi (Ortlepp, 1939)
n.gen., n.comb. avec les trichostrongylides Molineinae. Bulletin du Museum National dllistoire
Naturelle, Paris, 2 Ser, 42 (2), 415-418.
DURETTE-DESSET, M. C. (1971) Essai de classification des nematodes heligmosomes. Correlations avec
la paleobiogeographie des hotes. Memoires du Museum National cCllistoire Naturelle, Paris,
Nouvelle Serie, Ser. A. Zool. 69, 1-126.
DURETTE-DESSET, M. C. & CHABAUD, A. G. (1977) Essai de classification des nematodes
Trichostrongyloidea. Annales de Parasitologie humaine et comparee, 52, 539-558.
DURETTE-DESSET, M. C. & CHABAUD, A. G. (1981) Nouvel essai de classification des nematodes
Trichostrongyloidea. Annales de Parasitologie humaine et comparee, 56, 297-312.
FOTEDAR, D. N. & BAMBROO, N. (1971) New trichostrongylid nematodc, Neocoopcria magniovata gen.
et sp. nov. from sheep in Kashmir. Proceedings of the Indian Science Congress Association, 58 (111),
555.
GIBBONS, L. M. & KHALIL, L. F. (1982). Ccrvicaprastrongylus, a new genus proposed for the ncmatode
species Ostertagia skrjabini Singh & Pande, 1963 (Trichostrongyloidea, Trichostrongylidae).
Systematic Parasitology (in press).
GIBBONS, L. M. & KHALIL, L. F. (1982) The morphology of the genital cone in the ncmatode family
Trichostrongylidae and its value as a taxonomic character. In: Concepts in Ncmatode Systematics.
Stone, A. R., Platt, H. M. & Khalil, L. F. (Editors). The Systematics Association Special Volume
No. London: Academic Press (in press).
HINAIDY, H. K. & PROSL, H. (1981) In: Systematics of Ostcrtagia sens. lat. (Nematoda:
Trichostrongylidae) (Workshop 14, EMOP 3). Parasitology, 82, 181-183.
JANSEN, J. (1958) Lebmaagtrichostrongylidcn bij ncderlanse herten. Utrecht, The Netherlands: H. J. Smits,
100 pp.
233
JANSEN, J. (1981) In: Systematics and biology o/Ostertagia sens. lat. (Nematoda: Trichostrongylidae)
(Workshop 14, EMOP 3). Parasitology, 82, 183-185.
KREIS, H. A. (1945) Beitrage zur Kenntnis parasitischer Nematoden. XII. Parasitischen Nematoden aus den
Tropen. Revue Suisse de Zoologie, 52, 551-596.
LEIPER, R. T. (1908) An account of some helminths contained in Dr. Wenyon's collection from the Sudan.
Report (3rd) of the Wellcome Tropical Research Laboratory, pp. 187-199.
LEIPER, R. T. (1912) Check-list of helminths parasitic in cattle (Bos taurus, Buffelas indiats, Bibos indiais).
Journal of the London School of Hygiene and Tropical Medicine, 1, 115-123.
LOPEZ-NEYRA, C. R. (1947). Helmintos de los vertebrados Ibericos. Granada: Instituto Nacional de
Parasitologia de Granada, Vol. II, pp. 413-802.
NEVEU-LEMAIRE, M.(1934) Sur la classification des trichostrongylides. Annales de Parasitologie humaine
et comparee, 12, 248-252.
OLSEN, L. S. (1948) A new trichostrongylid nematode from the pika, Ochotona princepsfigginsi. American
Midland Naturalist, 39, 748-753.
ORLOFF, I. W. (1933) Sur la reconstruction de la systematique du genre Ostertagia Ransom, 1907. Annales de
Parasitologie humaine et comparee, 11, 96-114.
OSCHE, G. (1958) Die Bursa und Schvvanzstrukturen und ihre Aberrationen bei den Strongylina
(Nematoda). Morphologische Studien zum Problem der Pluriund Paripotenzer schneinungen.
Zeitschrift fur Morphologie und Okologie der Tiere, 46, 571-635.
RAILLIET, A. & HENRY, A. (1909). Classification des Strongylidae. Metastrongylinae. Compte Rendu des
Seances de la Societe de Biologie, 66, 85-88.
RANSOM, B. H. & HALL, M. C. (1912) A new nematode, Ostertagia bullosa, parasitic in the alimentary
tract of sheep. Proceedings of the United States National Museum, 42, 175-179.
ROETTI, C. (1941) Nematodirinae parassita della capra. Nuova specie. Annali di Patologia Tropicale e di
Parassitologia, 2, 244-250.
SARWAR, M. M. (1956) Studies on some trichostrongylids of ruminants from the Indo-Pakistan subcontinent. Biologia, 2, 145-215.
SCHWARTZ, B. & ALICATA, J. E. (1933) Description of two parasitic nematodes from the Texas peccary.
Proceedings of the United States National Museum, 82, 1-6.
SKRJABIN, K. I , SHIKHOBALOVA, N. P., SHULTS, R. S., POPOVA, T. I., BOEV, S. N. &
DELYAMURE, S. L. (1952) Key to parasitic nematodes. III. Strongylata. Moscow: Academy of
Sciences of the USSR. (English translation published by the Israel Program for Scientific
Translations, 1961, 890 pp.)
STRINGFELLOW, F. (1970) Comparative morphology of the genital cones of Cooperia (Nematoda,
Trichostrongylidae) from cattle and sheep in the United States with a key to the common species.
Journal of Parasitology, 56, 1189-1198.
STRINGFELLOW, F. (1971) Functional morphology and histochemistry of structural proteins of the
genital cone of Ostertagia ostertagi, with a comparison of the genital cones of other Ostertagia
common in cattle in the United States. Journal of Parasitology, 57, 823-831.
STRINGFELLOW, F. (1972) Comparative morphology of genital cones of Ostertagia from sheep in the
United States. Journal of Parasitology, 58, 265-270.
TRAVASSOS, L. P. (1920) Esboco de uma chave geral dos nematodes parasitos. Revista de Veterinaria y
Zootechnia, Rio de Janeiro, 10, 59-70.
TRAVASSOS, L. P. (1937) Revisao de familia Trichostrongylidae Leiper, 1912. Monografias do Instituto
Oswaldo Cruz, No. 1, 512 pp.
Y A M A G U T I , S. (1941). Studies on the helminth fauna of Japan. Part 35. Mammalian nematodes II.
Japanese Journal of Zoology, 9, 409-439.

YAMAGUTI, S. (1961) Systema Helminthum. Vol. III. The Nematodes of Vertebrates. New York & London:
Interscience Publishers, vi+ 1261 pp.
YORKE, W. & MAPLESTONE, P. A. (1926) The nematode parasites of vertebrates. London: J. & A.
Churchill, 536 pp.
Accepted 15th October, 1981.
LONDON SCHOOL OF HYGIENE AND TROPICAL MEDICINE

University of London
The following postgraduate courses are offered:
Full and part-time MSc and diploma:
MSc Clinical Tropical Medicine
MSc Community Health in Developing Countries
MSc Community Medicine
MSc Epidemiology
MSc Human Nutrition
MSc
MSc
MSc
MSc
MSc
MSc
Medical Demography
Medical Microbiology
Medical Parasitology
Medical Statistics
Occupational Hygiene
Occupational Medicine
Diploma in Industrial Health

Diploma in Tropical Medicine and Hygiene
Diploma of Professional Competence

in Comprehensive Occupational Hygie
Part-time courses not leading to an award are held in:

Medical Statistics and Epidemiology
Population Studies
Research degrees:
MPhil and PhD
Further information may be obtained from:
The Registrar
London School of Hygiene and Tropical Medicine
Keppel Street (Govver Street)
London WC1E 7HT
Bursaries
Courses at the School are designed for the training of health personnel in an
international context and several pay particular attention to the needs of developing
countries.
It is expected, however, that fees for overseas students will increase next year. These
will be announced as soon as possible.
To help students pay the higher fees the School will ofler a number of substantial
bursaries in 1982/83. Priority will be given to applicants who (a) are self-supporting; {b)
are from developing countries and (c) have been accepted for the more expensive
courses of study.
If you think that you may qualify for a bursary, you should write without delay,
stating your course preference and giving reasons to support your application to:
The Registrar (as above)

A Key For The Identification of Genera of The Nematode Family Trichostrongylidae Leiper, 1912 PDF

Загружено:

Сведения о документе

Оригинальное название

Авторское право

Доступные форматы

Поделиться этим документом

Поделиться или встроить документ

Параметры публикации

Этот документ был вам полезен?

Это неприемлемый материал?

Авторское право:

Доступные форматы

A Key For The Identification of Genera of The Nematode Family Trichostrongylidae Leiper, 1912 PDF

Загружено:

Авторское право:

Доступные форматы

Journal of Helminthology (1982) 56, 185-233

A key for the identification of genera of the nematode family

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

Family Trichostrongylidae Leiper, 1912

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

Spiculopteroides Jansen, 1958 is similar to Mazamastrougylus and that when specimens

A key for the genera of Trichostrongylidae

Family Trichostrongylidae Leiper, 1912

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

Paralibyostrongylus cassonei, dorsal ray.

L. M. GIBBONS and L. F. KHALIL

4-(3) Tip of anterolateral ray not modified (Fig. 19).

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

4-(5) Spicules long and thin (Fig. 25).

Subfamily Cooperiinae Skrjabin & Shikhobalova, 1952

A key for the genera of Trichostrongylidae

,L. M. GIBBONS and,L. F.-KHALIL

Parasites of African bovids {Nesotragus moschotus)

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for. the.genera of-.Trichostrongylidae

Subfamily Ostcrtagiinac Lopez-Neyra, 1947

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

"L. M. GIBBONS and'L.T. KHALIL

key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

21-(20) Branches of dorsal ray slender, usually curved (Fig. 84).

Subfamily Graphidiinae Travassos, 1937

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

. 104-106. 104. Apteragia quadrispicuhta. bursa. 105. Sptculopteragta asymmetric* spicules.

L. M. GIBBONS and L. F. KHALIL

key for the genera of Trichostrongylidae

Genera of uncertain affinities

* ^ararhabdonema Kreis, 1945 (Figs. 115-122)

L. M. GIBBONS and L. F. KHALIL

key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

key for the genera of Trichostrongylidae

L. M. GIBBONS and L.F. KHALIL

key for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

4. Obeliscoides Graybill, 1924 (Figs. 137-145)

- Travassosius Khalil, 1922 (Figs. 146-151)

L. M. GIBBONS and L. F. KHALIL

for the genera of Trichostrongylidae

L. M. GIBBONS and L. F. KHALIL

A key for the genera of Trichostrongylidae

- Neocooperia Fotedar & Bambroo, 1971