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Source : MNHN, Paris

MÉMOIRES DU MUSÉUM NATIONAL D’HISTOIRE NATURELLE

Série A, Zoologie, Tome 107

A TAXONOMIC REVISION OF THE GENERA CITTOTAENIA RIEHM,

1881, CTENOTAENIA, RAI LU ET, 1893, MOSGOVOYM SPASSKII

1951 AND PSEUDOCITTOTAENIA TENORA, 1976.

(CESTODA : ANOPLOCEPHALIDAE).

lan. BEVERIDGE*

SUMMARY

INTRODUCTION. 2

ACKNOWLEDGEMENTS. 3

ABSTRACT. 3

MATERIALS AND METHODS. 4

DESCRIPTIONS. 5

1. Subfamily ANOPLOCEPHALINAE Blanchard, 1891. 5

Genus CTENOTAENIA Railliet, 1893. 5

Genus MOSGOVOYIA Spasskii, 1951. 10

Genus PSEUDOCITTOTAENIA Tenora, 1976. 31

2. Subfamily MONIEZIINAE Spasskii, 1951. 36

Genus CITTOTAENIA Riehm, 1881. 36

3. Species referred to other généra of Anoplocephalidae. 42

Subfamily ANOPLOCEPHALINAE. 42

Genus STRINGOPOTAENIA gen. nov. 42

Subfamily MONIEZIINAE. 46

Genus MONIEZIA. 46

SPECIES INCERTAE SEDIS . 50

SPECIES INQUIRENDAE. 52

DISCUSSION. 53

REFERENCES. 57

DESSINS. 65

Source : MNHN, Paris

UN BEVERIDGE

INTRODUCTION

In an earlier révision of the anoplocephalid cestodes of Australian marsupials (Beveridge, 1976)

it was found difïicult to define the relationships of the généra Progamolaenia Nybelin, 1917, Para-

nwniezia Maplestone and Southwell, 1923 and Phascolotaenia Beveridge, 1976, revised or described

in that work with the généra of the “ Cittotaenia complex ” most similar to them, in particular the

généra Mosgovoyia Spasskii, 1951 and Ctenotaenia Railliet, 1893, as reviewed by Spasskii (1951). Prior

to Spasskii’s (1951) révision of the Anoplocephalidae, the species belonging to these two généra were

placed within the large and obviously composite genus Cittotaenia (Riehm, 1881), as in the monograph

by Baer (1927). Tenora (1976) indicated some of the shortcomings of Spasskii’s earlier work and

rearranged the species in order to overcome the obvious deficiencies.

However, the révisions of both Spasskii (1951) and Tenora (1976) suffer from the disadvantage

that they are based more on a study of the literature rather than of the relevant specimens. Conse-

quently, it was decided to undertake a full révision of the genus Cittotaenia sensu Baer (1927) working

from existing collections of cestodes and providing new descriptions for ail species involved, even

though some of them are extremely well known. In this way it was hoped that some of the difficulties

outlined earlier might be overcome.

Because many of the species formerly placed in the genus Cittotaenia by Baer (1927) hâve since

been transferred to a wide variety of new and resurrected généra in two subfamilies (Spasskii, 1951,

Tenora, 1976) as well a new species having been added to the genus the format of the présent révision is

somewhat unconventional. The généra Cittotaenia Riehm, 1881, Ctenotaenia Railliet, 1893, Mosgovoyia

Spasskii, 1951 and Pseudocittotaenia Tenora, 1976 are dealt with in full. In addition, although the Aus¬

tralian représentatives of the genus Paramoniezia were discussed by Beveridge (1976), Pa. psittacea

(Fuhrmann, 1904) Spasskii, 1951 is also described in the présent work as it originally belonged to the

genus Cittotaenia (see Fuhrmann, 1904,1921, Baer, 1927) and, as indicated by Beveridge (1976), does not

belong in the genus Paramoniezia. Species transferred to généra other than those revised in the

présent paper hâve been described in full and their reclassification discussed and justified in detail.

Species with inadéquate descriptions, incomplète or missing types which cannot be placed

satisfactorily in any known genus are listed as Cittotaenia sensu lato with their original nomenclatural

combinations, but are considered as species incertae sedis or inquirenda.

Source : AANHN, Paris

REVISION OF THE C1TTOTAEN1A COMPLEX

ACKNOWLEDGEMENTS

I should like to thank Prof. A. Aeschlimann for the provision of laboratory facilities and for

access to the parasite collection in his department. Spécial thanks are due to Prof. Cl. Vaucher for

his advice and help during my stay at Neuchâtel. I am indebted to Prof. Dr. B. Hôrning for advice,

specimens and most particularly for access to his very extensive personal library. Thanks are also

due to Dr. R. Lichtenfels for his forbearance and patience with my numerous requests for material

and data on types.

I should also like to acknowledge and thank the following people for the loan of material or for

access to collections in their care :

Dr. C. Bursey, Dr. A. W. Grundmann, Dr. M. Pritchard, Mr. S. Prudhoe, Dr. R. L. Rausch,

Dr. A. A. Rego, Dr. G. D. Schmidt, Mrs. P. M. Thomas and Dr. D. Voth.

The work was carried out whilst the author was in receipt of a C.S.I.R.O. Fellowship.

ABSTRACT

The anoplocephalid cestode généra Citlotaenia Riehm, 1881, Ctenotaenia Railliet, 1893 and Mosgovoyia

Spasskii, 1951 are revised. The genus Ctenotaenia Railliet, 1893 is upheld, with a single species, Ct. marmotae

(Frôhlich, 1802). Ct. citelli (Kirshenblatt, 1939) and Ct. avicola (Fuhrmann, 1897) fall as synonyms of it.

The genus Mosgovoyia Spasskii, 1951 is redefined with three valid species, M. pectinala (Goeze, 1782), M. cte-

noides (Railliet, 1890) comb. nov. and M. variabilis (Stiles, 1895) comb. nov. The species M. perplexa (Stiles,

1895), M. oitana Sawada and Kugi, 1974 and Citlotaenia wittei Baer and Fain, 1955 are treated as synonyms

of M. pectinata (Goeze, 1782). The genus Neoctenotaenia Tenora, 1976 is a synonym of Mosgovoyia Spasskii,

1951. The genus Pseudocittotaenia Tenora, 1976 is retained for P. praecoquis (Stiles, 1895) comb. nov., with

Cittotaenia megasacca Smith, 1951 as its synonym, and P. glandularis sp. nov. created for the species described

by Smith (1951) as Cittotaenia praecoquis (Stiles, 1895). The genus Cittotaenia Riehm, 1881 is retained provi-

sionally within the sub-family Monieziinae Spasskii, 1951, with C. denliculala (Rudolphi, 1804) as the type

species, and Cittotaenia viscaciae (Spasskii, 1951) comb. nov. Cittotaenia psittacea Fuhrmann, 1904 is made

the type and only species of a new genus Stringopotaenia gen. nov. Cittotaenia rhea Fuhrmann, 1904 and C. be-

quaerti Vigueras, 1943 are both transferred provisionally to the genus Moniezia Blanchard, 1891. C. tachyglossi

Johnston, 1913 is considered a species incertae sedis. C. quadrata von Linstow, 1904, C. dratchynskii Roma-

novitch, 1915 and C. krishnai Nama, 1972 are considered species inquirendae.

MATERIALS AND METHODS

The présent révision is based on cestode collections in various muséums and universities as well

as on specimens in the collections of private individuals. The following abbreviations of institution

names are used in the text :

* Institut de Zoologie, Université de Neuchâtel, Suisse.

Source : MNHN, Paris

IAN BEVERIDGE

BM British Muséum (Natural History).

HCIOC Helminth Collection of Instituto Oswaldo Cruz, Rio de Janeiro, Brasil.

LSTM Liverpool School of Tropical Medicine, collection currently on loan to the University

of Neuchâtel.

NUE Nara University of Education, Japan.

UAHC University of Adelaide, Department of Zoology, Helminth Collection currently held

in the South Australian Muséum, Adelaide, Sth. Australia.

UMVS University of Melbourne, School of Veterinary Science, Australia.

UN University of Neuchâtel, Switzerland.

UNMC University of Nebraska, Manter Collection, U.S.A.

USNM United States National Muséum Parasite Collection, U.S.D.A. Beltsville, Maryland,

U.S.A.

UU University of Utah, Dep. of Biology, Sait Lake City, U.S.A.

The number of specimens examined is shown where it is known. In the case of some large

species for which only slide material was available, the number of worms examined is not known.

Muséum numbers referred to in the text under material examined may designate a single specimen

or a sériés of specimens.

Host and geographical records hâve been tabulated for clarity when they are extensive. The

literature cited is not exhaustive as, in the case of the better known species, much of the literature

is of no taxonomie significance. Works with incomplète host or parasite identifications ( eg. Lepus sp.,

Cittotaenia sp.) hâve been disregarded as hâve those with no or vague locality records, unless the host

is reported as new. Works merely repeating earlier literature (where identifiable) hâve been omitted.

Published measurements are tabulated. Synonymies prior to 1896 are those of Stiles (1896),

and only principal references are given. Detailed synonymies of these parasites as far as 1912 can be

found in Stiles and Hassall (1912).

Records given to the présent day are not exhaustive. Works such as general parasitology texts

or papers in which only a passing reference is made to the parasite in question hâve been disregarded.

Because of numerous changes in both host and parasite names, tables summarising host and

geographical data for a parasite hâve original host déterminations listed as synonyms of the current

names and indicate the original name applied to the parasite recorded.

The following texts hâve been used as a basis of host nomenclature : — Ellerman and Morrison-

Scott (1951), Cabrera (1961) and Hall and Kelson (1959).

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMP1.EX

DESCRIPTIONS

Family ANOPLOCEPHALIDAE Kholodkovskii, 1902

1. — Subfamily ANOPLOCEPHALINAE Blanchard, 1891

Genus CTENOTAENIA Railliet, 1893

Ctenotaenia Railliet, 1893 : 278 ; Spasskii, 1951 : 256-275 ; Tenora, 1976 : 9-10.

Cittotaenia Riehm, 1881a : 200 pro parle ; Stiles, 1896 : 170 ; Baer, 1927 : 49 ; Yamaguti, 1959 : 376.

Types species. Ctenotaenia marmotae (Frôhlich, 1802). Only species.

Diagnosis. Cestodes of moderate size. Strobila broad, ribbon-like. Scolex small, unarmed.

Suckers unarmed. Proglottides numerous (over 100 in gravid strobilae), craspedote, greatly extended

transversely. Longitudinal osmoregulatory canals paired. Transverse canal connects left and right

ventral canals at posterior margin of each proglottis. Génital ducts cross longitudinal osmoregu¬

latory canals dorsally. Genitalia paired. Cirrus sac opens to génital atrium anterior to vagina. Cirrus

sac dorsal to vagina on both sides of strobila. Internai and external séminal vesicles présent. Testes

numerous, anterior and posterior to utérus in single band, médial to female genitalia. Séminal récep¬

tacle présent. Ovaries situated in latéral quarters of proglottis medulla. Single transverse utérus

in each proglottis. Utérus does not cross longitudinal osmoregulatory canals. Gravid utérus sac-

like with anterior and posterior diverticula. Pyriform apparatus présent. Parasites of Sciuridae

(Rodentia).

Remarks. The genus Ctenotaenia was resurrected by Spasskii (1951) for Ct. marmotae and

seven other species he whished to distinguish generically from Cittotaenia into which ail had previously

been placed (Baer, 1927). However, even Spasskii (1951) admitted that the taxonomy of the genus

was unsatisfactory and Tenora (1976) subsequently reduced the number of species removing most

to other généra. Ct. asiatica Tokobajev and Erkulov, 1966 was admitted as a valid species by Tenora

and Hôrning (1972) but was later transferred to the genus Paranoplocephala Lühe, 1910, a genus with

a single set of genitalia per proglottis, by Tenora in 1976. The présent révision reduces the number

of valid species to one, Ct. citelli (Kirshenblatt, 1939) and Ct. avicola (Fuhrmann, 1827) falling as

synonyms of Ct. marmotae.

The genus Ctenotaenia differs from Mosgovoyia and Pseudocittotaenia in that the testes are

not entirely posterior to the utérus but occur both anterior and posterior to it. Ctenotaenia differs

from Progamotaenia, Phascolotaenia and Paramonieza (sensu Beveridge, 1976) in that the testes in

Ctenotaenia are restricted to the area between the female genitalia.

Source : MNHN, Paris

IAN BEVERIDGE

Ctenotaenia marmotae (Frôhlich, 1802)

Figs 1-10, Tables 1-3.

Taenia marmotae Frôhlich, 1802 : 77-79, pl. 2, figs 17-20.

Moniezia marmotae (Frôhlich, 1802) Blanchard, 1891 : 187, 444, 461-467, figs 31-35.

Taenia pectinata Goeze, 1782 pro parte, Rudolphi, 1804 : 108.

Cittotaenia pectinata (Goeze, 1782) Stiles and Hassall, 1896b : 407 pro parte, Baer, 1927 : 55.

Cittotaenia pectinata citelli Kirshenblatt, 1939 : 116-128.

Cittotaenia citelli (Kirshenblatt, 1939) Kirshenblatt, 1947 : 115-118.

Ctenotaenia citelli (Kirshenblatt, 1939) Spasskii, 1951 : 263-267, fig. 123.

Cittotaenia avicola Fuhrmann, 1897 : 108-117, pl. 5, figs 1-8.

Ctenotaenia avicola (Fuhrmann, 1897) Spasskii, 1951 : 260-263, fig. 122.

Types. From Marmota marmota Linnaeus, 1758, location unknown.

Material examined.

From Marmota marmota Linnaeus, 1758 : 1 specimen, Briançon, France, September 1887 from

collection of R. Blanchard, now in collection of C. Joyeux, UN ; 2 specimens without collection data,

UN ; 80 specimens, Bevers, Switzerland, 14 July 1964, G. Zelenka, UN.

From Citellus major erythrogenys Brandt, 1841 : 6 specimens, Siberia, U.S.S.R., no other data,

UN.

From Ancw sp. : types of Cittotaenia avicola, no other data, UN.

Description. Large, broad, ribbon-like worms, anterior extremity tapering markedly. Scolex

small, globular, distinctly demarcated from strobila in contracted specimens, rounded anteriorly and

merging into strobila in relaxed specimens. Neck absent. Proglottides craspedote narrow, straight-

edged vélum overhanging adjacent proglottis. Mature proglottides much wider than long with

approximate length : width ratio 1 : 10-1 : 17. Gravid proglottides with approximate length : width

ratio 1 : 10-1 : 32. Longitudinal musculature weakly developed, arranged in two circles of muscle

bundles in cortex. Outer muscle bundles oval or circular in transverse section, small, composed of 5-35

muscle fibres. Bundles of inner ring smaller, less regularly arranged, composed of 2-10 muscle fibres.

Transverse muscles filiform, forming narrow band internai to longitudinal muscles. Dorso-ventral

muscle fibres fine, Crossing medulla and cortex at regular intervals. Ventral longitudinal osmoregul-

atory canal wider than dorsal canal, situated médial to dorsal canal. Transverse canal connects left

and right ventral canals at posterior margin of each proglottis. Dorsal and ventral canals of each

side of strobila loop laterally at level of suckers then return towards mid-line. Dorsal canals from

each side join in mid-line and continue anteriorly as pair of ducts, similarly ventral canals of both

sides. Annular anastomosis connects four ducts at anterior extremity of suckers. Génital atrium

prominent, forming large ovoid sinus, opening to exterior in posterior half of latéral proglottis margin,

dividing margin in ratio 1 : 1.5- 1 : 3. Cirrus sac elongate, pyriform, thick-walled, usually reaching

or Crossing longitudinal osmoregulatory canals. In immature proglottides, cirrus sac may not reach

osmoregulatory canals. Cirrus long, coiled within cirrus sac, distal région unarmed, proximal région

heavily armed with rows of spines. On everted cirri, spines longest at distal extremity, diminishing

in size near mid-region. Spines on proximal région very short. Cirrus surrounded by prostatic cells.

Large, ovoid internai séminal réceptacle présent. External séminal vesicle elongate, slightly coiled.

Vas deferens narrow. Testes numerous, distributed in 3-6 horizontal and 2-4 dorsoventral layers.

Testes invariably lie in single field between female génital glands on dorsal aspect of medulla. Occa-

sionally testes overlie médial extremity of ovary but never lie latéral to ovary. Testes situated both

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

anterior and posterior to utérus. Distal vagina forms ovoid atrium with hairy lining. Mid-region

of vagina tube-like, thin-walled. Junction of tube with atrium surrounded by muscular sphincter.

Vagina merges into elongate, pyriform séminal réceptacle, extending along posterior margin of proglot-

tis, dorsal to ovary, to poral margin of vitellarium. Ovary fan-shaped, composed of numerous clavate

lobules, situated on ventral aspect of medulla. Vitellarium ovoid or reniform, lobulate, posterior

and dorsal to ovary. Mehlis’ gland spherical, posterior to ovary, dorsal to vitellarium. Utérus tube-

like, transverse, single, in middle of proglottis, passing dorsal to ovary, anterior to vitellarium and

séminal réceptacle. Gravid utérus with numerous anterior and posterior diverticula. Utérus never

extends laterally beyond longitudinal canals. Egg spherical, thick-shelled. Pyriform apparatus

terminâtes in two elongate horns.

Vestigial, supernumerary vitellaria présent in many proglottides, of variable size, posterior to

utérus, connected to utérus in some instances by narrow duct. One to three, usually two, extra

vestigial vitellaria per proglottis.

Remarks. Although a comparatively well-known species, the descriptions and illustrations

of it in the literature are rather poor. Stiles (1896) gave a brief description of the parasite based on

Blanchard’s material, and his description was subsequently used by Spasskii (1951), though the figures

which Spasskii reproduced from Stiles’ work and labelled as Ct. marmotae (Fg. 120, p. 258) are in fact

Stiles’ drawings of Cittotaenia denticulata (Rudolphi, 1804). Spasskii and Shalajeva (1961) gave an

improved description of the parasite based on material from Marmota bobak, whilst Tenora and Hôrn-

ing (1972) gave a detailed bibliographie review of the species but did not deal in detail with its morpho-

logy. Tenora (1976) further discussed the parasite but some of the morphological observations he

offers appear to be in error. He commented on the distribution of the testes thus (Tenora, 1976, p. 9) :

“ The testes in the strobila of C. marmotae and C. citelli are situated always above the tubular utérus.

They fill the upper half of proglottides and in fully formed hermaphrodite proglottides they may even

reach under the utérus ”. The statement appears inconsistent and as iliustrated in fig. 10, testes

lie both anterior and posterior to the utérus, hence Tenora’s (1976) observations cannot be accepted.

Tenora (1976) also used this feature as a differentiating character in a key to the généra of the Ano-

plocephalinae (pp. 15-16). In some mature proglottides examined during the présent work, the trans¬

verse osmoregulatory canal was greatly distended, displacing the testes to the anterior half of the

proglottis, much as is shown in Plate il fig. 7 of Stiles (1896). In the case mentioned testes were

still distributed both anterior and posterior to the utérus, but the utérus was not readily seen and it

is possible that Tenora (1976) was looking at a similar abnormality in his material and mistook the

transverse osmoregulatory canal for the utérus.

Blanchard (1891) described and iliustrated additional osmoregulatory canals to those found

in the présent study. However, Berthoud’s (1966) description and drawing of the osmoregulatory

System agréé with the current description as do the observations of Tenora and Hôrning (1972).

In the current révision, Ct. citelli (Kirshenblatt, 1939) is considered to be a synonym of Ct. mar¬

motae. Originally described from Citellus citellus xanthoprymnus by Kirshenblatt (1939) the species

was subsequently found to be widely distributed in Russia, occurring also in Citellus pygmaeus and

C. citellus dauricus and was described in some detail by Spasskii (1951). Although the types were

not available for study, specimens from Citellus major erythrogenys from Siberia, together with Spasskii’s

detailed description permit the re-assessment of the taxonomie position of the species. Spasskii

(1951) concluded that Ct. citelli was very similar to Ct. marmotae but that they differed in two significant

respects, namely that the cirrus of Ct. citelli was heavily armed whilst that of Ct. marmotae was not,

and that there were more testes per proglottis in Ct. citelli. The cirrus of Ct. marmotae is in fact heavily

armed, as is shown in figs 2 and 5 though this feature was not mentioned by Stiles (1896) upon whose

description Spasskii (1951) relied. Spasskii and Shalajeva (1961) subsequently redescribed Ct. mar¬

motae from Marmota bobak and iliustrated a heavily armed cirrus. Consequently the supposed lack of

cirrus armature in Ct. marmotae cannot be used to distinguish it from Ct. citelli.

The number of testes per proglottis in Ct. citelli has been recorded as 150-200 compared with

100-160 for Ct. marmotae (Table 1). In the présent révision the number of testes per proglottis was

Source : MNHN, Paris

ii i

UH J] J

s«i=S «a . s g g 1 | - „

MÎ il! i îiîïîii i J Irai!

>nly available.

Table 2. — TESTIS NUMBER IN 5 PROGLOTTIDES FROM EACH OF 4 STROBILAE OF

CTENOTAENIA MARMOTAE.

HOST

Marmola marmota L. Citellus erythrogenys

(Brandt)

Strobila

3 4

No. testes

166

140

128 115

Per

133

135

163 138

Proglottis

129

125

187 116

140

159

166 104

151

116

165 145

Table 3. —

HOST RECORDS

OF CTENOTAENIA MARMOTAE

Host

Localit

REFERENCE

Marmola marmota

Briançon, France

Blanchard (1891)

Linnaeus, 1758

Vaud, Valais, Switzerland

Galli Valerio (1918, 1923, 1926, 1930a, b,

Vaud, Switzerland

Switzerland

Valais, Switzerland

Valais, Uri, Graübunde, Fribourg,

1931, 1933, 1935)

Fuhrmann (1926)

Schweitzer (1949).

Schweitzer and Burgisser (1949)

Bouvier et al. (1959)

Hôrning (1969)

Marmota marmota

Berne, Switzerland

Swiss Alps

Austria

Pôllatal, Austria

Czechoslovakia

Hôrning and Tenora (1971)

Tenora and Hôrning (1972)

Jettmar and Anschau (1951)

Sixl (1975)

Tenora (1961)

latirostris

Kratochil, 1961

Marmota marmota

Kazakhstan, USSR

Gvozdev and Kapitonov (1966)

baibacina

Brandt, 1843

Marmota bobak

Kazakhstan, USSR

Gvozdev and Kapitonov (1966)

Muller, 1776

Marmota bobak

Kazakhstan, USSR

Spasskii and Shalajeva (1961)

tshaganensis

Bashanov, 1930

Citellus citellus

Armenia, USSR

Kirshenblatt (1939)

xanthoprymnus

Armenia, USSR

Akumian (1948)

Bennett, 1835

Citellus citellus

USSR

Kirshenblatt (1947)

dauricus

Brandt, 1843

Citellus pygmaeus

USSR

Kirshenblatt (1947),

Pallas, 1179

Citellus suslicus

Ukraine, USSR

Spasskii (1951)

Kadenatsii (1957) (in Sharpilo, 1966)

Güldcnstaedt, 1770

Ukraine, USSR

Sharpilo, 1961

« Anas sp. » ?

Byelorussia, USSR

Switzerland

Arzamasov et al. (1966)

Fuhrmann (1897)

Source : MNHN, Paris

IAN BEVERIDGE

not found to differ significantly between the two forms. The testes in these cestodes are not always

easy to count accurately as they lie in several layers and the difliculties are increased by the thickness

of the cortex and hence the general opacity of the worms. The number of testes in a sériés of adjacent

proglottides from several strobilae indicates the high level of variation which can occur within

a single strobila (Table 2). Although statistical data for anoplocephalids are limited, Beveridge (1974)

has shown that in the case of Taenia pisiformis (Bloch, 1780), at least 50 % of the variability in testis

number in a population can be ascribed to within-strobila variation. Since a similar high level of variab¬

ility occurs in strobilae of Ct. marmotae it is not considered valid to distinguish Ct. citelli on the basis

of number of testes per proglottis. Table 1 also indicates that no other valid quantitative différences

occur between these taxa. Spasskii and Shalajeva (1961) argued that Ct. marmotae and Ct. citelli

should be maintained as separate since they inhabit different host généra. However, both the genus

Marmota and the genus Citellus belong to the family Sciuridae and the ranges of the two généra even

in the présent day are contiguous or overlap so that allopatry cannot be used as a criterion to distinguish

the species. Consequently Ct. citelli has been treated as a synonym of Ct. marmotae.

Cittotaenia avicola Fuhrmann, 1897 was described from specimens held in the Geneva Natural

History Muséum and labelled initially as “ Taenia lanceolala Goeze ” and as having been collected

from a duck, Anas sp. Spasskii (1951) transferred Fuhrmann’s species to the genus Ctenotaenia noting

its close similarity with Ct. citelli but maintaining it as a valid species because of différences in the

number of testes per proglottis. It is évident from a re-examination of the types that Ct. avicola

is a synonym of Ct. marmotae, a mistake presumably having been made in the original label. Fuhr¬

mann (1897) commented on the close similarity of his new species to Ct. marmotae (then Cittotaenia

marmotae) but decided, erroneously, that the cirrus sac was more like that of Mosgovoyia pectinata

(Goeze, 1782) (formerly Cittotaenia pectinata (Goeze, 1782)) and consequently placed Ct. avicola as a

new species intermediate between the “ marmotae ” and “ pectinata ” groups of the genus Cittotaenia.

There are two other records of Ct. avicola in the literature. Meggitt (1927) identified a cestode

from Sommett’s jungle fowl, Gallus sonnerati Temminck, 1813 as “ C. ? arvicola ” apparently intending

C. avicola, however the specimen was incomplète and could not be fully identified. Southwell (1930)

reported that he had referred to this species some specimens from an Impeyan pheasant, Lophophorus

impejanus (Latham, 1790), but that on subséquent examination an armed rostellum was detected

and the cestodes were re-identified as Cotugnia margareta Beddard, 1916. He suggested that Meggitt’s

specimens might also belong to this species. Both records can now be disregarded as Ct. avicola falls

as a synonym of Ct. marmotae.

Genus MOSGOVOYIA Spasskii, 1951

Mosgovoyia Spasskii, 1951 : 286-297 ; Yamaguti, 1959 : 383 ; Tenora, 1976 : 17.

Cittotaenia Riehm, 1881a pro parte Joyeux and Baer, 1961 : 550..

Neoclenotaenia Tenora, 1976 : 13.

Type Species. Mosgovoyia pectinata (Goeze, 1782).

Diagnosis. Cestodes of moderate size. Strobila broad, ribbon-like. Scolex small, unarmed.

Suckers unarmed. Proglottides numerous (over 100 in gravid strobilae), craspedote, greatly extended

transversely. Longitudinal osmoregulatory canals paired. Transverse canal connects left and right

ventral canals at posterior margin of each proglottis. Génital ducts cross longitudinal osmoregu¬

latory canals dorsally. Genitalia paired. Cirrus sac opens to génital atrium anterior to vagina.

Cirrus sac dorsal to vagina on both sides of strobila. Internai séminal vesicle présent. External

séminal vesicle absent. Testes numerous, entirely posterior to utérus in single band or two groups,

either restricted to area between female genitalia or extending laterally beyond them. Distal vagina

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

surrounded by glandular cells. Séminal réceptacle présent. Ovaries situated in latéral quarters of

proglottis medulla. One or two transverse, tubular uteri per proglottis, either restricted to medulla

or Crossing longitudinal osmoregulatory canals dorsally, terminating posterior to cirrus sac and vagina.

Gravid utérus sac-like with or without anterior and posterior diverticula Pyriform apparatus présent.

Parasites of Leporidae (Lagomorpha) and rarely Sciuridae (Rodentia).

Key to species of Mosgovoyia

1. Utérus in mature proglottides reaching but never Crossing longitudinal osmoregulatory canals.

peclinata

Utérus in mature proglottides invariably extending laterally beyond osmoregulatory canals.... 2

2 (1). Testes extend laterally beyond female genitalia, usually in two groups ; cirrus sac broad, ellipsoidal.

clenoides

Testes restricted to area between female genitalia, almost invariably in single band ; cirrus sac narrow,

elongate. variabilis

Remarks. The genus Mosgovoyia was established by Spasskii (1951) for Cittotaenia peclinata

(Goeze, 1782) and C. perplexa (Stiles, 1895). Spasskii (1951) also designated a specimen of C. pecti-

nata described from a viscaca by Joyeux and Dollfus (1931) as a new species, Mosgovoyia viscaciae.

The taxonomie history of the former two species is of some importance to the présent révision and

hence it will be reviewed in some detail.

In Stiles’ révision (Stiles, 1896), the species Cittotaenia pectinata, C. perplexa and C. variabilis

(Stiles, 1895) constituted the so-called “ pectinata ” group within the genus, being characterised by

the narrow, elongate cirrus sac. Hall in 1908 added a further species, C. mosaica, differentiated from

C. pectinata by the small size of the cirrus sac and by mosaic markings on the strobila. However,

Douthitt (1915) found that Stiles was mistaken in his measurements of the cirrus sac of C. perplexa

and that C. mosaica was a synonym of C. perplexa. Douthitt (1915) also established the subspecies

C. pectinata americana for American représentatives of the species.

Baer (1927) recognised only three valid species of Cittotaenia in lagomorphs, namely C. denti-

culata (Rudolphi, 1804), C. ctenoides (Railliet, 1890) and C. pectinata, considering the last named species

to be extremely variable and placing as synonyms to it not only C. variabilis and C. perplexa but

also C. marmotae and C. quadrata von Linstow, 1904. Baer (1927) also suppressed the subspecies

C. pectinata americana.

Rees (1933a) gave a detailed account of the morphology of C. pectinata collected in Wales and

designated her material as a new subspecies C. pectinata europea. Because of the confusion arising

from Baer’s (1927) révision of the “ pectinata ” group, Arnold (1938) undertook yet another review

based on new material and reinstated C. perplexa and C. variabilis as well as C. pectinata americana

as valid taxa.

Spasskii (1951) removed the type species of the genus Cittotaenia, C. denticulato, to the subfamily

Monieziinae because it possesses a reticulated utérus, and transferred C. ctenoides and C. variabilis

to the resurrected genus Clenotaenia, thereby breaking up the so-called “ pectinata ” group. The

genus Mosgovoyia, created to contain C. pectinata and C. perplexa, was distinguished from other généra

by the occurrence of testes posterior to the utérus, extending laterally to the female genitalia on either

side of the strobila and by the presence of accessory osmoregulatory canals connecting the transverse

canals of adjacent proglottides. Spasskii (1951) also noted that a nominal subspecies of M. pectinata

had never been designated and hence proposed the name C. pectinata pectinata for the European form

of the species.

The présent révision fails to support Spasskii’s (1951) arrangement of the genus Mosgovoyia although

retaining it as a valid genus. It is to Spasskii’s crédit that he recognised the composite nature of the

genus Cittotaenia sensu Baer, 1927 and removed C. denticulata making a révision of the genus possible.

However, the way in which Spasskii (1951) distributed the remaining species of Cittotaenia between

Source : MNHN, Paris

IAN BEVERIDGE

the généra Ctenotaenia and Mos'govoyia is inconsistent. In the type species of the genus Ctenotaenia,

Ct. marmotae, the distinguishing characteristic is the distribution of the testes both anterior and posterior

to the utérus but limited to the area between the female genitalia. By contrast, in members of the

genus Mosgovoyia sensu Sapsskii (1951), the testes lie in a band entirely posterior to the utérus but

extend laterally beyond the female genitalia. In support of this generic distinction, Spasskii (1951)

argues convincingly that the genus Mosgovoyia has been derived by a doubling of the genitalia from

species of the genus Schizorchis Hansen, 1948 in which the testes lie entirely posterior to the utérus

and the hosts of which are members of the genus Ochotona, also Lagomorpha. The genus Ctenotaenia

was considered to hâve arisen by doubling of the genitalia of species of the genus Paranoplocephala

Lühe, 1910 in which the testes lie aporal to the female genitalia. Species of both Ctenotaenia and

Paranoplocephala parasitise rodents. If these arguments from morphological features, host distribu¬

tions and possible phylogenetic afïlnities are valid, then the other lagomorph parasitising species

C. variabilis and C. ctenoides which Spasskii (1951) transferred to the genus Ctenotaenia but which also

hâve the testes situated entirely posterior to the utérus should also be transferred to the genus Mos¬

govoyia■ This change has been made in the présent révision. The revised genus Mosgovoyia is dis-

tinguished from related généra principally by the distribution of testes, but also by the morphology

of the female génital ducts since the distal vagina is surrounded by layers of glandular cells and the

vagina leads into a thin-walled, elongate séminal réceptacle. Other anoplocephaline généra ( Pro-

gamotaenia Nybelin, 1917, Phascolotaenia Beveridge, 1976) also hâve morphologically distinctive

female génital ducts which are of considérable taxonomie use.

M. pectinata differs from the other species currently placed in the genus by the possession of

accessory osmoregulatory canals and Spasskii (1951) used it as a generic character. Although the

features of the osmoregulatory System hâve been considered as having considérable phyletic, and

hence taxonomie significance (Wardle, 1932), giving them generic significance in the présent classifi¬

cation leads to the impossibility of successfully assimilating several other characters which may be

of at least equal importance. Although the existence of accessory canals in Schizorchis ochotonae

Hansen, 1948 as well as in M. pectinata was one factor involved in the choice of this feature as a generic

character, they do not occur in ail species of Schizorchis. They apparently exist in S. ochotonae and

S. yamashitai Rausch, 1963 but hâve not been described in either S. caballeroi Rausch, 1960 or S. altaica

Gvozdev, 1951 (Hansen, 1948, Spasskii, 1951, Rausch, 1960, 1963, Rausch and Ohbayashi, 1974).

In order to avercome the difficulty, Tenora (1976) created the genus Neoctenotaenia for the species

M. ctenoides and M. variabilis. In the présent révision the presence of accessory osmoregulatory

canals is not considered a valid generic criterion and hence the genus N eoctenotaenia is treated as a

synonym of Mosgovoyia.

Mosgovoyia pectinata (Goeze, 1782)

Figs 11-25, Tables 4-5

? Taenia acutissima Pallas, 1781 : 75-81, pl. 3, fig. 25.

Taenia leporina Rudolphi, 1810 : 82.

Taenia pectinata Goeze, 1782 : 363-368, pl. 27, figs 7-13.

? Alyselminthus pectinatus (Goeze, 1782) Zeder, 1800 : 246-249.

? Halysis pectinata (Goeze, 1782) Zeder, 1803 : 332.

Dipylidium pectinatum (Goeze, 1782) Riehm, 1881a : 200, 575-583, pl. 5, figs 4, 14, pl. 6, figs 4, 7.

Moniezia pectinata (Goeze, 1782) Blanchard, 1891 : 187, 445, 450-452, 457-460, figs 26-30.

Ctenotaenia pectinata (Goeze, 1782) Railliet, 1893 : 278-279.

Cittotaenia pectinata (Goeze, 1782) Stiles and Hassall, 1896b : 407.

Cittotaenia pectinata americana Douthitt, 1915 : 47, pl. 16, fig. 45.

Cittotaenia pectinata europea Rees, 1933a : 250.

Cittotaenia pectinata septentrionalis Romanovitch, 1915 : 453.

Mosgovoyia pectinata (Goeze, 1782) Spasskii, 1951 : 287-295, figs 135-139.

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Mosgoaoyia pectinata pectinata Spasskii, 1951 : 289-295.

Mosgoaoyia pectinata americana (Douthitt, 1915) Spasskii, 1951 : 295.

Mosgoaoyia pectinata europaea (Rees, 1933) Yamaguti 1959 : 383.

Ctenotaenia perplexa Stiles, 1895 : 345.

Cittotaenia perplexa (Stiles, 1895) Stiles and Hassall, 1896b : 407.

Mosgoaoyia perplexa (Stiles, 1895) Spasskii, 1951 : 295-296, fig. 140.

Cittotaenia bursaria von Linstow, 1906 : 164, 184, pl. 2, figs 39-40.

Cittotaenia mosaica Hall, 1908 : 691-699, figs 1-6.

Cittotaenia wittei Baer and Fain, 1955 : 11-13, figs 2-3.

Neoctenotaenia wittei (Baer and Fain, 1955) Tenora, 1976 : 13.

Mosgoaoyia oitana Sawada and Kugi, 1974 : 264-266, figs 9-15.

Types. Whereabouts unknown.

Material examined.

From Oryctolagus cuniculus Linnaeus, 1758. 5 specimens Milford, Surrey, England, 8 August

1974, A. Mead-Briggs, UN and UMVS ; 1 specimen Essex, England, 9 June 1963, R. J. Knowles,

BM 1963.7.8.75 ; Cardiganshire, Wales, G. Rees, BM ; Iniskea North, Mayo, Ireland, R. J. Knowles,

September 1967, BM 1970.4.17.27-28 ; 1 specimen, Newton, Mayo, Ireland, September 1969, R. J. Know¬

les, BM 1970.4.17.29 ; Germany, no other data, USNM 1234-1237 ; 1 specimen, N. Ireland, no date,

H. Shatowy, BM 1976.5.14.3.

From Lepus europaeus Pallas, 1778. 1 specimen, Nancy, France, 7 December, 1905, C. Joyeux,

UN.

From Lepus europaeus cravashayi de Winton, 1899 (syn. L. crawshayi). 1 specimen Mukana,

Zaïre, no date, UN (type of Cittotaenia wittei).

From Lepus europaeus saxatilis Cuvier, 1823. (syn. L. saxatilis). 2 specimens, Transvaal,

South Africa, 8 March 1957, F. Zumpt, BM 1958.8.4.135-137.

From Lepus timidus Linnaeus, 1758. 1 specimen Isle of Mull, Scotland, 22 June 1953, BM

1963.11.26.11-20; 1 specimen Anagh, Mullet, 28 August 1969, R. J. Knowles, BM 1970.4.17.30;

4 specimens, Bernese Oberland, Switzerland, 21 November, J. Codonrey, UN.

From Lepus capensis Linnaeus, 1758. Kenya, no other data, USNM 54538 ; Sudan, no other

data, USNM 54507.

From Lepus nigricollis Cuvier, 1823. Nedenkuni, Sri Lanka, no other data, UN (types of

Cittotaenia bursaria).

From Lepus nigricollis ruficaudatus Geoffroy, 1826. (syn. L. ruficaudatus). 10 specimens,

Songara, India, November 1920 ?, ? T. Southwell, LSTM.

From Lepus americanus (Erxleben, 1777). Maine, USA, no other data, USNM 44683 ; Alaska,

USA, no other data, USNM 30098 ; Ontario, Canada, 19 July 1932, D. G. Mackintosh, USNM 42519 ;

Frank’s Bay, Ontario, Canada, 20 June, 1932, D G. Mackintosh, USNM 42520 ; 1 specimen St. Edward’s

Is., Canada, C. Bursey, UN.

From Lepus americanus macfarlani Merriam, 1900. 1 specimen, Mile 81, Taylor Highway,

Alaska, 10 August 1963, R. L. Rausch ; 1 specimen, same locality and collector, 11 August 1963 both

in Colin. R. L. Rausch ; 1 specimen Willowlake, Mile 88 Richardson Highway, Alaska, 15 February

1964, R. L. Rausch, UN.

From Lepus townsendi Bachman, 1859. Oklahoma, USA, no other data, USNM 59076 ; Albany

County, Wyoming, USA, R. F. Honess, no other data, USNM 59076.

From Lepus californicus Gray, 1837. Idaho, USA, no other data, USNM 59259.

From Lepus californicus melanotis Mearns, 1890 ; 12 specimens, Colorado, USA, 1972-1973, P. C. Brit-

tain, D. R. Voth, UN, USNM 74287.

From Lepus californicus deserticola Mearns, 1898. 3 specimens, Dugway, Tooele County, Utah,

USA, 22 March 1960, A. W. Grundmann, UU 1022 ; 3 specimens, same locality, 14 November 1951,

Source : MNHN, Paris

IAN BEVERIDGE

J. M. Butler, UU 1040 ; 2 specimens, same locality, 21 March 1967, Foster and A. W. Grundmann,

UU 1042 ; 2 specimens, Stansbury Is., Great Sait Lake, Utah, USA, March 1960, A. W. Grundmann,

UU 8-2.

From Lepus othus othus Merriam, 1900. 2 specimens, Nome, Seward Peninsula, Alaska, 1 October

1960, F. H. Fay, Colin. R. L. Rausch.

From Sylvilagus floridanus (Allen, 1890). Maryland, USA, no other data, USNM 17449 ;

1 specimen, Bowie, Maryland, USA, A. Hassall, UN (cotype of Cittotaenia perplexa).

From Sylvilagus floridanus mearnsi (Allen, 1890). 1 specimen, East Lansing, Michigan, USA,

18 April 1946, R. L. Rausch in colin. R. L. Rausch.

From Sylvilagus nuttalli grangeri (Allen, 1895). Burch Creek, Montana, USA, no other data,

USNM 30957, 30960 ; 14 slides Albany County, Wyoming, USA, R. F. Honess, USNM 59073 ; 3 speci¬

mens, 30 miles east of Salina, Utah, USA, 17 May 1957, A. W. Grundmann, UU 157-6.

From Spermophilus variegatus (Erxleben, 1777). 4 specimens, Red Canyôn, 30 miles east of

Salina, Utah, USA, 15 May 1958, A. W. Grundmann, UU 157-1.

Host unknown. Slides of serial sections, Rutshuru, Zaïre, April 1937, J. Ghesquière, UN

(described by Mahon, 1954).

? Lepus sp. 1 slide, ? France, collection of R. Blanchard, now in colin, of C. Joyeux, UN.

Description. Large, broad, ribbon-like to small, leaf-shaped worms. Scolex small, rounded,

merging imperceptibly into unsegmented neck région of relaxed worms. Neck absent in contracted

specimens, scolex dilficult to distinguish. Proglottides greatly extended transversely, craspedote.

Mature proglottides with approximate length : width- ratio of 1 : 7-1 : 8. Gravid proglottides with

ratio 1 : 4-1 : 5. Longitudinal muscle poorly developed, forming two irregular rows of muscle bundles

around inner margin of cortex. Inner bundles much smaller and with fewer fibres than outer bundles.

Transverse muscle fibres filiform, in band internai to longitudinal muscles. Dorso-ventral muscles

fine, single, Crossing cortex and medulla at irregular intervals. Longitudinal osmoregulatory canals

paired. Ventral canal wider than dorsal canal, situated medially or directly ventrally to it. Transverse

canal connects left and right ventral osmoregulatory canals at posterior margin of each proglottis.

Numerous accessory canals connect transverse canals of adjacent proglottides. At level of suckers,

dorsal canals from each side of strobila join, as do ventral canals. Short dorso-ventral anastomosis

connects common dorsal and common ventral duct. Génital atrium of insignificant size, situated

in middle of latéral proglottis margin of mature proglottides, dividing margin in ratio 1 : 1 ; situated

in posterior third of margin in gravid proglottides dividing margin in ratio 2:1. Cirrus sac narrow,

elongate, highly variable in size, always reaching latéral nerve, usually reaching or extending beyond

longitudinal osmoregulatory canals. Cirrus narrow, frequently coiled, armed with minute bristles.

Armature very difïicult to detect except in sections. Small, elongate internai séminal vesicle présent.

Vas deferens narrow, thin-walled, coiled, following course of vagina and séminal réceptacle, dividing

into two branches at level of vitellarium. Each branch passes transversely dividing into vasa efferentia.

External séminal vesicle absent. Testes numerous, entirely posterior to utérus between longitudinal

osmoregulatory canals, on dorsal aspect of medulla, in 1-3 transverse and 1-3 horizontal rows. Testes

extend beyond female genitalia on both sides of strobila, rarely présent directly posterior to vitellarium.

Testes between female genitalia usually in unbroken band, sometimes in two separate groups. In

some strobilae most mature proglottides hâve testes in two groups. In other strobilae proglottides

with single field or two groups of testes alternate irregulary. Testes persist in near-gravid proglottides.

Vagina narrow, lined internally with bristles, surrounded externally by layers of glandular cells.

Vagina curves anteriorly and medially with cirrus sac, leads to elongate, thin-walled séminal réceptacle

without hairy lining or glandular cells. Ovary fan-shaped, composed of numerous clavate lobules,

on ventral aspect of medulla. Vitellarium oval or reniform, lobulate, posterior and dorsal to ovary.

Mehlis’ gland spherical, anterior to vitellarium, dorsal to ovary Utérus tube-like, transverse, usually

single, occasionally double. Utérus does not cross longitudinal canals in tubular stage. Lies in

middle of proglottis, dorsal to ovary, anterior to Mehlis’ gland. Gravid utérus sac-like with numerous

anterior and posterior diverticula. Rarely crosses longitudinal osmoregulatory canals, usually displaces

Source : MNHN, Paris

REVISION OF THE C1TTOTAEN1A COMPLEX 15

them towards poral margin. Egg spherical with thick refractile shell. Fine sub-shell and inner

membranes présent. Pyriform apparatus terminating in two elongate horns.

Remakes. Little can be added to the descriptive anatomy of M. pectinala following the

detailed accounts by Rees (1933a) and to a lesser extent by Spasskii (1951). However, some minor

différences do exist between descriptions. Both Rees (1933a) and Spasskii (1951) considered the

cirrus to be unarmed, but, in some whole mounts as well as in serial sections it is possible to distinguish

minute bristles. They were not seen on everted cirri and the difficulty in detecting them probably

accounts for the earlier descriptions of the cirrus being unarmed. An external séminal vesicle was

both described and illustrated by Spasskii (1951). The structure is présent only in occasional proglot-

tides and is due simply to dilatation of the distal vas deferens by sperm. In the above description,

the external séminal vesicle is stated to be absent.

Measurement are given by many authors for neck length in M. pectinata. There is no obvious

distinction between the scolex and the neck in this species, though in relaxed specimens there exists

an unsegmented région between the posterior margin of the suckers and the first signs of segmentation.

The length of this région was used in the présent révision as the “ neck ” length, however, it is of no

systematic use because of its dependence on the State of relaxation of the specimen. The anterior

end of a type specimen of Cittotaenia bursaria von Linstow, 1906 (now a synonym of M. pectinata)

illustrâtes (fig. 12) that in a severely contracted specimen, the neck is entirely absent and it is virtually

impossible to measure the size of the scolex.

A detailed description of the egg and of the development of the pyriform apparatus was given

by Rees (1933b). In one specimen of M. pectinata from Canada, irregular cavities were noted in the

pyriform apparatus at the base of the arms. (Figs 24, 25).

In spite of a detailed knowledge of the general anatomy of the parasite, considérable deficiencies

exist in the information available on variation, particularly in quantitative characters and their bearing

on the systematics of the species, as instanced in the discussion below of the various taxa placed as

synonyms of M. pectinata.

The subspecies of M. pectinata hâve, in the présent révision, been supressed. Douthitt (1915),

in erecting the sub-species americana, drew attention to différences in worm size and proglottis number

between it and the European form. Rees (1933a), designating the material she studied from Wales

as C. pectinata europea, noted several différences from the descriptions of American specimens, namely

the smaller scolex, the presence of a neck and the greater length of the strobila, the narrow unbranched

dorsal osmoregulatory canal, the greater number of testes and the lack of an external séminal vesicle.

Rees (1933a) however did not personally examine American specimens and some of the différences

she enumerated are due to errors in early descriptions. Arnold (1938) compared sériés of the two

forms and considered C. pectinata americana a valid sub-species though the only consistent différences

he gave were those of strobilar size and proglottis number. Both characters are notoriously variable

in cestodes, and examination of Table 4 shows that, according to the literature, European specimens

may achieve a greater size, but the majority of European specimens examined in the présent révision

were no bigger and did not contain more proglottides than American specimens. Since ail measure-

ments of internai organs also overlap, it is concluded that neither the two subspecies mentioned above,

nor M. pectinata pectinata Spasskii, 1951, created to replace C. pectinata europea as the nominal sub¬

species, can validly be maintained and they are therefore placed as synonyms of M. pectinata.

Cittotaenia wittei Baer and Fain, 1955 was differentiated from M. pectinata because of the

glandular investment of the vagina and the strongly lobed utérus in the new species. A detailed exa¬

mination of the types of C. wittei revealed no significant différences from M. pectinata and it is therefore

considered a synonym of the latter species.

Mosgovoyia oitana Sawada and Kugi, 1974 was separated from M. pectinata on account of the

greater sized scolex, the larger ovary, the position of the génital pores in the posterior half of the

latéral proglottis margin and the number of uterine diverticula. The position of the génital pores

can vary within a single strobila and can be found in the middle of the latéral proglottis margin in

mature proglottides and in the posterior third of gravid proglottides in the same strobila. It is there-

Source : MNHN, Paris

LAN BEVERIDGE

fore not a distinctive charactèr. Comparison of the quantitative data in Table 4 shows that none

of the measurements used by Sawada and Kugi (1974) is successful in separating M. oitana from

M. pectinata and hence M. oitana falls as a synonym.

M. perplexa (Stiles, 1895) is also regarded as a synonym of M. pectinata. Stiles (1896) differen-

tiated M. perplexa (then Cittotaenia perplexa) by the size of the cirrus sac, the length of the vagina

and the fact that in M. perplexa, the testes are arranged in two latéral groups in mature proglottides

rather than in a single field. Subsequently, Arnold (1938) maintained the validity of M. perplexa,

differentiating it also on total length, position of appearance in the strobila of the génital primordia

and the location of ovarian follicle development. The validity of the two species was also maintained

by Honess (1963) working with new collections, though he concluded that the criteria of séparation

involving length of strobilae and appearance of organ Systems were not reliable since they were highly

variable or depended upon the technique of préservation. He was, however, able to separate the

two species on the features of the scolex and neck.

A re-examination of the range of variability of most of these characters indicates that few

are as reliable as has been stated in the past. The features of the scolex and neck région of M. pec¬

tinata are dépendent upon the method of fixation and are extremely variable as shown in figs 11-12.

The “ width ” of the neck cannot be used, as Honess (1963) claimed, to separate the species as its

width will dépend upon the State of relaxation of the specimen or may be entirely absent. Honess

(1963) in fact claimed that Arnold’s (1938) drawings of the scoleces of M. pectinata and M. perplexa had

been misdesignated as they showed exactly the opposite features to those described by Honess (1963)

for the two species. However, Stiles (1896) indicated that the types of M. perplexa were contracted

and hence Arnold’s figure may be taken as correct. The extent of variability seen in specimens of M. pec¬

tinata in the présent review includes both forms used by Honess (1963) to separate the two species.

As stated by Honess (1963), the size of worms and the position of occurrence of genitalia in

the strobila were found to be too variable for use as spécifie characters.

Although in some specimens of M. perplexa, the testes are separated into two latéral groups,

insufflcient emphasis has been placed on the extent of variation in testis distribution. In the types,

as noted by Stiles (1896) and Arnold (1938), the testes in immature proglottides form a continuous

band across the medulla. They only separate into two groups in mature proglottides. Furthermore,

considérable variation may occur within a single strobila. In some of Honess’ specimens of M. per¬

plexa (USNM no. 59073) proglottides with a single band of testes or with two group3 alternate irregu-

larly in the mature région of the strobila. The degree of séparation of testes into two groups may

vary from the presence of a broad distinct gap to the situation in which the testes in the centre of the

proglottis are arranged in a single row with the space of one testis only between the two groups. In

Honess’ specimens from Lepus townsendi from Wyoming determined by him as M. pectinata (USNM

no. 59076), as well as in specimens from other hosts and localities (USNM nos. 59259, 30098) in which

the testes of most proglottides are arranged in a single band, occasional proglottides occur with two

groups of testes. A similar situation can occasionally be found in European specimens (BM no. 1693.

7.81). Consequently, as Douthitt (1915) pointed out, the testis distribution is far from invariable

and cannot be used successfully to separate M. pectinata from M. perplexa. Similar variability occurs

in other anoplocephlids, particularly in M. ctenoides (vide infra), the genus Moniezia (see Theiler,

1924) and in several species of Progamotaenia Nybelin, 1917 (see Beveridge, 1976).

The séparation of M. perplexa and M. pectinata on the basis of the size of the cirrus sac appears

justifiable from Stiles’ measurements of 0.29-0.32 mm and 1.0 mm respectively. Douthitt (1915)

remeasured the cirrus sac in the types of M. perplexa and found that Stiles was mistaken, the size

of the cirrus sac ranging from 0.475-0.640 mm. A différence in cirrus sac size was also found by

Arnold (1938) who gave 0.43-0.64 mm and 1.0-1.76 mm for the cirrus sacs of the two species respec¬

tively. Honess (1963) on the other hand gave measurements of 0.27-0.88 mm and 0.73-1.08 mm

respectively, showing overlap between the two groups. If the various sets of measurements are

compared with the data obtained from M. pectinata in the présent study, it is évident that the range

0.43-1.35 mm clearly encompasses the two supposed species, so that the cirrus sac size cannot be

used as a distinguishing feature. In the types of both M. perplexa and C. mosaica, the cirrus sac

Source : MNHN, Paris

REVISION OF THE CJTTOTAENIA COMPLEX

reaches the latéral nerve but does not cross the longitudinal osmoregulatory canals (Stiles, 1896, Hall

1908) whilst in M. pectinata the cirrus sac crosses the canals (Stiles, 1896). The différences in cirrus

sac size and position are certainly quite marked in extreme cases, however Honess (1963) found that

whilst the cirrus sac almost invariably crossed the longitudinal osmoregulatory canals in M. pectinata,

in specimens designated by him as M. perplexa, the cirrus sac terminated médial to the canals in 25 %

of cases, over the canals in 50 % of cases and latéral to the canals in 25 % of cases. Furthermore,

in some specimens examined by the writer, the cirrus sac reached the osmoregulatory canals in mature

proglottides, but in gravid proglottides of the same strobilae, did not reach half way to the osmoregu¬

latory canals. The position of the cirrus sac in relation to the osmoregulatory canals therefore shows

considérable variability and does not represent a valid distinguishing characteristic. Furthermore,

there is no corrélation between the occurrence of testes in two groups in a strobila and the size of the

cirrus sac. In some of Honess’ specimens of M. pectinata, the typically long cirrus sac was présent

in ail proglottides but 50 % of the mature proglottides had the testes in two groups.

Each of the morphological features formerly used to separate M. pectinata from M. perplexa

can therefore be shown to vary in such a fashion that neither single nor multiple characters can be

used successfully to separate the two. Whilst individual specimens at extremes of the range of variation

may be identifiable with one type or the other, ail spécifie parameters overlap and hence M. perplexa

must fall as a synonym of M. pectinata.

Different patterns of variability seem to exist in different parasite populations. In the Wyoming

population studied by Honess (1963) a proportion of the cestode population had the morphological

characteristics of M. perplexa. In other populations, (e.g. from Lepus americanus in Alaska, Canada

and the U.S.A.) the occurrence of two groups of testes is extremely rare and the cirrus sac is invariably

long, extenting beyond the osmoregulatory canals. The causes of this type of variation are unknown

but similar observations hâve been made on populations of Andrya macrocephala by Rausch and

Schiller (1949) and in Progamotaenia macropodis by Beveridge (1976). Voge (1952) retained as valid

species Hymenolepis diminuta and H. citelli solely on the basis of the frequency of occurrence of varia¬

tions in testis number and position, though she was subsequently able (1969) to support this distinction

with biological and fine-structural différences. A comparable study may lead to the re-establishment

of M. perplexa as a valid taxon, however, the material currently available is insufficient and in the

absence of reliable differentiating criteria, M. perplexa must fall as a synonym.

It has been possible to establish several cases of mistaken identifications which hâve appeared

in the literature. The cestode reported by Joyeux and Gaud (1945) as Cittotaenia pectinata has been

re-examined and found to be C. denticulata. Joyeux and Baer (1936) also recorded C. pectinata from

Oryctolagus cuniculus in the Carmargue, France, however, the specimens in Joyeux’s collection with

this collection data are C. denticulata.

Edelenyi (1965) reported C. denticulata from Lepus europaeus in Hungary, however, it is clear

from the description and figures that the parasite in question is M. pectinata.

Diaz-Ungria and Aleman (1955) described M. pectinata from Sylvilagus floridanus in Venezuela,

however, from the photographs they provide, it appears that the cirrus sac is very small and does

not reach halfway to the longitudinal osmoregulatory canals, that the utérus crosses the osmoregu¬

latory canals and that the testes do not extend laterally as far as the canals, ail of which are features

of M. variabilis comb. nov. The photographs also show paired uteri in proglottides, a feature most

frequently observed in M. variabilis and the host is the usual host of the latter species. Attempts

to obtain and examine the material were unsuccessful and for this reason the record is retained provi-

sionally as one of M. pectinata.

The présent report of M. pectinata in Spermophilus variegatus represents a new host record.

The parasite is only rarely found in hosts other than leporids, though Rankin (1946) recorded it from

Sciurus carolinensis, a rodent from the same family as S. variegatus and Gubanov (1964) has reported

it from another sciurid Citellus undulatus. In the présent case, several specimens were found in the

one host by the collector, Dr. A. W. Grundmann, together with M. variabilis. Some of the worms

were gravid. Although indicating that M. pectinata is capable of developing in rodents, the few

records available suggest that this is a rare occurrence.

Source : MNHN, Paris

IAN BEVERIDGE

Table 5. — HOST RECORDS

OF MOSGOVOYIA

PECTINATA

Host Locality

Original

Détermination

Reference

Oryctolagus cuniculus Linnaeus, 1785

Great Britain

C. pectinata

Mead-Briggs and Page (1975)

Surrey, England

C. pectinata

Mead-Briggs and Yaughan

England, Wales

C. pectinata

(1973)

Baylis (1939)

Aberystwyth, Wales

C. pectinata

Rees (1933a), Evans (1940)

Scotland

C. pectinata

Cameron and Parnell (1933)

I. of Eigg, Scotland

C. pectinata

Fahmy (1960)

Mackintosh (1955)

I. de St. Pierre,

Switzerland

M. pectinata

Hôrning (1974)

Granada, Spain

C. pectinata

Lôpez-Neyra (1947)

USSR

M. pectinata

Gvozdev et al. (1970)

Richelieu, France

? C. pectinata

Dollfus (1961)

France

C. pectinata

Joyeux and Baer (1936)

Czechoslovakia (Zoo)

M. pectinata

Jaros et al. (1966)

Lepiu

s europaeus Pallas, 1778

Great Britain *

C. pectinata

Mead-Briggs and Page (1975)

England

C. pectinata

Irvin (1970)

Sweden

C. pectinata

Burgaz (1970)

Lorraine, France

C. pectinata

Joyeux and Baer (1936)

Travers, Switzerland

C. pectinata

Bouvier (1965)

Poland

M. pectinata

Drygas and Piotrowski (1955)

Poznam, Poland

M. pectinata

Czapliiiksi et al (1965)

Bialystock, Poland

M. pectinata

Wieczorowski (1968)

Hungary

C. denticulata

Edelényi (1965)

Bulgaria

M. pectinata

Yanchev (1963a, b, 1970,1973)

Azerbaïdjan USSR

M. pectinata

Sadikhov (1958)

Ukraine, USSR

M. pectinata

Sharpilo (1966)

Georgia, USSR

M. pectinata

Rodonaya (1967)

Gruzia, USSR

M. pectinata

Rodonaya (1966)

Northern

Caucasus, USSR

C. pectinata

Naumov (1944)

USSR

M. pectinata

Gvozdev et al. (1970)

Azerbaïdjan, USSR

M. pectinata

Sadikhov (1962)

Byelorussia, USSR

M. pectinata

Merkusheva (1960)

Leput

s europaeus crawshayi de Winton, 1899

(syn. Lepus crawshayi de Winton, 1899)

Zaïre

C. wittei

Baer and Fain (1955)

Lepu

s europaeus saxatilis Cuvier, 1823

(syn. Lepus saxatilis Cuvier, 1823)

Mozambique

M. pectinata

Cruz E. Silva (1971)

* Host originally given as L. capensis occidenlalis, a change in host nomenclature having taken place since 1951.

Source : MNHN, Paris

REVISION

THE CITTOTAENIA COMPLEX

Host

Détermination

Reference

Lepus capensis Linnaeus, 1758

Afghanistan

M. pectinata

Tenora and Kullmann (1970)

Lepu.

s capensis granatensis Rosenhauer, 1856

(syn. Lepus granatensis Rosenhauer, 1856)

Granada, Spain

C. pectinata

Lôpez-Neyra (1947)

Lepus capensis tibetanus Waterhouse, 1841

(syn. Lepus tibetanus Waterhouse, 1841)

Kazakstan, USSR

C. pectinata

Gvozdev (1948)

Lepu.

s capensis tolai Pallas, 1778.

(syn. Lepus tolai Pallas, 1778)

Bartoga, USSR

M. pectinata

Gvozdev (1965)

Kirgizia, USSR

M. pectinata

Tokobaev (1960)

USSR

M. pectinata

Tokobaev and Erkulov (1966)

Gvozdev et al. (1970)

Kazakhstan, USSR

M. pectinata

Gvozdev (1964)

Tadzhikistan, USSR

M. pectinata

Gafurov et al. (1971)

Lepus limidus Linnaeus, 1758

(syn. Lepus variabilis Pallas, 1778)

Briançon, France

C. pectinata

Stiles (1896), Joyeux and Baer

Sweden

C. pectinata

(1936)

Burgaz (1970)

Scotland

C. pectinata

Irvin (1970)

Germany

C. pectinata

Stiles (1896)

Verkhoyansk, USSR

M. pectinata

Gubanov and Federov (1956)

Yakutia, USSR

M. pectinata

Gubanov et al. (1957)

USSR

C. pectinata

Naumov (1940)

Vaud, Switzerland

Ct. pectinata

Galli-Valerio (1909, 1930a)

Tobolk, USSR

C. pectinata

Romanovitch (1915)

Yakutia, USSR

M. pectinata

Gubanov (1964)

Kirov, USSR

C. pectinata

Naumov (1944)

Buriat, USSR

M. pectinata

Oshmarin (1965)

Moscow, USSR

M. pectinata

Maklakova (1973)

Azerbaidjan, USSR

M. pectinata

Sadikhov (1962)

Byelorussia, USSR

M. pectinata

Merkusheva (1960)

Lepus brachyurus Temmink, 1845

Kyushu, Japan

M. oitana

Sawada and Kugi (1974)

USSR

M. pectinata

Gvozdev et al. (1970)

Tohoku, Japan

M. pectinata

Inaba and Yagisawa (1973)

Lepus

i timidus scotius Hilzheimer

Great Britian

C. pectinata

Mead-Briggs and Page (1975)

Lepus

i nigricollis Cuvier, 1823

Sri Lanka

C. bursaria

von Linstow (1906)

India

C. pectinata

Southwell (1930)

Source : MNHN, Paris

IAN BEVERIDGE

urigmai

Locahty Détermination

Lepus nigricollis ruficaudatus Geoffroy, 1826

(syn. Lepus ruficaudatus Geoffroy, 1826)

India

C. pectinata

Southwell (1930)

India

C. pectinata

Katiyar and Pande (1965)

Lepus americanus Erxleben, 1777

Manitoba, Canada

C. pectinata

Boughton (1932)

Alaska, USA

C. p. americana

Philip (1937, 1948)

Wyoming, USA

M. pectinata

Honess (1963)

Minnesota, USA

C. pectinata

Erikson (1944)

Lepus americanus struthopus Bangs, 1898

Newfoundland, Canada

M. pectinata

Dodds and Mackiewicz (1961)

Lepus californicus Gray, 1837

New Mexico, USA

M. pectinata

Samson (1968)

California, USA

C. pectinata

Lechleitner (1959)

Lepus californicus melanotis Mearns, 1890

Kansas, Nebraska,

USA

C. pectinata

Lyman (1902), Douthitt

(1915), Arnold (1938)

Colorado, USA

C. pectinata

Brittain and Voth (1975)

Lepus californicus deserticola Mearns, 1898

Utah, USA

C. p. americana

Grundmann (1958)

Lepus townsendi campanius Hollister, 1915

North Dakota, USA

C. ? pectinata

Voth and James (1965)

Wyoming, USA

M. p. americana

Honess and Winter (1956)

* Caprolagus hispidus (Pearson, 1839)

(syn. Lepus hispidus Pearson, 1839)

India

C. pectinata

Southwell (1930)

Sylvilagus floridanus Allen, 1890

North Dakota, USA

M. p. americana

Novlesky and Dyer (1970)

Eastern USA

C. perplexa

Bell and Chalgren (1943)

Oklahoma, USA

C. perplexa

Douthitt (1915)

Maryland, USA

C. perplexa

Stiles (1896)

Sylvilagus floridanus mallurus Thomas, 1898

North Carolina, USA

C. pectinata

Harkema (1936)

Sylvilagus floridanus mearnsii Allen, 1894

Minnesota, USA

C. pectinata ,

C. perplexa

Erikson (1947)

Michigan, USA

C. pectinata

Haugen (1942)

* Host identification uncertain.

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Host

Locality

Original

Détermination

R eference

Sylvïlagus floridanus margarita Miller, 1898

Venezuela C. pectinata

(détermination doubtful)

Diaz-Ungria and Aleman

(1955)

Sylvilagus floridanus alacer Bangs, 1896

USA

M. p. americana,

M. perplexa

Honess and Winter (1956)

Sylvilagus nuttalli grangeri Allen, 1895

Wyoming, USA

C. pectinata,

C. perplexa

M. p. americana,

Honess (1935)

M. perplexa

Honess and Winter (1956)

Sylvilagus nuttalli pinetis Allen, 1894

Colorado, USA

C. mosaica

Hall (1908)

Wyoming, USA

M. perplexa

Honess and Winter (1956)

Sciurus carolinensis pennsylvanicus Ord, 1815

(syn. Sciurus leucotis Gapper, 1830)

Massachusetts, USA

C. p. americana

Rankin (1946)

Citellus undulatus Pallas, 1778

Yakutia, USSR

M. pectinata

Gubanov (1964)

Mosgovoyia ctenoides (Railliet, 1890) comb. nov.

Figs 26-34. Tables 6-7.

Dipylidium leuckarti Riehm, 1881a : 200, 566-577, pl. 5, figs 3, 11-13, 16, pl. 6, figs 5-6.

Taenia leuckarti (Riehm, 1881) Neumann, 1888 : 426.

Moniezia leuckarti (Riehm, 1881) Blanchard, 1891 : 187, 444, 450-451.

Ctenotaenia leuckarti (Riehm, 1881) Railliet, 1893, : 278.

Cittotaenia leuckarti (Riehm, 1881) Stiles and Hassall, 1896b : 407.

Taenia ctenoides Railliet, 1890 : 346.

Cittotaenia ctenoides (Railliet, 1890) Stiles 1896 : 179-181, pl. 14.

Ctenotaenia ctenoides (Railliet, 1890) Spasskii, 1951 : 268-269. fig. 125.

Neoctenotaenia ctenoides (Railliet, 1890) Tenora, 1976 : 13.

Types. From Oryctolagus cuniculus Linnaeus, 1758, Germany, coll. Riehm, cotype in USNM

1327.

Material examined.

From Oryctolagus cuniculus Linnaeus, 1758. 2 specimens, Milford, Surrey, England, 8 August,

1974, A. Mead-Briggs, UMVS ; 10 slides, Paris, France, 5 February 1921, 1930, C. Joyeux, UN ; 1 spe-

Source : MNHN, Paris

IAN BEVERIDGE

cimen, St. Quentin, France, 17 December 1929, C. Joyeux, UN ; 3 specimens, Algeria, 1 November

1926, C. Joyeux, UN ; 2 specimens, Ile de St. Pierre, Switzerland, 17 March 1976, B. Hôrning, UN ;

3 specimens, Graz, Austria, June 1976, E. Ebermann, in colin, of E. Ebermann.

Description. Large, broad, ribbon-like worms. Scolex small, rounded anteriorly, merging

imperceptibly into neck région or may be demarcated from neck by slight constriction posterior to

suckers. Proglottides craspedote, extended transversely. Mature proglottides with approximate

length : width ratio of 1 : 4-1 : 6. Gravid proglottides with similar ratio. Longitudinal musculature

weakly developed, arranged in two rows of muscle bundles. Bundles of inner row smaller with up

to 25 fibres per bundle. Bundles of outer row larger, elongate with more fibres. Transverse muscle

fibres fine, arranged in band internai to longitudinal muscles. Dorso-ventral muscles weakly developed,

composed of individual, irregularly arranged fibres. Longitudinal osmoregulatory canals paired.

Ventral canal straight, thin-walled, wider than dorsal canal. Dorsal canal narrow, surrounded by

layer of muscle cells. Transverse canal connects left and right ventral canals at posterior margin of

each proglottis. Dorsal canals from each side join one another at level of suckers, similarly ventral

canals fuse. Short common dorsoventral duct joins the fused canals from each side. Accessory

canals connecting adjacent transverse canals in strobila absent. Génital atrium of insignificant size,

situated in posterior half of latéral proglottis margin, dividing margin in ratio 1 : 2-1 : 3. Cirrus

sac very short, ellipsoidal, never reaching more than halfway from latéral proglottis margin to longi¬

tudinal osmoregulatory canals. Cirrus short, coiled, unarmed. Internai séminal vesicle présent

occupying up to two-thirds of volume of cirrus sac. Vas deferens thin-walled, coiled, follows course

of vagina and séminal réceptacle to vicinity of Mehlis’ gland and divides into numerous vasa efferentia.

Testes numerous, in 2-4 transverse and 1-5 horizontal rows entirely posterior to utérus. Testes densest

just to aporal side of ovaries, some overlying lobules of ovary dorsally. Testes extend posterior to

vitellarium, with 3-8 testes poral to females genitalia. Testes usually in two latéral groups with

médian space, number of testes towards centre of proglottis low. Occasionally testes form continuous

band across medulla. Both types of arrangement of testes may occur within one strobila. Testes

do not persist in gravid proglottides. Vagina narrow, surrounded externally by layers of glandular

cells. Vagina terminâtes near longitudinal osmoregulatory canals in narrow duct without surrounding

glandular cells which leads into elongate, thin-walled séminal réceptacle, also without glandular cells.

Séminal réceptacle terminâtes in narrow duct in vicinity of Mehlis’ gland. Ovary fan-shaped, composed

of numerous clavate lobules, situated on ventral aspect of medulla. Vitellarium reniform, lobulate,

posterior and dorsal to ovary. Mehlis’ gland spherical, anterior and dorsal to vitellarium. Utérus

tube-like, transverse, usually single, occasionally paired, extending across middle of medulla anterior

to Mehlis’ gland. Poral to Mehlis’ gland, utérus bends posteriorly, crosses longitudinal osmoregul¬

atory canals dorsally and terminâtes posterior to vagina and proximal extremity of cirrus sac. Gravid

utérus with numerous anterior and posterior diverticula. Egg spherical, shell thick, refractile, subshell

and inner membranes présent. Pyriform apparatus drawn into two elongate horns.

Remarks. The above description of M. ctenoides is similar in most respects to that of earlier

publications. It agréés with Dollfus’ description (1951) in that the testis distribution is somewhat

variable and that the testes may be distributed in a single band or in two groups posterior to the utérus.

Dollfus’ suggestion that the size and morphology of the cirrus sac are more reliable taxonomie characters

than testis distribution is also confirmed, though it is considered safest to use the fact that the cirrus

sac never reaches more than half way to the longitudinal osmoregulatory canals rather than the absolute

size of the cirrus sac which might vary between worms. M. ctenoides is readily distinguished from

M. pectinata, the species most closely related to it by the form of the cirrus sac and by the fact that

in M. ctenoides the utérus invariably extends beyond the osmoregulatory canals even in the early

stages of its development. This species is distinguishable from M. variabilis by the shape of the cirrus

sac and by the occurrence of testes poral to the female genitalia in M. ctenoides but not in M. variabilis.

Riehm (1881a) described a branched osmoregulatory System in the posterior proglottides of

this species, and his figure was subsequently reproduced by Stiles (1896) and Spasskii (1951). This

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX 25

type of System is not found in strobilae which are apolytic and is probably an abnormality which may

occur in the first formed proglottides of a strobila.

The descriptions of the egg and pyriform apparatus are similar to those of Obitz (1934).

Several authors hâve recorded this species from north American leporids (see Table 7). Unfor-

tunately their specimens could not be located and the occurrence of this species has not been confirmed

in the material to hand from North America.

Although the spécifie epithet commonly used since the révision of Stiles (1896) has been ctenoides,

recently, Dollfus (1951) and Yamaguti (1959) hâve revived the use of the earlier epithet leuckarti,

both without any explanation of its use. The species was first described under the name Dipylidium

leuckarti by Riehm (1881a), and as Stiles (1896) has shown, the original description is quite adéquate

and cannot be confused with other species. Railliet (1890) described the same parasite under the

name Taenia ctenoides, but subsequently (1893) recognised the synonymy of the two and treated

T. ctenoides as a synonym placing D. leuckarti in his new genus Ctenotaenia. The spécifie name was

also recognised as leuckarti by Stiles and Hassall (1896b) in their paper on the priority of the genus

Cittotaenia over Ctenotaenia in which they established the new combination Cittotaenia leuckarti. Stiles

(1896) however introduced the combination Cittotaenia ctenoides without any explanation and this

name has persisted in common use. The probable reason for this change was Stiles’ discovery that

in 1888, Neumann had proposed the name Taenia leuckarti, apparently unaware that the name was

preoccupied by Taenia leuckarti Krahbe, 1869. The name under these circumstances was probably

presumed lost by Stiles and the name first applied to the parasite after Neumann’s error, that of

Taenia ctenoides was taken as the correct name. However, if one applies current rules of zoological

nomenclature (Art. 59 and ammendments of 1972) then Blanchard’s removal of the species under

the combination Moniezia leuckarti ensures the survival of the spécifie epithet leuckarti. According

to current amendments, since Neumann’s transfer of the species and the resulting homonomy were

overlooked prior to Stiles’ (1896) révision, Blanchard’s (1891) transfer of the species to the genus Monie¬

zia means that leuckarti is the correct current epithet. However, at the time of Stiles’ work, the code

of nomenclature did not exist as such, and since Stiles’ action is quite logical and probably follows

usage of that period, the question of the correctness of the spécifie epithet would appear to hinge

upon whether one considers the rules of nomenclature can be ma de rétrospective to decisions taken

before their existence.

Furthermore, as mentioned earlier, the name ctenoides has been in common use since 1896 and

except for a paper by Smith in 1908, and those of Dollfus (1951) and Yamaguti (1959), the name

leuckarti has not been used. Although leuckarti cannot be considered a nomen oblitum because not

50 years has elapsed since its use, it is virtually such in practice. In the présent review, it has been

decided to allow common use to prevail and to continue to use the epithet ctenoides. A formai sub¬

mission will be made to the International Commission of Zoological Nomenclature on the correct name.

Table 6. — MEASUREMENTS OF MOSGOVOYIA CTENOIDES (in mm)

Stiles Arnold Lôpez-Neyra Dollfus Présent

(1896) (1938) (1947) (1951) description

Host O. cuniculus O. cuniculus O. cuniculus O. cuniculus O. cuniculus

Length 800

Width 10

Scolex diam.

Sucker diam. 0.176

Neck

No. proglottides

Mature

proglottides

460 200-300

10.5 7-7.5

0.32-.048 0.45-.046

0.12-0.25 0.11-0.164

0.25-0.42

560 300

430-550 173-243

7-10 10-14

0.30-0.52

0.14-0.162 0.12-0.22

0.38-0.65

410-510

3.5-5.5 X

0.75-1.0

Source : MNHN, Paris

IAN BEVERIDGE

Stiles

Arnold

Lôpez-Neyra

Dollfus

Présent

(1896)

(1938)

(1947)

(1951)

description

Host

0. cuniculus

O. cuniculus

O. cuniéulus

O. cuniculus

Gravid

4.5-8.3 X

proglottides

1.5-1.7

Cirrus sac

0.16 x 0.128

0.15-0.25

0.29 X 0.13

0.12-0.7 X

0.16-0.25 X

0.065-0.128

0.08-0.12

Internai

0.1-0.16 x

séminal vesicle

0.06-0.08

No. testes

120-160

70-150

50-70

100-160

Testis size

0.05-0.06

0.046-0.081

0.05-0.07

0.04-0.07

Ovary

0.40-0.88

0.45-0.55 X

0.35-0.50

Vitellarium

0.15-0.3 X

0.08-0.18

Mehlis’gland

Dorsal

0.12

osmoregul-

atory canal

Ventral

0.015

osmoregul-

atory canal

0.07

Egg

Pyriform

0.064

0.062-0.069

0.07-0.085

0.056

0.06-0.07

apparatus

Oncosphere

Table 7. — HOST RECORDS OF MOSGOVOYIA

CTENOIDES

Host Locality n Original

J Détermination

Reference

Oryclolagus cuniculus Linnaeus, 1785

Great Britain

Surrey, England

Wales

Netherlands

France

Paris, Toulouse,

St. Quentin,

France

Rhiems, France

Richelieu, Cada¬

rache, France

Germany

Hamburg,

Germany

Czechoslovakia

C. ctenoides

C. leuckarti

C. ctenoides

Ct. ctenoides

Mead-Briggs and Page (1975)

Mead-Briggs and Vaughan

(1973)

Stephens (1952)

van der Broek and Jansen

(1964)

Stiles (1896)

Joyeux and Baer (1936)

Courtehoux (1948)

Dollfus (1951, 1961)

Stiles (1896)

Arnold (1938)

Paéenovsky (1973)

Source : MNHN, Paris

REVISION OF THE C1TTOTAEN1A COMPLEX

Host Locality

Original

Détermination

Reference

Granada, Spain

C. ctenoides

Lôpez-Neyra (1947)

Huesca, Spain

C. ctenoides

Tarazona Vilas (1955)

Portugal

C. ctenoides

da Silva Leitâo (1964)

Morocco

C. leuckarti

Dollfus (1951)

Algena

C. ctenoides

Joyeux (1927)

Azores

C. ctenoides

Stiles (1896)

? Lepus europaeus Pallas, 1778

Czechoslovakia

Ct. ctenoides

Novak et al., (1966)

Lepus californiens Gray, 1837

Oklahoma, USA

C. ctenoides

Ward (1934)

Sylvilagus floridanus Allen, 1890

North Dakota,

USA

C. ctenoides

Novlesky and Dyer (1970)

Oklahoma, USA

Philadelphia,

C. ctenoides

Ward (1934)

USA

Ct. leuckarti

Smith (1908)

Sylvilagus floridanus alacer Bangs, 1896

Oklahoma, USA

C. ctenoides

Smith (1940)

Sylvilagus aquaticus Bachman, 1837

Oklahoma, USA

C. ctenoides

Ward (1934)

Sylvilagus aquaticus aquaticus Bachman, 1837

Oklahoma, USA

C. ctenoides

Smith (1940)

Mosgovoyia variabilis (Stiles, 1895) comb. nov.

Figs 35-40, Tables 8, 9.

Ctenotaenia variabilis Stiles, 1895 : 345.

Cittotaenia variabilis (Stiles, 1895) Stiles and Hassall, 1896b : 407.

Cittotaenia variabilis angusta Stiles, 1896 : 193, pl. 19, figs 13-14.

Cittotaenia variabilis imbricata Stiles, 1896 : 193.

Neoctenotaenia variabilis (Stiles, 1895) Tenora, 1976 : 13.

Cittotaenia pectinata (Goeze, 1782) pro parte Baer, 1927 : 55.

Types. From Sylvilagus floridanus (Allen, 1890). (syn. Lepus sylvestris Allen, 1890), Bowie,

Maryland, USA, A. Hassall, USNM, BM 95.9.11.1. For details see Stiles (1896).

Material examined.

From Sylvilagus floridanus (Allen, 1890). 1 specimen, New York, USA, C. Bursey, UN ;

3 specimens, Lincoln, Nebraska, USA, 4 October 1897, A. S. Pearse, from collection of H. B. Ward,

UN ; 1 specimen, cotype, Bowie, Maryland, USA, A. Hassall, BM 95.9.11.1 ; 1 specimen, Colorado,

USA, G. D. Schmidt, in colin, of G. D. Schmidt.

Source : MNHN, Paris

IAN BEVERIDGE

From Syloilagus floridqnus mearnsi (Allen, 1894). 1 specimen, East Lansing, Michigan, USA,

29 November, 1945, R. L. Rausch, in colin. R. L. Rausch.

From Syloilagus nuttalli grangeri (Allen, 1895). 1 specimen, Curacanti Res., Gunnison River,

Colorado, USA, 2 July 1961, A. W. Grundmann, UU 49 ; Wyoming, USA, no other data, USNM 59078 ;

Montana, USA, no other data, USNM 30954.

From Syloilagus palustris Bachman, 1837. Georgia, USA, no other data, USNM 30954.

From Syloilagus sp. Virginia, USA, USNM 39887 ; Louisiana, USA, USNM 28513.

From Oryctolagus cuniculus Linnaeus, 1758 dom. 1 specimen, Ohio, 1943-1944, R. Rausch,

in coll. of R. Rausch ; 1 specimen, Coshocton, Ohio, 28 November 1944, R. Rausch, in coll. of R. Rausch.

From Spermophilus oariegatus (Erxleben, 1777). 2 specimens, Red Canyon, 30 miles east of

Salina, Utah, USA, 15 May 1958, A. W. Grundmann, UU 157-1.

Description. Large broad ribbon-like worms. Scolex small, rounded anteriorly merging

imperceptibly into neck région, or in some specimens subglobular, distinctly demarcated from strobila

with neck absent. Proglottides craspedote, greatly extended transversely. Mature proglottides

with approximate length : width ratio of 1 : 6. Gravid proglottides with ratio of 1: 8. Longitudinal

musculature weakly developed, arranged in two rows of bundles around inner edge of cortex. Trans¬

verse muscles fine, arranged in band internai to longitudinal muscles. Dorso-ventral muscles fine,

few in number, irregularly arranged. Longitudinal osmoregulatory canals paired. Dorsal canal

narrow, surrounded by layer of muscle cells, médial to ventral canal. Ventral canal wider, thin-

walled. Transverse canal connects left and right osmoregulatory canals at posterior margin of each

proglottis. Accessory canals connecting transverse canals absent. Dorsal canals from each side of

strobila join in midline at level of suckers. Common duct passes anteriorly fusing with common ventral

duct by short dorso-ventral anastomosis. Génital atrium of insignificant size, situated in anterior

half of latéral proglottis margin, dividing margin in ratio 5 : 4. Cirrus sac small, elongate, thin-walled,

reaching only half way to longitudinal osmoregulatory vessels. Cirrus straight, unarmed. Small,

elongate internai séminal vesicle présent. Vas deferens narrow, thin-walled, coiled, fohowing course

of vagina and séminal réceptacle, decreasing in diameter. Vasa efferentia not seen. Testes numerous

almost invariably situated in single transverse band, always posterior to utérus, on dorsal aspect of

medulla, between female genitalia. Rarely testes in single proglottis almost separated into two groups.

Laterally, testes extend no further than aporal margin of vitellarium. Testes in 2-4 horizontal

and 1-2 transverse rows. Testes do not persist in gravid proglottides. Vagina narrow, surrounded

by layers of glandular cells. Two thirds of distance to osmoregulatory canals, vagina leads into

narrow thin-walled duct without surrounding glandular cells. Duct extends medially to elongate, thin-

walled séminal réceptacle dorsal to ovary. Ovary fan-shaped, composed of numerous clavate lobules,

situated on ventral aspect of medulla. Vitellarium reniform, lobulate, posterior and dorsal to ovary.

Mehlis’ gland spherical, anterior to vitellarium, dorsal to ovary. Utérus paired or single, transverse

extending across middle of proglottis anterior to Mehlis’gland, dorsal to ovary. Utérus passes poste

riorly, crosses longitudinal osmoregulatory canals dorsally, terminating posterior to vagina and cirrus

sac. Paired uteri may meet in midline or there may be additional small cavities between uteri, lined

with squamous epithelium rather than with cuboidal epithelium of uteri. Gravid utérus sac-like,

without prominent diverticula. Egg spherical, thick-shelled. Thick sub-shell membrane and very

fine inner membrane présent. Oncosphere surrounded by pyriform apparatus, latter forming two

elongate horns.

Remarks. Some confusion exists in the literature between this species and M. pectinata

probably due to Baer’s (1927) earlier attempt to synonymise the two. The two species differ markedly

in the extension of the utérus of M. oariabilis across the osmoregulatory canals in the tubular stage,

the utérus of M. pectinata being at ail times restricted to the cortex, and in the distribution of testes

which lie between the female genitalia in M. oariabilis but extend latéral to the female genitalia in

M. pectinata. M. oariabilis possesses in common with M. ctenoides a muscular coat to the dorsal osmore¬

gulatory canal which is absent in M. pectinata. M. oariabilis can be distinguished from M. ctenoides

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

by the shape of the cirrus sac and by the distribution of the testes. Although the testes in M. oaria-

bilis are nearly always distributed in a single band, occasional proglottides can be found in which

two groups are présent indicating similar patterns in variability of testis distribution to M. pectinata

and M. ctenoides.

Lyman (1902) described a sphincter surrounding the génital atrium in this species but his

observations hâve not since been confirmed and no such sphincter was seen in the material currently

available.

This species is virtually restricted to host species belonging to the genus Sylvilagus, apart from

the single record included here from the rodent Spermophilus variegatus, and from the domestic rabbit,

Oryctolagus cuniculus.

Table 8 . — MEASUREMENTS OF MOSGOVOYIA VARIABILIS (in mm)

Stiles

T.vman

Arnold

Honess

Présent

(1896)

(1902)

(1938)

(1963)

description

Sylvilagus

floridanus

Syl. floridanus

Sylvilagus

[L. sylvaticus

L. palustris )

( L. sylvaticus )

floridanus

nultalli

Length

100-180

170-180

450

111-324

104-940

Width

10.5

3-8

8-14

Scolex diam.

0.32-0.56

0.462-0.872

0.44-0.61

0.37-078

0.48-0.75

Sucker diam.

0.11-0.28 X

0 . 2 - 0.282 x

0.16-0.28

0.20-0.26

0.11-0.24

0.106

Neck

0.26-0.84

0-0.4

No. proglottides

750

313-519

340-660

Mature

4.0-6.9 x

proglottides

0.55-1.0

Gravid

5.5-7.5 X

proglottides

0.55-1.0

Cirrus s.c

0.4 x 0.05

0.32-0.45

0.25-0.70 X

0.04-0.05

No. testes

65-90

75-100

60-135

80-120

Testis size

0.08-0.10

0.053-0.071

0.07-0.08

Ovury

0.69 X 0.1

0.48-0.71

0.4-0.6 X

0.2-0.5

Vitellarium

0.266 X 0.088

0.15-0.22 x

0.13-0.18

Mehlis’gland

Dorsal

osmoregulatory

canal

Ventral

osmoregulatory

canal

0.09-0.13

0.005-0.03

0.1-0.25

0.06-0.064

0.058

0.052-0.068

0.045

Pyriform

apparatus

0.012-0.016

0.016

0.030

Oncosphere

0.015

Source : MNHN, Paris

IAN BEVER1DGE

Table 9. — HOST RECORDS OF MOSGOVOYIA VARIABILIS

Host Locality

Original

détermination

Reference

Sylvilagu.

s floridanus (Allen, 1890)

North Dakota, USA

North Carolina, USA

Eastern USA

Maryland, Long Is., USA

Nebraska Kansas, USA

Alabama, USA

Ct. variabilis

C. variabilis

Novlesky and Dyer (1970)

Stringer et al. (1969)

Bell and Chalgren (1943)

Stiles (1896)

Lyman (1902)

Price and Ingram (1959)

Sylvilagu;

i floridanus alacer (Bangs, 1896)

Kansas, USA

C. variabilis

Arnold (1938)

Sylvilagu;

t floridanus mallurus (Thomas, 1898)

Alabama, USA

Connecticut, USA

New York,

Pennsylvania, USA

Alabama, USA

C. variabilis

Moore and Moore (1947)

Clancy et al. (1940) Hosley

(1942)

Arnold (1938)

Moore (1939)

Sylvilagus floridanus mearnsii (Allen, 1894)

Minnesota, USA

Iowa, USA

C. variabilis

Erikson (1947)

Morgan and Waller (1940)

Sylvilagu;

! transitionalis (Bangs, 1895)

Massachusets, USA

Connecticut, USA

C. variabilis

Rankin (1946)

Clancy et al. (1940) Hosley

(1942)

Sylvilagu;

s auduboni (Baird, 1858)

Colorado, USA

C. variabilis

McCrease (1957)

Sylvilagu;

t auduboni vallicola Nelson, 1907

California, USA

C. ? variabilis

Herman and Jankiewicz (1943)

Sylvilagu;

! nuttalli grangeri (Allen, 1895)

Wyoming, USA

C. variabilis

Ci. variabilis

Honess (1935)

Honess and Winter (1956)

Sylvilagu.

s nuttalli pinetis (Allen, 1894)

Colorado, USA

C. variabilis

Stock (1962)

Sylvilagus palustris Bachman, 1837

North Carolina, USA

Florida USA

? Maryland, USA

C. variabilis

Stringer et al. (1969)

Stiles (1896)

Arnold (1938)

Lepus americanus (Erxleben, 1777)

Minnestoa, USA

C. variabilis

Erikson (1944)

Source : MNHN, Paris

REVISION OF THE CITTOTABNIA COMPLEX

Genus PSEUDOCITTOTAENIA Tenora, 1976 emend.

Pseudocittotaenia Tenora, 1976 : 14-15.

Types Species. Pseudocittotaenia praecoquis (Stiles, 1895).

Diagnosis. Small cestodes. Scolex rounded, unarmed. Suckers unarmed. Proglottides

numerous (over 50 in gravid strobilae), craspedote, extended transversely. Longitudinal osmoregul-

atory canals paired. Transverse canal connects left and right ventral canals at posterior margin of

each proglottis. Génital ducts cross longitudinal osmoregulatory canals dorsally. Genitalia paired.

Cirrus sac opens to génital atrium anterior to vagina. Cirrus sac dorsal to vagina on both sides of

strobila. Internai séminal vesicle présent. External séminal vesicle présent or absent. Testes

numerous, entirely posterior to utérus, either restricted to area between female genitalia or extending

laterally to female genitalia. Séminal réceptacle large, prominent. Ovaries situated in latéral quarters

of proglottis medulla. Single transverse, tubular utérus in each proglottis at anterior extremity of

proglottis, extending beyond longitudinal osmoregulatory canals ventrally, terminating anterior to

cirrus sac and vagina. Gravid utérus sac-like, with anterior and posterior diverticula. Pyriform

apparatus présent. Parasites of Geomyidae (Rodentia).

Remarks. The genus Pseudocittotaenia has been retained for two species, P. praecoquis (Stiles,

1895) comb. nov. and P. glandularis sp. nov., both parasites of geomyid rodents in North America.

The genus is distinguished from ail other related généra by the position of the utérus which

is close to the anterior extremity of the proglottis, crosses the longitudinal canals ventrally rather

than dorsally and in the tubular stage terminâtes anterior rather than posterior to the cirrus sac and

vagina. This combination of characters does not occur in any of the related cestodes with paired

genitalia, but it does occur in certain members of the genus Paranoplocephala Lühe, 1810 parasitising

North American rodents, in particular P. troeschi Rausch, 1946. It is therefore reasonable to suggest

that species of Pseudocittotaenia hâve arisen by duplication of the genitalia of North American species

of Paranoplocephala and that their évolution has been parallel to the duplication of genitalia in European

forms of Paranoplocephala which hâve given rise to the genus Ctenotaenia (see Spasskii, 1951, Tenora,

1976).

Pseudocittotaenia praecoquis (Stiles, 1895) comb. nov.

Figs 41-47, Table 10.

Ctenotaenia praecoquis Stiles, 1895 : 345 (as praecoquus).

Cittotaenia praecoquis (Stiles, 1895) Stiles and Hassall, 1896b : 407.

Neoctenotaenia praecoquis (Stiles, 1895) Tenora, 1976 : 13.

Cittotaenia megasacca Smith, 1951 : 313, figs 1-5.

Pseudocittotaenia megasacca (Smith, 1951) Tenora, 1976 : 15.

Types. From Geomys bursarius (Shaw, 1800), Ames, Iowa, USA, Osborne 1894, 2 specimens

in USNM 11372.

Material Examined.

From Geomys bursarius (Shaw, 1800). Holotype and paratype.

Source : MNHN, Paris

IAN BEVEREDGE

From Thomomys talpoides parowanensis Goldman, 1938. 8 specimens, Griffith Creek, Tushar

Mtns., Beaver County, Utah, USA, 12 August 1960, A. W. Grundmann, UU 14-20 ; 4 specimens, same

locality, 10 August 1960, A. W. Grundmann, UU 14-21.

From Thomomys talpoides clusius Coves, 1875 ; 10 specimens, Savoy, Carbon County, Wyoming,

USA, 1948, C. F. Smith ; paratypes of C. megasacca in USNM and UNMC.

From Thomomys talpoides tenellus Goldman, 1939. 7 specimens, Moran, Wyoming, USA,

25, 26 June 1948, R. L. Rausch in colin. R. L. Rausch and UN.

Description. Short, broad worms with less than 100 proglottides in gravid strobilae. Scolex

rounded anteriorly. Suckers oval, retracted within scolex. Neck absent. Proglottides craspedote,

extended transversely. Mature proglottides with approximate length : width ratio of 1 : 12. Gravid

proglottides with approximate ratio of 1 : 11. Terminal proglottides distinctly narrower than sub¬

terminal proglottides, with approximate ratio of 1 : 3. Muscular System not examined. Longitudinal

osmoregulatory canals paired. Ventral canal wider than dorsal canal, situated médial to it. Transverse

canal connects left and right ventral canals at posterior margin of each proglottis. Osmoregulatory

canals of scolex not seen. Génital atrium deep, prominent, situated in middle of latéral proglot¬

tis margin in mature proglottides, near posterior extremity of latéral proglottis margin in gravid

proglottides. Cirrus sac elongate, pyriform, thick-walled, extending just beyond ventral osmore¬

gulatory canal. Cirrus elongate, coiled, uniformly covered with rows of short spines, surrounded

by glandular cells. Ovoid internai séminal réceptacle occupies one quarter to one third volume

of cirrus sac. Ellipsoidal external séminal vesicle présent, overlying proximal pôle of cirrus sac.

Vas deferens narrow, passes anteriorly and medially from séminal vesicle. Testes ovoid or sphe-

rical, numerous, arranged in 2-3 dorsoventral and 1-5 horizontal rows on dorsal aspect of medulla,

in single band, entirely posterior to utérus. Few or no testes latéral to female genitalia. Considérable

variation exists in number of testes latéral to female genitalia within individual strobilae. Testes

occasionally extend as far laterally as longitudinal osmoregulatory canals and rarely lie latéral to

canals. Vagina narrow, tube-like opens to génital atrium posterior to cirrus sac. Body of vagina

lies ventral to cirrus sac on both sides of strobila. Vagina merges into enormous, pyriform séminal

réceptacle. Séminal réceptacle persists in gravid proglottides. Ovary fan-shaped, composed of

numerous clavate lobules, situated on ventral aspect of medulla. Vitellarium ovoid, posterior and

dorsal to ovary. Mehlis’ gland not seen. Single utérus per proglottis, developing as elongate, trans¬

verse tube close to anterior margin of proglottis. Utérus anterior to ovaries, séminal réceptacles,

crosses longitudinal osmoregulatory canals ventrally, terminating anterior to cirrus sac. Gravid

utérus sac-like with numerous anterior and posterior diverticula. Anterior diverticula larger than

posterior diverticula. Utérus eventually fills proglottis. Egg approximately spherical, thick-shelled.

Oncosphere surrounded by pyriform apparatus which terminâtes in two elongate horns.

Remarks. The species was first described by Stiles (1895, 1896) under the names Ctenotaenia

praecoquis and CiUotaenia praecoquis respectively, though the two specimens available to him were

insufficient to allow a full description to be made. Subsequently, Smith (1951) redescribed C. praeco¬

quis from Thomomys talpoides together with a new species which he named Cittotaenia megasacca.

Unfortunately, he apparently did not examine the types of C. praecoquis, and it is évident from the

re-examination of the types of both C. praecoquis and C. megasacca, made by the writer, that they

are identical. Although the types of C. praecoquis are not well preserved, and the cirrus sac is smaller

than in other specimens examined, the scolex, position of utérus and testis distribution are exactly

as those seen also in the types of C. megasacca. Smith’s (1951) specimens attributed to C. praecoquis

are a distinct species, and are described below under the new name Pseudocittotaenia glandularis.

Consequently, C. megasacca must fall as a synonym of C. praecoquis. However, the situation is

complicated by the fact thatrecently (Tenora, 1976), these two hâve been placedin separate généra in two

sub-families. C. praecoquis was transferred to a new genus Neoctenotaenia by Tenora (1976) having

Neo. ctenoides as the type species. In the présent révision, the genus N eoctenotaenia is placed as a

synonym of Mosgoooyia. The genus Pseudocittotaenia was created by Tenora (1976) within the sub-

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

family Monieziinae Spasskii, 1951 for the species P. megasacca (Smith, 1951), P. bequaerti (Vigueras,

1943) and provisionally P. rhea (Fuhrmann, 1904), under the assumption that the utérus of each of

these species was reticulate during the early part of its development. Smith’s (1951) original descrip¬

tion of C. megasacca indicated quite clearly that the utérus was initially tubular and that it subse-

quently developed anterior and posterior diverticula, an observation confirmed in the présent re¬

description. Because of this, the genus must be transferred to the Anoplocephalinae. P. bequaerti

and P. rhea are removed to the genus Moniezia Blanchard, 1891 (see below). Although C. megasacca

falls as a synonym of C. praecoquis, the genus Pseudocittotaenia can be retained for P. praecoquis.

The above description difîers from the original and that of Smith (1951) in a number of respects.

However, since Stiles’ material (1896) was poor, comparison with the original description will not be

made in detail. Smith (1951) stated that the cirrus was unarmed, however, at least one of the para-

types of C. megasacca has several cirri everted and on them the armature is readily visible. It is not

possible to see the cirrus armature on inverted cirri of the paratypes, however, it is readily seen on

inverted cirri in other specimens examined. The description of the external séminal vesicle given

by Smith (1951) implies that it is merely a passively produced dilation of the vas deferens when filled

with sperm. Re-examination of his material indicates that the external séminal vesicle is invariably

présent and is easily distinguished before the male reproductive System has commenced functioning.

Although Smith (1951) in his drawings of the species shows testes anterior to the utérus, this

situation was not found in re-examination. Occasional testes were found which overlapped the

posterior margin of the utérus but none were found entirely anterior to it.

The occurrence of testes latéral to the female genitalia is highly variable and there may be no

testes whatever latéral to the female genitalia (fig. 45). Although the testes extended to the longitu¬

dinal osmoregulatory canals quite frequently, only rarely did they lie latéral to the canals.

P. praecoquis is readily distinguished from the only other member of the genus P. glandularis

by the lack of glandular cells around the distal extremity of the vagina, by the much longer, narrower

cirrus sac, by the presence of an external séminal vesicle and by the occurrence of testes latéral to the

female genitalia.

Stiles’ (1896) and Smith’s (1951) measurements of this species are included in Table 10. Smith’s

measurements were checked and since they appear to be accurate are not repeated. Only additional

measurements made are tabulated.

Table 10. — MEASUREMENTS OF PSEUDOCITTOTAENIA PRAECOQUIS (in mm)

From

StUes (1896)

Geomys

bursarius

Smith (1951)

types of

C. megasacca

paratypes

of C.

megasacca

Présent description

From Thomomys talpoides

Utah and Th. tenellus

Wyoming

Length

14-22

14-51

Width

5.5

No. proglottides

150

68-89

71-130

Scolex diam.

0.43 X 0.32

0. 65-0.82

0.50-0.65

0.16-0.128

0.22 x 0.19

0 . 16 - 0.22 x

0.17-0.20

1.6-2.2 X

1.3-2.8 X

proglottides

0.15-0.20

0.2-0.4

Gravid

2.4-4.0 X

proglottides

0.37-0.80

0.4-0.7

100-135

Testis diam.

0.025-0.067

0.02-0.08

Source : MNHN, Paris

IAN BEVERIDGE

Fnm

Stiles (1896)

Geomys

bursarius

Smith (1951)

types of

C. megasacca

paratypes

of C.

megasacca

Présent description

From Thomomys talpoides

Utah and Th. tenellus

Wyoming.

Cirrus sac

0.24 x 0.096

0.52 X 0.15

0.3-0.4 X

0.08-0.013

Internai semical

0.07-1.9 x

0.16-0.27 x

vesicle

0.06-1.1

0.05-0.12

External séminal

0.2 x 0.07

0 .12-0.20 x

vesicle

0.05-0.1

Ovary

0.37-0.43 x

0.33-0.47 x

0 .11-0.12

0.14-0.16

Vitellarium

0.06 x 0.04

0.04-0.05 x

0.08-0.10

Séminal

0.72-0.19

0.80 X 0.25

0.35-0.65 x

réceptacle

0.14-0.17

Dorsal osmoregul-

atory canal

0.019

0.02

Ventral osmoregul-

atory canal

0.32

0.07-0.1

Egg

0.032-0.036

0.038

0.04

Oncosphere

0.01

0.015

Pyriform apparatus

0.025

0.035

Pseudocittotaenia glanduiaris sp. nov.

Figs 48-54, Table 11.

CiUotaenia praecoquis (Stiles, 1895) Stiles and Hassall, 1896b sensu Smith, 1951 : 312-313.

Types. From Thomomys talpoides clusius Coves, 1875. Savoy, Carbon County, Wyoming,

USA, 1948, C. F. Smith, holotype in USNM 4379.

Additional records. From Thomomys talpoides wasaichensis Durrant, 1946, Utah, USA

(det. as C. praecoquis) (Frandsen and Grundmann, 1961).

Material examined.

From Thomomys talpoides clusius Coves, 1875. 1 specimen, Holotype, USNM.

From Thomomys talpoides wasatchensis Durrant, 1846. 4 slides of fragments, Monte Cristo

Forest Camp, Cache County, Utah, USA, 10 July 1959, J. C. Frandsen and A. W. Grundmann, UU

41-3 ; 2 specimens, same locality, 11 July 1959, same collectors, UU 41-3.

Description. Scolex large, rounded, mildly four lobed. Proglottides craspedote, extended

transversely. Mature proglottides with approximate length : width ratio of 1 : 7-1 : 12. Gravid pro¬

glottides, longer, relatively narrower, with ratio of 1 : 1.5-1 : 4. Muscular System not examined.

Longitudinal osmoregulatory canals paired. Ventral canal wider than dorsal canal, situated médial

to it. Transverse canal connects left and right ventral canals at posterior margin of each proglottis.

Scolex osmoregulatory canals not seen. Génital atrium deep, narrow, situated in mid-region of latéral

Source : MNHN, Paris

REVISION OF THE C1TTOTAEN1A COMPLEX

proglottis margin. Cirrus sac ovoid or ellipsoidal, thick-walled, invariably extending just to aporal

side of ventral osmoregulatory canal. Cirrus short, uncoiled, armature not seen. Cirrus surrounded

by masses of intensely staining glandular cells. Ovoid internai séminal vesicle présent, occupying

up to one third to one half volume of cirrus sac. External séminal vesicle absent. Vas deferens

narrow, slightly coiled, extending medially and anteriorly from cirrus sac. Testes ovoid or spherical,

numerous, arranged in 2-3 dorso-ventral and 3-5 horizontal rows on dorsal aspect of medulla. Testes

entirely posterior to utérus, invariably in single band, between female genitalia. Few testes lie dorsal

to aporal part of ovary but none seen latéral to ovary. Vagina short, tube-like, opening to génital

atrium posterior to cirrus sac. Distal extremity of vagina surrounded by mass of intensely-staining

glandular cells. Vagina merges into enormous, pyriform séminal réceptacle. Séminal réceptacle

persiste in gravid proglottides. Ovary fan-shaped, composed of numerous clavate lobules situated

on ventral aspect of medulla. Vitellarium roughly ovoid, lobulate, posterior and dorsal to ovary

Mehlis’ gland not seen. Utérus tubular, single, transverse, close to anterior proglottis margin. Utérus

anterior to ovaries, séminal réceptacles and testes, Crossing longitudinal osmoregulatory canals ventrally,

terminating close to dorsal canal, anterior to proximal pôle of cirrus sac. Gravid utérus sac-like with

anterior and posterior diverticula which may subdivide. Diverticula reduced or absent in région

where utérus crosses osmoregulatory canals. Egg spherical thick shelled. Oncosphere surrounded

by pyriform apparatus. Pyriform apparatus terminâtes in two elongate horns.

Remarks. This species was first described, but not illustrated, under the name Cittotaenia

praecoquis by Smith (1951) and subsequently recorded from Utah by Frandsen and Grundmann (1961).

Through the kindness of Dr. Grundmann, the latter specimens hâve been examined, and their co-

identity with Smith’s (1951) specimens established.

The species is here placed in the genus Pseudocittotaenia since it shares in common with the

type species the situation of the utérus at the anterior extremity of the proglottis, passing the longi¬

tudinal canals ventrally and terminating at a level anterior to the cirrus sac. It is distinguished from

P. praecoquis by the limitation of the testes to the area between the female genitalia and by masses

of glandular cells surrounding the cirrus and the distal région of the vagina. The spécifie name is

given on account of this latter characteristic.

Table 11. MEASUREMENTS OF PSEUDOCITTOTAENIA GLANDULARIS (in mm)

Smith

(1951)

Thomomys talpoides

Wyoming

Smith’s material

Holotype

Thomomys talpoides

Wyoming

Holotype

Présent description

Thomomys talpoides

Utah

Length

75-150

Width

2.0

Scolex diameter

0.74-0.85

0.65-0.66

Sucker diameter

0.26-0.29

0.25-0.30 x

Neck

No. proglottides

176-262

0.20-0.30

Mature proglottis

1.0-1.6 X

Gravid proglottis

0.05-0.18

1 . 6 - 2.0 x

0.2-0.4

1.2-2.2 x

0.24 x 0.09

0.8-0.85

0.35-1.0

0.15-0.24 x

Internai séminal vesicle

0.07 x 0.04

0.06-0.09

0.08-0.14 x

0.03-0.09

Source : MNHN, Paris

LN BEVERIDGE

(1951)

Tliomomys talpoides

Wyoming

Smith’s material

Thomomys talpoides

Wyoming

Présent description

Thomomys talpoides

Utah

No. testes

45-60

110

Testis size

0.039-0.054

0.04-0.06

Séminal réceptacle

0.34-0.42 x

0.11-0.14

0.11-0.30 X

0.10-0.17

Ovary

0.32 X 0.17

0.27-0.40 X

0.20-0.25

Vitellarium

0.12 X 0.04

0.10-0.15 X

0.09-0.1

Dorsal osmoregulatory canal

0.015

0.01

Ventral osmoregulatory canal

0.05

0.04-0.12

Egg

Pyriform apparatus

0.035

0.04

Oncosphere

0.009

0.007

2. Subfamily MONIEZIINAE Spasskii, 1951

Genus CITTOTAENIA Riehm, 1881

Cittotaenia Riehm, 1881a : 200 ; Stiles and Hassall, 1896b : 407 ; Stiles, 1896 : 170 ; Douthitt, 1915 :

46-50 ; Baer, 1927 : 49-59 ; Fuhrmann, 1932 : 61 ; Lôpez-Neyra, 1947 : 227 ; Spasskii, 1951 : 413-420 ;

Wardle and McLeod, 1952 : 365 ; L<5pez-Neyra, 1954 : 109-116 ; Yamaguti, 1959 : 200, 375-376 ; Tenora,

1976 : 14.

Types species. Cittotaenia denticulata (Rudolphi, 1804).

Diagnosis. Cestodes of moderate size. Strobila ribbon-like. Scolexsmall, unarmed. Suckers

unarmed. Proglottides numerous (more than 100 in gravid strobilae), craspedote, extended transver-

sely. Longitudinal osmoregulatory canals paired, with or without accessory longitudinal vessels and

numerous anastomosing supplementary vessels connected to them. Transverse canal connects left

and right ventral canals at posterior margin of each proglottis. Genitalia paired. Génital ducts

pass longitudinal osmoregulatory canals dorsally. Cirrus sac dorsal to vagina on both sides of strobila.

Internai and external séminal vesicles absent. Testes numerous, in single band or two groups. Séminal

réceptacle présent. Ovaries situated in latéral quarters of proglottis medulla. Single transverse,

tubular utérus, very slightly reticulated, not extending laterally beyond longitudinal osmoregulatory

canals. Utérus develops anterior and posterior diverticula, finally becoming sac-like. Pyriform

apparatus présent. Parasites of Leporidae (Lagomorpha) and Chinchillidae (Rodentia).

Remarks. The définition of the genus given above is essentially similar to that of Spasskii

(1951). It is left provisionally within the sub-family Monieziinae, where it was placed by Spasskii,

because the utérus is slightly reticulated. However, the extent of the réticulation is very slight in

C. denticulata and usually consist of only 2 to 3 small loops in an otherwise simple tubular utérus,

Source : MNHN, Paris

REVISION OF THE C1TTOTAENIA COMPLEX

differing considerably from the complex net-like utérus of the more typical members of the sub-family

such as species of Moniezia and Andrya. In typical members of the subfamily Monieziinae, the net-

like utérus gradually fills with eggs and loses its reticulated character whereas in C. denticulata the

utérus develops anterior and posterior diverticula, a characteristic of the Anoplocephalinae. The

situation is further complicated by the second species of the genus, C. viscaciae comb. nov. the mor-

phology of which is still poorly known but which seems to hâve a slightly more reticulated utérus than

C. denticulata. It is therefore proposed to leave the genus Cittotaenia within the subfamily Monieziinae

pending further clarification of its taxonomie position.

The genus is differentiated from anoplocephaline généra parasitising rodents and lagomorphs

by the slightly reticulated utérus, and within the Monieziinae, from Diandrya Darrah, 1930 by the

slight réticulation of the utérus and the absence of a distinct prostate, and from the genus Moniezia

again by the utérus, the lack of interproglottidal glands, the strong development of the cirrus sac and

the relationships of vagina and cirrus sac on either side of the strobila.

Cittotaenia denticulata (Rudolphi, 1804)

Figs 55-65, Tables 12-13.

Taenia denticulata Rudolphi, 1804 : 81.

Alyselminthus denticulatus (Rudolphi, 1804) de Blainville, 1828 : 607.

Moniezia denticulata (Rudolphi, 1804) Blanchard, 1891 : 187.

Ctenotaenia denticulata (Rudolphi, 1804) Stiles and Hassall, 1896a : 6-9.

Cittotaenia denticulata (Rudolphi, 1804) Stiles and Hassall, 1896b : 407.

Taenia goezi Baird, 1853 : 78.

Moniezia goezi (Baird, 1853) Blanchard, 1891 : 444, 452-457, figs 21-25.

Ctenotaenia goezi (Baird, 1853) Railliet, 1893 : 278.

Cittotaenia latissima Riehm, 1881a : 200.

Dipylidium latissimum (Riehm, 1881) Riehm, 1881b : 583-590, pl. 5, figs 5, 15, 17, pl. 6, fig. 2.

Taenia latissima (Riehm, 1881) Neumann, 1888 : 426.

Types. In Berlin Muséum (not examined).

Material examined.

From Oryctolagus cuniculus Linnaeus, 1758. 1 specimen, no locality or date, BM 1957.5.1.9-10 ;

8 specimens, Milford, Surrey, England, 8 August 1974, A. Mead-Briggs, UMVS ; 2 specimens, Camargue,

France, January 1935, C. Joyeux, UN ; 1 specimen, Si Allai Tazi, Morocco, 15 April 1942, C. Joyeux,

UN ; 1 specimen, Paris, France, 1930, C. Joyeux, UN.

Description. Large, broad, ribbon-like worms. Scolex small, quadrate, distinctly separated

from strobila, four lobed with sucker on each lobe. Neck absent. Proglottides extended transversely,

craspedote with narrow, slightly undulating vélum overhanging adjacent proglottis. Mature proglot¬

tides with approximate length : width ratio of 1 : 8-1 : 10. Gravid proglottides with ratio of 1 : 3-

1 : 8. Longitudinal musculature moderately developed, arranged in two irregular rows of oval or

elongate bundles in cortex. Bundles of two rows of approximately equal size. Transverse muscles

filiform, in band internai to longitudinal muscles. Dorso-ventral muscles fine, Crossing medulla and

cortex at irregular intervals. Longitudinal osmoregulatory canals variable in number. Dorsal canal

narrow, thick-walled, surrounded externally by layer of muscle cells. Usually 2-4 major ventral

canals situated both latéral and médial to dorsal canal. 1-2 additional smaller canals may be présent.

Transverse canal connects ventral canals at posterior margin of each proglottis. Irregular anastomoses

connect various central canals as well as anastomosing network of canals connecting transverse canals

Source : MNHN, Paris

IAN BEVERIDGE

of adjacent proglottides. In scolex, canals consist of paired dorsal and paired ventral canals only.

Posterior to suckers, canals of each side describe prominent latéral loop, return to near midline and

extend anteriorly between suckers. Transverse anastomosis connects ventral canals from each side.

Similar anastomosis connects two dorsal canals. Four canals extend anteriorly from level of two

transverse anastomoses. Two short dorso-ventral loops connect dorsal and ventral canal on each

side of scolex. Génital atrium deep, narrow, situated in posterior half of latéral proglottis margin

dividing margin in ratio of 2 : 1. Génital papilla présent in fully mature proglottides as prominent

dome-shaped projection close to postero-lateral corner of proglottis. Cirrus sac large, oblong, thick-

walled, surrounded by layer of polygonal parenchymatous cells. Cirrus short uncoiled. Distal région

of wider diameter, heavily armed with rows of spines. Proximal région unarmed, slightly dilated

when filled with sperm. Internai and external séminal vesicles absent. Vas deferens narrow, thin-

walled, coiled, surrounded by layer of glandular cells. Vasa efferentia not seen. Testes numerous

in 2-3 transverse and 6-8 horizontal layers on dorsal aspect of medulla. Testes usually lie both anterior

and posterior to utérus, but in some immature proglottides may lie entirely posterior to utérus. Testes

lie between female genitalia, overlapping ovary and vitellarium dorsally, but never extending laterally

beyond them. Vagina narrow, uncoiled, opens to génital atrium posterior to cirrus sac. Vagina

lies ventral to cirrus sac on both sides of strobila. Séminal réceptacle oval or circular in dorso-ventral

view, dorsal to ovary. Ovary circular with central réservoir and numerous elongate clavate lobules

radiating anteriorly, posteriorly laterally, medially and dorsally. Vitellarium reniform to U-shaped,

lobulate, posterior and dorsal to centre of ovary. Mehlis’ gland spherical, lying between arms of

vitellarium, slightly dorsal to them. Utérus single in each proglottis, transverse, tube-like with small

number of réticulations and vestigial anterior and posterior diverticula. Developing utérus slightly

reticulate with 1-4 loops in each utérus, or rarely non-reticulate in the form of a simple transverse tube.

Numerous anterior and posterior diverticula présent. Utérus does not cross longitudinal osmoregul-

atory canals. Gravid utérus sac-like, filling proglottis medulla. Réticulations not visible. Egg

spherical, thick-shelled. Oncosphere surrounded by pyriform apparatus drawn out into two elongate

horns. Horns rarely subdivided giving appearance of three-horned pyriform apparatus.

Remarks. This species has been well described in the past (John, 1926) and is sufïiciently

characteristic morphologically that confusion with other species is not likely to occur. The présent

description differs little from that of John (1926). The number of ventral osmoregulatory canals

was found to be more variable and a network of accessory canals connecting transverse canals of

adjacent proglottides, similar to that in Mosgovoyia pectinata was found. John (1926) presumably

saw these accessory canals but simply stated that “ secondary longitudinal canals occur ”. He also

stated that whilst secondary canals were présent in mature proglottides, only two ventral canals were

présent in younger proglottides. In the material studied here, accessory canals were seen within

2 mm of the scolex and even proglottides with only génital anlagen in them were found with a complex

network of accessory canals.

Riehm (1881a) described the pyriform apparatus as terminating in elongate filaments. Although

not found in the présent study, it may be that the material available was not fully developed. One

embryo was found with a three-horned pyriform apparatus.

The relationships of uterine development to the systematic position of the parasite hâve been

discussed earlier.

Edelényi (1965) identified C. denticulata from Lepus europaeus in Hungary, but from the figure

provided, the parasite in question appears to be M. pectinata. In a subséquent paper (Edelényi,

1966), C. denticulata is again reported from L. europaeus in Hungary, but as there is no description

or diagram from which to check the identification, the record has been included in Table 13, but with

réservation.

Joyeux and Gaud (1945) reported M. pectinata from Oryctolagus cuniculus in Morocco, but

re-examination of their material has revealed that it is in fact C. denticulata.

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Table 12. — MEASUREMENTS OF CITTOTAENIA DENTICULATA (in mm)

Stiles

John

Baer

Arnold

Présent

(1896

(1926)

(1927)

(1938)

description

Length

400-500

30-70

400-500

260

Width

11-13

8.5

Scolex diam.

0.8 X 0.63

0.75-1.18 X

0.43-0.52

0 .8-1.0

0.10-0.76

0.75-0.90

Sucker diam.

0.23-0.29

0.2-0.3

0.23-0.30

0.24-0.26 X

0.21-0.24

Neck

0.21-0.92

No. proglottides

200

300

260

—

Mature proglottides

5.7 X 0.6-0.7

Gravid proglottides

4-11 x 1.7-3.0

Cirrus sac

1.12 x 0.32

0.75 x 0.26

1.12 x 0.3

0.50-0.97

0.6-0.95 X

0.23-0.25

No. testes

100

225-250

200

Testis size

0.115

0.129

0.100

0.041-0.12

0.04-0.12

Séminal

0.2-0.4 X

0.36-0.45 X

réceptacle

0.15-0.2

0.31-0.35

Ovary

1.4

0.32-1.42

0.1-1.0 X

0.4-0.8

Vitellarium

0.5

0.28-0.75 X

0.15-0.28

Mehlis’ gland

Dorsal osmore-

0.12

gulatory canal

Ventral osmore-

0.02-0.06

gulatory canals

0.06-0.18

0.052-0.060

0.054

0.052-0.060

0.046-0.075

0.065

Pyriform apparatus

0.038

Oncosphere

0.008

0.015

Table 13.

— HOST RECORDS OF CITTOTAENIA

DENTICULATA

Host

Locality

Reference

Oryctolagus cuniculus Linnaeus, 1785

Great Britain

Surrey, England

Northumberland, England

England, Wales

Aberystwyth, Wales

Wales

Scotland

Granada, Madrid, Spain

France

France, Germany

Mead-Briggs and Page (1975)

Mead-Briggs and Vaughan (1973)

Boag (1972)

Baylis (1939)

John (1926), Evans (1940)

Stephens (1952)

Fahmy (1960)

Lôpez-Neyra (1947, 1954)

Du Buysson (1904)

Stiles (1896)

Source : MNHN, Paris

IAN BEVERIDGE

Host

Locality

Reference

Hamburg, Germany

Arnold (1938)

Alsace, France

Galli-Valerio (1930a)

Richelieu, France

Dollfus (1961)

Ile de St. Pierre, Switzerland

Galli-Valerio (1910, 1930a)

Morocco (det. as M. pectinata)

Joyeux and Gaud (1945)

Lepus europaeus Pallas, 1778

Scotland

Cameron and Parnell (1933)

? Hungary

Edelényi (1966)

Lepus timidus Linnaeus, 1758

Scotland

Cameron and Parnell (1933)

Sweden

Burgaz (1970)

Cittotaenia viscaciae (Spasskii, 1951) comb. nov.

Figs 66-72, Table 14.

Mosgovoyia viscaciae Spasskii, 1951 : 296-297.

Cittotaenia pectinata (Goeze, 1782) pro parte Joyeux and Dollfus, 1931 : 155.

Types. Single specimen from ? Lagostomus maximus (Desmarest, 1817) (syn. Viscacia viscacia

(Molina, 1782)), Valdivia, Chile. Whole mount and serial sections in UN.

Material examined. Type.

Description. Moderate sized worm. Scolex small, rounded anteriorly. Suckers cup-shaped,

opening circular. Scolex merges imperceptibly into unsegmented neck région. Proglottides extended

transversely, craspedote with broad, straight-edged vélum overhanging adjacent proglottis. Mature

proglottides with approximate length : width ratio of 1 : 5. Gravid proglottides with ratio of 1 : 4.5.

Longitudinal musculature weakly developed, arranged in two rows of small, oval bundles containing

3-15 fibres per bundle, in ring around inner edge of cortex. Transverse muscles fine, in band internai

to longitudinal muscles. Dorso-ventral muscles fine, few in number, irregularly arranged. Longi¬

tudinal osmoregulatory canals paired. Ventral canal wider than dorsal. Transverse canal connects

left and right ventral canals at posterior margin of each proglottis. Small accessory longitudinal

canals associated with transverse canal. Broad génital papilla présent in many proglottides, in posterior

half of latéral proglottis margin. Génital atrium shallow. Cirrus sac large, oblong. Walls of cirrus

sac thick, muscular in mid and proximal régions, diminishing in thickness distally. Cirrus uncoiled,

armed with numerous, awl-like, spirally-arranged spines. Mid région of cirrus unarmed. Spines

were absent from everted cirri, presumably due to tardy fixation of specimen. Proximal région of

cirrus slightly dilated with sperm. Internai and external séminal vesicles absent. Vas deferens

thin-walled, slightly coiled, passing medially in arc, dorsal to ovary, adjacent to séminal réceptacle.

Vasa efferentia not seen. Testes numerous, situated in two entirely separate groups in posterior

part of proglottis, on dorsal aspect of medulla in 2-5 horizontal and 1-4 transverse rows. Testes extend

latéral to female genitalia, overlying vitellarium dorsally. Vagina opens to génital atrium posterior

to cirrus sac via narrow invaginated extension of atrium. Vagina narrow, thin-walled, posterior

and ventral to cirrus sac. Séminal réceptacle small, clavate, dorsal to centre of ovary. Ovary circular

in dorso-ventral view, situated on ventral aspect of medulla, with central réservoir and numerous

Source : MNHN, Paris

EVISION OF THE CITTOTAENIA COMPLEX

clavate lobules radiating from it. Vitellarium U-shaped, lobulate, dorsal to ovary. Anterior parts

of vitellarium lie either side of séminal réceptacle. Mehlis’ gland not seen. Testes, vitellaria and

séminal réceptacles persist after involution of ovary. Utérus slightly reticulated in early stage of

development, lying on ventral aspect of medulla. Ring of utérus surrounds each séminal réceptacle

with major branch anterior and one posterior to testes. Gravid utérus sac-like, non-reticulated,

filling proglottis. Egg spherical, thick-shelled. Oncosphere surrounded by pyriform apparatus

drawn into two elongate horns.

Remarks. This specimen was briefly described by Joyeux and Dollfus (1931) amongst a

collection of cestodes sent to them from the Münich Muséum. Its morphological features difîered

somewhat from those of M. pectinaia, particularly with respect to the occurrence of testes in two

groups, a feature which these authors noted was characteristic of M. ctenoides rather than M. pecti-

nata. However they cited Baer’s révision of the Anoplocephalidae as evidence that M. pectinata

was a highly variable species and placed their specimen under this name. Spasskii (1951) subsequently

erected a new species, Mosgovoyia viscaciae for the specimen in spite of the fact that he had only

Joyeux and Dollfus’ (1931) quite inadéquate description from which to judge its taxonomie position.

Although only the one type specimen was available for description, and the State of the specimen

is now such that a full description cannot be made, the features of the worm described above indicate

quite clearly that it does not belong to the genus Mosgovoyia but that it probably belongs to the genus

Cittotaenia. The cestode is similar to C. denticulata in many respects, particularly in possessing a

slightly reticulated utérus, a large, oblong, cirrus sac and heavily-armed cirrus as well as in the structure

of the ovary which in both species is circular with lobules radiating from a central réservoir, rather

than being fan-shaped as is the case with ail other species dealt with herein. C. viscaciae can be

distinguished from C. denticulata by the occurrence of testes in two groups in the posterior part of

the proglottis and by the lack or at least the small number of accessory osmoregulatory canals.

The host of this species cannot be stated with certainty. Joyeux and Dollfus (1931) gave

as the host “ Viscacia viscacia (Molina) syn. Lagostomus trichodactylus Brookes ” apparently deriving

the host nomenclature from Trouessart (1899). The current name of L. trichodactylus is Lagostomus

maximus (Desmarest, 1817) (see Cabrera 1961) however, it does not occur in Chile which was the

collection locality given by Joyeux and Dollfus (1931). Both Osgood (1943) (p. 137) and Cabrera

(1961), however, give Lepus viscacia Molina, 1782 as a synonym of Lagidium viscacia (Molina, 1782),

the common mountain viscacia of Chile, which occurs, according to Osgood (1943) in the provinces

of Aconcagua, Santiago and Valparaiso, which are not too distant from Valdivia. It is also possible

that the initial identification of the host was merely the vernacular name “ viscacia ” which was

taken to he the “ Argentinian or plans viscacia ”, L. maximus rather than a “ mountain viscacia ”

belonging to the genus Lagidium Meyen, which occur in Peru and Chile. Since there are several

species in the genus Lagidium, it is not possible to suggest which one is the most likely host. The

true host of C. viscaciae is of some importance since another anoplocephalid, Cittotaenia quadrata

von Linstow, 1904 has been described from Lagidium peruanum Meyen, 1833, in Peru. The descrip¬

tions and extant material of C. quadrata are so poor as to make the détermination of its taxonomie

position impossible, however, in the slides that do exist it appears to hâve a cirrus sac similar to C. vis¬

caciae. It is therefore not impossible that C. viscaciae might be a synonym of C. quadrata though

without new collections of cestodes from Lagidium peruanum the matter cannot he pursued further.

Source : MNHN, Paris

IAN BEVERIDGE

Table 14. — MEASUREMENTS OF CITTOTAENIA VISCACIAE (in mm)

Joyeux and Dollfus

(1931)

Présent

description

Length

106

Width

Scolex diameter

0.50

0.45

Sucker diameter

0.15-0.16

No. proglottides

157

Mature proglottis

5.5 X 1.0

Gravid proglottis

5.5 X 1.5

Cirrus sac

0.72 X 0.15

0.55-0.80 x

0.22-0.25

Internai séminal vesicle

—

No. testes

140-160

120

Size testes

0.06

Séminal réceptacle

0.22 x 0.17

Ovary

0.6

0.5 x 0.5

Vitellarium

0.225

0.30 x 0.26

Egg

0.06

0.075

Pyriform appparatus

0.020

Oncosphere

0.010

3. — Species Referred to other généra of Anoplocephalidae

Subfamily ANOPLOCEPHALINAE

Genus STRING0P0TAEN1A gen. nov.

Type species. Stringopotaenia psitlacea (Fuhrmann, 1904).

Diagnosis. Anoplocephaline cestodes with paired reproductive organs. Scolex unarmed.

Suckers unarmed. Proglottides numerous (greater than 100 in gravid strobila), craspedote, extended

transversely. Longitudinal osmoregulatory canals paired. Transverse canal connects left and right

ventral canals at posterior margin of each proglottis. Génital ducts cross longitudinal osmoregulatory

canals dorsally. Cirrus sac dorsal to vagina on right hand side of strobila, ventral to it on left hand

side. Internai séminal vesicle présent, external vesicle absent. Testes numerous, scattered throughout

medulla. Séminal réceptacle présent. Ovaries situated in latéral quarters of proglottis, utérus in

middle of proglottis with two U-shaped loops passing anteriorly over vitellaria. Gravid utérus of

similar shape with anterior and posterior diverticula. Egg with pyriform apparatus. Parasites of

Psittacidae (Aves).

Remarks. See below.

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Stringopotaenia psittacea (Fuhrmann, 1904) comb. nov.

Figs 73-78, Table 15.

Cittotaenia psiltacea Fuhrmann, 1904 : 358-386.

Paramoniezia psittacea (Fuhrmann, 1904) Spasskii, 1951 : 303-305, fig. 143.

Types. Single specimen, from Stringops habroptilus Gray, 1845 (Psittacidae : Stringopinae),

New Zealand, as 19 slides of histological sections, UN.

Material examined. Type.

Description. Scolex at anterior extremities of suckers square in transverse section. Sections

posterior to suckers cross-shaped, indicating four lobes to scolex. Proglottides craspedote with

narrow, straight-edged vélum, extended transversely. Mature proglottides with approximate length :

width ratio of 1 : 9. Gravid proglottides with ratio of 1 : 6. Longitudinal musculature strongly

developed, arranged in bundles of variable size around inner margin of cortex. Two to three layers

of bundles présent. Transverse muscles fine, forming band internai to longitudinal muscles. Dorso-

ventral muscles not seen. Longitudinal osmoregulatory canals paired. Ventral canal wider than

dorsal canal. Transverse canal connects left and right ventral canals at posterior margin of each

proglottis. Longitudinal nerve latéral to canals. Génital atrium insignificant, in posterior half of

latéral proglottis margin. Cirrus sac oblong, elongate, extending beyond osmoregulatory canals.

Cirrus coiled, thick-walled, muscular, armed with short stout bristles. Internai séminal vesicle elongate.

External séminal vesicle absent. Vas deferens prominent, greatly coiled, surrounded by pale-staining,

polygonal cells. Coils of vas deferens extend in transverse plane across entire width of medulla anterior

to cirrus sac, and in horizontal plane from latéral edge of ventral osmoregulatory canal to ovary.

Proximal coils of vas deferens smaller in diameter, without surrounding polygonal cells. Vas deferens

runs to dorsal aspect of medulla, divides into numerous vasa efîerentia. Testes numerous, in 2-3

transverse layers and 6-8 horizontal layers on dorsal aspect of medulla. Testes extend throughout

medulla and are limited laterally by osmoregulatory canals. Testes lie anterior and posterior to the

utérus extend latéral to female genitalia and overlie ovary, cirrus sac and vas deferens dorsally. Vagina

opens to génital atrium posterior to cirrus sac. Vagina lies ventral to cirrus sac on right hand side

of strobila and dorsal to it on left hand side. Distal vagina tube-like, narrow, lined internally with

hairs, vaginal wall thick, muscular, surrounded by layers of glandular cells. Proximal vagina (médial

to osmoregulatory canals) thin-walled, ending in elongate, pyriform, thin-walled sac without hairy

lining or glandular cells. Séminal réceptacle circular in dorso-ventral view, dorsal to ovary. Ovary

in latéral part of proglottis medulla fan-shaped, composed of numerous clavate lobules, situated on

ventral aspect of medulla. Vitellarium oblong or reniform, dorsal and posterior te ovary. Mehlis’

gland spherical, anterior and dorsal to vitellarium. Oocapt présent at origin of oviduct. Oviduct

joins duct from séminal réceptacle forming short fertilisation duct which joins with vitelline duct

and common duct formed passes anteriorly to Mehlis’ gland, emerges on dorsal side of gland and

continues anteriorly and ventrally to utérus. Utérus single, transverse, tube-like, situated in pos¬

terior half of proglottis close to transverse osmoregulatory canal. Utérus bends anteriorly forming

loop over vitellarium, dorsal to ovary, ventral to séminal réceptacle and Mehlis’ gland. Junctions

of transverse body of utérus with loops marked by short, posteriorly-directed diverticula. In gra¬

vid proglottides, utérus crosses longitudinal canals dorsally, extends to postero-lateral corners of

proglottis. Gravid utérus sac-like, with anterior and posterior diverticula but retaining anterior

loops over sites of vitellaria. Egg approximately spherical, thick shelled. Inner membrane pré¬

sent. Oncosphere surrounded by pyriform apparatus terminating in two elongate horns. One or

two sets of rudimentary genitalia présent near centre of each proglottis. If one set présent, situated

Source : MNHN, Paris

IAN BEVERIDGE

to one side of mid-line. Uterine duct usually présent, with ovarian lobules and/or vitellarium. Mehlis’

gland occasionally présent. Coiled portion of vas deferens présent in one proglottis.

Remarks. Although repeating in many instances the observations made by Fuhrmann (1904,

1908, 1921) on this species, the additional features described significantly alter the taxonomie position

of the parasite. Minor différences exist between the présent observations and those of Fuhrmann.

Fuhrmann (1921) gave as the testis diameter 0.08-0.16 mm compared with the présent finding of

0.065 mm, a différence for which there is no obvious explanation. He also stated that testes lie enti-

rely anterior to the utérus, yet a small number can be found overlapping the utérus dorsally with

others entirely posterior to it.

The major morphological feature upon which the taxonomie position of the parasite appears

to hinge and which Fuhrmann did not describe in detail is the structure of the utérus. The double

U-shape of the utérus in both mature and gravid proglottides indicates a much doser affinity with

the genus Paronia Diamare, 1900 than with Cittolaenia, Ctenotaenia or Paramoniezia into ail of which

généra the parasite has formerly been placed. In the genus Paronia, the mature proglottis possesses

two uteri each in the form of an inverted U over the female génital complex. In the gravid proglottis,

the uteri may fuse to give a double U-shaped structure. A similarly-shaped utérus is présent in

Stringopolaenia psittacea. The morphology of the utérus was not easy to study as only serial sections

were available. Most were oblique and showed only one of the U-shaped components, however, a

few sections were found which indicated the shape of the entire utérus and one of these has been drawn

(Fig. 74). The few sections that show the entire length of the utérus are not the most convincing

for demonstrating the U bend over the female genitalia, so a drawing is also included (Fig. 76), showing

half the utérus with a prominent hend over the remnants of the vitellarium. The same general uterine

shape is also évident in mature proglottides, the U loop over the vitellarium being demarcated on

either side by two short, posteriorly-directed diverticula. S. psittacea differs from typical représenta¬

tives of the genus Paronia in having a single utérus per proglottis instead of two uteri which may

fuse, in the relative positions of cirrus sac and vagina on either side of the strobila and in the accessory

female génital organs présent in the centre of the proglottis. Since S. psittacea cannot readily be

accommodated in the genus Paronia, a new genus has been proposed, the name being derived from

that of the host.

Spasskii (1951) used the position of the vagina dorsal to the cirrus sac on one side of the strobila

and ventral on the other side to place S. psittacea in the genus Paramoniezia Maplestone and South-

well, 1923, apparently overlooking the fact that the type species of the genus, P. suis Maplestone

and Southwell, 1923 was described as having the vagina on one side of the cirrus sac on the right

hand side of the strobila and either dorsal or ventral on the other side of the strobila. The situation

of the génital ducts in S. psittacea is in fact the same as in the genus Moniezia. In addition, Beveridge

(1976) re-examined the type of P. suis and found that Maplestone and Southwell (1923) had erred

in their description of the relationships of cirrus sac and vagina in this species and that the cirrus

sac lay on the same side of the vagina on both sides of the strobila. Although Beveridge (1976) retained

and redefined the genus Paramoniezia, the présent redescription indicates clearly that S. psittacea

does not belong within it.

Baer (1927) placed Paramoniezia suis as a synonym of S. psittacea (then Cittotaenia psittacea),

saying that he could find no différences between them. This statement must be due in part to the

extremely poor State of the type and only specimen of P. suis. So poor are the slides that an adé¬

quate redescription cannot be given, however, the two species differ markedly in the morphology

of the séminal réceptacle as well as in the vas deferens which in S. psittacea is an enormous coiled struc¬

ture surrounded by prostatic cells and in P. suis a simple coiled tube.

The Australian représentatives of the genus Paramoniezia were discussed by Beveridge (1976).

Two valid species were admitted, the type species, P. suis and a new species P. johnstoni Beveridge,

1976. The only other species in the genus, P. phacochoeri was considered a species inquirenda. The

removal of P. psittacea to a new genus leaves in the présent révision the genus Paramoniezia as a

wholly Australian one with only two species.

Source : MNHN, Paris

REVISION OF THE CITT0TAEN1A COMPLEX

Apart from morphological similarities, the généra Paronia and Stringopotaenia are both found

in Psittacidae. Ail of the other anoplocephalid généra found in Psittacidae hâve single genitalia.

Aporina Fuhrmann, 1902, Bioporuterina Burt, 1973, Hemiparonia Baer, 1927, Paronia Diainare, 1900

and Triulerina Fuhrmann, 1921 are distributed through Africa, America, Asia and Australia whilst

Pulluterina Smithers, 1954 is known only from New Zealand. When compared with Pullulerina

nestoris Smithers, 1954, the resemblance between the mature proglottis and half of that of S. psittacea

is striking. The morphology of the utérus in mature proglottides is very similar, although the gravid

utérus in P. nestoris has elongate anterior and posterior diverticula unlike S. psittacea. Several instances

are known in which two généra differ only in possessing single or paired genitalia and it is supposed

that the former arose from the latter (Baer, 1955). Pulluterina and Stringopotaenia may be related

in the same way and hence similarities between Paronia and Stringopotaenia may be due to convergence.

It is possible to interpret the extra sets of female genitalia which occur so regularly in the centre of

the proglottides of S. psittacea as a legacy of the duplication of genitalia. Similar vestigial genitalia

occur less frequently in species of Ctenotaenia and Mosgovoyia.

New Zealand has two généra of indigenous and quite unique parrots, the keas, Nestor spp.

(subfamily Nestorinae) which constitutes the only genus in the subfamily, parasitised by Pulluterina

nestoris (see Smither, 1954) and the kakapo, Stringops habroptilus (subfamily Stringopinae) again the

only genus in the subfamily, parasitised by S. psittacea. The long séparation of New Zealand from

other continents and its unique parrot fauna parallel the différences in the two endemic généra of

parrot cestodes when compared with the anoplocephalid généra of Psittacidae elsewhere in the world.

It is to be regretted that the description of S. psittacea must be based on a single specimen,

however, the host is now almost extinct (Merton, 1975) and the likelihood of obtaining additional

material is small.

Table 15. — MEASUREMENTS OF STRINGOPOTAENIA PSITTACEA (in mm)

Fuhrmann

(1904)

Fuhrmann

(1921)

Présent

description

Length

100

130

Width

6.0

6.5

—

Scolex diameter

0.23

0.57

0.55

Sucker diameter

0.23 X 0.26

0.18-0.19

Mature proglottis

3.9 X 0.4

Gravid proglottis

Cirrus sac

0.52 X 0.02

0.5 x 0.05

0.58 X 0.10

Internai séminal vesicle

—

No. testes

approx. 200

Testis size

0.08-0.16

0.065

Séminal

réceptacle

—

Ovary

0.8 X 0.28

Vitellarium

0.28

0.23 X 0.13

Mehlis’ gland

0.08

Dorsal osmoregulatory

canal

Ventral osmoregulatory

canal

Egg

0.06

Pyriform apparatus

—

Oncosphere

Source : MNHN, Paris

IAN BEVERIDGE

Subfamily MONIEZIINAE

Moniezia rhea (Fuhrmann, 1904) comb. nov.

Figs 79-90, Table 16.

Cittotaenia rhea Fuhrmann, 1904 : 386-387.

Ctenotaenia rhea (Fuhrmann, 1904), Spasskii, 1951 : 272-273, fig. 128.

Pseudocittotaenia rhea (Fuhrmann, 1904) Tenora, 1976 : 15.

Types. From Rhea americana Linnaeus, 1758, no collection data, alcohol specimens, whole

mounts and serial sections, in UN.

Material examined. Types. 4 specimens from Rhea americana, Brazil, in HCIOC and UN.

Description. Small, narrow worm. Scolex very large, quadrate, forming four bulbous

lobes with very large, oval, muscular suckers. Neck long, narrow in relaxed specimens, absent in

contracted specimens. Proglottides craspedote with narrow, straight-edged vélum overhanging

adjacent proglottis, extended transversely. Mature proglottides with approximate length : width ratio

of 1 : 5-1 : 9. Gravid proglottides with ratio of 1 : 7-1 : 9. Cortex extremely thick, approximately

5 times as wide as medulla. Longitudinal musculature very powerfully developed. Inner longi¬

tudinal muscle fills two-thirds thickness of cortex, arranged in bundles of variable size with 2-40 fibres

per bundle. Bundles arranged in 6-8 irregular layers around inner part of cortex. Outer longitudi¬

nal muscles arranged in ring of small bundles, 2-3 deep, with 2-10 fibres per bundle. Two rings of lon¬

gitudinal muscle separated by narrow band of parenchyma. Transverse muscles fine, arranged in nar¬

row band internai to longitudinal muscles. Dorso-ventral muscles well developed, single, irregularly

arranged, Crossing medulla at irregular intervals, passing between bundles of inner longitudinal muscles.

Longitudinal osmoregulatory canals paired. Ventral canal straight-sided, of considérable internai dia-

meter. Transverse canal connects left and right ventral canals at posterior margin of each proglottis.

Dorsal canal much narrower than ventral canal, présent in scolex and neck only. Absent in strobila

posterior to level of proglottides with génital analgen. Genitalia paired. Génital ducts cross longitu¬

dinal osmoregulatory canals dorsally. Génital atrium very small, situated in posterior half of latéral

proglottis margin. Cirrus sac elongate, extremely narrow, extending beyond osmoregulatory canal.

Cirrus very slightly coiled, narrow, heavily armed with short stout bristles. Armature seen only in sec¬

tions of cirri in gravid proglottides. Internai and external séminal vesicles absent. Vas deferens thin-

walled, greatly coiled. Coils extend in transverse plane from médial border of ventral osmoregulatory

canal to poral border of vitellarium, dorsal to ovary. In longitudinal plane, coils extend from trans¬

verse osmoregulatory canal at anterior margin of proglottis to vitellarium. Following réduction in dia-

meter, vas deferens runs along dorsal edge of medulla with vasa efîerentia branching from it. Testes

numerous, occupying whole dorsal plane of medulla in single transverse layer and 3-5 longitudinal rows.

Testes fill entire space between transverse osmoregulatory canals and extend laterally beyond female

genitalia to ventral osmoregulatory canals. Vagina very narrow, surrounded by loose layer of cells.

Opens independently to génital atrium posterior to cirrus sac. Cirrus sac dorsal to vagina on right hand

side of strobila, ventral to it on left hand side. On left hand side, vagina crosses cirrus sac médial

to ventral osmoregulatory canal and passes to ventral side of medulla. Vagina terminâtes in elon¬

gate, thin-walled séminal réceptacle, immediately dorsal to ovary. Séminal réceptacle sinuous,

widest at poral end, diminishing in size aporally. Ovary fan-shaped, composed of numerous

clavate lobules, close to ventral osmoregulatory canal on ventral aspect of medulla. Vitellarium

ovoid, lobulate, posterior and dorsal to ovary. Hemispherical réservoir présent on dorsal surface

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

at apex of ovary. Ovarian duct curves dorsally joining with aporal end of séminal réceptacle

to form short, dorsally-directed fertilisation duct. Vitelline duct joins fertilisation duct and common

duct coils medially and dorsally, passes along dorsal margin of medulla then anteriorly and ventrally

to utérus. Mehlis’ gland greatly reduced. Represented only by layer of spindle-shaped cells sur-

rounding uterine duct as it passes along dorsal border of medulla. Utérus net-like, restricted to

medulla in early stages of development. During filling, utérus becomes complex tubular network

extending across ventral aspect of medulla, Crossing ventral osmoregulatory canal dorsally. Gravid

utérus sac-like, extending to latéral margins of proglottis, with only vestiges of initial diverticula

remaining. Egg spherical, thick-shelled. Inner membrane présent. Pyriform apparatus with two

short polar horns in early stage of development. Fully developed pyriform apparatus with two

narrow, apposed horns ending in small hollow cône.

Remarks. Only a limited amount of material of this species was available for study. The

type material was found to be in a very poor State of préservation as whole specimens stained very

poorly and histological sections showed the internai organs in an advanced State of autolysis. Fortu-

nately, four further specimens from Rhea americana from Brazil were obtained. Although contracted

and impossible to study as whole mounts, serial sections showed the specimens to be in an excellent

State of préservation and it was possible to establish the morphology of the species from them.

Fuhrmann’s descriptions (1904, 1021) were necessarily incomplète as they were based on rather

unsatisfactory specimens, however some additional morphological details were obtained from the

types. Fuhrmann (1921) described and illustrated a poorly developed pyriform apparatus with two

short polar horns, as shown in fig. 80. Spasskii (1951) speculated that this was merely a stage in the

development of the pyriform apparatus as identical forms occurred in the development of the structure

in Moniezia spp. His spéculations are correct as in other parts of the type material there are fully

developed eggs in which the pyriform apparatus is drawn into two parallel, elongate horns terminating

in a small conical “ cap ”, identical to that of Moniezia expansa (Rudolphi, 1810).

In spite of a relatively full account of the morphology of the parasite, its taxonomie position is

not easy to establish. Using Spasskii’s keys (1951), it belongs to the genus Moniezia Blanchard, 1891

and does in fact hâve numerous features in common with this genus, namely that the utérus is highly

reticulated, internai, and external séminal vesicles are absent, the vagina lays dorsal to the cirrus sac

on one side of the strobila and ventral to it on the other and in the structure of the pyriform apparatus.

A number of différences exist however. The scolex and suckers are extraordinarily strongly developed

as is the longitudinal musculature, the cirrus sac is extremely long and slender, the vas deferens is a

very large, greatly coiled organ, the séminal réceptacle is sinuous but has its greatest internai diameter

at its poral extremity (this is similar to Moniezia mettami Baylis, 1934 redescribed by Mahon, 1954),

and Mehlis’ gland which is a prominent spherical organ in species of Moniezia is reduced to a single

layer of cells. Yet none of these criteria in themselves appear to be sufilcient for the érection of a

new genus. Furthermore, as is évident from this révision, the Monieziinae in Central and South

American animais appear to be in need of révision from further collections. Apart from Moniezia

rugosa (Diesing, 1850) in South American primates, the présent révision has added M. bequaerti (Vigue-

ras, 1943) comb. nov. from a Central American rodent to the genus. In addition, C. viscaciae is also

found in a South American rodent. Consequently, Cittotaenia rhea is here assigned provisionally to

the genus Moniezia pending a detailed review of the group, when, hopefully, the true associations of

the parasite can be established.

The spécifie name given by Fuhrmann in 1904 was rhea however, in 1908 and 1932 Fuhrmann

gave it as rheae, leading to some confusion in the subséquent literature. The original name, rhea,

is used in the current review.

Following the initial collection, the parasite has been reported only from rheas in an American

zoo (Keahey and Trapp, 1969).

Source : MNHN, Paris

IAN BEVERIDGE

Table 16. — MEASUREMENTS OF MONIEZIA RHEA (in mm)

Fuhrmann

Présent description

(1904)

(1921)

Types

New material

Length

50-90

115-228

Width

—

3-6

Scolex diam.

1.2

1.03-.146

1.2-1.3

Sucker diam.

0.7 X 0.55

0.62-0.78 X

0.70-0.75 X

0.52-0.58

0.50

Neck

1.7-2.3

Mature

proglottis

1.9 x 0.4

2.3 x 0.13

Gravid proglottis

3.0 x 0.4

3.0 x 0.32

Cirrus sac

0.6 x 0.02

0.4 x 0.06

0.29-0.9 x

0.026-0.1

0.38 x 0.02

No. testes

approx. 110

approx. 100

approx. 120

Testis size

0.05-0.06

0.05

0.02-0.06 X

0.1

Séminal réceptacle

0.06-0.3 X

0.03-0.13

Ovary

0.5

0.26-0.36 X

0.32-0.40 x

0.16-0.21

0.05-0.10

Vitellarium

0.19

0.12 x 0.08

0.01-0.15 X

0.03-0.04

Dorsal osmoregulatory

canal

0.013 (neck)

—

Ventral osmoregulatory

canal

0.06-0.08

0.026-0.039

0.05-0.14

Egg

0.070

0.055

0.06

Pyriform apparatus

0.010-0.012

0.035

—

Oncosphere

0.012-0.013

0.01

Moniezia bequaerti (Vigueras, 1943) comb. nov.

Figs 91-97, Table 17.

Cittotaenia bequaerti Vigueras, 1943 : 11-13 (as bequarti or bequaerti), figs 12-15.

Cittotaenia becquarti Lopez-Neyra, 1954 : 113.

Pseudocittotaenia bequaerti (Vigueras, 1943) Tenora, 1976 : 15.

Types. From Capromys pilorides (Say, 1882) (Rodentia : Capromyidae), Havana, Santa Clara

and Matanzas, Cuba. Paratype in USNM.

Matériel examined. 6 specimens from Capromys pilorides, Cuba, 1959, J. G. Baer, UN ;

paratype.

Description. Small, narrow cestodes. Scolex large, conical in dorso-ventral view, with base

of cône directed anteriorly. Suckers large, muscular, cup-shaped, anteriorly directed, situated at

anterior extremity of scolex. Scolex merges imperceptibly into elongate neck. Proglottides cras-

pedote, with narrow, straight-edged vélum overhanging adjacent proglottis, extended transversely.

Source : AANHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Mature proglottides with approximate length : width ratio of 1 : 4. Gravid proglottides with ratio

of 1 : 2. Longitudinal musculature weakly developed, consisting of two rings of large diameter fibres

around inner région of cortex. Fibres occasionally grouped in twos or threes but never in larger

bundles. Transverse muscles fine, in band internai to longitudinal muscles. Dorso-ventral muscles

fine, few in number, irregularly arranged. Longitudinal osmoregulatory canals paired. Ventral

canal wider than dorsal canal, lying internai to dorsal canal. Transverse canal connects left and right

ventral canals at posterior margin of each proglottis. Genitalia paired. Génital ducts pass longitudinal

osmoregulatory canals dorsally. Génital atrium shallow, situated in middle of latéral proglottis

margin. Cirrus sac short, oblong, thick-walled, reaching longitudinal osmoregulatory canals. Cirrus

uncoiled, unarmed. Internai and external séminal vesicles absent. Vas deferens thin-walled, highly

convoluted. Vasa efferentia not seen. Testes numerous, extending through most of proglottis medulla.

Distribution of testes limited laterally by longitudinal osmoregulatory canals. Testes absent anterior

to ovaries and génital ducts, arranged in 1-2 transverse and 5-8 longitudinal rows on dorsal aspect

of medulla. Vagina narrow, thin-walled, opening to génital atrium posterior to cirrus sac. Vagina

lies ventral to cirrus sac on right hand side of strobila, dorsal to it on left hand side. Séminal réceptacle

elongate, clavate, dorsal to ovary. Ovary fan-shaped, composed of numerous clavate lobules, situated

on ventral aspect of medulla in anterior half of proglottis close to longitudinal osmoregulatory canals.

Vitellarium oval or sub-triangular, posterior and dorsal to ovary. Mehlis’ gland spherical, situated

between ovary and vitellarium. Utérus net-like at commencement of development, extending over

entire area of medulla. Islands of parenchyma between uterine strands disappear as utérus fills.

Gravid utérus sac-like, filling proglottis. Egg spherical, thick-shelled. Oncosphere surrounded by

pyriform apparatus, terminating in two elongate horns.

Remarks. A single type specimen was available for study, as well as the material described

above from the type host and locality. The description differs from that given by Vigueras (1943) in

that the cirrus armature he described was not found. The original description States that the testes do

not lie external to the longitudinal osmoregulatory canals (“ pero no traspasan los vasos excretores ”)

however fig. 13 accompanying his description clearly shows some 15 testes latéral to the osmoregu¬

latory canals on each side of the strobila. The drawing is apparently in error as no testes were found

latéral to the osmoregulatory canals in the material redescribed above.

In spite of the fact that Vigueras (1943) described the developing utérus as reticulate (“ utérus

con ramificaciones simples haciendos sacciformes con lobulaciones ”) he still placed the new species

in the genus Cittotaenia. The structure of the utérus in this species is typical of the généra Moniezia

and Diandrya Darrah, 1930, and is assigned to the former as it lacks séminal vesicles and a prostate

gland, the presence of which characterises the genus Diandrya. Using Spasskii’s key (1951), M. be-

quaerti belongs to the sub-genus Baeriezia Skrjabin and Schulz, 1937 since it lacks interproglottidal

glands. It is easily distinguished from M. baeri Skrjabin, 1931 by its small size and from M. mettami

Baylis, 1934 by the lack of a vaginal sphincter. M. bequaerti most closely resembles M. rugosa (Diesing,

1850) a parasite from South American primates, however the two species can be distinguished from

one another by the number of testes per proglottis and by the presence of a vaginal sphincter in

M. rugosa.

M. bequaerti is the first species of this genus to be recorded from rodents. With the exception

of M. rugosa and M. rhea comb. nov. ail other species of the genus are parasites of herbivora.

In Vigueras’ original description (1943), the spécifie epithet is spelt bequaerti in the discussion

and in the captions to the figures, in honour of a Dr. Bequaert, however, at the heading of the descrip¬

tion it is given as Cittotaenia bequarti sp. nov. Since this is a typographical error rather than an

intentional omission of the e, the name bequaerti has been used above.

Source : MNHN, Paris

IAN BEVERIDGE

Table 17. — MEASUREMENTS

OF MONIEZIA

BEQUAERTI (in mm)

Vigueras

Présent

(1943)

description

Length

120-210

up to 210

Width

4.7

Scolex diameter

1.5 X 1.8

1.10-.135

Sucker diameter

0.42

0.45-0.60

Neck

1.0-1.5

Mature proglottis

4.0-4.8 x

2.8-3.2 x

1.0-1.3

0.7-0.8

Gravid proglottis

2.0 x 2.5-3.0

2.6-4.4 x

1.5-2.2

Cirrus sac

0.24-0.28 x

0.2-0.3 x

0.065-0.080

0.06-0.07

No. testes

200-240

approx. 150

Testis size

0.48-0.56

0.07

Séminal receptable

0.20-0.45 x

0.09-0.25

Ovary

0.20-0.25 x

0.09-0.16

Vitellarium

0 . 11 - 0.16 x

0.10-0.15

Mehlis’ gland

Dorsal osmoregulatory canal

0.04

Ventral oremoregulatory canal

0.07

Egg

0.054-0.058

0.07

Pyriform apparatus

0.045

Oncosphere

0.012-0.016

0.020

SPECIES INCERTAE SEDIS

Cittotaenia tachyglossi

Figs 98-102, Table 18.

Cittotaenia tachyglossis Johnston, 1913 : 75, 77-78, figs 1-3.

Types. From Tachyglossus aculeatus Shaw, 1792, Townsville, Queensland, Australia, 1911,

whole mounts, spirit material and serial section, UAHC 1115, 1122.

Material examined.

From Tachyglossus aculeatus Shaw, 1792. Types ; fragments of several specimens, Townsville

Queensland, 1912, P. A. Maplestone, LSTM.

Source : MNHN, Paris

REVISION OF THE CITTOTAENIA COMPLEX

Description. Small cestodes. Scolex unarmed, approximately hemispherical, merging into

neck région. Suckers hemispherical, thick-walled, embedded in scolex, openings directed anteriorly

and laterally. Proglottides craspedote with narrow, mainly straight-edged vélum overhanging adjacent

proglottis, extended transversely. Mature proglottides with approximate length : width ratio of 1 : 4.

Longitudinal musculature moderately developed, arranged in two rings of muscle bundles around

cortex. Bundles small with few fibres. Fibres of inner bundle ring larger than those of outer ring.

Transverse muscles fine, in band internai to longitudinal muscles. Dorso-ventral muscles very fine,

sparse, Crossing cortex and medulla at irregular intervals. Longitudinal osmoregulatory canals paired.

Ventral canal slightly wider than dorsal canal, lying internai to dorsal canal. Transverse canal connects

left and right ventral canals at posterior margin of each proglottis. Anterior to suckers, transverse

anastomosis connects ventral canals from either side. Similar anastomosis joins dorsal canals. Four

canals continue anteriorly from anastomoses. On each side of strobila, short dorso-ventral loop

connects dorsal and ventral canals. Genitalia paired. Génital ducts cross longitudinal osmoregu¬

latory canals ventrally. Génital atrium small, situated in middle of latéral proglottis margin. Cirrus

sac narrow elongate, extending beyond osmoregulatory canals. Cirrus slightly coiled, unarmed.

Proximal extremity of cirrus sometimes slightly dilated. Internai and external séminal vesicles

absent. Vas deferens thin-walled, greatly coiled near proximal pôle of cirrus sac, passing posteriorly,

dorsal to ovary. Vasa efîerentia not seen. Testes numerous, scattered throughout proglottis medulla

on dorsal aspect, overlying female genitalia and occasionally lying just latéral to longitudinal osmore¬

gulatory canals. Testes in single dorso-ventral layer. Vagina opens to génital atrium posterior to

cirrus sac. Prominent, discrète, muscular sphincter présent at distal extremity of vagina. Séminal

réceptacle absent. Ovaries situated in latéral quarters of proglottis medulla, on ventral aspect, fan-

shaped, composed of numerous clavate lobules. Vitellarium posterior and dorsal to ovary, narrow

elongate, extending medially almost to midline of proglottis. Mehlis’ gland anterior to vitellarium

at apex of ovary. Uterine duct emerges from Mehlis’ gland on dorsal aspect, runs anteriorly to beyond

anterior margin of ovary, terminating in slight enlargement. Utérus not seen. Gravid proglottides

not présent.

Remarks. Although incomplète, the présent description extends that of Johnston (1913) as

well as recording Maplestone’s collection from the type host and locality of apparently the same species.

Johnston’s description is in error in stating that the génital ducts pass the osmoregulatory canals

dorsally, but the error is probably due to the fact that Johnston did not préparé sections of the parasite.

Apart from the fact that C. tachyglossi has paired genitalia, it is very similar to the single-pored

species Linstowia echidnae (Thompson, 1893) from the same host species. The position of the génital

ducts ventral to the osmoregulatory canals and the aporally extended vitellarium are both character-

istics of L. echidnae, the latter feature occurring only in this species. The similarities pointed out

suggest that C. tachyglossi probably represents a new genus in the family Linstowiidae, liowever, the

form of the utérus is not known and it is considered best to leave the parasite under its présent name

pending collection of gravid specimens and the completion of the description.

If the above suggested taxonomie position is correct, it is probable that C. tachyglossi developed

from L. echidnae by duplication of the genitalia, as has been suggested for other cestodes (Baer, 1927,

Spasskii, 1951, Baer, 1955).

Source : MNHN, Paris

IAN BEVERIDGE

Table 18. — MEASUREMENTS OF CITTOTAENIA TACHYGLOSSI (in mm)

Johnston, Présent study

1913 (From types)

Length (fragments)

Width

0.56

2.0

Scolex diameter

0.56

0.65-0.79

Sucker diameter

0.13

0.21-0.28

Neck

0.6-1.1

Mature proglottis

2.2 x 0.4

Cirrus sac

0.2 X 0.033-0.05

0.43-0.50 X

0.04-0.06

No. testes

per proglottis

approx. 200

Testis size

0.015

0.03-0.05

Ovary

0.24 X 0.16

Vitellarium

0.45 X 0.04

Mehlis’ gland

0.06

Dorsal osmoregulatory

canal

0.01

Ventral osmoregulatory

canal

0.02

SPECIES INQUIRENDAE

Cittotaenia quadrata

Cittotaenia quadrata von Linstow, 1904, : 680-681, figs 3-4.

Ctenotaenia quadrata (von Linstow, 1904) Spasskii, 1951 : 271, fig. 127.

Neoctenotaenia quadrata (von Linstow, 1904) Tenora, 1976 : 13.

Cittotaenia pectinata (Goeze, 1782) pro parte Baer, 1927 : 55.

Types. From Lagidium peruanum Meyen, 1833, Peru. Slides of serial sections in UN.

Material examined. Types.

Remarks. The type material is in too poor a State to permit redescription and the original

description is inadéquate to détermine satisfactorily the taxonomie position of the parasite. As

indicated earlier, C. quadrata may be similar to C. viscaciae and they may even corne from the same

host, however, the matter cannot be taken further without fresh material.

Cittotaenia dratchynskii

Cittotaenia dratchynskii Romanovich, 1915 : 451.

Moniezia ? dratchyuskii (Romanovich, 1915) L<5pez-Neyra, 1954 : 121.

Types. From Rangifer tarandus Linnaeus, 1758, Western Siberia, présent whereabouts unknown.

Source : MNHN, Paris

REVISION OF THE C1TTOTAENIA COMPLEX

Cittotaenia krishnai Nama, 1972

Cittotaenia krishnai Nama, 1972 : 52-53, fig. 1.

Types. From Capra hircus Linnaeus, 1758, Jodhpur, India.

Remarks. Unfortunately, the types of this species hâve been lost (H. S. Nama, personal

communication), and as it is impossible to détermine the position of this species from the original

description, it must be placed as a species inquirenda.

DISCUSSION

The présent review, together with an earlier révision of the anoplocephalid cestodes of Australian

marsupials (Beveridge, 1976) constitute a révision of most of the généra of anoplocephaline cestodes

with paired genitalia parasiting mammals. Only two généra hâve been neglected, Ectopocephalium Rausch

and Ohbayashi, 1974 and Diuterinotaenia Gvozdev, 1961, both of which are monotypic, recently and

adequately described and with characteristic morphological features which would prevent their confusion

with other généra. A third genus Eranuides Semenova, 1972 based on two incomplète specimens

from a reindeer was initially placed in the subfamily Anoplocephalinae (Semenova, 1972) though the

morphology of the mature proglottis is remarkably similar to that of species of Moniezia. Tenora

(1976) transferred the genus to the sub-family Monieziinae.

The various généra described in this review, their component species and related généra together

with their distinguishing characteristics are set out in Table 19 and a key to them is provided below.

As suggested by Beveridge (1976), provided patterns of infra-specific variation are taken into

account, the distribution of the testes within the proglottis provides important generic criteria within

the sub-family under considération. The structure of the vagina and séminal réceptacle was also

found to be characteristic for a genus in several instances and may be of greater taxonomie value

than realised in the past.

The présent révision has shown that the parasites of the “ Cittotaenia complex ” are exclusively

parasites of mammals. Species previously placed in this group from avian hosts hâve invariably

had to be removed to other généra or subfamilies following redescription. As indicated in Table 19,

each genus is, for the most part restricted to one family of host species, although minor exceptions

do exist. Although not taken into account in the taxonomy of the parasites the finding that related

parasites share similar hosts suggests that the classification presented conforms to some extent to

what one might expect of a “ natural classification

In view of the relatively extensive literature on the phylogeny of the Anoplocephalinae (see

Baer, 1927, Spasskii, 1951, Tenora, 1976) the présent révision requires that some comment be made

on the validity of earlier théories. The possible phylogeny of the Australian généra has been discussed

by Beveridge (1976).

Spasskii (1951) argued that the généra Mosgovoyia and Ctenotaenia arose by duplication of the

genitalia of members of the généra Schizorchis Hansen, 1948 and Paranoplocephala Lühe, 1910 respec-

tively. The argument was based on the fact that in the généra Schizorchis and Mosgovoyia, both parasites

of lagomorphs (Ochotonidae and Leporidae respectively), the testes lie entirely posterior to the utérus.

In spite of the substantial changes to the genus Mosgovoyia made in the présent révision, the rétention

of the occurrence of testes posterior to the utérus as a key generic character in no way contradicts

Spasskii’s hypothesis. The présent inclusion of ail of the anoplocephaline cestodes of leporids in one

genus rather strengthens the hypothesis.

Similarly, the changes in the composition of the genus Ctenotaenia in no way invalidate the

basic hypothesis. Although species of the rodent généra Marmota and Citellus are found in North

Source : MNHN, Paris

IAN BEVERIDGE

America as well as Europe and Asia, Ctenotaenia marmolae appears to be a strictly Eurasian species

as investigations of the parasites of marmots in North America hâve not revealed its presence there

(Darrah, 1930, Philip, 1938, Rausch, 1951). Species of Paranoplocephala occur in Eurasian marmots

(P. transversaria (Krabbe, 1879) and P. ryjikovi Spasskii, 1950) which hâve a utérus limited to the

cortex during the entire course of its development and in which the testes are disposed aporally to

the female genitalia but both anterior and posterior to the utérus, characteristics which are also présent

in Ct. marmotae. In fact, duplication of a proglottis of the type found in these species of Paranoplo¬

cephala gives precisely the morphological features of Ct. marmotae.

In certain North American species of Paranoplocephala , such as P. troeschi Rausch, 1946 the

testes are again limited to the région aporal to the female genitalia, but they are entirely posterior

to the utérus which is at the anterior extremity of the proglottis and which crosses the longitudinal

osmoregulatory canals ventrally, terminating anterior to the cirrus sac. Duplication of such a pro¬

glottis would give almost exactly that of a species of Pseudocittotaenia and since the features of the

utérus in this latter genus are totally different to those found in related généra with paired genitalia,

it seems reasonable to suggest that the genus Pseudocittotaenia arose in this fashion, by a duplication

of genitalia, in a manner parallel to Ct. marmotae in Europe and Asia.

Spasskii (1951) and Tenora (1976) considered the subfamily Monieziinae to be “ phylogenetically

younger ” than the sub-family Anoplocephalinae and in particular, the genus Cittotaenia was considered

to hâve arisen by duplication of the genitalia of species of Andrya Railliet, 1893 (see Tenora, 1976).

This hypothesis seems unlikely as the utérus of C. denticulata is scarcely reticulated whilst that of species

of Andrya is net-like. Furthermore, the genus Diandrya Darrah, 1930 seems to be a far more logical

product of the duplication of the genitalia of species of Andrya. The présent révision gives little

or no due as to the phylogenetic affinities of the genus Cittotaenia. In the writer’s opinion, it would

be better to refrain from speculating upon the matter until the systematics are better understood

and until a reasonable hypothesis présents itself.

Table 19. — RELATIONSHIPS OF THE GENERA OF ANOPLOCEPHALINE CESTODES

WITH PAIRED GENITALIA PARASITISING MAMMALS AND THE MONIEZIINE

GENUS CITTOTAENIA.

Genus and Species

Hosts

Char acier istic

feature(s )

Reference

Cittotaenia Riehm, 1881

C. denticulata

C. viscaciae

Lagomorphs

(Leporidae)

and rodents

(Chinchillidae)

Utérus slightly reticulate,

with anterior and posterior

diverticula

Ctenotaenia Railliet, 1893

Ct. marmotae

Rodents

(Sciuridae)

Testes anterior and poste¬

rior to utérus, between fe¬

male genitalia

Diuterinotaenia Gvozdev, 1961

D. spasskii

Lagomorphs

(Ochotonidae)

Uteri paired, arranged lon-

gitudinally

Gvozdev (1961)

Ectopocephalium Rausch and Ohbayashi, 1974

E. abci

Lagomorphs

(Ochotonidae)

Long neck, scolex covered

in glands

Rausch and

Ohbayashi (1974)

Source : MNHN, Paris

REVISION OF THE C1TTOTAEN1A COMPLEX

Genus and Species

Host

Characteristic

fealure[s )

Reference

Mosgovoyia Spasskii, 1951

M. pectinata

Lagomorphs

Testes entirely posterior to

M. clenoides (Leporidae)

M. variabilis

Paramoniezia Maplestone and Southwell, 1923

utérus, utérus restricted to

medulla or terminating in

cortex posterior to cirrus sac

P. suis

? Pigs (Suidae)

Testes scattered throughout

P. johnstoni

Phascolotaenia Beveridge, 1976

and marsupials

(Vombatidae)

medulla, séminal réceptacle

large, pyriform, merging in-

to vagina.

Beveridge (1976)

P. comani

Marsupials

Most testes posterior to ute-

Progamotaenia Nybelin, 1917

(Vombatidae)

rus, two small groups ante¬

rior to utérus near osmore¬

gulatory canals. Séminal

réceptacle as in Progamo¬

taenia

Beveridge (1976)

P. bancrofti

Marsupials

Testes anterior to utérus,

P. aepyprymni

(Macropodidae

extending latéral to female

P. diaphana

P. effigia

P. ewersi

P. festiva

P. lagorchestis

P. proterogyna

P. villosa

P. zschokkei

Pseudocittotaenia Tenora, 1976

and Vombatidae)

genitalia. Vagina narrow,

séminal réceptacle circular

Beveridge (1976)

P. praecoquis

Rodents

Utérus anterior to testes,

P. glandularis

(Geomyidae)

crosses osmoregulatory ca¬

nals ventrally, terminâtes

anterior to cirrus sac.

KeY TO GENERA OF ANOPLOCEPHALINE CESTODES WITH PAIRED GENITALIA PARASITISING MAMMALS

1. Utérus paired, longitudinal. Diuterinotaenia

Utérus single or paired, transverse. 2

2 (1). Neck extremely long, scolex embedded in diverticulum of host’s intestine, scolex covered with glands.

Ectopocephalium

Neck short or absent, parasite restricted to lumen of intestine or biliary System, scolex without glands.

3 (2). Utérus in tubular stage extends beyond osmoregulatory canals ventrally, terminating anterior tolevel

of cirrus sac. Utérus at anterior extremity of proglottis. Pseudocittotaenia

Source : MNHN, Paris

IAN BEVERIDGE

Utérus not at anterior extremity of proglottis. Either restricted to medulla in tubular stage, or if

not, then crosses osmoregulatory canals dorsally, terminating in cortex posterior to cirrus sac

and vagina. 4

4 (3). Testes restricted to région between female genitalia, lying both anterior and posterior to utérus. .. .

Ctenotaenia

Some testes lying poral to female genitalia, or if restricted to area between genitalia, then entirely posterior

to utérus. 5

5 (4). Testes in single band or two groups entirely posterior to utérus. Mosgovoyia

Testes entirely or in part anterior to utérus. 6

6 (5). Testes arise in single band or two groups anterior to utérus. Progamotaenia

Some testes posterior to utérus. 7

7 (6). Testes distributed throughout medulla. Paramoniezia

Testes in band posterior to utérus with two small groups anterior to utérus, in latéral extremities of

proglottis medulla. Phascolotaenia

Source : MNHN, Paris

REVISION OF THE C1TTOTAENIA COMPLEX

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Source : MNHN, Paris

Ctenotaenia marmotae. Fies 1-10.

1 "“'“.U 2 : ““S

ing musculature ; 8. Scolex showihg osmoregulatorv System, 9. Egg

Source : MNHN, Paris

10. Mature proglotti».

Mosgovoyia pectitala. Figs 11-25.

11. Scolex of relaxed specimen; 12. Scolex from type specimen of Citlotaenia bursaria von Linstow, 1906 (now a synonvm

of M. nectinata) showing efTects of severe "contraction on scolex; 13. Transverse histological section through strohila

showing nuisenlatiire ; 14. Fcmale génital comple.v: 15. Cirrus sac and distal vagina from specimen from Oryctolagus

caniailiix, England; 16. Cirrus sae ami vagina from specimen from Le/ms americanus front North America showing

extern of variation in lengtli of cirrus sac with respec t to vagina.

Source : MNHN, Paris

17-19. Specimens from Oryctolagus ctiniculns , England. 17. Mature proglottis; 18. Pre-gravid proglottis with

19. Gravid proglottis showing accessorv osmoregulatory canals.

•om Lepus townsendi , USA, USNM 59076, mature proglott.s showing slight break in band of testes ; 21. From Lepus

californiens , USA, USNM 59259, mature proglottis showing complété séparation of the testes into 2 latéral groups;

22 From Svlvilagus nutalli granyeri , USA, USNM 59073, two adjacent proglott.des from strob.la sho^ng varmt.on

in testis distribution between proglottides of one worm;23. Normal embryo and pynform apparatus; 24. Pynform

apparatus showing (as in fig. 25) vacuolations within arms ; 25. Egg.

Scale lines : figs 11-23, 0.1 mm; figs 23-25, 0.01 mm.

Source : MNHN, Paris

moagovoyia ctenmdei. Figs 26-34

*■ Sco1 " ol > , '"ï ««•«' *»*,.

plex; 30. Egg. ’ n8verse 8et,lon °* strobila showing musculature ; ;

Source : MNHN, Paris

31-33. Mature proglottides showing variation in testis

or paired utérus.

Scale Unes : figs 26-29, 31-34, 0.1 mit

distribution ; 34. Gravid proglottides showing proglottides with

i ; fig. 30, 0.01 mm.

Mosgovoyia variabilis. Fies 35-40.

3d. Scolex; 36. Egg; 37. Transverse histological section of strobila showing n

tal complex; 39. Gravicl proglottis; 40. Mature proglottis.

Scale Unes : figs 35, 37, 38-40, 0.1 mm; fig. 36, 0.01 mm.

scuiature ; 38. Génital ducts and female geni-

Source : MMHN, Paris

Pseudocittotaenia praecoe/uix. Fies 41-47.

41. Scolex ; 42. Cirrus sac and distal vagina. dorsal view, showing utérus Crossing the osmoregulatorv canals ventraUy and tcrminating anterior

to cirrus sac; 43. Pre-mature proglottis showing distribution of testes posterior to ute.-us; 44. Egg: 4a. Latéral part of two proglottides,

ventral view, showing variation in testis distribution within woriu.

Source : MNHN, Paris

v /r

Pseiidiia'lloltieiiiii ylaii(lttlari.i. Fu;> 48-54.

From holotvpe from Thomnmys talpair/es. Fig. 48. From other material from sanie liost speries.

48. Scolex ; 49. Cirrus sac and distal vagina; 50. Egg; 51. Gravid proglottis.

Source : MNHN, Paris

Jglottis ; 53. Mature proglottis; 54. Post-mature pri

Cittotaenia denticulata. Fies 55-65.

55. Scolex showing osmoregulatory System ; 56. Cirrus sac ; 57. Female génital complex ; 58. Egg ; 59. Embrvo with three

armed pvriform apparatus; 60. Latéral région of proglottis showing osmoregulatory system; 61. Transverse histolo-

gical section of strobila showing musculature.

Source : MNHN, Paris

62. Mature proglottis; 63. Prématuré proglottis shouing utérus: 64. L'te

ami the formation of divertieula : 63. Gravid utérus.

Seale lines : figs 33-57, 60-63, 0.1 mm; figs 58. 59, 0.01

■rus during development

sliowing sliglu retictdations

Source : MNHN, Paris

Ciflolaenia viscaciae. Fies 60-72. Ali drawings froin type.

66. Scolex; 67. Cirrus sac: 68. Mature proglottis; 69. Egg; 70. Génital ducts and female génital cotnplex.

Source : MNHN, Paris

71. Longitudinal section of proglottis showing reticulated utérus; 72. Transverse section of proglottis showing musculature

and disposition of variôus génital organs.

Scale lines : Figs 66-68, 70-72, 0.1 mm ; fig. 69, 0.01 mm.

Source : MNHN, Paris

Slrinyopotaenia .psittacea. Fies 73-78. Ail drawings from type.

73. Latéral région of mature proglottis reconstructed from serial sections; 74. Histological section of gravid utérus; 75. Egg.

Source : MNHN, Paris

76. Histological section of portion of gravid utérus ; 77. Cirrus sac and distal vagina ; 78.

showing musculature and génital organs.

Scale Unes : figs 73, 74, 76-78, 0.1 mm; fig. 75, 0.01 ram.

Transverse histological section

Source : MNHN, Paris

Moniezia rhea. Figs 79-90. Figs 79-83 from types,

79. Scolex; 80. Egg showing partial development of pyrifor

®2- Egg with fully developed pvriform apparatus : 83. Cirrus sac; 84. Transverse histological section showing musculature;

85. Transverse histological section of neck showing présente of two pairs of longitudinal osmoregulatorv canals;

86. Longitudinal histological section of mature proglottis showing reticulated utérus; 87. Longitudinal histological

section of mature proglottis showing reticulated utérus; 87. Longitudinal histological section of post mature proglot¬

tis showing developing utérus ; 88. Longitudinal histological section through utérus of gravid proglottis ; 89. Trans¬

verse histological section of mature proglottis showing genitalia, right hand side of proglottis ; 90. Transverse histo¬

logical section of left hand side of proglottis showing génital ducts.

S cale lines : figs 79-80, 83-90, 0.1 mm; figs 81, 82, 0.01 mm.

Source : MNHN, Paris

Moniezia bequaerti. Figs 91-97.

91. Scolex; 92. Egg; 93. Mature proglottis; 94. Latéral région of post-mature proglottis with developing, reticulated uté¬

rus ; 95. Cirrus sac and distal vagina ; 96. Subséquent stage of uterine filling with réduction in the number of réticu¬

lations ; 97. Gravid utérus.

Scale lines : figs 91, 93-97, 0.1 mm; fig. 92, 0.01 mm.

Source : MNHN, Paris

'■ SC °b 9 . 9 ' C ‘ rrUS f C “ ld d ‘ Stal , Vag ' na; 10 °- Mature Promis; 101. Female génital complex; 102. Transverse histo-

logical section showing musculature and génital ducts. H iransverse tusto-

Source : AANHN, Paris

IMPRIMERIE NATIONALE

7 564 016 6 02

Source : MNHN, Paris

2 9 JUIL io?o

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