Systematic Parasitology 54: 81–88, 2003.

© 2003 Kluwer Academic Publishers. Printed in the Netherlands.

81

Viscachataenia n. g., with the redescription of V. quadrata (von Linstow,

1904) n. comb. (Cestoda: Anoplocephalidae) in Lagidium viscacia
(Rodentia: Chinchillidae) from Argentina

G. Denegri1,2 , M. C. Dopchiz2 , M. C. Elissondo1,2 & I. Beveridge3

1 Departamento de Biologia, Facultad de Ciencias Exactas y Naturales, Universidad Nacional de Mar del Plata,
Funes 3250, Mar del Plata, Buenos Aires, Argentina
2 Consejo Nacional de Investigationes Cientı́ficas y Técnicas, Ciudad de Buenos Aires Argentina
3 Department of Veterinary Science, University of Melbourne, Parkville, Victoria, 3052, Australia

Accepted for publication 19th July, 2002

Abstract
A new genus, Viscachataenia, is erected to accommodate the anoplocephalid cestode Cittotaenia quadrata von
Linstow, 1904, based on a redescription of the species from material collected from the chinchillid rodent Lagidium
viscacia in Argentina. The new genus is characterised by paired genitalia, a reticulate uterus and the vagina entering
the genital atrium anterior to the cirrus-sac. Viscachataenia therefore has similarities with Monoecocestus Beddard,
1914, a genus which is common in South American rodents but which has a single set of genitalia in each segment.
Cittotaenia viscaciae (Spasskii, 1951) and Bertiella findlayi Mazza, Parodi & Fiora, 1932, also from viscachas, are
considered synonyms of V. quadrata.

Introduction M. pectinata as its type-species, and also placed in it

Two species of anoplocephalid cestodes with paired
genitalia have been described from viscachas belong-
ing to Lagidium Meyen (Rodentia: Chinchillidae)
from South America. von Linstow (1904) provided
a brief description of Cittotaenia quadrata, collected
from Lagidium peruanum Meyen in Peru, based on a
specimen sent to him by Sir Arthur Shipley in Cam-
bridge. No additional details associated with the col-
lection were provided. The species name was applied
because of the rather characteristic egg, which was
four-lobed (von Linstow, 1904). Subsequently, Joyeux
& Dollfus (1931) reported a specimen of Cittotaenia
pectinata (Goeze, 1782) probably from L. viscacia
from Valdivia, Chile, held in the collections of the
Munich Museum. They noted that the species dif-
fered somewhat from C. pectinata, a species found
normally in leporids (Beveridge, 1978), but consid-
ered that C. pectinata was a morphologically variable
species based on Baer’s review (1927) of the fam-
ily Anoplocephalidae and allocated their specimen to
this species. Spasskii (1951) erected Mosgovoyia, with
the specimen described by Joyeux & Dollfus (1931) as
a new species, M. viscaciae.
Beveridge (1978) redescribed M. viscaciae from
the single incomplete specimen, showing that it did not
belong within Mosgovoyia, as it had a reticulate uterus
and was therefore more similar to Cittotaenia Riehm,
1881 to which it was transferred. Beveridge (1978)
also examined the extant type-material of C. quadrata
and noted a number of similarities with C. viscaciae.
He concluded however that, although C. quadrata and
C. viscaciae might be the same species, the existing
specimens did not allow any firm conclusions to be
drawn.
The other species of anoplocephalid cestodes
known from viscachas have a single set of genitalia in
each segment. Mazza et al. (1932) described Bertiella
finleyi Mazza, Parodi & Fiora, 1932 from C. viscaciae
in Argentina, a species which has not been reported
since its original description, while Parra (1953) re-
ported Monoecocestus thelkeldi (Parra, 1952) from
Lagidium peruanum in Peru.
82
The collection of new material from viscachas
from Argentina has allowed a re-examination of this
problem. In the present paper, C. quadrata is re-
described from new material and C. viscaciae is placed
tentatively as a synonym of C. quadrata. Bertiella
finleyi is in part a synonym of C. quadrata. A new
genus is erected to accommodate C. quadrata, be-
ing allied with Monoecocestus Beddard, 1914, which
occurs commonly in South American rodents (Rego,
1961).
Materials and methods
Fourteen specimens of Lagidium viscacia boxi
Thomas were collected from Las Bayas, Ñorquincó
Department, Río Negro Province, Argentina. Details
of the collection methods and locality are provided by
Led et al. (1979). Cestodes were washed in running
water, fixed in 5% formaldehyde and subsequently
stored in 70% ethanol. Five cestodes were stained
with Celestine blue, dehydrated in ethanol and cleared
in methyl salicylate. The outer muscle layers were
removed using fine forceps and cestodes were subse-
quently mounted in Canada balsam. Mature regions of
cestodes were embedded in paraffin, and serial trans-
verse sections cut at a thickness of 5 µm were stained
with haematoxylin and eosin. Gravid segments were
teased apart in water to release eggs and, following
examination in water, eggs were cleared in lactophenol
and subsequently mounted permanently in polyvinyl
lactophenol. Coverslip pressure applied to some eggs
resulted in expulsion of the pyriform apparatus, allow-
ing examination of its morphology and surrounding
membranes.
Drawings were made with the aid of a drawing
tube attached to an Olympus BH microscope. Mea-
surements were made using an ocular micrometer and
are presented in millimetres, unless otherwise speci-
fied, as the range of 10 measurements followed by the
mean in parentheses.
Specimens have been deposited in the Museo
de la Plata (MLPA), Argentina and in the Muséum
d’Histoire Naturelle, Geneva (MHNG).
Viscachataenia n. g.
Diagnosis
Anoplocephalidae Cholodkovsky, 1902. Strobila
large. Segments craspedote, wider than long. Geni-
talia paired. Genital ducts cross osmoregulatory canals
dorsally. Cirrus-sac powerfully developed; internal
seminal vesicle present; external seminal vesicle ab-
sent. Testes arranged in two laterally-extended groups
on posterior border of segment. Ovaries poral. Vagina
enters genital atrium anterior to cirrus-sac. Seminal
receptacle absent. Uterus initially transverse, tubular,
becoming reticulate, then saccate. Pyriform apparatus
present. Parasitic in chinchillid rodents; South Amer-
ica. Type-species: V. quadrata (von Linstow, 1904) n.
comb.
Viscachataenia quadrata (von Linstow, 1904) n.
comb.
Syns Cittotaenia quadrata von Linstow, 1904;
C. pectinata (Goeze, 1782) sensu Joyeux & Dollfus
(1931); Mosgovoyia viscaciae Spasskii, 1951; C. vis-
caciae (Spasskii, 1951) Beveridge, 1978; Bertiella
finlayi Mazza, Parodi & Fiora, 1932 (in part)
Specimens examined
Type-specimen of Cittotaenia quadrata von Linstow,
1904, 8 slides of serial sections, 5 frontal, 3 transverse,
MNHG 126/64-71.
Type-specimen of Cittotaenia viscaciae (Spasskii,
1951), 12 slides of serial sections, MNHG 126/16-27.
Five specimens from Lagidium viscacia boxi, Las
Bayas, Argentina, MLPA 5066, MHNG INVE 33493-
4.
Description (Figures 1-12)
Large cestodes, 185-320 (250) long, maximum width
10. Scolex 0.40-0.53 (0.39, n=5) in diameter, lacking
rostellum (Figure 1); 4 sub-circular suckers, 0.13-0.17
(0.16) × 0.12-0.17 (0.16). Neck absent. Segments
transversely elongate, prominently craspedote. Im-
mature segments much wider than long, 0.32-1.00
(0.56, n=5) × 3.6-6.5 (5.2, n=5). Mature segments
0.35-1.10 (0.75) × 4.4-7.1 (5.7).
Genitalia paired. Genital atrium in middle of lat-
eral margin of segment, deep, with thick corrugated
walls. Cirrus-sac robust, with thick muscular walls,
broadest near distal extremity (Figure 4), 0.67-0.90
(0.77) × 0.22-0.39 (0.31) in mature segments, 0.70-
1.10 (0.91) × 0.34-0.39 (0.37) in gravid segments;
distal cirrus straight, thick walled, armed with spines
20 µm long (Figures 2, 4); proximal cirrus slightly
sinuous, with few spines; internal seminal vesicle
subcircular, 0.16-0.25 (0.20) × 0.10-0.19 (0.15) in

mature segments, 0.41-0.52 (0.46) × 0.21-0.32 (0.26)
in gravid segments. When everted, genital atrium
forms conical projection or genital papilla at base of
cirrus; internal seminal vesicle not evident in everted
cirri (Figure 3). Strong retractor muscle leads medially
from proximal pole of cirrus-sac (Figure 4). Exter-
nal seminal vesicle absent; vas deferens coiled, passes
posteriorly and medially towards testes; in vicinity of
testes, vas deferens branches, giving off fine vasa ef-
ferentia to individual testes. Testes small, spherical,
0.03-0.05 (0.04) in diameter, distributed in 3-4 dorso-
ventral layers on dorsal aspect of medulla (Figure 5);
testes arranged in 2 groups per segment, with 80-107
(90) testes per group. Testes extend from vitellarium
medially towards mid-line of segment; medial termi-
nation of testis groups variable, with some individual
testes or small groups of testes scattered between
major fields; considerable variation occurs between
adjacent segments (Figure 9).
Vagina simple, tubular, entering genital atrium an-
terior to cirrus-sac (Figure 4). Vagina runs initially
along anterior margin of cirrus-sac, then runs poste-
riorly and medially, terminating between ovary and
vitellarium; proximal vagina variously distended by
sperm (Figures 4, 5), not forming distinctive semi-
nal receptacle. Ovary flabelliform, on ventral aspect
of medulla, 0.23-0.45 (0.36) × 0.35-0.45 (0.41), with
numerous clavate lobules radiating anteriorly and pos-
teriorly from central oöcapt. Vitellarium reniform,
slightly lobulate, dorsal and slightly poral to ovary, at
level of oöcapt, 0.14-0.19 (0.16) × 0.17-0.22 (0.20).
Short fertilisation canal joins vitelline duct immedi-
ately before entry to Mehlis’ gland. Mehlis’ gland
anterior to vitellariun, not visible in most whole-
mounts, subcircular, 90 µm in diameter. Uterine duct
leads dorsally from Mehlis’ gland (Figure 5), then
turns ventrally and runs anteriorly to uterus. Uterus
initially a single transverse cord extending from os-
moregulatory canals, between ovary and vitellarium,
then medially, ventral to testes (Figure 5). Uterus sub-
sequently develops dorsal and ventral branches from
median tube, eventually forming reticulated network
extending across segment, except at distal extremi-
ties where broad tubular extensions develop, extending
to but not beyond osmoregulatory canals (Figure 10).
Subsequent uterine development involves expansion
of tubular network, with most posterior part of uterus
consisting of single transverse tube from which other
reticulations extend; uterus confined within osmoregu-
latory canals. Gravid segments 0.70-1.10 (0.91) × 5.6-
10.3 (8.5). Gravid uterus saccate, with numerous inter-
83
nal subdivisions, filling medulla and extending lateral
and dorsal to osmoregulatory canals (Figure 12). Egg
thick-shelled, generally 4-lobed in shape, with size of
individial lobes varying considerably (Figures 6, 7);
some eggs 3-lobed (Figure 8). External surface of eggs
with filamentous projections arranged in 4, or more
rarely 3 groups, depending upon number of lobes to
egg (Figures 6-8). Oncosphere 15-19 (17)µm in di-
ameter, surrounded by elongate pyriform apparatus,
33-38 (35) × 18-23 (20)µm; distal extremity of pyri-
form apparatus attached to inner envelope by strands
of tissue, sometimes suggesting presence of elongate
horns extending from pyriform apparatus, which in
fact represent foldings of inner envelope.
Osmoregulatory system consists of simple ring an-
terior to suckers (Figure 1) into which paired dorsal
and ventral canals empty; dorsal canals narrower, ex-
ternal to ventral canals, diameter 0.03-0.07 (0.06);
ventral canals medial to dorsal canals, wider, 0.06-
0.16 (0.13) in diameter; transverse canal 0.07 in di-
ameter, connects ventral canals at posterior margin of
each segment. Musculature strongly developed. Lon-
gitudinal musculature consists of 2 layers of fibre
bundles, each with up to 10 fibres per bundle; trans-
verse muscles present in band internal to longitudinal
musculature; dorso-ventral muscles well developed,
individual fibres cross medulla at irregular intervals.
Discussion
Although von Linstow’s description of C. quadrata is
brief, the cestodes described above conform with his
description in most respects. In particular, the eggs are
four-lobed, apparently a characteristic of the species
(von Linstow, 1904). In addition, the specimens agree
with all significant features visible in the type-slides of
C. quadrata in the MNHG. A minor difference which
exists is the description and illustration of a seminal
receptacle by von Linstow (1904). In the new material,
the proximal end of the vagina is variously dilated, but
as the shape of the dilatation is not consistent, it has
not been identified as a seminal receptacle. von Lin-
stow (1904) illustrated (Figure 3) the vagina emptying
into the genital atrium dorsal to the cirrus-sac. This
situation could not be confirmed in the type-slides and
is possibly an error. The position of the ovary was
described by von Linstow (1904) and illustrated (Fig-
ure 3) as dorsal to the vitellarium and Mehlis’ gland.
Spasskii (1951) commented that such an arrangement
was most unlikely and is almost certainly an error.
84

Figures 1-8. Viscachataenia quadrata (von Linstow, 1904) n. g., n. comb. from Lagidium viscacia. 1. Scolex. 2. Histological section through
distal part of cirrus-sac showing vagina anterior to cirrus-sac. 3. Everted cirrus showing peduncle formed by everted genital atrium and lack
of obvious internal seminal vesicle. 4. Genitalia, dorso-ventral view. 5. Transverse histological section through genitalia in mature segment,
dorsal aspect towards top of page. 6-8. Eggs, showing variation in shape and number of lobes of egg shell. Scale-bars: 1-5, 0.1 mm; 6-8,
0.01 mm. Abbreviations: c. cirrus; do, dorsal osmoregulatory canal; isv, internal seminal vesicle; m, Mehlis’ gland; o, ovary; t, testis; u, uterus;
v, vitellarium; va, vagina; vd, vas deferens; vo, ventral osmoregularory canal.
Figures 9-12. Viscachataenia quadrata (von Linstow, 1904) n. g., n. comb. from Lagidium viscacia. 9. Mature segments showing variation in the distribution of testes. 10. Post-mature segment
showing reticulate uterus which forms following the dissolution of the female genitalia; the testes persist. 11. Pre-gravid segment in which the uterus is filling with eggs, losing its reticulate
appearance, but is confined within the osmoregulatory canals. 12. Gravid segment with fully developed uterus obscuring the osmoregulatory canals. Scale-bars: 0.1mm.
85
86
In the new material, these structures were found in
their ususal dorso-ventral relationships. Despite these
differences, which are probably simple errors in von
Linstow’s (1904) account, the cestodes described here
are considered to be C. quadrata.
C. viscaciae is treated tentatively as a synonym of
C. quadrata following comparison of the new speci-
mens with the type-slides of C. viscaciae. Beveridge
(1978) noted that the structure of the cirrus-sac was
similar in the two species, and suggested that they
might be synonyms. However, because of the poor
quality of the type-material of C. quadrata and the
incomplete nature of the type-specimen of C. visca-
ciae, Beveridge (1978) concluded that new material
was required before any judgement could be made on
the matter. The current description agrees with that of
Beveridge (1978) in most respects. Beveridge (1978)
stated that an internal seminal vesicle was absent, but
in the new material it is evident that the vesicle may
be slender or inapparent in some segments, partic-
ularly when the cirrus is everted. Beveridge (1978)
also described a seminal receptacle but, as explained
above, in the new material, the structure is quite vari-
able in shape. The egg of C. viscaciae was described
by Beveridge (1978) as spherical rather than four-
lobed as is the case in C. quadrata, a difference which
is not readily explained, but could be related to the
degree of maturity of the eggs. The eggs of C. visca-
ciae are observable only in utero, and in comparable
eggs of C. quadrata, while the lobed nature of the
egg is evident, the fine, hair-like projections on the
surface of the egg are not visible. Beveridge (1978)
also described the vagina entering the genital atrium
posterior to the cirrus-sac. However, this is shown in
only one figure (Beveridge, 1978, figure 71) in which
the posterior margin of the genital papilla, formed by
evagination of the genital atrium, may have been mis-
taken for the termination of the vagina. In the new
material, the position of the distal vagina is extremely
difficult to see in most segments and is only clearly
visible in a few segments in which the cirrus-sac has
been displaced while removing the outer muscles with
forceps. In sections, the vagina is readily detectable as
a very slender tube anterior to the cirrus-sac (Figure 2).
The difficulty of tracing the distal vagina probably ac-
counts for the conflicting descriptions of von Linstow
(1904) and Beveridge (1978). C. viscaciae is therefore
treated provisionally as a synonym of C. quadrata.
It is possible that the differences in the shape of the
egg indicate the existence of two distinct but very
closely related species of cestodes in Lagidium, but
the evidence at present is not substantive.
Mazza et al. (1932) described a new species of
anoplocephalid cestode, Bertiella finlayi, collected
from L. viscacia ( as L. tucumanus Thomas) in Jujuy
Province, Argentina. They described collecting both
adult and juvenile cestodes. Attempts to locate type-
specimens were unsuccessful. However, examination
of their published figures and photographs suggests
that while the adult specimens they reported were
probably C. quadrata, the ‘juvenile’ specimens were
a related cestode, Monoecocestus threlkeldi (Parra,
1953). Figures 2-4 of Mazza et al. (1932) are of ma-
ture and gravid specimens of a small cestode which
is indistinguishable from the original description of
M. thelkeldi given by Parra (1953). The large cestodes
they reported are of a similar size to C. quadrata, and
although Mazza et al. (1932) gave a description of
the internal morphology of the segments which cor-
responds closely with M. threlkeldi, they did include
drawings and photographs of the eggs, most of which
are four-lobed and have groups of filamentous pro-
jections on the shell. Their figure 9, of a five-lobed
egg, represents a feature not seen in the material de-
scribed above. There seems little doubt therefore that
the large cestodes described by Mazza et al. (1932),
despite of the absence of types, are C. quadrata. For
this reason, B. finlayi has been included as a synonym
of C. quadrata.
The development of the uterus of C. quadrata has
not been described previously. von Linstow (1904) de-
scribed it simply as filling the segment, with dorsal and
ventral diverticula. Beveridge (1978), when describ-
ing C. viscaciae, noted that the uterus was reticulate
and, on these grounds, removed the species from Mos-
govoyia which has a simple tubular uterus (Spasskii,
1951; Beveridge, 1978), placing it tentatively in Cit-
totaenia, the type-species of which, C. denticulata
(Rudolphi, 1904), has a slightly reticulate uterus.
The uterus of C. quadrata forms initially as a sin-
gle transverse cord and subsequently develops dorsal
and ventral branches, producing a complex reticulate
uterus stretching across the segment, with a series of
parallel branches extending laterally at each extremity.
The fully-developed uterus is saccate. This form of
development differs from those described by Spasskii
(1951) and Rausch (1976), but resembles the form of
uterine development found in Paranoplocephala om-
phalodes (Hermann, 1783), as described by Rausch
(1976), in which the uterus forms as a transverse cord-
like aggregation of cells, developing into a transverse

band which becomes fenestrated and eventually fills
the segment as a sac with diverticula. It is likely
that, as uterine development is described in detail in
a greater number of species, more variations on the
currently accepted types of uterine development will
be recorded.
The presence of a reticulate uterus and paired geni-
talia indicates similarities with Cittotaenia, Moniezia,
Blanchard, 1891 and Diandrya Darrah, 1930. How-
ever, C. quadrata differs from all of these genera in
having the vagina enter the genital atrium anterior to
the cirrus-sac, a characteristic observed only in Mo-
noecocestus, a genus which has a single set of genitalia
per segment. The unusual egg of C. quadrata, with
groups of fine hair like projections, has parallels in
species of Monoecocestus with the eggs of M. amer-
icanus (Stiles, 1895), M. thomasi Rausch & Maser,
1977 and M. variabilis (Douthitt, 1915) which also
have their surfaces uniformly covered with fine projec-
tions (Freeman, 1949; Rausch & Maser, 1977). Other
species in which the egg has been described, do not
appear to exhibit this characteristic (Spasskii, 1951;
Rego, 1961).
The egg of C. quadrata has a pyriform apparatus
which is attached at its narrower pole to the inner egg
membrane by a small mass of granular tissue. In whole
eggs, it frequently appears as if the pyriform apparatus
extends at this narrower pole into elongate recurved
horns. However, if the pyriform apparatus is removed
from the egg and can be rotated under the pressure of
a cover-slip, it is apparent that the ‘horns’ are simply
foldings of the inner membrane of the egg. It is impor-
tant therefore that the pyriform apparatus is described
from free eggs some of which can be crushed to ex-
pel internal structures, rather than from whole eggs in
utero.
The unique features of C. quadrata, with paired
genitalia and the vagina entering the genital atrium
anterior to the cirrus-sac suggest that it should be ac-
commodated in a separate genus and hence the new
genus Viscachataenia is proposed. The biogeograph-
ical relationships of the new genus suggest that it is
derived from Monoecocestus, a genus parasitic pri-
marily in South American rodents, by duplication of
the genitalia. The phenomenon of duplication of the
genitalia appears to be common in the Anoplocephali-
nae (Baer, 1927; Spasskii, 1951; Beveridge, 1978).
The unusual features of the eggs of Viscachataenia
and some species of Monoecocestus further support an
association between the genera.
87
The original host of V. quadrata was identified by
von Linstow (1904 ) as Lagidium peruanum, a species
occurring mainly in Peru (Nowak, 1999). The host
of C. viscaciae is uncertain (Beveridge, 1978), but is
probably L. viscacia, which occurs in Chile (including
the area around Valdivia), Peru, Bolivia and Argentina
(Nowak, 1999). The current report confirms the occur-
rence of the species in L. viscacia, although L.v. boxi
is considered by some authors (Rowlands, 1974) to
represent a distinct species. Nowak (1999) followed
Cabrera (1961) in recognising only three species of
Lagidium, L. peruanum, L. viscacia and L. wolffsohni
Thomas, the latter species occurring in the extreme
south of Chile and Argentina. The opinions of Cabrera
(1961) and Nowaks (1999) are followed here, suggest-
ing that V. quadrata occurs in both L. peruanum and
L. viscaciae.
Acknowledgement
We wish to thank Dr C. Vaucher for kindly allowing
us to examine type-specimens.
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