Article available at http://www.parasite-journal.org or http://dx.doi.org/10.1051/parasite/2003104309
PROGAMOTAENIA
CAPRICORNIENSIS
SP. NOV. ( C E S T O D A : ANOPLOCEPHALIDAE)
F R O M WALLABIES (MARSUPIALIA: M A C R O P O D I D A E ) FROM Q U E E N S L A N D , A U S T R A L I A BEVERIDGE I.* & TURNI C.**
Summary: Progamotaenia capricorniensis sp. nov. (Cestoda: Anoplocephalidae) is described from the wallabies Macropus dorsalis (Gray, 1837] and Petrogale assimilis Ramsay, 1877 from Queensland, Australia. The new species is characterised by a fimbriated velum composed of 26-32 digitiform to triangular projections on each side of the proglottis, paired uteri and 140190 testes distributed in a single band across the medulla. Minor variation occurs in the distribution of the testes. The above characters distinguish the new species from its most closely related congeners P. lagorchestis (Lewis, 1914), P. proterogyna (Fuhrmann, 1932), P. spearei Beveridge, 1980 and P. villosa (Lewis, 1914). P. capricorniensis appears to exhibit a highly disjunct distribution within its usual host, A/1, dorsalis.
Résumé : PROGAMOTAENIA CAPRICORNIENSIS SP. NOV. (CESTODA: ANOPLOCEPHALIDAE) PARASITE DE WALLABIES (MARSUPIALIA: MACROPODIDAE) PROVENANT DU QUEENSLAND, AUSTRALIE
Description de Progamotaenia capricorniensis sp. nov. (Cestoda: Anoplocephalidae), parasite de l'intestin grêle des wallabies Macropus dorsalis (Gray, 1837) et Petrogale assimilis Ramsay, 1877. la nouvelle espèce se distingue par le nombre de projections (26-32) du velum sur chaque côté du proglottis, la possession de deux utéri dans chaque proglottis et la possession de 140-190 testicules distribués dans une bande antérieure de l'utérus. Ces caractères distinguent la nouvelle espèce des espèce voisines : P. lagorchestis (Lewis, 1914), P. proterogyna (fuhrmann, 1933), P. spearei Beveridge, 1980 et P. villosa (Lewis, 1914). La distribution géographique de la nouvelle espèce semble être extrêmement dispersée à l'intérieur de l'aire de répartition de KEY WORDS : Progamotaenia, Cestoda, Anoplocephalidae, new species, l'hôte, M. dorsalis. marsupials, wallabies, Australia. MOTS CLÉS : Progamotaenia, Cestoda, Anoplocephalidae, nouvelle espèce, marsupiaux, wallabies, Australie.
Spratt et al. ( 1 9 9 1 ) and Beveridge et al. ( 1 9 9 8 ) , is des-
INTRODUCTION
T
cribed in this paper and its relationships with c o n g e -
h e anoplocephalid genus Progamotaenia
Nybe-
ners are e x a m i n e d .
lin, 1917 consists o f a n u m b e r o f species o f c e s -
todes which parasitise the small intestine and
bile ducts o f kangaroos, wallabies, rat-kangaroos and wombats (Beveridge, 1976, 1978, 1980, 1985; Beveridge & T h o m p s o n , 1979; Turni & Smales, 1 9 9 9 ) . Currently, 20 species are recognised within the g e n u s (Table I ) , although additional u n d e s c r i b e d species have
been
reported (Spratt et al., 1 9 9 1 ) . In addition, allozyme studies ( B a v e r s t o c k et al., least o n e
1 9 8 5 ) have suggested that at
currently recognised
s p e c i e s , P.
festiva
(Rudolphi, 1 8 1 7 ) , consists o f at least ten sibling species raising the possibility that other s p e c i e s
com-
p l e x e s exist within the genus. As part o f a revision o f the genus, the u n n a m e d species from the blackstripe wallaby, Macropus
dorsalis
(Gray, 1 8 3 7 ) , reported b y
MATERIALS AND METHODS
S
pecimens collected by the authors were relaxed in water and fixed in either 10 % formalin or AFA (Pritchard & Kruse, 1982). Cestodes were stained in car-
mine or celestine blue, dehydrated in ethanol, cleared in clove oil or methyl salicylate and mounted in Canada
balsam. After clearing, the tegument and longitudinal muscles o f most specimens were removed using fine forceps. Some proglottides were sectioned by hand using a scalpel blade to provide cross sections of the proglottis. Additional proglottides were e m b e d d e d in paraffin, sectioned at a thickness o f 5 µm and the sections were stained with haematoxylin and eosin. Eggs were expelled from gravid proglottides and examined in water before
* Department of Veterinary Science, University of Melbourne, Veterinary Clinical Centre, Werribee, Victoria, 3030, Australia. ** Department of Microbiology and Parasitology, University of Queensland, St Lucia, Queensland. 4072, Australia. Correspondence: Ian Beveridge. Tel.: 61 3 97312285 - Fax 61 3 97312263. E-mail:
[email protected] Parasite, 2003, 10. 309-315
clearing in lactophenol. Drawings w e r e made with the aid o f a drawing tube attached to an Olympus BH-2 microscope. Measurements w e r e made with a ruler or with an ocular micrometer. All measurements are presented in millimetres as the mean followed, in parentheses, by the range and number measurements.
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BEVERIDGE I. & TURNI C.
Host genera
Species name abietiformis Turni & Smales, 1999 aepyprymni Beveridge, 1976 bancrofti (Johnston, 1912) capricorniensis sp. n. diaphana (Zschokke, 1907) dorcopsis Beveridge, 1985 effigia Beveridge, 1976 ewersi (Schmidt, 1975) festiva (Rudolphi, 1817)
gynandrolinearis Beveridge & Thompson, 1979 johnsoni Beveridge, 1980 lagorchestis (Lewis, 1914) macropodis Beveridge, 1976 proterogyna (Fuhrmann, 1932) queenslandensis Beveridge, 1985 ruficola Beveridge, 1978 spearei Beveridge, 1980 thylogale Beveridge & Thompson, 1979 villosa (Lewis, 1914) wallabiae Beveridge, 1985 zschokkei (Janicki, 1906)
Velum: straight (s) or laciniate (no/ t
Onycbogalea Aepyprymnus Onycbogalea Macropus Lasiorhinus Dorcopsis Macropus Macropus, Wallabia Macropus, Wallabia, Onycbogalea, Setonix, Vombatus, Lagorchestes Lagorchestes Lagorchestes Lagorchestes Macropus, Wallabia Macropus Thylogale Macropus Thylogale Thylogale Lagorchestes Dorcopsis Macropus, Onycbogalea, Lagorchestes. Dendrolagus
12-16
S S 26-32 s 13-17 S S s
s s 20-35 S 25 17-20 s 25-35 15-18 24-30 s 12
No of ut
Testis groups
Testis no.
2 1 2 1(2)* 2 (1) 2 1 (2)
11-13 35-46 c. 200 142-191 39-64 34-50 67-96 102-140 22-65
2 2
c. 50 100-190 70-90 80-149 65-80 43-50 114-140 30-40 60-100 20-28 c. 100 60-80
2 2 2 2
1 1
(2)
1 2 2 1, 2 1 2 (1) 2 (1) 2 2 2 2 1
1
2 1 1
1 1 2
1 1 1
2 1 1 1
1
* Figures in parentheses indicate less common arrangements of testis distribution. Table I. - Species of Progamotaenia
with host genera and principal morphological features.
S p e c i m e n s have b e e n deposited in the South Austra-
Additional material e x a m i n e d : from Macropus
lian Museum, Adelaide, South Australia (SAM), the
two specimens, Rossmoya, Q u e e n s l a n d (23° 0 1 ' S, 150°
dorsalis-.
Muséum National d'Histoire Naturelle, Paris (MNHN)
29' E ) , coll. I. Beveridge, 2 6 . X . 2 0 0 0 , SAM 2 8 4 7 9 ; two
and the Natural History Museum, London ( B M N H ) .
specimens, Milman, Q u e e n s l a n d (23° 0 8 ' S, 150° 24', E), coll. I. Beveridge, 26.ix. 1977, SAM 28478, serial sections SAM 2 8 4 8 0 , spirit material SAM 2 2 8 0 ) . From
RESULTS
assimilis
ynonymy: "Progamotaenia sp. n." of Spratt et al., 1991, p. 4 9 ; Beveridge et al. Macropus
landensis
Petrogale
1877: two specimens, "The
Pin-
nacles", 5 k m south o f Charters T o w e r s , Q u e e n s l a n d
PROGAMOTAENIA CAPRICORNIENSIS SP. N O V . (Figs 1-11)
S
Ramsay,
dorsalis).
1998, p. 4 7 9 (host:
Progamotaenia
cf.
queens-
Beveridge, 1985 o f Beveridge et al. ( 1 9 8 9 ) ,
p. 2 7 6 (host: Petrogale Type-host: Macropus Other hosts: Petrogale
assimilis). dorsalis assimilis
(20° 0 8 ' S, 145° 16' E ) , coll. R . Close, 15.viii.1982, SAM 2 2 0 8 5 (slides), 11937 (spirit material).
DESCRIPTION Large cestodes, 242 ( 1 9 9 - 2 6 5 , n = 8 ) long, m a x i m u m width 7.1 ( 6 . 0 - 9 . 0 , n = 10), c o m p o s e d o f c. 5 6 0 proglottides. S c o l e x lacking prominent l o b e s (Fig. 1), 1.01
(Gray, 1 8 3 7 ) .
( 0 . 8 6 - 1 . 1 1 , n = 10) in diameter; suckers ovoid, 0.40
Ramsay, 1877.
( 0 . 3 2 - 0 . 4 5 , n = 10) long, 0.34 ( 0 . 2 9 - 0 . 3 6 , n = 10) w i d e .
Site in host: small intestine.
Neck absent in incompletely relaxed s p e c i m e n s , 1.10
Type-locality: T a u n t o n National Park, Dingo, Q u e e n s -
( 0 . 4 4 - 1 . 7 3 , n = 3 ) long in relaxed s p e c i m e n s . Proglot-
land, Australia (23° 3 3 ' S, 149° 13' E ) .
tides greatly e x t e n d e d transversely, with b r o a d velum
T y p e material: holotype, coll. C. Turni, 12.viii. 1996,
forming 29 ( 2 6 - 3 2 , n = 10) digitiform l o b e s o n e a c h o f
SAM 2 8 4 7 5 ; paratypes: s a m e data, three s p e c i m e n s ,
dorsal and ventral aspects o f proglottis; l o b e s e x t e n -
SAM 2 8 4 7 6 ; serial sections, SAM 2 8 4 8 1 ; o n e s p e c i m e n
ding over 1/2 to 3/4 o f s u c c e e d i n g proglottis (Figs 7-
BMNH, 2 0 0 3 . 5 . 6 . 6 ; three s p e c i m e n s , coll. C. Turni,
11). Premature proglottides 0.42 ( 0 . 3 7 - 0 . 5 2 , n = 1 0 )
28.ii.1996, 15.vi.1996, 24.x. 1996, SAM 28477, 32146; o n e
long, 4 . 7 9 ( 4 . 2 1 - 5 . 6 2 , n = 10) wide; length o f velum
specimen, coll. C. Turni, 15.vi.1996, MNHN 63 HG CIX,
0.32 (0.22-0.44, n = 10). Mature proglottides 0.48 (0.37-
slides 184-186.
0.59, n = 10) long, 5.80 (5.00-6.55, n = 10) wide; length
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N E WSPECIESOF CESTODES FROM »
WALLABIES
Figs 1-6. - Progamotaenia capricorniensis sp. nov. from Macropus dorsalis (Marsupialia). Fig. 1. Scolex, dorsoventral view. Fig. 2. Egg. Fig. 3. Lateral margin of premature proglottis showing circular genital atrium not opening to exterior and vagina extending from seminal receptacle to genital atrium. Fig. 4. Lateral margin of mature proglottis showing patent genital atrium and cirrus sac with sperm in internal seminal vesicle. Fig. 5. Lateral margin of postmature proglottis showing uterus starting to extend across osmoregulatory canals towards posterolateral margin of proglottis and vagina reduced to blind tube extending laterally from seminal receptacle but scarcely reaching osmoregulatory canals. Fig. 6. Transverse histological section of proglottis showing musculature and genital ducts passing osmoregulatory canals dorsally. Scale-lines: Figs 1, 3-6, 0.1 mm; Fig. 2, 0.01 mm. Abbreviations: AO, accessory osmoregulatory canal; DO, dorsal osmoregulatory canal; ES, external seminal vesicle; GA, genital atrium; IS, internal seminal vesicle; LM, inner longitudinal muscles; M, Mehlis' gland; O, ovary; SR, seminal receptacle; TM, transverse muscle; U, uterus; V, vagina; VI, vitellarium; VO, ventral osmoregulatory canal. Parasite, 2003, 10, 309-315
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Figs 7-11. - Progamotaenia capricorniensis sp. nov. from Macropus dorsalis, showing sequential development of genitalia. Fig. 7. Premature proglottis prior to patency of genital atria. Fig. 8. Premature proglottides with patent genital atria but with testes distributed either in a single row in middle of proglottis or divided into two groups. Fig. 9. Mature proglottis with extranumerary vestigial vitellaria medial to principal vitellarium. Fig. 10. Postmature proglottis showing uterus extending posterior to cirrus sac but with remnants of testes still visible. Fig. 11. Gravid proglottis. Scale- lines: 0.1 mm. Abbreviations: A. accessory, vestigial vitellaria.
o f velum 0.41 ( 0 . 2 6 - 0 . 5 6 , n = 10). Postmature proglot-
Longitudinal osmoregulatory canals paired. In s c o l e x ,
tides 0.53 ( 0 . 4 7 - 0 . 5 9 , n = 10) long, 6.66 ( 6 . 1 6 - 7 . 2 5 , n =
dorsal and ventral canals o f e a c h side fuse at level o f
10) wide; length o f velum 0.44 ( 0 . 2 0 - 0 . 5 5 , n = 1 0 ) .
anterior part o f suckers; single canal joins c o m m o n
Gravid proglottides 0.66 ( 0 . 4 8 - 0 . 9 4 , n = 10) long, 5.81
canals from e a c h side o f strobila near anterior extre-
( 4 . 9 1 - 7 . 1 8 , n = 10) wide; length o f velum 0.55 (0.37-
mity o f s c o l e x (Fig. 1). In strobila, dorsal canal external
0.86, n = 1 0 ) .
to ventral canal (Fig. 5 ) . Dorsal canal, narrow, sinuous,
Inner longitudinal musculature well d e v e l o p e d , c o m -
0.045 ( 0 . 0 3 9 - 0 . 0 6 2 , n = 10) in diameter; ventral canal
p o s e d o f large bundles; b u n d l e s circular in cross s e c -
broader, almost straight, 0.23 (0.16-0.37, n = 10) in dia-
tion, e a c h c o m p o s e d on n u m e r o u s individual fibres;
meter. At posterior margin o f e a c h proglottis, ventral
bundles closer to medulla larger, c o m p o s e d o f greater
but not dorsal canals joined b y single transverse canal,
n u m b e r o f individual fibres than bundles furthest from
0.04 ( 0 . 0 3 - 0 . 0 5 , n = 10) in diameter. Tiny a c c e s s o r y
medulla (Fig. 6 ) ; outer longitudinal muscles consist o f
osmoregulatory canals present, medial and either dorsal
individual fibres. Transverse muscles well d e v e l o p e d ,
or ventral to ventral canal, 0 . 0 1 5 in diameter; a c c e s -
forming b a n d i m m e d i a t e l y internal to
longitudinal
sory canals occasionally have transverse canals joining
muscles. Dorsoventral muscles arranged as individual
those from b o t h sides. Genital ducts cross o s m o r e g u -
fibres; most prominent in cortex, separating rows o n
latory canals dorsally (Fig. 6 ) .
longitudinal muscle bundles; few, scattered dorsoven-
Genitalia paired. Genital atrium shallow, o p e n i n g in
tral muscle fibres cross medulla.
middle o f lateral proglottis margin. Genital atrium in
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NEW SPECIES OF CESTODES FROM WALLABIES
premature proglottides circular in dorsoventral views, n o n - p a t e n t (Fig. 3 ) . In mature proglottides,
atrium
receptacle, terminating near medial margin o f ventral o s m o r e g u l a t o r y canal, anterior to external
seminal
patent, consisting o f shallow depression with few folds
vesicle (Fig. 8 ) . Uteri subsequently d e v e l o p both ante-
in margin (Fig. 4 ) . Cirrus sac well developed, with thick
rior and posterior diverticula while paired digitiform
muscular wall, extending across osmoregulatory canals,
outgrowths cross osmoregulatory canals dorsally and
entering medulla, 0.68 ( 0 . 5 6 - 0 . 7 6 , n = 10) long, 0.16
e x t e n d to posterolateral aspect o f proglottis, posterior
( 0 . 1 3 - 0 . 1 8 , n = 10) wide in mature proglottides (Figs 4,
to cirrus sac (Figs 10, 11). Eggs ovoid with rough shells,
5). Distal region o f withdrawn cirrus sinuous or straight,
0.035 (0.031-0.039, n = 10) long, 0.027 (0.025-0.027, n =
a r m e d with small s p i n e s ; p r o x i m a l
region
coiled,
unarmed; cirrus surrounded b y masses o f deeply stai-
10) wide; o n c o s p h e r e 0 . 0 1 8 ( 0 . 0 1 4 - 0 . 0 2 0 , n = 10) in diameter; pyriform apparatus absent (Fig. 2 ) .
ning glandular cells. Internal seminal vesicle prominent, usually but not invariably divided into two sections (FIgs 4, 5 ) ; distal section 0.20 ( 0 . 1 2 - 0 . 3 1 , n = 10) long, 0.11 ( 0 . 0 9 - 0 . 1 3 , n = 10) wide; proximal section 0.18 (0.13-0.27, n = 10) long, 0.11 ( 0.07-0.14, n = 10) wide. External seminal vesicle fusiform, c o v e r e d with single layer o f glandular cells. 0.37 ( 0 . 2 7 - 0 . 5 9 , n = 10) long, 0.07 (0.04-0.12, n = 1 0 ) wide, extending from proximal p o l e o f cirrus sac to midway anterior to seminal receptacle (Fig. 3 ) . Testes spherical, 0.07 ( 0 . 0 5 - 0 . 0 9 , n = 10) in diameter, extending in b r o a d b a n d anterior to uteri from median border o f ventral osmoregulatory canals (Fig. 7 ) ; b a n d 1-4 testes b r o a d anteroposteriorly, deep
dorsoventrally;
testes more
densely
1-3
packed
towards lateral extremities o f medulla; testes n u m b e r 170 ( 1 4 2 - 1 9 1 , n = 10); in small n u m b e r o f proglottides, b a n d o f testes thins in middle to single row (Fig. 8 ) ; in very small n u m b e r o f proglottides, row b r o k e n in middle o f medulla with testes forming two groups (Fig. 8 ) ; all patterns visible within individual c e s t o d e ; n o individual has large n u m b e r o f proglottides with testes divided into two groups.
DISCUSSION
T
h e species o f Progamotaenia
described above
is characterised b y paired uteri, a fimbriated or
laciniate velum with 26-32 digitiform projections
and testes usually distributed in a single b a n d anterior to the uteri, numbering 1 4 0 - 1 9 0 . It is therefore differentiated from all c o n g e n e r s since those species with two uteri and a velum c o m p o s e d o f large numbers o f posteriorly directed digitiform p r o c e s s e s (P. chestis
(Lewis, 1 9 1 4 ) , P. spearei
lagor-
Beveridge, 1980, P. vil-
losa (Lewis, 1 9 1 4 ) ) have testes distributed in two distinct g r o u p s a n d n u m b e r i n g
20-90 per
proglottis
( T a b l e I ) . O f the species with a fimbriated velum, two uteri and testes distributed in a single b a n d , P. proterogyna
only
(Fuhrmann, 1 9 3 2 ) resembles the spe-
cies described a b o v e , but it has only 6 5 - 8 0 testes per proglottis c o m p a r e d with 140-190 in the species described. S o m e variation occurs within the genus in the sepa-
Vagina narrow tube, 0.02 ( 0 . 0 1 - 0 . 0 5 , n = 10) in dia-
ration o f the testes into two lateral groups. Most spe-
meter, o p e n s to genital atrium posterior to cirrus sac,
cies are clearly separable into those with two distinct
lacking surrounding investment of glandular cells (Figs 3,
groups and those with a single continuous b a n d o f
4 ) ; runs posterior to cirrus sac to ovoid seminal r e c e p -
testes. However, s o m e variation d o e s o c c u r within
tacle, 0.38 (0.25-0.57, n = 10) long, 0.22 ( 0 . 1 4 - 0 . 2 9 , n =
P. diaphana
10) wide. Distal vagina clearly visible in premature pro-
1817), P. macropodis
glottides, prior to patency o f genital atrium (Fig. 3 ) ; fol-
densis
( Z s c h o k k e , 1 9 0 7 ) , P. festiva
Beveridge,
(Rudolphi,
Beveridge, 1976, P. 1985 and P. ruficola
queenslanBeveridge,
lowing patency o f atrium, distal vagina atrophies; in
1978, the significance of which was discussed by B e v e -
post-mature proglottides vagina consists o f short duct
ridge ( 1 9 8 5 ) . In the species described above, the slight
extending from seminal receptacle to osmoregulatory canals but not b e y o n d (Fig. 6 ) . Ovary flabelliform, on medial margin o f seminal receptacle, 0.56 ( 0 . 4 8 - 0 . 6 2 , n = 10) long, 0.27 ( 0 . 1 9 - 0 . 3 9 , n = 10) wide, c o m p o s e d o f n u m e r o u s clavate lobules, on ventral aspect
of
medulla (Fig. 4 ) . Vitellarium c o m p a c t , U-shaped, posterior and dorsal to ovary, 0.33 (0.27-0.36, n = 10) long, 0.15 ( 0 . 1 1 - 0 . 1 9 , n = 10) wide. Mehlis' gland spherical, dorsal to and e n c l o s e d by recurrent arm o f vitellarium, visible in f e w proglottides only, 0.14 (0.10-0.17, n = 5) by 0.09 ( 0 . 0 6 - 0 . 1 2 , n = 5 ) . Supranumerary, vestigial vitellaria medial to principal vitellarium in s o m e proglottides (Fig. 9 ) .
separation o f testes into two groups occurs in a f e w proglottides only and in these cases, the majority o f proglottides in the s a m e strobila exhibit testes distributed in a single b a n d . Consequently, P. niensis
capricor-
is considered to have the testes distributed in
a single b a n d in spite o f the fact that in a small n u m b e r o f proglottides, there may b e a slight break in testis distribution in the centre o f the proglottis. T h e species described here therefore clearly differs from all previously described species and is considered to b e n e w . T h e n a m e is given in recognition o f the fact that most collections have b e e n m a d e in the Capricornia region o f central Q u e e n s l a n d .
Uteri paired, transverse, tubiform, e x t e n d from near
Several aspects o f the m o r p h o l o g y o f the n e w species
centre o f proglottis, anterior to ovary and
warrant c o m m e n t . T h e
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in
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BEVERIDGE I. & TURNI C.
many, but not all specimens, appears to consist o f t w o dilatations. T h e proximal dilatation, the true internal seminal vesicle, is invariably present, while in most proglottides the proximal region o f the cirrus b e c o m e s extremely dilated s o as to form an additional c o m p o nent to the internal vesicle. As it is not invariably present, it has n o t b e e n included in t h e characters defining t h e species. It d o e s , however, provide a useful character in separating most s p e c i m e n s o f P. capricorniensis from c o n g e n e r s .
( D i n g o ) . East o f Rockhampton, a significant n u m b e r o f s p e c i m e n s o f M. dorsalis has b e e n e x a m i n e d ( B e v e ridge et al., 1 9 9 8 ) without finding P. capricorniensis. Consequently, the distribution o f this cestode is thought to b e highly disjunct. This hypothesis is supported b y the finding o f a single infection in the rock wallaby Petrogale assimilis. Beveridge et al. ( 1 9 8 9 ) e x a m i n e d 3 6 rock wallabies o f this species, finding infection o f P. capricorniensis (reported as "P. cf. queenslandensis" in a single s p e c i m e n only, collected near Charters T o w e r s , Q u e e n s l a n d . Additional examination o f s p e cimens o f P. assimilis have failed to reveal e x a m p l e s of this c e s t o d e (unpublished observations) supporting the original observations that the species is u n c o m m o n in this host. Equally interesting is the fact that a substantial proportion o f the sample o f M. dorsalis examined b y Beveridge et al. ( 1 9 9 8 ) w a s from t h e Charters T o w e r s region, but n o P. capricorniensis were found. T h e s e data further support the hypothesis that the distribution o f the c e s t o d e is highly disjunct.
A pyriform apparatus w a s not o b s e r v e d in eggs from the specimens examined. It is possible that the cestodes described w e r e not fully gravid, though this is highly unlikely given the n u m b e r o f large s p e c i m e n s o f the species available. A pyriform apparatus is also absent in P. bancrofti (Johnston, 1912), P. lagorchestis, P. queenslandensis and P. wallabiae Beveridge, 1985 (Beveridge & T h o m p s o n , 1979; Beveridge, 1980, 1 9 8 5 ) . However, Beveridge ( 1 9 7 6 ) o b s e r v e d that in P. ewersi (Schmidt, 1 9 7 5 ) , a pyriform appparatus m a y d e v e l o p only after the proglottides have b e e n shed. T h e s a m e m a y apply to the species noted, as well as to P. capricorniensis. T h e o c c u r r e n c e o f vaginal atrophy is a characteristic ACKNOWLEDGEMENTS o f m a n y species Progamotaenia, with the temporal relationships o f insemination and atrophy o f the vagina This w o r k w a s supported financially b y the Ausbeing o f specific value (Beveridge, 1 9 7 6 ) . In P. capritralian Biological Resources Study. T h a n k s are corniensis, the precise s e q u e n c e o f relationships could due to Dr L. Warner for assistance in collecting not b e determined. In premature proglottides, the full wallabies, to Dr D.M. Spratt for c o m m e n t s o n a draft extent o f the vagina from the seminal receptacle to the of the manuscript and two anonymous referees for suggenital atrium w a s clearly observable. In these progested improvements to t h e submitted manuscript. glottides, t h e genital atrium w a s subcircular in dorsoventral view (Fig. 3 ) but did not o p e n to the exterior. In subsequent proglottides, in which the genital atrium REFERENCES o p e n e d to t h e exterior and consisted o f folded walls, the vagina could not b e traced to the atrium, suggesBAVERSTOCK P.R., ADAMS M. & BEVERIDGE I. Biochemical difting that insemination occurs prior to the genital atrium ferentiation in bile duct cestodes and their marsupial o p e n i n g to the exterior. This observation n e e d s to b e hosts. Molecular Biology and Evolution, 1 9 8 5 , 2, 3 2 1 - 3 3 7 . e x a m i n e d in detail using longitudinal serial sections o f BEVERIDGE I. A taxonomic revision of the Anoplocephalidae the anterior regions o f cestodes. If confirmed, it repre(Cestoda: Cyclophyllidea) of Australian marsupials. Aussents a novel form o f development within the genus. tralian Journal of Zoology, Supplementary Series, 1 9 7 6 , 44, In most s p e c i e s o f Progamotaenia, t w o transverse 1-110. osmoregulatory canals are present (Beveridge, 1 9 7 6 , BEVERIDGE I. Progamotaenia ruficola sp. n. (Cestoda: Ano1978, 1980, 1 9 8 5 ) . In P. capricorniensis, a single transplocephalidae) from the red kangaroo, Macropus rufus verse canal c o n n e c t s left a n d right ventral canals in (Marsupialia). Journal of Parasitology, 1 9 7 8 , 64, 2 7 3 - 2 7 6 . e a c h proglottis. In this respect, P. capricorniensis BEVERIDGE I. Progamotaenia Nybelin (Cestoda: Anoplocepharesembles P. ewersi (Schmidt, 1 9 7 5 ) , P. gynandrolilidae): new species, redescriptions and new host records. nearis Beveridge & Thompson, 1979, P. johnsoni BeveTransactions of the Royal Society of South Australia, 1 9 8 0 , ridge, 1980, P. lagorchestis a n d P. ruficola. 104, 6 7 - 7 9 . Beveridge et al. ( 1 9 9 8 ) reported helminths from 3 9 BEVERIDGE I. Three new species of Progamotaenia (Cestoda: M. dorsalis from central a n d northern Q u e e n s l a n d , Anoplocephalidae) from Australasian marsupials. Systemawith P. capricorniensis (as "Progamotaenia sp. n . " ) tic Parasitology, 1 9 8 5 , 7, 9 1 - 1 0 2 . occurring at only a single locality, Milman north o f BEVERIDGE I., CHILTON N.B., JOHNSON P.M., SMALES L.R., SPEARE R. R o c k h a m p t o n . S u b s e q u e n t collecting ( u n p u b l i s h e d & SPRATT D.M. Helminth communities of kangaroos and observations) has supported this observation, with the wallabies (Macropus spp. and Wallabia bicolor) from c e s t o d e b e i n g found in M. dorsalis only to the north north and central Queensland. Australian Journal of Zooof R o c k h a m p t o n (Milman, R o s s m o y a ) and to the west logy, 1 9 9 8 , 46, 4 7 3 - 4 9 5 . 314
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