Journal of Helminthology (1979) 53, 133-140

New species of Moaciria Freitas 1956 (Nematoda: Heterakoidea) from Australian Reptiles H. I. JONES Department of Parasitology, University of Queensland, St. Lucia, Brisbane 4067 Queensland, Australia ABSTRACT

Moaciria sphenomorphi n. sp. is described from the skink Sphenomorphus gracilipes, Moaciria butleri n. sp. from the elapid snake Pseudechis australis (both from Western Australia), and Moaciria etnae n. sp. from the python Liasis childreni from Queensland. These species differ from one another and from M. aharengai and M. icosiensis principally in the number and disposition of body and caudal papillae, and from M. freitasi in possessing caudal alae. M. butleri and M. etnae also differ from the other species in possessing a conspicuous three-pointed internal lip tooth, and all three new species possess a small median papilla within the posterior rim of the precloacal sucker.

The genus Moaciria was erected by Freitas (1956) to accommodate M. aharengai. In a revision of the Heterakoidea Inglis (1967) defined the genus in the subfamily Spinicaudinae as being characterised by an anterior cuticular flange projecting slightly anterior to the lips, and in the male narrow caudal alae supported by a few stout papillae, a gubernaculum, and a deep, posteriorly-directed precloacal sucker. Inglis (1957, 1967) placed Spinicauda icosiensis (Seurat 1917) Travassos 1920 in the genus Moaciria, and made it synonymous with Aplectana pharyngeodentata Belle 1957, which from published descriptions had been placed in the genus Bellaplectana by Skrjabin, Schikhobalova and Lagodovskaja (1961). Chabaud and Brygoo (1960) described Spinicauda {Moaciria) freitasi from a number of species of chameleons in Madagascar. There have been no further reports of worms belonging to this genus. The present paper describes three new species which were recovered during a study of the parasites of Australian snakes. MATERIALS AND METHODS All specimens were recovered from snakes in the Western Australian and Queensland Museums which had been fixed in formalin and preserved in 70% alcohol. The worms were cleaned, cleared in chlorolactophenol for examination and stored in 70% alcohol with glycerine. Figures were drawn using a drawing tube attached to a Leitz Wetzlar Dialux Microscope. RESULTS Family Heterakidae Subfamily Spinicaudinae

MOACIRIA SPHENOMORPHI n. sp. (Figs 1-4) Two males recovered from the intestine and rectum of an adult Sphenomorphus gracilipes (Scincidae) which was removed from the stomach of Drysdalia coronata (Elapidae) (Western Australian Museum No. R17119), collected eight kilometers west of Denmark, Western Australia, in October 1962. 133

H. I. JONES

.1

.1

Moaciria sphetwmorphi n. sp. FIG. 1. posterior end male, lateral; FIG. 2. posterior end male, ventral; FIG. 3. anterior end male, dorsal; FIG. 4. posterior oesophageal bulb, lateral.

Description (See Table 1) Small worms, in fair condition, less than five mm long, tapering anteriorly, maximum width about midlength and widening again slightly posteriorly so that the posterior end, before the small spiky tail, appears club-shaped ventrally. Five longitudinal rows of equally spaced papillae, three ventrally and one each laterally, most conspicuous at posterior end but visible throughout the length of the worms. Lateral alae commence near the anterior end and extend posteriorly to the level of the head of the spicules. Three lips, small, not offset from the body; a small flange projects anteriorly from the inner surface of each lip. 134

New species of Moaciria from reptiles

No papillae visible. Excretory pore conspicuous, about two-thirds along length of oesophagus, with posteriorly-directed excretory duct. Nerve ring not seen. Cervical papillae small, anterior to bulb. Pharynx short, leading into narrow oesophagus of constant width which extends posteriorly to a large triangular posterior valved bulb. Poor condition of intestine precluded examination. Posterior end thick, tail short, tapering, finely-tipped, with three pairs of ventral sessile papillae, the anteriormost very large, and one pair of lateral papillae. Precloacal sucker deep, thick-walled, posteriorly-directed, with a small median papilla in the posterior rim, surrounded by narrow alae. Five pairs of small papillae around sucker and cloaca, and three pairs of short, thick, pedunculated papillae anterolateral and lateral to alae. Spicules simple, equal, similar with slight ventral curve and poorly chitinized blunt tips. Gubernaculum large, well-chitinized, thick anteriorly and tapering to blunt tip near cloacal opening. Moaciria sphenomorphi

Host: Habitat: Locality: Type:

Sphenomorphus gracilipes Rectum and intestine near Denmark, South-Western Australia. To be deposited in the Queensland Museum.

MOACIRIA BUTLERI n. sp. (Figs. 5-10) One male and five females collected from the rectum of Pseudechis australis (WAM R28698), collected on Barrow Island, Western Australia, January 1967. Description (See Table 1) Small worms, tapering anteriorly. Lateral alae commence about one-third of way along oesophagus and extend in female to one-third of length of tail and in male to head of spicules. Lateral field cells large and very prominent. Three small lips, not offset from body in male, but offset from body and retractile in female. A small flange extends just anterior to each lip from its inner surface, which is lined from the pharynx. Inside each lip is a conspicuous three-pointed tooth. Two minute papillae on outer surface of each lip. Pharynx short, leading into long narrow oesophagus with large globular valved posterior bulb. Anterior to this bulb is a slight swelling which also appears to be valvular. Intestine at its inception wider than the bulb, without diverticula. Excretory pore about two-thirds of distance along oesophagus; excretory ducts wide and conspicuous extending posteriorly. Cervical papillae and nerve ring not seen. Small papillae regularly spaced in the cuticle of the male in eight longitudinal rows, more conspicuous in the posterior three-quarters of the worm, not seen in females. Male: Tail tapering to a fine point, with four pairs of approximately equally-spaced ventral or ventrolateral papillae and one pair of lateral papillae situated between second and third ventral pair. Precloacal sucker deep, large, chitinized, posteriorly-directed, with a small median papilla in the posterior rim, surrounded by narrow caudal alae commencing just anterior to the sucker and extending past cloaca. Alae meet anteriorly but do not meet posteriorly, supported by six pairs of short, thick pedunculated papillae. At least eight pairs of mostly very small sessile papillae scattered on ventral and lateral aspects of worm at level of sucker and spicules, and one dorsal papilla at level of head of gubernaculum. Spicules equal, similar, simple, with blunt tips. Gubernaculum large, about two-thirds length of spicules, heavily chitinized, club-shaped, tip doubled and extending from cloacal aperture. In addition a chitinized plaque in posterior rectal wall, against which the tips of the spicules abut. 135

* from anterior end

Length Width (maximum) Oesophagus length (without bulb) Oesophagus width Oesophagus bulb length Oesophagus bulb width Pharynx length Excretory pore* Cervical papilla* Commencement of lateral alae* Tail length Spicule length Gubernaculum Sucker diameter Vulva* Eggs 4.95 0.26 0.49 0.03 0.10 0.10 0.05 _ 0.05 0.18 0.15 0.29 0.18 0.04 _ -

4.65 0.24

0.50 0.03 0.13 0.13 0.04 0.45 0.06

0.20 0.13 0.30 0.20 0.04 _ -

Holotype Paratype

Moaciria sphenomorphi

-

0.34 0.33 0.45 0.29 0.08

0.66 0.04 0.16 0.16 0.05 0.66 _

6.37 0.41

_ _ 46% 84 x 56um

0.42 0.63

0.70 0.06 0.18 0.20 — 0.56 _

7.12 0.33

Holotype Allotype

0.76 0.07 0.18 0.20 0.06 0.57 _

0.57 0.05 0.15 0.14 0.05 0.50 0.09

0.77-0.84 0.06-0.07 0.17-0.20 0.20-0.22 — 0.56-0.58 _ 0.42-0.54 0.69-0.80 _ — _ 44%-47% 82 x 54-86 x 58um

0.20 0.10 0.35 0.22 0.06 — -

6.82-7.35 0.43-0.56

6.37 0.38

4.95 0.28

7.50-9.52 0.35-0.49

0.54-0.55 0.49 0.58-0.67 0.59 _ — — _ — 47% 44%-49% 87x59um 84 x 59-92 x 62um

0.80-0.84 0.06-0.07 0.19-0.22 0.21-0.22 0.06-0.07 0.67-0.73 0.09-0.13

Female Paratypes (5)

Female Paratypes (4)

Allotype

Moaciria etnae Holotype

Moaciria butleri

Body measurements of M. sphenomorphi, M. butleri and M. etnae (in mm)

TABLE 1

2

New species of Moaciria from reptiles

Moaciria butleri n. sp. FIG. 5. posterior end, male, lateral; FIG. 6. posterior end, male, ventral; FIG. 7. posterior end, female, lateral; FIG. 8. anterior end, female, lateral; FIG. 9. anterior extremity, female, ventral; FIG. 10. lips, en face. 137

H. I. JONES Female: Vulva slightly protruding, just anterior to mid-length of body. Amphidelphic. Eggs numerous, slightly elongated, smooth-shelled, unembryonated. Tail thin and tapering to a fine point, with two or possibly three pairs of small inconspicuous ventral papillae and one pair of dorsal papillae. Moaciria butleri Host: Pseudechis australis Habitat: Rectum Locality: Barrow Island, North-Western Australia. Types: To be deposited in the Queensland Museum. This species has been named after Mr. Harry Butler, who collected the type host.

MOACIRIA ETNAE n. sp. (Figs. 11-14) One male and twelve females recovered from rectum of Liasis childreni (Queensland Museum No. J26067), collected near Mt. Etna, Queensland (21°42' S, 142°15' E), August 1975. Three females in poor condition, collected from rectum of L. childreni (QM J14231), from Gympie, Queensland, January 1967, are considered to be the same species. Description (See Table 1) Small rather robust worms, tapering anteriorly and posteriorly. Cuticle finely striated transversely, with a few inconspicuous papillae. Maximum width about one-third of distance along body. Narrow lateral alae extend from about one-third of distance along oesophagus to one-third of distance along tail in female, and to head of spicules in male. Three lips, not offset from body in male, but clearly offset and retractile in females. Threepointed tooth at anterior tip of each lip, as in M. butleri. Pharyngeal lining extends onto internal lip surface. Pharynx short, with slight swelling at junction with oesophagus. Oesophagus long and thin, with slight swelling which appears to be valvular followed by a slight constriction immediately anterior to the large globular posterior valvular bulb. This was slightly longer than wide in the male, but slightly wider than long in all females. Intestine wider than oesophagus at its origin. Excretory pore conspicuous, about one-third of distance along oesophagus. Nerve ring not seen. Very small inconspicuous cervical papillae near anterior end. Lateral field cells large and conspicuous. Eight longitudinal rows of equally-spaced papillae in cuticle of male, mainly in posterior two-thirds of the worm. Male: Shorter and narrower than females. Tail slender, short, tapering to a fine point, with three pairs of ventral sessile papillae (the posteriormost two pairs close together), one lateral pair at level of second and third ventral pairs, and two dorsal papillae. Precloacal sucker deep, with chitinized rim, posteriorly directed, with a median papilla in the inner posterior rim, surrounded by short alae which appear to terminate posteriorly at level of cloaca. Alae supported by four pairs of short, stout papillae and two pairs of smaller papillae. Three pairs of ventrolateral sessile papillae anterior to sucker, and one pair of large lateral papillae caudal to sucker. Gubernaculum large, club-shaped, chitinized, extending to cloacal aperture, two-thirds length of spicules. Spicules equal, simple, similar, with blunt tips. Female: Vulva about midlength of worm, flush with body surface and difficult to see. Amphidelphic. Eggs numerous, elongated, smooth, unembryonated. Tail long and tapering, terminating in a fine point, with two pairs of very small ventral and one pair of very small dorsal papillae. 138

New species of Moaciria from reptiles

.1 Moaciria etnae n. sp. FIG. 11. posterior end, male, lateral; FIG. 12. posterior end, male, ventral; FIG. 13. lips, en face; FIG. 14. anterior end, female, ventral.

Moaciria etnae Host: Liasis childreni Habitat: Rectum Locality: near Mt. Etna, Batsville, Queensland, Australia. Types: To be deposited in the Queensland Museum. DISCUSSION The three lips, bulbed oesophagus and male cloacal sucker put these three species in the superfamily Heterakoidea. The offset nature of the lips (in the female), the prominent lateral line cells, relatively long narrow oesophagus and the similar, simple spicules place them in the subfamily Spinicaudinae Travassos 1920. The presence of caudal alae supported by papillae is found only in the genera Moaciria and Strongyluris, but of these two only the former possesses a gubernaculum; this, and the large and deep posteriorly-directed sucker put these worms into the genus Moaciria Freitas 1956. M. sphenomorphi differs from the other two species in being relatively stouter with a relatively smaller tail, in the absence of the tripled internal lip tooth, in the relatively more conspicuous body papillae in five rows round the body, and in the smaller papillae supporting the cloacal alae. M. etnae and M. butleri are very similar to one another. M. etnae 139

H. I. JONES

differs from M. butleri principally in the smaller size and more slender build, in the absence of a posterior cloacal thickening posterior to the gubernaculum, and in the presence of two papillae on the dorsal surface of the male tail. The only perceptible difference between the females of these two species was that the vulva was on a slight protrusion in M. butleri, whereas it was flush with the body wall and difficult to see in M. etnae; this difference may however be an artifact of preservation. M. butleri and M. etnae differ from M. alvarengai Freitas 1956 and M. icosiensis (Seurat, 1917) Inglis 1957 in possessing the tripled tooth on the anterior internal lip surface. All three new species differ from M. alvarengai and M. icosiensis in possessing a small papilla in the ventral rim of the sucker: this structure is also present in M.freitasi. In addition, in M. alvarengai and M. icosiensis the excretory pore is relatively further anterior than in any of these three species. In M. icosiensis the lateral alae extend further along the tail in the female which appears to possess more papillae than these new species. In M. alvarengai the caudal appendage possesses a pair of lateral papillae at its base, the two unpaired papillae on its length, whereas these species all possess four pairs (at least) of ventral or ventrolateral papillae on the tapering tail. M.freitasi (Chaubaud and Brygoo, 1960) has no caudal alae, and has a more slender gubernaculum than these species. M. alvarengai was found in the skink Mabuya maculata in Brazil, M. icosiensis from the skinks Gongylus ocellatus in Algeria and Chalcides delislii, C. sepoides and Scincus officinalis in Egypt (Belle 1957), and M.freitasi from eight species of chameleon, a skink, Mabuya sp., and from a gecko, Uroplatus fimbriatus, in Madagascar (Chabaud and Brygoo, 1960; Brygoo, 1963). The present records thus extend the known geographical range of the genus, and the host range, M. butleri and M. etnae being the only species so far reported from snakes. However, skinks figure largely in the diet of both these species of snake, and it is possible that both these hosts were accidental, though in none of the three infected snakes were there any identifiable food residues. No Moaciria sp. were found in another nine L. childreni examined from Queensland, nor in another eleven Pseudechis australis collected from Western Australia. ACKNOWLEDGEMENTS I would like to thank Dr. G. M. Storr, Curator of Reptiles and Birds at the Western Australian Museum, and Ms. J. Covacavitch, Curator of Reptiles at the Queensland Museum, for allowing me to examine the reptiles in their care and for providing facilities.

REFERENCES BELLE, E. A. (1957) Helminth parasites of reptiles, birds and mammals in Egypt. IV. Four new species of Oxyurid parasites from reptiles. Canadian Journal of Zoology, 35, 163-169. BRYGOO, E. R. (1963) Contribution a la connaissance de la parasitologie des Cameleons malgaches. Annales de Parasitologie Humaine et Comparee, 38, 149-334. CHABAUD, A. G. and BRYGOO, E. R. (1960) Nematodes parasites de Cameleons malgaches. Memoires de I'Institut Scientifique de Madagascar. Ser. A. Biologie Animate, 14, 125-159. FREITAS, J. F. Teixeira de. (1956) Novo parasito de reptil da Ilha Fernando de Noronha: Moaciria alvarengai g. n., sp. n. (Nematoda, Subuluroidea). Revista Brasileira de Biologia, 16, (3), 335-359. INGLIS, W. G. (1957) A review of the nematode superfamily Heterakoidea. Annals and Magazine of Natural History, ser. 12, 10, 905-912. INGLIS, W. G. (1967) The evolution, host relationship and classification of the nematode superfamily Heterakoidea. Bulletin of the British Museum (Natural History), Zoology, 15, (1), 1-28. SEURAT, L. G. (1917) Sur les Oxyures des sauriens du Nord Africain. Archives de Zoologie experimental et generate, 56, 401-444. SKRJABIN, K. I., SCHIKHOBALOVA, N. P. and LAGODOVSKAJA, E. A. (1961) Essentials of Nematology. X. Oxyurata of Animals and Man, Part 2. Moscow Academy of Science: USSR (in Russian). TRAVASSOS, L. (1920) Contribuisoes para o conhecimento da fauna helmintologica brazileira. IX. Sobre as especias do genera Spinicauda, n. g. Memorias do Instituto Oswaldo Cruz, 12, 44-50. Received 29'JDecember, 1977. 140