Breinlia (Johnstonema) annulipapillata (Johnston & Mawson, 1938) Chabaud & Bain, 1976

Spratt, David M., 2011, New records of filarioid nematodes (Nematoda: Filarioidea) parasitic in Australasian monotremes, marsupials and murids, with descriptions of nine new species 2860, Zootaxa 2860 (1), pp. 1-61 : 41-43

Breinlia (Johnstonema) annulipapillata (Johnston & Mawson, 1938) Chabaud & Bain, 1976

Dipetalonema annulipapillatum Johnston & Mawson, 1938a, pp117–118 , figs 26–29 ( Onychogalea fraenata , Macropus dorsalis ) Burnett R., Queensland; Johnston & Mawson 1938b, pp. 189–190, figs. 6–7 ( Macropus agilis ) (host identified in Johnston & Mawson, 1939a, p. 122).

Johnstonema annulipapillatum (Johnston & Mawson) Yeh, 1957, p. 202 ; Mackerras 1962, pp. 427–428, ( Macropus dorsalis (= Protemnodon dorsalis ); Spratt & Varughese 1975, pp. 12–14.

Breinlia annulipapillata (Johnston & Mawson, 1938) Beveridge, Speare, Johnson & Spratt 1992, p.367 ( Onychogalea unguifera ) (spelling emendation)

Breinlia (Johnstonema) annulipapillata (Johnston & Mawson) Chabaud & Bain, 1976 ; Beveridge et al. 1985, p. 379; Spratt et al. 1991, pp. 44, 49–50, 56, 61, 69.

Johnstonema andersoni Spratt & Varughese, 1975, pp. 14–17 , figs. 1–12, ( Macropus rufus (syn. Megaleia rufa )) Quilpie, Queensland., ( Macropus robustus , M. giganteus , Wallabia bicolor ) new synonymy.

Breinlia (Johnstonema) andersoni (Spratt & Varughese) Chabaud & Bain, 1976, p. 377 ; Spratt et al. 1991, pp. 48, 53, 58–59, 61, 69.

Type host. Onychogalea fraenata (Marsupialia: Macropodidae )

Other hosts. Macropus agilis , Macropus antilopinus , Macropus dorsalis (Gray) , Macropus giganteus , Macropus parryi , Macropus robustus erubescens , Macropus robustus robustus , Macropus robustus woodwardi , Macropus rufus , Onychogalea unguifera , Wallabia bicolor (Marsupialia: Macropodidae ).

Site in host. Subcutaneous connective tissue.

Other material examined. From Onychogalea fraenata: QLD : 1♀, (no N #) Dingo .

From Onychogalea unguifera: QLD : 2♂, 3♀, ( AHC 45883) , 1♀, ( N1022 ) , 1♂, 9♀, ( QM G232534 ) ; 1♀, ( N1035 ) , 2♀, ( N1037 ) Wernadinga Station ; 2♀, ( N1016 ) Chadshunt Station .

From Macropus agilis: QLD : 2♀, ( N5124 ), 3♀, ( N5220 ). Townsville .

From Macropus antilopinus: QLD : 2♀, ( N1037 ) Burlington Station .

From Macropus dorsalis: QLD : 9♂, 16♀, ( AHC 45884; 1♀, ( N4558 ) Rockhampton.

From Macropus parryi: QLD : 1♀, ( N588 ) Inkerman Station ; 3♀, ( N699 ) Townsville Crematorium .

From Macropus robustus erubescens: QLD : 1♂, 6♀, ( N33 ) Charleville. WA: 10♂ 34♀, ( N5475 ) 6 km N, 1♂, ( N5474 ), 45 km N Fortescue River Roadhouse, Pilbara.

From Macropus robustus robustus: QLD : 3♂, 5♀, ( QM G232535 ) “Warrawee” via Charters Towers ;

From Macropus robustus woodwardi Thomas, 1901 : NT: 1♀, ( N5402 ) Stuart Highway 40 km N Katherine. WA: 4♀, ( AHC 45882) Kimberley province .

From Macropus rufus: QLD : 3♂, 6♀, ( AHC 45881) , 1♂ ( AHC 45890) Adavale ; 3♀, ( N32 ) Richmond. WA : 1♀, ( N5478 ), 10 km N , 4♀, ( N5472 ) 22 km N Fortescue River Roadhouse, Pilbara.

From Wallabia bicolor: QLD : 1♀ fragment, ( N3157 ) Rockhampton. NSW: 1♂ ( UAZ, HC3032 ) Milson Is. VIC: 1♀ ( SAM 10198) Warby Range .

Distribution and hosts. Breinlia (Johnstonema) annulipapillata like B. (B.) spelaea is, with two exceptions, restricted to the north of the continent and is known from a number of macropodid species in northern Queensland, the Northern Territory and the Kimberley and Pilbara regions of Western Australia. There is a single male specimen recorded from W. bicolor from Milson Is. New South Wales in the collection of the Zoology Department, University of Adelaide ( Spratt & Varughese 1975) and a single female from Warby Range Victoria identified by the author.

Remarks. Johnston and Mawson (1938a) originally described this species from male worms recovered from the subcutaneous tissue of Onychogalea fraenata and from the knee region of Macropus dorsalis from Burnett River, and the coelome of Wallabia bicolor from coastal New South Wales (locality mentioned only in Johnston and Mawson (1939b), p. 189 and probably referring to Milson Is.). They later doubtfully assigned two immature worms, recovered from the dorsal aorta of Macropus sp. (probably M. agilis ) form the Gulf of Carpenteria, to this species ( Johnston & Mawson 1938b). On the basis of the smooth cuticle and stout, subequal spicules, Yeh (1957) erected but did not define the genus Johnstonema (Onchocercinae) for Dipetalonema annulipapillata . Chabaud and Anderson (1959) transferred Johnstonema to the subfamily Splendidofilariinae, Mackerras (1962) suggested that the immature female worms of Johnston and Mawson (1938b) probably belonged to a species of Breinlia . Chabaud and Bain (1976) placed the genus Johnstonema as a subgenus of Breinlia Yorke & Maplestone, 1926 .

Spratt and Varughese (1975) examined the damaged holotype (right spicule missing, body broken and badly discoloured) of B. (J.) annulipapillata and reported that paratypes could not be located. They also noted that the labelling of the material from Macropus sp. as allotype was in error. They described and illustrated B. (J.) andersoni sp. nov. indicating that it differed from B. (J.) annulipapillata by the exceptional complexity of the spicular morphology, the presence of a cervical dilatation and the presence of caudal alae in the male. Having now examined many more specimens from a range of host species including all of those previously reported under both species names I conclude that B (J.) andersoni is in fact a synonym of B. (J.) annulipapillata and that the previous characters thought to differentiate the two species are erroneous and based on the poor state of the holotype and absence of an allotype. The characteristic features of B. (J.) annulipapillata are those noted for B. (J.) andersoni by Spratt and Varughese (1975) and in particular the complex morphology of the robust spicules, bifurcate tip of the right spicule and three, large, fleshy papillae on the tail tip of males and females, two lateral and one median. In addition, there is generally a cervical dilatation in both sexes, the maximum width of males and particularly females frequently occurs in the anterior one fifth to one third of the body, in females often at or near the vulva, and the male tail is invariably helically coiled.

Just as Breinlia (B.) spelaea was the most morphometrically variable species of the subgenus Breinlia , so too Breinlia (J.) annulipapillata is the most morphometrically variable species of the subgenus Johnstonema both between genera and species of hosts and within a single host individual. Greatest variation occurs in length and width of body, length of muscular and especially glandular portions of oesophagus and length of tail in males and females, size of spicules in males and distance of vulva from anterior end in females. Spicule ratio (1: 1.1) is con- sistent across host species with the exception of M. dorsalis (1:1.4). Breinlia (J.) annulipapillata is distinguished from the only other species known in the subgenus, B. (J.) woerlei , by the presence of three conspicuous fleshy tail papillae, absence of longitudinally elongate, refractile cuticular bosses in both sexes, much shorter length of males, complexity and much smaller size of spicules and restriction of cloacal papillae to the cloacal region.

All of the material examined has come from the subcutaneous connective tissues of hosts. Thus, I suspect that the original record of this species from the coelome of Wallabia bicolor in coastal New South Wales (probably Milson Is.) ( Johnston & Mawson 1938a) is in error and much more likely to be B. (B.) mundayi which occurs commonly in pleural and peritoneal cavities of this host in southeastern Australia. Their record of two immature females of this species from the dorsal aorta of Macropus sp. from Inverleigh, near the Flinders River, Gulf of Carpenteria remains a mystery and the whereabouts of the material is unknown.