Contributions to Zoology, 86 (1) – 2017Isabel T Hyman; Irantzu de la Iglesia Lamborena; Frank Köhler: Molecular phylogenetics and systematic revision of the south-eastern Australian Helicarionidae (Gastropoda, Stylommatophora)

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Appendix

Zoobank Registration: lsid:zoobank.org:pub:0CEB38 99-B108-48FD-96D5-0076A3966DB1

 

Taxonomic account and descriptions

Helicarionidae Bourguignat, 1877

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Diagnosis. Shell present, complete or reduced, 5-35mm in diameter; usually thin-walled, glossy; spiral grooves present on protoconch and teleoconch. Mantle with accessory lobes lying over body and shell lappets of variable size lying over shell. Sole of foot tripartite; caudal apparatus present, formed from curled up sole. Kidney unilobed; minor venation on roof of mantle cavity absent or present; mantle gland absent. Genital system oviparous; oviduct glandular. Bursa copulatrix variable in length; inserted on vagina or, if vagina absent, at junction of free oviduct and penis. Stimulator absent. Epiphallus enters penis through simple pore, fleshy lips or verge; interior of penis variable. Penial tunica present, open at proximal end, attached by muscle fibres to epiphallus. Epiphallic retractor caecum absent or present; epiphallic flagellum absent or present; where present, flagellum contains an axial filament. Spermatophore a soft capsule with hard-walled tail-pipe.

Remarks. Belongs to Helicarionoidea along with Ariophantidae and Urocyclidae; a superfamily unified by the presence of a flagellum with an axial filament, an epiphallic caecum, and mantle lobes (Hausdorf, 1998; Hyman and Ponder, 2010). However, flagellum and caecum are absent, presumably secondarily lost, in some members of all three families (e.g. Tarocystis, Sheaia, Levidens; Hausdorf, 1998; Hyman and Ponder, 2010). The boundaries of the three families are poorly defined; one character, presence of a proximally open penial tunica, distinguishes Helicarionidae from the other two families (Hausdorf, 1998; Hyman and Ponder, 2010).

Brevisentis Hyman, 2007

Brevisentis Hyman, 2007: 93.

Type species: Helix jacksoniensis Gray, 1834, by original designation; masculine

 

Diagnosis. Shell. Medium-sized, discoidal to depressedly globose with low spire; whorls rounded; protoconch with fine, shallow spiral grooves, teleoconch with spiral rows of very fine pustules; umbilicus narrow (Fig. 7-8).

Animal. Grey to black with red mucus in at least two species. Mantle lobes moderately small; shell lappets elongate, triangular (Fig. 9).

Genitalia. Ovotestis of 4-5 lobes embedded in digestive gland. Talon usually embedded in digestive gland. Spermoviduct not folded. Distal portion of free oviduct with elongate capsular gland; free oviduct internally smooth. Bursa copulatrix relatively long, reaching half to two thirds of the way along spermoviduct, inserted on vagina, internally with longitudinal pilasters in duct; sac oval to tear-shaped. Vagina short, internally with longitudinal pilasters. Epiphallus enters penis through verge; epiphallic caecum absent; epiphallic flagellum with axial filament present, containing small internal cryptae. Spermatophore is a soft-walled capsule with hard tail-pipe, with short branching spines in spiraling pattern on tail-pipe.

Remarks. Brevisentis originally contained two species that were previously included in Melocystis Iredale, 1937, a genus now considered as a synonym of Nitor Gude, 1911 (Hyman, 2007). A third species, B. kaputarensis, was subsequently included for strong similarity in shell features (Stanisic et al., 2010). The monophyly of Brevisentis as so delineated is confirmed herein. The three allopatric species have very similar shells, but differ in body colouration and genital anatomy. Samples from Wollemi National Park, Great Dividing Range, do not group with any described Brevisentis species in mtDNA phylogeny but form the sister group of (B. atratus + B. kaputarensis), probably representing an undescribed species. However, because the single lot consists of immature specimens only, we delay description of this species pending collection of suitable material.

Brevisentis jacksoniensis (Gray, 1834)

Figs 7A-B, 8A-C, 9A-B, 10

FIG2

Fig. 7. Shells of Brevisentis. A-B. B. jacksoniensis: A. Syntype of Helix jacksoniensis NHMUK 20160421 (photographer: NHMUK). B. Holotype of Expocystis exclusus AM C.101143 (size indicative; photographer: AM). C. B. atratus AM C.455107. D. B. kaputarensis holotype QM MO49155 (size indicative; photographer: Michael Murphy). Scale bar = 10 mm.

FIG2

Fig. 8. Scanning electron micrographs showing shell microsculpture in Brevisentis. A-C. B. jacksoniensis AM C.205293: A. Protoconch. B. early teleoconch. C. mid-teleoconch. D-F. B. kaputarensis QM MO78909: D. Protoconch. E. early teleoconch. F. mid-teleoconch. Scale bars = 100 µm.

FIG2

Fig. 9. Live specimens of Brevisentis. A-B. B. atratus: A. AM C.517469, Camden. B. AM C.403813, Coolah Tops National Park. C. B. kaputarensis, QM MO78908, Mt Kaputar. Scale bars = approx. 10 mm.

Helix jacksoniensis Gray, 1834: 65; Reeve, 1854: pl. 207, sp. 1462.

Helix (Microcystis) jacksoniensis: Cox, 1868: 7, pl. 9, fig. 6.

Nanina (Microcystis) jacksoniensis: Pfeiffer and Clessin, 1881: 36.

Nanina (Subg. Xesta Sect. Microcystis) jacksoniensis: Tryon, 1886: 113, pl. 37, fig. 21.

Nanina (Microchlamys) jacksoniensis: Cox, 1909: 7.

Melocystis jacksoniensis: Iredale, 1937a: 5.

Expocystis exclusus Iredale, 1941: 4, fig. 7.

Melocystis exclusus: Smith, 1992: 238.

Brevisentis jacksoniensis: Hyman and Ponder, 2010: 44-45, figs 7J-K, 8J-K, 9J-K, 10B, 11B, 12G-H, 13G, 14D, 15P-R; Stanisic et al., 2010: 280-281.

 

Material examined. Types. 2 syntypes of Helix jacksoniensis Gray, 1834 (NHMUK 20160421, Port Jackson, leg. Lamb; Fig. 7A), holotype of Expocystis exclusus Iredale, 1937 AM C.101143, Broken Bay, N of Sydney; Fig. 7B).

Non-type material. See Table S1. Additional material: AM C.319501, AM C.348670, AM C.356645, AM C.162857, AM C.320010.

 

Diagnosis. Shell. Medium-sized, 4.65-5.25 whorls, amber to golden brown, depressedly subglogose with a low spire; whorls rounded, sutures impressed; protoconch with fine spiral grooves, teleoconch with fine spiral grooves becoming pustulose towards the last whorl. Umbilicus narrow (Table 3, Figs 7A-B, 8A-C).

FIG2

Table 3. Shell dimensions. Abbreviations: n = number of measured specimens, SH = shell height, SD = shell diameter, NW = number of whorls, AH = aperture height, AW = aperture width.

Animal. Grey with red mucus (staining body red); sole with a slightly paler mid field; caudal gland small. Mantle lobes moderately small, not fused; shell lappets narrow, triangular, right lappet moderately long, left lappet moderately short (Fig. 9A-B).

Genitalia. Penis moderately short, enclosed in penial tunica, slightly swollen at proximal end. Penis interior with longitudinal pilasters at distal end, breaking up into pustules at proximal end. Epiphallus relatively short, approximately 1.5 times penis length, entering penis apically or slightly to one side through rounded, pustulose verge of variable length, ranging from one third to three quarters penis length. Epiphallic flagellum with small internal cryptae near epiphallus, tail of flagellum without visible cryptae. Spermatophore with smooth capsule; one complex branching spine on tail-pipe near capsule; opposite, eight increasingly smaller and simpler short branching spines spiraling around tail-pipe only (Fig. 10).

FIG2

Fig. 10. Reproductive anatomy of Brevisentis jacksoniensis, AM C.205293. A. Genitalia. B. Spermatophore. C. Penial complex. D. Penial interior. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 2 mm, B-D = 1 mm.

Remarks. This species was considered to be a synonym of Nitor circumcinctus for some years (Cox, 1909; Iredale, 1937a). Subsequently, its type locality was treated as doubtful and it was placed in incertae sedis (Iredale, 1941; Smith, 1992; Smith et al., 2002). After examination of the holotype the species was resurrected by Hyman (2007) with Expocystis exclusus as a synonym; this treatment is followed here.

Brevisentis jacksoniensis is found from Kiama to the Hawkesbury Valley in dry vine thicket, sclerophyll forest and woodland, living in litter and under logs and rocks (Fig. 11). It was recorded by Hyman and Ponder (2010) as ranging to the Warrumbungle National Park in the north, and in the west to Bathurst and the Blue Mountains, but this appears to be in error: specimens from all three localities have been identified herein as belonging to B. atratus. Specimens from Mulgoa and Richmond (W of Sydney), Palm Beach and Hornsby (N Sydney), Berowra Waters and Berowra Creek (N of Sydney) and Kurrajong (Hawkesbury Valley) have all been confirmed by sequencing or dissection to belong to B. jacksoniensis.

FIG2

Fig. 11. Occurrence records of Brevisentis from the malacological collection of the Australian Museum, Sydney. Symbols: ● = B. jacksoniensis, Δ = B. atratus, + = B. kaputarensis , = B. n. sp. Wollemi NP.

While the three Brevisentis species all have very similar shells, the shell of B. jacksoniensis is slightly larger than B. kaputarensis, with more impressed sutures, and B. jacksoniensis has a higher spire than either of its congeners (Fig. 1A). This species can also be identified by its paler body colouration stained with red mucus. Brevisentis atratus also produces red mucus, but its black body colour renders it less visible. Anatomically, the species are easily distinguished by the longer penis with a blind tip seen in B. atratus and the absence of pustules in the penis interior of B. kaputarensis.

Brevisentis atratus Hyman, 2007

Figs 7C, 12

Brevisentis atratus Hyman, 2007: 94-95, fig. 2; Stanisic et al., 2010: 280-282, 324.

 

Material examined. Holotype. AM C.446473 (Coolah Tops NP, at lookout 1km S of Pinnacle Lookout, end of Pinnacle Rd, NSW, 31°41.8’ S, 150°1.1’ E, basalt rock pile at edge of escarpment, leg. Shea, 26/5/2000).

Paratypes. AM C.44645 (same data as holotype).

Non-type material. See Table S1. Additional material: AM C.446477, AM C.360695, AM C.164160, AM C.135218, AM C.446459, AM C.517469.

 

Diagnosis. Shell. Medium size, 4.65-5.5 whorls, pale amber, depressedly globose to discoidal; whorls rounded; protoconch with very fine spiral grooves; teleoconch with spiral rows of tiny pustules; umbilicus narrow (Table 3, Fig. 7C).

Animal. Body dark grey to black with red mucus; tiny specks of red pigment visible in living animal; sole pale grey with a paler mid field; caudal horn small. Mantle lobes moderately small, not fused; shell lappets narrow, triangular, right lappet moderately long, left lappet moderately short.

Genitalia. Penis relatively long, sometimes not fully enclosed in penial tunica, not swollen proximally. Penis interior with primarily longitudinal pilasters at distal end, breaking into wavy pilasters then into pustules at proximal end; interior with a sharp narrowing near proximal end that is not visible externally, effectively dividing penis into two separate chambers. Epiphallus relatively short, approximately 1.5 times penis length, entering penis to one side through short verge (about ¼ penis length), leaving a small blind penis tip. Epiphallic flagellum with small internal cryptae near epiphallus, tail of flagellum without visible cryptae. Spermatophore not observed (Fig. 12).

FIG2

Fig. 12. Reproductive anatomy of Brevisentis atratus, AM C.360695. A. Genitalia. B. Spermatophore. B. Penial interior. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 2 mm, B = 1 mm.

Remarks. When first described, B. atratus was recorded from Mt Gibraltar (Bowral, SE NSW) to Coolah Tops National Park in the Hunter region (Hyman, 2007). Specimens from the Blue Mountains, Bathurst and Warrumbungle National Park, mistakenly attributed to B. jacksoniensis, are here shown to belong to B. atratus (Fig. 11). This species is found in rocky outcrops in vine thicket, sclerophyll forest and open woodland, living under rocks, logs and litter.

Brevisentis atratus can be distinguished from B. jacksoniensis by its black body colouration and lower-spired shell, and from B. kaputarensis by its larger and paler shell with a flatter spire. It has a very distinctive penial anatomy, with a longer penis than its congeners, distinguished by a narrowing near the proximal end that gives the impression of two separate chambers and a small blind tip.

Brevisentis kaputarensis Shea and Griffiths, 2010

Figs 7D, 8D-F, 9C, 13

Brevisentis kaputarensis Shea and Griffiths, 2010 in Stanisic et al., 2010: 280-281.

 

Material examined. Holotype. QMMO32629 (Dawsons Spring, Mt Kaputar, NE NSW, 30°16’S, 150°10’E, fern gully/remnant rainforest, under logs, in litter, leg. Stanisic et al., 8/11/1983; Fig. 7D).

Paratypes. QMMO49155, QMMO78908.

Non-type material. See Table S1.

 

Diagnosis. Shell. Medium size, 4.6-5.2 whorls, pale amber, discoidal with a low spire; whorls rounded; protoconch with very fine spiral grooves, teleoconch with spiral rows of very fine pustules; umbilicus narrow (Table 3, Figs 7D, 8D-F).

Animal. Body dark grey; sole with a paler mid field; caudal horn moderately small. Mantle lobes moderately small, not fused; Shell lappets narrow, triangular, right lappet moderately long, left lappet moderately short (Fig. 9C).

Genitalia. Penis relatively short, thick, tubular; penis and sometimes part of epiphallus enclosed in penial tunica. Penis interior with anastomosing ridges, primarily longitudinal. Epiphallus relatively short, approximately twice penis length, entering penis apically through a verge of variable length (1/3 to nearly complete penis length). Epiphallic flagellum with small internal cryptae near epiphallus, tail of flagellum without visible cryptae. Spermatophore with smooth capsule; one complex branching spine on tail-pipe near capsule; opposite, fourteen increasingly smaller and simpler short branching spines spiraling around tail-pipe only (Fig. 13).

FIG2

Fig. 13. Reproductive anatomy of Brevisentis kaputarensis, QM MO78908. A. Genitalia. B. Penial complex. C-D. Penial interior. E. Spermatophore. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars = 1 mm.

Remarks. Known only from Nandewar Range, northeastern NSW (Fig. 11), B. kaputarensis is the smallest species of this genus, found in dry woodland, in rock piles and leaf litter. This species has a larger penial verge than its congeners, and the penis interior has anastomosing ridges rather than pustules.

Mysticarion Iredale, 1941

Mysticarion Iredale, 1941: 7.

Type species: Mysticarion leucospira insuetus Iredale, 1941, by original designation; masculine

 

Diagnosis. Shell. Small to medium-sized, thin, pale golden to golden amber, glossy, globose with a low spire, whorls rounded. Protoconch with notched spiral grooves, teleoconch with very fine spiral grooves or smooth (Figs 14-15).

FIG2

Fig. 14. Shells of Mysticarion. A-B. M. insuetus: A. Syntype of Mysticarion leucospira insuetus AM C113776 (size indicative; photographer: AM). B. M. insuetus AM C.517468. C. Paratype of Mysticarion obscurior sp. Nov. AM C.524914. D. M. porrectus (probable holotype of Helicarion porrectus Iredale, 1941 AM C.101136 (size indicative, photographer: AM). E-F. M. hyalinus: E. Syntype of Vitrina hyalina Pfeiffer, 1855 NHMUK 1983075 (photographer: NHMUK). F. Holotype of Fastosarion staffordarum Stanisic, 2010, QM MO62372 (size indicative, photographer: QM). Scale bar = 10 mm.

FIG2

Fig. 15. Scanning electron micrographs showing shell microsculpture in Mysticarion. A-C. M. insuetus, QM MO49377: A. Protoconch. B. early teleoconch. C. mid-teleoconch. D-F. M. hyalinus, AM C.512424: D. Protoconch. E. early teleoconch. F. mid-teleoconch. Scale bars = 100 µm.

Animal. Pale cream with minimal markings. Mantle lobes usually small, not fused; shell lappets wide at base, rapidly tapering, triangular (Fig. 16).

FIG2

Fig. 16. Live specimens of Mysticarion. A. M. insuetus, AM C.506269, Oxley Wild Rivers NP. B. M. obscurior, AM C.524913, Wambina NR. C. M. porrectus, AM C.5000935, Mt Tomah. D. M. porrectus, resting pose, AM C.517188, Dorrigo. E-F. M. hyalinus: E. AM C.512424, Cunningham’s Gap. F. AM C.512438, Bunya Mountains. Scale bars = approx. 5 mm.

Genitalia. Spermoviduct of 2-4 lobes, embedded in digestive gland. Talon and carrefour embedded in albumen gland. Spermoviduct not folded into foot but with 1-2 small folds in proximal half. Free oviduct with spherical capsular gland in distal portion; internal walls of capsular gland smooth, remainder of free oviduct with weak longitudinal pilasters (absent in one species). Bursa copulatrix moderately long, approximately half spermoviduct length; sac portion oval; inserted at junction of penis and free oviduct. Vagina absent. Penis tunica attached tightly by muscle fibres to middle of epiphallus; epiphallus enters penis through verge; epiphallic caecum absent; near flagellum, epi­phallus containing internal cryptae; epiphallic flagellum with axial filament present, containing spiraling rows of internal cryptae with a long slender tail. Spermatophore a soft-walled capsule with hard tail-pipe; relatively long, complex branching spines present in spiraling pattern along tail-pipe.

Remarks. Mysticarion was introduced for two globose semislugs, Vitrina leucospira Pfeiffer, 1857 and its nominal subspecies M. leucospira insuetus (Iredale, 1941). Vitrina leucospira has since been excluded from the genus based on an examination of the type material, which does not appear to be that of an Australian helicarionid; and the taxon insuetus has been elevated to a full species. Helicarion porrectus Iredale, 1941 and Vitrina hyalina Pfeiffer, 1855 were both subsequently included in Mysticarion by Hyman and Ponder (2010) and followed by Stanisic et al. (2010). This treatment is confirmed here by molecular phylogenetics.

This genus contains arboreal semislugs with a golden, globose shell with a relatively high whorl count and a pale body. The genital features unifying this group include a moderately long penis with a penial verge and longitudinal pilasters, a short epiphallus with an accessory flagellum containing internal cryptae but no epiphallic caecum, a moderately long bursa copulatrix, and the absence of a vagina. The spermatophore is spiraling with branched spines present on the base of the capsule and continuing on the tail-pipe.

Mysticarion insuetus Iredale, 1941

Figs 14A-B, 15A-C, 16A, 17

Mysticarion leucospira insuetus Iredale, 1941: 7.

Mysticarion insuetus: Stanisic et al., 2010: 308-309, 330.

 

Material examined. Syntype. AM C.113776 (Scone, 32° 03’ S, 150° 52’ E, pre-1912; Fig. 14A).

Non-type material. See Table S1. Additional specimens: AM C.164470, AM C.456541, AM C.204742, AM C.204744.

 

Diagnosis. Shell. Small, thin, transparent golden amber, 3.2-3.9 whorls, glossy, globose with a low spire, whorls rounded. Protoconch with slightly notched, deep spiral grooves, teleoconch with very fine, shallow spiral grooves (Table 3, Figs 14A-B, 15A-C).

Animal. Body white with pale brown speckling on sole, tail tip, neck and shell lappets; eye tentacles grey, grey stripe down each side of tail, black line along mantle edge. Caudal horn large. Right mantle lobe moderately large, other lobes small, none fused. Right shell lappet long, left lappet moderately long, both lappets wide at base, rapidly tapering, white with black markings on underside (Fig. 16A).

Genitalia. Bursa copulatrix moderately short, less than half spermoviduct length, duct uniform in diameter, sac tear-shaped. Penis moderately long, slender, slightly swollen proximally, enclosed in penial tunica. Penis interior with two main longitudinal pilasters and multiple longitudinal ridges. Epiphallus short, approx. penis length, containing internal cryptae near flagellum; entering penis through rounded verge with a pointed tip of about half penis length. Epiphallic flagellum with few internal cryptae and slender tail. Spermatophore with 7 branched spines in a spiraling pattern: three widely spaced, complex branched spines at base of capsule, followed by four closely spaced, increasingly short and simple spines on tail-pipe (Fig. 17).

FIG2

Fig. 17. Reproductive anatomy of Mysticarion insuetus, AM C.483756. A. Genitalia. B. Penial complex. C. Penial interior. D. Spermatophore. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 2 mm, B-D = 1 mm.

Remarks. Mysticarion insuetus has a wide, disjunct range that stretches over 400 km, from Razorback, south of Sydney to the Macleay Valley in northern NSW (Fig. 18). Throughout much of its range, M. insuetus is sympatric with both M. porrectus and M. obscurior sp. Nov. However, M. insuetus is generally found at lower altitudes, in drier habitats, such as vine thickets, while M. porrectus and M. obscurior are present in rainforest or wet sclerophyll forests, and in the case of M. porrectus, usually at higher altitudes.

FIG2

Fig. 18. Occurrence records of Mysticarion species from the malacological collection of the Australian Museum, Sydney. Symbols: ● = M. porrectus, Δ = M. insuetus, + = M. hyalinus, ♠ = M. obscurior.

The type locality for M. insuetus was given simply as ‘Scone’ (Iredale, 1941). The material closest to this locality in the research collections of the AM and the QM comes from Barrington Tops National Park, where three Mysticarion species are present: M. insuetus, M. porrectus, and M. obscurior. Mysticarion porrectus can easily be distinguished from its congeners by its larger size and less globose shell with a flatter spire and fewer whorls. Mysticarion insuetus and M. obscurior have much more similar shells, but the shell of M. insuetus is slightly higher-spired and the last whorl descends more rapidly. The two species can be distinguished much more readily alive, since the body of M. obscurior is considerably darker with orange and brown pigmentation. The fourth species in the genus, M. hyalinus, is found further to the north, and while it is very similar in appearance to M. insuetus it can be distinguished by its larger size and paler body colour. M. insuetus has a unique genital anatomy consisting of a short epiphallus and moderately long, narrow penis with a long verge and numerous longitudinal pilasters.

Mysticarion obscurior sp. nov.

Figs 14C, 16B, 19

Material examined. Holotype. AM C.524913 (Matcham, Matcham Road, Wambina Nature Reserve, NSW, 33°24’10” S, 151°26’24” E, leg. Hyman et al., 13/5/2016).

Paratype. AM C.524914 (same data as holotype; Fig. 14C).

Non-type material. See Table S1. Additional material: AM C.340671, AM C.334330, AM C.456540, AM C.500932.

 

Diagnosis. Shell. Small, thin, transparent, pale gold, 3.5 whorls, glossy, globose with a very low spire, whorls rounded. Protoconch with notched spiral grooves, teleoconch with very fine, shallow spiral grooves (Fig. 14C).

Animal. Body beige, pale orange along neck and middle of tail, brown on sides of tail, eye tentacles grey, black line along mantle edge, right shell lappet discontinuously edged with black. Caudal horn large. Right mantle lobe moderately large, other lobes small, none fused. Right shell lappet long, left lappet moderately long, both lappets wide at base, rapidly tapering; with black markings (Fig. 16B).

Genitalia. Bursa copulatrix moderately long, reaching half way along spermoviduct; bursa duct broad at base, then narrowing; sac small, oval-shaped. Penis moderately long, broad, slightly swollen at proximal end, enclosed in penial tunica. Penis interior with two main longitudinal pilasters and multiple wavy ridges. Epiphallus short, approx. 1-1.5 times penis length, containing very few (2-3) internal cryptae near flagellum; entering penis through pyramidal verge of about one third penis length. Epiphallic flagellum slender, with no internal cryptae. Spermatophore not observed (Fig. 19).

FIG2

Fig. 19. Reproductive anatomy of Mysticarion obscurior sp. Nov. A. Genitalia (AM C.374301). B. Penial interior (AM C.524913). Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars = 1 mm.

Remarks. Mysticarion obscurior has a very similar range to M. insuetus, stretching from just north of Gosford on the Central Coast to Bellingen in northern NSW, in subtropical rainforest and wet sclerophyll forest (Fig. 18). The two species co-occur in some areas (e.g. Little Jilliby State Conservation Preserve), although in general M. insuetus appears to be found in drier habitats. Mysticarion obscurior can be distinguished from its congeners by its darker body colouration of brown and pale orange pigmentation and by the discontinuous black edge on the right shell lappet, which is visible in preserved specimens even when the rest of the body pigmentation has faded. This species also has a paler shell with a slightly lower spire than M. insuetus. Unique features of its genital anatomy include a flagellum with no internal cryptae and only a very few cryptae in the epiphallus, indicating that the spermatophore very fewer branching spines. It also has a distinctive pyramidal penial verge and a bursa copulatrix with a long duct with broad and narrow portions and a small, distinct, rounded sac.

Etymology. From obscurior (Latin = darker; adjective), referring to the darker colour of the animal relative to its congeners.

Mysticarion porrectus (Iredale, 1941)

Figs 14D, 16C-D, 20

Helicarion porrectus Iredale, 1941: 6; Smith, 1992: 234.

Mysticarion porrectus: Hyman and Ponder, 2010: 36-37, Figs 5B, 6C, 7F, 8F, 9F, 12C, 13C, 15D-F, 16A; Stanisic et al., 2010: 308-309, 329.

 

Material examined. Probable holotype. AM C.101136 (NW of Sydney, Blue Mountains, Mt. Irvine, NSW, 33°29’ S, 150°28’ E, leg. Ward, pre-1941; Fig. 14D).

Non-type material. See Table S1.

 

Diagnosis. Shell. Small to medium-sized, thin, glossy, pale golden to golden amber, globose with a flat spire. Protoconch with notched spiral grooves, teleoconch with fine, almost indistinct spiral grooves (Fig. 14D).

Animal. Body pale cream with no markings. Caudal horn moderately large. Sole slightly paler than body. Mantle lobes moderately small; shell lappets moderately small, wide at base, rapidly tapering, joined by a narrow mantle collar (Table 3; Fig. 16C-D).

Genitalia. Penis of moderate length, proximally swollen, penis and part of epiphallus enclosed in penial tunica. Penis interior with two main longitudinal pilasters, one proximally swollen, and multiple smaller longitudinal pilasters breaking up into pilasters at proximal end. Epiphallus very short, approx. equal to penis length, containing internal cryptae near flagellum; entering penis through very short verge. Epiphallic flagellum with internal cryptae and slender tail. Spermatophore not observed (Fig. 20).

FIG2

Fig. 20. Reproductive anatomy of Mysticarion porrectus, AM C.462828. A. Genitalia. B. Penial interior. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A= 2mm, B = 1 mm.

Remarks. Hyman and Ponder (2010) revised this species and concluded that it had a very wide range containing several disjunct populations that were all identical anatomically (Fig. 18). Our DNA study confirms that populations from Robertson, the Royal NP, the Blue Mountains, Barrington Tops, Dorrigo and the Gibraltar Range are genetically not very distinct and since they are also indistinguishable anatomically, should be treated as a single species. Material from Narooma in southeastern NSW was unable to be conclusively allocated to this species since the single specimen available was immature and proved unsuitable for DNA extraction. Even if this population is excluded, the range of this species still bridges a linear distance of over 500 km. Mysticarion porrectus is generally found at higher altitudes than M. obscurior and M. insuetus, both of which are found throughout parts of its range, and occupies rainforest or wet sclerophyll forest.

Mysticarion porrectus is larger and less globose than its congeners, with a lower whorl count. It also has the shortest epiphallus of any Mysticarion species. Other distinctive features of the reproductive system include a proximally swollen penis with two main longitudinal pilasters and a very short penial verge, and very little folding present in the ovotestis.

Mysticarion hyalinus (Pfeiffer, 1855)

Figs 14E-F, 15D-F, 16E-F, 21

Vitrina hyalina Pfeiffer, 1855: 296-297; Reeve, 1862: pl. 9, sp. 68; Cox, 1868: 85, pl. 14, figs 7, 7a; Pfeiffer, 1876: 24; Smith, 1992: 244 (in Incertae Sedis).

Vitrina hyalinus: Cox, 1909: 6.

Helicarion hyalinus: Tryon, 1885: 172, pl. 39, figs 75-76; Hedley, 1888: 50; Iredale, 1937a: 8.

Mysticarion hyalina: Hyman and Ponder, 2010: 38-40, figs 5C, 7G, 8G, 9G, 12D, 13D, 15G-I.

Mysticarion hyalinus: Stanisic et al., 2010: 308-309.

Fastosarion staffordorum Stanisic, 2010 (in Stanisic et al., 2010): 304-305.

 

Material examined. Types. Syntype of Vitrina hyalina Pfeiffer, 1855 NHMUK 1983075 (Moreton Bay, leg. Strange, coll. Cuming; Fig. 14E), Holotype of Fastosarion staffordorum Stanisic, 2010 QMMO62372 (Dandabah, Bunya Mts NP, SE Qld, 26°53’S, 151°36’E, notophyll vine forest, leg. Bishop, 5/3/1976; Fig. 14F).

Non-type material. See Table S1.

 

Diagnosis. Shell. Medium-sized, 3.9-4.6 whorls, thin, glossy, golden amber, globose with a low spire. Protoconch with notched spiral grooves, continuing onto early teleoconch; last whorls with no sculpture (Table 3, Figs 14E-F, 15D-F).

Animal. Body very pale cream with faint brown specks of pigment on shell lappets and mantle edge. Caudal horn moderately large. Right mantle lobe moderately small, other lobes small, none fused. Shell lappets moderately small, wide at base, rapidly tapering (Fig. 16E-F).

Genitalia. Penis moderately long, tubular, slightly swollen proximally, penis and sometimes part of epiphallus enclosed in penial tunica. Penis interior with one main longitudinal pilaster and multiple smaller longitudinal pilasters. Epiphallus long, approx. twice penis length, containing internal cryptae near flagellum; entering penis through verge of about one quarter to one third penis length. Epiphallic flagellum with internal cryptae and slender tail. Spermatophore with 11-13 moderately long, complex branched spines in a spiraling pattern; initial 3-4 spines situated on base of capsule, remainder on tail-pipe; initial 5-6 spines have two major branches; remaining spines are increasingly short and simple (Fig. 21).

FIG2

Fig. 21. Reproductive anatomy of Mysticarion hyalinus A-C. AM C.512424, Cunninghams Gap, SE Qld: A. Genitalia. B. Penial complex. C. Penial interior. D-G. AM C.512438, Bunya Moutains, SE Qld: D. Genitalia. E. Penial complex. F. Penial interior. G. Spermatophore. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A, D = 2 mm, B-C, E-G = 1 mm.

Remarks. This species was revised by Hyman and Ponder (2010). Here we include another species, Fastosarion staffordorum, in synonymy with Mysticarion porrectus. Three specimens from each species were sequenced, and while the two populations form monophyletic groups in the phylogenetic tree, the genetic distances between them are clearly within the range of intraspecific divergence (Table 1). Dissections revealed that the genitalia of the two taxa are identical; hence F. stafforodum is here considered as a younger synonym of M. hyalinus. This means that like M. porrectus and M. insuetus, M. hyalinus has a disjunct range including populations from Casino in northern NSW to Cunningham’s Gap in southern QLD, and 200 km further north, one population in Bunya Mountains National Park (QLD) (Fig. 18). This species is found in rainforest habitats, living on tree trunks and leaves.

Mysticarion hyalinus is larger than M. insuetus but slightly smaller than M. porrectus (Fig. 1B). It has a globose shell into which it can fully retract and a very pale body. Unique genital features of this species include a longer epiphallus than its congeners, a moderately long penis with a moderately small verge and one main longitudinal pilaster.

Parmavitrina Iredale, 1937

Parmavitrina Iredale, 1937c: 8.

Type species: Helicarion planilabris Cox, 1866, by monotypy; feminine

Desidarion Iredale, 1941: 8.

Type species: Desidarion dispositus Iredale, 1941, by monotypy; masculine

 

Diagnosis. Shell. Medium to large, ear-shaped, last whorl large and sometimes widely flared, umbilicus closed, shell glossy, protoconch and teleoconch sculptured with fine to very fine spiral grooves or smooth (Figs 22, 23).

FIG2

Fig. 22. Shells of Parmavitrina. A. P. planilabris, AM C.103630. B. P. rubrica AM C.376379 (size indicative; photographer; AM). C. P. disposita, probable holotype of Desidarion rubricus Iredale, 1941, AM C.101147 (size indicative; photographer: AM). D. P. megastoma, AM C.448244. E. P. maculosa, AM C.448344. F. P. flavocarinata, AM C.512373. Scale bar – 10 mm.

FIG2

Fig. 23. Scanning electron micrographs showing shell microsculpture in Parmavitrina. A-C. P. planilabris, AM C.460262: A. Protoconch. B. early teleoconch. C. mid-teleoconch. D. P. rubrica, AM C.205515 protoconch. E. P. disposita, AM C.460261, early teleoconch. F. P. flavocarinata, AM C.512373, protoconch. Scale bars = 100 µm.

Animal. Grey to dark brown, usually with spots or mottling on sides of tail and mantle lobes and shell lappets. Tail keeled. Mantle lobes small to moderately large, cephalic shield formed from median lobe or from fused left and median lobes, shell lappets moderately small to moderately large, connected by narrow collar; right lappet rounded (Fig. 24).

FIG2

Fig. 24. Live specimens of Parmavitrina. A. P. planilabris, MV F219267, Boorganna Nature Reserve (photographer: A. Moussalli). B. P. rubrica, AM C.524922, Forest of Tranquility. C. P. megastoma, C.512383, Washpool National Park. D. P. flavocarinata, C.512374, Washpool National Park. Scale bars = approx. 10 mm.

Genitalia. Ovotestis of 3-5 lobes embedded in digestive gland. Carrefour and talon usually embedded in albumen gland. Spermoviduct folded 3-4 times. Distal portion of free oviduct with elongate capsular gland with no internal sculpture, remainder of free oviduct with longitudinal pilasters internally. Bursa copulatrix relatively short, internally with longitudinal pilasters (duct) and weak transverse ridges (sac), inserted on vagina. Vagina short to moderately long, internally with longitudinal pilasters. Epiphallus enters penis through a verge or thickened ring; epiphallic caecum absent; near flagellum, epiphallus often swollen with internal cryptae; epiphallic flagellum with axial filament present, containing spiraling rows of internal cryptae with a long slender tail. Spermatophore a soft-walled capsule with hard tail-pipe; branching spines present in spiraling pattern on base of capsule and along part of tail-pipe; sometimes with a single long spine towards end of tail-pipe.

Remarks. Parmavitrina was introduced as a monotypic subgenus of Helicarion based on its expansive mouth, flattened upper surface and degenerate base (Iredale, 1937a). Another large semislug, Desidarion dispositus, was described a few years later as being similar to Parmavitrina, but due to its less flattened shell and complete base it was placed in a separate genus (Iredale, 1941). The anatomy of P. planilabris was described by Hyman and Ponder (2010) but no anatomical details have previously been known for Desidarion, beyond a few brief notes given to justify taking Desidarion out of synonymy with Helicarion (Hyman and Ponder, 2010). In the current study, shell shape analysis reflected the groupings shown by Stanisic et al. (2010), with D. dispositus and D. rubricus showing a significantly lower H/D ratio compared to P. planilabris and P. megastoma. However, mitochondrial phylogeny and comparative anatomy indicated that these similarities are superficial and that the two genus names are synonymous with the older Parmavitrina having priority. Anatomical comparisons show that Parmavitrina and Desidarion are unified by their large size, flattened shell with a wide aperture, and spermatophore with spines situated primarily on the capsule rather than the tail-pipe (corresponding to internal cryptae in the epiphallus and very few in the flagellum).

In the phylogenetic tree, Parmavitrina megastoma clustered with two undescribed species. Individuals from the three populations differed in shell size and aperture width: the largest species (from Bruxner Park, north of Coffs Harbour) had a very widely flared last whorl and wide aperture, the smallest species (from Gibraltar Range National Park) had a less flared last whorl and narrower aperture, and the third species was intermediate. The three species also differed anatomically. The intermediate species most closely matches the holotype of P. megastoma and the other two species are described below as P. maculosa sp. nov. and P. flavocarinata sp. nov. These three closely related species differ from the rest of Parmavitrina in several ways; they have a more reduced shell of fewer than 3 whorls, a more slender penis with a verge, a longer vagina and bursa copulatrix, and a more slender flagellum forming a spermatophore with short, highly branched spines. However, this clade of three species groups with the other members of Parmavitrina based on both morphology and molecular phylogeny and does not warrant recognition as a separate genus.

Parmavitrina planilabris (Cox, 1866)

Figs 22A, 23A-C, 24A, 25

Vitrina planilabris Cox, 1866: 45-46 [Holotype missing, presumed lost].

Vitrina macgillivrayi Cox, 1868: 86, pl. 14, figs 8, 8a; Pfeiffer, 1876: 23; Cox, 1909: 6 [unnecessary replacement name for Vitrina planilabris Cox, 1866].

Helicarion macgillivrayi: Tryon, 1885: 171, pl. 39, figs 67-68.

Parmavitrina planilabris: Iredale, 1937a: 8, fig.8; Smith, 1992: 240; Hyman and Ponder, 2010: 40-41, figs 5D, 7H, 8H, 9H, 12E, 13E, 15J-L; Stanisic et al., 2010: 312-313.

 

Material examined. Non-type material. See Table S1.

 

Diagnosis. Shell. Large, 3.4-3.8 whorls, amber, ear-shaped, flattened; last whorl large and flared with membraneous base; shell glossy, protoconch sculptured with fine spiral grooves, teleoconch sculptured with very fine, almost indistinct spiral grooves (Table 3, Figs 22A, 23A-C).

Animal. Grey with darker spots and orange-red mucus, mid field of sole paler than outer fields, sides of sole striped, slime network prominent, tail strongly keeled, caudal gland small. Mantle lobes moderate size, left and median lobes fused; shell lappets broad but moderately small, rounded, pustulose (Fig. 24A).

Genitalia. Bursa copulatrix short, equal in length to free oviduct, sac spherical to tear-shaped. Penis large, often very swollen at proximal end, sometimes not fully covered by penial tunica. Penis interior with irregular pilasters, ridged, primarily longitudinal at distal end, breaking up into irregular pilasters or pustules at proximal end. One pilaster often greatly swollen towards proximal end. Epiphallus relatively short, approx. 1-1.5 x penis length, containing internal cryptae near flagellum; entering penis through thickened ring. Epiphallic flagellum with internal cryptae and long, slender tail. Spermatophore not observed (Fig. 25).

FIG2

Fig. 25. Reproductive anatomy of Parmavitrina planilabris, QM MO33052: A. Genitalia. B. Penial interior. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 4 mm, B = 2 mm.

Remarks. The species has been described from “Mitchell River, NSW”; the holotype is presumed lost. Parmavitrina planilabris is distributed from Buladelah in mideastern NSW to the New England Tableland in northern NSW in rainforest and wet sclerophyll forest (Fig. 26). This species has a unique genital anatomy consisting of a large, proximally swollen penis with no penial verge and irregular penial pilasters. It can be distinguished from the similarly sized P. disposita, with which it is sympatric around Chichester, by its grey colouration and shell with a membraneous base.

FIG2

Fig. 26. Occurrence records of Parmavitrina species from the malacological collection of the Australian Museum, Sydney. Symbols: □ = P. planilabris, ● = P. megastoma,  = P. rubrica, + = P. flavocarinata, ♠ = P. maculosa, ♦ = P. disposita.

This species is phenotypically similar and phylogenetically closely related to P. rubrica. Both taxa differ from each other by p-distances of 2.4-3.6% in 16S and 5-6% in COI, which are near the boundary between intra- and interspecific distances. It is possible that P. rubrica should not be retained as a separate species and merely represents a southern range extension of P. planilabris. However, there are several differences between the two taxa. Parmavitrina rubrica is slightly smaller, with a more orange-brown colouration; it does not have fused mantle lobes; it has a smaller, more tubular penis with a penial verge, internally with deep, zigzagging or wavy longitudinal pilasters. In contrast, P. planilabris is larger, more grey in colouration, with fused mantle lobes; it has a proximally swollen penis with a short penial tunica, no penial verge, and less distinct, broad, shallow, ridged pilasters breaking up into irregular pilasters or pustules proximally. Given the small but distinct morphological differences between the two taxa we retain P. rubrica as separate species.

Parmavitrina rubrica (Shea and Griffiths, 2010) comb. nov.

Figs 22B, 23D, 24B, 27

Desidarion rubricus Shea and Griffiths, 2010 in Stanisic et al., 2010: 310-311, 331.

 

Material examined. Holotype. AM C.376379 (W of Wyee, NW of Dooralong, Watagan road above Lowers Gully, NSW, 33°10.150’S, 151°19.100’E, on rock near road edge, 13/1/2000).

Paratypes. AM C.333639; AM C.373660; AM C.373672; AM C.373717; QMMO29727.

Non-type material. See Table S1.

 

Diagnosis. Shell. Large, 3.4-3.6 whorls, greenish amber, ear-shaped, flattened, protoconch slightly raised, last whorl large with membranous base; shell glossy, protoconch with fine spiral grooves, teleoconch with very fine, indistinct spiral grooves (Table 3, Fig 22B, 23D).

Animal. Orange-brown with orange-red mucus, faintly spotted on tail and shell lappets; mid field of tail paler than outer fields, slime network well-developed, tail keeled, caudal horn small. Mantle lobes moderately large, none fused, median lobe forming cephalic shield; shell lappets moderately large, connected by a narrow collar, right lappet rounded with a small point (Fig. 24B).

Genitalia. Bursa copulatrix short, slightly longer than free oviduct, sac oval-shaped. Penis thick, moderately short, cylindrical, penis and part of epiphallus enclosed in penial tunica; penis interior with approx. five longitudinal, wavy pilasters. Epiphallus moderately short, about twice penis length, with internal cryptae adjacent to flagellum; epiphallus enters penis through short to medium length verge, ranging from one sixth to half penis length. Epiphallic flagellum with spiraling rows of internal cryptae and slender tail. Spermatophore with branching spines present in spiraling pattern on base of capsule; two rows of spines present, one of thick spines with multiple branches and the second of thinner, shorter, less branched spines; tail pipe short, smooth, without branches (Fig. 27).

FIG2

Fig. 27. Reproductive anatomy of Parmavitrina rubrica. A. Genitalia, C.364763. B. Penial interior, C.364763. C. Genitalia, AM C.334201. D. Penial interior, QM MO49406. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A, C = 4 mm, B, D = 2 mm.

Remarks. Parmavitrina rubrica is found in the Hunter region, in rainforest and wet sclerophyll forest from Ourimbah to north of Buladelah, with a population in Gordon (northern Sydney) that may be introduced (Fig. 26; Stanisic et al., 2010). This species is slightly smaller than P. planilabris and P. disposita. This size difference and its degenerate shell edge distinguishes it from P. disposita, which shares its orange-brown body colour. This species can also be distinguished by the combination of a moderately short, thick penis with wavy longitudinal pilasters and a short to moderately long penial verge.

A population south of Taree is anatomically distinct from the rest of the species. It has a much longer epiphallus and a penis with a shorter verge. Only a single sequence represents this population in the tree and while it is distinct from the rest of the species, it is not highly divergent, and so we retain this population as part of P. rubrica.

Parmavitrina disposita (Iredale, 1941) comb. nov.

Figs 22C, 23E, 28

Desidarion dispositus Iredale, 1941: 8; Stanisic et al., 2010: 310-311, 330.

Helicarion dispositus: Smith, 1992: 233.

 

Material examined. Probable holotype. AM C.101147 (Barrington Tops, NSW, 32°01’30” S, 151° 27’00” E, pre-1941; Fig. 22C).

Non-type material. See Table S1.

 

Diagnosis. Shell. Large, 3.7-4.4 whorls, amber, ear-shaped, flattened, protoconch slightly raised, last whorl large; shell glossy, protoconch with fine spiral grooves, teleoconch with very fine, indistinct spiral grooves (Table 3, Figs 22C, 23E).

Animal. Orange-brown with orange-red mucus, paler on sides of neck and along tail; sole cream, slime network well-developed, tail keeled, caudal horn small. Mantle lobes moderately large, none fused, median lobe forming small cephalic shield; shell lappets moderately large, connected by a narrow collar, right lappet rounded.

Genitalia. Bursa copulatrix short, approximately equal in length to free oviduct, sac spherical. Penis large, slightly swollen at proximal end, enclosed in penial tunica, penis interior with numerous longitudinal pilasters, becoming more irregularly shaped at proximal end with longitudinal threads between them. Epiphallus very short, less than penis length, swollen with internal cryptae; enters penis through a short verge. Epiphallic flagellum with spiraling rows of internal cryptae and slender tail. Spermatophore with branching spines on base of capsule and tail-pipe, tail-pipe very short, one very long, slender spine with no accessory branches attached near the junction of capsule and tail-pipe; spermatophore golden brown, tips of branches dark brown (Fig. 28).

FIG2

Fig. 28. Reproductive anatomy of Parmavitrina disposita. A. Genitalia, AM C.165923. B. Penial interior, AM C.460261. C. Spermatophore, AM C.165923. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 4 mm, B-C = 2 mm.

Remarks. This species is found from Barrington Tops and the Mt Royal Range to the Liverpool Ranges. It is also recorded from the Comboyne Plateau (Stanisic et al., 2010) (Fig. 26).

Parmavitrina disposita can be distinguished from P. planilabris, which is similar in size, by its orange-brown body colour and complete shell. Its genital morphology differs from that of its congeners by the presence of a very short epiphallus and a large penis with a small verge and numerous longitudinal pilasters.

Parmavitrina megastoma (Cox, 1868)

Figs 22D, 24C, 29

Vitrina megastoma Cox, 1868: 87.

Helicarion megastoma: Tryon, 1885: 170, pl. 38, figs 52-53.

Parmavitrina megastoma: Iredale, 1941: 8; Smith, 1992: 240; Stanisic et al., 2010: 312-313, 331.

 

Material examined. Holotype. AM C.101145 (Clarence River, NSW).

Non-type material. See Table S1.

 

Diagnosis. Shell. Amber, moderately large, 2.5-3.0 whorls, ear-shaped, flattened; last whorl very large and flared with membraneous base; shell glossy, protoconch with fine spiral grooves, teleoconch with very fine, indistinct spiral grooves (Table 3, Fig. 22D).

Animal. Brown with purple mucus, faintly spotted, yellow line along neck, foot border edged with orange, mid field of sole slightly darker than outer fields, slime network prominent, tail with a strong, reddish keel, caudal horn small. Mantle lobes moderately large, left and median lobes fused to form a large cephalic shield; shell lappets broad, moderate size, rounded, edged with black and with faint black markings (Fig. 24C).

Genitalia. Bursa copulatrix long, more than twice length of free oviduct, sac elongate. Penis relatively slender, cylindrical; penis and part of epipahllus enclosed in a thick penial tunica. Penis interior with one longitudinal pilaster and numerous weak longitudinal ridges. Epiphallus long, more than twice penis length; epiphallis enters penis through a moderately long, slender verge. Epiphallic flagellum with spiraling rows of internal cryptae and slender tail. Spermatophore not observed (Fig. 29).

FIG2

Fig. 29. Reproductive anatomy of Parmavitrina megastoma, AM C.377461. A. Genitalia. B. Penial complex. C. Penial interior. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 4 mm, B-C = 2 mm.

Remarks. This species is listed by Stanisic et al. (2010) as ranging from Coffs Harbour to Grafton (Fig. 26). Specimens confirmed as P. megastoma by dissection or sequencing range from just north of Coffs Harbour to the Gibraltar Range and the Grafton area, in dry woodland to rainforest habitats. Parmavitrina megastoma is intermediate in size between P. maculosa and P. flavocarinata and is the only species known to exude bright purple mucus when disturbed (mucus colour is not known for P. maculosa). It is sympatric with P. flavocarinata in the Gibraltar Range National Park and Washpool National Park. It may also be sympatric with P. maculosa since the two species occur very close together just north of Coffs Harbour, P. megastoma in Bagawa State Forest and P. maculosa in Bruxner Park Flora Reserve.

Parmavitrina maculosa sp. nov.

Figs 22E, 30

Material examined. Holotype. AM C.448344 (near Coffs Harbour, Orara East SF, Bruxner Park Flora Reserve, cnr Bruxner Park Rd and Sealy Lookout road, NSW, 30° 15’ 02” S, 153° 06’ 30” E, tall open Blackbutt forest, Imperata/Themeda/Lomandra groundcover, under burnt hardwood logs, 14/9/2005).

Paratypes. AM C.532840. Same data as holotype.

Non-type material. See Table S1.

Description. Shell. Large, 2.5 whorls, golden-amber, strongly ear-shaped, flattened; last whorl extremely large and flared with membraneous base; shell glossy, very faint, indistinct spirals visible on protoconch and teleoconch (Fig. 22E).

Animal (in spirit). Grey, with prominent grey spots on tail, mantle lobes and shell lappets; mid field of sole slightly darker than outer fields, slime network prominent, tail keeled, caudal horn small. Mantle lobes large, left and median lobes fused to form a large cephalic shield; shell lappets moderate size, right lappet rounded.

Genitalia. Bursa copulatrix long, more than twice length of free oviduct, sac slender. Penis very short, penis and part of epiphallus enclosed in thin penial tunica, penis interior with longitudinal pilasters, two strong and six weaker. Epiphallus approximately 2-2.5 times penis length; enters penis through a short verge. Epiphallic flagellum with spiraling rows of internal cryptae and slender tail. Spermatophore not observed (Fig. 30).

FIG2

Fig. 30. Reproductive anatomy of Parmavitrina maculosa. A. Genitalia, AM C.532840. B. Penial interior, AM C.344211. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 4 mm, B = 1 mm.

Remarks. Parmavitrina maculosa, preliminarily identified as ‘Helicarionidae sp. NN 6’ through curatorial work, is only known from in or around Bruxner Park Flora Reserve, north of Coffs Harbour in northern NSW, in open blackbutt and moist sclerophyll forests. It is represented by very few specimens in the collections of the AM and the QM and has not been observed live, so comparative comments on its external appearance are based only on preserved specimens.

This species is larger than its congeners, with a more widely flared aperture and the shortest penis length relative to body size.

Etymology. From maculosa (Latin = spotted; adjective), referring to spots on both sides of the tail.

Parmavitrina flavocarinata sp. nov.

Figs 22F, 23F, 24D, 31

Material examined. Holotype. AM C.512373 (Washpool National Park, off Gwydir Highway, NSW, 29° 29’ S, 152° 20’ E, 23/2/2016).

Paratypes. AM C.532841 (Same data as holotype).

Non-type material. See Table S1.

Description. Shell. Moderately large, 2.6 whorls, orange-amber, ear-shaped, flattened; last whorl large and flared with membraneous base; shell glossy, protoconch with fine spiral grooves, not closely spaced, and teleoconch with very fine, faint spiral grooves (Figs 22F, 23F).

Animal. Dark brown, spotted, yellow line along neck, foot border edged with crimson, slime network moderately strong, tail with a strong, yellow keel, caudal horn small. Mantle lobes moderately small, none fused, median lobe forms small cephalic shield; shell lappets broad, moderate in size, right lappet rounded, edged with black and with black markings (Fig. 24D).

Genitalia. Bursa copulatrix moderately long, approximately 1.5 times length of free oviduct. Penis short, appearing cylindrical when enclosed in penial tunica, penial tunica thick at distal end and thin at proximal end; penis swollen at proximal end; penis interior with longitudinal, zig-zagging pilasters, two present at distal end and numerous present at proximal end. Epiphallus short, less than 1.5 times penis length; enters penis through thickened ring. Epiphallic flagellum with spiraling rows of internal cryptae and slender tail. Spermatophore with spiraling rows of branching spines present on base of capsule with a few continuing on to tail-pipe; spines split into multiple branches aligned longitudinally (Fig. 31).

FIG2

Fig. 31. Reproductive anatomy of Parmavitrina flavocarinata, AM C.532841. A. Genitalia. B. Penial complex. C. Penial interior. D. Spermatophore. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars: A = 4 mm, B-C = 1 mm.

Remarks. Parmavitrina flavocarinata, preliminarily identified as ‘Helicarionidae sp. NN7’ through curatorial work, is known from the Gibraltar Range National Park to the Girard State Forest in northern NSW (Fig. 26), living under logs in rainforest habitats, and is the smallest Parmavitrina species. It is most similar to the sympatric P. megastoma but can be distinguished by its smaller size, darker colouration with small but distinct spots and lack of purple mucus. Its shell is darker in colour and has a less widely flared aperture. Anatomically it is distinguished by its proximally swollen penis with a penial tunica that is thick distally and thin proximally, zig-zagging pilasters and the lack of a penial verge.

Etymology. From flavus (Latin = yellow; adjective) and carinata (Latin = keeled; adjective), referring to the strong yellow keel.

Cucullarion Stanisic, 1998

Cucullarion Stanisic, 1998: 597.

Type species: Cucullarion parkini Stanisic, 1998, by original designation; masculine

 

Diagnosis. Shell. Medium sized, reduced, ear-shaped, flattened; base and internal whorls completely membraneous, dorsal calcified area plate-like; shell glossy, no sculpture.

Animal. Beige to orange- or red-brown; caudal horn small. Mantle lobes large, left and median lobes fused to form cephalic shield; shell lappets large, fused at both sides, covering most of shell (Fig. 32A-B).

FIG2

Fig. 32. Live specimens of Cucullarion, Peloparion and Ubiquitarion gen. nov. A. Cucullarion parkini, AM C.512610, Lamington National Park. B. C. albimaculosum, AM C.103209 (photographer: G. Millen). C. Peloparion helenae, MV F219249, Barrington Tops National Park (photographer: A. Moussalli). D. Ubiquitarion iridis, AM C.512394, Pikapene National Park. Scale bars = approx. 10 mm.

Gentitalia. Ovotestis of 3-4 lobes grouped together and embedded in tip of digestive gland. Talon and carrefour embedded in albumen gland. Spermoviduct folded 5 times. Free oviduct long with no internal sculpture; capsular gland small, spherical. Bursa copulatrix moderately long, reaching two thirds of the way along spermoviduct, consisting of swollen base, slender duct and elongate sac; internally with longitudinal pilasters; inserted on vagina. Vagina very short. Penial tunica tightly attached to epiphallus by muscle fibres; epiphallus enters penis through verge; epiphallic caecum absent; epiphallic flagellum with axial filament present; single internal crypt present in part of epiphallus adjacent to flagellum but not in flagellum. Spermatophore a soft-walled capsule with hard tail-pipe; one spine present at junction of capsule and tail-pipe; tail-pipe sculptured with a spiraling row of tiny teeth.

Remarks. Cucullarion was introduced for a semislug from southern Queensland, C. parkini (Stanisic, 1998) and was later expanded to include a second species, C. albimaculosum (Stanisic et al., 2010). The two species are notable for their degenerate, plate-like shells with membranous sides, bases and internal whorls.

Cucullarion parkini Stanisic, 1998

(Figs 32A, 33A-B)

Cucullarion parkini Stanisic, 1998: 598, figs 1-4; Stanisic et al., 2010: 314-315, 332.

 

Material examined. Holotype. QMMO60128 (The Knoll NP, Tamborine Mtn, SE QLD, 27° 55’ S, 153° 11’ E, leg. Stanisic et al., 26/2/1985).

Non-type material. See Table S1.

 

Diagnosis. Shell. Medium sized, 2.8 whorls, yellow-amber, ear-shaped, flattened; base and internal whorls completely membraneous, dorsal calcified area plate-like; shell glossy, no sculpture.

Animal. Beige with reddish-brown colouring deepening on tail and neck, speckled with tiny spots of yellow pigment, faintly spotted with darker brown; sides of sole striped; tail with a strong reddish-brown keel; caudal horn small. Mantle lobes large, left and median lobes fused to form cephalic shield; shell lappets large, fused at both sides, covering most of shell, edged with reddish brown and with reddish brown longitudinal ridges (Fig. 32A).

Gentitalia. Bursa copulatrix long, swollen at base, duct slender, sac elongate. Penis moderate size, cylindrical, completely encased in penial tunica; penis interior with one raised penial pilaster and one smaller penial pilaster, each leading up to a rounded penial pilaster proximally; both pilasters pustulose; remainder of penis interior with longitudinal ridges becoming pustulose proximally. Epiphallus short, approximately penis length, enters penis through short verge of about a quarter penis length. Epiphallic flagellum moderately long, slender, without internal cryptae; one crypt present in adjacent part of epiphallus. Spermatophore with elongate capsule and long, thin tail-pipe sculptured with a single row of tiny teeth; a single short spine present at junction of tail-pipe and capsule (Fig. 33A-B).

FIG2

Fig. 33. Reproductive anatomy of Cucullarion. A-B. C. parkini, AM C.512610 A. Genitalia. B. Penial interior. C-D. C. albimaculosum, QM MO25791. C. Penial interior. D. Genitalia. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens. Scale bars = 2 mm.

Remarks. Cucullarion parkini was previously known only from The Knoll NP in SE QLD. Here we extend the range to include Binna Burra in Lamington NP. This rainforest species has only ever been found a handful of times, despite relatively frequent collecting in these locations. This may be due to its highly reduced shell and slender body, which would allow it to hide deeply in the base of palm trees (individuals have been observed emerging from palm trunks during rainstorms) or other crevices. However, it is also likely that this species is present in very low numbers.

The congener of this species, Cucullarion albima­culosum, is also found in Binna Burra, although the two species have not been collected together. Cucullarion parkini can be distinguished from C. albimaculosum by its slightly smaller size and paler colour, and by the lack of white pigment on its body. Anatomically, the two species differ in the length of the flagellum and the penis interior.

Cucullarion albimaculosum Stanisic, 2010

(Figs 32B, 33C-D)

Cucullarion albimaculosa Stanisic, 2010 (in Stanisic et al., 2010): 314-315, 332.

 

Material examined. Holotype. QMMO17249 (Collins Ck crossing, Border Ranges NP, N NSW, 28°26’ S, 153°09’ E, on trees, leg. Stanisic and Potter, 10/3/1987).

Non-type material. See Table S1.

 

Diagnosis. Shell. Medium sized, 3.0 whorls, amber, ear-shaped, flattened; base and internal whorls completely membraneous, dorsal calcified area plate-like; shell glossy, no sculpture (Table 3).

Animal. Orange brown with spots of thick white pigment on tail, mantle lobes and shell lappets; tail keeled; caudal horn small. Mantle lobes large, left and median lobes fused to form cephalic shield; shell lappets large, fused at both sides, covering most of shell (Fig. 32B).

Genitalia. Bursa copulatrix long, swollen at base, duct slender, sac elongated. Penis moderate size, cylindrical, completely encased in penial tunica; penis interior with diamond shaped pustules and four longitudinal pilasters at distal end. Epiphallus moderately short, approximately 1.5 times penis length, enters penis through short verge of about one fifth to one third penis length. Epiphallic flagellum very long, slender, without internal cryptae, tip attached by connective tissue to interior of left body wall; one crypt present in adjacent part of epiphallus. Spermatophore not observed (Fig. 33C-D).

Remarks. This species is found in the Border Ranges NP (NE NSW) and Lamington NP (SE QLD) (Fig. 34), from arboreal habitats in subtropical rainforest. It has been collected more frequently than C. parkini, despite its similarly cryptic nature. Cucullarion albimaculosum can be distinguished from other semislugs by the white pigment spots present on the tail, mantle lobes and shell lappets.

FIG2

Fig. 34. Occurrence records of Cucullarion, Peloparion, and Ubuiquitarion from the malacological collection of the Australian Museum, Sydney. Symbols: ● = U. iridis,  = P. helenae, + = C. albimaculosum, □ = C. parkini.

Peloparion Iredale, 1937

Peloparion Iredale, 1937a: 8.

Type species: Helicarion helenae Godwin-Austen, 1883, by subsequent designation of Iredale (1941); masculine

 

Diagnosis. Shell. Small, ear-shaped, flattened, last whorl large, umbilicus closed, shell glossy, protoconch with fine spiral grooves, teleoconch smooth (Table 3, Fig. 35).

FIG2

Fig. 35. Shells of Peloparion and Ubqiuitarion. A. Peloparion helenae: holotype of Peloparion submissus Iredale, 1941, AM C.101142 (size indicative; photographer: AM). B. Ubiquitarion iridis holotype AM C.456552. Scale bar = 3 mm.

Animal. Pale grey with bright red mucus. Mantle lobes and shell lappets well developed.

Genitalia. Ovotestis of four lobes, embedded in digestive gland. Talon and carrefour embedded in albumen gland. Spermoviduct folded twice. Free oviduct very long with no internal sculpture; capsular gland not externally visible. Bursa copulatrix moderately short, reaching less than halfway along spermoviduct; internally smooth; inserted on vagina. Vagina very short. Epiphallus enters penis through moderately long verge; epiphallic caecum absent; epiphallic flagellum with axial filament present; numerous internal cryptae present in part of epiphallus adjacent to flagellum but not in flagellum. Spermatophore a soft-walled capsule with hard tail-pipe; branching spines present on capsule; tail-pipe with only a single spine.

Remarks. Peloparion was originally introduced as a subgenus of Helicarion for Vitrina hyalina Pfeiffer, 1855 (Iredale, 1937c), but subsequently Iredale (1941) stated that the name was intended for Helicarion helenae Godwin-Austen, 1883. The genus was revised by Hyman (2007) and Hyman and Ponder (2010).

Peloparion helenae (Godwin-Austen, 1883)

(Figs 32C, 35A, 36)

Helicarion helenae Godwin-Austen, 1883: 146, pl. 41, figs 1-8; Tryon, 1885: 170, pl. 38, figs 50-51; Cox, 1909: 6.

Peloparion submissus Iredale, 1941: 7, fig. 8; Smith, 1992: 240.

Peloparion helenae: Iredale, 1941: 7; Smith, 1992: 240; Hyman and Ponder, 2010: 42; Stanisic et al., 2010: 310-311, 330.

 

Material examined. Holotype (Peloparion submissus). AM C.101142 (Scone, NSW, 32°3’ S, 150°52’ E, pre-1941; Fig. 35A).

Non-type material. See Table S1.

 

Diagnosis. Animal. Pale grey with faint spots on sides of body and lappets; appearing red from mucus. Sole uniformly pale, foot border red; caudal horn large, edged with dark grey. Slime network visible. Mantle lobes and shell lappets well developed, lappets stained red and joined by a narrow collar; right lappet moderately large, rounded, edged with black, with a raised ridge on lower side; left lappet slightly shorter, rounded at tip, with a raised ridge on lower side (Fig. 32C).

Genitalia. Bursa copulatrix moderately short, sac tear-shaped and distinguishable from duct. Penis cylindrical, fully enclosed in penial tunica, internally with two main longitudinal pilasters broken up at proximal end into several smaller zigzagging threads; additional smaller longitudinal pilasters present; transverse ridges at proximal end; epiphallus moderately short, less than twice penis length, with numerous internal cryptae adjacent to flagellum; entering penis through thick, moderately long verge sculptured with transverse ridges. Spermatophore with approximately six branching spines present on capsule in a spiraling pattern and one spine on tail-pipe (Fig. 36).

FIG2

Fig. 36. Reproductive anatomy of Peloparion helenae, AM C.456546. A. Genitalia, scale bar = 4 mm. B. Penial interior, scale bar = 2 mm. C. Spermatophore, scale bar = 1 mm. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens.

Remarks. We follow Hyman (2007) and Hyman and Ponder (2010) in recognising the synonymy of Peloparion helenae and P. submissus, based on an examination of the type material.

Peloparion helenae is only known from Barrington Tops are in the Hunter Valley, living arboreally in beech forest and snow gum woodland. It is easily distinguished from other helicarionid semislugs by its small size and bright red mucus. It was previously grouped with Ubiquitarion iridis, another small semislug with rounded, dark-edged shell lappets. However, while some elements of the genitalia appear to be quite similar, there are considerable differences in the penial complex and spermatophore. In Peloparion helenae there are numerous internal cryptae in the epiphallus but none in the flagellum, giving rise to a spermatophore with long branching spines on the capsule but very few on the tail-pipe. In contrast, U. iridis has a thick muscular tunica obscuring the flagellum and adjacent part of the epiphallus, and a spermatophore with only a single branching spine on the capsule but a double row of numerous short, branching spines on the tail-pipe. The two species are also not grouped together based on mitochondrial DNA sequences, and as a consequence we have erected a new genus for Peloparion iridis. Peloparion helenae appears to be most closely related to Mysticarion, but is too anatomically divergent to be included in this group.

Ubiquitarion gen. nov.

Type species: Peloparion iridis Hyman, 2007

Description. Shell. Small, ear-shaped, flattened, last whorl large, underside membranous, umbilicus closed, shell glossy, protoconch and teleoconch with no visible sculpture (Table 3, Fig. 35B).

Animal. Beige with yellowish and black markings and iridescent specks of pigment. Mantle lobes and shell lappets well developed.

Genitalia. Ovotestis of one lobe, embedded near apex of digestive gland. Talon and carrefour embedded in albumen gland. Spermoviduct folded three times. Distal portion of free oviduct with small spherical capsular gland with no internal sculpture, remainder of free oviduct with longitudinal pilasters internally. Bursa copulatrix moderately long, reaching halfway along spermoviduct; internally with longitudinal pilasters (duct only); inserted on vagina. Vagina short, internally with longitudinal pilasters. Epiphallus enters penis through thickened ring; epiphallic caecum absent; descending arm of epiphallus (near flagellum) encased in a thick muscular sheath; epiphallic flagellum with axial filament present, also encased in thick muscular sheath. Spermatophore a soft-walled capsule with hard tail-pipe; one branching spine present on capsule; double row of branching spines present in spiraling pattern along tail-pipe.

Remarks. This genus is created for Peloparion iridis. This species is superficially similar to Peloparion helenae, but differs anatomically, particularly in the penial complex and spermatophore. Mitochondrial DNA sequences also show that the two species do not group together.

Etymology. Combination of ubique (Latin = ‘everywhere’; arion (Latin) referring to a ‘kind of snail or slug’; masculine.

Ubiquitarion iridis (Hyman, 2007) comb. nov.

(Figs 32D, 35B, 37)

Helicarion helenae; Godwin-Austen, 1883: 146, pl. 41, fig. 1 [live drawings only] (not Godwin-Austin, 1883: 146, pl. 41, figs 1-8).

Peloparion iridis Hyman, 2007: 90; Stanisic et al, 2010: 279, 310-311.

 

Material examined. Holotype. AM C.456552 (Sydney Harbour, Elizabeth Bay, NSW, pre-1877).

Non-type material. See Table S1.

 

Diagnosis. Animal. Beige, with a paler stripe along tail with dark stripes to either side. Mid field of sole cream; outer fields grey. Mantle lobes and shell lappets well developed, right lappet rounded and edged with black; left lappet short, elongate with rounded tip, bisected by black line and raised cream-coloured ridge; black spots on lobes and lappets (Fig. 32D).

Genitalia. Bursa copulatrix moderately long, sac oval-shaped and distinguishable from duct. Penis cylindrical, fully enclosed in penial sheath, internally with two main longitudinal pilasters and additional shorter longitudinal pilasters at proximal end, connected by transverse threads; epiphallus long, between two and three times penis length, entering penis through thickened ring; descending arm of epiphallus and epiphallic flagellum encased in a thick muscular sheath. Spermatophore a soft-walled capsule with hard tail-pipe; one branching spine present on capsule; double row of branching spines present in spiraling pattern along tail-pipe (Fig. 37).

FIG2

Fig. 37. Reproductive anatomy of Ubiquitarion iridis, AM C.512399. A. Genitalia, scale bar = 2 mm. B. Penial interior, scale bar = 1 mm. Abbreviations: alb, albumen gland; bc, bursa copulatrix; cg, capsule gland; ep, epiphallus; fl, flagellum; her, hermaphroditic duct; ov, ovotestis; p, penis; pp, penial pilasters; pr, prostate; prm, penial retractor muscle; pt, penial tunica; pv, penial verge; ut, uterus; vd, vas deferens.

Remarks. Two species of Peloparion were accepted for many years (Iredale, 1941; Smith, 1992; Smith et al., 2002), Peloparion helenae for specimens ranging from Sydney to southern Queensland and Peloparion submissus for specimens from Barrington Tops, until the discovery that the type specimens of both were identical (Hyman, 2007). The names were synonymised with the senior name, Peloparion helenae being retained for the Barrington Tops species and a new name, P. iridis, introduced for the more widely ranging species (see Hyman, 2007 for a more detailed account).

Given its unusually wide range, spanning over 750 km from Sydney to north of Brisbane in habitats ranging from dry vine thicket to rainforest, this species was extensively sampled in order to ensure that it was not a species complex. Both molecular and morphological investigations confirmed the presence of a single widespread species, with a disjunct range including populations around Sydney (where this species is thought to have been introduced around 1865; Hyman, 2007), a single known locality at Seal Rocks in central NSW and then populations ranging between Grafton in northern NSW and Gympie in southern QLD (thought to be the original range of the species; Hyman, 2007).