taxonID	type	description	language	source
03E487A3FF98C477FF01B2BBFCA5FD67.taxon	diagnosis	Diagnosis: Shell present, complete or reduced, 5 – 35 mm 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 upcurled 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 a 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: The Helicarionidae is distributed in Australia, islands of the Pacific, Southeast Asia, Madagascar and the Mascarene Islands. It is grouped in superfamily Helicarionoidea, along with the Asian Ariophantidae and the African Urocyclidae. The three families are unified by the presence of a flagellum with an axial filament, an epiphallic caecum, and mantle lobes (Hausdorf, 1998; Hyman & Ponder, 2010); however, the flagellum and epiphallic caecum are absent in some members of all three groups. The only character that reliably distinguishes Helicarionidae from the other two families is the presence of a penial tunica that is open at the proximal end rather than being fused to the penis (Hausdorf, 1998; Hyman & Ponder, 2010).	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF98C476FC34B480FA5BFED7.taxon	diagnosis	Differential diagnosis External morphology: Small to medium-sized shell, ear shaped, flattened, thin, golden, sometimes with a greenish tinge, glossy, 3.2 – 3.9 whorls, whorls rounded, base membraneous. Protoconch with fine spiral grooves, teleoconch with very fine spiral grooves. Body colour varying from black to pale grey, fawn or cream; sole often a contrasting colour to body. Mantle lobes and shell lappets of moderate size, none fused; mucous network prominent; caudal horn well developed. Genital anatomy: Ovotestis of two to four lobes, embedded in digestive gland. Talon and carrefour embedded in albumen gland. Spermoviduct curves in a U shape towards tail then folds behind and descends towards head. Free oviduct with indistinct capsular gland in proximal portion; internal walls of capsular gland smooth; remainder of free oviduct with longitudinal pilasters. Bursa copulatrix moderately long, approximately half spermoviduct length; sac portion often tear shaped with elongated tip; inserted on free oviduct. Vagina short. Penis long, slender, with internal sculpture of V-shaped rows of papillose lamellae. Penis tunica attached by muscle fibres to middle of epiphallus; epiphallus enters penis through simple pore; epiphallic caecum absent; epiphallic flagellum with axial filament present, containing spiralling rows of internal cryptae. Spermatophore a soft-walled capsule with hard tail-pipe; branching spines present in spiralling pattern along tail-pipe. Remarks	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF99C469FC28B72AFAE6F8B3.taxon	description	(FIGS 6 – 18)	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF87C464FF06B0E3FB99FA86.taxon	materials_examined	Material examined Types: Neotype of H. cuvieri: MNHP no number (Needles picnic ground, S of Strathgordon Road, SW TAS) (Kershaw, 1979). Lectotype of Vitrina nigra: MNHP no number (Western Port, Vic) (Kershaw, 1981). Holotype of H. leopardinus: AM C. 101141 (Ourimbah Scrubs, NSW). Holotype of H. mastersi callidus: AM C 101138 (Twofold Bay, NSW). Non-type material: See Supporting Information (Table S 1). Description External morphology: Shell greenish gold to orange-brown, 2.7 – 3.5 whorls. Body colour variable, ranging from cream to black, often speckled with orange, cream or brown. Shell lappets sometimes dark bordered, sometimes with two to three pigmented warts. Tail with a moderately strong keel, usually paler than body colour. Genital anatomy: Penis generally slender, tubular, of variable length, occasionally slightly swollen proximally. Penial interior sculptured with nine to 50 rows of papillose ridges arranged in a deep V shape. Penis-to-epiphallus ratio variable, 0.18 – 1.31; epiphallus entering penis apically or laterally. Flagellum with internal cryptae. Spermatophore variable, with nine to 20 spines. Remarks Helicarion cuvieri has previously been recorded only from Tasmania, where it is widespread (Fig. 6). Here we expand the delineation of this species to include populations from eastern Victoria and southeastern NSW. This species and its congener, H. mastersi, are both highly variable in body size and colour, making it difficult to distinguish them reliably. Helicarion cuvieri generally has a uniform body and sole colour in darker shades ranging from grey to orange-brown and black, with dark eye tentacles. In specimens with a contrasting pale sole (clades E, F and G), the tail tip often has a dark vertical stripe. Three pigmented warts are often (but not always) present on the shell lappets, and the tail has a longer keel. Helicarion mastersi has no pigmented warts, generally paler coloration, with pale eye tentacles in shades of cream, pink, grey and brown, deepening on the tail, and has a pale sole contrasting with its body colour. The two species can be distinguished morphologically by the spacing of the internal penial lamellae; H. cuvieri has fewer, coarser lamellae arranged in a deep V shape, whereas H. mastersi has more numerous, finer lamellae arranged in a shallower V shape.	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF87C464FF06B0E3FB99FA86.taxon	description	Within H. cuvieri there are eight morphologically distinct subclades (clades A – H), which are morphologically distinct; these are described below. specimens with 13 – 16 spines, formula (2, 2, 9, 3); Brown Mountain specimens with 20 spines, formula (4, 11, 1, 4) (Figs 10, 11 A – C). Remarks Clade A comprises all Tasmanian specimens along with one population from the Otway Ranges in Victoria and another population from Brown Mountain in NSW. There is a wide range of colour variation seen in clade A, including the only known orange forms of Helicarion. The two geographically isolated populations differ anatomically from the rest of the clade. Semislugs from the Otway Ranges have a much simpler spermatophore, with nine simple spines, in contrast to the ten to 16 more complex spines seen in Tasmanian specimens. Likewise, specimens from Brown Mountain in NSW have a much more complex spermatophore than Tasmanian specimens, with 20 spines in total, including a higher number of branching spines; this population also has a slightly longer penis. These differences probably reflect the isolation of these geographical areas. There may also be a connection with altitude, because both regions have a relatively high elevation. Members of this clade are microsympatric with Attenborougharion rubicundus in southeastern Tasmania but can be distinguished by their smaller size and generally uniform cream to dark brown coloration, in contrast to the two-toned burgundy and green coloration of A. rubicundus. CLADE A (FIGS 6, 7 A, 8 A – C, 9, 10, 11 A – C) Diagnosis External morphology: Shell golden amber, 2.7 – 3.5 whorls (Figs 7 A, 8 A – C). Body cream, orange, grey or dark brown, often speckled with orange, cream or brown. Eyestalks blue-grey to black. Shell lappets dark bordered, with three pigmented warts, two on right lappet, one on left lappet, with black pigmentation in grey specimens and orange pigmentation in orange-brown specimens. Tail with a moderately strong keel, usually paler than body colour (Fig. 9). Genital anatomy: Penis medium length, tubular, occasionally slightly swollen proximally. Penial interior with ~ 14 – 16 lamellae. Penis-to-epiphallus ratio of 0.41 – 0.84; epiphallus enters penis apically or slightly laterally. Spermatophore variable; Otway Ranges specimens with nine spines, formula (0, 7, 2, 0); Tasmanian CLADE B (FIG. 12) Diagnosis External morphology: Shell golden amber, 3.1 whorls. Body and sole a uniform dark grey (in alcohol). Shell lappets darker, black bordered, with three pigmented warts, two on right lappet, one on left lappet. Tail keeled. Fig. 12 Genital anatomy: Penis moderately long, tubular, slightly swollen proximally. Penial tunica covering approximately four-fifths of penis. Penial interior with ~ 12 lamellae. Penis-to-epiphallus ratio of 0.53; epiphallus enters penis slightly laterally. Spermatophore with 15 spines, formula (3, 5, 5, 2). Remarks Clade B is represented by only a single specimen from Alpine NP in Victoria. This specimen is distinguished by its very dark body colour, the presence of pigmented warts on its shell lappets, and the moderately long penis with very few penial lamellae. Further collecting in the Alpine NP and adjacent high-altitude parts of Victoria and southern NSW is necessary to understand this clade better. SPECIES LIMITS IN HELICARION SEMISLUGS 955 CLADE C (FIGS 6, 7 B, 8 D – F, 13 A, 14) Diagnosis External morphology: Shell golden, 3.3 whorls (Figs 7 B, 8 D – F). Body and sole dark grey-brown (in alcohol); in life with a speckled appearance. Lappets narrow, lacking pigmented warts, dark bordered. Left lappet with dark streaks on inside. Mid field of sole slightly paler (Fig. 13 A). Genital anatomy: Penis long, slender, proximally swollen, swollen portion folded over. Penial tunica covering approximately three-quarters of penis. Penial interior with ~ 25 lamellae. Penis-to-epiphallus ratio of 0.74; epiphallus also folded. Spermatophore of 12 spines, formula (1, 4, 6, 1) (Fig. 14). Remarks Clade C is known only from Mitchell River NP (Fig. 6). No material was available to measure; however, based on the limited material available members of this clade appear to be relatively small, similar in size to members of clade G from the Blue Mountains. Clade C semislugs can be distinguished from other clades of H. cuvieri by their longer penis and epiphallus, both of which are folded. CLADE D See clade H below. CLADE E (FIGS 6, 7 C, 8 G – I, 11 D – F, 13 B, C, 15) Diagnosis External morphology: Shell amber to orange-brown, 2.8 – 3.4 whorls (Fig. 7 C, 8 G – I). Body grey or pale brown to black, colour deepening on tail and neck, with a vertical dark stripe at tail tip; sole generally paler than body. Shell lappets with three pigmented warts in Victorian populations (Fig. 13 B, C). Genital anatomy: Penial interior of few (nine to 13) lamellae arranged in deep V-shaped rows. Penis size and epiphallus entry variable: penis-to-epiphallus ratio of 0.44 – 0.50, epiphallus entering penis apically (Victorian populations and Flinders Island) ’ penis-to-epiphallus ratio of 0.18 – 0.36, epiphallus entering laterally (Nadgee NR, Burragorang, Mt Kelgoola and Mt Coricudgy); or penis-to-epiphallus ratio of 0.83, epiphallus entering laterally (Wyong, Wombeyan Caves). Flagellum with prominent internal cryptae, corresponding to ten to 13 robust branching spines on spermatophore; spermatophore formula (2 – 3, 4 – 10, 0 – 7, 0 – 2) (Figs 11 D – F, 15). Remarks Clade E includes material from Victoria, including the type locality of H. niger. However, H. niger as previously understood contained all Victorian populations and Flinders Island (Tas). In contrast, clade E does not three pigmented warts on their shell lappets and in the presence of a medium-length penis with the epiphallus entering apically. Populations from Nadgee Nature Reserve (the type locality of H. mastersi callidus), Burragorang, Mt Kelgoola and Mt Coricudgy all have a very short penis, the shortest observed in Helicarion to date, with the epiphallus entering slightly laterally. Finally, specimens from Wyong and Gosford (representing H. leopardinus) and from Wombeyan Caves have a relatively long, broad penis with a slightly lateral epiphallus entry point. These differences are significant; however, there is only slight genetic differentiation, and one of the three groups thus delimited (the short penis group) is paraphyletic. Furthermore, there are some strong characters uniting the three groups, including a penial interior with a small number of relatively large internal lamellae, and the presence of a robust spermatophore with 13 – 16 spines, most highly branched. It is also possible that the penial differences are attributable to reinforcement, because the clades exhibiting a very short penis are all in close proximity to another clade (see Taxonomic Preamble). Kershaw (1981) described a small penial papilla in H. niger; this has not been observed in any specimens in the present study. contain all Victorian populations; in fact, from Victoria, only populations from Wilsons Promontory (the type locality) and the Central Highlands are included. Several populations from south-eastern NSW are also included (Fig. 6). The resulting distribution, the largest of any H. cuvieri subclade, contains several large gaps, notably between Wilson’s Promontory in Victoria and Nadgee Nature Reserve in NSW, and between Nadgee Nature Reserve and Wombeyan Caves. It is probable that further collecting will extend the known range of clade E still further and may somewhat reduce the size of the gaps in its distribution. In addition to the largest distribution, this taxon also exhibits the most anatomical variation seen in any Helicarion clade. The Victorian populations (including Flinders Island; equivalent to H. niger s. s. and forming a monophyletic group) are significantly larger than the NSW specimens, and also differ in exhibiting CLADE F (FIGS 6, 7 D, 8 J – L, 11 G – I, 16 A – C, 17) Diagnosis External morphology: Shell golden amber, 3.0 – 3.5 whorls (Figs 7 D, 8 J – L). Body cream to pale grey or brown, colour deepening on tail and neck, with a white to pale pink sole. Shell lappets with one to three pigmented warts (one or two on right lappet, one on left), less visible in pale specimens (Fig. 16 A – C). Genital anatomy: Penis long, slender, penis-to-epiphallus ratio of 0.29 – 0.87, epiphallus entering laterally or apically. Penial interior of 12 – 21 lamellae arranged in deep V-shaped rows. Spermatophore of 11 – 17 branches, with a gap after the second or third branch (Figs 11 G – I, 17). Remarks Clade F is distributed in southeastern NSW and eastern Victoria. Members of this clade are very similar in external morphology to members of the neighbouring clade E and to pale specimens of clade A. All three taxa are large semislugs with two or three pigmented warts on their shell lappets. Anatomically, semislugs belonging to clade F can be distinguished from members of clade E by their longer penis relative to the epiphallus, with a larger number of penial lamellae. This clade contains four well-supported, morphologically distinct subclades. The major differences lie in the spermatophore and flagellum, particularly in the presence of a large gap in the pattern of spermatophore spines in populations from Mt Gulaga and Tallaganda NP, with some variation also present in the penis shape and length. However, the genetic distances between the subclades are low and indicative of their rather recent evolutionary origin and potentially incomplete lineage differentiation. We are also unsure whether the anatomical differences are truly indicative of reproductive incompatibility. CLADE G (FIGS 6, 7 E, 8 M – O, 16 D, E, 18) Diagnosis External morphology: Shell golden, 3.2 – 3.6 whorls (Figs 7 E, 8 M – O). Body cream to dark grey, darker on tail; lappets dark edged, lacking pigmented warts (Fig. 16 D, E). Genital anatomy: Penis very long, slender, swollen medially; proximal end similar in width to epiphallus; penis-to-epiphallus ratio of 1.08 – 1.31, epiphallus entering penis laterally through simple pore. Epiphallus with an extra fold. Interior of penis with numerous lamellae (~ 50), slightly deeper and more folded in narrow proximal portion. Bursa copulatrix with a very long duct, duct longer than bursa. Spermatophore with 14 evenly spaced branching spines of decreasing complexity; formula (1, 9, 2, 2) (Fig. 18). Remarks This clade is distributed in the Blue Mountains NP, including Mt Wilson, Mt Tomah Botanic Gardens and Katoomba (Fig. 6). Specimens from Jenolan Caves probably also belong to this clade. A separate taxon, clade E, is found further south in Burragorang. Members of clade G are significantly smaller than members of clade A and H. mastersi. These semislugs can be distinguished from other subclades of H. cuvieri by their extremely long penis, with a distinctive shape and relatively numerous penial lamellae. CLADE H (FIGS 6, 7 F, 8 P – R, 16 F, 19) Diagnosis External morphology: Shell greenish gold, 3.0 – 3.4 whorls (Figs 7 F, 8 P – R). Body grey, with deeper grey-black wrinkles and black eyestalks. Shell lappets and mantle lobes speckled grey, dark bordered, without pigmented warts. Sole a uniform grey (Fig. 16 F). Genital anatomy: Penis long, slender, subdivided into two portions, with distal portion covered by penial tunica. Penial interior with wavy, pustulose lamellae arranged in a V shape in distal portion; proximal portion with deep, distinct, less pustulose lamellae arranged in much deeper V shape (close to longitudinal). Total number of lamellae ~ 14 – 16. Penis-to-epiphallus ratio of 0.79; two arms of epiphallus twisted around one another. Flagellum with internal cryptae. Spermatophore of 19 spines, formula not recorded, most spines with multiple branches (Fig. 19). Remarks Clade H is known only from the Brindabella Range (Fig. 6). Helicarion specimens have also been collected at other sites in and around the ACT, including Tidbinbilla Nature Reserve, Namadgi National Park and Tantangara Mountain, but the identity of these specimens has not been confirmed. Specimens from Wee Jasper at the northern tip of the Brindabella Range are very similar in external morphology and in genital morphology, with the same overall pattern of a two-chambered penis but with a shorter penial complex and fewer spines on the spermatophore. This degree of variation is consistent with the population-level differences observed in other clades, and it is likely that the Wee Jasper population forms part of clade H despite not grouping together on the molecular tree. Members of clade H are unique in their grey speckled appearance and greenish golden shell. They are significantly smaller than all other Helicarion semislugs. Their penial anatomy is highly distinct, particularly in the presence of very deep, nearly longitudinal lamellae in the proximal portion of the penial interior. The large number of spermatophore spines also helps to distinguish this clade.	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF8BC458FC18B3DEFECFF891.taxon	description	(FIGS 6, 20 – 24)	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF8BC458FC18B3DEFECFF891.taxon	materials_examined	Material examined Holotype: AM C. 101139 (Kiama, 34 ° 40 ′ S, 150 ° 51 ′ E, leg. G. Masters, pre- 1868). Paratypes: AM C. 103648 (same data as holotype). Non-type material. See Supporting Information (Table S 1). Description External morphology: Shell golden brown to pale amber, 3.1 – 3.7 whorls. Body cream, greyish or fawn, with pink to orange-brown pink coloration, deepening on the tail; sole cream. Mantle lobes and shell lappets cream, sometimes lined with grey, with dark markings on underside. Genital anatomy: Penis tubular, sometimes slen- der with a small bulge at proximal end, sometimes broad and slightly curving; epiphallus 1.5 – 3 times penis length, entering penis either apically or laterally (leaving a small blind tip) through a simple pore. Penial interior with> 20 closely spaced lamellae. Spermatophore with nine to 14 evenly spaced spines (N = 5) (in specimens from Kangaroo Valley, first branch is separated from remainder by a small gap). Remarks This species is delimited herein in a different manner from previous taxonomic studies, in that several populations from throughout NSW previously subsumed under the name H. mastersi are now recognized as members of H. cuvieri. This also applies to the two nominal species-taxa ‘ H. leopardinus ’ and ‘ H. callidus ’ introduced by Iredale (1941). See the description of H. cuvieri above for features that can be used to distinguish H. cuvieri and H. mastersi. Helicarion mastersi is found from Kangarooo Valley in southern NSW to southern Sydney (Fig. 6). This species encompasses three well-differentiated subclades (I, J and K), described below. Differences between these three subclades include size and coloration, the presence or absence of a small blind penis tip and a varying number of spines on the spermatophore. To clarify the distinct status of these three forms further, denser sampling throughout the Upper Nepean dam catchment region and the Kangaroo Valley is necessary, particularly along potential zones of contact.	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
03E487A3FF8BC458FC18B3DEFECFF891.taxon	description	CLADE I (FIGS 6, 20 C, 21 G – I, 22 C, 23 C, F, I, 24 B) Diagnosis External morphology: Shell golden brown to pale amber, 3.1 – 3.5 whorls (Figs 20 C, 21 G – I). Body greyish or orange brown, darkening on the tail; sole cream. Shell lappets often lined with grey, with dark markings on underside (Fig. 22 C). Genital anatomy: Penis broad; penis-to-epiphallus ratio of 0.67 – 0.89, epiphallus entering penis laterally through simple pore. Spermatophore with nine to 11 evenly spaced branching spines of decreasing complexity; formula (1, 2, 6 – 8, 0 – 2) (Figs 23 C, F, I, 24 C). Remarks The range of this clade stretches from Macquarie Pass to southern Sydney (Fig. 6). The phylogenetic tree shows two clear groups: one from Razorback and Mt Kembla, and the other from Royal NP and Bulli Pass, but there are no significant anatomical differences. Members of clade I can be distinguished from other Helicarion by their smaller size (they are similar in size to members of clades G and H but smaller than the other members of the genus) and from other NSW taxa by their grey-brown to orange-brown coloration. Like semislugs belonging to clades J and K, they have an interior penial sculpture of numerous rows of fine lamellae, but differ from these taxa in their broader penis, with a slightly lateral entry point, and spermatophore with fewer, less highly branched spines. CLADE J (FIGS 6, 20 B, 21 D – F, 22 B, 23 B, E, H, 24 B) Diagnosis External morphology: Shell golden, 3.2 – 3.7 whorls (Figs 20 B, 21 D – F). Body cream, with pale yellowish brown coloration on tail; sole cream. Mantle lobes and shell lappets pale (Fig. 22 B). Genital anatomy: Penis slender, tubular, with small bulge at proximal end; penis-to-epiphallus ratio of 0.40 – 0.52, epiphallus entering penis apically through a simple pore. Spermatophore with 13 – 14 branching spines; formula (1 – 2, 12, 0, 0); first branch separated from remainder by a small gap (Figs 23 B, E, H, 24 B). Remarks Clade J is known only from near Bellawongarah, north of Nowra (see Fig. 6). The majority of specimens have been collected along Kangaroo Valley Road as it crosses Berry Mountain. Very little material exists, reflecting the lack of collecting in this area; additional collecting might reveal a broader range. Specimens belonging to clade J are similar in size to those of clade K but are slightly flatter in shape and can be distinguished by their paler body colouring. Individuals are similar in appearance to those belonging to clade F (part of H. cuvieri) but can be distinguished by the lack of pigmented warts on the shell lappets. Members of this taxon have a unique genital anatomy of a relatively short penis (penis-to-epiphallus ratio 0.4 – 0.52) with a small swelling at the proximal end and no blind tip, and a robust spermatophore with 13 – 14 heavily branched spines and a gap between the first and second spines. CLADE K (FIGS 6, 20 A, 21 A – C, 22 A, 23 A, D, G, 24 A) Diagnosis External morphology: Shell golden, 3.2 – 3.7 whorls (Figs 20 A, 21 A – C). Body fawn, with pink coloration, deepening on the tail; sole cream (Fig. 22 A). Genital anatomy: Penis broad, curved, with small blind tip; penis-to-epiphallus ratio of 0.50 – 0.81, epiphallus entering penis laterally through a simple pore. Spermatophore with 11 evenly spaced spines (N = 4), formula (0, 3, 7, 1) (Figs 23 A, D, G, 24 A). Remarks Clade K is found in and around Kiama (the type locality of H. mastersi), at localities including M i n n a m u r r a N P a n d B a r r e n G r o u n d s N a t u r e Reserve (Fig. 6). Its nearest relatives are distributed close by (clade I in Macquarie Pass to the north and clade J in Kangaroo Valley to the west), but the three taxa are allopatric. Members of clade K can be distinguished from other members of H. mastersi by their pink body coloration and pale sole. These semislugs are similar in size to clade J but slightly larger than clade I. Anatomically, this taxon differs in the shape and size of the penis (particularly in the presence of a large blind penis tip) and the spermatophore type. BIOGEOGRAPHY The genus Helicarion is widespread and comparatively abundant in south-eastern Australia; a probable testimony to its fairly well-developed dispersal ability. It occurs throughout the zone of temperate forests and woodlands with mild to warm summers, but generally not in the zones of hot summers to the north and west of its current range and the zone of cold summers in the alpine areas of Tasmania (Fig. 6). This entire region was exposed to more xeric conditions during the Plio-Pleistocene glacial cycles. A review of phylogeographical analyses has revealed that climatic fluctuations in this region have caused range contractions of mesic biota during the Quaternary, followed by expansions during wetter periods (Byrne et al., 2011). Furthermore, it has been demonstrated that these contractions have produced high levels of intraspecific differentiation in a range of species that are restricted to wet forests and alpine areas, which has resulted from the long-term isolation of populations in refuges and from local extirpations of populations in areas with more xeric conditions (Byrne et al., 2011). A concordant history of historical range contraction into climatic refuges and subsequent range expansions is the probable cause of the observed genetic structuring in Helicarion. Such a scenario may also explain the existence of morphologically well-differentiated high-altitude groups in Helicarion, which may have undergone a long history of isolation, whereas lowland forms are deemed to have expanded their ranges relatively recently. Based on the phylogeographical relationships documented here (e. g. Fig. 3), it is reasonable to postulate that Helicarion has originated on the Australian mainland and has colonized Tasmania subsequently. Quaternary climatic cycles have impacted Tasmania even more profoundly than the continental mainland as they caused repeated glaciations and created links to the continental Australia owing to sea level fluctuations. These processes have promoted both the isolation of populations in glacial refugia and an exchange between Tasmanian and mainland biota (e. g. McKinnon et al., 2004). The signature of such fluctuations is seen in the phylogeny of Helicarion revealing significant lineage divergence among the Tasmanian populations and links with the continental fauna (subclade A).	en	Hyman, Isabel T, Köhler, Frank (2018): Reconciling comparative anatomy and mitochondrial phylogenetics in revising species limits in the Australian semislug Helicarion Férussac, 1821 (Gastropoda: Stylommatophora). Zoological Journal of the Linnean Society 184: 933-968, DOI: 10.1093/zoolinnean/zly017, URL: https://academic.oup.com/zoolinnean/advance-article/doi/10.1093/zoolinnean/zly017/4998202
