Contributions to Zoology, 83 (4) – 2014Francesco Criscione; Frank Köhler: Molecular phylogenetics and comparative anatomy of Kimberleytrachia Köhler, 2011 – a genus of land snail endemic to the coastal Kimberley, Western Australia with description of new taxa (Gastropoda, Camaenidae)

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Appendix

FIG2

Fig. 4. Shells. (A) K. jacksonensis n. sp. holotype WAM S66548 (B) K. leopardus n. sp. paratype WAM S66543; (C) K. nelsonensis n. sp. holotype WAM S66547; (D) K. serrata n. sp. paratype WAM S49616; (E) K. setosa n. sp. paratype WAM S49617; (F) K. silvaepluvialis n. sp. holotype WAM S66545; (G) Succochlea maretensis n. gen. n. sp. paratype WAM S83041. Scale bars = 1 cm. Note that mantle pattern of preserved animal is partly visible through transparent shells and that foot tissue protrudes from some shells.

FIG2

Fig. 5. SEM micrographs showing microsculpture of protoconch and teleoconch. (A-C) K. crawfordi Solem, 1979 WAM S49624; (D-F) K. jacksonensis n. sp. paratype WAM S66549. Scale bars: A, C, D, F = 500 µm; B, E = 1 mm.

FIG2

Fig. 6. SEM micrographs showing microsculpture of protoconch and teleoconch. (A-C) K. leopardus n. sp. paratype WAM S49628; (D-F) K. nelsonensis holotype WAM S66547. Scale bars: A, C, D, F = 500 µm; B, E = 1 mm.

FIG2

Fig. 7. SEM micrographs showing microsculpture of protoconch and teleoconch. (A-C) K. setosa n. sp. paratype WAM S49617; (D-F) K. serrata n. sp. paratype WAM S49616. Scale bars: A, C, D, F = 500 µm; B, E = 1 mm.

FIG2

Fig. 8. SEM micrographs showing microsculpture of protoconch and teleoconch. (A-C) K. silvaepluvialis n. sp. holotype WAM S66545; (D-F) Succochlea maretensis n. gen. n. sp. paratype WAM S83041. Scale bars: A, C, D, F = 500 µm; B, E = 1 mm.

FIG2

Fig. 9. Penial anatomy. (A) K. crawfordi Solem, 1979 WAM S49624; (B) K. jacksonensis n. sp. holotype WAM S66548; (C) K. leopardus n. sp. holotype WAM S66543; (D) K. nelsonensis n. sp. holotype WAM S66547 (E) K. setosa n. sp. holotype WAM S66544; (F) K. silvaepluvialis n. sp. holotype WAM S66545. Scale bars = 5 mm (refer to p. 248 for abbreviations).

FIG2

Fig. 10. Genital anatomy of Succochlea maretensis n. gen n. sp. holotype WAM S66554. (A) Whole genitalia; (B) Penis, opened. Scale bars = 5 mm (refer to p. 248 for abbreviations).

Taxonomic account and descriptions


Family Camaenidae Pilsbry, 1895

Genus Kimberleytrachia Köhler, 2011.

Torresitrachia – Solem, 1979: 45–92 (partim); 1985: 922–933 (partim); 1991: 178–185

(partim), Solem and McKenzie, 1991: 247–263 (partim).

Kimberleytrachia - Köhler, 2011b: 377-400.

Type species. Kimberleytrachia somniator Köhler, 2011.

Diagnosis. Shell. Medium sized (D = 12–24 mm; Table 2), thin or mostly very delicate, discoidal to broadly conical with flat to moderately elevated spire; umbilicus narrowly open, may be partly concealed; protoconch with crowded pustules arranged in radial pattern; teleoconch with or without well-developed thin radial ribs (faint radial growth lines may be present), with well-developed microsculpture of periostracal setae or projections (particularly visible in juvenile shells); whorls moderately to slowly increasing, separated by deep suture, periphery well rounded. Shell colour light to yellowish brown, no colour banding. Aperture wide, with relatively thin, weakly to moderately expanded and weakly reflected lip, palatal node absent, basal node weak or absent, parietal wall inconspicuous.

Genital anatomy. Penis without penial sheath, embedded in connective tissue; epiphallus well-developed, with well-developed epiphallic flagellum. Inner penial wall supporting simple to rather complex armature of longitudinal, oblique or transverse pilasters or lamellae, frequently with pad-like swelling underneath opening to epiphallus, pustulation, lateral lamellae or more complex armature may be present. Penial retractor muscle attached at epiphallus between mid-portion to near posterior end. Epiphallus opens to penial lumen through constricting circular vergic ring. Inner epiphallic wall supporting longitudinal pilasters that vary in number and development. Vas deferens rather thin, entering epiphallus laterally near distal end or terminally through slit-like opening or often hose-like extended into lumen of epiphallus. Bursa copulatrix elongate, tubular, reaching anterior end of albumen gland.

Remarks. The original description of Köhler (2011b) is here emendated to include the presence of radial ribs and fine pustules on the shell of some species. For comparative reasons and in order to complete the documentation of the morphology of all species, we provide illustrations of K. crawfordi (Fig. 5A-C, 9A), whose genital anatomy has not been figured before. New records are reported for the following species: K. umbonis (Solem, 1979), K. crawfordi (Solem, 1979), K. canopi Köhler, 2011.

Kimberleytrachia jacksonensis n. sp. (Figs 4A, 5D-F, 9B)

Material examined

Holotype. Australia, WA, Bonaparte Archipelago, Brunswick Bay, small islet 0.2 km E of Jackson Island, 15°10’23” S, 124°39’00” E; coll. 11 Jul 1988 (WAM S66548 wet, dissected).

Paratypes. Same as holotype (FMNH 219086, 5 wet; FMNH 219085, 1 dry; WAM S66549, 2 wet).

Other material. Australia, WA, Bonaparte Archipelago, Brunswick Bay, Jackson Island, 15°10’28”S, 124°38’43” E (11 Jul 1988), 4 wet (FMNH 219088), 4 dry (FMNH 219087), 2 wet (WAM S66551), 3 dry (WAM S66550).

Etymology. For Jackon Island, Latinised adjective of feminine gender.

Description. Shell (Figs 4A, 5D-F). Large to moderate in size (Table 2), delicate, broadly conical with moderately high spire; periphery well rounded; umbilicus 30-60% concealed; colour yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with very fine pustules. Teleoconch with faint growth lines and no radial ribs; entirely covered either with dense, fine, short periostracal projections (setae) extending onto inner umbilical whorls or fine pustules. Aperture moderately wide with moderately expanded, moderately reflected, thin lip; palatal node absent, basal node weakly developed.

Genital anatomy (Fig. 9C). Penis elongate, twice as long as vagina. Distal half of inner penial wall with densely packed transversal folds, broadening proximally; proximal half with longitudinal wide pilasters. Epiphallus almost one third shorter than penis, with thin epiphallic flagellum (ten times shorter than epiphallus); inner epiphallic wall supporting several longitudinal folds along its median portion, terminating into a broad swelling supporting regular pustulations. Penial retractor muscle rather short. Free oviduct as long as vagina, two times shorter than bursa copulatrix.

Comparative remarks. Shell size and shape similar to many congeners, differing from K. alphacentauri by larger size, from K. aequum, K. chartacea, K. leopardus, K. nelsonensis and K. serrata by more elevated spire. Shell sculpture differing from K. achernaria, K. canopi, K. silvaepluvialis and K. umbonis by presence of setae on whole teleoconch; not readily differentiated from remaining species based on shell features only. Inner penial wall sculpture similar to K. canopi, K. deflecta and K. leopardus, differing by sculpture of the inner epiphallic wall.

Distribution. Known only from Jackson Island.

Kimberleytrachia leopardus n. sp. (Figs 4B, 6A-C, 9C)

Material examined

Holotype. Australia, WA, NW Kimberley, 15 km NE of Youwanjela Creek, Prince Regent River NR; 15°31’36” S, 125°21’24” E; coll. V. Kessner, 26 Jan 2010, gully with vine thicket, sandstone boulders, scree, running stream (WAM S66543, 1 wet, dissected).

Paratypes. Same as holotype (WAM S49628, 12 wet; WAM S49711, 1 dry).

Other, non-type material. Australia, WA, NW Kimberley, Prince Regent River NR: 5 km N of Mt. York, rainforest patch on creek with massive sandstone boulders, under slabs; 15°26’21” S, 125°32’23” E (WAM S49648, 1 wet; WAM S49717, 3 dry); about 1.7 km S of Youwanjela Creek, rainforest in gully on upper slopes; 15°36’09” S, 125°26’33” E (WAM S49619, 35 wet; WAM S49702, 9 dry); 5.5 km NE of Youwanjela Creek, sandstone terraces with small patches of vine thicket; 15°33’00” S, 125°28’51” E (WAM S49620, 10 wet; WAM S49703, 5 dry); about 72 km NNE of Bachsten Creek Camp, sandstone terraces, small vine thicket patches; 15°17’34” S, 125°29’50” E (WAM S49622, 5 wet; WAM S49708, 1 dry); 4 km SE of Mt. Brookes, sandstone pavement, on boulders in protected notch in platform; 15°13’15” S, 125°25’50” E (WAM S49641, 5 wet; WAM S49716, 2 dry); Garimbu Creek, sandstone gorge, vine thicket, under boulders; 15°21’37” S, 125°30’17” E (WAM S49666, 1 wet); 4 km SE of Mt. Brookes, in slot in sandstone pavement, under boulders; 15°13’12” S, 125°25’55” E (WAM S49662, 3wet; WAM S49783, 1dry; WAM S49713, 3dry); Prince Regent River Nature Reserve, vine thicket in sinkhole, on ceilings of overhangs of massive sandstone boulders, 15°17’32” S, 125°29’43” E (WAM S49709, 5 dry).

Etymology. For the colour pattern of the mantle roof, resembling leopard fur, visible through transparent shell; characteristic feature of Kimberleytrachia species. Derived from ‘leopardus’ (Latin = leopard), noun in apposition.

Description. Shell (Figs 4B, 6A-C). Large to moderate in size (Table 2), delicate, broadly conical with low spire; periphery well rounded; umbilicus 60-90% concealed; colour yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with very fine pustules. Teleoconch with marked, very thin radial ribs and well-developed dense, fine pustules, extending on entire shell surface and inner umbilical whorls; periostracal projections not present. Aperture moderately wide with strongly expanded, moderately reflected, thin lip; palatal node absent, basal node well developed.

Genital anatomy (Fig. 9C). Penis elongate, twice as long as vagina. Distal two thirds of inner penial wall with fine and densely packed transversal lamellae, extending onto relatively narrow pad-like swelling, proximal third with longitudinal wide pilasters. Epiphallus almost twice shorter than penis, with thin epiphallic flagellum (five times shorter than epiphallus); inner epiphallic wall supporting one broad longitudinal pilaster along its median portion. Penial retractor muscle rather short. Free oviduct one quarter shorter than vagina, five times shorter than bursa copulatrix.

Comparative remarks. Shell size and shape similar to K. aequum, K. chartacea, K. nelsonensis and K. serrata, differing from those species by presence of radial ribs and absence of periostracal projections. Inner penial wall sculpture similar to K. canopi, K. crawfordi, K. setosa, K. jacksonensis and K. umbonis, differing from the former three species by transverse lamellae covering a larger area and from the latter two by sculpture of the inner epiphallic wall.

Distribution. Northern part of the Regent River catchment within the Prince Regent Nature Reserve (Fig. 1).

Kimberleytrachia nelsonensis n. sp. (Figs 4C, 6D-F, 9D)

Material examined.

Holotype. Australia, WA, Bonaparte Archipelago, Port Nelson, 5 km N of Mt. Knight, ca. 3 km E of Careening Bay, 15°06’41” S, 125°01’53” E; coll. 13 Jul 1988 (WAM S66547).

Paratypes. Same as holotype (FMNH 219150, 2 wet; FMNH 219149, 6 dry; WAM S66546, 4 dry).

Etymology. For Port Nelson, Latinised adjective of feminine gender.

Description. Shell (Figs 4C, 6D-F). Moderate in size (Table 2), delicate, broadly conical with moderately low spire; periphery well rounded; umbilicus less than 30% concealed; colour yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with very fine pustules. Teleoconch with faint growth lines and no radial ribs; entirely covered with dense, fine, short periostracal projections (setae) extending onto inner umbilical whorls. Aperture moderately wide with weakly expanded, weakly reflected, thin lip; palatal node absent, basal node weakly developed.

Genital anatomy (Fig. 9D). Penis elongate, almost twice as long as vagina. Distal portion of inner penial wall with median pad-like swelling supporting fine tubercles and with densely-packed transversal lamellae, becoming less fragmented proximally; proximal portion with wide longitudinal pilaster originating from swelling and narrower pilaster reaching atrium. Epiphallus more than twice shorter than penis, with thin epiphallic flagellum (five times shorter than epiphallus); inner epiphallic wall supporting one longitudinal pilaster along its median portion, proximally thin and distally widening into a broad swelling supporting regular pustulations. Penial retractor muscle wide and short. Free oviduct as long as than vagina, five times shorter than bursa copulatrix.

Comparative remarks. Shell size and shape similar to K. aequum, K. alphacentauri, K. chartacea, K. leopardus and K. serrata, differing from K. achernaria, K. canopi, K. silvaepluvialis and K. umbonis by presence of setae on whole teleoconch; not readily differentiated from other congeners based on shell characters only. Inner penial wall sculpture rather distinct from all other species by presence of fragmented lamellae.

Distribution. Known from the type locality only.

Kimberleytrachia serrata n. sp. (Figs 4D, 7A-C)

Material examined

Holotype. Australia, WA, NW Kimberley, Prince Regent River Nature Reserve, 14.5 km SSW of Boongaree Is, rainforest patch in a gully, sandstone boulders, scree, 15°14’00” S, 125°06’47” E; coll. V. Kessner, 27 Jan 2010, under rocks (WAM S66553).

Paratypes. Same as holotype (WAM S49616, 5 wet; WAM S49780, 6 dry).

Other, non-type material. Australia, WA, NW Kimberley, upper reaches of Prince Regent River NR, Sandstone outcrops in open woodland, 15°17’03” S, 125°14’25” E (WAM S49796, 1dry).

Etymology. For the distinctive shape of periostracal projections on the body whorl, resembling saw teeth. From ‘serratus’ (Latin = serrated, toothed like a saw), adjective in feminine gender.

Description. Shell (Figs 4D, 7A-C). Moderate in size (Table 2), delicate, almost discoidal with low spire; periphery well rounded; umbilicus almost not concealed; colour light brown, base of shell lighter, yellowish-brown. Protoconch with very fine pustules. Teleoconch with faint growth lines and no radial ribs; entirely covered with dense, fine, short periostracal projections, extending onto inner umbilical whorls; projections denticle-shaped on body whorl and with smoother, rounded tips on remaining teleoconch whorls. Aperture narrow with strongly expanded, weakly reflected, thin lip; palatal node absent, basal node moderately developed.

Comparative remarks. Shell size and shape similar to K. aequum, K. alphacentauri, K. chartacea, K. leopardus and K. nelsonensis, by unique shape of periostracal projections. No adult was collected. Hence, the genital anatomy of this species remains unknown.

Distribution. North of the Prince Regent Reserve, north-east of Prince Frederick Harbour.

Kimberleytrachia setosa n. sp. (Figs 4E, 7D-F, 9E)

Material examined

Holotype. Australia, WA, NW Kimberley, S of Bachsten Creek Base Camp, 15°59’26” S, 125°19’40” E; coll. V. Kessner, M. Maier, 14 Jan 2010, rocky outcrops, small pockets of vine thicket on top of sandstone escarpment, under overhangs (WAM S66544, wet, dissected).

Paratypes. Same as holotype (WAM S49617, 7 wet).

Other, non-type material. 4.7 km NE of Bachsten Creek Base Camp, sandstone platforms and gully with boulders and small patches of vine thicket, 15°59’33” S, 125°19’44” E (WAM S49618, 2 wet); Bandicoot Creek, ca 24 km SE of Bachsten Creek Camp, small pockets of vine thicket on sandstone outcrops, 16°10’31” S, 125°19’48” E (WAM S49625, 5 wet); 1.3 km W of Bachsten Creek Base Camp, sandstone outcrops, sandy patches, boulders, 15°59’22” S, 125°19’00” E (WAM S49707, 2 dry); Pitta Gorge, rainforest over sandstone scree and soil, 15°52’59” S, 125°35’05” E (WAM S49786, 1 dry); 3.5 km S of Bachsten Creek Base Camp, gully with boulders and small patches of vine thicket, 16°01’15” S, 125°19’41” E (WAM S49633, 6 dry).

Etymology. In reference to the periostracal projections (setae) on the shell. Derived from ‘setosus’ (Latin = covered by hairs), adjective of feminine gender.

Description. Shell (Figs 4E, 7D-E). Moderate in size (Table 2), delicate, broadly conical with relatively high spire; periphery well rounded; umbilicus 30-60% concealed; colour yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with very fine pustules. Teleoconch with faint growth lines and no radial ribs; entirely covered with dense, fine, short periostracal projections (setae) extending onto inner umbilical whorls. Aperture wide with strongly expanded, weakly reflected, thin lip; palatal node absent, basal node well developed.

Genital anatomy (Fig. 9E). Penis elongate, almost twice as long as vagina. Distal third of inner penial wall with fine and densely packed transversal lamellae, extending onto relatively narrow and long pad-like swelling, longitudinal wide pilasters at proximal two thirds. Epiphallus twice as long as penis, with thin epiphallic flagellum (ten times shorter than epiphallus); inner epiphallic wall proximal part with two broad longitudinal pilasters, median part proximally essentially smooth, distal part with three longitudinal pilasters (one reflexing into flagellum). Penial retractor muscle long. Free oviduct one quarter shorter than vagina, five times shorter than bursa copulatrix.

Comparative remarks. Shell size and shape very similar to all congeners, differing from K. alphacentauri by larger size and from K. aequum, K. chartacea, K. leo­pardus and K. serrata by more elevated spire. Shell sculpture differing from K. achernaria, K. canopi, K. silvaepluvialis and K. umbonis by presence of setae on whole teleoconch; not readily differentiated from remaining species based on shell features only. Inner penial wall sculpture similar to K. leopardus, K. canopi, K. crawfordi and K. umbonis, differing from the former species by smaller area covered by transverse lamellae and from the latter three by sculpture of the inner epiphallic wall.

Distribution. Together with K. umbonis and K. crawfordi, this species occupies the southernmost parts of the genus’ range.

Kimberleytrachia silvaepluvialis n. sp. (Figs 4F, 8A-C, 9F)

Material examined

Holotype. Australia, WA, NW Kimberley, Prince Regent River NR, 27.4 km SE of Boongaree Is, 15°18’56” S, 125°21’10” E; coll. V. Kessner, R. Barrett, M. Maier, 23 Jan 2010, in vine thicket patch in narrow gully, sandstone cliffs and boulders, running stream, under overhangs (WAM S66545, wet, dissected).

Paratypes. Same as holotype (WAM S49631, 2 wet).

Etymology. In reference to the affinity of species of this genus to rainforest ecosystems (vine thickets), derived from ‘silva’ (Latin = forest) and ‘pluvialis’ (Latin = of rain), noun and adjective of feminine gender declined to genitive case.

Description. Shell (Figs 4F, 8A-C). Large in size (Ht: D = 25.3 mm, H = 14.3 mm, W = 5.3), delicate, broadly conical with moderately high spire; periphery well rounded; colour yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with rather coarse pustules. Teleoconch with marked, very thin radial ribs and well-developed dense, fine pustules, extending on entire shell surface and inner umbilical whorls but restricted to narrow subsutural area on body whorl; periostracal projections not present. Aperture wide with strongly expanded, weakly reflected, thin lip; palatal node absent, basal node well developed.

Genital anatomy (Fig. 9F). Penis elongate, twice as long as vagina. Distal third of inner penial wall with wide pad-like corrugated swelling, tapering into thin corrugated pilasters along wall median third, proximal third with smooth longitudinal pilasters. Epiphallus almost twice as long as penis, with thin epiphallic flagellum (ten times shorter than epiphallus); inner epiphallic wall with abundant, fine, transversal lamellae and several thin, regularly spaced, longitudinal pilasters. Penial retractor relatively short. Free oviduct one third shorter than vagina, five times shorter than bursa copulatrix.

Comparative remarks. Shell size and shape very similar to all congeners, except K. alphacentauri, K. chartacea and K. leopardus, differing from K. amplirhagadoides Köhler, 2011, K. crawfordi, K. crucis Köhler, 2011, K. deflecta, K. setosa and K. somniator by absence of setae; not readily differentiated from remaining species based on shell features only. Inner penial wall sculpture very distinctive and similar only to that of K. amplirhagadoides, differing from it by sculpture of the inner epiphallic wall. Data available on the morphology of this species come from the holotype, the only adult specimen collected.

Distribution. Known from the type locality only.

Succochlea n. gen.

Type species. Succochlea maretensis n. sp.

Etymology. Contraction derived from ‘succinum’ (Latin = amber) and ‘cochlea’ (Latin = snail); for ‘amber snail’ referring to its shell colour; noun of female gender.

Diagnosis. Shell (Figs 4G, 8D-F). Moderate, moderately elevated, moderately deep suture. Umbilicus open, wide, partially concealed by outer lip reflection. Protoconch sculpture of axial ridgelets. Teleoconch microsculpture of thin axial ribs. Aperture wide, not deflected from axis of coiling; outer lip thin.

Genital anatomy (Fig. 10A-B). Penis without penial sheath, embedded in connective tissue; epiphallus well-developed, with well-developed epiphallic flagellum. Inner penial wall supporting armature of pilasters or lamellae, with pad-like swelling underneath opening to epiphallus. Penial retractor muscle attached at epiphallus between mid-portion to near posterior end. Epiphallus opens to penial lumen through constricting circular vergic ring. Vas deferens rather thin, entering epiphallus laterally near distal end through slit-like opening or often hose-like extended into lumen of epiphallus. Bursa copulatrix elongate, tubular, reaching anterior end of albumen gland.

Comparative remarks. Shell easily distinguished from most of other camaenids genera in the region by combination of thinness, lack of teleoconch macro- and micro- sculpture. Shell also very similar to Kimberleytrachia, differing from the latter by having a distinctive protoconch microsculpture of radial ribs, less pronouncedly descending body whorl, reduced apertural lip, lacking basal nodule, periostracal projections and pustulation. Genitalia differ from Kimberleytrachia by vagina longer than penis.

Succochlea maretensis n. sp. (Figs 4G, 8D-E, 10A-B)

Material examined

Holotype. Australia, WA, Bonaparte Archipelago, North Maret Island, 14°23’56” S, 124°58’23” E; coll. V. Kessner, A. Longbottom, 15 Jul 1988 (WAM S66554, wet, dissected).

Paratypes. Same as holotype (WAM S83042, 8 wet; WAM S83041, 23 dry; FMNH 219185, 7 wet; FMNH 219184, 22 dry).

Other, non-type material. Australia, WA, Bonaparte Archipelago, SW corner of Berthier Island, 14°30’45” S, 124°58’50” E (WAM S83040, 2 dry, 1 wet); South Maret Island, 14°26’27” S, 124°58’38” E (WAM S83043, 4 dry; FMNH 219187, 4 dry); unnamed island between Maret and Berthier Islands, 14°28’53” S, 125°00’13” E (WAM S83045, 5 wet; WAM S83044, 6 dry; FMNH 219202, 5 wet; FMNH 219201, 7 dry).

Etymology. For Maret Island, Latinised adjective of feminine gender.

Description. Shell (Figs 4G, 8D-F). Large to moderate in size (Table 2), delicate, broadly conical with high spire; periphery well rounded; umbilicus 30% concealed; colour golden yellowish-brown, base of shell lighter, pale yellowish-horn. Protoconch with axial ridgelets. Teleoconch with faint, very thin radial ribs and no pustulations; periostracal projections not present. Aperture moderately wide with weakly expanded, not reflected, thin lip; palatal and basal nodes absent.

Genital anatomy (Fig. 10). Penis elongate, slightly shorter than vagina. Distal half of inner penial wall with relatively narrow pad-like swelling, covered by fine and densely packed transversal lamellae, proximal half with longitudinal wide pilasters. Epiphallus almost as long as penis, with thin epiphallic flagellum (three times shorter than epiphallus); inner epiphallic wall supporting three broad longitudinal pilasters. Penial retractor muscle rather short. Free oviduct as long as vagina, twice shorter than bursa copulatrix.

Distribution. Known from the Maret and Berthier island group only.