Amplirhagada albertiana, Köhler & Johnson, 2012

Köhler, Frank & Johnson, Michael S., 2012, Species limits in molecular phylogenies: a cautionary tale from Australian land snails (Camaenidae: Amplirhagada Iredale, 1933), Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 165 (2), pp. 337-362 : 356-358

AMPLIRHAGADA ALBERTIANA SP. NOV.

FIGURES 7V , 18 , 19

Type locality: Western Australia, Kimberley, Bonaparte Archipelago, Albert Island , 14°31′09.1″S, 124°55′48.4″E .

Material examined: Holotype WAM S42947 (dissected specimen) . Paratypes WAM S33000 (four preserved specimens) AM C.470992 (one preserved body) ( Table 1). Additional, non-type material . FMNH 219169-70 , FMNH 219670 from Albert Island (KC-043, coll. 14.vii.1988, V. Kessner) .

Etymology: For Albert Island, adjective of feminine gender.

Description: Shell ( Fig. 7V ) globose to broadly conical with medium high spire, thick, periphery evenly rounded ( Table 3); background and ventral whitish colour crème, with brown, sub-sutural, and mid-whorl bands, shell tip may be covered in brownish suffusion; outer lip white; teleoconch with axial growth lines only. Umbilicus 50–100% concealed by columellar reflection ( Table 4). Aperture simple, slightly to well expanded, slightly reflected, with weak or wanting basal node. Genital anatomy ( Figs 18 , 19 ). Penis inflated, long, coiled within penial sheath; vas deferens entering penial sheath within anterior third, penial retractor muscle moderate in length; penial verge short (<1/10 length of penial chamber), pointed; inner penial wall without main pilaster; posterior third of wall with densely packed, elongate, roundish, irregularly arranged pustules; anterior half of inner wall with irregularly shaped and arranged, corrugated, indistinct, oblique pilasters. Vagina long (about same length as penis), tubular; bursa copulatrix moderately long, tubular, extending to anterior end of spermoviduct, with inflated head.

Comparative remarks: Shell most similar to A. berthierana in shape and colour but smaller (similar to and A. turbinensis ), differing from the former two species by closed instead of open umbilicus. Unlike A. intermedia , A. berthierana , and A. turbinensis penial wall not entirely covered with pustules. Typical features are coiled penis and absence of main pilaster. Bursa copulatrix conspicuously elongated (even more so than in former species). Material kept in the FMNH is labelled as ‘ Amplirhagada sp 69’.

Distribution: Albert Island only ( Fig. 6 ).

AMPLIRHAGADA GEMINA KÖHLER, 2010

FIGURE 20A–D

Amplirhagada gemina Köhler, 2010a: 270 , figs 82–84.

Type locality: Western Australia, Kimberley mainland near Camden Sound, 4.5 km north of Prior Pt , 1.5 km south of Hall Pt , 15°40′45″S, 124°22′45″E .

Material examined ( Table 1): Holotype WAM S34611 ( Fig. 20A ) . Paratypes AM C.463754, WAM S33029 . Non-type material. WAM S42955 -7 ( Fig. 20B–D ).

Diagnosis: Shell ( Fig. 20A–D ) semi-globose to broadly conical, with moderately elevated spire; periphery evenly rounded to slightly angulate; with regular growth lines only. Umbilicus 60–100% concealed by columellar reflection ( Table 4). Colour brown, base beige, lip colour whitish. Aperture simple, slightly expanded, weakly reflected, basal node absent. Genital anatomy. Penis slightly inflated, moderately long, straight; vas deferens entering penial sheath apically, penial retractor muscle moderate in length; penial verge tiny; inner penial wall entirely covered with narrowly packed, elongate, rather small pustules, no main pilaster differentiated. Vagina short (shorter than penis), inflated; bursa copulatrix moderately long, wide, extending to anterior end of spermoviduct, with weakly inflated head ( Köhler, 2010a). Comparative remarks: Amplirhagada gemina and A. kimberleyana are difficult to differentiate. Their shells look almost identical, with specimens from Lulim Island being more similar to A. kimberleyana than to conspecific specimens from the mainland ( Fig. 3 ). Main differentiating feature is slightly different penial wall sculpture: A. gemina has more crowded, smaller, undifferentiated pustules while A. kimberleyana has larger and fewer pustules, arranged in rows, some of them with hooks forming a weak main pilaster.

Distribution: South-west Kimberley coast between south of Halls Pt and Wilsons Pt, including Lulim Island, close to mainland coast ( Fig. 6 ).

AMPLIRHAGADA KIMBERLEYANA KÖHLER, 2010

FIGURE 20E–F

Amplirhagada kimberleyana Köhler, 2010a: 268–9 , figs 79–81.

Type locality: Western Australia, northern Slate Island, Camden Sound, 2 km north-west of Wilsons Pt , 15°32′10″S, 124°23′56″E .

Material examined ( Table 1): Holotype WAM S34611 ( Fig. 20E ) . Paratypes AM C.463754, WAM S33029 . Non-type material. WAM S42955 -7 ( Fig. 20F ).

Taxonomic remarks: The type material was collected in 1988 on the ‘northern Slate Island’, which is identical to Wailgwin Island, named so subsequently.

Diagnosis: Shell ( Fig. 20E, F ) semi-globose to broadly conical, with moderately elevated spire; periphery evenly rounded to slightly angulate; with regular growth lines only. Umbilicus 60–100% concealed by columellar reflection ( Table 4). Colour brown, occasionally banded, base and background colour beige, lip colour whitish. Aperture simple, slightly expanded, weakly reflected, basal node absent. Genital anatomy. Penis slightly inflated, moderately long, straight; vas deferens entering penial sheath within anterior third, penial retractor muscle moderate in length; penial verge tiny; inner penial wall entirely covered with narrowly packed, elongate, rather small pustules, main pilaster weakly differentiated, consisting of 2–3 parallel rows of inconspicuously enlarged pustules that support little hooks. Vagina short (shorter than penis), inflated; bursa copulatrix moderately long, wide, extending to anterior end of spermoviduct, with weakly inflated head ( Köhler, 2010a).

Comparative remarks: Material studied here is topotypic. See above for comparison with A. gemina .

Distribution: Wailgwin Island only ( Fig. 6 ).

Figures

Figure 3. Island-based comparison of specimens of Clade 3 by means of box and whisker plots for shell parameters H, D, H/D, and N.

Figure 6. Map of the study area with known occurrences of Amplirhagada species as documented by Solem (1981) and Köhler (2010a, 2011a, c). Triangles (in italics): Localities of known species: and, A. anderdonensis; bas, A. basilica; ber, A. berthierana; boo, A. boongareensis; buf, A. buffonensis; cam, A. camdenensis; cof, A. coffea; dec, A. decora; dis, A. discoidea; epi; A. epiphallica; for, A. forrestiana; gem, A. gemina; gib, A. gibsoni; glo, A. globosa; int, A. intermedia; inu, A. inusitata; kes, A. kessneri; kim, A. kimberleyana; lam, A. lamarckiana; lin, A. lindsayae; mon, A. montalivetensis; pue, A. puescheli; reg, A. regia; sph, A. sphaeroidea; tri, A. tricenaria; uwi, A. uwinsensis; via, A. vialae; yor, A. yorkensis. Circles (in bold): material studied herein: alb, A. albertiana sp. nov. (Albert Island); emo, A. ambulator sp. nov. (East Montalivet Island group); lul-wip, A. gemina; nma, A. intermedia (North Maret Island); prr, A. tealei sp. nov. (Mainland View, Prince Regent Reserve); sb, A. imperialis sp. nov. (Boongaree Island); sma, A. intermedia (South Maret and Natfi Island); tur, A. turbinensis sp. nov. (Turbin island); wal, A. montalivetensis (Walker Island); wmo, A. fitzpatricki sp. nov. (West Montalivet Island).

Figure 7. Shells of CLADES 1 and 2 (columns showing front view, top view, basal view each). A–C, Amplirhagada montalivetensis (Smith, 1894). A, lectotype of Helix (Hadra) montalivetensis Smith, 1894 NHMUK 1890.12.30.138–142, Montalivet Island. B, paralectotype AM C33896. C, WAM S33021 (Walker Island). D–H, Amplirhagada ambulator sp. nov. D, paratype WAM S33019 (East Montalivet Island). E, East Montalivet Island (WAM S33018). F, Patricia Island (WAM S33027). G, East Montalivet Island (WAM S42934). H, Don Island (WAM S33025). I–P, Amplirhagada intermedia (Solem, 1981). I, North Maret Island (WAM S42940). J, North Maret Island (WAM S33007). K, North Maret Island (WAM S33007). L, North Maret Island (WAM S42939). M, South Maret Island (WAM S42943). N, South Maret Island (WAM S42942). O, South Maret Island (WAM S42941). P, Natfi Island (WAM S42944). Q–R, Amplirhagada fitzpatricki sp. nov. paratypes West Montalivet Island (WAM S33010). S, Amplirhagada turbinensis sp. nov. paratype, Turbin Island (WAM S42946). T–U, Amplirhagada berthierana Köhler, 2010a, Berthier Island. T, paratype WAM S33029. U, holotype WAM S34611. V, Amplirhagada albertiana sp. nov. paratype Albert Island (WAM S33000). Note that foot tissue protrudes from some of the shells. Scale bar = 10 mm.

Figure 18. Genital anatomy of Amplirhagada albertiana sp. nov. holotype WAM S42947 (Albert Island). Abbreviations: ag, albumen gland; at, atrium; bc, bursa copulatrix; hd, hermaphroditic duct; p, penis; rm, penial retractor muscle; so, spermoviduct; vd, vas deferens; vg, vagina. Scale bar = 10 mm.

Figure 19. Penial anatomy of Amplirhagada albertiana sp. nov. holotype WAM S42947 (Albert Island). A, detail of inner penial wall. B, schematic representation of coiled penis within penial sheath (not to scale). Abbreviations: pv, penial verge; sh, penial sheath. Scale bar = 5 mm.

Figure 20. Shells of CLADE 3. A–D, Amplirhagada gemina. A, holotype WAM S34626 (4.5 km north of Prior Pt.). B, WAM S42955 (Wilsons Point). C, WAM S42957 (Wilsons Point). D, WAM S42954 (Lulim Island). E–F, Amplirhagada kimberleyana. E, holotype WAM S34625 (northern Slate Island). F, WAM S42952 (Wailgwin Island). Note that foot tissue protrudes from some of the shells. Scale bar = 10 mm.