Amplirhagada intermedia, (SOLEM, 1981)

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 : 352-354

AMPLIRHAGADA INTERMEDIA ( SOLEM, 1981)

FIGURES 7I–P , 14 , 15

Hadra (Hadra) imitata Smith, 1894: 92 (partim).

Amplirhagada alta intermedia Solem, 1981: 270– 271 , fig. 59e (partim).

Type locality: Western Australia, Kimberley, Bonaparte Archipelago, South Maret Island (~ 14°26′S, 124°59′E) .

Material studied: Holotype WAM S14444 (dried, worn shell) . Non-type material. WAM S33003 , WAM S33005 -7 , WAM S33013 -4 , WAM S42939 -4 , AM C.470986-90 ( Table 1) .

Taxonomic remarks: Based on similar shells, Solem (1981) subsumed material from various places on remote islands and the mainland under one polytypic species, Amplirhagada alta . Next to the nominate form from Corneille Island, Admiralty Gulf, he assigned specimens from Crystal Creek, Mitchell species are island endemics with ranges smaller than 20 km in diameter ( Köhler, 2010a, 2011a, b), these localities are considerably distant from each other (e.g. Corneille Island – South Maret Island – 90 km, with several islands in between that harbour different species). Therefore, the three subspecies named by Solem (1981) are elevated to the rank of distinct species. They are probably restricted to the immediate vicinity of their respective type localities.

Plateau, Kimberley mainland, to A. alta crystalla Solem, 1981 and specimens from South Maret (type locality), North Maret, and East Montalivet Island to A. alta intermedia Solem, 1981 . As demonstrated herein, a similar shell does not necessarily indicate close relationships. Although most Amplirhagada Diagnosis: Shell ( Fig. 7I–P ) variable in elevation and colour; broadly conical with low to moderately elevated spire; periphery rounded to slightly compressed ( Table 3). Teleoconch smooth with regular, faint axial growth lines only. Umbilicus 50–100% concealed by columellar reflection. Background colour and base crème, with well-developed brown subsutural and mid-whorl bands, rarely bands blending into each other, covering upper part of shell in brown suffusion; lip colour white ( Table 4). Aperture simple, slightly to well expanded, slightly reflected, with weak or wanting basal node. Genital anatomy ( Figs 14 , 15 ). Penis tubular, long, straight; vas deferens entering penial sheath near apical end, penial retractor muscle short; penial verge moderate in length (1/4 length of penial chamber), pointed; inner penial wall with narrow, rounded, regularly corrugated main pilaster, comprising more than posterior half of penial wall, segments resembling completely fused pustules, supporting two to three little hooks per segment; entire surface of inner penial wall with dense, elongated, rhomboid pustules. Vagina rather short (shorter than penis), inflated; bursa copulatrix moderately long, wide, extending to anterior end of spermoviduct, with weakly inflated head.

ferentiation in shells and genital anatomy, which is more or less consistent. Shells from Natfi Island much flatter with fewer whorls (similar to A. montalivetensis ). However, Natfi Island specimens not differentiated in genital anatomy or mitochondrial markers from topotypic specimens. Shell similar to A. turbinensis and A. berthierana , but penial wall anatomy different (see below).

Distribution: South Maret, North Maret, and Natfi Islands only ( Fig. 6 ).

AMPLIRHAGADA BERTHIERANA KÖHLER, 2010

FIGURE 7T–U

Amplirhagada berthierana Köhler, 2010a: 244 , figs 39–41.

Type locality ( Fig. 1): Western Australia, Kimberley, Bonaparte Archipelago, Berthier Island , 14°29′49″S, 124°59′34.1″E .

Comparative remarks: Shell similarly variable in shape and colour as A. ambulator , but shell never beehive-shaped with whorls not as flattened in crosssection but more rounded, separated by deeper suture; differing from A. fitzpatricki by more conspicuous banding, absence of axial ribs. Present species particularly differing from the former species in inner penial wall structure. Supporting regular, dense pustulation. Considerable genetic structuring between North and South Maret Islands not accompanied by similar dif-

Material studied: Holotype WAM S34611 ( Fig. 7U ) . Paratypes AM C.463754, WAM S33029 ( Table 1) .

Diagnosis: Shell ( Fig. 7T–U ) globose to broadly conical with medium high spire, thick, periphery evenly rounded ( Table 3); background and ventral colour crème, with well-marked brown, sub-sutural, and mid-whorl bands; outer lip white; teleoconch with axial growth lines only. Umbilicus 50–80% concealed by columellar reflection ( Table 4). Aperture simple, slightly to well expanded, slightly reflected, with weak or wanting basal node. Genital anatomy. Penis inflated, moderately long, straight; vas deferens entering penial sheath apically, penial retractor muscle moderate in length; penial verge rather short (1/6 length of penial chamber), elongately rounded; inner penial wall with narrow, regularly corrugated main pilaster, tapering in prominence and thickness towards both ends; entire inner penial wall with dense, indistinct, small pustules. 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. Large, almost globose shell not conspicuously differing from other species, such as A. ambulator or A. intermedia . However, penial charaters such as large, smooth, corrugated main pilaster and fine pustulation of entire penial wall are characteristic for only this species. Only in A. intermedia has the entire penial wall much larger, longer and rhomboid pustules.

Distribution: Berthier Island only ( Fig. 6 ).

Figures

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 14. Genital anatomy of Amplirhagada intermedia (WAM S42943, South Maret Island). Abbreviations: ag, albumen gland; at, atrium; bc, bursa copulatrix; hd, hermaphroditic duct; p, penis; rm, penial retractor muscle; so, spermoviduct; vd, vas deferens. Scale bar = 10 mm.

Figure 15. Penial anatomy of Amplirhagada intermedia (WAM S42943, South Maret Island). Abbreviations: mp, main pilaster of inner penial wall; pv, penial verge; rm, penial retractor muscle; sh, penial sheath; vd, vas deferens. Scale bar = 5 mm.