Amalda hilgendorfi (von Martens, 1897 )

Kantor, Yuri I., Castelin, Magalie, Fedosov, Alexander & Bouchet, Philippe, 2020, The Indo-Pacific Amalda (Neogastropoda, Olivoidea, Ancillariidae) revisited with molecular data, with special emphasis on New Caledonia, European Journal of Taxonomy 706, pp. 1-59 : 14-16

Amalda hilgendorfi (von Martens, 1897)

Figs 2–4 , 5–6 , 7 A–G, 8I–J

Ancillaria hilgendorfi von Martens, 1897: 167 , pl. 16 figs 26–27.

Amalda hilgendorfi richeri Kilburn & Bouchet, 1988: 285–287 , figs 18–22 [not figs 23–24 = Amalda cacao sp. nov.].

Amalda vernedei herlaari van Pel, 1989: 14 , figs 1–3.

Amalda hilgendorfi vezzaroi Cossignani, 2015: 24–25 , text figures.

Baryspira hilgendorfi – Kira 1962: 208, pl. 71 fig. 1. — Kuroda et al. 1971: 300 [Japanese text], 196 [English text], pl. 54. fig. 1.

Amalda aureomarginata Northern New Caledonia form – Kilburn & Bouchet 1988: 284, Fig. 13 .

Amalda aureomarginata Chesterfield form – Kilburn & Bouchet 1988: 284, figs 14–17.

? Amalda (Alcospira) hilgendorfi – Ninomiya 1990: pl. 2 fig. 5.

Amalda hilgendorfi – Okutani 2000: 523, pl. 260 fig. 2 (erroneously labelled A. hayashii on the plate) [not pl. 260 fig. 3 = A. hayashii ]. — Gratecap 2016b: 4, 12, figs 3–4. — Tsuchida 2017: 995, pl. 288 fig. 3.

Amalda vernedei herlaari – Poppe 2008: pl. 546, figs 7–9.

Amalda hilgendorfi richeri – Gratecap 2014 (part): 14, 19, figs 19–23 [not figs 24–25 = Amalda cacao sp. nov.]

Type material

JAPAN • Holotype of Ancillaria hilgendorfi ; ZMB 31632 .

NEW CALEDONIA • Holotype of Amalda hilgendorfi richeri ; North of New Caledonia , Grand Passage; 18°57′ S, 163°14′ E; 480 m deep; MNHN IM-2000-1378.

ARAFURA SEA • Holotype of Amalda vernedei herlaari ; Arafura Sea; 150 fathoms [270 m]; formerly ZMA Moll. no. 3.88.040, now NBC .

PHILIPPINES • Holotype of Amalda hilgendorfi vezzaroi ; Philippines, Aliguay island; MMM.

Description

SHELL. Variable, narrowly to broadly fusiform with high, acuminate spire; thin to thick microshagreened primary callus impressed at suture, forming distinct to very distinct, obtuse keel above suture. Protoconch exposed, medium large (n =2, first whorl diameter 0.75–0.92 mm), border with teleoconch indistinct. Secondary callus very thin to distinct, tongue-shaped, broadening in middle part and rounded adapically, extending on last three whorls, seemingly less shagreened than primary callus. Plication plate with 2–7 weak ridges, sometimes nearly obsolete. Olivoid groove from narrow and distinct to broad and shallow or nearly obsolete, labral tooth distinct, obtuse. Upper anterior band usually thickened, sometimes heavily. Colour variable, from pale yellowish to brown, olivoid band always lighter than body whorl cloak, anterior band the darkest anterior zone of shell, plication plate usually nearly white, primary callus bordered by lighter narrow band (or line). Height to 81 mm, exceptionally up to 98 mm.

RADULA (n =16 of different morphs). Lateral teeth very similar in all species of Amalda , unicuspid, hook-shaped ( Fig. 7A ). Central tooth normally tricuspid with central cusp shortest. Some variation in central tooth shape ( Fig. 7 B–G); central cusp from extremely short, nearly obsolete, to subdivided in two cusplets, or with additional denticles on side. Cusps of the central tooth either with smooth edges, or serrated ( Fig. 7G ). Morphological variation not related to geographical distribution.

Remarks

Molecular analysis demonstrated high heterogeneity of this species with several subclades, mostly geographically determined ( Fig. 2 ), with groupings from Japan, the Philippines, Papua New Guinea and the Solomon Islands, north of New Caledonia and the Coral Sea, and south of New Caledonia and Norfolk Ridge. This structure is, however, not clear-cut; for example, individuals with identical haplotypes are found in northern New Caledonia (MNHN IM-2009-33734) and Papua New Guinea (MNHN IM-2009-22244) (Haplotype #18, Fig. 3 ). At the same time, the specimens collected in the south of New Caledonia are grouped in two different subclades, one of which also includes one specimen from Papua New Guinea, while the second one includes specimens from southern and northern New Caledonia and the Coral Sea, as well as one specimen from Papua New Guinea.

This suggests that despite a non-planktotrophic larval development, larval dispersal in A. hilgendorfi over time can take place across large geographic distances. The presence in New Caledonia of two highly divergent genetic groups (marked by green oval and green rectangle on Fig. 3 ), both with affinities with haplotypes from Papua New Guinea, suggests that larval exchange between these distant regions might have been quite frequent.

The species is variable in shell shape and colouration. Several morphs could be distinguished, mostly geographically determined and not necessarily distinct molecularly. This has resulted in the description of three subspecies. Moreover, morphologically very similar sympatric specimens may belong to different subclades. For a better characterization of the variability, we describe these morphs separately.

General distribution

Amalda hilgendorfi is broadly distributed in the West Pacific, recorded from Japan, the Philippines, Papua New Guinea and the Solomon Islands, eastern Australia, the Coral Sea, and New Caledonia.

Figures

Fig. 2. Molecular phylogeny of Amalda hilgendorfi (von Martens, 1897) produced by a Bayesian analysis of the Cox1 gene dataset. Outgroups and other species of Amalda have been removed from the tree. Only posterior probabilities ± 90% are shown. The unit for the scale is “substitutions/site”. The geographical distribution and morphological morphs are coded for each specimen: Phil =Philippines; Sol =Solomon Islands; PNG= Papua New Guinea; sNC = South New Caledonia; nNC = North New Caledonia; CS=Coral Sea. Numbers in parentheses at the tips of the tree link the MNHN IM voucher number to the corresponding photo on the right.

Fig. 3. Median-joining haplotype network based on Cox1 sequences of Amalda hilgendorfi (von Martens, 1897). Circles are colour coded by general location reported on the map and sized according to each haplotype’s relative abundance. Lengths of the lines connecting haplotype circles are proportional to the number of nucleotide differences among haplotypes. Green rectangle and oval encircle assemblages of haplotypes from New Caledonia (each including a specimen from Papua New Guinea).

Fig. 4. Amalda hilgendorfi (von Martens, 1897). A–D. Typical morph, Japan. A–C. MNHN IM-2019- 616, SL 53.1 mm. D. MNHN IM-2019-621, SL 50.0 mm. E–G. Philippine morph. E. MNHN IM-2007- 31950, SL 38.7 mm. F–G. MNHN IM-2007-33427, SL 40.3 mm. H–L. Solomon Islands and Papua New Guinea (PNG) morph. H. Solomons, MNHN IM-2007-33286, SL 52.6 mm. I. PNG, MNHN IM-2013- 44995, SL 51.3 mm. J. Solomons, MNHN IM-2009-11961, SL 32.9 mm. K. PNG, MNHN IM-2009- 22244, SL 63.1 mm. L. PNG, MNHN IM-2009-23330, SL 69.3 mm. M. Amalda tankervillii (Swainson, 1825), Venezuela, Isla de Margarita, 30–35 m, collection of Ron Voskuil, RV 0175, SL 80 mm. Photo by Ron Voskuil et al. (2011) (http://olivirv.myspecies.info/en/taxon-pages/ancillaria-tankervillii). Shells (except M, reduced) at the same scale.

Fig. 5. Amalda hilgendorfi (von Martens, 1897). A–F. Coral Sea morph. A-B. MNHN IM-2007- 33313, SL 66 mm. C. MNHN IM-2007-33289, SL 51.3 mm. D. MNHN IM-2007-33288, SL 60.3 mm. E. MNHN IM-2013-63901, SL 27-3 mm. F. MNHN IM-2007-33312, SL 25.9 mm. G–K. Northern New Caledonia morph = A. hilgendorfi richeri Kilburn & Bouchet, 1988. G. Holotype, MNHN IM-2000- 1378, SL 66.9 mm. H. MNHN IM-2009-11916, SL 65 mm. I. MNHN IM-2009-11918, SL 43.2 mm. J. MNHN IM-2007-42675, SL 48.7 mm. K. MNHN IM-2009-11966, SL 29 mm. Shells at the same scale.

Fig. 6. Amalda hilgendorfi (von Martens, 1897). Southern New Caledonia morph. A–D and E–H belong to two molecularly distinct subclades. A–B. MNHN IM-2007-43664, SL 30.6 mm. C. MNHN IM- 2007-43673, SL 34.2 mm. D. MNHN IM-2009-11964, SL 15.2 mm. E–F. MNHN IM-2007-43671, SL 33.8 mm. G. MNHN IM-2007-43674, SL 34.7 mm. H. MNHN IM-2009-11945, SL 31.6 mm. I. Norfolk Ridge, Banc Éponge, NORFOLK 1, stn DW 1693, 24°55′ S, 168°21′ E, 564–1144 m, SL 61 mm, specimen not sequenced, shell not to scale. Shells (except I, reduced) at the same scale.

Fig. 7. Radulae of species of Amalda H. Adams & A. Adams, 1853. A–G. A. hilgendorfi (von Martens, 1897). A–B. MNHN IM-2019-621 (shell on Fig. 4D). C. MNHN IM-2009-11961 (shell on Fig. 4J). D. MNHN IM-2013-58045, Papua New Guinea. E. MNHN IM-2013-19909, Papua New Guinea. F. MNHN IM-2013-63430, southern New Caledonia. G. MNHN IM-2007-33312 (shell on Fig. 5F). H. A. aureomarginata Kilburn & Bouchet, 1988, MNHN IM-2013-63421, specimen not sequenced, southern New Caledonia. I–J. A. miriky sp. nov. I. Miriky, stn CP3210, specimen not sequenced, Madagascar. J. Miriky, stn DW3215 (shell on Fig. 10J). K–L. A. cacao sp. nov., holotype, MNHN IM- 2000-35296 (shell on Fig. 11A–E).

Fig. 13. Radulae of species of Amalda H. Adams & A. Adams, 1853. A–C. A. montrouzieri (Souverbie, 1860). A–B. Île Ouen – Baie du Prony, LAGON, stn DW239, 22°24.3′ S, 166°57.8′ E, 42–43 m, SL 20.2 mm. C. Grand Récif Sud, LAGON, stn DW544, 22°50.8′ S, 166°48.5′ E, 25 m, SL 25.4 mm (shell on Fig. 12I). D–E. A. alabaster sp. nov. D. Holotype, MNHN IM-2007-43627, SL 26.7 mm (shell on Fig. 15A–B). E. MNHN IM-2007-43636, SL 17.6 mm. F. A. cf. alabaster, MNHN IM-2013-63897, SL 18.1 mm (shell on Fig. 15H–I). G. A. allaryi Bozzetti, 2007, MNHN IM-2013- 68272 (shell on Fig. 17A). H. A. coriolis Kilburn & Bouchet, 1988, MNHN IM-2013-63899 (shell on Fig. 18H–I).