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Neogene Fossil Tonnoidean Gastropods of Indonesia

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Neogene Fossil Tonnoidean Gastropods of Indonesia Neogene fossil tonnoidean gastropods of Indonesia A.G. Beu Beu, A.G. Neogene fossil tonnoidean gastropods of Indonesia. Scripta Geologica, 130: 1-186, 327 figs., Leiden, September 2005. A.G. Beu, Institute of Geological and Nuclear Sciences, P O Box 30368, Lower Hutt, New Zealand ([email protected]). Key words – Gastropoda, Tonnoidea, Neogene, Indonesia, taxonomy, biogeography. Tonnoidean gastropods in K. Martin’s and other collections in the Nationaal Natuurhistorisch Museum, Leiden (and a few other minor collections) are reidentified and classified. The resulting fauna of 99 species is very similar to that of the Indo-West Pacific today, 70% of species still inhabiting the Indone- sian region. Species endemic to the Miocene and Pliocene rocks of Indonesia are Bursa sangirana sp. nov., and two new (unnamed) species similar to Bursina ignobilis (Beu); Cassis depressior Martin and C. preange­ rensis Martin; Cypraecassis denseplicata (Martin) and an unnamed species of Cypraecassis; Sconsia martini van Regteren Altena and S. pulchra Pannekoek; Echinophoria vandervlerki Martin (possibly a synonym of E. wyvillei (Watson)); Phalium menkrawitense Beets and P. rembangense (Martin); Distorsio denseplicata van Regteren Altena and D. djunggranganensis (Martin); Biplex magnifica (Martin), B. pamotanensis (Martin) and B. perliberalis (Beets); Cymatium (Monoplex) gembacanum (Martin) (?=C. exaratum (Reeve)), C. remban­ gense (Wanner & Hahn), and C. tjaringinense (Martin); Cymatium (Ranularia) pseudopyrum (Martin) and an unnamed species of C. (Ranularia); Cymatium (Septa) dharmai sp. nov.; Sassia (Cymatiella) fennemai (Martin), S. (C.) menkrawitensis (Beets) and an unnamed species of S. (Cymatiella); Eudolium erbi (Haanstra & Spiker), E. errabundum (Beets) and E. pamotanense (Martin); Malea(?) papuana (Beets); and Sconsodolium (gen. nov.) rembangense (Pannekoek). These 30 species (30% of the recorded fauna), and the generic groups Sconsia, Sconsodolium and Sassia (Cymatiella), are all “additional” tropical western Pacific taxa that became extinct before the present day (Sconsia and Sassia (Cymatiella) only locally), as a result of Pleistocene climate change. Species previously included in Bufonaria Schumacher prove to belong in two distinct genera; spe- cies closely related to “Bursa” nobilis have a subcentral (rather than mid-left edge) opercular nucleus and are reclassified in Bursina Oyama. “Sconsia” rembangensis Pannekoek is an elongate, axially ridged cassid with a coarsely plicate inner lip; the new genus Sconsodolium is proposed for it. Galeodea bituminata (Martin) and G. carolimartini Beets are both earlier names for the western Pacific species (originally de- scribed in the Recent fauna) previously known as G. echinophorella Habe. Eudolium javanum (Martin) is an earlier name for the Indo-West Pacific species (originally described in the Recent fauna) previously known as E. pyriforme (G.B. Sowerby 3rd), whereas E. bituminata Martin is a synonym of the near-cosmo- politan species E. bairdii (Verrill & Smith). Ranella spinosa var. granosa Martin is either an earlier name for the western Pacific species (originally described in the Recent fauna) previously known as Bufonaria perelegans Beu, or a closely similar, but distinct, species. Purpura bantamensis Martin, Cassis tegalensis Martin, Dolium losariense Martin, and Tritonium verbeeki Boettger are all synonyms of Cymatium (Linatella) cingulatum (Lamarck). Contents Introduction ................................................................................................................................................................. 2 Localities ........................................................................................................................................................................ 3 Pliocene-Pleistocene extinctions of Indonesian fossils ...................................................................... 5 Abbreviations ............................................................................................................................................................. 7 Systematic palaeontology ................................................................................................................................... 8 Beu. Neogene tonnoideans of Indonesia. Scripta Geol., 130 (2005) Acknowledgements ........................................................................................................................................... 127 References ................................................................................................................................................................ 128 Appendix: Taxa removed from the Tonnoidea .................................................................................. 150 Index of species names and synonyms .................................................................................................. 153 Introduction The Neogene molluscan fauna of Indonesia is important for the understanding of world molluscan biogeography, as it provides the only extensive, diverse fossil record of the tropical Neogene Indo-West Pacific fauna. For tonnoidean gastropods, with their very wide distributions conferred by a long planktotrophic larval life, this fossil record provides the only evidence of whether or not a species was present in the tropical Indo- West Pacific area relatively early in Neogene time and whether any “additional” Pacific taxa have become extinct before the present day. Of course, it safely can be assumed that not all species that inhabited Indonesia during Neogene time are represented in the fossil record, but the Indonesian fauna is the only record there is. Taxonomists also routinely should investigate the possibility that species living in the Indo-West Pacific region received names earlier as fossils in Indonesia than they did in the Recent fauna. Three examples are provided here of species of which the earliest name was provided for an Indonesian fossil, supplanting a name based on Recent material; Galeodea bitu­ minata (Martin, 1933) (= G. echinophorella Habe, 1961), Eudolium javanum (Martin, 1879) (= E. pyriforme G.B. Sowerby 3rd, 1914) and Bufonaria granosa (Martin, 1884) (?= B. perel­ egans Beu, 1987). A popular introduction to some of the richly diverse molluscan fossils of Indonesia was provided by Dharma (2000). The aims of this review are to provide a classification for the endemic tonnoidean species named from the Indonesian Neogene, and to clarify the fossil record of the Re- cent tropical Indo-West Pacific fauna. It includes the material described in many papers by Karl Martin, the “Kendeng beds” (not a currently used stratigraphical division) material reviewed by van Regteren Altena (1942), much material described by Beets, the Nias Island fauna described by Wissema (1947), and some undescribed collections, in other words, all the material available in the Nationaal Natuurhistorisch Museum/ Naturalis, in Leiden (RGM). Extensive additions have been made from the collection of Bunjamin Dharma, in Jakarta, Indonesia, and from the Indonesian collections of Profes- sor Elio Robba (Università di Milano-Bicocca). The family Ficidae is not included here, as Riedel (1994) has removed it to a superfamily of its own. A very clear distinction is obvious in Indonesia (as in most other parts of the world) between the Paleogene and Neogene molluscan faunas. The Paleogene one consists almost entirely of extinct species, most of which have no obvious relatives in the Recent fauna. In contrast, the Neogene fauna (at least, of tonnoideans) consists largely of living species, along with a relatively small number of Indonesian Neogene endemic species and one genus, i.e., additional Indo-West Pacific taxa that did not survive into Recent times. In addition to the species listed here, many referred by Martin (in numerous papers) to Hindsia, “Argobuccinum”, etc., have been referred to the Ranellidae in the past, but actually are Buccinidae belonging in or closely related to Nassaria Link, 1807. These incorrectly assigned species are not included in the present catalogue. Most of the Indonesian molluscs included here also were included in the large computer catalogue Beu. Neogene tonnoideans of Indonesia. Scripta Geol., 130 (2005) 3 by Skwarko & Sufiati (1994), but their list is highly uncritical in classification, taxonomic position and synonymy. The material has been reidentified for the present paper by examination of the actual collections. An appendix revises the taxonomic position of a few of the Buccinidae, Turbinellidae(?) and Conidae described in the Indonesian fauna (originally proposed within the Tonnoidea) by Martin and Beets, and lists the Indone- sian fossil species of Colubraria (Buccinidae) that I am aware of. Localities Localities for the “Kendeng beds” faunas were listed by van Regteren Altena (1938, pp. 248-262), in two large tables that documented, respectively, the Mijnwezen Collec- tion localities and Cosijn’s Collection localities for Pleistocene molluscs from the upper Kendeng beds. These localities are referred to in the present work only by the numbers cited by van Regteren Altena and (as in van Regteren Altena’s lists) are prefixed by M for Mijnwezen localities. Cosijn’s localities have not been cited, as this section of the RGM collection at present bears registration numbers, but no localities (the localities can be found from the register); only the registration numbers and numbers of specimens are cited here. This is of little
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