Molluscan Research 32(2): 61–120 http://www.mapress.com/mr/
ISSN 1323-5818 Magnolia Press
Tonnoidean gastropods of French Polynesia A.G. BEU1*, P. BOUCHET2 & J. TRÖNDLÉ3 GNS Science, P O Box 30368, Lower Hutt, New Zealand 5040 2 Muséum National d’Histoire Naturelle, 55 rue Buffon, F-75231 Paris cedex 5, France 3 Attaché au Muséum National d’Histoire Naturelle, 55 rue Buffon, F-75231 Paris cedex 5, France *Corresponding author–Email:
[email protected] 1
Abstract The tonnoidean gastropod fauna of French Polynesia (54 species) includes 26 species recorded from the Austral Islands (including 10 from Rapa), 33 species from the Marquesas Islands, 39 from the Society Islands, 32 from the Tuamotu Islands, and 3 from the Tarava Seamounts. Most species have planktotrophic larval development and are distributed from East Africa to eastern Polynesia, but many common western Pacific species are not present. With the possible exception of Semicassis salmonea n. sp. (Cassidae), described from the Marquesas, and Gyrineum pusillum (Ranellidae), restricted to the Austral (and Tuamotu?) Islands in southeastern-most Polynesia, no species is endemic to any individual island groups, but several species with broad overall ranges are known from only one archipelago within French Polynesia. Three species (Monoplex intermedius, Septa peasei, Ranellidae; Distorsio graceiellae, Personidae) are much more common in the Marquesas Islands than further westwards. Three species of Bursidae (Bursa lamarckii, Bursina nobilis, Tutufa tenuigranosa) are recorded only from the Marquesas Islands, whereas the only record of Bursina fijiensis is from the Austral Islands. The two very similar species Bursa asperrima and B. cruentata have a complex distribution; only B. cruentata is common west of Hawaii, and only B. asperrima occurs east of Hawaii, but only B. cruentata has been collected at the Marquesas Islands. Ranella venustula is a synonym of Bursa rhodostoma. Neotypes are designated for Buccinum ponderosum Gmelin, 1791, B. nodulosum Gmelin, 1791, Cassis caputequinum Röding, 1798, C. denticulata Röding, 1798, C. glabra Röding, 1798, C. hamata Röding, 1798, Phalium edentulum Link, 1807, P. quadratum Link, 1807, Buccinum biarmatum Dillwyn, 1817, B. pantherina Dillwyn, 1817, Cassis tenuilabris Menke, 1828, and Dolium rufum Blainville, 1829, and lectotypes are designated for Buccinum cornutum Linnaeus, 1758, Murex bufonius Gmelin, 1791 and Cassis torquata Reeve, 1848. Key words: biogeography, Caenogastropoda, Gastropoda, distribution, new species, planktotrophic larvae, Semicassis, taxonomy
Introduction For several years one of us (AGB) has been identifying the collections of gastropods of the superfamily Tonnoidea gathered by the Muséum National d’Histoire Naturelle (MNHN), Paris, during expeditions in the Indo-West Pacific (e.g., Beu 1998; cruises listed by Bouchet et al. 2008, appendix 1). Most specimens have been collected in relatively deep water, and many represented new taxa or new records when they first came to light. However, in the Southwest Pacific, at least, the tonnoidean fauna is now well-known, by contrast with the rest of the South Pacific. The present report records the tonnoideans collected in French Polynesia (Fig. 1), with an assessment of their biogeographical significance, based on several expeditions of the Tropical Deep-Sea Benthos programme, with additions from material compiled by local collectors. These combined collections are much more thorough and representative than previous ones from French Polynesia. Apart from the one new species of Semicassis Mörch, 1853, most species recorded here occur widely throughout the Indo-West Pacific region. The nomenclature, type material, and geographical ranges of most ranellid, personid and bursid species are merely summarised here for the taxa covered previously by Beu (1998). The main interest of the fauna therefore lies in the biogeographical patterns, with composition and proportions of common and uncommon species differing in French Polynesia from those in the more familiar faunas of the main western Pacific island groups.
Tonnoideans in the history of molluscan exploration in French Polynesia It is probable that the European 18th century navigators who called in to Tahiti collected seashells there, but by the time such specimens reached the hands of the conchologists of the time
all trace of their geographical origin had been lost or had become the generalized ‘South Seas’. The first geographically accurate records of Tonnoidea from French Polynesia originated from Hugh Cuming's 1827–1828 visit to South Marutea and Anaa in the Tuamotus (St John 1940; Dance 1966: 147–157). This was the source of Broderip's (1833) Ranella affinis (= Bursa granularis) and R. pusilla (= Gyrineum pusillum), and Reeve's (1844) Ranella livida (= Bursa granularis) and R. venustula (= Bursa rhodostoma). Very few, if any, records seem to have followed in the 19th century, although G.B. Sowerby III (1897) described Lotorium armatum from the ‘Marquesas Islands?’, almost certainly in error (see below). Dautzenberg and Bouge (1933) published the first checklist of the molluscs of French Polynesia. Although their nomenclature is at times difficult to reconcile with the modern one, a critical evaluation of their list shows that 21 species of Tonnoidea were recorded. Again, few records were published in the decades that followed, until Bernard Salvat's foundation in 1971 of the field station that has now become CRIOBE in Moorea. This began a new era of acquiring and disseminating knowledge on the marine biodiversity of French Polynesia, starting with Salvat and Rives's (1975) Coquillages de Polynésie and Richard's (1985) Compendium, published on the occasion of the Fifth International Coral Reef Symposium. Richard (1985) recorded 33 tonnoidean species, of which 30 are confirmed here. A catalogue of the molluscs recorded from the Marquesas Islands was compiled by Tröndlé and Cosel (2005), summarising published records by Jardin (1858), Rehder (1969), Lavondes et al. (1973), Salvat and Rives (1975), Richard (1985, 1987), and Planes et al. (1995), and we refer to their catalogue below for earlier Marquesan records. Besides French scientists, the Smithsonian Institution's Harald Rehder (1907–1996) also made French Polynesia one of his favourite field destinations (see, e.g., Rehder 1969), and his field work resulted, among other things, in the discovery of Septa peasei in the Marquesas. New records continued to be
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62 accrued by local amateurs and visiting scientists, and Tröndlé and Boutet (2009) provided a critically compiled checklist of the marine molluscs of French Polynesia, including some new records based on MNHN expedition material (already with
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some identifications by AGB) and specimens in local private collections. Such records have been verified and are fully documented in the present review; 45 of their 46 species records are here considered valid.
FIGURE 1. Islands and main island groups of French Polynesia. Islands (upright type) and seamounts (Italics) from which tonnoideans are listed are shown in larger type; all others are in smaller type.
Material and methods The material that forms the backbone of this review originates from various deep-sea expeditions and coastal surveys. Detailed station data are listed in the Appendix for those stations where tonnoideans were collected, and with a few exceptions only the station numbers and abbreviated localities are included in the main text. Apart from some USNM material
and the material in private collections, CRIOBE and EPHE listed below, all material listed here is in MNHN. Three deep-sea cruises on board RV Alis were part of the MNHN–IRD Tropical Deep-Sea Benthos programme (see Bouchet et al. 2008 for context) and explored, in chronological order, the Marquesas (MUSORSTOM 9 Expedition, 18 Aug–11 Sept 1997; cruise leader B. Richer de Forges; see Richer de Forges et al. 1999), the Austral Islands (BENTHAUS Expedition,
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29 Oct–28 Nov 2002; cruise leader B. Richer de Forges), and the Tarava Seamounts, Society Islands, and the western part of the Tuamotu chain (TARASOC Expedition, 21 Sept–26 Oct 2009; cruise leader P. Bouchet). (The Tarava seamounts form a chain to the southwest of Tahiti, extending over 700 km, rising from a 4,000–4,500 m deep seafloor to a summit area 500–800 m below sea level; Clouard et al. 2003). Other sources that provided tonnoidean specimens from offshore waters include a geology cruise REMARQ to the Marquesas Islands (RV Alis, 23 Sept–7 Oct 2002; cruise leader G. Cabioch) and collections made in the 1980s throughout French Polynesia by Joseph Poupin on board RV Marara for the Service Mixte de Contrôle Biologique (SMCB). The sources of the records of coastal species include a couple of shore-based biodiversity surveys: ‘Atelier Marquises 1997’ (Ua Huka Island only, 16 Sept–19 Oct, R. Cosel, J. Tröndlé, and J. Tardy), and ‘Atelier Rapa 2002’ (Rapa Island only, 27 Oct–10 Dec 2002) (Lozouet et al. 2003, 2004). Collections made at the Marquesas Islands by a party including Clay Bryce (Western Australian Museum) and Kirstie Kaiser (Santa Barbara Museum of Natural History) during 17 Oct–1 Nov 1999 (Bryce 2000), of which part was presented to MNHN, include additional records of tonnoideans. However, the present review owes much to the private collections of Michel Boutet, Robert Gourguet, Jean Letourneux and Jean Tröndlé. Robert Gourguet kindly sent us photographs of at least one representative specimen of each of the recorded shallowwater species (5 March 2011), confirming their identification; several photographs have been received of the less common species and of subtly distinct taxa such as Gyrineum species. Robert Gourguet also sent us details of a few specimens (listed in text below) of rare species in other private collections. Full specimen data (number, and whether collected dead or alive) is not available for this material in private collections, but empty shells are specified in the lists provided, so most records apparently refer to a single live-collected specimen. The locality records have also been amplified by the material in the collections of CRIOBE (Centre de Recherches Insulaires et Observatoire de l’Environnement, Moorea), identified recently by JT, and EPHE (École Pratique des Hautes Études, Université de Perpignan, France), also identified by JT. A few further records, without individual localities but all collected in Moorea, are listed on the Moorea Biocode website (http://mooreabiocode.org), with specimen photographs on the Calphotos Berkeley website (http://calphotos.berkeley.edu/ fauna/sci-invertebrate.html), but no additional species records are listed on these sites, and we have not included them here. The Museo Naturalistico ‘Libero Gatti’ (2009) website also lists a number of records of tonnoideans (among many other molluscs) collected at Aratika, Fakarava, Niau and Toau Islands in the Tuamotu Group. These records have been added below where relevant. This website includes a later (2010) photographic listing of the species collected, allowing a minor correction to the tonnoidean identifications. Outside French sources, some material recorded from the Marquesas Islands by Rehder (1969) was also borrowed from the USNM for confirming (or, in some cases, not confirming) the identities of his specimens. A few other specimens have also been cited from the USNM collection. Conventions adopted in the text and abbreviations for institutions housing specimens referred to are as follows: Abbreviations AMNH—American Museum of Natural History, New York,
USA; AMS—Australian Museum, Sydney, Australia; ANSP—Academy of Natural Sciences of Philadelphia, USA; CRIOBE—Centre de Recherches Insulaires et Observatoire de l’Environnement, Moorea, French Polynesia; EPHE—École Pratique des Hautes Études, Perpignan, France; GNS—GNS Science, Lower Hutt, New Zealand; IRD—Institut de Récherche pour le Développement, Marseille, France; JL—Private collection of Jean Letourneux, Papeete, Tahiti; JT—Private collection of Jean Tröndlé, La Force, France; LACM—Los Angeles County Museum of Natural History, Los Angeles, USA; MB—Private collection of Michel Boutet, Papeete, Tahiti; MCZ—Museum of Comparative Zoology, Harvard University, Cambridge, USA; MNLG—Museo Naturalistico ‘Libero Gatti’, Italy (abbreviated reference to website; see References); MHNG—Muséum d'Histoire Naturelle de la Ville de Genève, Switzerland; MNHN—Muséum National d'Histoire Naturelle, Paris, France; NHMUK—Natural History Museum, London, UK; NMNZ—Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand; NMP—Natal Museum, Pietermaritzburg, Republic of South Africa; NMV—Museum Victoria, Melbourne, Australia; RG—Private collection of Robert Gourguet, Papeete, Tahiti; RGM—Netherlands National Museum of Natural History, Leiden, the Netherlands, geological collections; SMCB—Service Mixte de Contrôle Biologique, France (material in MNHN); UPSZTY—Uppsala University Museum of Evolution, Uppsala, Sweden; USNM—United States National Museum of Natural History, Washington, DC, USA. Dimensions of specimens are stated in the order H (= height), D (= maximum diameter). All specimens are assumed to be adult and collected as empty shells, unless the preserved or dried animal and/or the operculum is retained, in which case they are identified as ‘live’ in the lists of MNHN expedition material examined. Unless otherwise stated, all types listed under ‘Type material’ have been examined by AGB.
Systematics Class Gastropoda Caenogastropoda Superfamily Tonnoidea Suter, 1913 (1825) Family Bursidae Thiele, 1925 Genus Bursa Röding, 1798 Bursa Röding 1798: 128. Type species (by subsequent designation, Jousseaume 1881: 174): Bursa monitata Röding, 1798 (= Murex bufonius Gmelin, 1791, by first reviser’s action by Winckworth 1945: 137), present-day, Indo-West Pacific.
Bursa asperrima Dunker, 1862 Fig. 2A Bursa asperrima Dunker 1862: 238; Dunker 1863: 57, pl. 19, figs 3–4; Beu 1998: 146, fig. 45f; Tröndlé and Boutet 2009: 22; Beu 2010: 43, pl. 1, fig. 9; Severns 2011: 150, pl. 59, fig. 1.
Type material Bursa asperrima, lectotype NHMUK 1968566/1, designated by Yen (1942: 217); two paralectotypes NHMUK 1968566/2–3, all supposedly from ‘China’. The lectotype and paralectotypes were illustrated by Cossignani (1994: 43) and
64 include one specimen of B. asperrima and two of the restricted western Atlantic species B. grayana Dunker, 1862. The lectotype (Yen 1942: pl. 19, fig. 114; Cossignani 1994: 43, left fig. in lower row) is the specimen of B. asperrima illustrated by Dunker (1863: pl. 19, figs 3–4). The most probable type locality is Hawaii. French Polynesian material examined Austral Islands: BENTHAUS: stn DW1880, Marotiri Islets, 90–94 m (1 live); stn DW1894, W of Rapa, 100 m (1); stn DW1919, Neilson Reef, 140–390 m (1 live? juvenile); stn DW1926, President Thiers Bank, 50–90 m (2); stn DW1927, as last, 95–105 m (1); stn DW1931, as last, 200 m (1); stn DW1937, as last, 469–500 m (1); stn DW1996, Rurutu: Avera, 489–1050 m (2, with fragments of B. latitudo and B. rhodostoma). Rapa: Atelier Rapa: stn 6, Baie de Haurei, 42 m (1); stn 22, Cape Rukuaga, 18–22 m (1 live); stn 31, Point Mei, 6 m (2 live); stn 32, Vavai, 15–20 m (1 live); stn 36, Point Kauira, 27 m (1 + 1 live); stn 81, baie Pake, intertidal (1). Specimens were collected alive around the Austral Islands in 6–90 m, and possibly in 140 m. Society Islands: B. asperrima was reported from the Society Islands by Tröndlé and Boutet (2009: 22); Tahiti: Afaahiti, under dead coral (MB); Faaone, reef, 0.5–1 m, under coral (JT); Mahaiatea (JT); Mahina, coastal reef, under coral (JT); Papeete (JT); Pueu, barrier reef,
