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Marine Mollusca of isotope stages of the last 2 million years in New Zealand. Part 4. Gastropoda (Ptenoglossa, Neogastropoda, Heterobranchia) AG Beua a GNS Science, Lower Hutt, New Zealand

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To cite this Article Beu, AG(2011) 'Marine Mollusca of isotope stages of the last 2 million years in New Zealand. Part 4. Gastropoda (Ptenoglossa, Neogastropoda, Heterobranchia)', Journal of the Royal Society of New Zealand, 41: 1, 1 — 153 To link to this Article: DOI: 10.1080/03036758.2011.548763 URL: http://dx.doi.org/10.1080/03036758.2011.548763

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Marine Mollusca of isotope stages of the last 2 million years in New Zealand. Part 4. Gastropoda (Ptenoglossa, Neogastropoda, Heterobranchia). AG Beu* GNS Science, Lower Hutt, New Zealand (Received 1 December 2009; final version received 10 June 2010)

Three new : (Pliciscala) flemingi (late Nukumaruan-early Castlecliffian, OIS71- 29?, Wanganui); Kuroshioturris putere (Opoitian-Waipipian, Westland and Hawke’s Bay), Antimelatoma waimea (Kapitean-Opoitian, Westland); two new genera: Cryptofusus (Turbi- nellidae) for New Zealand species formerly referred to Finlay (Australian); Onoketoma (, ) for ‘‘Insolentia’’ solitaria King, bathyal Nukumaruan, Palliser Bay. Xymene (sensu stricto; separated again from Axymene, Xymenella and Zeatrophon) is useful biostratigraphically in near-shore facies: X. drewi (Hutton) (based on Marwick’s lectotype), Mangapanian-late Nukumaruan; X. expansus (Hutton), late Nukumaruan-early Castlecliffian (to OIS19); X. plebeius (Hutton), OIS17-Recent. New fossil records from Pleistocene (OIS7) deposits at Hokianga Harbour, Northland: Agnewia tritoniformis (Blainville), physis (Linne´ ), Philine tepikia Rudman. Other biostratigraphically useful species: Zeatrophon bonneti (Cossmann) (Nukumaruan-OIS9; 7?); delicatulus Powell, caudatum Powell (both OIS25-19, 17?). New synonymy: Otahua Marwick, 1948 Iredale, 1936; (Eucominia) marlboroughensis Powell C. elegantula (Finlay), Nukumaruan- Recent; Antiguraleus Powell, 1942 Iredale, 1918; Antiguraleus depressipirus (Beu) Propebela ula (Watson); Mo¨ rch, 1853 is possibly an earlier name for Liracraea Odhner, 1924, but Liracraea is retained; Antimelatoma ahiparana and A. otagoensis of Powell A. buchanani (Hutton); Dall, 1924Asperdaphne Hedley, 1922; most Australasian ‘‘Asperdaphne’’ species are assigned to Verrill. revised: zelebori (Dunker); Opalia () nympha (Hutton) (Mangapanian-Nukumar- uan); cookiana (Dell, 1956) (OIS23-Recent); Cominella (Eucominia) incisa (Hutton), Nukumaruan, Hawke’s Bay; pangoides (Beu), Waipipian; Iredalula striata (Hutton), Kapitean(?)-Castlecliffian (-OIS11) distinguished from I. alticincta (Murdoch & Suter), Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Nukumaruan-Recent; zealandica (Hutton) (OIS13-Recent); () olsoni Beu, Nukumaruan; Kuroshioturris angustata (Powell), Mangapanian-Recent; Austrotoma aguayoi (Carcelles) (Recent, SW Atlantic); Struthiolariopsis ferrieri (Philippi), , Quiriquina, Chile (); Mitrellatoma angustata (Hutton), Nukumaruan; gypsata (Watson), Recent, E New Zealand, removed from Paracomitas; Paracomitas protransenna (Marshall & Murdoch) (Nukumaruan), P. gemmea (Murdoch) (Castlecliffian) and P. augusta (Murdoch & Suter) (P. augusta powelli Dell; Recent); Aoteadrillia wanganuiensis (Hutton) (chordata Suter and many other synonyms; Nukumaruan-Recent), A. finlayi (Powell) and A. apicarinata (Marshall & Murdoch) (Nukumaruan); Bulla quoyii Gray (OIS7-Recent); Bulla australis Fe´ russac, Atlantic,B. mabillei Locard; Philippia lutea (Lamarck), Nukumaruan-Recent. Keywords: biostratigraphy; Castlecliffian; dispersal; Haweran; Holocene; Mollusca; new ; new species; New Zealand; Nukumaruan; oxygen isotope stages; Pleistocene; Pliocene; time scale; .

*Email: [email protected]

ISSN 0303-6758 print/ISSN 1175-8899 online # 2011 The Royal Society of New Zealand DOI: 10.1080/03036758.2011.548763 http://www.informaworld.com 2 AG Beu

Introduction The obvious new, larger taxa and some This paper is the fourth part of a revision of the previous misidentifications have been revised in taxonomy and biostratigraphy of New Zealand these four papers, but there is little doubt that fossil Mollusca of the last two million years, this is merely the beginning of recognising the with comments on related Recent and earlier true relationships of the late Neogene mollus- Pliocene taxa. The first report (Beu 2004) can fauna of New Zealand. Further taxonomic reconsidered the generic positions or specific revisions and reidentifications undoubtedly will determinations of some New Zealand late bring to light an increasing list of molluscs that Neogene to Recent molluscs, including mi- extended their ranges to Wanganui Basin grants from eastern and the tropical briefly during the many glacial-interglacial Pacific that appeared only briefly in New oscillations of late Pliocene-Pleistocene time. Zealand during interglacial periods. The second (Beu 2006) reconsidered time ranges and Abbreviations and conventions taxonomy of bivalves over this period, includ- ing further warm-water migrants. Part 3 (Beu Collections and institutions housing specimens 2010) commenced reconsideration of the time listed here are abbreviated: ranges and taxonomy of some gastropods over S22/f123, etc. Locality numbers in the New the same period.The present report is a recon- Zealand Fossil Record sideration of the time ranges and taxonomy of Electronic Database (FRED; the remaining gastropods over the same period, http://data.gns.cri.nz/fred/), with time ranges recorded in oxygen isotope administered by the Geologi- stages (OIS) where possible. Because of space cal Society of New Zealand; considerations, a number of further gastropods map sheet number in NZ Map Series 260 (1: 50,000) requiring revision have been deleted from this followed by the individual work (partially listed below). A summary paper fossil locality number within will follow. Time ranges have been compiled that sheet. (Offshore samples from the detailed faunal lists provided for the are assigned record numbers formations of Wanganui Basin by Fleming based on their latitude and (1953), assigned to individual oxygen isotope longitude.) stages in the correlation diagram of cycles in R11/742729, etc. Grid references to fossil

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 localities on map sheets of Wanganui Basin compiled by Carter & Naish NZ Map Series 260 (1: (1999). The faunas included in this overview, 50,000); map sheet number their ages and their oxygen isotope stage followed by grid reference correlations were summarized in Part 1 of this on that sheet. series of papers (Beu 2004), modified slightly in AIM Auckland War Memorial the introduction to Part 2 (Beu 2006), and are Museum, Auckland. not repeated here. Once again this work is AMSAustralian Museum, Sydney. founded on the seminal works on Wanganui AUGD Geology Programme, School of Geology, Geography and Basin by Fleming (1947,1953), and much of Environmental Sciences, Uni- what the present ‘Wanganui Basin working versity of Auckland group’ has published (particularly Beu 2004, BMNH Natural History Museum, 2006; Abbott et al. 2005; Naish et al. 2005; London. Pillans et al. 2005) is based on Fleming’s CMC Canterbury Museum, Christ- brilliant foundation. church [New Zealand]. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 3 GNSInstitute of Geological & Dimensions of specimens are cited in milli- Nuclear Sciences, Lower Hutt; metres, in the consistent order H (height), individual collection abbrevia- D(diameter). tions are used throughout the The standard abbreviations for points of the text: compass (N north, S south, E east, GSbulk collection locality W west, NE, SSW, etc.) are used to abbre- numbers; RM New Zealand Recent viate locality details.Unless otherwise stated, I Mollusca registration have examined the type material cited below numbers; under ‘Type material’. TM type Mollusca registra- tion numbers; WM non-New Zealand Moll- Locality details usca registration numbers. Detailed locality records are listed below for the MNHN Muse´ um National d’Histoire less common species, for which localities help to Naturelle, Paris. establish accurate time range. Full locality data NHMW Naturhistorisches Museum are not listed for common, widespread species. Wien, Vienna. NMNZ Museum of New Zealand Te Papa Tongarewa, Wellington. Biostratigraphy NMV Museum Victoria, Melbourne. SMF Senckenberg Museum, Fran- Figure 1 is a time-scale diagram for the last 2 kfurt (Main). million years in New Zealand, repeated from VM Victoria University of Wellin- Beu (2004, fig. 4; 2006, fig. 1), apart from gton Geology Department, deleting the now out-dated PlioPleistocene Mollusca Specimen registra- boundary. Figure 2 shows the time ranges in tion numbers. oxygen isotope stages of most of the gastropods V311, etc. Victoria University of discussed in the present paper and previous Wellington, School of Earth papers (Beu 2010). Some taxa with earlier time Sciences and Geography, ranges or no biostratigraphical utility are dis- bulk collection locality num- cussed in the text but not included in Fig. 2. As bers. in Beu (2006), I adopt the time scale of Cooper Other abbreviations and conventions: (2004) and the definitions of local stages adopted by me (Beu in Cooper 2004; Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 MPT Mid-Pleistocene transition 2006:155): HaweranCastlecliffian boundary (the change from predomi- at Rangitawa Tephra, 0.34 Ma, in OIS10 (in nantly 41 ka to roughly 100 Rangitawa Stream, Rangitikei valley); ka temperature and sea-level CastlecliffianNukumaruan boundary at Oto- cycles, between 1 and 0.65 toka Tephra, 1.63 Ma, in OIS57 (on the Ma). Wanganui coast east of Ototoka Stream OISOxygen isotope stage (OISis mouth); NukumaruanMangapanian bound- preferred here to the alter- ary at the base of Hautawa Shellbed, 2.4 Ma, native, MIS, ‘marine isotope in OIS97 (proposed but not formally defined, stage’, as the isotopes involved are exclusively of oxygen and, on Hautawa Road, between Turakina and at least by correlation, the Murimoto valleys, north of ; Beu ‘stages’ can be applied to non- in Cooper 2004:217); and Mangapanian marine as well as marine Waipipian boundary at the base of Mangapani rocks). Shellbed, 3.0 Ma (in Mangapunipuni Stream, 4 AG Beu

GEOMAGNETIC OXYGEN ISOTOPE STAGE POLARITY TIME-SCALE NZ STAGE EPOCH AGE,Ma 0 2 HOLOCENE 4 5

6 HAWERAN 7

LATE 8

9 0.34Ma 10 RangitawaTephra 11 12

13 0.5 14

MIDDLE BRUNHES, C1n 15

16 17

18 mid-Pleistocene 20 transition, c.920-640ka 21 22 CASTLECLIFFIAN 1r 23 24 26 1.0 28 PLEISTOCENE Jaramillo 30 1n 32 Cobb 34 Mountain 36 37 38 40 42

2r 44 EARLY 46 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 48 1.5 50 52 53 54 55 MATUYAMA, C1r 1.63Ma 58 Ototoka Tephra 60 61 62 63 64 65 Réunion NUKUMARUAN 69 C2n 70 (part) 71 72 2.0 Figure 1 New Zealand Pleistocene-Holocene time scale (Beu 2006, fig. 1). Correlation of oxygen isotope stages with the geomagnetic polarity time-scale modified slightly from Carter & Naish (1999). Pliocene- Pleistocene boundary now at 2.59 Ma, below base of diagram; Late Pleistocene is applied only to the Last Glaciation, OIS2. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 5

SPECIES STAGE HAWERAN CASTLECLIFFIAN NUK

Stiracolpus huttoni, Xymene drewi, frazeri, Paracomitas protransenna, Aoteadrilliafinlayi, A.apicarinata, Taxonia suteri Opalia flemingi ? Calliostoma nukumaruense ? Waipipian Maoricrypta profunda ? zealandica,Taniella planisuturalis ? Zeatrophon bonneti Xymene expansus ? Xymene plebeius Paracomitas gemmea ? ? Paracomitas augusta Stiracolpus vigilax Maoricrypta costata Maoricrypta youngi Zelippistes benhami, Stephopoma roseum Acirsa cookiana labiata, Agnewia tritoniformis, pustulosum, , Bulla quoyii

Ponderia zealandica Agnewia kempae Aeneator delicatulus, Buccinulum caudatum Iredalula striata Philine tepikia Cantharidella tessellata ? varia ?

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Pelicaria vermis, Stiracolpus symmetricus, Aoteadrllia wanganuiensis

OXYGEN ISOTOPE STAGE 3 5 7 9 11 13 15 17 19 21 23 25 29 33 37 41 45 49 53 57 61 65 69 71 mpt AGE, m.y. 0 0.5 1.0 1.5 2.0 PERIOD PLEISTOCENE

Figure 2 Time ranges of Pleistocene-Recent gastropods discussed in this paper and in Beu (2010), in oxygen isotope stages (additional to Beu 2004, fig. 5; Beu 2006, fig. 2). Time scale as in Fig. 1. Time ranges derived from correlation of Wanganui formations (Fleming 1953; Abbott & Carter 1999; Abbott et al. 2005) with oxygen isotope stages, slightly modified from Carter & Naish (1999) following Pillans et al. (2005). Approximate duration of the Mid-Pleistocene Transition (c. 920-640 ka) from predominantly 41 ka cycles to 100 ka cycles shown by bold bar labelled ‘‘MPT’’. NUK=Nukumaruan Stage; Castlecliffian/Nukumaruan boundary at 1.63 Ma. 6 AG Beu

Waitotara valley). As I have pointed out before Figure 2 is constructed to demonstrate: (Beu 2004:155), these New Zealand local stage boundary definitions are critical for allowing 1. A group of species (Stiracolpus huttoni, the recognition of the stages by the criteria Struthiolaria frazeri, Xymene drewi, Para- identified in the present paper and in previous protransenna, Aoteadrillia finlayi, papers in this series. The stages would not be A. apicarinata) and the genus Taxonia, all identifiable by biostratigraphical proxies if of which became extinct at the end of different boundary positions were adopted. Nukumaruan time, and so provide reliable The PlioPleistocene boundary, as defined criteria for identifying the Nukumaruan in 1984 in the Vrica section, southern Italy, falls Castlecliffian boundary (with the proviso within OIS65, at 1.81 Ma, but has been that a form closely resembling Aoteadrillia removed from the figure, as the International finlayi still occupies bathyal depths along Union of Geological Sciences has recently (July central eastern New Zealand). 2009) ratified a new definition of the Pliocene 2. Longer-ranging species that became extinct Pleistocene boundary at the Global Stratotype during CastlecliffianHaweran time: Section and Point (GSSP) of the Gelasian Stage Calliostoma nukumaruense, in OIS17; in the Monte San Nicola section in Sicily, Maoricrypta profunda, in OIS19 or possibly within OIS103, with a date of 2.588 Ma, only 17; Trivia () zealandica and Ta- 1 m above the GaussMatuyama magnetic niella planisuturalis, which both retreated reversal in the Monte San Nicola section. The from a Nukumaruanearly Castlecliffian Gelasian Stage is now the lowest stage of the last appearance in central New Zealand to Pleistocene, and the former PliocenePleisto- be last known in OIS15(?) at Ohope Beach, cene boundary at Vrica is now defined as the Whakatane; Iredalula striata in OIS11; GSSP of the Calabrian Stage. The Pliocene Zeatrophon bonneti in OIS9. Pleistocene boundary (as judged by the position 3. The evolving lineages of Xymene drewi of the GaussMatuyama reversal) now falls (MangapanianNukumaruan), X. expansus between the Parihauhau and Te Rama Shell- (late NukumaruanOIS19) and X. plebeius beds of Fleming (1953), as shown in the (OIS17 Recent), and of Paracomitas correlation diagram by Carter & Naish protransenna (Nukumaruan), P. gemmea (1999), high in the Mangapanian succession in (Castlecliffian; OIS23 14?, limited by its Wanganui Basin. This position is about 200 ka, restriction to offshore siltstone facies) and

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 or four or five glacialinterglacial cycles, earlier P. augusta (Recent). than the first obvious cooling in the New 4. Short-ranging species of biostratigraphical Zealand succession, identified by the appear- utility during CastlecliffianHaweran time: ance of the Zygochlamys delicatula (Hutton, Stephopoma roseum (OIS13 and 9 at Wan- 1873) fauna in Wanganui Basin (Fleming 1944), ganui; OIS7 and Recent in NE North defining the base of the Nukumaruan Stage. It Island); Stiracolpus vigilax (OIS11 9at should be noted that this new definition equates Wanganui; OIS7 at Te Piki, near East Cape), the PliocenePleistocene boundary with the Maoricrypta costata (OIS29 5a at Wanga- Neogene boundary, but the term nui, Recent in NE ), M. youngi Neogene is used throughout this paper in the (OIS11 Recent), Zelippistes benhami (OIS usual sense of Cenozoic stratigraphers working 13 and 9 at Wanganui; Recent in NE North with marine successions (e.g. McGowran et al. Island), Semicassis labiata, Argobuccinum 2009), and as usually used in New Zealand, that pustulosum, Agnewia tritoniformis, Hydatina is, extending from the end of time physis, and Bulla quoyii (all OIS7 Recent), until the present day. Acirsa cookiana (OIS23 Recent), Opalia Marine Mollusca of isotope stages of the last 2 million years in New Zealand 7

flemingi (late NukumaruanCastlecliffian, (2004, 2006) indicate that only seven events OIS31, 29?), Ponderia zealandica (OIS13 occurred during the MPT. Most obvious origi- only; Recent in Cook Strait and near East nation and events during the last 2 m Cape), Agnewia kempae (OIS7 only, at Te yr occurred either at the end of Nukumaruan Piki), Aeneator delicatulus and Buccinulum time or since the MPT (23 events), during the caudatum (both OIS25 17), and Philine period of high-amplitude glacial-interglacial tepikia (OIS9 and 7 only). cycles. The end-Nukumaruan extinction appar- 5. Species previously thought to have short ently resulted from closure of the Manukau (HaweranRecent) time ranges, now seaway and the consequent decrease in larval known to have appeared earlier and to have transport to Wanganui Basin. Data from unreliable, facies-controlled time ranges: gastropods (Fig. 2) demonstrate that seven Cantharidella tessellata (earliest record late further events occurred during the MPT, Nukumaruan); Risellopsis varia (earliest whereas a further 20 occurred since then, record OIS25?, Teer Formation, Cascade, making totals of 14 MPT and 43 post-MPT SWestland). events. These numbers confirm that the biota 6. Species previously thought to consist of was more drastically affected by the large-scale numerous taxa with short ranges of high glacial-interglacial cycles of the late middle and biostratigraphical utility during this period, late Pleistocene than by the more gradually but each interpreted in these papers as changing temperature regime of the MPT. consisting of one species that ranged right through the period and provides no bios- tratigraphical utility: Stiracolpus symmetri- Other taxa useful in biostratigraphy cus, Pelicaria vermis, and Aoteadrillia Still further molluscan taxa actually or poten- wanganuiensis. tially are useful for biostratigraphy in Nuku- maruanHaweran rocks. They have not been The biostratigraphically useful taxa in Fig. investigated in these papers, because of space 5 supplement those shown and discussed pre- and time constraints, or because further taxo- viously by me (Beu 2004, fig. 5; 2006, fig. 2), nomic revision is required before they are useful together providing the means of distinguishing in biostratigraphy (e.g. and Splen- Nukumaruan from Castlecliffian faunas, and , Conoidea; Mesopeplum, Pectinidae). of identifying a number of the oxygen isotope Some have well-established biostratigraphical

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 stages during CastlecliffianHaweran time. utility (e.g. Aethocola species). They are listed briefly here, as an indication of further work required; no doubt others remain unrecognized. Mid-Pleistocene transition Several of the listed species (identified with an As noted previously (Beu 2006:156), one of the asterisk*) are small molluscs, under ca 6 mm in aims of this series of papers is to recognize greatest dimension, constituting the majority of evidence for origination and extinction caused the Mollusca, but ignored in the present works by temperature change during the last 2 m yr in favour of taxa identifiable in the field. and, in particular, to what degree origination and extinction coincided with the mid-Pleisto- cene transition (MPT) (ca 1.00.65 Ma; taken Polyplacophora here as occupying OIS23 17), when glacial- interglacial cycles changed gradually from the 1. Cryptoconchus marwicki Bucknill, 1928 41 ka period of Plioceneearly Pleistocene time (Beu & Maxwell 1990:333, pl. 43j, l), limited to the eccentricity-driven, roughly 100 ka to Castlecliffian rocks (records from OIS period of the last Â800 ka. Data from Beu 15, 13). 8 AG Beu

Gastropoda (early NukumaruanRecent) (Beu & Max- well 1990:347, pl. 47l, p), the largest and 2. *Scissurona fossilis Laws, 1940, Mangapa- lowest-spired form (diameter 1523 mm), nian and Nukumaruan; recorded from almost smooth, with no subsutural row of Wilkies Shellbed at Wilkies Bluff and beads and a smooth base. This genus seems Nukumaru Brown Sand at Nukumaru to represent gradual evolution from the Beach (Laws 1940); occurs also in Waipuru strongly sculptured shell of the monileine Shellbed. genus Antisolarium to be able more readily 3. Haliotis (Sulculus?) powelli Fleming, 1952 to live infaunally and to occupy the mobile (Beu & Maxwell 1990:347, pl. 47c), occurs sand habitat occupied, convergently, by in Castlecliffian rocks at Ohope Beach Umbonium in the N Pacific. Zethalia was (OIS15?) and in the Komako area, Po- included in tribe Monileini, rather than hangina, eastern Wanganui Basin, Nuku- Umboniini, by Hickman & McLean maruan (Carter 1972). (1990:126) on anatomical grounds, and 4. Emarginula haweraensis Powell, 1931, the although Williams et al. (2010) did not largest New Zealand Emarginula species; recognize these tribes, they demonstrated occurs in Waipipian to Nukumaruan rocks that Umbonium and Zethalia are distinct (Waipipi shellbeds, type; Okauawa Stream genera on the basis of molecular phylo- and Kaiwaka SRoad, Hawke’s Bay, late geny. Nukumaruan). 7. Pliconacca denticulifera (Marwick, 1924), 5. *Brookula funiculata Finlay, 1924, re- apparently ranging from early Nukumar- corded only from late Nukumaruan to uan (Kuripapango, inland Hawke’s Bay, Haweran rocks (Waipuru Shellbed to GS12687, U20/f10A) to Recent (see Maji- Landguard Sand, OIS 9). Numerous un- ma [1989:63] for reference to Pliconacca named Recent species of Brookula are Cossmann & Martin in Martin, 1914; known from the NZEEZ (Spencer et al. Majima specifically mentioned the Mio- 2009:201) and it is quite feasible that B. cene species ‘Uberella’ cicatrix Marwick, funiculata is still living. 1931, which seems to have been ancestral 6. Zethalia species (, Umboniinae): to P. denticulifera). an excellent example of a long-continued, 8. plauta Beu (2008:313, fig. anagenetic lineage, with arbitrary subdivi- 17EG) (Castlecliff, OIS19, to Recent);

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 sion into species. Currently accepted spe- living records are from off W Northland cies are Z. russelli Marwick, 1965 only. (Opoitian) (Beu & Maxwell 1990:289, pl. 9. * treadwelli (Hutton, 1885), lim- 36e, h), a small species (diameter 67 mm) ited to Castlecliffian rocks at Wanganui; with the tallest in the genus, a but apparently no specimens have been subsutural row of small beads, another collected for more than 100 years. How- periumbilical row of small beads, and the ever, as with Brookula species, numerous umbilicus not quite filled by a lobe of the unnamed species of are known inner callus forming a spiral plug; Z. from the Recent fauna of the NZEEZ coronata Marwick, 1948 (Waipipianearly (Spencer et al. 2009:205) and one of these Nukumaruan), intermediate in size (dia- could well be M. treadwelli. meter 1017 mm) and with small beads in a 10. Margineulima christyi (Marwick, 1924) subsutural row, but with a completely (Beu & Maxwell 1990:328, pl. 41h; Max- filled umbilicus and smooth base; and Z. well 1992:120), supposedly a Mangapa- zelandica (Hombron & Jaquinot, 1854) nianNukumaruan restricted species, this Marine Mollusca of isotope stages of the last 2 million years in New Zealand 9

large, distinctive species was recorded ough Formation in the Awatere valley. from Castlecliffian rocks at Wanganui by The superabundant living species A. glans Fleming (1953:188, 199): Kaimatira Pu- (Ro¨ ding, 1798) (Beu & Maxwell 1990, 364, mice Sand (OIS 25; in GS 4148, R22/f7414) pl. 49c) is not known fossil earlier than and Kaikokopu Shellbed (OIS 19; in Castlecliffian, but the significance of this is GS4156, R22/f7417). uncertain, as specimens very similar to its 11. species: Marshall & Burch (2000) supposed ‘form’ A. chathamensis (Finlay, revised Murexsul species (as )in 1928) occur uncommonly in Nukumaruan the modern fauna and some late Plioce- rocks of Wanganui and Hawke’s Bay, in nePleistocene fossil records. The rela- near-shore facies. A. taitae Marwick, 1924 tively small species M. espinosus (Hutton, (Beu & Maxwell 1990:324, pl. 42k) is 1885) is apparently limited to Mangapa- limited to late Nukumaruan rocks of nian and Nukumaruan rocks, and the Hawke’s Bay, A. cottoni (King, 1933) is earliest definite record of the larger species limited to Nukumaruan rocks in SWair- M. octogonus (Quoy & Gaimard, 1833) arapa, and A. clavicula (King, 1933), an (Beu & Maxwell 1990:359, pl. 48 l) is from unusual species with a narrowly rounded Castlecliff (Kaikokopu Shellbed, OIS 19; rather than angulate margin to the sutural Fleming 1953:199), but Marshall & Burch ramp, is limited to early Nukumaruan (2000) recorded specimens similar to both bathyal rocks at Palliser Bay. M. octogonus and M. espinosus, along with 14. dubius (Hutton, 1878), now limited a possible third species, from Ashcott to the NE North Island warm province; Road, W of Waipukurau, Hawke’s Bay Fleming (1953:188, 199) recorded ‘Taron n. (GS10858, U22/f 9588, late Nukumaruan). sp.’ from Kaimatira Pumice Sand (OIS 25; Late Neogene species require further in GS4163, R22/f6459) and T. dubius from study. Kaikokopu Shellbed (OIS 19; in GS4156, 12. Cominella (Eucominia?) hamiltoni (Hut- R22/f7417). Taron possibly represents an- ton, 1885) (Beu & Maxwell 1990:324, pl. other rare warm-water migrant from the N 42g), a distinctive small species limited to North Island in Wanganui Basin, but T. Nukumaruan rocks. dubius has direct development, so any 13. Aethocola species: many species of this transport presumably would have been by genus have highly useful, restricted ranges, rafting.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 most notably the Kapitean index species A. 15. planata (Hutton, 1885), coerulescens (Finlay, 1930) (Beu & Max- apparently limited to Mangapanian well 1990:266, pl. 30e). Several Pliocene Castlecliffian rocks. However, the Recent species have rather short ranges also: A. species A. lawsi Finlay, 1930 seems to be pagoda Finlay, 1924 (Beu & Maxwell conspecific (Oaro, Marlborough, Nuku- 1990:298, pl. 37g) and A. (Zelandiella) maruan [Beu 1979]; Waipuru Shellbed to pliocenica (Powell, 1931) (Beu & Maxwell Upper Castlecliff Shellbed [OIS 11], as 1990:299, pl. 37d), both limited to Opoitian Proximitra n. sp., etc., of Fleming’s [1953] early Mangapanian rocks; A. (Zelandiella) lists). Taxonomic revision required; again, propenodosa (Bartrum, 1919), recorded many unnamed species are recorded from only from Opoitian rocks at Kaawa Creek, the NZEEZ by Spencer et al. (2009:209) and SW Auckland; and A. (Zelandiella) allani probably include A. planata. (King, 1934) ( conoidea Zittel, 1864), 16. species: the restricted ranges re- limited to OpoitianWaipipian Starbor- corded by Beu & Maxwell (1990) and 10 AG Beu

Maxwell (2009) for some late Neogene Nukumaruan; last recorded uncommonly Alcithoe species suggest they are useful in late Nukumaruan Petane Group in for biostratigraphy, but in practice they central Hawke’s Bay. are uncommon and variable, and difficult 21. Tomopleura subalbula (Murdoch, 1900), to use: A. arabica (Gmelin, 1791) (Beu & NukumaruanHaweran; recorded from Maxwell 1990:366, pl. 49d), first record Waipuru Shellbed (late Nukumaruan) to Nukumauan; A. exigua Marwick, 1926 and Landguard Sand (OIS 9) and at Te Piki, A. nukumaruesis (Marshall & Murdoch, near East Cape (OIS7); apparently 1920) (an ecophenotype of A. arabica?), ancestral to the Recent species T. albula both limited to Nukumaruan; A. (Hutton, 1873) but the distinction needs to (Quoy & Gaimard, 1833) (Beu & Maxwell be verified from more material. 1990:367, pl. 49i), CastlecliffianRecent. 22. *Zenepos lacunosa (Hutton, 1885); Powell 17. Three large, long, narrow, short-spired, (1942) recorded this from Castlecliff (ho- distinctive species of seem lotype) only; I am not aware of any other to be limited to Nukumaruan rocks of records. Hawke’s Bay: hectori (Kirk, 1882), 23. *Ringicula uniplicata Hutton, 1885, re- S. kirki (Marwick, 1924), and S. marwicki corded only from the late Nukuma- (Finlay, 1927) (Beu & Maxwell 1990:326, ruan Petane Group of Hawke’s Bay, but pl. 42j; Maxwell 2009:245), but these again apparently not collected for more require revision and comparison with the than 100 years, and the type material is Recent species renamed by Marshall lost. (2004). 18. Zeadmete kumeroa Fleming, 1943, appar- ently limited to Nukumaruan rocks of S Hawke’s Bay, and Z. pliocenica (Finlay, 24. *Cosa wanganuica Finlay, 1930, apparently 1930), apparently limited to Castlecliffian limited to Castlecliffian rocks of Wanganui rocks at Wanganui (recorded by Fleming (Kaimatira Pumice Sand, OIS 25, to Tai- (1953:226, 230) only from Pinnacle Sand nui Shellbed, OIS 13; Fleming 1953). (OIS14; GS4098,R22/f6425) and Tainui 25. *Hamacuna nukumaruensis (Laws, 1940), Shellbed (OIS 13; GS4013, R22/f6353, & limited to MangapanianNukumaruan GS4097, R22/f6516) at Wanganui, but rocks (Wilkies Shellbed at Wilkies Bluff

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 recently collected from Upper Castlecliff and Nukumaru Brown Sand at Nukumaru Shellbed (OIS 11; GS13590, R22/f0100). Beach; Laws 1940). NeogeneRecent species require revision. 26. Talabrica bellula (A. Adams, 1854) and 19. Bonellitia lacunosa (Hutton, 1885) (Beu & related species all require investigation. Maxwell 1990:326, pl. 42 l) is another The earliest New Zealand record of the species apparently limited to late Nuku- genus seems to be Mangapanian. Fleming maruan rocks of Hawke’s Bay; collected (1953:178) recorded T. bellula only from recently at localities in Petane Group near Lower Okehu Siltstone (Castlecliffian, OIS Napier and at Maharakeke Road SW of 29; Mowhanau Formation of Abbott & Waipukurau. B. lacunosa seems to reach a Carter 1999). T. nummaria Powell, 1931 is larger size and have coarser than recorded only from Nukumaruan and, the Recent species B. superstes Finlay, possibly, Castlecliffian rocks (recorded by 1930. Fleming (1953:180) from Okehu Shell Grit 20. Comitas allani Powell, 1942 (Beu & (OIS27), but in brackets, indicating Maxwell 1990:327, pl. 42p), Opoitian to thought reworked from older rocks). T. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 11

inornata Wild & King, 1932 apparently is a 30. Warm-water taxa recorded by Fleming rare species recorded only from Nukumar- (1953:272274) from Oturi shellbed, Wai- uan rocks. T. senecta Powell, 1931 (Beu & pipi (OIS5a): a final subject that deserves Maxwell 1990:310, pl. 40b, f) apparently is fuller treatment than the cursory glance limited to Nukumaruan rocks at Castle- given it here is the taxa recorded by Fleming point (although both T. nummaria and from the basal Oturi shellbed of the Hauriri T. inornata possibly are synonyms of Terrace cover beds (OIS5a, ca 80 ka) at T. senecta). An unnamed species occurs Waipipi, mouth of Wairoa Stream, W end in Castlecliffian (OIS15) rocks at Matata, of Waverley beach, W of Wanganui. Re- collections have shown that this is a ‘fossil’ Bay of Plenty (NMNZ M.137685), and boulder beach, with Barbatia novaezelan- another is present in the Recent fauna of diae (Smith, 1915) abundant among the in the Three Kings Islands (NMNZ situ boulders, infaunal boring bivalves such M.137651; BA Marshall NMNZ pers. as Pholadidea present in their bore-holes in comm. October 2009; Spencer et al. the boulders, and a few specimens of 2009:198). Lamprell (2003) treated Talab- Cleidothaerus albidus (Lamarck, 1819) at- rica Iredale, 1924 as a of Crassa- tached to the boulders. The small amount tina Kobelt, 1881 (West Africa), but judged of shelly sand surrounding the boulders from illustrations of African species contains the following warm-water species, (e.g. Ardovini & Cossignani 2004, according to Fleming’s (1953) list: Mono- figs. pp. 277278) this synonymy is most dilepas monilifera (Hutton, 1873), Maori- unlikely to be correct. The shape crypta costata (G. B. Sowerby I, 1824), M. and sculpture of Crassatina species are cf. youngi Powell, 1940, Murexsul mariae both strongly distinct from those of Talab- Finlay, 1930, and cf. mustelina rica species. Angas, 1871 (that is, Serrata fasciata (G. B. 27. Marama murdochi Marwick, 1927 (Beu & Sowerby II, 1846); Marshall [2004:8]). Maxwell 1990:316, pl. 41c, d), species and These almost all have taxonomic problems genus last recorded in late Nukumaruan or could refer to other, similar species, e.g. rocks; a distinctive, inflated relative of Fleming’s material identified as Murexsul Dosina with a large, pouting lunule mariae appears to me to be M. octogonus and fine commarginal sculpture, limited (Quoy & Gaimard, 1833), which still lives in

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 to offshore, shelf, mudstone facies. W Cook Strait; Monodilepas contains a 28. ‘Parilimya’ neozelanica (Suter, 1914) (Bore- number of similar species with distinct ham 1965:73, pl. 20, figs. 2, 3, as ‘Thracia’ ranges around New Zealand at present; neozelanica; Beu & Maxwell 1990:346, pl. and Maoricrypta youngi still lives in W 46g, i); recorded rarely from Wanganui Cook Strait (see below). The only certain Castlecliffian rocks and the Recent fauna; warm-water, NE North Island species pre- collected recently from Upper Castlecliff sent is Maoricrypta costata, although if Shellbed (OIS 11). Fleming’s identification is correct, Serrata 29. Thracia magna Marshall & Murdoch, 1921 fasciata is possibly another, as it is not (Beu & Maxwell 1990:288, pl. 34h), limited recorded south of Mahia Peninsula (Mar- to WaipipianNukumaruan rocks. This shall 2004:9). This fauna deserves further distinctive, large species is last recorded attention, although recent re-collections from late Nukumaruan rocks (Nukumaru have not brought any apparently significant Brown Sand) at Wanganui. species to light. 12 AG Beu

Taxonomy (Cossmann 1912, pl. 4, figs. 33, 34) (new Phylum Mollusca synonym). Class Gastropoda Informal group Ptenoglossa Tumidiacirsa de Boury 1911:221. Replacement name for Pseudoacirsa de Boury, 1909, suppo- Family Epitoniidae sedly a junior homonym of Pseudacirsa Kobelt, 1903 (although under the current ICZN Code Genus Acirsa Mo¨ rch, 1857 Pseudoacirsa is not a homonym of Pseudacirsa, their synonymy with Acirsa was implied by Acirsa Mo¨ rch 1857:77. Type species (by sub- Bouchet & Ware´ n (1986:526) so the question is sequent designation, Bouchet & Ware´ n irrelevant) (new synonym). 1986:526): Scalaria eschrichti Holbo¨ ll in Mo¨ l- ler, 1842, Recent, Arctic & W N Atlantic Notacirsa Finlay 1926a: 231. Type species (by (N Canada & W Greenland to Massachusetts, original designation): Turbonilla (Pyrgiscus) USA; Bouchet & Ware´ n 1986:528, fig. 1225). oamarutica Suter, 1917, early , New Zealand (new synonym). Acirsella de Boury 1886:xxi. Type species (by original designation): Scalaria inermis Plastiscala Iredale 1936:302. Type species (by Deshayes, 1861, Eocene (Lutetian), Paris Basin original designation, Iredale 1936:336): Scala (Cossmann 1912, pl. 4, figs. 29, 30; synonymy morchi Angas, 1871, Recent, SE Australia (new implied by Bouchet & Ware´ n 1986:526). synonym).

Hemiacirsa de Boury 1890:268. Type species Remarks. Bouchet & Ware´ n (1986:526) re- (by original designation): Turbo lanceolatus garded all the subgenera of Acirsa recognized Brocchi, 1814, Pliocene, Mediterranean by Cossmann (1912) based on Recent type (Cossmann 1912, pl. 4, figs. 23, 24; synonym species as synonyms of Acirsa, and commented according to Bouchet & Ware´ n 1986:526). that ‘Other genera based on fossil type species may be synonyms’, implying that they thought Pseudacirsa Kobelt 1903:26. Type species (by the other very similar subgenera listed in the subsequent designation, Nordsieck 1968:75): synonymy above are also synonyms. Another Scalaria coarctata Jeffreys, 1884, Recent, N synonym may well be Proacirsa Cossmann

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Norway (Bouchet & Ware´ n 1986:526, fig. 1224; (1912:96; type species inornata synonym according to Bouchet & Ware´ n Terquiem & Jourdy, 1869, Bathonian, France) 1986:526). but as this almost identical shell is in age, whereas all other members of the genus are Plesioacirsa de Boury 1909:256. Type species Cenozoic to Recent, the synonymy deserves (by original designation): Scalaria subdecussata more careful evaluation than is possible here. It Cantraine, 1835, Recent, SEurope (Bay of was discussed in some detail by de Boury Biscay to Madiera and Canaries; W Mediterra- (1917:58), who thought the age difference nean) (Bouchet & Ware´ n 1986:528, fig. 1226; unimportant. In his critique of Cossmann’s synonym according to Bouchet & Ware´ n (1912) classification, de Boury (1917:57) ob- 1986:526). jected to Cossmann’s (1912:96) comments on the similarity of these subgenera, pointing out Pseudoacirsa de Boury 1909:256. Type species that the of some of these groups (by original designation): Acirsa bezanconi de (particularly Acirsa sensu stricto) is obtuse and Boury, 1909, Eocene (Lutetian), Paris Basin submammillate, whereas that of others (notably Marine Mollusca of isotope stages of the last 2 million years in New Zealand 13

Plesioacirsa) is pointed and elongate. However, Tate, 1888. However, Finlay obviously inten- this distinction between tall, narrow, plank- ded his ‘I provide this for’ statement to be a totrophic protoconchs and shorter, wider, type species designation, and this is accepted lecithotrophic ones is no longer regarded as a here. generic character. De Boury (1917:59) also admitted that distinguishing Acirsella from Acirsa cookiana (Dell, 1956) (Fig. 3F,G) Plesioacirsa was extremely difficult. Bouchet & Ware´ n’s (1986) synonymy is well justified, Mathildona cookiana Dell 1956b:39, fig. 30; particularly as the three similar shells they Maxwell 1966:447 (position queried); Bieler illustrated (Bouchet & Ware´ n 1986, figs. 1995:632, fig. 92 (holotype; removed from 12241226) are the type species of the subge- Mathildidae). nera Acirsa (fig. 1225), Pseudacirsa (fig. 1224) Mathildona? cookiana. Powell 1979:250. and Plesioacirsa (fig. 1226). In view of the range Acirsa cookiana. Weil et al. 1999:128, fig. 395; of taxa included by Bouchet & Ware´ n (1986) it Spencer et al. 2009:205. is accepted here that the New Zealand fossil species assigned to Notacirsa by Finlay (1926a) Type material. Mathildona cookiana, holotype belong in Acirsa. One of the few apparently (Bieler 1995, fig. 92; Weil et al. 1999, fig. 395) distinctive characters of Notacirsa, compared NMNZ M.8842 (Marshall 1996:23), Recent, with typical N Atlantic species of Acirsa, is the Cook Strait, in 137 m, 40852.6?S, 174849.5?E; relatively prominent, wide, widely spaced axial two lots of paratypes in NMNZ, also from ridges (e.g. in N. oamarutica (Suter, 1917); Cook Strait, in 113 and 148 m (Dell 1956b:39). Finlay 1926a, pl. 56, fig. 16; N. vetusta Maxwell 1992, pl. 14 l), but some other species illustrated Other material examined. Opoitian: A specimen by Cossmann have much coarser ridges (e.g. A. in GS1567, W19/f7516 (Waiau River at junc- drevermanni de Boury in Cossmann 1912:196, tion with Pakihiwi Stream, inland from Wair- pl. 6, figs. 46; A. duvergieri de Boury in oa, N Hawke’s Bay) is a small spire from a shell Cossmann 1912:197, pl. 6, fig. 21). Other very similar to TM8602 (see below). This genera proposed for Australian Recent taxa, specimen was identified as ‘ cf. planostoma in particular Plastiscala Iredale (1936:302) (Hutton, 1885)’ by J Marwick in the GNS (type species Scala morchi Angas, 1871, Recent, collection, but is considerably wider and more SAustralia; Weil et al. 1999:140, fig. 433, strongly sculptured than other specimens re-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 holotype illustrated, BMNH 1871.7.5.10) seem ferred to A. planostoma, and appears to be an highly likely to be further synonyms of this incomplete spire from a much larger shell almost cosmopolitan, speciose genus. The ho- similar to Acirsa cookiana. Castlecliffian: lotype of Scala morchi (examined, AGB) is very Omapu Shellbed (OIS 23), Castlecliff coastal similar to Acirsa cookiana (Dell, 1956) in most section E of Mowhanau Beach, coll. C. A. characters other than its smaller size, its Fleming (TM8601, GS4060, R22/f6391, 1, Fig. markedly weaker spiral sculpture and its few 3F, G); Pinnacle Sand (OIS 15/14) at ‘the low varices at irregular, sparse intervals, and pinnacles’ gully, Castlecliff coastal section generic separation does not seem warranted. (one, TM8602), collected and presented by the Finlay (1926a:231) gave a very vague type late Mrs Jean Boswell, a keen collector of species designation for Notacirsa, stating only fossils from the Castlecliff section, near her ‘Notacirsa n. gen./I provide this for Turbonilla home in Wanganui. Jean donated several valu- oamarutica Suter [reference]’. He also included able specimens to GNS’s collection over many two other named New Zealand species and an years. Another specimen of A. cookiana from Australian fossil, Scalaria (Hemiacirsa) lampra the same locality was present in her collection, 14 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 3 (AE,H) Opalia (Pliciscala) flemingi n. sp.; A, B, holotype, TM8590, GS4107, R22/f6430, Ototoka Siltstone (Castlecliffian, OIS 31), E of Ototoka Beach, Wanganui; height 36.2 mm; C, paratype, TM8593, locality in doubt (stratigraphically higher than Ototoka Siltstone?), illustrated by Beu & Maxwell (1990, pl. 47k); height 31.3 mm; D,E,H, paratype, TM 8591, all data as holotype; D, sculpture of last , SEM; E, whole shell, height 32.3 mm; H, SEM, punctae enlarged. (F,G) Acirsa cookiana (Dell), TM8601, GS4060, R22/f6391, Omapu Shellbed (Castlecliffian, OIS 23), E of Mowhanau Beach, Wanganui, SEM; F, protoconch; G, whole specimen, height 14.2 mm.

but I do not know its present location. Recent: likely to be conspecific, and is quite similar to A specimen in Suter’s collection from the the Omapu Shellbed specimen. It was not famous ‘110 fathoms [200 m] off Great Barrier mentioned by Murdoch & Suter (1906), but a Island’ sample (Murdoch & Suter 1906) seems number of other unrecorded and unidentified Marine Mollusca of isotope stages of the last 2 million years in New Zealand 15

taxa from this station are present in Suter’s prominent axial costae of planktotrophic pro- collection. Others: NMNZ M.8839, Cook toconchs of Epitoniidae Nystiellinae (Bouchet Strait, 148 m, 40849.6?S, 174836.8?E (five, & Ware´ n 1986, figs. 11311138) (; paratypes of M. cookiana); NMNZ M.287949, Nu¨ tzel 1998). Okutani’s (2000, pl. 160, fig. 16) SE slope of Ranfurly Bank, 153143 m, illustration under the name Plastiscala morchi 37838.5?S, 178856.4?E (two); NMNZ M.287 (Angas, 1871) shows a specimen closely resem- 948, N of Alderman Islands, 108113 m, bling Acirsa cookiana, and much larger than P. 36847.5?S, 176800.0?E (one); NMNZ morchi. However, it has still more prominent, M.287950, NZOI Stn P515, 37 km NE of Great strongly convex, polished spiral cords and Island, Three Kings Islands, 550 m, 33858.0?S, weaker axial ridges than New Zealand speci- 172830.6?E (one). mens. Hasegawa & Nakayama (2009) recently confirmed this by recording what is apparently Distribution. Acirsa cookiana is uncommon the same species under the name Acirsa morsei around northern and central New Zealand at (Yokoyama, 1926) from off Noto Peninsula, present and a few fossil specimens have now Sea of Japan. Iredale’s (1936, pl. 22, figs. been recognized at Castlecliff, Wanganui. Pos- 2123) illustrations of Plastiscala morchi show sibly its uncommonness at Castlecliff results a shell with aligned ventral varices that are from the deposition of most Nukumar- absent from New Zealand specimens, and with uanCastlecliffian formations in Wanganui much weaker spiral cords (Scala morchi Angas Basin in no more than about 5070 m of water, 1871:15, pl. 1, fig. 7; May 1923:63, pl. 29, fig. that is, significantly shallower than Recent 13; Cotton 1956:4, fig. 13; Macpherson records of A. cookiana, which are from ca 1958:33, pl. 29, fig. 13; Weil et al. 1999:140, 100550 m. The very similar Opoitian specimen fig. 433). Australian specimens are also much from N Hawke’s Bay suggests that this is a smaller (H 78 mm for the apparent synonym widespread, if uncommon, species group in Plastiscala verconis Cotton [1939:172]; H 9.7, D New Zealand late Neogene rocks, but its range 2.8 mm for the holotype of P. morchi, BMNH is poorly known at present. 1871.7.5.10). New Zealand specimens are also significantly larger and have more numerous, Dimensions. Holotype: H 9.8, D 3.7 mm (Dell narrower, more closely spaced spiral cords than 1956b:39); GS4060, TM8601, Omapu Shellbed: Australian ones: 12 very closely spaced ones on H 14.2, D 4.8 mm; TM8602, ‘the pinnacles’: H spire whorls and about 1516 on the last whorl

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 16.1 (slightly incomplete), D 5.4 mm. on TM8601, nine more widely spaced ones on spire whorls and about 1214 on the last whorl Remarks. Acirsa cookiana has more prominent on TM8602 (low, narrow, closely spaced cords spiral cords than most species assigned to on the ramp and base are difficult to count). Acirsa previously, but relatively weak, uniform However, it appears that the relative promi- spiral sculpture is present over the entire nence of secondary cords (all prominent on exterior on most Acirsa species (Bouchet & TM8601, some suppressed over much of the Ware´ n 1986, figs. 12241226), and some species height but strengthening on the last whorl on previously assigned to the synonym Acirsella de TM8602) affects the overall shell appearance Boury, 1886 have prominent spiral cords (e.g. greatly. New Zealand specimens also lack the Cossmann 1912, pl. 4, figs. 31, 32). The true, raised, strongly aligned varices that are protoconch is present on one of the Castlecliff present on Australian material of P. morchi, specimens (Fig. 23I) and is a normal epitoniine although low ridges marking growth pauses are lecithotrophic protoconch, closely similar to present at irregular intervals on both Castlecliff those illustrated by SEM by Bouchet & Ware´ n specimens. Despite the deservedly critical re- (1986, figs. 11651181, 12271235), lacking the views of Weil et al. (1999) by Beechey (2009) 16 AG Beu

and Petit (2009a), I agree with Weil et al. Tewkesbury Formation, Nukumaru Beach, W (1999:128) that A. cookiana is a prominently of Ototoka Stream mouth (one incomplete);?O- sculptured species of Acirsa. totoka Siltstone, poorly localized (TM8593, Fig. 23D, specimen illustrated by Beu & Genus Opalia H. Adams & A. Adams, 1853 Maxwell (1990:348, pl. 47k); collected by OJ Marston from a locality well to the W of Kai- Opalia H. Adams & A. Adams 1853:222. Type Iwi). The last paratype was incorrectly stated species (by subsequent designation, de Boury by Beu & Maxwell (1990:348, pl. 47k) to be 1886:xxvi): Scalaria australis Lamarck, 1822, from ‘the pinnacles’ gully, but actually came Recent, southeastern Australia. from well to the west. The collector is unfami- liar with Wanganui stratigraphy, but the speci- Psychrosoma Tapparone-Canefri 1876:154. Un- men is clearly from much lower in the section necessary new name for Opalia Carpenter, than Pinnacle Sand. However, its preservation 1865, supposedly not of H Adams & A Adams, is better than that of the rest of the type 1853. material, it bears more prominent axial ridges than the other paratypes, and it is also Subgenus Pliciscala de Boury, 1887 possible that it is from significantly higher in the section than Ototoka Siltstone. The full Pliciscala de Boury 1887:19. Type species (by stratigraphic range of O. flemingi n. sp. is, original designation): Scalaria gouldi Deshayes, therefore, unclear at present. 1861, Eocene, Paris Basin. Distribution. NukumaruCastlecliff coastal sec- Remarks. Kilburn (1985:271) tentatively re- tion, only, in Wanganui Basin; early Castleclif- ferred the only other species I am aware of fian (OIS31; possibly later) and latest that resembles Opalia flemingi n. sp., O. aglaia Nukumaruan; only seven specimens seen. The (Bartsch, 1915), to Opalia (Pliciscala), and is record of ‘Acrillospira’ (sic,Acrilloscala) from followed here, again tentatively. Tewkesbury Formation at Nukumaru Beach (Abbott et al. 2005, appendix table, p. 149) was Opalia(Pliciscala)flemingin.sp.(Fig.3AE,H). based on Fleming’s (1953:158) record of this species (RM Carter, James Cook University Acrilloscala n. sp. Fleming 1953:158,174. pers. comm. 6 October 2005), which in turn was Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Opalia n. sp. Beu & Maxwell 1990:348, pl. 47k. based on Marwick’s collections from Nuku- maru Beach (Fleming’s [1953:158] list for Type material. Holotype (Fig. 3A, B) TM8590, GS1164-1202, R22/f6348, R22/f6309). The GS4107, R22/f6430, Ototoka Siltstone (OIS only other record by Fleming (1953:174) is 31), coast E of mouth of Ototoka Stream, from GS4107, R22/f6430, ‘Ototoka Siltstone W of Wanganui, collected by CA Fleming; with Tongue’ of Fleming, in the coastal section E of two paratypes, TM85912. Ototoka Stream mouth (Ototoka Siltstone; Abbott & Carter 1999:96). Unfortunately, this Other material examined, all paratypes. GS4107 part of the Wanganui coastal section is heavily re-collection, R22/f6430 (one incomplete); overgrown, deeply weathered, and inaccessible GS1198, R22/f6305, ‘sandstone immediately at present. above coal [that is, a lignite bed in Maxwell Formation], coast between Ototoka Stream Dimensions. Holotype: H 36.2, D 11.1 mm; and Okehu Stream’ (upper Tewkesbury For- paratype TM8591: H (incomplete) 39.9, D 11.2 mation; one incomplete); GS1164, R22/f6348, mm; paratype TM8592: H 32.3, D 9.9 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 17

Description. Shell tall and narrow, large for interspaces on spire whorls, but numerous, genus (height 3240 mm), weakly sculptured closely spaced and covering entire surface with narrow, closely spaced axial and spiral including basal disc (but not apertural lips) on ridges; rather thin and fragile; apparently last few whorls of well-preserved specimens. mostly composed of aragonite (white in most elliptical, with weak posterior sinus, specimens), but with a thin, pinkish to pale weakly angled at anterior end; lips smooth, brown (presumably calcite) outer layer that thickened, forming continuous peristome and a detaches rather easily (an intritacalx?). Proto- terminal ; small, narrow, sharp-edged, not seen. Teleoconch evenly tapered, of weakly ridged fasciole bordering lower half of 1213 lightly convex whorls with moderately inner lip on holotype, weakly developed on impressed , spire angle 17188. Peribasal other specimens. Low, indistinct varices present cord prominent, smooth on most specimens, at irregular intervals of about 2/3 to one whorl faintly nodulose on a few, narrowly rounded, on lowest 57 whorls. issuing from suture on last whorl, continuous with suprasutural cord when that is present, Remarks. The taxonomic position of this spe- delineating basal disc; termination faintly an- cies is a little uncertain. The regular pitting of gling lower right profile of outer lip. Axial the whorl surface (Fig. 3D, H) was interpreted sculpture of low, narrow, rather closely spaced, by Beu & Maxwell (1990:348) as evidence for a slightly irregular ridges with indistinctly defined position in Opalia, following Kilburn edges, weakly curved, inclined slightly forwards (1985:241,266). The presence of only low, towards upper suture; prominent on early spire irregular varices also supports a position in whorls, weakening down teleoconch; narrower Opalia. I have searched long and hard for a and more closely spaced over upper part of Recent analogue of this species, with little whorl (a weakly defined sutural ramp) than success; nothing similar was described by Weil lower down; each axial interspace slightly wider et al. (1999), Nakayama (2003) or Garcia (2003, than one ridge; 32 on last whorl and 33 on 2004). It is most nearly similar to the South penultimate whorl of holotype, 26 on last whorl African Recent species O. (Pliciscala) aglaia and 28 on penultimate whorl of paratype (Bartsch, 1915) (Kilburn 1985:271, figs. 51, 52) TM8593. Spiral sculpture of one low suprasu- but the few known specimens of O. aglaia are tural and one low subsutural fold (subsutural much smaller (reaching 206.4 mm, but most fold formed by closely spaced, weakly nodulose specimens are half this size) and a little more

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 tops of axial ridges) slightly more prominent finely sculptured than O. flemingi, with nar- than remainder of spiral sculpture, rest of rower, more numerous (3040 per whorl), more whorl (between suture and peribasal cord on closely spaced, and slightly more strongly last whorl) entirely covered with 1218 low, curved axial ridges. Kilburn (1985:272) de- weak, closely spaced cords per whorl, crossing scribed O. aglaia as having ‘micropunctate axial ridges without forming nodules; relatively interstices’. This is the only really similar few, prominent, uniform and wide on some species I am aware of, and a close phylogenetic specimens (e.g. paratype TM8593) but lower, relationship with O. aglaia appears feasible, more closely spaced, more varied and more implying that this is a further example of numerous on others, including holotype; with transport as planktotrophic larvae in the Ant- numerous very fine interstitial spiral threads on arctic Circumpolar Current. some specimens, including holotype. Basal disc Two Recent lots in NMNZ (M.171153, off sculptured much as on whorl sides, but all Lord Howe Island, 565960 m, 31845.73?S, sculpture much more subdued. Exterior surface 159820.93?E, RV ‘Tangaroa’, one large; bearing microscopic rectangular pits in regular M.172283, W Norfolk Ridge, W of Cape spiral rows; pit rows mostly few and in spiral Reinga, 785800 m, 34817.09?S, 168825.82?E, 18 AG Beu

RV ‘Tangaroa’, one large and seven small) are twentieth century. Fleming’s seminal work on superficially similar to Opalia flemingi n. sp., the stratigraphy and paleontology of Wanganui but have more regular, even spiral cords and Basin underpins all later research on this axial ridges, lack obvious punctae, and belong magnificent succession. in the group of epitoniids usually assigned to Acrilla. The large specimen in M.171153 is Subgenus Nodiscala de Boury, 1890 particularly similar in size and shape, but has lightly angled early spire whorls not seen in O. Nodiscala de Boury 1890:12. Type species (by flemingi n. sp. A Grebneff (Dunedin pers. original designation): Scalaria bicarinata G. B. comm.) pointed out similarities to O. leeana Sowerby II, 1844, Recent, tropical Indo-West (Verrill, 1883) and O. () andrewsii Pacific (Hawaii and the Red Sea to South (Verrill, 1882) (Abbott 1974, figs. 1211, 1214) Africa; Weil et al. 1999:106, fig. 339, as Opalia from the NE United States. None of the Recent (Nodiscala) bicarinata). Australian species of Opalia is similar; all have much fewer, lower, more widely spaced, less Remarks. The taxonomy of this group of sharply defined axial costae, weaker spiral Epitoniidae is complex and undecided even at sculpture, and a more prominent peribasal the generic level, and I use Nodiscala here as a ridge. The regularly pitted surface of Opalia subgenus of Opalia (1) to underscore the need flemingi n. sp. cannot be a preservational for a reassessment of these epitoniid genera, (2) artefact, so the new species is not related to because the genera and subgenera in this group, such taxa as Acrilla acuminata (G. B. Sowerby discussed below, all have punctate teleoconchs II, 1844) (Wilson 1993, fig. p. 274; Indo-West resembling that of Opalia, and (3) because the Pacific to Keppell Bay, Queensland). Another Nodiscala species illustrated by Cossmann Australian Recent species that resembles O. (1912, pl. 4, figs. 5, 6; pl. 5, figs. 23, 2527; flemingi n. sp. quite closely is Propescala valida pl. 6, figs. 9, 15, 20, 24) most closely resemble (Verco, 1906) (Verco 1906a:147, pl. 4, fig. 7; the New Zealand species under consideration Cotton 1956:1, fig. 19), which, however, is here, O. (Nodiscala) nympha (Hutton, 1885), much smaller (H 7 mm). A final Australian(?) O. (Nodiscala) ahiparana (Powell, 1930), O. species resembling the new species is ‘Scalaria’ (Nodiscala) zelandica (Finlay, 1930) and distincta E. A. Smith (1891:441, pl. 35, fig. 15; O. (Nodiscala) maxwelli (Finlay, 1930). Iredale 1936:305, pl. 22, fig. 29), which Iredale These species (at least Opalia nympha and

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 included in the ‘not rediscovered’ list of possi- O. maxwelli, in most previous classifications) ble Atlantic species from the muddled HMS have been placed by most previous authors in ‘Challenger’ Station 164B, supposedly dredged Funiscala de Boury (1891:205; type species [as off Sydney, New South Wales, in 750 m. reidentified by de Boury 1911:219]: Scalaria ‘Scalaria’ distincta is particularly similar to O. speyeriana Sacco, 1891, Oligocene, Italy). The flemingi, but has regular varices in the centre of type species originally designated for Funiscala the venter of each whorl that are not present on by de Boury (1891:205) was Scalaria pusilla O. flemingi n. sp., and appears to be related to Philippi, 1844 (Oligocene or Miocene, N Ger- Acrilla acuminata. Whatever its relationships, many) but Cossmann (1912:86, footnote 2) Opalia flemingi n. sp. is a very distinctive stated that the type species was misidentified. addition to the New Zealand epitoniid fauna. Cossmann’s treatment followed that of de Boury (1911:219), who discussed the problem Etymology. It is a pleasure to name this finest of Funiscala in more detail. He concluded that of the new Wanganui PlioPleistocene molluscs in at least two earlier publications he had in honour of Charles Fleming KBE FRS, the misidentified the shell he now called Scalaria leading New Zealand paleontologist of the late speyeriana Sacco as S. pusilla. S. speyeriana Marine Mollusca of isotope stages of the last 2 million years in New Zealand 19

then, in his opinion, became the true type Not (Confusiscala) nympha. Suter species of Funiscala, whereas he now regarded 1917:85 (Miocene). S. pusilla as a species of Bifidoscala. The situation is that covered by ICZN Article Type material. Scalaria nympha, two syntypes 70.3. As de Boury’s (1911) identification of Canterbury Museum M-3098, M-3099 (Brad- the type species of Funiscala has been accepted shaw et al. 1992:33), from Petane, Hawke’s by subsequent authors, the type species is here Bay, Nukumaruan, that is, Tangoio Limestone fixed (under Article 70.3) as Scalaria speyeriana or Mairau Mudstone, high in Petane Group, Sacco, 1891, misidentified as Scalaria pusilla hillside N of Esk River and junction of High- Philippi, 1841 in the original designation by de ways 3 and 5, 30 km N of Napier. Boury (1891:205). De Boury (1917:50) made this case still more complex: ‘unfortunately, Other material examined. Waipipian: a record in Funiscala must be called Rugatiscala’. He FRED of ‘Epitonium aff. nympha’ from GS876, repeated the type-species problem and consid- Q21/f6493, Tangahoe Formation, mouth of ered that the type species of Rugatiscala is Scala Waihi Stream, Waihi Beach, Hawera, S Tar- levesquei de Boury, 1887 (Cossmann 1912, pl. 3, anaki (identification published by Suter fig. 43). He also stated (de Boury 1917:51) that 1921:25; as ‘Epitonium aff. (Confusiscala) the main distinguishing character of Bifidoscala nympha (Hutt)’) is not matched by a specimen is the presence of a single row of punctae in the in the GNScollection. Mangapanian: Manga- [spiral] furrows, whereas Nodiscala species illu- pani Shell Conglomerate, Mangapani Bluff, strated by Cossmann (1912, pl. 5, figs. 23, Mangapunipuni Stream, Waitotara valley, 2527; pl. 6, figs. 9, 15, 20, 24) are more Mangapanian (late Pliocene) (Laws 1940:53; prominently and coarsely punctuate, and spe- Fleming 1953:119; recorded from GS4227, cies assigned to Funiscala by Cossmann R21/f8478; but again specimen not found in (1912:5, figs. 8, 9, 24) lack punctae altogether. GNS). Nukumaruan: otherwise recorded only The presence or absence and degree of puncta- from Nukumaruan rocks, although not re- tion in various species of Opalia (sensu lato) corded from Wanganui Basin Nukumaruan varies greatly, and it is as difficult to use this rocks by Fleming (1953). The single specimen character as all others in Epitoniidae. At attributed to this species in the GNSreference present the name Funiscala (and possibly also collection is from GS6985 (T24/f6489, shelly Bifidoscala Cossmann, 1888, Rugatiscala de conglomerate at Ashhurst Domain, cliff 600 m

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Boury, 1913, and several others) is regarded W of railway bridge over Manawatu River, E as a probable synonym of Nodiscala, although of ; Fig. 4E, F). it is unclear whether Nodiscala really can be maintained as separate from Opalia and/or Distribution. (Waipipian?); Mangapanian Pliciscala in the long run. Nukumaruan, Wanganui Basin and Hawke’s Bay. Opalia (Nodiscala) nympha (Hutton, 1885) (Fig. 4A, B, DF) Dimensions. Syntypes, M-3098: H (incomplete) 6.1, D 2.6 mm; M-3099: H (incomplete) 6.2, D Scalaria nympha Hutton 1885a:321; Hutton 2.9 mm; GS6985: H 7.9, D 2.8 mm. 1893:67, pl. 8, fig. 71. Epitonium nympha. Suter 1915:13. Remarks. Opalia (Nodiscala) nympha is a small Funiscala nympha. Finlay 1930b:233; Laws species (H 68 mm) with a teleoconch of at 1940:53; Fleming 1966:49; Beu & Maxwell least seven whorls (the teleoconch spire 1990:411. and protoconch are not preserved on the Funiscala (?) nympha. Maxwell 2009:244. material examined; naturally decollate?) with 20 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 4 (A,B,DF) Opalia (Nodiscala) nympha (Hutton), Nukumaruan, SEM; A,B,D, syntype, CMC M3098, ‘‘Petane’’, Hawke’s Bay; A, incomplete specimen, height 5.8 mm; B, punctae on base of A; D, surface of penultimate whorl; E,F, GS6895, T24/f6489, Ashurst Domain, E of Palmerston North; E, whole specimen, height 7.9 mm; F, surface of penultimate whorl. (C) Opalia (Nodiscala) maxwelli (Finlay), RM4020, Recent, Portobello sta. Mu6845, c. 130200 m, near head of Papanui Canyon, off Otago Peninsula, RV ‘‘Munida’’; SEM, height 5.0 mm. 1415 wide, prominent, essentially smooth imperceptibly into the intercostal intervals, axial costae per whorl. The costae have narrow which are each about twice the width of one crests, concave sides and wide edges, merging costa. The intercostal intervals appear under Marine Mollusca of isotope stages of the last 2 million years in New Zealand 21

the light microscope to be crossed by many illustrated here with the aim of allowing wider even, fine, low, closely spaced spiral lirae, but recognition of the species, and so assessment of under SEM examination these are seen to be its possible utility as a Mangapanian vague, partially obliterated, irregular ridges, Nukumaruan index species. As noted above, possibly resulting from partial abrasion of a its generic position also is unclear. O. nympha former intriticalx. A prominent peribasal spiral resembles the specimen illustrated as Funiscala rib marks off the apparently smooth, flat basal levesquei (de Boury, 1887) (middle Eocene disc, and high, weakly spirally lirate varices are [Cuisian], Paris Basin) by Cossmann (1912, present at roughly one whorl intervals down the pl. 3, fig. 43). The two species have similarly entire spire. The apertural lips are wide, double prominent axial sculpture, interrupted at irre- (that is, grooved), smooth on the inner edge, gular intervals by more prominent varices, a but raised into a prominent, spirally lirate varix correspondingly thickened, varicate outer lip, a on the exterior, and the inner lip is margined by similar rather restricted basal disc, and similar a narrow fasciole. SEM examination showed weak spiral sculpture. If F. levesquei were a that the surface between the axial ridges bears typical Funiscala species, the generic position of irregular, small, subcircular pits, and the basal O. nympha would be clear. However, in their surface inside the peribasal cord is coarsely and catalogue of Paris Basin Paleogene molluscs, Le regularly pitted. Renard & Pacaud (1995:93) referred F. leves- The most nearly similar of Recent species is quei to Opalia (Rugatiscala) de Boury, 1913, Opalia maxwelli (Powell 1979:252, fig. 58.3), proposed after Cossmann (1912) was pub- which has very similar axial sculpture but has a lished. This suggests that Rugatiscala de Boury more obviously concave base and less promi- (1913:176177; type species Scalaria levesquei) nent varices than O. nympha (Fig. 4C). Its might be a more suitable position. However, surface also bears pits in the base, inside the the better illustration of a complete specimen of peribasal ridge, as in O. nympha. SEM exam- R. levesquei by de Boury (1913, pl. 8, fig. 6) ination also showed that Recent specimens bear demonstrates that this is a more typically a finely reticulate intritacalx, and the unusual, Opalia-like shell than the incomplete, more irregularly ridged surface of both fossil speci- finely sculptured (evidently distinct) one illu- mens of O. nympha examined possibly results strated by Cossmann (1912, pl. 3, fig. 43), from the abrasion of an intritacalx. O. ahipar- having coarser axial and spiral sculpture than ana (Powell 1979:262, fig. 58.1) has less promi- in the earlier illustration and a series of regular

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 nent axial costae, two more prominent varices almost aligned down the ventral face of peribasal cords (one in the usual peripheral the teleoconch. The situation is further con- position, the other below it on the base) bearing fused by the characters of the Recent species large, rounded nodules that are not present on O. maxwelli, which lacks the finely reticulate any other New Zealand Epitoniidae, and lacks surface sculpture (an intritacalx?) of species obvious varices before the terminal one, which referred to Funiscala by Cossmann (1912), and is more prominent than in O. nympha. The final is much closer in appearance to species Coss- species needing comparison, O. zelandica (Po- mann referred to Nodiscala de Boury, 1890. well 1979:252, fig. 58.2) is more Epitonium-like Finlay (1930b:232, pl. 44, fig. 35) proposed than O. nympha, with lower, narrower and Nodiscala zelandica at the same time as Funis- more numerous axial costae, and no peribasal cala maxwelli, and evidently thought them not spiral cord; ‘the labial varix is heavy, and to be congeneric because of the more prominent previous varices are prominent on the spire axial ridges of O. maxwelli. As noted above, a 3 at ⁄4 whorl intervals’ (Powell 1979:252). position in O. (Nodiscala) is adopted for all the Opalia (Nodiscala) nympha has remained New Zealand species discussed here, but this little-known since it was described, and is group is urgently in need of major revision. The 22 AG Beu

major generic overhaul of all epitoniids sug- Remarks. The New Zealand PlioceneRecent gested by the excellent new research by Gitten- species Scalaria zelebori Dunker, 1866 has berger & Gittenberger (2005) and Gittenberger usually been assigned to Cirsotrema, but its et al. (2006) indicates that none of the species generic position has been debated recently. considered here is likely to be placed in its Weil et al. (1999:128, figs. 399, 400) placed C. phylogenetically correct genus at present, but a zelebori in Boreoscala Kobelt, 1902 (type spe- huge amount of further research is required cies: Scalaria greenlandica Perry, 1811, Recent, before the genera are resolved. North AtlanticNorth Pacific), and this has been followed by a few New Zealand authors Genus Cirsotrema Mo¨ rch, 1853 (Morley 2004:101). The New Zealand fossil and Recent species are definitely less elaborately Cirsotrema Mo¨ rch 1853:49. Type species (by sculptured than the more extreme Recent monotypy): Scalaria varicosa Lamarck, 1822, tropical species of Cirsotrema. However, the Recent, Indo-West Pacific (Wilson 1993:274, rather different nature of the wide, simple, pl. 44, fig. 1a,b). closely spaced spiral cords, lack of a basal disc, and wide, low, smooth, convex-crested Caloscala Tate 1885:3. Type species (by sub- axial ridges of Boreoscala greenlandica (see sequent designation, Tate 1890:230): Caloscala Bouchet & Ware´ n 1986:518, figs. 12101213), compared with the complex orders of spiral mariae Tate, 1885, late Eocene, Tortachilla cords and threads and complex, multilamellate Limestone, Aldinga, (synonym varices in New Zealand species assigned to of Cirsotrema according to de Boury [1887:40] Cirsotrema, makes a position in Boreoscala and Tate [1890:230]). unlikely to be correct. The original assignment of Australasian fossil species to Cirsotrema was Pseudostenorhytis Sacco 1891:72. Type species made by the well-known epitoniid specialist de (by monotypy): Cirsotrema (Pseudostenorhy- Boury (1887:40), who regarded Tate’s Adelaide tis) stenorhytoides Sacco, 1891, late Miocene, Eocene species Caloscala mariae Tate, 1885 and Italy (synonym according to Cossmann the other species included in Caloscala by Tate, 1912:51). Scalaria lyrata Zittel, 1864 (Oligocene Miocene, New Zealand) as species of Cirsotre- Elegantiscala de Boury 1911:216. Type species ma. As de Boury’s (1887) work is not widely Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (by original designation): Scalaria elegantissma available, his brief statement is quoted in full Deshayes, 1861, Eocene, Paris Basin (synonym here: ‘The subgenus Caloscala has been estab- according to Cossmann [1912:51], but not lished by M. Ralph Tate for C. lyrata (Zittel) according to de Boury [1917:39]). and C. mariae, Ralph Tate. The author has had the generosity to send us a very beautiful Cirsotremopsis Thiele 1928:92. Type species (by example of this latter species, which has many monotypy): Scalaria cochlea G. B. Sowerby II, similarities [‘‘beaucoup de rapports’’] with S. 1844, Recent, Mediterranean Sea to Angola acuta, Sow. We have demonstrated that this (Weil et al. 1999:32, fig. 790) (synonym accord- species cannot be separated from the true ing to Clench & Turner 1950:226). Cirsotrema’ (free translation from de Boury 1887:40). Later, de Boury (1911:215) restated Dannevigena Iredale 1936:303. Type species (by this case: ‘I persist in thinking that there is no original designation): Dannevigena martyr Ire- place to conserve this subgenus [Caloscala], dale, 1936, Recent, southeastern Australia (new which appears to include shells belonging synonym). to Cirsotrema Mo¨ rch. These forms cannot be, Marine Mollusca of isotope stages of the last 2 million years in New Zealand 23

as I had thought for a while, confounded with regarded as synonyms of Cirsotrema. If the very our subgenus Coroniscala, which differs com- similar New Zealand, Australian and South pletely from Cirsotrema by its ribs, which are American fossil species usually assigned to not at all crisped and frilled like those of the Cirsotrema should require a distinct genus or last subgenus’ [Cirsotrema]. The illustration of subgenus, Caloscala Tate, 1885 is available for C. mariae by Tate (1893, pl. 12, fig. 2) shows a them. Like Recent tropical Cirsotrema species, shell very similar to such large, prominently but few other epitoniids, the fossils have a sculptured New Zealand fossil species as C. calcitic teleoconch and preserve well in lime- lyratum (Zittel, 1864) and the Argentinean stone faunas from which the aragonitic mol- fossil species C. rugulosum (G. B. Sowerby I, luscs have been dissolved. Epitoniidae is 1846) (Griffin & Nielsen 2008:305, pl. 23, perhaps the most obvious group crying out figs. 8, 9). Cossmann (1912:52) and de Boury for resolution by molecular techniques, because (1913:170) also expressly included C. zelebori in of its poorly understood phylogeny, staggering Cirsotrema. The NukumaruanCastlecliffian diversity and few, intergrading taxonomic char- specimens reported below, with thicker varices acters, and a position in Cirsotrema should be than C. zelebori, are intermediate in many maintained for C. zelebori and similar species characters between C. zelebori and the large until molecular studies can be carried out to OligoceneMiocene species such as C. lyratum confirm or deny this position. and C. caelicola Finlay, 1926, helping confirm a Tate (1885:3) included two species in position in Cirsotrema for C. zelebori. On the Caloscala, but did not designate a type species. other hand, it should be noted that Cossmann, I am grateful to TA Darragh (Museum Victoria ‘avec la collaboration de M. de Boury’ (1912, pers. comm. September 2005) for pointing out pl. 3, figs. 5, 6), placed the large Patagonian that Tate (1890:230) subsequently designated Oligoceneearly Miocene species here identi- the type species himself, with the statement fied as Cirsotrema rugulosum (G. B. Sowerby I, (under the heading Scalaria (Cirsotrema) mar- 1846) in genus Boreoscala, significantly confus- iae Tate) that ‘This species is the type of my ing the generic position, as these South Amer- subgenus Caloscala...’. ican shells are exceedingly similar to the large New Zealand fossils of the same age. The large Cirsotrema zelebori (Dunker, 1866) (Fig. 5AE) Recent southern Australian species Dannevi- gena martyr Iredale (1936:303, pl. 22, fig. 25; Scalaria zelebori Dunker in Dunker & Zelebor

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Wilson 1993:275, pl. 44, fig. 4; named for the 1866:912; Hutton 1873a:21; Hutton 1880:69; drowned captain of FIS‘ Endeavour’, HC Tryon 1887:78, pl. 15, fig. 75; Hutton 1893:66; Dannevig; Verco 1935, pl. 16) is particularly Clessin 1897:50, pl. 13, fig. 5; Suter in Hutton closely similar to the large New Zealand and 1904:80; Suter 1905a:72; Moss 1908:30, pl. 7, South American OligoceneMiocene species fig. 17. assigned to Cirsotrema, although its spiral Scala (Opalia?) zelebori. Frauenfeld 1867:7, sculpture (‘faintly ...striate’; Wilson 1993:275) pl. 1, fig. 6. is weaker than on most fossil species, other Scalaria (Opalia) zelebori. Martens 1873:29. than C. gagei Maxwell (1978:36, fig. 44). There Scalaria intermedia Hutton 1873b:10. is little doubt that C. martyr is a Recent species Scalaria reevei Clessin 1897:63, pl. 15, fig. 9; of the same clade as the Australian, New Suter 1899a:54. Zealand and South American fossil species, Scalaria (Opalia) zelebori. Suter 1899a:54. that is, Dannevigena is a further synonym of Scala zelebori. Verco 1906a:145 (South Austra- Caloscala and Cirsotrema. It seems likely that lian record dismissed). more of the greatly over-split subgenera pro- Scala (Cirsotrema) huttoni de Boury in Coss- posed in Epitoniidae by de Boury should be mann 1912:52 (unnecessary new name for 24 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 5 Cirsotrema zelebori (Dunker), Wanganui fossils. (A) GS15338, S22/f0164, Maoricrypta-dominated shellbed, Turakina Valley Road, E of Wanganui; early Castlecliffian, c. OIS43 41; short, coarsely sculptured, abraded specimen, possibly an unnamed species; height 14.9 mm. (B,C,E) C. zelebori, GS4003, R22/f7394, Landguard Sand (Haweran, OIS 9), Landguard Bluff, Wanganui, SEM; B, surface of penultimate whorl, with multilamellate varices; C, centre right area of B enlarged; E, lower right area of C enlarged, showing micropunctae. (D) C. zelebori, holotype of Scalaria intermedia Hutton, TM8285, Tainui Shellbed (Castlecliffian, OIS13), ‘‘ShakespeareCliff’’, Wanganui; height 33.2 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 25

Scalaria intermedia Hutton, 1873, not a junior unimportant, as S. intermedia is clearly a homonym). synonym of C. zelebori. Scala (Cirsotrema) zelebori. de Boury 1913:170, pl. 8, fig. 2. Other material examined. Not listed; wide- Epitonium (Opalia) zelebori. Suter 1913:322, pl. spread and not uncommon. The earliest record 46, fig. 8 (emended to Epitonium (Cirsotrema) in of Cirsotrema zelebori seems to be from Kaawa corrigenda). Creek, SW Auckland, Opoitian (Laws Epitonium zelebori. Bucknill 1924:58, pl. 7, 1936:111). It occurs at most younger fossil fig. 12. localities throughout New Zealand, including Cirsotrema zelebori. Finlay 1928:246; Laws almost all formations in Wanganui Basin. 1936:111; Powell 1937:75, pl. 8, fig. 25; Powell I have seen the apparently distinct Turakina 1946a:77, pl. 8, fig. 25; Powell 1958a:98, pl. 8, fig. valley species (OIS43 41?; see below) only 25; Powell 1962:90, pl. 8, fig. 25; Fleming from the one early Castlecliffian shellbed. 1966:49; Powell 1976a:107, pl. 15, fig. 15; Powell Recent specimens of C. zelebori occur through- 1979:252, pl. 48, fig. 17; Beu & Maxwell out New Zealand, usually found washing 1990:411; Spencer & Willan 1996:24; Nu¨ tzel ashore on sandy beaches; particularly common 1998:93, pl. 13IK, OQ (showing punctate on Orewa Beach, Northland. Its host cnidar- sculpture, pl. 13Q). ian(s) have not been identified. Cirsotrema (Cirsotrema) zelebori. Spencer et al. 2009:205; Maxwell 2009:244. Distribution. Opoitian (early Pliocene) to Boreoscala zelebori. Weil et al. 1999:128, fig. Recent, throughout New Zealand. 399, 400; Morley 2004:101. Dimensions. ‘Type’ of Scalaria zelebori: H 24, D 8 mm (Suter 1913:322); Scalaria intermedia, Type material. The type material of Scalaria holotype: H 33.2 (incomplete, originally ca 36), zelebori was stated by Suter (1913:322) to be in D 12.4 mm; Recent, RM4759, Waihi Beach, NHMW. Anita Eschner (NHMW pers. comm. Bay of Plenty: H 27.2, D 9.6 mm; H 26.3, D 5 October 2007) reported that 11 syntypes of 9.6 mm; RM319, Mount Maunganui, Bay of Scalaria zelebori are present: NHMW 89781 Plenty: H (incomplete) 27.0, D 10.2 mm; H (one syntype, labelled as ‘Dunker’s original’), 26.2, D 9.5 mm; NMNZ M.18724, Waihi NHMW 89782 (nine syntypes), and NHMW Beach, Bay of Plenty: H 30.4, D 11.5 mm;

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 89783 (one syntype, from J Zelebor’s collec- NMNZ M.277711, beach, Maioro, SW of tion), all from ‘New Zealand’ (not seen). The Waiuku, SW Auckland: H 29.8, D 10.6 mm; type material of Scalaria reevei was stated by H 28.3, D 11.1 mm. Clessin (1897:63) to be in ‘Mus. Stuttgt.’ [Stuttgart], not seen. Scalaria intermedia, holo- Remarks. Suter (1899a:54) and de Boury type TM8285 (Fig. 5D), from Shakespeare (1913:170) attributed the name Scalaria zelebori Cliff, Wanganui; almost certainly from Tainui to Frauenfeld, as Frauenfeld (1867:7) seems to Shellbed (OIS 13). have considered that he described the species Cossmann (1912:52) listed the name first. However, the name was first published by ‘Scalaria huttoni de B. (Sc. intermedia Hut- Dunker (in Dunker & Zelebor 1866:912), as ton)’, so presumably Cossmann and de Boury Suter (1905a:72) later pointed out. thought that S. intermedia is a junior homo- Fleming (1953:144) listed ‘Cirsotrema n. sp. nym, but did not state the earlier usage, and I aff. zelebori (Dunk.)’ in the fauna from Nuku- have been unable to find one. De Boury (1913) maru Brown Sand in the Nukumaru Beach returned to the name Scalaria intermedia Hut- coastal section, from GS4117, R22/f6437, ton, so apparently it is not preoccupied. This is ‘unlocalised as to member’. Reconsideration 26 AG Beu

of this record, among the many other ‘n. sp.’ sculpture. The very small punctae occur in records of molluscs by Fleming (1953), raised spiral rows in the interspaces of the spiral cords the question of the status of Scalaria intermedia (Fig. 5C,E), although Nu¨ tzel (1998:93) noted Hutton, 1873, and more generally the taxon- that the punctae are not visible on all speci- omy of New Zealand late Neogene species of mens. SEM examination showed that the Cirsotrema. punctae are much smaller and sparser than The holotype of Scalaria intermedia has not those of Opalia species, seem to be at least been illustrated previously, and an illustration partially revealed by corrosion of the shell therefore is provided here (Fig. 5D) as surface, and vary in visibility over the surface an example of a typical, large specimen of of one specimen. This newly revealed character C. zelebori. The question of its status was requires comparison in a range of other New considered by de Boury (1913:170171), who Zealand material to determine whether it is examined a ‘magnifique’ specimen identified as present, but is likely to be present on all C. intermedia, 27 mm high, which MNHN Cirsotrema species. acquired from M Bonnet (a fossil, and so Two specimens from a Maoricrypta-domi- presumably from Castlecliff, Wanganui; de nated shellbed exposed on Turakina Valley Boury did not state a locality, but Cossmann Road, east of Wanganui (Fig. 5A; GS15338, [1912:52] cited Wanganui). De Boury thought S22/f0164, S22/096385, ca 600 m S of Mangara C. intermedia distinct from C. zelebori, Stream bridge and ca 200 m N of Glenroy although he admitted that he had compared Station; early Castlecliffian, ca OIS 4341) Scala intermedia with only a few abraded belong in a species distinct from Cirsotrema Recent specimens of C. zelebori, and that zelebori. They are similar in appearance to a comparison with better specimens was desir- moderate-sized specimen of C. zelebori, but able. The holotype is merely a large specimen of have a slightly wider spire angle, very wide, C. zelebori. The spiral sculpture, the number almost smooth axial ridges (heavy varices), up and position of the complex axial lamellae and to five times as thick as on any specimens of the ridges and nodules formed on them by the C. zelebori I have seen, and a very wide, spiral cords, the peribasal cord, and the aper- prominent fasciolar ridge alongside the inner ture, including the fasciolar ridge alongside the lip. However, it is unclear to what extent the inner lip, all agree exactly. The only slight smoothness of the outer surfaces of the lamellae difference, also pointed out by de Boury (apart is due to abrasion. If it is naturally smooth,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 from the slightly larger size of C. intermedia at these specimens conform to the description the same number of whorls*27 vs 24 mm), provided by A Grebneff (Otago Uuniversity is the slightly higher and thinner axial lamellae pers. comm. May 2008) of Cirsotrema forresti of the fossil. However, the varices are com- Dell (1956b:40, figs. 33, 47, 48), originally pound axial ridges as in Recent material, and proposed in C. (Tioria) Marwick, 1928. Greb- the holotype is closely comparable with many neff has concluded that the Cirsotrema species Recent specimens. The specimen identified by in deep, cool water in southern New Zealand is Fleming (1953:144) as ‘C. n. sp. aff. zelebori’ C. forresti, with a slightly lower spire than in C. (GS4117, R22/f6437) also agrees in almost all zelebori, smooth rather than spirally ridged characters with Recent specimens, and again variceal surfaces, a more prominent basal has slightly higher and thinner axial lamellae cord and fasciole than in C. zelebori, and in than most Recent specimens. There is little many specimens a nodule or spine on the doubt that this difference is trivial. shoulder (adapical extremity) of each axial Nu¨ tzel (1998:93, pl. 13IK,OQ) illustrated ridge. It seems possible that this species occurs specimens of Cirsotrema zelebori, including at Wanganui. However, it is likely that the SEM enlargements showing the micropunctate Turakina Valley Road specimens are smooth Marine Mollusca of isotope stages of the last 2 million years in New Zealand 27

because they have been abraded during rework- Beu, 1973 is correctly referred to Euthria (as is ing from Nukumaruan or older rocks, and the suggested below), it seems likely to be a member wider varices suggest that they are a species of the mostly Northern Hemisphere family distinct from both C. zelebori and C. forresti.A . It should be noted, however, that few other, relatively large specimens of C. Hayashi (2005) did not distinguish Buccinulidae zelebori from Nukumaruan localities (GS4259, from a monophyletic Buccinidae on the grounds R22/f6489, muddy sandstone below Kuranui of molecular phylogeny (determined from Limestone, Waitotara desert [Fleming complete 16SrRNA gene sequences). The 1953:130]; Nukumaru Limestone, bluff along- family position of Iredalula has long been in side Nukumaru Stream on New Plymouth- doubt, and in recent years it has usually been Wanganui highway, in GNS) include specimens referred to the Buccinidae (sensu lato), but with much thicker axial ridges than any Recent following Ponder (1968b) and Beu & Maxwell specimens seen, but are not as extreme as the (1987), it seems likely to be a member of the Turakina Valley Road early Castlecliffian ones. , which was recognized as a family The latter probably represent an unnamed by Bouchet et al. (2005). species, but better material is required to confirm their distinctive characters. An attempt Genus Aeneator Finlay, 1926 at re-collection of this material did not reveal any further specimens. Aeneator Finlay 1926b:414. Type species (by Cirsotrema zelebori is also similar to Epito- original designation): Verconella marshalli Mur- nium (Boreoscala) blainei Clench & Turner doch, 1924, Pleistocene and Recent, New Zeal- (1953:361, pl. 180), from Florida (A Grebneff and. pers. comm. May 2008). The illustrations by Clench & Turner (1953) show a specimen with ?Ellicea Finlay in Marwick 1928:432. Type thicker, slightly more fluted axial ridges and species (by monotypy): orbita Hut- coarser spiral cords than C. zelebori, although ton, 1873, Pliocene (and Recent?), New Zealand. otherwise similar in size, shape and the presence of a peribasal spiral cord. Clench & Turner ?Pittella Marwick 1928:486. Type species (by (1953:362) mentioned the similarity of E. blai- original designation): Ellicea (Pittella) valida nei to ‘E. magellanicum Philippi [1845] from Marwick, 1928, Pliocene, Pitt Island, Chatham Patagonia’. C. magellanicum is another species Islands.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 of the same group as C. zelebori, but has weaker spiral sculpture and a slightly lower Remarks. Although it has been customary in spire than C. zelebori, and is possibly descended recent years to treat Ellicea Finlay, 1928 as a from the Patagonian fossil species related to C. synonym of Aeneator Finlay, 1926 (e.g. Powell rugulosum. 1979:201; Spencer & Willan, 1996:26; Spencer et al. 2009:208), their fossil records seem to indicate that they had distinct times of origin in Clade Neogastropoda New Zealand and, therefore, possibly different Family Buccinulidae ancestors. Ellicea has the earlier first record (Tongaporutuan, early late Miocene; three Remarks. I follow Harasewych & Kantor (2004) bathyal species described) whereas Aeneator in recognizing on anatomical grounds several of huttoni Finlay, 1930 (Kapitean, latest Miocene) the Southern Hemisphere families proposed by is the earliest species recorded in Aeneator Powell (1929). In particular, most New Zealand (sensu stricto) in New Zealand. Both are ‘buccinids’ probably belong in Buccinulidae otherwise Pliocene to Recent (sub?)genera, at Powell, 1929 whereas, if ‘Buccinulum’ pangoides least in New Zealand. succinctus 28 AG Beu

Tenison Woods (1897:16, pl. 4, fig. 6, 6a) was Type material. Aeneator delicatulus holotype referred to Ellicea by Darragh (1970:198) and AIM AK72003, from between Kai-Iwi and Goudey (2006:32, figs. 6, 6a; 49, fig.1) and Okehu Streams, Castlecliff coastal section, superficially resembles Ellicea species closely, Wanganui. Powell (1929:92) also recorded a especially E. valida (Marwick) (1928:486, fig. ‘large specimen’ (here regarded as a paratype) 130) from Waipipian (Pliocene) Whenuataru in GS1163, R22/f6498, between Ototoka and Tuff, Pitt Island, . However, Okehu Streams, Wanganui coastal section, but TA Darragh (Museum Victoria pers. comm. 6 this specimen cannot be located in GNS. August 2010) informed me that T. succinctus belongs in Dennantia Tate, 1888. This removes Other material examined. Castlecliffian: the stratigraphically lowest record of Aeneator an apparently anomalously shallow-water re- delicatulus is in Kaimatira Pumice Sand (OIS cord of what is an entirely bathyal (sub)genus 25; Fleming 1953:188) in the Castlecliff coastal in New Zealand. At least three species living section. Other records are from Lower Kai-Iwi along the Chilean coast at present have been Siltstone (OIS 23), particularly common (Flem- Aeneator assigned to (McLean & Andrade ing 1953:192), Omapu Shellbed (OIS 23; Flem- 1982:12-15; Osorio & Romajo 2007) and one ing 1953:194) and Lower Westmere Siltstone referred to A. (Ellicea) (A. loisae Rehder (OIS21; Fleming 1953:196). Records from 1971:593, figs. 7, 8). Most of these species Kaikokopu Shell Grit (OIS 19; Fleming would be better referred to another (unnamed?) 1953:199, ‘Aeneator aff. delicatula’) and Kupe genus, but A. loisae does seem to be a species of Formation (OIS17) in at least two collections Ellicea, suggesting the possibility that Ellicea (Fleming 1953:203, gastropod shellbed and was a Neogene migrant to New Zealand. The Mactra tristis layer, ‘A. aff. delicatula’) are Seymour Island Eocene species referred to based on small, abraded, equivocal specimens, Aeneator by Stilwell & Zinsmeister (1990) possibly all reworked from older units. A. seem more likely to belong in the Antarctic delicatula therefore essentially is limited to genus Thiele, 1912 (Beu 2009). Ellicea Kaimatira Pumice Sand and the siltstone beds and Aeneator also possibly had an earlier, of OIS27 21 in cycles exposed in the cliffs NW unrecorded history in New Zealand. Further and SE of Mowhanau Beach, Castlecliff. Speci- research is needed on these obviously similar mens in the GNSreference collection are: Kai- generic groups, but it seems likely that they are Iwi, Castlecliff, CA Fleming collection (three);

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 not as closely related, in a phylogenetic sense, basal conglomerate member, Upper Okehu as the similar teleoconchs and radulae of Siltstone (OIS 27), coast W of Kai-Iwi, Wan- Recent species seem to indicate. Once again, ganui (GS4075, R22/f6406, one, slightly coarser this is a question that will be resolved by DNA spiral sculpture than others); upper part of Lower Kai-Iwi Siltstone (OIS 23), Castlecliff studies. section (GS4061, R22/f6392, six); Omapu Shellbed (OIS 23), coast E of Omapu Stream, Aeneator (Aeneator) delicatulus Powell, 1929 E of Mowhanau, Wanganui (GS4060, R22/ (Fig. 6E, J) f6391, and GS10909, R22/f6391A, eight); fossi- liferous silt member, Lower Westmere Siltstone Aeneator delicatula Powell 1929:91, pl. 4, fig. (Lower Westmere Shellbed; Abbott & Carter 75. 1999; OIS21), coast E of Mowhanau, Castle- Aeneator delicatulus. Fleming 1966:59; Beu & cliff (GS4058, R22/f6389, one). Maxwell 1990:361, pl. 48o. Aeneator (Aeneator) delicatulus. Maxwell Distribution. Early Castlecliffian, OIS25 21; 2009:244. 1917?, Wanganui Basin only; there are no Marine Mollusca of isotope stages of the last 2 million years in New Zealand 29 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 6 (A) Aeneator (Aeneator) marshalli (Murdoch), type species of Aeneator Finlay, 1926, lectotype, TM6837, ‘‘Castlecliff, blue sandy clays’’, Castlecliffian; height 71.7 mm. (B,C,F,H,I) Cominella (Eucominia) elegantula (Finlay); B, GS4013, R22/f6353, Tainui Shellbed (Castlecliffian, OIS 13), ‘‘the buttress’’, Castlecliff; height 30.5 mm; C,I, Recent, 2 specimens of ‘‘marlboroughensis form’’, NMNZ M.50340, RV ‘‘Acheron’’ stn. 76481, W of Makara, Cook Strait, 119132 m; C, height 31.4 mm; I, height 34.5 mm; F, GS11465, V21/f6015A, Mairau Mudstone (late Nukumaruan), ‘‘the Watchman’’ hill, Napier; height 31.8 mm; H, GS4140, R22/f6450, LowerCastlecliffShellbed(Castlecliffian,OIS15),Castlecliffcoast;height33.1mm.(D,G)Buccinulumcaudatum Powell, 2 specimens, GS4060, R22/f6391, Omapu Shellbed (Castlecliffian, OIS 23), E of Mowhanau beach, Wanganui; D, height 48.1 mm; G, height 38.8 mm. (E,J) Aeneator (Aeneator) delicatulus Powell, GS10909, R22/ f6319A, Omapu Shellbed (Castlecliffian, OIS 23), E of Mowhanau Beach, Wanganui; height 64.4 mm. 30 AG Beu

records from localities away from the excellent 1924 (Fig. 6A; GNSTM6837, ‘Castlecliff, blue exposures in the coastal section. sandy clays’). One of the other, taller syntypes was also illustrated by Murdoch (1924, pl. 10, Dimensions. Aeneator delicatulus, holotype: H figs. 2, 3) as ‘Verconella marshalli (juv.)’. 42, D 21.5 mm; large specimen in GS1163: H Dimensions of lectotype: H 71.7, D 33.5 mm. 68, D 31 mm (Powell 1929:92); GS10909, R22/ f6319A, Omapu Shellbed: H 54.4, D 27.5 mm Genus Buccinulum Deshayes, 1830 (Fig. 26A,B; specimen illustrated by Beu & Maxwell 1990, pl. 48o). Buccinulum Deshayes 1830:143. Type species Remarks. Aeneator delicatulus is a very distinc- (by subsequent designation, Iredale 1921:208): tive, small Aeneator species with a short spire, a lineatus Gmelin, 1791 ( linea long, very straight, left-inclined anterior - Martyn, 1784; valid, ICZN Opinion 479 1957), nal canal, and consistently very fine, closely Pleistocene and Recent, New Zealand. spaced, silky-looking spiral sculpture. The max- imum size achieved is smaller and the sculpture Evarne H Adams & A Adams 1853:79. Type of A. delicatulus is quite obviously much finer species (by monotypy): Buccinum linea Martyn, than that of the much more common and wide- 1784. spread, more long-ranging type species of the genus, A. marshalli (Murdoch, 1924) (Nukumar- Euthrena Iredale 1918:34. Type species (by uanRecent) and there is no doubt that these are original designation): Fusus vittatus Quoy & distinct species. A. marshalli also is not found in Gaimard, 1833, Pleistocene and Recent, New siltstone beds, and seems essentially limited to Zealand. Type B shellbeds (high-stand systems tract shellbeds) in the Wanganui succession. As Re- cent specimens occasionally wash ashore on Tasmeuthria Iredale 1925:262. Type species beaches (e.g. Ohope Beach, Whakatane), (by original designation): Siphonalia clarkei at least part of the range of A. marshalli is Tenison Woods, 1875, Recent, southeastern evidently further inshore than that formerly Australia. inhabited by A. delicatulus. The type material of Aeneator marshalli Evarnula Finlay 1926b:415. Type species (by

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 includes four syntypes representing two possi- original designation): Cominella striata Hutton, ble species; two are the relatively large, short- 1875 (Buccinum linea Martyn, 1784). spired species usually known by this name, whereas two are the smaller, more elongate, Chathamina Finlay 1928:252. Type species (by possibly distinct species A. attenuatus Powell, original designation): Tritonidea fuscozonata 1927. These all seem best regarded as syntypes Suter, 1908, Recent, New Zealand. despite Murdoch’s (1924:159,160) statements ‘Type in the Wanganui Museum’ and ‘A fossil Buccinulum caudatum Powell, 1929 (Fig. 6D, G) specimen is chosen for the type...’, as the illustrated specimen is not identified as ‘type’ in the caption, and no syntype is now labelled Buccinulum (Evarnula) caudatum Powell as ‘type’ or any other qualifying word or 1929:75, pl. 2, fig. 50; Fleming 1966:60. phrase. To ensure that this name is maintained Buccinulum lineum caudatum. Ponder 1971:243, in its accustomed usage, Murdoch’s (1924) fig. 5, nos. 13. larger illustrated syntype is here designated Buccinulum caudatum. Beu & Maxwell the lectotype of Verconella marshalli Murdoch, 1990:362, pl. 49b; Maxwell 2009:244. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 31

Type material. Buccinulum (Evarnula) cauda- within Wanganui Basin. Ponder (1971:243) trea- tum, holotype AIM AK72018, from ‘between ted B. caudatum as a subspecies of B. linea Kai Iwi and Okehu, Wanganui, Upper Plio- (Martyn, 1784). However, the species status of cene’ (Powell 1929:76), that is, early Castleclif- B. caudatum is demonstrated by both the fian beds of the Wanganui coastal section (not strongly marked morphological differences and seen). the partial syntopy of the two forms. Specimens of B. caudatum differ from those of B. linea in Other material examined. Castlecliffian: Bucci- their much larger size, in their strongly concave nulum caudatum was recorded by Fleming rather than straight or weakly concave sutural (1953:188, 192, 194, 196, 199, 203) from Kai- ramp, in their much more prominent axial matira Pumice Sand (‘B. aff. caudatum’; OIS ridges, in their longer and more prominent 25), Lower Kai-Iwi Siltstone and Omapu transverse (spiral) ridges inside the outer lip, Shellbed (OIS 23), fossiliferous silt member of and in their more flared outer margin of the outer Lower Westmere Siltstone (Lower Westmere lip, outside the transverse ridges. The proto- Shellbed; Abbott & Carter 1999; OIS 21), conch also is correspondingly larger than that of Kaikokopu Shell Grit (OIS 19), and Kupe B. linea. The stratigraphic range of B. caudatum Formation (‘derived’; OIS17). As with Aenea- is virtually identical to that of Aeneator delica- tor delicatulus, the youngest records are based tulus, and specimens of the two species occur on equivocal, abraded specimens, possibly syntopically. However, specimens of B. cauda- derived from older formations, and the true tum are more common in Kaikokopu Shellbed time range seems to be OIS25 19. Specimens than is A. delicatulus, and it seems likely that at in the GNSreference collection are: Kai-Iwi, least some of these are in situ, that is, its Castlecliff, CA Fleming collection, labelled time range more certainly extends up to include ‘topotypes’ (15); fossiliferous silt member, Low- OIS19. er Westmere Siltstone, Castlecliff coast (GS4058, R22/f6389, two); Omapu Shellbed, Genus Cominella Gray, 1850 coast E of Omapu Stream, E of Mowhanau, Wanganui (GS4060 re-collection, R22/f6391, Cominella Gray 1850:72. Type species (by sub- 30). sequent designation, Iredale 1918:34; confirmed by ICZN Opinion 479 1957): Buccinum testudi- Distribution. Early Castlecliffian, OIS25 19 neum Bruguie` re, 1798 (Buccinum (17?), in the Wanganui coastal section only. Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 maculosum Martyn, 1784; valid, ICZN Opinion Dimensions. Buccinulum caudatum, holotype: H 479 1957), Pleistocene and Recent, New 57.5, D 27 mm; paratype: H 61, D 27.5 mm Zealand. (Powell 1929:76); GS4060, R22/f6391, Omapu Shellbed: H 48.1, D 23.2 mm; H 38.8, D 18.9 Acominia Finlay 1926a:240. Type species (by mm; ‘Kai-Iwi, topotypes’, CA Fleming collec- original designation): Buccinum adspersum Bru- tion: H 52.0, D 27.3 mm; H 47.6, D 25.3 mm. guie` re, 1789, Pleistocene and Recent, New Zealand. Remarks. As with Aeneator delicatulus, Bucci- nulum caudatum is limited to early Castlecliffian Subgenus Eucominia Finlay, 1926 rocks (OIS25 19; 17?) and as it is a very large, highly distinctive species, the largest species Eucominia Finlay 1926a:239. Type species (by referred to Buccinulum (the still larger B. pan- original designation): Buccinum nassoides goides Beu, 1973 is transferred below to Euthria Reeve, 1846, PlioceneRecent, southern New Gray, 1850), it is useful for biostratigraphy Zealand. 32 AG Beu

Zephos Finlay 1926a:239. Type species (by Other material examined. Mangapanian: a sin- original designation): cingulata Hutton, gle specimen from Weka Pass, N Canterbury 1885, Pliocene, New Zealand. Synonym of Eu- (GS6215, M34/f7660, 50 m above Waiauan cominia according to Ponder (1968a:35). limestone, Weka Creek, 160 m downstream from junction with Weka Pass Stream; with Cominella (Eucominia) elegantula (Finlay, 1926) Taxonia suteri, waipipiensis (Marwick, (Fig. 6B, C, FI) 1924), and Glycymerita (Manaia) manaiaensis (Marwick, 1923)) resembles Cominella elegan- Eucominia elegantula Finlay 1926a:240, pl. 57, tula and C. virgata closely in most characters, figs. 14, 15. but is a little larger than any specimens from Eucominia elegantula subspecies verrucosa Fin- Wanganui or Hawke’s Bay and has rather more lay 1926a:241, pl. 57, fig. 16. prominent spiral cords. It possibly represents Cominella (Eucominia) elegantula. Powell an unnamed species. Nukumaruan: 30 m above 1929:95; Fleming 1966:62; Ponder 1968a:35; Torlesse basement, Makohine Stream, Pohan- Beu & Maxwell 1990:364, 412, pl. 49e; Maxwell gina, E Wanganui Basin, early Nukumaruan 2009:244. (GS2763, T23/f6486; one); Tangoio Limestone, Cominella (Eucominia) elegantula verrucosa. roadside quarry on Waipunga Road, 1 km N of Powell 1929:95. junction with Turnbull Road, central Hawke’s Cominella (Eucominia) marlboroughensis Bay (GS12711, V20/f045; 12 incomplete); Mair- Powell 1946b:143, pl. 12. fig. 10 (new synonym). au Mudstone, ‘the watchman’ hill, Ahuriri Eucominia marlboroughensis. Dell 1956b:52, lagoon, Napier (GS11465, V21/f6015A; eight); fig. 60. Okawa Stream, Matapiro syncline (GS1063, Eucominia cf. elegantula. Dell 1956b:53, fig. 62. V21/f8476; four); limestone underlying shellbed Cominella (Eucominia) verrucosa. Fleming on Trig. Y, 1 km NW of Paeroa Road, 2 km 1966:62. SW of Whetukura on OrmondvilleTe Uri Cominella (Eucominia) elegantula marlborough- Road, ca 15 km ENE of Dannevirke . Ponder 1968a:35; Powell 1979:195, pl. 40, (GS2365, U23/f6410; two); Mairau Mudstone, fig. 5; Spencer & Willan 1996:27; Spencer et al. walkway W side of Roro-o-kuri, Ahuriri, 2009:208. Napier (GS12386, V21/f05; two); Maharakeke Mudstone, Maharakeke Road, W of Waipu- kurau, Hawke’s Bay (GS10857, U23/f7049;

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Type material. Eucominia elegantula, holotype one); Tirohia Station, opposite Maraekakaho, AIM AK70218, from Castlecliff, Wanganui, Matapiro syncline (GS12424, V21/f013; 10); horizon unknown, possibly from Tainui Darkys Spur Formation?, lowest limestone Shellbed (OIS 13); with one paratype bed in sequence, Waipunga Road, N side of AK71669, same data as holotype. Eucominia hairpin bend on ascent from Esk Valley to- elegantula verrucosa, holotype AIM AK70282 wards Kaiwaka Road, N of Napier, central (O Lee, AIM, pers. comm. 11 March 2008); Hawke’s Bay (GS12816, V20/f147; three, al- Finlay (1926a:241) stated the locality only as most smooth); Tangoio Limestone, cutting on ‘same as last, with the species itself, but rarer’, track 500 m N of Petane Pa, N of Napier, that is, he used ‘subspecies’ in the sense of a central Hawke’s Bay (GS12790, V20/f038; five, sculptural variant rather than the modern usage most incomplete and abraded, almost smooth); of a geographically isolated form. Cominella Mairau Mudstone, ‘Petane corner’, junction of (Eucominia) marlboroughensis, holotype AIM Highways 3 and 5, opposite Esk River bridge, AK71453, Recent, from 150 m, 41816?S, N of Napier (GS12501, V20/f017; one, smooth, 173817?E, ca 40 km E of Cape Campbell, more inflated than all others, referred uncer- Cook Strait (all not seen). tainly). Recorded also by Fleming (1953:151) Marine Mollusca of isotope stages of the last 2 million years in New Zealand 33

from GS3093, S22/f9489, Waipuru Shellbed (?), common. Most specimens of C. elegantula are Hunterville-Turakina Valley Road 5 km W of abraded, whereas C. excoriata is not abraded Hunterville, Rangitikei valley. Castlecliffian: when collected from offshore siltstone units, so common in most later Castlecliffian highstand apparently C. elegantula lived in relatively systems tract shellbeds at Wanganui, not listed; shallow water, and C. excoriata lived signifi- recorded by Fleming (1953:203204, 209, 218, cantly further offshore in Hawke’s Bay. This 230, 239) from gastropod shellbed, Cyclomac- probably reflects a facies limitation rather than tra tristis, and bivalve shellbed members of the real time range, reducing any possible Kupe Formation (OIS17), undifferentiated biostratigraphical utility. Kai-Iwi Group, W side valley (GS4235, S22/f6457, ‘cf. verrucosa’), Lower Dimensions. Holotype of Cominella marlbor- Castlecliff Shellbed (OIS 15), Tainui Shellbed oughensis: H 29.3, D 13.4 (Powell 1946b:143); (OIS13), and Upper Castlecliff Shellbed(OIS Cook Strait: H 26.8, D 12.7 mm; H 25.4, D 12.3 11). Recent: the holotype of Cominella marlbor- mm; H 32.7, D 17.7 mm (Dell 1956a); GS4013, oughensis and the material reported by Dell R22/f6353, Tainui Shellbed at ‘the buttress’, (1956b:52, 53) as ‘Eucominia marlboroughensis’ Castlecliff: H 30.5, D 14.6 mm; GS4140, R22/ and ‘Eucominia cf. elegantula’ are from Cook f6450, Lower Castlecliff Shellbed, Castlecliff: Strait (150 m, 40 km E of Cape Campbell, H 33.1, D 15.0; GS2763, Makohine Stream, Cook Strait, holotype; 275 m, 41828.5?S, Pohangina, Nukumaruan: H 32.2, D 16.5 mm; 174850?E, & 110 m, between Island Bay Bank GS10857, Maharakeke Road, Waipukurau, and East Bank [Dell 1956b:52], and 137 m, Nukumaruan: H 33.8, D 18.5 (unusually wide 40852.6?S, 174849.5?E [Dell 1956a:53]); 15 other specimen); GS12424, Tirohia Station, central lots in NMNZ, ranging from Kahurangi Hawke’s Bay: H 35.0, D 17.6 mm; H 28.2, D Shoals, N Westland, 91 m (M.59050, four 15.1 mm; H 28.4, D 14.5 mm; GS11465, ‘the small) and off Westhaven Inlet, to off Kapiti watchman’, Napier: H 31.8, D 16.3 mm; Island (M.7681, two), throughout Cook Strait, Recent, NMNZ M.50340, W of Makara, W to Port Gore, Marlborough, 139144 m Cook Strait, 119132 m, 41811.00?S, (M.53717, one small) and to Palliser Bay, E 174839.00?E, RV Acheron sta. 76481 (Fig. Cook Strait, in 274 m (M.11543, one). The 26I,K): H 34.5, D 16.8 mm (dorsal view); H largest lot is M.135047, Nicholson Canyon, off 31.4, D 16.2 mm (ventral view). Turakirae Head, 640 m (24 stained, dead

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 specimens). Remarks. Ponder (1968a:37) described the new Recent ‘subspecies’ Cominella (Cominella) ex- Distribution. Mangapanian, N Canterbury?; coriata tolagaensis on the basis of a large early (rarely) and late Nukumaruan in Hawke’s number of specimens from and Bay and (rarely) Wanganui Basin; common in nearby areas in the eastern North Island, north later Castlecliffian rocks of Wanganui Basin; of Gisborne, where it occurs as far N as the still living offshore in Cook Strait in about point E of Te Araroa on the N coast of East 110280 m. Available records suggest that in Cape. It was stated expressly to be a ‘chron- Hawke’s Bay, Cominella elegantula is limited to osubspecies’ of C. (Cominella) excoriata,a upper beds of Petane Group (Darkys Spur highly distinctive species occurring in high Formation to Te Ngaru Mudstone, and possi- Nukumaruan formations of the Petane Group bly a few units above and below these), of late in central Hawke’s Bay. One of the themes of Nukumaruan age. However, I have not seen this series of papers is that chronosubspecies do specimens from the highest, richly fossiliferous not exist; speciation (or origination) produces unit, Devils Elbow Mudstone, where Comi- discrete species and, if no speciation event nella (Cominella) excoriata (Finlay, 1926) is occurred, members of a gradual anagenetic 34 AG Beu

series are conspecific. In some ways C. elegan- Bay (Fig. 26J). C. elegantula occurs fairly tula more nearly resembles C. tolagaensis than commonly in shellbeds in the Castlecliff sec- does C. excoriata. It is clear, though, that these tion, Wanganui, and not uncommonly in actually are three distinct species. Nukumaruan siltstone units in central Hawke’s Cominella excoriata is distinguished readily Bay. C. elegantula is a small, weakly sculptured from all other New Zealand species by its northern end-member of C. nassoides (Reeve, prominent, narrow, almost smooth, widely 1846), with a less prominent sutural ramp than separated, obviously sinuous axial costae and other forms and only weak or, in some speci- its low, weakly defined spiral cords. Its outer lip mens, no peripheral nodules on its regular, is sinuous in conformity with the axial costae. It even, axially elongate ridges. Specimens re- has no clearly defined sutural ramp, but the corded from the Recent fauna in Cook Strait area below the suture is slightly flattened by Powell (1946b) and Dell (1956b) as C. through the posterior (adapical) weakening of (Eucominia) marlboroughensis and further ma- the axial costae. C. tolagaensis has wider but terial examined in NMNZ demonstrate that less prominent, more closely spaced, and much closely similar, small, weakly sculptured speci- less sinuous axial ridges than C. excoriata, but mens intergrading with C. elegantula still live in has much more prominent, wide, closely spaced the Cook Strait area. The main point for the spiral cords than any other New Zealand present paper is that C. elegantula is a single, Cominella species, with each spiral interspace rather variable species (or geographical sub- filled with 12 much narrower, closely spaced species of C. nassoides; BA Marshall NMNZ cords. The prominent, wide, closely spaced pers. comm.) of Nukumaruan and possibly axial ridges and outer lip are nearly straight, Mangapanian to Recent age, offering no bios- as in almost all other Cominella species, but in tratigraphical criteria through the distinction stark contrast with their obvious, strong sinu- between a fossil species C. elegantula and a osity in C. excoriata. It also differs markedly Recent species C. marlboroughensis. Also, C. from C. excoriata in its taller, more obvious, (Eucominia) elegantula, C. (Cominella) tola- more deeply concave sutural ramp. Its pattern gaensis and C. (Cominella) excoriata definitely of ‘chequerboard’ colour maculations also is are three distinct species. unique for a New Zealand Cominella species, other than the still more ‘square-patterned’, Cominella (Eucominia) incisa (Hutton, 1885) essentially smooth species C. maculosa (Fig. 7AG)

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (Martyn, 1784), although it is not unlike that of Australian species such as C. lineolata incisa Hutton 1885a:328. (Lamarck, 1809) (Wilson 1994, pl. 9, fig. Nassa incisa. Hutton 1887:210. 5AC) and C. acutinodosa (Reeve, 1846) (Wil- incisus. Suter 1915:26, pl. 4, fig. 13. son 1994, pl. 9, fig. 8AB). Some specimens in Zephos incisa. Finlay 1926a:240; Fleming Hawke’s Bay shallow-water late Nukumaruan 1966:62. rocks differ from both C. tolagaensis and C. Cominella (Eucominia) incisa. Beu & Maxwell excoriata in their slightly smaller size, lower 1990:412; Maxwell 2009:244. spire, weaker spiral sculpture, weakly devel- oped sutural ramp, and in most specimens Type material. Clathurella incisa holotype CMC almost completely lacking axial sculpture. M3259 (again, not included in the catalogue by However, careful comparison has demon- Bradshaw et al. 1992), from ‘Te Aute, Hawke’s strated that these are merely abraded specimens Bay. Collected by Mr. A. Hamilton’ (Hutton of C. (Eucominia) elegantula, and more coar- 1885a:328; the collector is stated only in the sely sculptured specimens of C. elegantula also original publication). Age stated by Suter occur at nearby, coeval localities in Hawke’s (1915:26) as ‘Miocene’. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 35 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 7 (A,G) Cominella (Eucominia) incisa (Hutton), holotype, CMC M3259, ‘‘Te Aute, Hawke’s Bay’’, between Raukawa Range and Mason Ridge?, early Nukumaruan?; height 20.0 mm. (B,D) Euthria pangoides (Beu), GS4253, Q22/f7544, Upper Waipipi Shellbed (Waipipian), Waverley Beach, W of Wanganui; A, holotype, TM5092, height 50.5 mm; D, paratype, TM5093, height 61.5 mm. (C,I) Antizafra speighti Marwick, GS10858, U22/f9588, Ashcott Road, W of Pukeora hill, W of Waipukurau, Hawke’s Bay, Nukumaruan; H, whole specimen, height 10.6 mm; I, protoconch of H. (E) Iredalula striata (Hutton), ‘‘Wanganui, Pliocene’’, Suter collection no. 50; height 30.8 mm. (F,H,J) Antizafra pisanopsis (Hutton), type species of Antizafra Finlay, 1926, all data as for Figs. C,I; SEM, 2 specimens; F, whole specimen, height 9.7 mm; H, broken specimen showing columellar plait, height 8.1 mm; J, protoconch of F. 36 AG Beu

Other material examined. Road cut E side of is well-preserved on the holotype and, as in C. Raukawa Road, on curve ca 300 m Sof onokeana, it is large (1.75 mm in diameter) and junction with Burma Road, W of Raukawa consists of two smooth, upright, evenly inflated Range, ca 4 km NW of Te Hauke and Te Aute whorls with a flattened apex and a minute College, Hawke’s Bay; in early Nukumaruan planorboid initiation. The protoconch termi- siltstone between Te Onepu Limestone nates in a short section bearing six narrow, (Mangapanian) and Mason Ridge Limestone closely spaced, smooth but shouldered axial (early Nukumaruan) (GS12694, V22/f351; one ridges, after which the normal spiral sculpture incomplete). and more widely spaced axial ridges commence. An identical protoconch is present on the Distribution. Recorded only from the two listed specimen in GS12694. The closest similarity is specimens. Too rare to have any biostratigra- with C. onokeana, which differs from C. incisa phical utility. in its slightly taller spire, its taller, more steeply sloping and less well-defined sutural ramp, and Dimensions. Holotype: H 20.0, D 11.1 mm. in the spiral cords around the peripheral, maximally inflated area of the last whorl being Remarks. Cominella (Eucominia) incisa is al- slightly but obviously more widely spaced. most as little-known and poorly understood as Hutton’s original ‘Te Aute’ locality for Mitrellatoma angustata (Hutton, 1886) (see Cominella incisa initially seemed unlikely to be below). However, the taxonomic position of correct, as there are few localities with fossil C. incisa is much clearer than that of M. molluscs, other than Pectinidae in Pliocene Te angustata. Suter’s (1915, pl. 4, fig. 13) illustra- Aute facies limestone, where this species might tion (the only previous one) and examination of have been collected close to Te Aute College the holotype (Fig. 7A, G) make it clear that it is (alongside Highway 2, near Te Hauke in south- a small Cominella (Eucominia) species in the central Hawke’s Bay). Some years ago I col- group typified by C. onokeana (King) lected a single incomplete spire of this species (1933:345, pl. 37, fig. 14) and C. otagoensis from siltstone cropping out along Raukawa (Finlay, 1926). Finlay (1926a:240) included all Road, between the Raukawa Range and the E these in his genus Zephos, now regarded as a side of Mason Ridge, NW of Te Aute College, synonym of Cominella (Eucominia). As the apparently explaining the previously enigmatic specimen is turned slightly to the left in Suter’s provenance of Hutton’s holotype. If this is

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 illustration, it is not clear that it actually has a indeed near the type locality of C. incisa,itis narrow and moderately contracted about the same age (earliest Nukumaruan) as base of the last whorl, and a very prominent, C. onokeana. ridge-margined siphonal fasciole. With its pro- Most material assigned to Cominella (Eu- minent sculpture of narrow, well raised, closely cominia) onokeana is from the Palliser Bay cliffs spaced spiral cords (nine on the penultimate between Lake Ferry (on Lake Onoke) and the and 13 on the last whorl, becoming markedly mouth of Whangaimoana Stream, South Wair- coarser and more widely spaced over the base) arapa, and further east along the same cliffs. A crossing high, relatively narrow, evenly specimen from Okawa Stream, Matapiro syn- rounded axial folds (14 on the penultimate cline, Hawke’s Bay (GS1063, V21/f8476) la- and 16 on the last whorl), it resembles C. belled ‘Zephos onokeana’ by J Marwick is powelli (Fleming, 1948) and C. cantuariensis severely abraded, and may well be a poor speci- (Dell, 1956), although it differs from this men of C. incisa. However, I have also previously species group in its more rounded axial ridges; recorded specimens of C. onokeana from bathyal they are narrowly angular and more widely mudstone in the Ohara depression, east of the spaced in the C. powelli group. The protoconch Ruahine Range in inland Central Hawke’s Bay Marine Mollusca of isotope stages of the last 2 million years in New Zealand 37

(Beu et al. 1977, table 1) (GS11464, U21/f8558, and, indeed, it was placed there ‘with some overlying Sentrybox Limestone (with Zygochla- hesitation’ originally, mainly because Buccinu- mys delicatula) in small outcrop on Mangleton lum is the traditional New Zealand genus in Road; GS11781, U21/f02, Jumped-up Stream which to include relatively small, elongate, downstream from Mangleton Road), so these weakly sculptured buccinoideans. B. pangoides species are probably distinct. Cominella (Euco- differs from species of Buccinulum in its larger minia) cingulata (Hutton, 1885) (Nassa (Tritiar- size (H 5061.5, D 24.832.3 mm), its wider ia) cingulata Hutton 1885a:327, ‘Greta, shape, its much weaker sculpture (little sculp- Canterbury’, ‘perhaps a Cominella’), a similar ture other than low, smooth, widely spaced species common in bathyal mudstone (Greta spiral cords around the neck of the last whorl), Formation) near Waikari, North Canterbury its unusually weakly impressed suture produ- (early Nukumaruan), differs from C. incisa and cing an almost straight spire outline, its tall, C. onokeana in its much wider, lower spiral steep, concave sutural ramp riding up to the cords, separated only by linear grooves periphery of the previous whorl, its short, (GS3297, N33/f9112, Greta Creek at main north widely open anterior with little road, 5 km Sof Hurunui River, N Canterbury; constriction on the right (outer) side, its tall, 14 specimens, labelled by J Marwick and CA narrow posterior canal produced by the poster- Fleming ‘topotypes of cingulata’ and ‘also topo- iorly elongate sutural ramp, its row of short but types of purchasi’). An almost equally poorly prominent spiral lirae inside the outer lip and, known species is Cominella (Eucominia) purcha- above all, the obvious, sharp, but relatively si Suter (1917:34, pl. 4, fig. 20), also from weak angulating ridge on the base of the ‘Waikari Valley, near the Greta Railway-station, columella, forming a low columellar fold that right on the bank of the stream’ (Suter 1917:34), is not present in any Buccinulum species or, that is, possibly from the same locality as C. indeed, in any similar New Zealand Buccinidae. cingulata. Suter’s illustration shows a shell with The columellar fold is accompanied by a few slightly narrower spiral cords than most material small, narrow nodules nearby on the inner lip, of C. cingulata, but agreeing in all other char- and the parietal ridge is unusually small, short, acters, and C. purchasi is taken here to be a narrowly pointed and prominent. These char- synonym of C. cingulata. The relationships of all acters are distinctive, and produce a shell with these similar species, C. incisa, C. onokeana and an overall appearance that is much heavier, C. cingulata, require clarification from further, thicker, wider and smoother than any Buccinu-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 well-localized material. lum species. The most similar genus I am aware of is Euthria, which is very similar to Buccinulum Family Buccinidae? pangoides in shape, general appearance, and relatively weak sculpture. Most European and, Genus Euthria Gray, 1850 in particular, West African Euthria species (Cosel 1982; Cosel & Burnay 1983; Rola´ n Euthria Gray 1850:67. Type species (by mono- 1985; Rola´ n et al. 2003, 11 Cape Verde Islands typy): Fusus lignarius ‘Chiaje’ (Fusus lignarius species discussed; most illustrated by Ardovini Lamarck, 1816,Murex corneus Linne´ , 1758), & Cossignani 2004:162163; Robin 2008:185) Pliocene to Recent, MediterraneanSPortugal. as well as South African Euthria species (E. ponsonbyi GB Sowerby III 1889:149, pl. 3, fig. 3; E. quecketti EA Smith 1901:110, pl. 1, fig. 1; Remarks. Several colleagues have expressed an Barnard 1959:169172; but E. filmerae G. B. opinion to me that ‘Buccinulum’ pangoides Beu Sowerby III, 1900 was transferred to (1973:323) is not correctly placed in Buccinulum (sensu lato) by Kilburn 1970:41) and a species 38 AG Beu

from Indonesia (E. effendyi Fraussen & Dhar- due to abrasion) and the tall, narrow posterior ma [2002]; Bali Strait, in 3040 m) have a canal. The exceedingly battered holotype of slightly to markedly longer anterior siphonal Pangoa mira Marwick, 1931 cannot be placed canal, and most lack the parietal nodule of in a family with any confidence. Attempts to re- Buccinulum pangoides and have only a low, collect further material from the type locality weak ridge on the base of the columella. The have been unsuccessful (‘Tutamoe conglomer- two Philippine Islands species referred to Eu- ate’, that is, basal, coarse-grained, fossiliferous thria by Fraussen (2003) were transferred to beds of turbidite sequences, in ‘James creek’ () by Snyder & Bou- (informal local name), a tributary of Pangopan- chet (2006). Most Euthria species also have a go Stream, north of Gisborne). The fauna at this prominent to low but obvious siphonal fasciole, site (GS13955, Y16/f564, grid ref. Y16/558152; 1 which is lacking in Buccinulum pangoides. km E of Te Rimu station, Tauwhareparae, inland However, the range of characters of the 17 or between Tolaga and Tokomaru Bays) consists of more Recent species now referred to Euthria is poorly preserved, calcite neomorphs of a Lill- very wide, and E. quecketti and some of the burnian (middle Miocene) bathyal fauna. It species named recently from the New Caledo- includes a common large, apparently unnamed, nian region have a small, narrow parietal angular Neilo species, and probably includes the nodule, a nodule on the base of the columella, specimens of unidentified Zemacies, Gemmula and almost no siphonal fasciole, and despite and Comitas species. It is admixed with battered their longer anterior siphonal canals they fragments of a shallow-water fauna (, resemble Buccinulum pangoides more than other Eucrassatella, Struthiolaria (Callusaria) callosa Euthria species do. Particularly similar New Marwick, 1924, , Zeacolpus, Cominella Caledonian species are Euthria philpoppei hendersoni Marwick, 1926) transported down- Fraussen (2002) and E. solifer Fraussen & slope into the deposition site. The holotype of Hadorn (2003); the latter resembles E. quecketti Pangoa mira apparently has been severely closely. Most Euthria species also have a abraded by this transport, and would have been relatively large, wide, essentially smooth, pau- better left unnamed. It has been placed in cispiral (lecithotrophic) protoconch, and that of by convention in recent catalogues, the holotype of B. pangoides seems to have been but seems just as likely to be a fasciolariid. similar, although with a smaller initiation than Ironically, numerous authors in the past most other named Euthria species. E. philpoppei referred many other New Zealand buccinoi-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 is similar to B. pangoides in overall shape as deans to Euthria (e.g. see comments by Finlay well, but differs in its more strongly concave 1928:250) as well as such muricids as Axymene sutural ramp, its low, rounded but obvious aucklandicus (Smith, 1902) (see below), but all nodules on the subsutural fold and on the proved to belong in other genera. In particular, periphery of all but the last whorl, and its many early authors referred species to Euthria slightly longer anterior siphonal canal. It that are now included in Buccinulum. However, has similar apertural armature, a similarly Buccinulum differs from Euthria in its smaller weak siphonal fasciole and, in particular, size, smaller protoconch, relatively larger par- similar short transverse lirae inside the outer ietal ridge, more elongate shape, and consistent lip to those of B. pangoides.‘Buccinulum’ spiral sculpture and colour pattern. Buccinulum pangoides seems to be the first authentic New is limited to New Zealand and southern Aus- Zealand species of Euthria. tralia. The resemblance is probably, again, The resemblance of Euthria pangoides to superficial, as Euthria presumably is a genus of Pangoa Marwick (1931:119, pl. 12, figs. 232, the mostly Northern Hemisphere family Bucci- 233) is entirely superficial, and is produced nidae, whereas Buccinulum was recognized by largely by the weak sculpture (perhaps entirely Harasewych & Kantor (2004) as belonging in a Marine Mollusca of isotope stages of the last 2 million years in New Zealand 39

distinct southern subfamily (or family?) Bucci- coiled and planorbid and the later whorl nulinae/idae. The recognition that Euthria oc- inflated’, significantly different from the smal- curs in South Africa and extends at present as far ler, paucispiral, mammillate protoconch with a east as Indonesia and New Caledonia suggests large initiation (evidently reflecting lecitho- that this is another of the groups that entered the trophic development) present in Buccinulum, Pacific from the Atlantic via South Africa, and and similar to that of some species of Euthria. leaves no great surprise that a species of Euthria extended down the Norfolk Ridge to New Zealand during the warm middle Pliocene. Family Colubrariidae

Euthria pangoides (Beu, 1973) (Fig. 7B, D) Genus Iredalula Finlay, 1926

Buccinulum pangoides Beu 1973:323, figs. 27, Iredalula Finlay 1926a:231. Type species (by 3033; Beu & Maxwell 1990:298, pl. 37s. original designation): striata Hutton, Buccinulum (?) pangoides. Maxwell 2009:244. 1873, early PliocenePleistocene (OIS11), New Zealand. Type material. Holotype TM5092, GS4253 re- collection, Q22/f7544, lower part of Upper Wai- Remarks. The family position of Iredalula has pipi Shellbed, 200 m E of mouth of Wairoa long been in doubt as, although Hutton Stream, Waipipi Beach, that is, W end of (1873b:5) and several later workers referred Waverley Beach, W of Wanganui; with five the type species to various genera of Conoidea, paratypes,TM5093-5097,fromthesamelocality. traditionally it has been referred to the Bucci- nidae in recent years. Ponder (1968b) placed it Other material examined. A few other speci- in family Colubrariidae based on its close mens from the same locality and shellbed in the similarity to genera such as Schu- collection of FD Chambers. macher, 1817 and, in particular, H. Adams & A. Adams, 1853, and described some Distribution. Euthria pangoides is still recorded anatomical characters. Beu & Maxwell only from the lowest, soft, matrix-supported (1987:62) followed a verbal suggestion by WF part of Upper Waipipi Shellbed, with a fauna Ponder in placing the group in Family Bucci- dominated by relatively small gastropods (un- nidae, Subfamily Pisaniinae. Iredalula species

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 like the upper, specimen-supported part of the differ from Metula species mainly by their finer shellbed with its fauna dominated by large sculpture dominated by spiral cords, with bivalves), at Waipipi, Waverley Beach. no axial elements other than faint growth lines, their larger, wider and more obviously shoul- Dimensions. Holotype: H 50.5, D 24.8 mm; dered protoconchs, and their almost complete tallest paratype, TM5093: H 61.5, D 28.9 mm; lack of varices. The outer lip is sinuous as in widest paratype, TM5094: H 60.2, D 32.3 mm Metula, but only lightly thickened inside in I. (Beu 1973:324). striata, although a slight terminal varix is formed by the Recent species I. alticincta Remarks. Little can be added to the description (Murdoch & Suter, 1906). Oliverio & Modica of Euthria pangoides given under the generic (2010) demonstrated on molecular phylogenetic heading above. The protoconch remains largely grounds that Colubraria and Metula belong in a unknown. I (Beu 1973:323) described the pro- separate family Colubrariidae, and are followed toconch as ‘perhaps only slightly damaged on here. A position for Iredalula near Metula in holotype where it appears to consist of one and the Colubrariidae seems likely, bur requires a half whorls, the initial half-whorl tightly DNA confirmation. 40 AG Beu

Iredalula striata (Hutton, 1873) (Fig. 7E) immature ‘undoubted examples’ of ‘’ striata. Bela striata Hutton 1873b:5. Siphonalia(?) cingulata Hutton 1885a:315; Hut- Other material examined. Finely sculptured ton 1893:41. specimens attributed to Iredalula striata: Opoi- Daphnella striata. Hutton 1887:214; Hutton tian: GS2891, J32/f9089, Shamrock Creek, near 1893:52, pl. 7, fig. 33; Harris 1897:63; Suter Kapitea Creek, ca 1 km upstream from Gold- 1913, appendix p. 1084 (in part not Bela striata sborough, Westland (one incomplete); Hutton, 1873); Suter 1915:42. GS12521, X19/f033, mudstone just below striata. Suter 1907:265; Suter Opoiti Limestone, 100 m N of prominent 1910:11; Suter 1913:488, pl. 46, fig. 27 (in part limestone outcrop on Mangapoike Valley not Bela striata Hutton, 1873). Road, N Hawke’s Bay (one small). Waipipian: Iredalula striata. Finlay 1926a:231; Fleming GS1586, X19/f7480, sandstone above Tahaenui 1966:60; Beu & Maxwell 1987:62; Beu & Limestone, Makeretu Stream, tributary of Maxwell 1990:364, 412, pl. 49f. Mangapoike River above Haupatanga Gorge, Not Iredalula striata. Ponder 1968b:221, figs. 9, N Hawke’s Bay (one fragmentary). Castleclif- 10; Powell 1979:204; Spencer & Willan 1996:27; fian: recorded by Fleming (1953:188, 192, 194, Spencer et al. 2009:208 (I. alticincta Murdoch 200, 203204, 207, 209, 226, 230, 232, 239) & Suter, 1906). from Kaimatira Pumice Sand (OIS 25), Lower Kai-Iwi Siltstone and Omapu Shellbed (OIS 23), Upper Westmere Siltstone (Upper West- Type material. Bela striata, type material miss- mere Shellbed, OIS 19), Kupe Formation and ing according to Suter (1915:42), but two Upper Kai-Iwi Siltstone (OIS 17), undifferen- syntypes are present in GNS, TM8686-7, from tiated Kai-Iwi Group, Pinnacle Sand (OIS 15- ‘Wanganui (U), Shakespeare Cliff’, Castleclif- 14), Tainui Shellbed and Shakespeare Cliff fian (almost certainly from Shakespeare Cliff Siltstone (OIS 13) and Upper Castlecliff Siltstone, OIS 13). The syntypes are labelled Shellbed (OIS 11). I have not seen specimens ‘Daphnella striata’ and ‘Acamptochetus stria- from higher in the section at Wanganui. tus’, ‘the larger sp. of 23 mm is the lectotype’, More coarsely sculptured specimens attrib- both by J Marwick and on an earlier label, but I uted to Iredalula alticincta: Nukumaruan: am not aware of a published lectotype designa- GS5216, V20/f8517, road cutting NE of Puke-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 tion. They also have a very early hand-written tapu, inland from Napier, central Hawke’s Bay, label (by FW Hutton?) reading ‘Bela striata’, late Nukumaruan (one spire, moderately finely and bear the old dark brown insoluble glue sculptured); GS12386, V21/f05, Mairau Mud- characteristic of specimens that have been stone (late Nukumaruan), N end of Roro-o- removed from Hutton’s wooden type-specimen kuri, N end of Ahuriri lagoon, Napier (one tablets. The two specimens are conspecific, and moderately finely sculpturedtwo apices); a lectotype is not required. Siphonalia(?) cingu- GS10612, T27/f6021, mudstone beneath Puke- lata four syntypes CMC M-3206 (not seen) nui Limestone (early Nukumaruan), E of (Suter 1915:42; Bradshaw et al. 1992:36; but summit of Admiral Road, Maungaraki Range, note that Bradshaw et al. listed them under SE (one large, coarsely sculptured); Bela striata, whereas these are definitely syn- NMNZ M.40402, Lophelia coral thicket (early types of Siphonalia(?) cingulata, and not of Bela Nukumaruan) in cliffs E of Lake Ferry, Palliser striata), from ‘Wanganui’, that is, siltstone beds Bay (two coarse apices); M.59368, mudstone of the Castlecliff coastal section. Suter beneath Pukenui Limestone (Mangapanian/ (1915:42) examined the syntypes and confirmed early Nukumaruan), cutting on Tupurupuru- that Siphonalia(?) cingulata was based on Te Wharau Road, E of Gladstone, SWairarapa Marine Mollusca of isotope stages of the last 2 million years in New Zealand 41

(six, coarsely sculptured); M.117614, large gully remaining in Suter’s collection (GNS TM943): between Lake Ferry and Whangaimona, Palli- H 18.8, D 7.75 mm (H:D2.43) (accompanied ser Bay, early Nukumaruan (one coarse apex). by three small syntypes, TM944-945); holotype Castlecliffian: GS13707, W15/f9583A, one; of I. venusta: H 23.75, D 8.0 mm (H:D2.97) GS14551, W15/f9583B, many; NMNZ (Powell 1934b:158); large specimen of I. alti- M.40277, many; all from cutting on Wainui cincta, GS10612, Admiral Road, S Wairarapa: Road, W side Ohiwa Harbour, Whakatane H 25.9, D 8.9 mm (H:D2.91). (OIS15?). Recent: 53 lots examined in NMNZ, ranging from off Ahipara, around Remarks. Comparison of fossils and the Recent the E North Island to off Palliser Bay, in specimens in NMNZ revealed a surprising 10550 m. taxonomy of Iredalula. No Recent specimens are as finely and evenly sculptured as Castlecliff Distribution. Finely sculptured specimens of specimens of I. striata, which have about 5060 Iredalula striata are rare in Opoitian and very narrow, low, closely spaced (almost un- Waipipian rocks of Westland and N Hawke’s countably fine) spiral threads on the penulti- Bay, and common in Kaimatira Pumice Sand mate whorl. Recent specimens have 620 wider (OIS25) and siltstone formations of OIS23 11 and more widely spaced spiral cords on the at Castlecliff, Wanganui, particularly in Shake- penultimate whorl. The sculpture of Recent speare Cliff Siltstone (OIS 13), although there specimens is exceedingly variable and, although are no records from OIS21. No Recent speci- coarsely sculptured specimens similar to the mens are now identified as I. striata. The syntypes of I. alticincta (Murdoch & Suter Recent species I. alticincta lives from the NW 1906:285, pl. 22, figs. 12, 13) tend to occur North Island (off Ahipara) to Palliser Bay, most commonly towards the southern limit of Cape Campbell and into E Cook Strait; the few its range (from Gisborne to Palliser Bay and specimens from the W North Island are shorter into E Cook Strait) and in deeper water (in ca and possibly distinct. Fossils of I. alticincta are 100550 m), even northern shallow-water speci- recorded only from Castlecliffian rocks at mens can be very coarsely sculptured, and most Whakatane and from Nukumaruan rocks of specimens are intermediate in sculptural coar- Hawke’s Bay and Wairarapa where, as in the seness between the types of I. venusta Powell Recent fauna, specimens in shallow-water for- (1934b:157, pl. 22, fig. 9) (with eight spiral mations tend to be finely sculptured whereas cords on the penultimate whorl) and I. alticinc-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 those in bathyal deposits are all very coarsely ta (with five cords). Others, particularly in the sculptured. north of its range from E Northland to the Bay of Plenty, are intermediate in coarseness be- Dimensions. Syntypes of Bela striata: H 22.5, D tween the holotype of I. venusta and Castlecliff 9.3 mm (H:D2.42) (marked ‘T’ inside outer specimens of I. striata, but none is anywhere lip); H 21.0, D 8.2 mm (H:D2.56); large, near as finely sculptured as Castlecliff speci- relatively narrow specimen of Iredalula striata mens. Some samples include a range of sculp- (Fig. 27J), ‘Wanganui, Pliocene’, Suter collec- ture within one collection (e.g. NMNZ tion no. 50: H 30.3, D 11.3 mm (H:D2.68); M.67634, W of Plate Island, 5964 m, six largest in NMNZ: M.26767, Castlecliff: H 30.8, relatively finely sculptured I. venusta form, D 12.4 mm (H:D2.48); H 27.4, D 12.1 mm one I. alticincta form). The holotype of I. (H:D2.26); largest specimen of I. alticincta, venusta is from off Cape Campbell, Marlbor- M.61081, ENE of Castlepoint, 278 m: H 35.3, ough, the southernmost record of the genus. D 11.2 mm (H:D3.15); figured syntype of I. Very coarsely sculptured fossils do occur, but alticincta: H 15.7, D 5.75 mm (H:D2.73) they are all in either Hawke’s Bay Nukumaruan (Murdoch & Suter 1906:286); largest syntype rocks of shallow facies (all relatively finely 42 AG Beu

sculptured specimens resembling the holotype I. striata after OIS11 by suppression of groups of I. venusta, with 815 spiral cords on the of spiral threads. They are also all old, chalky penultimate whorl), or in SWairarapa bathyal shells with a corroded surface, and more localities (coarsely sculptured specimens resem- material is required to determine whether they bling the syntypes of I. alticincta, with 56 constitute a third species. spiral cords on the penultimate whorl). Other The abundant specimens of Iredalula striata differences between Castlecliff specimens and at Wanganui (Castlecliffian, OIS25 11) all others also seem constant. Castlecliff shells seemed at first sight to provide a further are slightly wider (height: diameter2.22.5, example of a northern species or form that rarely as great as 2.7, compared with Recent extended its range southwards to Wanganui specimens: 2.83.2, rarely as little as 2.4), during interglacial periods (e.g. Beu 2004). consistently more evenly inflated (particularly Some aspects of its range at Wanganui, such the area between the outer margin of the as the absence from OIS21, probably result sutural ramp and the neck; the lateral outline from its preference for offshore siltstone habi- is almost flat on most Recent shells), the sutural tats rather than a true absence from the basin. ramp is not defined by the obvious margining However, it now seems likely that the history of ridge of Recent specimens, fine axial ridges on Iredalula is more complex. The finely sculp- the first teleoconch whorl are weak on I. striata tured species I. striata seems to have occurred but well-developed on Recent shells, and the widely in New Zealand early in Pliocene time, outer lip is significantly thinner on Castlecliff but then retreated to become an endemic Cook specimens than on Recent shells, most large Strait species by Pleistocene time*although the ones of which have a lightly thickened but very small amount of OpoitianWaipipian obvious, slightly developed external varix that material makes identification uncertain, and is scarcely discernible on Castlecliff fossils. In the older specimens have a slightly more my opinion Castlecliff fossils, and probably the obvious terminal varix than Castlecliff shells. few OpoitianWaipipian fossils from Westland I. striata became extinct at some time during or and N Hawke’s Bay, are the extinct species I. after OIS11 (there are no records from striata, whereas Recent specimens, Ohope Wanganui Basin younger than OIS11). Con- Beach Castlecliffian fossils, and Hawke’s currently, the more coarsely sculptured and BaySWairarapa Nukumaruan fossils are more flat-sided species I. alticincta occupied another species, for which the earlier name is the E North Island, including in deeper envir-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 I. alticincta. There is certainly no doubt that I. onments than I. striata is known from, and venusta intergrades completely with I. alticincta remains living there. The many Castlecliffian in sculptural prominence, the one character specimens from Ohope Beach, Whakatane (OIS that distinguishes these two forms. 15?) reach a smaller maximum size, are nar- The only Recent specimens I have seen that rower and flatter-sided, and have fewer, slightly diverge from this pattern are from the W North more prominent spiral cords than Castlecliff Island (NMNZ M. 52642, W of Rangitikei shells, and seem best identified as I. alticincta. River mouth, 82 m, three specimens; M.153427, Therefore, three species seem to be recog- WNW of Cape Egmont, 146 m, one; M.153431, nizable in Iredalula: I striata (Opoitian SW of Manukau Harbour, 502 m, one). These Castlecliffian; Castlecliffian at Wanganui specimens are smaller, shorter and wider than E only, last recorded in OIS11), I. alticincta North Island specimens and have fewer, lower, (NukumaruanRecent, E North Island), and wider, more widely spaced spiral cords that an apparently unnamed species living off the W fade out over the last half-whorl. They also North Island. This is a very preliminary have 35 fine spiral threads on each wide, conclusion, requiring at least the examination low spiral cord, and could have evolved from of radulae and anatomy of many Recent Marine Mollusca of isotope stages of the last 2 million years in New Zealand 43

specimens from E and W North Island to tell Fleming (1953:131, 140, 144, 146, 148,151, whether they are indeed all conspecific, as their 158, 174) recorded specimens of Antizafra sculptural variation seems haphazard. pisanopsis from Wanganui Nukumaruan rocks (Hautawa Shellbed, Nukumaru Limestone, Nukumaru Brown Sand, Mangamako Family Shellbed, undifferentiated Nukumaru group, Waipuru Shellbed, and Tewkesbury Forma- Genus Antizafra Finlay, 1926 tion). He also recorded A. speighti from Nuku- maru Limestone (Fleming 1953:140) and from Antizafra Finlay 1926b:421. Type species Okehu Shell Grit (p. 181; Castlecliffian, OIS 27; (by original designation): pisanopsis as ‘A. aff. speighti’, possibly reworked?). As far Hutton, 1885 (Fig. 7H), Pliocene, New as I am aware, all three species of Antizafra are Zealand. limited to Nukumaruan and possibly Manga- panian rocks, and the extinction of the genus occurred at about the end of Nukumaruan time. There are no records of A. and Remarks. Antizafra is a distinctive genus, with only the two of A. speighti listed here from one or two wide, low plaits on the base of the localities outside Hawke’s Bay. columella (Fig. 7H). Both A. pisanopsis (Hut- ton, 1885) (Fig. 7F, H, J) (originally spelled ‘pisaniopsis’, but corrected in the volume’s Family Muricidae corrigenda sheet to ‘pisanopsis’ [Beu & Maxwell Subfamily Tripterotyphinae 1990:325], although Hutton himself [1893:45] later spelled it ‘pisaniopsis’ again) and the Genus Ponderia Houart, 1986 markedly taller and narrower species A. can- cellaria (Hutton, 1885) have two low plaits Ponderia Houart 1986:88. Type species (by occupying the lower two-thirds of the columel- original designation): zealandica la, with a weak groove between them, whereas Hutton, 1873, Pleistocene and Recent, New A. speighti (Marwick, 1924) (Fig. 27H, I) has Zealand. only a single relatively narrow plait on the base of the columella. However, only the lower plait

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 seems to extend down the whole columella in A. Remarks. Houart (1986) demonstrated that ‘Ty- pisanopsis and A. cancellaria. All three also is’ zealandica Hutton, 1873, usually placed in have unusually graceful shapes and narrow, PterynotusorPrototyphisinrecentyears,belongs cancellate spiral and axial sculpture, although in a distinctive genus otherwise occurring in the sculpture is much weaker on A. speighti the subtropical to tropical Southwest Pacific, than on the others. A. pisanopsis and A. designatingT.zealandicaasthetypespeciesofthe cancellaria also differ from A. speighti in having new genus Ponderia. He included Ponderia abies a shorter and more mammillate protoconch Houart, 1986 (off New South Wales, in 154 (Fig. 7I, J), although it is equally smooth in all 164m)andPonderiacanalifera(G.B.SowerbyII, three species. Only the Nukumaruan and pos- 1841), an earlier name for the restricted Norfolk sibly Mangapanian species A. pisanopsis, A. IslandspecieslaternamedPterynotuszealandicus cancellaria and possibly A. speighti belong in iredalei by Fleming (1962a). Houart (1988, 1990) this genus; the other species previously referred has since added the species P. caledonica, P. here are transferred tentatively to Macrozafra magna and P. elephantina, all from New Caledo- Finlay, 1926, and this might well be a better nia and the Coral Sea. Houart & He´ ros position than Antizafra for A. speighti. (2008:471) included Ponderia in the subfamily 44 AG Beu

Tripterotyphinae, along with 103106 m (three); M.60467, Ranfurly Saddle, Pilsbry & Lowe, 1932, Jousseaume, off East Cape, 126128 m (one fragment); 1880, Prototyphis Ponder, 1972 and possibly M.60026, Rangatira Knoll, NW of White Island, Semityphis Martin, 1931. Whether this group 166169 m (one small dead); M.64788, Ranga- really deserves separation from subfamily tira Knoll, NW of White Island, Bay of Plenty, was not resolved by Barco et al.’s 164 m (one small dead). (2010) phylogeny, as they did not include any purported genera of Tripterotyphinae. Distribution. Castlecliffian to Recent; OIS13 (Tainui Shellbed), only, at Wanganui; living in Ponderia zealandica (Hutton, 1873) (Fig. 8AC) the Cook Strait region from Cape Farewell to offshore from Wanganui to the Rangitikei Typhis zealandica Hutton 1873b:2. River, and on Ranfurly Bank, East Cape. The Murex zealandica. Hector 1886:50, fig. 7, no. 4 two small dead specimens from Rangatira (not Murex zelandica Quoy & Gaimard, 1833, Knoll, Bay of Plenty, are possibly late Pleisto- type species of ). cene fossils, as have been reported from a Murex (Alipurpura) angasi. Suter 1913:404; nearby locality previously (37,00040,000 yrs, Suter 1914:2 (not Murex angasi Crosse, 1863). at Rungapapa Knoll; Beu 2004:155). (Poropteron) zealandicus. Finlay 1926b:419, fig. 56; Fleming 1961:1315. Dimensions. Typhis zealandica, holotype: H Pterynotus () zealandicus zealandi- 27.3, D 18.7 mm (spines slightly incomplete); cus. Fleming 1962a:115, pl. 1, figs. 15, 16; Dell & GS4013, R22/f6353, Tainui Shellbed at ‘the Fleming 1964:1, 2, fig.; Fleming 1966:57, figs. buttress’, Castlecliff: H 31.8, D 17.9 (spines 1262, 1263. incomplete); H 30.0, D 21.1 mm; Recent, Ponderia zealandica. Houart 1986:88, figs. 1, 1a; M.17977, N of Cape Farewell: H 22.7, D Maxwell 2009:245. 18.9 m; M.50080, SW of Wanganui: H 23.4, Pterotyphis (Ponderia) zealandicus. Beu & Max- D 15.5 mm (spines slightly incomplete). well 1990:358, pl. 48i; Spencer & Willan 1996:26; Spencer et al. 2009:211. Remarks. The very distinctive little muricid Ponderia zealandica is entirely restricted to Type material. Typhis zealandica, holotype Tainui Shellbed (OIS 13) in the Wanganui TM8294 (Fig. 8C), from Shakespeare Cliff, succession, and therefore is useful for identify-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Wanganui; undoubtedly from Tainui Shellbed, ing this shellbed in isolated outcrops. It has a the only formation this species occurs in. completely closed siphonal canal and shoulder (anal) spine, and three wing-like varices similar Other material examined. Castlecliffian: limited to those of Pterynotus. It also has a classic to Tainui Shellbed (OIS 13) at Wanganui; muriciform aperture, smooth and evenly oval moderately common throughout Wanganui Ba- with a continuous peristome and a well-raised sin. Recent: 10 lots in NMNZ: M.17977, due N of rim, a prominent shoulder angulation delimit- Cape Farewell, 8295 m (one good, closely ing a narrow, strongly concave sutural ramp, a similar to Wanganui fossils); M.90254, NE of narrow peribasal angulation formed by a Cape Farewell, 95102 m (one); M.50593, Sof relatively prominent spiral cord, and sculpture Waverley, Wanganui, 7782 m (two); M.50080, of a few other weak spiral cords crossing SW of Wanganui, 5864 m (two); M.50113, SW several low, narrow axial ridges in each inter- of Turakina River mouth, 5557 m, on bryozoa variceal interval, forming low nodules at their and shells (10 small dead); M.53311, W of intersections with the peripheral cord. The Rangitikei River mouth, 75 m (one small terminal varix closely resembles that of the dead); M.70662, Ranfurly Bank, off East Cape, similar-sized, less common but more widely Marine Mollusca of isotope stages of the last 2 million years in New Zealand 45 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 8 (AC) Ponderia zealandica (Hutton), type species of Ponderia Houart, 1986, Tainui Shellbed (Castlecliffian, OIS13), Wanganui; A,B, GS4013,R22/f6353, ‘‘the buttress’’, Castlecliff; height 30.0 mm; C, holotype, TM8294, ‘‘Shakespeare Cliff’’, Wanganui; height 27.3 mm. (D,M) Axymene aucklandicus (Smith), type species of Axymene Finlay, 1926, coarsely sculptured form, GS10963, R22/f6619A, Tainui Shellbed (Castlecliffian, OIS13), ‘‘the buttress’’, Castlecliff, SEM;D, whole specimen, height 13.6 mm; M, protoconch of D. (E,K) Xymenella pusilla (Suter), type species of Xymenella Finlay, 1926, RM4389, Recent, MV ‘‘Wahine’’ wreckage, Taranaki Street wharf, Wellington, SEM; E, whole specimen, height 8.9 mm; K, protoconch of E. (F,H) Xymene expansus (Hutton); F, GS15338, S22/f0164, Maoricrypta-dominated shellbed, Turakina Valley Road, early Castlecliffian, c. OIS43 41; height 24.1 mm; H, Hutton’s figured syntype, CMC M3301, ‘‘Wanganui’’, ex Drew collection; height 22.8 mm. (G,J,L) Xymene plebeius (Hutton), type species of Xymene Iredale, 1915; G, juvenile, SEM, GS14698, Q31/f0431, Kaikoura Peninsula high terrace (last interglacial, OIS 5e-5c?); L, protoconch of G; J, RM1950, Recent, Laingholme, Manukau Harbour, Auckland; height 11.5 mm. (I) Xymene drewi (Hutton), GS12711, V20/f045, Kaiwaka Limestone (late Nukumaruan), quarry near junction Turnbull and Waipunga Roads, above Esk Valley, N of Napier; height 14.7 mm. 46 AG Beu

stratigraphically distributed species Prototyphis Bay (late Nukumaruan; GS10849, V20/f8572) eos (Hutton, 1873) in being thin, flange-like and are much larger than Ponderia zealandica, but almost flat, bearing low, wavy lamellae in otherwise similar (H 40.0, D [incomplete] 21.4 conformity with three low radial frills that are mm; H [incomplete] 34.2, D 20.4 mm).They not well preserved on most specimens. How- possibly represent a large, unnamed species of ever, the long shoulder spine and closed canals Ponderia reaching about 50 mm in height. make Ponderia zealandica readily recognized. However, more prominent spiral cords around Dell & Fleming (1964) recorded a specimen the base and secondary spines on the face of the of Ponderia zealandica dredged in Cook Strait, terminal varix suggest a closer relationship with on a rough bryozoan bottom (M.17977, listed Prototyphis eos. As usual, only more material above). Nearby areas in Cook Strait have since can resolve the status of this form. yielded 17 further specimens. It appears that this species has occupied bryozoan-dominated substrates in the same area for at least the last Subfamily half-million years. Fleming (1962a) surmised that the Norfolk Island endemic species he Genus Xymene Iredale, 1915 called ‘Pterynotus’ zealandicus iredalei (Ponderia canalifera) indicated that the simi- Kalydon Hutton 1883b:576; Hutton 1884:222 lar form P. zealandica was among the warm- (Marshall 1995:495). Type species (by mono- water migrants that appeared briefly in New typy): Fusus plebeius Hutton, 1873, Pleistocene Zealand from the north during interglacial and Recent, New Zealand (suppressed, ICZN periods, but actually it is a temperate Cook Opinion 911 1970). Iredale (1915:471) pointed Strait and East Cape species in an otherwise out that Kalydon is a junior homonym of subtropical to tropical SW Pacific genus. The Calydon Thomson, 1864; this is still the case limitation of this species to Tainui Shellbed in under ICZN Article 58.5. the Wanganui succession presumably results from a narrow ecological tolerance, and the Xymene Iredale 1915:471. Type species (by origi- rare preservation of the offshore bryozoan- nal designation): Fusus plebeius Hutton, 1873, community in the Wanganui succession. PleistoceneandRecent,NewZealand.(Although The other tripterotyphine in the Wanganui essentially a replacement name for Kalydon fauna, Prototyphis eos (P. angasi powelli Hutton, 1883, Xymene was proposed specifically

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Fleming, 1962) occurs more widely in Waipuru as a new genus, with a type species designation.) Shellbed, Kaikokopu Shellbed (two collec- tions), Kupe Formation, and Lower Castlecliff Shellbed (Fleming 1953:151, 199, 203, 218), and Remarks. Xymene was used by Ponder (1972) as was collected from Pinnacle Sand by Mrs Jean an all-encompassing genus for most New Zeal- Boswell, reflecting its wider tolerance of a and Ocenebrinae, including several taxa that, variety of rocky shallow-water and offshore considering the level of genera currently recog- environments at present. Houart & He´ ros nized in the Muricidae, are clearly not conge- (2008:473) listed the species referred to Proto- neric. The complex is being revised by BA typhis, nearly all of which are limited to New Marshall and R Houart (pers. comm.), and Zealand. The exceptions are P. angasi (Crosse, most taxonomic revision will be covered by 1863), which occurs in southeastern Australia, them, but as an initial step the genera Axymene, and P. gracilis Houart & He´ ros, 2008 from Fiji. Xymenella and Zeatrophon of Finlay (1926b) Three fragmentary specimens from Devils are separated again here from Xymene. Xymene Elbow Mudstone at the top of Devils Elbow plebeius (Fig. 8G, J, L) and other species of hill, Highway 2 ca 35 km N of Napier, Hawke’s Xymene (sensu stricto) have a distinctive, very Marine Mollusca of isotope stages of the last 2 million years in New Zealand 47

short, smooth, conical, paucispiral protoconch obvious than that of Z. bonneti, and this passes of ca 2.2 whorls, with a low, wide, hemispherical gradually into the first teleoconch spiral cord, initiation with an in-rolled, slightly hetero- and gradually acquires teleoconch axial sculp- strophic appearance. The junction with the ture. This, therefore, seems to be a lecitho- teleoconch is rather obscure, suggesting that trophic protoconch, and the distinction from this is a lecithotrophic protoconch. In combina- that of Z. bonneti probably once again merely tion with the distinctive, wide spiral cords on the reflects the developmental difference. Com- teleoconch, the weak axial ridges producing low bined with the prominently lamellate, clathrate nodules at sculptural intersections, and the teleoconch sculpture of species assigned to simple aperture with a lightly flared outer lip, Zeatrophon, the genus is easily recognized with- this protoconch allows recognition of a separate in the New Zealand context. However, its genus Xymene. Axymene aucklandicus (Fig. relationships to such genera as Trophon, Tro- 8D,M), type species of Axymene Finlay, 1926, phonopsis and need to be defined. also has a paucispiral, smooth protoconch, but Xymenella pusilla (Suter, 1907) (Fig. 8E, K), it consists of little more than one whorl, and is type species of Xymenella Finlay, 1926, has a much more upright and dome-shaped than that similar protoconch to that of Xymene species, of Xymene (sensu stricto), with a large, swollen but a little taller and more obviously conical, of initiation and a deeply impressed suture, similar ca 2.5 whorls, joining the teleoconch abruptly to protoconchs of many . The after 23 prominent, curved axial ridges. The junction with the teleoconch is abrupt, as the teleoconch is distinctive because of its promi- shell acquires normal teleoconch axial ridges. nent, convex-crested spiral cords, small size but This is more certainly a lecithotrophic proto- relatively tall shape, and very prominent no- conch than that of Xymene, and indicates dules inside the outer lip. Not all the small New possible direct development. Combined with Zealand ‘trophonine’ taxa fit neatly into these the more elongate teleoconch shape, the more genera, however, and e.g. the generic position of strongly shouldered whorls, the more promi- ‘Zeatrophon’ huttoni (Murdoch, 1900) is not nent, sparser teleoconch sculpture, foliose on obvious (although this is a reasonably common some specimens, and the less flared aperture fossil at Castlecliff). These genera require clar- with nodules inside the outer lip, the much more ification and confirmation, including radular paucispiral protoconch allows distinction of the study. genus Axymene from Xymene. In contrast, the Some of the younger fossil species of

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 other similar genera, Zeatrophon and Xymenel- Xymene (sensu stricto) are reviewed here, as la, have taller protoconchs with more numerous they are useful in NukumaruanCastlecliffian whorls. Zeatrophon bonneti (Fig. 9D) has a tall, biostratigraphy, as pointed out by Fleming conical, multiwhorled protoconch of ca 3.2 (1955:521) when correlating the Castlecliffian whorls, sculptured entirely with fine, widely rocks at Ohope Beach, Whakatane. Because of spaced granules, more obvious on the first its intertidal environment, Xymene appears to whorl than lower down, and with a distinctive have a mostly late Neogene time range, but this spiral keel around the last whorl. The junction is a classic case of the poor preservation of the with the teleoconch is abrupt and obvious, and near-shore environment in the fossil record. development clearly is planktotrophic. Z. ambi- Named species included in Xymene (sensu guus (Fig. 9J has a similar tall, conical proto- stricto): conch, but smooth and a little shorter, of only Xymene coctor Marwick, 1948, Waipipian, ca two whorls, with a larger initiation; the Otahuhu well, Auckland (based on abundant, junction with the teleoconch is much less abrupt very small, apparently immature specimens than that of Z. bonneti. The protoconch devel- with fewer spiral cords and wider spiral inter- opes a keel on the last half-whorl, but less spaces than X. drewi; status unclear). 48 AG Beu

Xymene drewi (Hutton, 1882) (Fig. 8I), Man- lectotype; ‘paratype’ (valid lectotype of Mar- gapanianlate Nukumaruan (to ca OIS73-71?). wick 1924b:198) CMC M3223, also from Xymene expansus (Hutton, 1882) (Fig. 8F, H), ‘Wanganui’ (Bradshaw et al. 1992:63). Both late Nukumaruanearly Castlecliffian (ca OIS are probably from Nukumaru Brown Sand at 7319). Nukumaru Beach, although the horizon of Xymene moniliferus (Hutton, 1885), Opoitian?, CMC M3222 is uncertain. ‘Shepherd’s hut, Waipara’, N Canterbury The status of the type material of Cominella (WaiauanWaipipian according to Ponder’s drewi is confusing. Suter (1915:23) treated one (1972:478) synonymy; early forms of Xymene of the two syntypes present in CMC as the require further study, based on better material; ‘holotype’ and the other as the ‘paratype’ (that they are more coarsely sculptured than X. is, inadvertently designating them as lectotype drewi). and paralectotype under ICZN Article 74.6), Xymene plebeius (Hutton, 1873) (Fig. 8G, J, L), although he did not illustrate them, it is not Castlecliffian (OIS19/17)Recent. known what evidence he used to make this Xymene pulcherrimus (Suter, 1917), Waipipian, choice, and it is not possible to tell from Suter’s Starborough Creek, Awatere (possiblyX. text which specimen was designated as which; moniliferus and/or X. coctor). This name is a Suter’s (1915) text is a bald description, with no senior secondary homonym of ‘Xymene’ pul- qualifying statements, and there is no definite cherrimus (Finlay, 1930), proposed in Zeatro- evidence that Suter’s ‘holotype’ was among phon, but listed in Xymene by Ponder Hutton’s type specimens. Indeed, there is no (1972:468), Powell (1979:174) and Spencer & way of demonstrating that the labels are not Willan (1996:26). Returning to a position in now associated with the wrong specimens. Zeatrophon would remove the necessity for a Marwick (1924b:198), possibly on the basis of replacement name, so one is not proposed here, Suter’s now missing labels, stated that Suter, although it should have been proposed by apparently not realizing that the ‘holotype’ and Ponder (1972). ‘paratype’ represent two distinct genera and species, had chosen the larger specimen as Xymene drewi (Hutton, 1882) (Fig. 8F) ‘lectotype and wrongly called ‘‘holotype’’ by him’, although the ‘holotype’ does not agree Cominella drewi Hutton 1882:278 (Marshall with Hutton’s description, Hutton’s stated 1995:497); Hutton 1883a:410. dimensions, or Hutton’s (1893, pl. 6, fig. 13)

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 drewei (sic). Hutton 1893:42, pl. 6, fig. illustration, which is a drawing of the specimen 13. regarded by Suter as the ‘paratype’. Marwick Euthria drewi. Suter 1915:23 (in part). (1924b:198) noted that ‘the smaller one is what Xymene drewi. Marwick 1924b:198, pl. 17, fig. has generally been considered as Euthria drewi, 8; Finlay 1926b:424; Fleming 1966:232, fig. and is the specimen figured in the Macleay 1240. Memorial Volume [Hutton 1893]. It has there- Xymene plebeius expansus. Ponder 1972:477, fore a better claim to be considered the type of fig. 3.6, 7 only; Beu & Maxwell 1990:415 (in E. drewi than does the shell chosen by Suter’. part not Xymene expansus Hutton, 1882). The problem is that Suter’s lectotype is a poor, Xymene expansus. Maxwell 2009:246 (in part abraded specimen of Buccinulum cf. littori- not Xymene expansus Hutton, 1882). noides (Reeve, 1846), family Buccinidae (illu- strated by Fleming 1966, pl. 106, figs. 1289, Type material. Cominella drewi, ‘holotype’ (in- 1290, from a sketch by J Marwick; Fleming valid lectotype of Suter 1915:23) CMC M3222, (1966) identified it as ‘paralectotype of Comi- from ‘Wanganui’; this specimen (a specimen of nella drewi Hutton, wrongly called ‘‘Holotype’’ Buccinulum) is concluded here to be the para- by Suter’), whereas the specimen identified as Marine Mollusca of isotope stages of the last 2 million years in New Zealand 49

the paralectotype by Suter is the Xymene indicating that he thought it reworked from species that was illustrated by Hutton (1893, older rocks) in Wanganui Basin. It also occurs pl. 6, fig. 13) and consequently has borne this widely and commonly in Nukumaruan rocks of name consistently up until the revision by Hawke’s Bay, Wairarapa and North Canter- Ponder (1972) (who evidently did not realize bury, in shallow-water sandstone units with the complications underlying this name). Mar- common Zethalia and other shoreface taxa. wick’s solution was to regard Suter’s lectotype selection as invalid, although he did not state so Distribution. The earliest occurrence is unclear, explicitly. He did, however, illustrate the Xy- but Xymene drewi certainly occurs in early mene specimen (CMC M3223) and identified it Mangapanian rocks; it presumably evolved as the lectotype (Marwick 1924b, pl. 17, fig. 8). from still more coarsely sculptured species Acceptance of Suter’s lectotype designation would necessitate adopting a different (possibly such as X. pulcherrimus (X. coctor?) in new) name for this well-known species, and Waipipian rocks. It extends as young as Nuku- consigning the accepted name to the synonymy maru Brown Sand and Tewkesbury Formation of one or other of several Buccinulum species in the Wanganui succession, that is, latest (the specimen is so abraded that it is not clear Nukumaruan, early Pliocene. It is common in which species it belongs in). I therefore follow shoreface rocks throughout Wanganui Basin, Marwick (1924b) in treating Suter’s (1915) Hawke’s Bay, Wairarapa and North Canter- inadvertent lectotype designation as invalid, as bury. X. pulcherrimus (Suter 1917:38, pl. 12, fig. it did not select the specimen and species 8), from Starborough Formation (Waipipian), illustrated by Hutton (1893) and does not agree Starborough Creek, Awatere valley, differs little with the species described by Hutton (1882, from X. drewi in most characters other than its 1883a), and I regard the lectotype designation coarser spiral sculpture and may well intergrade by Marwick (1924b, pl. 17, fig. 8) as the valid gradually*that is, this might well be an anage- one. This case seems to be covered by ICZN netic series*and would then indicate a range Article 74.2: ‘If it is demonstrated that a from at least Waipipian to Nukumaruan. specimen designated as a lectotype was not a syntype, it loses its status as a lectotype’. Suter’s Dimensions. Hutton (1893:42) stated the dimen- inadvertently designated lectotype is not con- sions as ‘length 20 mm; breadth 11 mm’. Suter specific with the species described and illu- (1915:23) stated the dimensions as H 17, D

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 strated by Hutton, and so seems unlikely to 8 mm, probably referring to the Buccinulum have been a syntype, and under Article 74.2 specimen he wrongly selected as the ‘holotype’. cannot be considered to be the lectotype. Marwick (1924b:199) stated the dimensions of Other material examined. Not listed; abundant. the lectotype as H 13, D 7 mm, and pointed out Xymenedrewi was recorded by Fleming that Hutton usually seems to have provided the (1953:119, 140, 144, 146, 148, 151, 158, 163, dimensions of the largest specimen available, 174) from Mangapani Shellbed, Nukumaru regardless of the dimensions of his type Limestone, Nukumaru Brown Sand (along material; GS12711, V20/f045, Kaiwaka Lime- with ‘Xymene cf. expansus’), Mangamako stone (Nukumaruan), Kaiwaka South Road, Shellbed, undifferentiated shellbeds of Nuku- Hawke’s Bay: H 13.7, D 8.1 mm; H 14.7, D maru Group, Waipuru Shellbed, Tewkesbury 7.6 mm; H 17.1, D 8.9 mm; H 13.5, D 7.4 mm; Formation (along with ‘Xymene aff. expansus’), GS1984, V22/f6496, Maraekakaho Creek, S Pukekiwi Shell Sand (‘Xymene aff. drewi’, along side Ngaruroro River, central Hawke’s Bay, with ‘Xymene expansus, derived’) and Butler’s Nukumaruan: H 17.2, D 9.2 mm; H 16.9, D Shell Conglomerate (in square brackets, 9.2 mm. 50 AG Beu

Remarks. Xymene drewi is the common, wide- show two specimens of X. drewi (nos. 6, 7) and spread, typical Xymene species of Nukumaruan one of X. expansus (no. 8). This, then, is a rocks. As pointed out by Marwick (1924b:199), biostratigraphically useful lineage in shallow- ‘Generically this shell cannot be separated form water rocks, X. drewi occurring in Mangapanian Xymene plebeius (Hutton), from which it differs to late Nukumaruan rocks, X. expansus occur- in the non-carination of the whorls’, that is, ring in late Nukumaruan and early Castlecliffian X. drewi lacks the shoulder angulation, demar- rocks (Nukumaru Brown Sand to about cating the obvious sutural ramp, that is so Kupe Formation [OIS17] at Wanganui), and prominent on X. plebeius and X. expansus. X. plebeius occurring in late Castlecliffian to Although it is similar in size to X. plebeius, X. Recent faunas. The evolution from X. drewi to drewi also differs from X. plebeius in its wider X. expansus was not a simple one, and if these and shorter shape, thicker shell, and wider, species had an ancestor-descendent relationship, more closely spaced spiral cords. From X. X. drewi survived for a short period after the expansus, apparently the immediate descendent evolution of X. expansus. The evolutionary of X. drewi, it differs in its smaller size, thicker change from X. expansus to X. plebeius also shell, narrower shape, coarser spiral sculpture seems to have been gradual over a short but and lack of the characteristic expanded last appreciable period (OIS19 17), although it is whorl mid-section developed by adult X. ex- possible that specimens reworked from older pansus. X. expansus also has a markedly thinner units confuse this picture. outer lip, with less obvious internal ridges than X. drewi, has a more markedly left-directed Xymene expansus (Hutton, 1882) (Fig. 8F, G) anterior siphonal canal, and develops a shallow sinus in the outer lip, corresponding to the low Trophon expansos (sic) Hutton 1882:278 (Mar- carina at the shoulder angle, that is absent from shall 1995:497). X. drewi. X. plebeius also differs from X. drewi Trophon expansus. Hutton 1883a:410; Hutton in having the more markedly left-directed 1893:39, pl. 6, fig. 5. anterior siphonal canal of X. expansus, but Trophon () expansus. Cossmann has narrower, more clearly separated spiral 1903:52, pl. 3, fig. 6; Suter 1915:27. cords than either similar species. Xymene expansus. Finlay 1926b:424; Fleming Ponder (1972) was impressed by the apparent 1966:57; Beu & Maxwell 1990:359, pl. 48e; intergradation between Xymene drewi and X. Maxwell 2009:246 (in part not Xymene expansus

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 expansus, and synonymized these names. How- Hutton, 1882). ever, the collection he examined was an early, Xymene plebeius expansus. Ponder 1972:427, fig. mixed one, from both Nukumaru Brown Sand 3, no. 8 only (in part not Trophon expansus and Tewkesbury Formation, and more careful Hutton, 1882). localization of specimens has demonstrated that these are separate species, stratigraphically seg- Type material. Trophon expansus, holotype regated to some extent (X. drewi in Nukumaru CMC M-3301 (Fig. 8H), with two paratypes, Brown Sand, X. expansus in Tewkesbury For- M-3302, from ‘Wanganui’, ex Drew collection mation), and apparently occupying slightly dif- (Bradshaw et al. 1992:64) (these are probably ferent environments. X. expansus seems to have better considered as three syntypes). The type inhabited more strongly estuarine environments horizon is unknown, but seems likely to be than X. drewi, presumably part of a gradual Tewkesbury Formation. Marshall (1995:497) adoption of a more estuarine environment pointed out that the original spelling ‘expansos’ through the lineage, as X. plebeius occurs only (a typographical error?) is an incorrect Latin on hard substrates in estuaries at present. termination, and adoption of ‘expansus’is Ponder’s (1972, fig. 3, nos. 68) illustrations mandatory. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 51

Other material examined. Xymene expansus was X. drewi or X. plebeius, and in most larger recorded by Fleming (1953:144, 158, 163, 174, specimens of X. expansus the last whorl is 178, 181, 182, 188, 199, 203204) from Nuku- expanded markedly more widely than in either maru Brown Sand (as X. cf. expansus), Tewkes- similar species (presumably inspiring the species bury Formation (as X. aff. expansus), Pukekiwi name) and the outer lip is flared weakly. The Shell Sand (‘derived’), Butler’s Shell Conglom- low, closely spaced spiral cords and weak axial erate (OIS31), basal conglomerate member of ridges are also obviously less prominent than in Lower Okehu Siltstone (OIS 29; Mowhanau either X. drewi or X. plebeius. This is therefore Formation of Abbott & Carter 1999), Okehu quite a distinctive species, resembling an unu- Shell Grit (OIS 27), basal conglomerate member sually large, wide, weakly sculptured specimen of Upper Okehu Siltstone (OIS 27; Ostrea- of X. plebeius with a more sharply angled outer bed of Abbott & Carter 1999), Kaima- lip profile. Very large specimens of the ‘inferus tira Pumice Sand (OIS 25), Kaikokopu Shell form’ of X. plebeius are as tall as normal Grit (along with ‘Xymene aff. expansus aff. specimens of X. expansus, but differ from X. plebeius’; OIS19) and Kupe Formation (OIS expansus in their narrower shape and narrower 17; in square brackets, ‘derived’, along with X. sutural ramp. Most large specimens of X. cf. plebeius). It also occurs widely in late Nuku- plebeius are corroded so severely that their maruan shoreface sands in central and southern spiral cords are not visible. Hawke’s Bay. Fleming & Marwick (in Lillie 1953, table 22) recorded X. expansus from Xymene plebeius (Hutton, 1873) (Fig. 8G, J, L) GS2324, U23/f6376 and GS2457, U23/f8355, in Nukumaruan Upper Kumeroa Formation in Fusus plebeius Hutton 1873a:9; Hutton Dannevirke district, southern Hawke’s Bay, 1873b:3. and commented: ‘... Xymene expansus (Hutt.), Fusus inferus Hutton 1873a:9. a common lower Castlecliffian fossil which clathrata. EA Smith 1874:3, pl. 1, ranges down into the uppermost Nukumaruan fig. 21 (Beu 2010, Fig. 1N) (in part not rocks’ (Fleming & Marwick in Lillie 1953:139). Trichotropis clathrata GB Sowerby II August 1874, pl. 2, fig. 10). Distribution. Late Nukumaruanearly Castle- Kalydon plebeius. Hutton 1884:220. cliffian (ca OIS71 to ca OIS19/17), in Kalydon inferus. Hutton 1884:220. Wanganui Basin and Hawke’s Bay. Trophon (Kalydon) inferus. Suter 1913:414, pl.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 46, fig. 17. Dimensions. Trophon expansus, Hutton’s figured Trophon (Kalydon) plebejus (sic). Suter syntype: H 22.8, D 13.3 mm; GS15338, S22/ 1913:416, pl. 46, fig. 19. f0164, Maoricrypta-dominated shellbed, Tura- Xymene plebeius. Iredale 1915:471; Fleming kina Valley Road, near Mangara Stream, early 1966:57; Powell 1979:173, pl. 37, fig. 16; Castlecliffian: H 24.9, D 14.5 mm; H 22.0, D Spencer & Willan 1996:26; Spencer et al. 13.2 mm; H 23.5, D 13.7 mm; H 21.2, D 11.4 mm. 2009:210; Maxwell 2009:246. Xymene inferus. Iredale 1915:471; Finlay Remarks. Xymene expansus is characterized by 1926b:424. its larger, wider teleoconch than either X. drewi Xymene plebejus. Finlay 1926b:424. or X. plebeius, and by its thinner-shelled, light- Xymene plebeius plebeius. Ponder 1972:475, fig. er-weight appearance produced by the lower, 1, no. 1; fig. 2, no. 1; fig. 3, nos. 15; Beu & wider spiral cords and thinner outer lip than in Maxwell 1990:415. either similar species. The outer lip is also angled sharply outwards at the base of Type material. Fusus plebeius, lectotype (of the sutural ramp more obviously than in either Ponder 1972, fig. 3) NMNZ M.80, with two 52 AG Beu

paralectotypes M.1820 (Marshall 1996:24), Dimensions. GS3513, Henley, Taieri Plains, from ‘New Zealand’; no locality was stated by Holocene: H 16.4, D 9.1 mm; H 16.5, D 9.3 Hutton (1873a:9). According to Ponder mm; H 12.7, D 6.9 mm; H 10.8, D 5.7 mm; H (1972:475) the type material of Fusus inferus is 10.9, D 5.8 mm; RM1950, Recent, Langholme, lost; Hutton (1973a:10) stated the locality as Manukau Harbour, Auckland: H 14.9, D ‘Stewart’s Island’, southern New Zealand, and 6.9 mm; H 11.2, D 5.5 mm; H 11.6, D similar unusually large, severely corroded speci- 5.9 mm; ‘inferus’ form, Stewart I., Suter collec- mens have been collected recently on tidal flats tion no. 4021: H 26.6, D 12.7 mm. near the head of Patterson Inlet. In order to associate the name unambiguously with a Remarks. Comparisons with Xymene drewi and species, so that it cannot threaten the stability X. expansus are provided above. Xymene of other names, the lectotype of Fusus plebeius plebeius is easily distinguished from all earlier Hutton, 1873, NMNZ M.80, is here designated species by its narrower shape and the more the neotype of Fusus inferus Hutton, 1873. The prominent appearance of its sculpture, pro- synonymy list by Ponder (1972:475) provided duced by the narrower, more widely separated the first reviser’s action selecting the name spiral cords. It also has a sutural ramp defined Fusus plebeius as the valid one of the two clearly by an obvious shoulder angulation, names proposed by Hutton (1873a), Fusus as in the larger and thinner-shelled species X. plebeius and Fusus inferus. expansus, but the angulation is absent from or much weaker on all earlier species. Other material examined. Xymene plebeius was Genus Axymene Finlay, 1926 recorded by Fleming (1953:199, 203204, 207, 214, 218, 226, 230, 242, 245, 261, 273) from Axymene Finlay 1926b:424. Type species (by Kaikokopu Shell Grit (as ‘Xymene aff. expansus original designation): Axymene turbator Finlay, aff. plebeius’; OIS19), Kupe Formation (as 1926 (Euthria aucklandica E. A. Smith, 1902), ‘X. cf. plebeius’) and Upper Kai-Iwi Siltstone central and southern New Zealand, Mangapa- (OIS 17), Seafield Sand and Lower Castlecliff nianRecent (Ponder 1972:494495, fig. 8). Shellbed (as ‘X. cf. plebeius’; OIS15), Pinnacle Sand and Tainui Shellbed (OIS 13), Mosstown Axymene aucklandicus (E. A. Smith, 1902) (Fig. Sand (OIS 11?), Landguard Sand and Brunswick 8D, M) Formation (OIS9), Denby Shellbed,at Hawera, Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 and Waipuna Conglomerate (OIS7) in Wanga- Euthria aucklandica E. A. Smith 1902:203, pl. nui Basin. It is not a common fossil elsewhere, 24, figs. 12, 13. but occurs in some young Pleistocene terrace Trophon (Kalydon) columnaris Suter 1908:178, cover beds throughout New Zealand; Holocene, pl. 7, fig. 1 (?junior primary homonym of 1 m below surface, 150 m N of Henley railway Trophon columnarius Hedley & May, Sept. station, Taieri Plains, Otago (GS3513, H45/ 1908). f9479, abundant), and other Holocene fossils Trophon (Kalydon) erectus Suter 1909a:27 throughout New Zealand. (replacement name for T. columnaris Suter, 1908); Suter 1913:414, pl. 19, fig. 7. Distribution. Mid-Castlecliffian (ca OIS19 17) Trophon (Kalydon) aucklandicus Suter to Recent; most fossil records are from Wan- 1909a:27; Suter 1909b:254; Suter 1913:411, pl. ganui Basin. Recent specimens live throughout 46, fig. 18. New Zealand on stones and shells*that is, any ?Buccinulum pertinax. Finlay 1926b:422 (not available hard substrate*lying on soft sub- Euthria lineata pertinax von Martens, 1878). strates in estuaries. Axymene erectus. Finlay 1926b:424. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 53

Axymene turbator Finlay 1926b:426, pl. 23, figs. Shellbed at ‘the buttress’ (35); GS4022 re- 127, 128. collection, R22/f6360, head of ‘the pinnacles’ Axymene traversi aucklandica. Fleming gully (one). Recorded also by Fleming 1951:137, pl. 18, fig. 4; Powell 1955:105. (1953:230) from Tainui Shellbed in GS4031, Axymene traversi erectus. Fleming 1951:137; R22/f6369; GS4097 and GS4102, both allo- Powell 1955:104. cated R22/f6516; all from the Castlecliff coastal Xymene aucklandicus. Ponder 1972:494, figs. section. 1.7, 2.16, 17, 8.18; Powell 1979:175; Beu & Fleming (1953:203204, 207, 218, 226, 239) Maxwell 1990:415; Spencer & Willan 1996:26; also recorded ‘Aymene n. sp.’ from two mem- Spencer et al. 2009:210; Maxwell 2009:246. bers of Kupe Formation at Castlecliff, and from Upper Kai-Iwi Siltstone, Lower Castle- cliff Shellbed, Pinnacle Sand, and Upper Type material. Euthria aucklandica, lectotype Castlecliff Shellbed, but examination of these designated by Fleming (1951:137, caption to pl. specimens has shown that his identifications 18, fig. 4), BMNH 1902.5.16.18, with three were based on more normal specimens of either paralectotypes BMNH 1902.5.16.19-21, all A. aucklandicus or A. traversi (Hutton, 1873). from Auckland Islands, 20 m. Trophon colum- Fleming (1953) also listed ‘Xymenella n. sp.’ naris Suter, lectotype TM957, with 10 para- from several units, but again these are based on lectotypes, TM958-967, designated by Boreham only slightly unusually sculptured specimens of (1959:47), from near the Bounty Islands, 100 m. X. pusilla (Suter, 1907). A range of other Boreham expressly designated the lectotype for MangapanianHaweran localities for A. auck- Axymene traversi erectus (Suter, 1909), but as landicus was listed by Ponder (1972:495). Trophon erectus was a replacement name for the preoccupied Trophon columnaris Suter, Distribution. Axymene aucklandicus occurs 1908, Suter’s type material was identified only widely in New Zealand in rocks of Mangapa- as ‘Trophon erectus’ by Suter (the original label nian to Castlecliffian age, at a few Haweran evidently having been discarded), Boreham did localities, and in the Recent fauna throughout not mention the replaced name, and there is no central and southern New Zealand. The more separate type material for T. columnaris, Bore- strongly spirally sculptured, axially foliate form ham’s designation applies equally to the re- described here is restricted to Tainui Shellbed placed and replacement names, in the spirit of (OIS13), as far as I am aware. Most material is

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 ICZN Article 72.7. Axymene turbator, holotype from the Castlecliff coastal section, because of AIM AK70088, with two paratypes AK72906, its excellent exposure. AK72907 (not seen). Finlay (1926b:426) pub- lished the type locality as ‘Dunedin Harbour, Dimensions. Lectotype of Euthria aucklandica: under stones at low tide’, but the type material H 8.6, D 4.1 mm; paralectotypes: H 9.3, D is labelled ‘Taieri Beach, 8 km Sof Taieri 4.2 mm; H 8.8, D 4.0 mm; H 7.4, D 3.2 mm; River’, SOtago coast (Owen Lee, AIM pers. GS4013, Tainui Shellbed at ‘the buttress’, comm. Nov. 2007). Castlecliff: H 14.2, D 6.5 mm (TM5523); H 18.0, D 7.5 mm; H 15.6, D 6.7 mm; H 14.8, D Other material examined. Tainui Shellbed 6.2 mm. strongly sculptured form: GS10963, R22/ f6619A, re-collection of GS4013, Tainui Remarks. Ponder (1972:494, figs. 1.7, 2.16, 17, Shellbed (OIS 13) at ‘the buttress’, Castlecliff 8.18) described the variation and range of coastal section, Wanganui (nine, including Axymene aucklandicus (under Xymene). His TM5523, specimen illustrated by Ponder description of the large range of variation [1972, fig. 8.7]); GS4013, R22/f6353, Tainui exhibited by both A. aucklandicus and the other 54 AG Beu

common, widespread species, A. traversi, allows end-members of other populations. I therefore recognition that the many forms identified by conclude that Ponder (1972:495) was correct to Fleming (1953) as ‘Axymene n. sp.’ are part of rank this form as part of the variation of A. the variation of one or the other of these aucklandicus rather than as a distinct species. species. Ponder also commented that ‘A fossil Although this form seems to be limited to population from the Tainui Shellbed at Castle- Tainui Shellbed, and will have some minor cliff, Wanganui, has a stronger shoulder and utility in biostratigraphy, its reliability is very more distinct secondary axial foliation than low. most Recent shells (fig. 8:7). Other Castleclif- Typical specimens of Axymene traversi fian shells resemble Recent specimens so that occur in small numbers in Tainui Shellbed the Tainui form is here regarded as merely an syntopically with A. aucklandicus, and are aberrant population of aucklandicus. Somewhat immediately separable by retaining their coral- similar shells are found in deep water off the line algal encrustation, as well as by their much Otago coast’ (Ponder 1972:495). As this dis- weaker spiral sculpture, sharper peripheral tinctive form is limited to Tainui Shellbed, I nodules, and taller protoconch with more examined it to assess its utility in biostratigra- numerous whorls than in A. aucklandicus. phy. The prominent, narrow, well-separated They also bear a prominent spiral carina on spiral cords distinguish Tainui Shellbed speci- the last protoconch whorl that is absent from mens from most other specimens of A. auck- A. aucklandicus. Similar-sized specimens of landicus. However, they are not as distinctive as ‘Zeatrophon’(?) huttoni (Murdoch, 1900) also the more clearly defined, wider, almost smooth occur in small numbers syntopically with both sutural ramp having only weak spiral sculpture, A. traversi and A. aucklandicus. the narrowly rounded peripheral angulation, and the well-defined axial foliation, which Genus Zeatrophon Finlay, 1926 separate the Tainui Shellbed form from all other specimens of A. aucklandicus. The most Zeatrophon Finlay 1926b:424. Type species (by extremely coarsely sculptured specimens of original designation): Fusus ambiguus Philippi, A. aucklandicus I have examined from other 1844). localities (RM19, Lyall Bay, Wellington, one; GS12290, E38/f015, boulders in Whiskey Remarks. The genus Zeatrophon is used here Creek, N coast of Cascade Point, SWestland; without prejudice, as a group to contain the

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Castlecliffian, ca OIS25; three) differ from following species deserving comment in this Tainui Shellbed specimens in their smooth, paper, and requiring removal from Xymene. lightly polished surface, without any axial The correct generic position must await the foliation, and in having a consistently narrower revision by Marshall & Houart (pers. comm.). sutural ramp and less obvious peripheral angu- Zeatrophon bonneti (Cossmann, 1903) has a lation, despite their prominent, narrow, widely distinctive, tall and relatively narrow, granulose spaced spiral cords, similar to those of Tainui protoconch of ca 3.2 whorls with a prominent Shellbed specimens. The smooth, paucispiral, spiral keel on the last whorl (Fig. 29F), whereas dome-shaped protoconch of Tainui Shellbed the type species, Z. ambiguus (Philippi, 1844), specimens is also indistinguishable from that of has shorter, smooth protoconch of ca 2.5 other specimens of A. aucklandicus. Also, a few whorls (Fig. 29E,G) with a less obvious keel. large specimens in the Tainui Shellbed popula- However, it appears that the protoconch of Z. tion develop a more rounded periphery, less bonneti reflects planktotrophic development, sharply defined sutural ramp and weaker whereas that of Z. ambiguus reflects lecitho- spiral cords over the last half-whorl, and trophic development, probably with at least a closely resemble the more coarsely sculptured brief demersal larval life. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 55

Zeatrophon bonneti (Cossmann, 1903) (Fig. excellent preservation, completely free of ma- 9AD, H) trix, suggest it is possibly from Landguard Sand (OIS9) at Landguard Bluff, but the sculpture Trophon stangeri. Hutton 1893:38 (not Trophon suggests it is more probably a Nukumaruan stangeri Gray, 1843). specimen from a locality such as Nukumaru Trophon () bonneti Cossmann Beach, or from Waipuru Shellbed or a similar 1903:200, pl. 3, fig. 7. stratigraphic horizon in the Rangitikei valley. Zeatrophon bonneti. Finlay 1926b:424; Fleming The two non-type specimens bear Cossmann’s 1966:57. label reading: ‘9670 N. Z. Wanganui Trophon Xymene bonneti bonneti. Ponder 1972:486, fig. bonneti Cossm. dedit. M. Bonnet’. They are 6: 14; Beu & Maxwell 1990:359, pl. 48m. more typical grey-stained, coarsely cancellate Xymene bonneti. Maxwell 2009:246. specimens, almost certainly from Castlecliffian Not Trophon (Trophonopsis) bonneti. Suter rocks of the Wanganui coastal section, and 1913:417, pl. 45, fig. 23 (incorrectly synony- probably from Tainui Shellbed, but it is im- mized with T. ambiguus pumila Suter, 1899, possible to be sure of the provenance of any of Recent). Bonnet’s specimens. Recognition (below) that Z. lassus Marwick, 1928 and Z. murrayi Flem- ing, 1943 intergrade with Z. bonneti removes Type material. The holotype of Trophon (Tro- some of the difficulties presented by the holo- phonopsis) bonneti (Cossmann 1903:200, pl. 3, type of Z. bonneti, but it is unfortunate that the fig. 7) has been recognized recently in MNHN, holotype is so unlike the usual Castlecliff De´ partement Sciences de la Terre, and regis- specimens. tered as J09323 (Fig. 9A,B; photos kindly sent by Didier Merle MNHN pers. comm. 29 June Other material examined. Nukumaruan: ‘Wai- 2009). It is clear that this is indeed the specimen kopiro’, Suter collection (two); hillside above illustrated by Cossmann (1903, pl. 3, fig. 7), Glenstrae Station, Oaro, Marlborough although the sculptural details have not been (GS10337, O32/f8843; four); Devils Elbow appreciated from Cossmann’s small figure. It is Mudstone, top of Devils Elbow hill, Highway from the collection of Bonnet, and is from 2, N of Napier (GS10849, V20/f8572; 60; ‘Wanganui’, New Zealand, presumably origin- GS2220, V20/f8462, two); Kaiwaka Limestone, ally sent by a New Zealand collector such as R Kaiwaka South Road, ascent from Esk Valley,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Murdoch. There is no doubt that Cossmann N of Napier (GS12712, V20/f047A, five); originally had only one specimen, so two others Mairau Mudstone, walkway W of Roro-o- remaining in the Cossmann collection, De´ par- kuri, Bayview, Napier (GS12386, V21/f05, tement Sciences de la Terre, MNHN (J08923) one); Mairau Mudstone, ‘the watchman’ hill, (Jean-Michel Pacaud MNHN pers. comm. Ahuriri lagoon, Napier (GS11465, V21/f6015, August 2007) presumably were added from two); Waipunga Road, 1.5 km Sof junction Bonnet’s collection at a later date, and have with Kaiwaka Road, W of Devils Elbow no type status. The holotype raises a slight (GS1983, V20/f8459, one); Maharekeke Mud- problem, as it is not a typical coarsely cancel- stone, Maharakeke Road, W foot of Pukeora late, grey-stained specimen from Castlecliff, but Hill, Waipukurau (GS10857, U23/f7049, four); has a brownish yellow stain, a weaker periph- Waipuru Shellbed, Mangawhero valley eral spiral cord than typical specimens, and (GS4360, S22/f6485, two); shellbed 150 m narrow, secondary spiral cords are present in above Waipuru Shellbed, upstream from Li- all spiral interspaces. The sculpture is similar to vingstone Bridge, Rangitikei River (GS3098, that of specimens of the lassus and murrayi T22/f8494, two). Castlecliffian: Ototoka Silt- forms of Z. bonneti. The yellow stain and stone (OIS31), Ototoka beach, Wanganui 56 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 9 (AD,H) Zeatrophon bonneti (Cossmann), Castlecliff, Wanganui; A, B, Cossmann’s illustrated syntype, MNHN De´ pt Science de la Terre, J09323, ‘‘Wanganui, dedit. M. Bonnet’’; height 25 mm; D. Merle photo, MNHN; C, D, H, GS4102, R22/f6516, 3 specimens, Tainui Shellbed (Castlecliffian, OIS 13), ‘‘the buttress’’, Castlecliff; C, height 22.5 mm; D, protoconch, SEM; H, height 19.6 mm. (E,G) Bedeva bartrumi (Marwick), type species of Otahua Marwick, 1948, paratype, TM8301, Otahuhu well, Auckland, Waipipian, ex Laws collection; height 43.2 mm. (F,I) (Crosse), WM12047, Dunwich, Stradbroke Island, Queensland, Australia; height 22.2 mm. (J,K) Zeatrophon ambiguus (Philippi), type species of Zeatrophon Finlay, 1926, GS4121, R22/f6521, Kupe Formation (OIS 17), Castlecliff coast, SEM; J, protoconch of K; K, small specimen, height 13.2 mm. (L) Agnewia kempae Powell, protoconch of specimen in Fig. 10J, GS10911, Y14/f7505C, Te Piki Member (Haweran, OIS7), Te Piki, near East Cape; SEM. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 57

(GS4107, R22/f6430, one); Kaimatira Pumice occurrence at Te Piki, near East Cape (OIS Sand (OIS 25), mouth of Kai-Iwi Stream, 7). Specimens are rare other than at Castlecliff, Castlecliff (GS4068, R22/f6399, two); as last, Wanganui, but it is common in Devils Elbow Mowhanau Stream between bridge and mouth, Mudstone at Devils Elbow, Napier-Wairoa Kai-Iwi (GS4163, R22/f6459, four); as last, Highway, and a few specimens occur in other Kai-Iwi Beach Road, Wanganui (GS4064, Nukumaruan rocks in central Hawke’s Bay. R22/f6395, three); as last, Tuckers pumice However, this form intergrades with specimens workings, Brunswick Road, W of Wanganui identified as Z. lassus, and it appears that this is (GS4148, R22/f7414, one); Lower Kai-Iwi Silt- another single, anagenetically evolving species stone (OIS23), Castlecliff (GS4061,R22/f6392, (see below). The earlier time range is, therefore, 25); Omapu Shellbed (OIS 23), Castlecliff poorly known, but Z. bonneti in the broad sense (GS4060, R22/f6391, 14); gastropod shellbed, probably extends from Opoitian to OIS7. Kupe Formation (OIS17), Castlecliff (GS4121, R22/f6521, five); Kupe Fm, Castlecliff Dimensions. Holotype, MNHN J09323, ap- (GS4041, R22/f6373, one); Kupe Formation, proximate dimensions (determined from photo Castlecliff (GS4050, R22/f6381, one); bivalve scale): H 25, D 12.5 mm; non-types, MNHN shellbed, Kupe Formation, Castlecliff (GS4045, J08923: H 19.4, D 8.9 mm; H 15.8, D 7.4 mm; R22/f6376, one); gastropod shellbed, Kupe GS4027, Seafield Sand, Castlecliff: H 29.0, D Formation, Castlecliff (GS4047, R22/f6378, 15.1 mm; H 21.3, D 10.0 mm; GS4102, Tainui one; GS4121, R22/f6521, one); undifferentiated Shellbed, Castlecliff: H 21.4, D 10.9 mm; H Kai-Iwi Group, ridge off end of No. 3 Line, E 22.4, D 9.8 mm; GS5833, Te Piki, East Cape: H of Wanganui (GS4361, S22/f6486, three); 35.7, D 19.3 mm. uppermost sand member, Seafield Sand (OIS 15), Castlecliff (GS4027, R22/f6365, two); Low- Remarks. The small, beautifully and elabo- er Castlecliff Shellbed (OIS 15), ‘the pinnacles’, rately sculptured species Zeatrophon bonneti, Castlecliff (GS4104 re-collection, R22/f6518, with high, narrow, fenestrate axial and spiral three); Lower Castlecliff Shellbed, Castlecliff ridges, is particularly distinctive, and has (GS4140, R22/f6450, two); Tainui Shellbed a biostratigraphically useful extinction time. (OIS13) at ‘the buttress’, Castlecliff (GS4102, Typical Castlecliff specimens have narrow, R22/f6516, eight); Tainui Shellbed, N side well-raised spiral cords with few interstitial Waimutu Road, Turakina valley (GS15140, cords, crossed by narrow axial ridges. Most

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 S23/f084, one). Haweran: GS4003, R22/f7394, specimens have no interstitial cords at all, but Landguard Sand (OIS 9), Landguard Bluff, some specimens have one interstitial cord in Wanganui (one). A particularly large, fine one or a few interspaces. The upper of the two specimen and three smaller ones from Te Piki, spiral cords on spire whorls is as prominent as 6 km E of Whangaparaoa, near East Cape (OIS or even more prominent than the lower cord, 7) are similar to Z. bonneti, but have inter- whereas on the Recent apparently descendent mediate, secondary spiral cords and seem to species X. caudatinus (Finlay, 1930) the upper intergrade with Z. caudatinus, although they spiral cord is less prominent than the lower one, have a prominent shoulder angle (GS5833, producing a more ‘rounded’ effect to the Y14/f505A, two; GS15443, Y14/f505B, two). margin of the sutural ramp, and interstitial spiral cords are more numerous (Ponder 1972). Distribution. The first occurrence of typical Comparison of a range of specimens specimens of Zeatrophon bonneti is relatively demonstrated that Ponder (1972) correctly late in Nukumaruan time (approximately OIS described the gradual development of the ex- 86?) and the last occurrence I am aware of is in treme late NukumaruanCastlecliffian form of OIS9 (Landguard Sand),with a probable Zeatrophon bonneti from Z. lassus (Marwick 58 AG Beu

1928:487, fig. 143) throughout middle to late intermediate between these named forms. In this Pliocene time, encompassing the type material of case, the elegans form seems merely to be an Z. murrayae (Fleming 1943:205, pl. 31, fig. 30) uncommon sculptural variant of Z. ambiguus. (Nukumaruan, GS3093, S22/f9489, Hunterville- The nominal taxa Z. elegans and Z. murrayi Turakina Valley Road, Wanganui; recorded also clearly are not restricted Nukumaruan species of by Fleming from several localities in Dannevirke biostratigraphical utility, and a distinction be- Subdivision, S Hawke’s Bay; holotype TM8611, tween an earlier species Z. lassus (Opoi- with four paratypes, TM8612-15; eight para- tianNukumaruan) and a later species Z. types from GS2312, U22/f9488, Ashcott Road, bonneti (late NukumaruanCastlecliffian) is of W of Waipukurau, TM8616-23). Most Nuku- very low reliability. maruan populations from Hawke’s Bay (com- In contrast, another synonymy proposed by mon at Ashcott Road and in Devils Elbow Ponder (1972) that I cannot support is of Mudstone at the top of Devils Elbow hill) have Zeatrophon mutabilis Marwick (1928:488, figs. numerous interstitial secondary spiral cords, 144, 146, 148) with Z. ambiguus. The many particularly on the last whorl, that are reduced specimens of Z. mutabilis now available from on or absent from Castlecliff shells, and a more the type locality, Titirangi Sand at Titirangi flared outer lip than typical Castlecliff shells, and Point, Te Whanga Lagoon, Chatham Island are indistinguishable from the holotype and the (early Nukumaruan), differ from all mainland few other specimens seen from the type New Zealand specimens in their consistently formation of Z. lassus, Whenuataru Tuff more predominant sculpture of wide, flat spiral (WaipipianMangapanian), Pitt Island, Cha- cords and their consistent short, wide shape, tham Islands (holotype TM8624, Whenuataru with a shorter siphonal canal than in Z. Tuff, Taruwhenua Peninsula, Pitt I). The proto- ambiguus. Contrary to Ponder’s (1972:486) conch differences described by Fleming statement, they are separable at all ages. There (1943:205) are slight and seem to result from is no doubt that Z. mutabilis is a distinctive damage; protoconchs are essentially identical on species restricted to Titirangi Sand (early specimens I have examined that were assigned to Nukumaruan). It also formerly occurred com- Z. bonneti, Z. lassus and Z. murrayae. Z. monly in the disused Moutapu quarry at the E murrayae is here regarded as a synonym of Z. end of Karewa Peninsula (GS12170, CH/f11; lassus. However, Z. lassus and Z. bonneti inter- GS13190, CH/f261; listed as ‘Xymene mutabilis’ grade very gradually with time, that is, they are in Campbell et al. 1994:158). More signifi-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 part of an anagenetically changing series, and it is cantly, abraded, brown-stained specimens of probably best to regard them as conspecific. Z. mutabilis occur also in small numbers on the However, further study of pre-Nukumaruan beach N of Owenga, SE Chatham Island, specimens is warranted before this synonymy is demonstrating that one reason for the distinc- accepted and the time range is clear. tive nature of the ‘Owenga shell bed’ fauna Comparison of specimens of Zeatrophon (Dell 1960:154), that is, the fauna cast ashore ambiguus (Fig. 9J, K) and Z. elegans Fleming on the beach north of Owenga, is that it (1943:205, pl. 31, fig. 32; Nukumaruan, holotype includes a contribution from Titirangi Sand. TM8625, GS2687, T24/f7415, Kumeroa, E of Although no outcrops of Titirangi Sand have Woodville, SHawke’s Bay) demonstrated that been observed along this coast, the formation is Ponder (1972) also was correct to regard Z. now known to be widespread, occurring as far elegans as falling within the range of variation of W as in the cliff-face above the beach N of Red Z. ambiguus. Specimens from Mangapani Bluff, Petre Bay, N of Waitangi, where it Shellbed (GS4227, R21/f8478, Mangapunipuni contains an ocean beach fauna dominated by Stream, Waitotara Valley; Mangapanian) illu- subtriangulata (Wood, 1828), and on strated by Ponder (1972, figs. 5.4, 5.5) clearly are the shore of Lake Te Wapu, Sof Kaingaroa Marine Mollusca of isotope stages of the last 2 million years in New Zealand 59

Harbour, NE Chatham Island (discovered in well as in its distinctive foliose axial sculpture, 2009 by Dr Kat Holt, Massey University). and seems better regarded as a genus distinct Erosion of former Titirangi Sand outcrops from . I also preferred to follow Kool (apparently now overgrown) along the coast (1993) and retained Bedeva and Haustrum in N of Owenga would explain the occurrence of Subfamily Ocenebrinae. A further recent con- this and other taxa at Owenga that have not tribution to the molecular phylogeny of the been found elsewhere in the modern beach Muricidae (Claremont et al. 2008) retained fauna, such as Cominella ellisoni consobrina the same genera in the and Oceneb- Powell (1933:206, pl. 36, fig. 8), a probable rinae as were recognized by Kool (1993) and, synonym of the Titirangi Sand species C. more recently, Barco et al. (2010) established ellisoni Marwick, 1928. Haustrinae firmly as a muricid subfamily, on the basis of molecular phylogeny, and showed that Lepsiella and Lepseithais can be accepted Subfamily Haustrinae Tan, 2003 as synonyms of Haustrum (although, unfortu- nately, they did not include Bedeva in their Genus Bedeva Iredale, 1924 analysis). Its highly distinctive shell characters demonstrate that Bedeva should be retained as Bedeva Iredale 1924:273. Type species (by a genus separate from Haustrum at present. original designation): Trophon hanleyi Angas, 1867, Recent, southeastern Australia. Bedeva bartrumi (Marwick, 1948) (Fig. 9E, G)

Otahua Marwick 1948:32. Type species (by Otahua bartrumi Marwick 1948:32, pl. 7, figs. 5, original designation): Otahua bartrumi Mar- 6, 12, 13; Fleming 1966:57. wick, 1948, Pliocene (and Pleistocene?), New Bedeva bartrumi. Beu & Maxwell 1990:295, pl. Zealand (new synonym). 37h; Maxwell 2009:245.

Type material. Otahua bartrumi holotype AIM Remarks. Tan (2003) provided a detailed com- AK71074 (not seen), five paratypes TM8297- parison of the anatomical, radular and oper- TM8301, all from GS3528, R11/f7014, Otahu- cular characters of Australian and New hu well, Auckland, Waipipian (Pliocene). Zealand species previously classified in Bedeva,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Haustrum Perry, 1811, Lepsiella Iredale, 1912, Other material examined. Waipipian: Otahuhu and Lepsithais Finlay, 1928 and, in view of their well, Auckland, Waipipian (GS3528, R11/ close similarity, proposed that they be placed f7014, six specimens); Greenwood Formation together in a new Subfamily Haustrinae*apart (Waipipian), Lower Waipara Gorge, North from ‘Lepsiella’ reticulata (Quoy & Gaimard, Canterbury (GS4997, N34/f6192, two 1832) and ‘L’. botanica (Hedley, 1918), which abraded). Nukumaruan: shellbeds above and Tan transferred to Phycothais Tan, 2003 in below Vinegar Hill Tephra (OIS61; Pillans Subfamily Rapaninae. In contrast, I pointed et al. 2005:79, figs. 5A, 11), Tewkesbury For- out (Beu 2004:214216) that species referred to mation (late Nukumaruan), Brunswick Road, Lepsiella and Lepsithais (other than L. botanica E side Kai Iwi Valley, Wanganui (GS15348, and L. reticulata) are indistinguishable from R22/f6542A, one, severely abraded). Castleclif- Haustrum in most characters, and are better fian: shellbed low in Maraetotara Formation, transferred there. Bedeva differs strongly from NW of high terraces of Maraetotara River, Haustrum in teleoconch characters, with a south of Cape Kidnappers, Castlecliffian (OIS markedly taller spire, a smaller aperture and a 29-31?; GS5315, W21/f8522, one, severely markedly longer anterior siphonal canal, as abraded). 60 AG Beu

Distribution. Waipipianearly Castlecliffian nurse eggs by B. hanleyi in New South Wales. (OIS29 31?), in Auckland, Wanganui Basin, B. hanleyi produces crawl-away miniatures of North Canterbury, and Hawke’s Bay (Cape the adult, that is, it has direct development, and Kidnappers). Bedeva bartrumi is a rare fossil, the protoconch of the initial hatched juvenile presumably reflecting its exposed intertidal (Anderson 1966, fig. 6) consists only of a half- rocky shore habitat (to judge from that of whorl hemispherical ‘cap’ and an initial half- Recent Bedeva species in Australia) and the whorl of teleoconch. The New Zealand species poor preservation of this environment in the B. bartrumi therefore had similar development fossil record. It is unlikely to have any bios- to B. hanleyi, despite the larger (slightly more tratigraphical utility. multispiral) protoconch of B. bartrumi and, assuming it was a Pliocene arrival from SE Dimensions. Otahua bartrumi, figured paratype: Australia, the genus can have arrived in New H 45.7, D (incomplete) 20.3 mm; largest para- Zealand only by rafting. The distinct proto- type: H 46.8 (incomplete, originally ca 55 mm), conchs demonstrate that B. bartrumi is a species D (incomplete) 22.8 mm; almost complete distinct from B. paivae. Dr Emily Vokes (for- paratype (TM8301, ex CR Laws collection; merly of Tulane University pers. comm.) sev- Fig. 9E,G): H 43.2, D 20.2 mm. eral years ago pointed out the similarity of B. bartrumi to B. blosvillei (Deshayes, 1832). That Remarks. When proposing the genus Otahua, larger and more strictly tropical species has Marwick (1948) remarked that it was probably slightly more elaborate sculpture and has been closest to the eastern Australian genus Bedeva referred to Jousseaume, 1883 (Wil- Iredale, 1924, and the differences are in my son 1994:22, pl. 5, fig. 27A,B). However, the opinion of specific rather than generic rank. latest classification returned B. blosvillei to New Zealand specimens are particularly similar Bedeva; the 46-mm-high specimen illustrated to Bedeva paivae (Crosse, 1864) (Fig. 9F,I) as (Houart in Poppe 2008:198, pl. 394, fig. 10) is illustrated by Wilson (1994, pl. 5, fig. 28AC), similar to B. bartrumi. The Otahuhu specimens having similar proportions and sculpture, a are particularly similar to a specimen labelled similar sutural ramp without spiral cords and ‘Bedeva vapida Woolacott, 1957’ in GNS a similar sharp peripheral angulation, and a (WM9323, Yeppoon, Queensland, ex W. J. very similar aperture, anterior sculpture and Paul collection; H 34.1 mm) as it lacks the siphonal fasciole. The only obvious differences foliose axial sculpture on the sutural ramp of B.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 are the larger size, thicker shell, slightly weaker paivae; this name is listed by Wilson (1994:22) sculpture and more weakly armed aperture of as a synonym of B. blosvillei. Otahuhu specimens. Wilson (1994:21) pointed Specimens from Lower Waipara Gorge out the difficulty of distinguishing B. hanleyi (Waipipian), shellbeds on Brunswick Road, from B. paivae ‘in field samples’, and stated Kai-Iwi, Wanganui (late Nukumaruan) and that B. hanleyi reaches 40 mm in height. the early Castlecliffian succession (OIS Marwick (1948, pl. 7, figs. 5, 6) illustrated the 2931?) at Cape Kidnappers are all severely low, dome-shaped, lecithotrophic protoconch abraded and not certainly referred to this of about 1.5 whorls in B. bartrumi, possibly one species, but they show no differences to distin- of the reasons for his assuming that Otahua guish them from B. bartrumi, and have a must be a genus distinct from Bedeva, as the distinctive yellowish, calcitic outer shell layer. protoconch makes it unlikely that Bedeva can It appears that the ‘Australian’ genus Bedeva have dispersed as larvae from Australia to New became extinct in New Zealand only recently, Zealand. Anderson (1966:243244, figs. 36) during early Castlecliffian time, but was never described the brooding of within a common as a fossil, because of its intertidal brood pouch and their feeding on the yolks of rocky shore habitat. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 61

Hutton (1880:49) and Suter (1913:415, pl. 45, 9a,b), based on a ‘Sinusigera ’, Cheletropis fig. 22) mistakenly recorded the abundant south- huxleyi Forbes, 1852, collected in a towing net ern Australian species Bedeva paivae from the off Cape Howe, SE Australia, during the voyage New Zealand present-day fauna. Suter of the ‘Rattlesnake’. The same or a very similar (1913:416) cited its range as ‘North and South larva apparently had earlier been named Islands, but not common, between tide-marks to Struthiolaria microscopica by Gray (1839:108; about [40 m]; Chatham Islands’. However, attributed to Blainville, ‘Indian Ocean, Paris Finlay (1926b:421) stated unequivocally of Be- Museum’, but never published by Blainville), deva that ‘this genus does not occur in New and was identified again as Sinusigera micro- Zealand’. The two lots of specimens in Suter’s scopica by Brazier (1878:144) from ‘360 miles collection identified as ‘Trophon (Kalydon) [580 km] north-east of Sydney’. This synonymy hanleyi’ (none is identified as T. paivae) are is stated unequivocally in Charles Hedley’s from Titahi Bay, Wellington (Suter’s no. 3101, unpublished, loose-leaf bibliographic catalogue one; collected by MK Mestayer) and ‘Chatham in AMS. H Adams & A Adams (1854, vol. 2:89) Is.’ (Suter’s no. 2534, 13; no source stated); all are also suggested that Struthiolaria microscopica beach-abraded specimens of Axymene traversi. Gray was the same species as Macgillivrayia The entire record of Bedeva as a member of the (sic,Macgillivraya) pelagica Forbes, 1852 Recent fauna of New Zealand was based on (type species of Macgillivraya Forbes, 1852; misidentifications. The present fossils therefore ‘taken in the towing-net by Mr. Macgillivray represent the only authentic records of the genus off the East coast of Australia, fifteen miles in New Zealand. [24 km] from the shore’; H Adams & A Adams 1854, vol. 2:89). However, illustrations of M. pelagica (Forbes 1852, pl. 3, fig. 8a,b; H Subfamily Rapaninae Adams & A Adams 1854, pl. 69, fig. 3ac) show a very simplified, smooth-shelled with four Genus Agnewia Tenison Woods, 1878 long, narrow velar lobes and a triangular oper- culum, clearly a larva of rather than a Cheletropis Forbes 1852:385. Type species (by muricid, whereas Cheletropis huxleyi in Forbes’ monotypy): Cheletropis huxleyi Forbes, 1852, (1852, pl. 3, fig. 9a,b) illustration is a strongly Recent, SE Australia (larva of Agnewia tritoni- bicarinate, taller-spired shell with a typical formis (Blainville, 1832) (nomen oblitum). ‘sinusigera claw’ on the outer lip; H Adams &

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 A Adams’s suggested synonymy is incorrect. Adamsia Dunker 1857:357. Type species (by Sinusigera cancellata d’Orbigny (H Adams & original designation): (Adamsia) typica A Adams 1858, pl. 137, fig. 4) is much more like Dunker, 1857 ( tritoniformis Blain- C. huxleyi and the larva of A. tritoniformis. ville, 1832), Pleistocene and Recent, southeast- H Adams & A Adams (1858:613614) also noted ern Australia and northern New Zealand that Cheletropis huxleyi Forbes was a synonym (junior homonym of Adamsia Forbes, 1840, of Sinusigera microscopica (Gray), and that Tunicata). Cheletropis is a synonym of Sinusigera. Although the identity of Sinusigera cancellata Agnewia Tenison Woods 1878:29. Replacement Orbigny, 1841 (named in Orbigny’s [1841] work name for Adamsia Dunker, 1857, junior homo- on Cuba) seems not to have been established, it nym (nomen protectum). will have been based on the larva of quite a different muricid species from the Caribbean Sea, not congeneric with A. tritoniformis.Tomy Remarks. The earliest name for this genus seems knowledge, Cheletropis has not been used as the to be Cheletropis Forbes (1852:385, pl. 3, fig. valid name for a genus since 1899, and as 62 AG Beu

Agnewia tritoniformis is a well-known name 1962:179, fig. 216; Iredale & McMichael satisfying the requirements of ICZN Code 1962:73; Hodgkin et al. 1966:45, pl. 17, fig. 5; Article 23.9.1, Agnewia is declared a nomen Macpherson 1966:254; Grant-Mackie & Chap- protectum here, and Cheletropis is declared a man-Smith 1971:675; Fleming 1975:52, 74, fig.; nomen oblitum. The requisite list of names to p. 78, fig.; Powell 1976b:99; Fleming 1978:723; justify this status (at least 25 usages of Agnewia Fleming 1979:76, 97, 108, fig.; p. 112, fig.; Powell as the valid name for this genus by at least 10 1979:181, pl. 36, fig. 8; Rehder 1980:71, pl. 3, fig. authors during the last 50 years) is provided in 2 (); Trew & Oliver 1981:33; Monger the following synonymy. 1984:50, unnumbered fig.; Richmond 1992:47, fig. 67; Wilson 1994:40, pl. 5, fig. 19; Jansen Agnewia tritoniformis (Blainville, 1832) (Fig. 1995:57, fig. 218; Spencer & Willan 1996:26; 10D, F, G, H) Richardson 1997:9; Jansen 2000:46, fig. 175; Hayward et al. 2001:61, fig. 8; Spencer et al. Purpura tritoniformis Blainville 1832:221, pl. 10, 2002, on-line list; Tan 2003:967, figs. 3E,F, fig. 10; Kiener 1836, Purpura, p. 37, pl. 8, fig. 8; 5U,V, 18AK, 19; Herbert et al. 2007:21, fig. Kesteven 1902:533, pl. 29, figs. 2, 3, 5, 7; Hedley 4C,D; Spencer et al. 2009:210; Maxwell 1903:384; Suter in Hutton 1904:72; Pritchard & 2009:246. Gatliff 1906:44; Lamy 1918:357; Fischer-Piette Purpura tritoniformis levidensis Kesteven & Beigbeder 1943:433. 1902:534, pl. 29, figs. 2, 5, 7. Struthiolaria microscopica. Gray 1839:108 (lar- Thais () tritoniformis. Suter 1913:424, val shell). pl. 45, fig. 19. Cheletropis huxleyi Forbes 1852:385, pl. 3, fig. Thais (Agnewia) tritoniformis. Thiele 1929:298; 9a,b (larval shell). Wenz 1941:1122, fig. 3188. Adamsia typica Dunker 1857:357; Angas Agnewia nautica Thornley 1952:43, fig. 2. 1867:192; Kobelt 1878:236; Gillies 1882:171. Sinusigera microscopica Brazier 1878:144 (larval Type material. Purpura tritoniformis Blainville, shell). ‘probable holotype’ in MNHN, as noted by Agnewia typica. Tenison Woods 1878:29. Lamy (1918:357); without locality; severely tritoniformis. Tryon 1880a:156, pl. abraded and evidently a beach shell, but 39, fig. 491 (not including figs. 488, 496). certainly conspecific with the species usually Cominella (Adamsia) typica. Watson 1886:214. known by this name; illustrated by Tan

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Cominella (Agnewia) tritoniformis. Brazier (2003:967, fig. 18A,B). The type locality is 1888:995; Whitelegge 1889:248; Brazier here designated as Sydney Harbour (Port 1894:167; Pritchard & Gatliff 1898:275. Jackson), New South Wales. Cheletropis hux- Agnewia tritoniformis. Tate & May 1901:353; leyi, holotype not found in BMNH, location Iredale 1915:475; Hedley 1918:M93; Cooke unknown (K Way, BMNH pers. comm. 14 1919:100, fig. 26 (radula); May 1921:87; Gatliff January 2008); probably such a minute, largely & Gabriel 1922:131; May 1923:85, pl. 40, fig. 16; conchiolin shell (a larval shell of A. tritonifor- Finlay 1926b:427, pl. 21, fig. 82; Powell 1937:80; mis) has not survived storage in ethanol since it Powell 1940:214; Powell 1946a:83; Dell 1950:26; was collected during the cruise of the ‘Rattle- Macpherson & Chapple 1951:130; Richardson snake’ in 18461850. Adamsia typica, four 1953:39, figs. 35; Bennett & Pope 1953:131; syntypes BMNH 1991575, labelled ‘Philippi- Kershaw 1955:316; Powell 1958a:103; Allan nen. Adelaide?’ in Dunker’s handwriting (Tan 1959:147, text-fig. 33.2; McMichael 1960:112, 2003:965) (not seen), almost certainly from SE fig. 252; Bennett & Pope in Dakin 1960:257, pl. Australia (although records from as far west as 65, fig. 2; Powell 1962:95; Fleming 1962b:86, 98, Adelaide are not confirmed by modern collec- fig.; p. 102, fig.; Macpherson & Gabriel tions; probably from near Sydney, New South Marine Mollusca of isotope stages of the last 2 million years in New Zealand 63 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 10 (A,I) ‘‘Adamsia’’ adelaidae A. Adams & Angas, holotype, BMNH 1870.10.26.76, Recent, ‘‘Adelaide’’, South Australia; H 16.7, D 7.5 mm. (B,C,E,J) Agnewia kempae Powell, GS10911, Y14/ f7505C, Te Piki Member (Haweran, OIS7), Te Piki, near East Cape; B,C,E, specimen lacking protoconch, all data as for Fig. 10J; 3 views of one specimen, with prominent nodule on outer lip, height 20.0 mm; J, apex of specimen in Fig. 9L. (D,F,G,H) Agnewia tritoniformis (Blainville), type species of Agnewia Tenison Woods, 1878, Recent; D,H, WM6364, Sydney, New South Wales, Australia; D, spire apex; H, protoconch of D; F,G, RM4784, dived, 5 m, Cape Karikari, Northland, New Zealand; height 19.7 mm; F, unwhitened; G, whitened with MgO. 64 AG Beu

Wales). Purpura tritoniformis levidensis Keste- Port Phillip and Westernport to Lorne and ven, holotype AMSC.13179 (Tan 2003, fig. Apollo Bay (Macpherson & Gabriel 1962:179). 18C,D), said by Kesteven to be from ‘New I am not aware of authentic records from South South Wales and Lord Howe Island’, but the Australia. holotype is labelled ‘Port Jackson, Sydney’, a strongly axially ridged specimen of A. tritoni- Dimensions. Purpura tritoniformis, ‘probable’ formis. Agnewia nautica Thornley, holotype holotype: H 21.4, D 12.4 mm; Purpura tritoni- AMSC.103331, with 14 paratypes, C.305363 formis levidensis holotype: H 28.3, D 14.9 mm; (nine), C.346697-8 (five), all from Port Ste- Agnewia nautica holotype: H 24.6, D 14.0 mm; phens, New South Wales (holotype: Tan 2003, GS14778, Hokianga, OIS 7: H 22.1 (incom- fig. 18E,F), a rather short and weakly sculp- plete; originally ca 27 mm), D (incomplete) tured form of A. tritoniformis. I am not aware 12.4 mm; RM4784, Cape Karikari: H 21.0, D of the repository of any type material of 11.3 mm; H 19.7, D 11.8 mm; H 18.8, D 10.0 Struthiolaria microscopica. mm; NMNZ M.212303, Lord Howe Island: H 35.3, D 18.9 mm; NMNZ M.39766, intertidal, Other material examined. Haweran: Akera-Dia- Otarawairere Bay, Ohope Beach, Bay of Plenty: la-Anadara locality, bank behind tidal flats H 29.5, D 15.7 mm; H 24.0, D 15.1 mm. between Te Rewa Point and Waiparera, Ho- kianga Harbour, Northland, OIS7 (GS14778, Remarks. The restricted New Zealand species O06/f064, one). Recent: RM2018, Whangapar- Agnewia kempae Powell, 1933 occurs uncom- apara, Great Barrier Island, CA Fleming monly at Te Piki, inland from Whangaparaoa, collection (one); RM4784, SCUBA, 5 m, Cape near East Cape (OIS7) (see below). It seems Karikari, Northland (seven); 19 lots in NMNZ, likely that A. kempae evolved through genetic from Cape Maria van Diemen to as far SE drift from a founder population of the very as Cape Runaway, and to Koakanui Point, similar species A. tritoniformis that appeared in Tokomaru Bay, N of Gisborne (NMNZ New Zealand as planktonic larvae from eastern M.21639) (listed by Tan 2003:968). Australia (Grant-Mackie & Chapman-Smith 1971:675; Beu & Maxwell 1990:360). A. tritoni- Distribution. OIS7 and Recent in New Zealand; formis is common on SE Australian shores at living in the northeastern North Island warm present, and is well known for its classical province only. Although Gillies (1882:171) ‘Sinusigera’ protoconch (Fig. 10D, H) (Kesteven

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 recorded Agnewia tritoniformis from New Zeal- 1902; Powell 1979:181), implying a long plank- and in his comments on Hutton’s (1880) cata- totrophic larval life. A. tritoniformis now lives logue with the bald statement ‘Adamsia uncommonly in several remote localities, sub- typica*Add’, Finlay (1926b:427) was the first tidally on extremely exposed rocky shores, in to provide accurate locality records from New northeastern New Zealand, from Cape Maria Zealand (two specimens in the collection of W van Diemen to Tokomaru Bay, N of Gisborne La Roche). Powell (1940:214) recorded speci- (Richardson 1953; Tan 2003). A single poorly mens from Cape Maria van Diemen and Tom preserved fossil specimen of A. tritoniformis Bowling Bay, N Northland, and Dell (1950:26) collected from the Anadara--Akera locality recorded further specimens from four localities. at Te Rewa Point, northern Hokianga Harbour, Richardson (1953) added 17 further records, as indicates that this species appeared in New far south as Oruaiti Bay, Whangaparaoa, E Bay Zealand at least as early as OIS7, approximately of Plenty. In Australia, it lives from New South coeval with the Te Piki records of A. kempae.It Wales (where it is common) to Bass Strait and seems likely that Agnewia arrived in New Zeal- (Wilson 1994:40), and at Lord Howe and at some time earlier than OIS7 (presumably and Norfolk Islands; in Victoria, recorded from during OIS11 9) and evolved into the endemic Marine Mollusca of isotope stages of the last 2 million years in New Zealand 65

A. kempae, and that OIS7 and later records of metres. A concentration I excavated recently A. tritoniformis are pseudopopulations recruited had an uppermost layer ca 5080 mm thick of as planktotrophic larvae from eastern Australia. concentrated, broken Atrina shells, with mod- erately common specimens of the ranellid Agnewia kempae Powell, 1934 (Figs. 9L, 10B, C, caudata (Gmelin, 1791), whereas L. E, J) caudata was not collected further inside the shell concentration. This indicates that a pin- Agnewia kempae Powell 1934a:273, pl. 59, figs. nid-dependent community was developed 19, 20; Richardson 1953:41, figs. 1, 2; Fleming across the estuarine flat, where fed 1966:58; Grant-Mackie & Chapman-Smith on Pinnidae, as has been observed at several 1971:675; Beu & Maxwell 1990:360, pl. 48h; localities in estuaries in the tropical Pacific, Richardson 1997:9; Maxwell 2009:246. along the South African coast, and at Whan- garei Heads in Northland at present. The Type material. Agnewia kempae, holotype AIM concentrations evidently represent tidally exca- AK70011, with one paratype AK72423 (neither vated ‘pot-holes’ in the estuarine flats that seen), from Te Piki, near Whangaparaoa, trapped shells washed into the small Te Piki East Cape, Haweran (penultimate interglacial, embayment from the present Waihau Bay and OIS7). the open coast outside it. The commonness of some species, including Agnewia kempae, varies Other material examined. GS10911, Y14/ from concentration to concentration, and A. f7505C, Te Piki Member (OIS7), Te Piki kempae has not been collected during recent road cutting, ca 6 km E of Whangaparaoa, visits. near East Cape (one); same locality, CA Flem- ing collection (presented by AWB Powell?) Remarks. Agnewia kempae is compared here (five); same locality, NMNZ M.26395 (one); with A. tritoniformis,asA. kempae seems to be M.40126 (two). a restricted New Zealand species. As pointed out by Beu & Maxwell (1990:360), the two Dimensions. GS10911, Te Piki: H 20.0 (proto- species are very similar. However, A. kempae is conch incomplete), D 9.8 mm (Fig. 10B, C, E); slightly smaller (H 20 rather than up to at least H 19.7, D 10.3 m; H 18.2, D 10.0 mm; H 35 mm) and most specimens are slightly nar- 16.9, D 8.8 mm; H 15.5, D (slightly incomplete) rower than A. tritoniformis. Also, A. kempae is

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 7.9 mm; M.26395: H 20.1, D 9.8 mm. consistently more thin-shelled than A. tritoni- formis, and also differs in consistently having Distribution. Agnewia kempae is recorded only weaker axial folds and primary spiral cords, from the Te Piki Member (OIS7) at Te Piki, fading out before the last whorl, revealing a inland from Whangaparaoa, near East Cape. relatively narrow, prominent peribasal spiral Most of the more unusual fossils at Te Piki cord, sculptured with several narrower, closely (that is, other than roseus, Dosina spaced spiral cords, and bearing a shallow zelandica Gray, 1835, Dosinia (Kereia) greyi groove along its crest. The surface is otherwise Zittel, 1864, and D. (Fallartemis) lambata sculptured with single, fine, moderately widely (Gould, 1850), which are common throughout spaced, sharply raised, slightly wavy spiral the member) occur in small but exceedingly cords and fine axial ridgelets in A. kempae. diverse, closely packed fossil concentrations The cords are single over most of the surface (ca122 m) greatly dominated by and rather evenly spaced over the entire exter- novaezelandiae (Reeve, 1853). The concentra- ior, with a few narrow secondary threads on the tions are scattered sparsely through the mem- centre of the last whorl, whereas they ber, separated from each other by some tens of are significantly more closely spaced into 66 AG Beu

groups that form the major spiral cords in A. slightly taller and narrower and is well pre- tritoniformis, with many secondary and finer served on most specimens of A. kempae, cords; the major spiral interspaces bear more whereas that of A. tritoniformis is slightly widely spaced spiral cords and otherwise weak- shorter and in most specimens is more abraded, er sculpture. One of the Te Piki specimens in suggesting that A. kempae occupied a more M.40126 retains wide, evenly rounded axial sheltered habitat than the rocky subtidal one of folds to the end of the last whorl, but its much A. tritoniformis. It seems feasible that Agnewia finer spiral cords than in A. tritoniformis and its kempae occupied the shallow, sheltered, sub- peribasal cord are readily distinguished. The tidal Atrina beds described under ‘Distribu- raised peribasal cord ends at an anteriorly tion’, above. protruding nodule on the outer lip, quite Consideration was given to the possibility prominent in some specimens (Fig. 30C,F,G), that Agnewia kempae might be another species although it is not well preserved on all speci- previously known from Australia, rather than a mens. Freshly collected specimens of A. kempae restricted New Zealand species. The only other are rather a bright reddish brown, reflecting Recent Australian species that has been as- their calcitic outer shell layer (Tan 2003:969) signed to Agnewia in recent years is Adamsia and possibly in part the original exterior adelaidae A Adams & Angas (1864:421, pl. 37, colour. Comparison of the nine specimens of fig. 2) (not to be confused with A. kempae listed above directly with the seven adelaidensis Crosse & Fischer [1865:50, pl. 2, of A. tritoniformis in RM4784, from Cape fig. 1], which was listed as a synonym of Karikari, and with numerous specimens from Haustrum vinosum (Lamarck, 1822) by Tan Sydney Harbour showed that they can be [2003:998]). The poorly known species Adamsia distinguished consistently by relative sculptural adelaidae tentatively was assigned to Agnewia prominence (much weaker in A. kempae than by Tan (2003:965), but definitely is not con- in A. tritoniformis), spire height (taller in generic with A. tritoniformis and is unlike any most specimens of A. tritoniformis than in other Australian muricid I know of (holotype A. kempae), the peribasal spiral swelling of BMNH 1870.10.26.76; Fig. 10A, I). It is a small A. kempae and the anteriorly protruding nodule (H 16.7, D 7.5 mm), tall and narrow, tall-spired on the outer lip of A. kempae, contrasting shell with 1214 moderately prominent, widely strongly with the finely and narrowly sinuous spaced, narrow axial ridges per whorl, two lip edge in all specimens of A. tritoniformis. wide, prominent spiral cords around the per-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Most specimens of A. kempae that lack the iphery, a further narrower one on the ramp and nodule on the outer lip have a thickened, rough four on the neck of the last whorl, a thin outer area on the anterior edge of the lip, apparently lip, and a very short, widely open anterior the base of the former nodule, now broken off. siphonal canal. The entire exterior is sculptured Microscopic examination showed that the axial with many low, thin, widely spaced, irregular growth ridges are curved around a formerly axial lamellae, and the pale brown exterior, protruding lip extension over the peribasal cord with paler spiral cords, indicates the presence of of all specimens, even when the nodule is not a calcite outer shell layer. The protoconch is preserved on the lip itself. Evidently A. kempae blunt and paucispiral, smooth, of about 11.5 fed by opening (edge-chipping?*as described whorls; that is, similar to those of Bedeva paivae for the muricid trunculus (Linne´ , and Haustrum vinosum (Tan 2003, fig. 2A,B) 1758) by Morton et al. 2007) other molluscan although not clearly differentiated from the shells with its lip nodule in a manner not teleoconch, very different from the plankto- adopted by A. tritoniformis, so the nodule trophic ‘sinusigera’ of A. tritoniformis. A rela- presumably was broken and regrown during tionship with Bedeva and Haustrum (subfamily life by some specimens. The protoconch also is Rapaninae; Claremont et al. 2008:218) seems Marine Mollusca of isotope stages of the last 2 million years in New Zealand 67

likely, but only examination of the radula and aperture (in C. otaioensis); and with sculpture anatomy will confirm the generic position of dominated by prominent, well-rounded axial Adamsia adelaidae. No similar species have ridges with evenly concave interspaces each been illustrated in any recent identification equal in width to one ridge, extending from guides to Australian gastropods. just below the upper suture (forming a narrow, The type material (holotype and 14 para- concave sutural ramp above the axial ridges) to types) of Agnewia nautica Thornley, 1952 also the lower suture on spire whorls and fading out was compared carefully with A. kempae, as this over the base on the last whorl, crossed by rather small, short-spired, weakly sculptured prominent, sharply defined, widely spaced, form is most similar to A. kempae of Australian convex-crested spiral cords over the entire material I have examined. However, all materi- teleoconch surface, with narrow secondary al assigned to A. nautica has the same spiral threads in some interspaces, forming only low sculpture as A. tritoniformis and lacks the nodules at sculptural intersections. The proto- protruding small rounded nodule on the outer conch is low, dome-shaped, of about two lip, formed by a peribasal spiral cord, that whorls, with its termination indistinct; the characterizes A. kempae. A. nautica is correctly initial tip is minute, sharp, and followed by a assigned to the synonymy of A. tritoniformis. rather wide, flattened upper surface; the second Apparently A. kempae evolved from A. tritoni- whorl is taller, bearing low axial ridges; incom- formis in New Zealand, rather than being a plete or corroded on all material seen. migrant from Australia. Remarks. Finlay (1930a:60) proposed the genus Pleia for shells resembling a small species of Family but having one low, narrow rather than three conspicuous columellar plaits, and Genus Cryptofusus n. gen. selected the Australian middle Miocene species decipiens Tate (1888:150, pl. 8, fig. 1; a synonym of P. tenisoni (Tenison Woods, 1879); Type species. Pleia cryptocarinata Dell, 1956, TA Darragh Museum Victoria pers. comm. NukumaruanRecent, bathyal, southern and October 2007) as the type species. He referred central New Zealand. his new species Pleia otaioensis (from Bluecliffs, Otaio River, New Zealand; Otaian, early Mio-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Diagnosis. A genus of Turbinellidae containing cene) to Pleia, but it seems likely that Finlay small, tall, narrow shells (2035 mm high) with misinterpreted the scale of Tate’s (1888) illustra- a tall, narrow spire, a long, straight, narrow, tion, as he somehow failed to realize the size anterior siphonal canal without a siphonal difference between the New Zealand and Aus- fasciole (at least, in later, MioceneRecent tralian species. Pleia tenisoni (Balcombian, Mid- species), the spire about equal in height to the dle Miocene; Muddy Creek, Victoria and River aperture and canal; a small, oval, weakly armed Murray cliffs, South Australia; Darragh aperture with a single low, narrow columellar 1970:165, 200; Fig. 11E, H) is a much larger, plait terminating at the base of the columella, more coarsely sculptured species than the New and two much weaker plaits decreasing in Zealand fossil and Recent species assigned to prominence above in some species (Cryptofusus Pleia (illustrated specimen, Fig. 11E,H; NMV otaioensis (Finlay, 1930)), the outer lip moder- P.314675, Muddy Creek Formation [Balcom- ately thin and crenulated by the exterior spiral bian, early middle Miocene], Muddy Creek, near cords (in C. cryptocarinatus), or slightly thick- Hamilton, western Victoria: H 70.8, D 25.4 mm, ened and bearing low transverse ridges, and rather than 2035 mm as in Cryptofusus with 56 low, short lirae well within the species). It also has markedly wider whorls, 68 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 11 (A,K) Kuroshioturris angustata (Powell), RM4033, Recent, 90 m, Doubtless Bay, Northland, SEM; A, whole specimen, height 9.4 mm; K, protoconch in A. (B) Cryptofusus cryptocarinatus (Dell), type species of Cryptofusus n. gen., GS10844, U26/f6037, Castlepoint, E Wairarapa, early Nukumaruan; height 23.3 mm. (C,D,F) Kuroshioturris putere n. sp., GS1567, W19/f7516, right bank Waiau River at junction Pakihiwi Stream, inland from Wairoa, N Hawke’s Bay, Opoitian; C, paratype, TM8673, height 14.0 mm; D,F, holotype, TM8672, height 13.7 mm. (E,H) Pleia tenisoni (Tenison Woods), type species of Pleia Finlay, 1930, NMV P.314675, Muddy Creek Formation (Balcombian, Middle Miocene), Muddy Creek, near Hamilton, W Victoria, Australia; height 70.8 mm. (G,J) Amalda (Baryspira) olsoni Beu, GS11225, V20/f8002, Darkys Spur Formation (late Nukumaruan), Darkys Spur, W of Devils Elbow, Hawke’s Bay; G, height 31.9 mm; J, height 34.3 mm (see also Fig. 17E). (I) Cryptofusus otaioensis (Finlay), GS9569, I38/f7748, Mount Harris Formation (Waitakian), upper Tengawai River, Canterbury; height 25.8 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 69

more prominent peripheral nodules defining a 12, fig. 2a,b; P. bakeri: Wilson 1994:74 [as a form much more obvious, strongly concave sutural of P. australasiae], pl. 12, fig. 2c). The new genus ramp, a second row of peribasal nodules defining Cryptofusus is proposed for the New Zealand a lower angulation, a sharply contracted neck, a species previously referred to Pleia, with Pleia very long, narrow, twisted siphonal canal, an cryptocarinata Dell, 1956 selected as type species unusually widely flared outer lip, and a single so that Recent specimens can be used to deter- very low, narrow columellar plait stopping well mine phylogenetically important characters. inside the aperture but placed significantly high- In most characters Cryptofusus n. gen. is er up than in Cryptofusus n. gen. The new genus more similar to Bellardi, 1884 Cryptofusus, in contrast, contains much smaller (type species: Murex lancea Gmelin, 1791, shells with rounded axial ridges having only Recent, Indo-West Pacific) (Wilson 1994:67, weak nodules, a more weakly defined sutural pl. 11, figs. 69, 15) than to Pleia tenisoni, but ramp, a rather more gently contracted base, and differs in its shorter shape, its weaker sculpture a shorter siphonal canal. The protoconch of P. and its much less obvious columellar plait, tenisoni is relatively large (1.8 mm wide) and terminating inside the aperture and only just blunt, of little over one whorl, smooth and visible at the columellar base of most species. polished, with a large, rounded but low initia- Latirogona ornata (Marshall, 1918) (Otaian tion, merging gradually into the teleoconch over Altonian, Pakaurangi Point, Kaipara Harbour, a quarter of a whorl. It is clearly a lecithotrophic Northland; Beu & Maxwell 1990:230, pl. 25s), one, so the larva of Pleia is unlikely to have been type and only species of Latirogona Laws, 1944, dispersed from Australia to New Zealand. The is similar to Dolicholatirus and Cryptofusus in large Australian species of Pleia clearly are more size (height 2030 mm) and general appear- nearly related to Pleuroploca Fischer, 1884 ance, but is a little wider, with more inflated (particularly some of the smaller Australian whorls, more prominent axial folds, a more Recent species such as P. bakeri (Gatliff & complex series of fine, closely spaced spiral Gabriel, 1912); Wilson 1994, pl. 12, fig. 2a,c) cords, and a much more complexly and promi- and, in particular, Latirus Montfort, 1810 and nently armoured aperture: ‘columella with two Hemipolygona Rovereto, 1899 (Wilson 1994, pl. or three definite plaits, and 36 additional short 11; Vermeij & Snyder 2006, figs. 1, 2) than to the ridges and tubercles; parietal region with 24 small, elongate, axially ridged but otherwise short ridges. Outer lip with 911 narrow lirae weakly sculptured New Zealand species included within’ (Beu & Maxwell 1990:230). The Eur-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 by Finlay (1930a) in Pleia. Indeed, Pleia tenisoni opean Cenozoic genera Latirulus Cossmann, seems closely related to Latirus, differing mainly 1899 (Lathyrulus Cossmann, 1901) (type in its much weaker columellar plait and much species: L. subaffinis (Orbigny, 1850), Eocene, longer, narrower siphonal canal. Darragh (1970) Paris Basin) and Dertonia Bellardi, 1884 (type referred to Pleia the three Australian middle species: D. iriae Bellardi, 1884, Pliocene, Italy) Miocene species Fasciolaria concinna Tate, 1888, are similar in shape and size to Cryptofusus, but F. decipiens and F. tenisoni, although in his differ in sculpture, have a shorter anterior opinion P. tenisoni and P. decipiens are conspe- siphonal canal, and have three prominent cific. Iredale & McMichael (1962:68) also re- columellar plaits (Wenz 1943:12421243). ferred the two Recent New South Wales species Maxwell (1992:123, pl. 16mq) assigned his ‘Pleia coronata (Lamarck, 1822)’ and ‘P.’ bakeri new species Latirulus(?) fraudator (Kaiatan, late (Gatliff & Gabriel, 1912) to Pleia, but these are Eocene, McCulloch’s Bridge, South Canter- typical species of Pleuroploca Fischer, 1884, with bury) to Latirulus quite hesitantly, and it 23 much more prominent columellar plaits is certainly much smaller than European spe- (‘Pleia’ coronata is a synonym of Pleuroploca cies, with a still shorter siphonal canal, and australasiae (Perry, 1811); Wilson 1994:74, pl. possibly represents a further unnamed genus. 70 AG Beu

Vermeij & Snyder (2006:419) also doubted that likely belongs in the Turbinellidae and is not L. fraudator is correctly assigned to Latirulus, a fasciolariid’ (Vermeij & Snyder 2006:414). On which they considered to be limited to Eocene this basis, it appears likely that Pleia belongs in rocks of Europe. Still further genera probably the Fasciolariidae, subfamily Peristerniinae, will be required for New Zealand Turbinellidae. near Latirus, whereas Cryptofusus belongs in Maxwell (1992:124, fig. 9g, pl. 18m) referred the Turbinellidae near Dolicholatirus. A posi- Latirofusus optatus Marshall & Murdoch, with tion in Turbinellidae tentatively is accepted some hesitation, to Fractolatirus Iredale, 1936 here, for both Cryptofusus n. gen. and Latir- (type species: F. normalis Iredale, 1936, Recent, ogona Laws, 1944. Similar small, elongate Sydney Harbour) but F. normalis has two equal tubinellids were referred to Subfamily Ptycha- (if weak) plaits, is prominently lirate deep tractinae by Beu & Maxwell (1990:414) but within the outer lip, and is a shorter, more Family Ptychatractidae Stimpson, 1865 was solid shell resembling a small species of Latirus referred to Superfamily Pseudolivoidea de Gre- (H 26 mm). Judging from Maxwell’s (1992:124) gorio, 1880 by Bouchet et al. (2005:256). comparisons a position in Streptochetus Coss- Dolicholatirus, Cryptofusus, Latirogona and mann, 1899 seems more reasonable, as the the New Zealand ‘ptychatractines’ Egestas protoconch differences pointed out by Maxwell Finlay, 1926, Conrad, 1860 (referred to are not now considered to be a generic char- Turbinellidae by Kantor et al. 2001), Latiromi- acter (paucispiral, that is, lecithotrophic in tra Locard, 1897 (see Bouchet & Kantor 2000) Streptochetus; of four whorls, conical, that is, and Paleopsephea Wade, 1926 are referred here planktotrophic in ‘L’. optatus). However, Strep- to Turbinellidae Turbinellinae Swainson, 1835. tochetus intortus (Lamarck, 1803) has a very Species referred to Cryptofusus n. gen. are: prominent, flared siphonal fasciole on a shell Cryptofusus cryptocarinatus (Dell) (1956b:89, otherwise quite like Cryptofusus new genus, pl. 14, fig. 137) (Fig. 11B), bathyal, Nukumar- although with weaker spiral cords and axial uanRecent. Fossils in bathyal fauna at Palliser ridges and less strongly convex whorls (Wenz Bay (Beu 1967:111, pl. 2, fig. 17), and at 1943:1264, fig. 3598). Streptochetus incertus (Deshayes, 1835) in the coloured illustration Castlepoint, E Wairarapa coast (GS10844, by Merle et al. (2008, pl. 27, fig. 3) lacks the U26/f6037, one specimen, Fig. 11B, collected widely flared fasciole, but is significantly larger and presented by Trevor Ritsema). and more finely sculptured than L. optatus and Cryptofusus otaiaoensis (Finlay) (1930a:60, pl. 3, fig. 30) (Fig. 11I), WaitakianAltonian, Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Cryptofusus species. L. optatus is possibly an early species of Cryptofusus, in a genus that Mount Harris Formation, South Canterbur- later lost planktotrophy, although its spiral yNorth Otago (GS9569, I38/f7748, upper sculpture is finer than in all other species Tengawai River, Waitakian, Fig. 11I; GS458, assigned here. The weak siphonal fasciole also J39/f7476, Pareora River, Otaian; GS11154, could have been lost by later species. Better J39/f026, Pareora River below Mt Horrible, material will help resolve the position of C. (?) Otaian; GS11149, X16/f8050, ‘The Peaks’, optatus. Darragh (1970) referred Fusus acifor- Waihao Downs, Altonian). mis Tate (1888:139, pl. 7, fig. 5a,b) to Fracto- Cryptofusus n. sp., axial ridges obsolete on last latirus also, but it is a typical species of two whorls, columellar plaits as in O. otaioen- Dolicholatirus. sis; one specimen from Pukeuri road cutting, A final question is the family position. Oamaru, Altonian (GS1912, J41/f9499);‘N. Vermeij & Snyder (2006) revised the genera gen. of Fasciolariidae’ in list by Olson in Gage related to Latirus, with excellent coloured (1957:125) (possibly an aberrant specimen of C. figures, and concluded that ‘Dolicholatirus otaioensis). Marine Mollusca of isotope stages of the last 2 million years in New Zealand 71

Cryptofusus(?) n. sp., small (height 1314 mm), stortha Olson, 1956, early and middle Miocene with two equal, weak columellar plaits, sipho- (AltonianWaiauan), New Zealand. nal canal relatively short; several specimens from Target Gully Shellbed, Oamaru, Altonian (labelled ‘Lathyrulus’ in early GNScollections). Remarks. The distinction between such taxa as Cryptofusus(?) optatus (Marshall & Murdoch) Amalda (Spinaspira), with a narrow apex (1923:123, pl. 12, fig. 2), Kaiatan (late Eocene), apparently reflecting a narrow, multispiral McCulloch’s Bridge, South Canterbury. protoconch, and Amalda (Baryspira), with a broader apex, seems to be based solely on a Etymology. Composed from (Greek, developmental difference, and is not recognized secret, hidden) and from the name of the type here. The ‘subgenera’ (listed above) proposed species, Pleia cryptocarinata, and from fusus by Olson (1956) intergrade in shell shape and (Latin, a spindle), used in the names of many have very similar radular and anatomical slender, siphonostomatous gastropods. Gender characters, and are synonyms in my opinion. masculine. In contrast, the distinctive radula demonstrates that Gracilispira Olson, 1956 is probably a separate genus. Family Olividae Subfamily Ancillariinae Amalda (Baryspira) olsoni Beu, 1970 (Figs. 11G,J, 17E) Genus Amalda H. Adams & A. Adams, 1853 Amalda (Gracilispira) olsoni Beu 1970a:132, pl. Amalda H Adams & A Adams 1853:148. Type 5, figs. 3941. species (by subsequent designation, Vokes Amalda (Baryspira) olsoni. Beu & Maxwell 1939:131): Ancillaria tankervillii Swainson, 1990:416; Maxwell 2009:246. 1825, Recent, tropical western Atlantic (Vene- zuela to N Brazil). Type material. Holotype VM316, with five paratypes VM317321, in Geology Depart- Subgenus Baryspira Fischer, 1883 ment, Victoria University of Wellington; VM316320 from Devils Elbow Mudstone, Baryspira Fischer 1883:600. Type species (by top of Devils Elbow hill, Highway 2, ca 35

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 subsequent designation, Cossmann 1889:64): km N of Napier; VM321 from Te Ngaru Ancillaria australis G. B. Sowerby I, 1830, Mudstone (the mudstone formation underlying PlioceneRecent, New Zealand. Devils Elbow Mudstone) at the same locality; all late Nukumaruan. Pinguispira Finlay 1926b:433. type species (by original designation): (Baryspira) opi- Other material examined. GS11225, V20/f8002, ma Marwick, 1924, Pleistocene (Nukumaruan), Darkys Spur Formation, long NShorizontal New Zealand. cutting on Darkys Spur road, between Waipunga and Kaiwaka, 5 km WNW of Devils Gemaspira Olson 1956:14. Type species (by Elbow, late Nukumaruan (abundant; 86 original designation): Ancilla (Baryspira) ro- collected). busta Marwick, 1924, Oligoceneearly Miocene (DuntroonianAltonian), New Zealand. Distribution. I (Beu 1970a:133) originally knew of Amalda olsoni only from the two uppermost Spinaspira Olson 1956:17. Type species (by siltstone units exposed on Devils Elbow hill original designation): Baryspira (Spinaspira) road. However, it has since proven to be very 72 AG Beu

much more common and to attain a much callus (in conventional ventral view) enveloping larger size in shallow-water sandy mudstone of the uppermost part of the outer lip and curving Darkys Spur Formation (Haywick et al. slightly backwards from the outer lip to extend 1991:219) exposed on Darkys Spur road, on up in an almost straight line to cross the spire the steep face above Waipunga, 5 km (straight apex, and the left edge of the spire callus line) WNW of the exposure at the top of Devils curving around to the left to the suture, and Elbow hill. I have still seen specimens from only then back to the right to merge with the parietal these two localities, so it apparently is limited to callus. The prominent, well raised, semicircular late Nukumaruan rocks of central Hawke’s pad thus produced on the left side of the spire is Bay. The uncommon specimens at Devils unique and renders A. olsoni easily recognized, Elbow evidently represent the fringe of the as it contrasts strongly with the slightly but population of a species that preferred shallower evenly convex outlines and narrowly pointed water nearer to shore. apex of the spire of A. australis and A. mucronata. The top fasciolar band around the Dimensions. Holotype: H 20.5, D 9.2 mm; base also protrudes more obviously (to the left largest paratype, VM317: H 27.7, D 12.3 mm in standard apertural view) than in all other (Beu 1970a:133); GS11225, Darkys Spur: H species I have compared it with. A. olsoni is, 42.8, D (incomplete) 18.4 mm; H 38.0, D 18.2 therefore, a distinctive species limited to mm; H 35.6, D (incomplete) 16.8 mm; H 34.4, Hawke’s Bay late Nukumaruan rocks, D 17.8 mm; H 32.1, D 15.5 mm; H 30.0, D 14.8 although apparently common only at Darkys mm; H 27.7, D 14.6 mm. Spur. Remarks. Because of its unusually tall, rela- tively narrow spire, with straight outlines apart Superfamily Conoidea from the distinctive spire callus protruding to the left (in conventional apertural, spire-up- Remarks. As noted by Bouchet (1990), some ward view), and because the type specimens earlier taxonomists (particularly Powell 1942, were relatively small and narrow, I (Beu 1944, 1966; Laseron 1954; Shuto 1969, 1971, 1970a:132) placed Amalda olsoni in the sub- 1983, and in several other papers) subdividing genus A. (Gracilispira) Olson, 1956 (type the huge ‘family’ recognized ‘pairs’ of species: Ancillaria novaezelandiae G. B. Sower- genera with distinct protoconchs but identical by II, 1859, PlioceneRecent, New Zealand). teleoconch characters in several groups, origin- Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Collection of the much larger specimens (3042 ally based on Finlay’s (1931) outdated concept mm rather than 2027 mm high) that are that one genus could not contain more than one abundant at Darkys Spur immediately made protoconch type. Turridae is now recognized as it obvious that A. olsoni is actually a species of Superfamily Conoidea, composed of several A. (Baryspira) resembling the Pliocene to families (Taylor et al. 1993; Puillandre et al. Recent shallow-water species A. australis (G. 2007, 2008), containing more than 11,000 B. Sowerby I, 1830) in most characters, parti- named species (Tucker 2004; Bouchet et al. cularly the size and shape of the last whorl, and 2009). One genus in each of these ‘pairs’ of a change of subgenus was adopted by Beu & genera has a blunt, paucispiral, relatively sim- Maxwell (1990:416). A. olsoni differs from A. ply sculptured protoconch, reflecting lecitho- australis in its narrower shape, caused by the trophic development, whereas the other has a taller and narrower spire, its evenly hemisphe- narrower, pointed, multiwhorled, more elabo- rical spire apex heavily coated with successive rately sculptured protoconch, reflecting plank- layers of spire callus, the right edge of the spire totrophic development. The various types of Marine Mollusca of isotope stages of the last 2 million years in New Zealand 73

lecithotrophic development are not distinguish- required to delete ‘genera’ based solely on able from protoconch characters alone, so all developmental differences. Medinskaya & Sy- non-planktotrophs are referred to here as soev (2001) and Kantor et al. (2008) have also lecithotrophs, whether development is direct demonstrated that teleoconch characters do not (spending their whole larval life protected with- always reflect radular differences in Conoidea, in the parent’s brood pouch, as in Pelicaria,or so the molecular phylogeny of the subfamily within an egg capsule as in volutes such as (Heralde et al. 2007) and of the Alicithoe, in either case emerging as a small, superfamily begun by Puillandre et al. (2008) crawling adult), demersal, or short-lived plank- and Bandyopadhyay et al. (2008) is desperately tonic (in either case with a brief free-living in need of expansion with many further taxa. larva, feeding on yolk as in direct developers, Many New Zealand Conoidea need to be rather than feeding in the ). The point reclassified to delete meaningless ‘genera’ based of Bouchet’s (1990) comment is that plankto- only on developmental type. Some obvious cases trophy is easily lost within one genus, and are Lirasyrinx Powell, 1942Parasyrinx Fin- developmental mode alone is not a character lay, 1924; Austroclavus Powell, 1942Splen- distinguishing genera. Marshall (1978:50; drillia Hedley, 1922; Micantapex Iredale, 1983:3) discussed the same question for the 1936 Harris & Burrows, 1891 (a generic classification of and long-accepted synonymy); and as noted above, , in which the same range of pro- Maoritomella Powell, 1942Tomopleura Casey, toconch types is encountered but, luckily, 1904, if indeed Tomopleura is appropriate for generic splitting has not occurred. He noted New Zealand species. (The type species of of Triphoridae that ‘planktotrophic and le- Tomopleura, T. nivea (Philippi, 1851) [Kilburn cithotrophic larval development occur through- 1986, figs. 6265; Sysoev in Poppe 2008:754, pl. out the family, often in the same genus or 627, figs. 8, 9; height 2629 mm] is larger and has species pair’. In some genera, such as New a much less well-defined sutural ramp and mid- Zealand species traditionally referred to Tomo- whorl angulation than are so obvious on species pleura, it has been realized for at least 20 years Tomopleura Cryptomella (initially pointed out to me by the late Phillip referred to . Finlay, Maxwell) that species with lecithotrophic pro- 1924 [type species: transenna Suter, toconchs (‘subgenus Maoritomella’ Powell, 1917, early Miocene, New Zealand] may be 1942) evolved by a number of different species required for New Zealand taxa previously in- cluded in Tomopleura and Maoritomella, other Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 losing planktotrophy independently. That is, Maoritomella is not a clade, but a grade, a than those transferred to by Maxwell group of unrelated species of Tomopleura with [2009]. This is one of many conoidean questions short, blunt protoconchs. Maxwell (2009:247) that will only be resolved by molecular studies.) has also convincingly transferred some species Some other synonymous generic ‘pairs’ were previously placed in this grade, along with pointed out by Bouchet (1990), and many others some placed by Powell (1942) in , recognized by Powell (1966) require synonymy. to Drilliola Cossmann, 1903. However, both he Some new generic placements are suggested here (Maxwell 2009:2478) and Spencer et al. for New Zealand NeogeneRecent Conoidea, (2009:209) catalogued both Tomopleura (Maor- although many other New Zealand Conoidea itomella) and Bathytoma (Micantapex) as still require re-evaluation. In particular, taxo- subgenera, when recognition of the solely nomic clarification of Neoguraleus sensu lato, developmental difference these ‘subgenera’ the most speciose and taxonomically ‘difficult’ of were based on indicates their synonymy. New Zealand conoidean genera, is a large task Much rationalization of conoidean genera is beyond the scope of this paper. 74 AG Beu

Family Turridae Petane (NMNZ M.27004, one); road cut, Subfamily Turrinae Puketapu, inland from Napier (GS5216, V20/ f8517, six); Castlepoint Fm, Castlepoint, E Genus Kuroshioturris Shuto, 1961 Wairarapa (GS10844, U26/f6037, one). Castle- cliffian: several early GNSlots labelled ‘Castle- Kuroshioturris Shuto 1961:83. Type species (by cliff, Wanganui’; Lower Castlecliff Shellbed original designation): Gemmula (Kuroshiotur- (OIS15) (GS4104,R22/f6518, re-collection, ris) hyugaensis Shuto, 1961, Pliocene to Recent, two); Upper Castlecliff Shellbed (OIS 11), Japan (Hasegawa, Okutani & Tsuchida in Castlecliff (GS4185, R22/f6502, three); re- Okutani 2000:627, pl. 312, fig. 40). corded also by Fleming (1953:199, 218, 226, 230) from Kaikokopu Shell Grit (OIS 19; Kuroshioturris angustata (Powell, 1940) (Fig. GS4134, R22/f7408, Kaikokopu Road, and 11A, K) GS4156, R22/f7417); Lower Castlecliff Pleurotoma (Hemipleurotoma) nodilirata Mur- Shellbed (OIS 15; GS4104, R22/f6518), Pinna- doch & Suter 1906:284, pl. 22, fig. 11 only (in cle Sand (OIS 13; in four collections on the part, Castlecliff specimen only; junior primary coast), and Tainui Shellbed (OIS 13; in three homonym of Pleurotoma nodilirata E. A. collections; GS10963, R22/f6619A, re-collec- Smith, 1878). tion of GS4013, two); NMNZ M.40275, cutting Micantapex angustatus Powell 1940:245, pl. 31, on Wainui Road, SW shore of Ohiwa Harbour, fig. 7; Powell 1942:56; Dell 1956a:130, pl. 19, Ohope, Bay of Plenty, OIS15? (13); M.158073, fig. 194; Fleming 1966:70. Herepuru Road, Matata, Bay of Plenty, OIS Lucerapex angustatus. Powell 1966:50, text-fig. 15? (14). Haweran: Landguard Sand (OIS 9), C49; Powell, 1979:228, fig. 53.1; Beu & Max- Landguard Bluff, opposite Wanganui City well 1990:418; Spencer & Willan 1996:29; (GS4003, R22/f7394, re-collection, three; Tucker 2004:72; Spencer et al. 2009:211. WRB Oliver collection, NMNZ, M.15715, ‘Lucerapex’ angustatus. Maxwell 2009:249. one); also recorded by Fleming (1953:273) from Waipuna Conglomerate (OIS7) at Land- Type material. Micantapex angustatus holotype guard Bluff, opposite Wanganui City (GS4001, AIM AK70470, Recent, from 48 m, off Wai- R22/f7392). Recent: RM72, 68 m, NW of

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 kuku Beach, North Cape (not seen). Cuvier I, Coromandel (one); RM78, 200 m, off Great Barrier Island, from the sample Other material examined. Mangapanian or ear- reported by Murdoch & Suter (1906) (two); liest Nukumaruan: beneath Pukenui Limestone RM4033, 90 m, off Doubtless Bay, Northland in large cutting on Tupurupuru Road, E of (four); 87 lots in NMNZ, ranging from (south- Gladstone, SWairarapa (NMNZ M.42870, one ernmost) off Oamaru, 55120 m (M.110910, typical), probably late Mangapanian rather one) and the Chatham Rise, through Cook than early Nukumaruan; Waihua River, 2.5 Strait as far W as Kapiti Island and Stephens km up Waihua Valley Road from Highway 2, 200 m N of sharp horseshoe bend in road, N Island, and up the east coast of the North Hawke’s Bay, early Nukumaruan (GS1409, Island as far N as North Cape (M.158067, E of W19/f8477, one). Nukumaruan: Petane, North Cape, 357447 m, one; M.158067, E of Hawke’s Bay, ex Suter collection (GNS, one); North Cape, 257327 m, one) and around as ‘Petane, cliffs opposite Eskdale Bridge’, that is, far W as Ahipara; most material is from the Mairau Mudstone, cliff at Esk River bridge, warm-water province between East Cape and junction of Highways 3 and 5 (GNS, one); North Cape. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 75

Distribution. Kuroshioturris angustata lives in a taller and narrower than the species considered wide range of depths (ca 30 300 m) all here, with either a completely smooth, polished around northern and eastern New Zealand. teleoconch surface apart from the peripheral There are no records from the southern South nodule row, or the rest of the teleoconch Island, around the Chatham Islands, around sculptured with thin, closely spaced axial la- the Three Kings Islands, or from the W coast of mellae. The New Zealand fossil and living either main island, other than in W Cook Strait species are not congeneric with these. and off Ahipara. It is most common on off- The illustration of a representative species shore, soft, fine-grained substrates, so its ap- of Kuroshioturris Shuto, 1961 by Powell parent scarcity in early Nukumaruan and early (1966:48, pl. 6, figs. 7, 8) has only weak nodules Castlecliffian rocks probably reflects this facies at the periphery, and does not appear to be preference. As a fossil, K. angustata has been related to ‘Micantapex’ angustatus. However, collected mainly from CastlecliffianHaweran the Japanese Recent species illustrated by rocks in Wanganui Basin and Nukumaruan Hasegawa et al. in Okutani (2000) have a single rocks in central Hawke’s Bay, although there row of prominent peripheral nodules, and are records also from late Mangapanianearly closely resemble M. angustatus: K. hyugaensis Nukumaruan rocks in Wairarapa and Castle- (Shuto, 1961), that is, the type species (Hase- cliffian rocks in the Bay of Plenty. gawa et al. in Okutani 2000:627, pl. 312, fig. 40); K. albogemmata Kuroda & Oyama in Dimensions. GS5216, Puketapu, Hawke’s Bay: Kuroda, Habe & Oyama (1971:221, pl. 57, H (incomplete) 14.8, D 6.3 mm; H 15.6, D 6.2 fig. 8; pl. 111, fig. 3; Hasegawa et al. in Okutani mm; GS4185, Upper Castlecliff Shellbed, Cas- 2000, pl. 312, fig. 38); K. nipponica (Shuto, tlecliff: H 12.8, D 4.6 mm; Castlecliff, early 1961) (Hasegawa et al. in Okutani 2000, pl. 312, collection: H 16.5, D 5.6 mm; Recent, NMNZ fig. 39); and K. kurodai (Makiyama, 1927) M.112114, Deepwater Cove, Bay of Islands, 55 (Hasegawa et al. in Okutani 2000, pl. 312, fig. m: H 13.3, D 4.8 mm; H 13.4, D 4.6 mm; 41). K. albogemmata and K. nipponica resemble NMNZ M.67615, W of Plate I, Bay of Plenty, M. angustatus particularly closely. The apex of 5964 m: H 15.8, D 5.3 mm; H 13.2, D 5.1 mm. the sinus trace forms the peripheral nodule row, Remarks. The generic position of ‘Micantapex’ as in Bathytoma, Gemmula and M. angustatus. angustatus has long been in doubt. It differs The protoconch was described by Powell (1966:48) as ‘rather large and composed of a Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 from species of Bathytoma (Micantapex)in its small size and much narrower shape, and so depressed, rounded and smooth first volution resembles several other genera, including Gem- and a convex and inflated second one, which is mula, at least superficially. Powell (1966) trans- smooth except for the last quarter whorl, with ferred it to Lucerapex Iredale, 1936, but the brephic axials’. However, this is clearly a type species (Pleurotoma casearia Hedley & lecithotrophic protoconch; at least K. albogem- Petterd, 1906, Recent, deep water off New mata and K. kurodai have a smaller, narrower South Wales) lacks all spiral sculpture other apex and are possibly planktotrophic. The than the peripheral nodulose carina formed by teleoconch shape, with a weak peribasal angu- the trace of the sinus apex, and instead has the lation, the presence or absence of a weak surface crossed by wavy axial lines (low, thin subsutural cord, the spiral sculpture of several axial lamellae). Species of Lucerapex recog- narrow cords, and the presence of one periph- nized in the New Caledonian fauna by the eral nodule row in K. albogemmata and K. MNHN informal ‘Turridae group’ (P Bouchet, nipponica agree exactly with M. angustatus. Y Kantor, RN Kilburn, A Sysoev; CD of Kuroshioturris appears to be the appropriate unpublished colour catalogue examined) are genus for M. angustatus. 76 AG Beu

The OpoitianWaipipian specimens as- Epidirona nodulosa Laseron (1954:9, pl. 1, figs. signed below to Kuroshioturris putere n. sp. 2123). The same is also true of some of the differ from Gemmula species in their much small, narrow New Zealand fossil species smaller size, shorter shape, much shorter ante- traditionally included in Bathytoma, such as rior siphonal canal, lack of a stromboid notch B. filaris (Marwick) (1931:134, pl. 15, fig. 279). in the lower margin of the outer lip, and Some of the smaller, more finely sculptured shallower posterior sinus at the peripheral species referred by Marwick (1931) to Gemmu- keel. It also has a simpler, low protoconch, la, such as G. peraspera (Marwick 1931:133, pl. which is quite unlike the protoconch of Gem- 15, figs. 276, 277), are also similar to Kuroshio- mula, described by Powell (1966:47) as ‘tall, putere n. sp. in their paucispiral proto- conical, polygyrate, and axially costate’, conch, narrowly triangular peripheral keel, and although that of Gemmula is probably plankto- few prominent spiral cords on the base of an trophic. K. angustata has a low, rounded, otherwise finely sculptured shell, and their smooth, mammillate protoconch of little more taxonomic position deserves reassessment. than 1.5 whorls (Fig. 11K), reflecting lecitho- There may well be still earlier occurrences of trophic development, and K. putere n. sp. this poorly known genus in New Zealand. apparently has an identical one. Powell I (Beu 1970b:232) followed Powell’s place- (1966:48) ranked Kuroshioturris as a subgenus ment of ‘Micantapex’ angustatus in Lucerapex of Ptychosyrinx Thiele, 1925 (Turrinae), but the when I suggested that ‘Micantapex’ pulcherri- shells illustrated by Hasegawa et al. in Okutani mus Vella, 1954 also belongs in Lucerapex (2000) show that it is a distinct genus, smaller (Tongaporutuan, Bell’s Creek bathyal fauna, and more weakly sculptured than Ptychosyrinx. near White Rock Road, SWairarapa). ‘ M.’ Powell also placed Lucerapex nearby, and pulcherrimus, with its numerous similar rows of illustrated the radula of ‘L.’ angustatus as large, sharp nodules, also resembles species of representing the genus Lucerapex, commenting Gemmuloborsonia Shuto, 1989. Sysoev & Bou- that ‘If angustatus ... is correctly assigned to chet (1996) revised the Pacific Recent species of Lucerapex, then the radula ...is very similar to Gemmuloborsonia, showing that it is a wide- that of acuta [(Perry, 1811)], con- spread Miocene to Recent bathyal turrine sisting of a pair of marginals, with the base genus, although it was included in a clade spread like a pair of calipers’ (Powell 1966:50, separate from Turrinae (sensu stricto) by Puil- text-fig. C49). The turrine radula indicates a landre et al. (2008). However, all species re-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 position in Turrinae (and in Kuroshioturris) for ferred to Gemmuloborsonia have a low, wide ‘M.’ angustatus, as compared with the clathur- columellar plait in at least some specimens, and elline radula described for Bathytoma by Powell this has not been observed in ‘M.’ pulcherrimus. (1966:64, as Micantapex). Neither Kuroshiotur- Species of Gemmuloborsonia also are much ris nor Lucerapex were studied but Bathytoma larger than ‘M’. pulcherrimus. These characters, is included in ‘’ clade 20 by therefore, suggest that ‘M.’ pulcherrimus should Puillandre et al. (2008). The radula, and pre- be referred to Kuroshioturris rather than Gem- ferably the DNA, of Lucerapex should be muloborsonia. Gemmuloborsonia species have a examined from a more typical species, to re- turrine radula (Sysoev & Bouchet 1996, fig. 1A) evaluate its subfamilial position, which is not unlike that of Kuroshioturris angustata, completely unknown at present. Some rela- rather than the clathurelline one that would tively small Recent Australian species that have been expected from the -like traditionally have been placed in Bathytoma teleoconch. More typical, unnamed species of or Epidirona also possibly belong in Kuroshio- Gemmuloborsonia are represented by material in turris, such as ‘Micantapex’ hecatorguia (Verco, GNSfrom middle Miocene deep-water siltstone 1907) (Cotton 1947, pl. 3, unnumbered fig.) and in Westland*a large, typical, robustly sculp- Marine Mollusca of isotope stages of the last 2 million years in New Zealand 77

tured species from Marsden-Kumara Road, lip slightly incomplete). Protoconch low, Clifdenian (GS4784, J32/f9586) and Eight- rounded, smooth, mammillate, of little more Mile Creek, Waiauan (GS4277, K32/f8521) than 1.5 whorls, with a large, subspherical and a smaller species from Alexander Street, initiation, apparently as in Kuroshioturris angu- Greymouth, Clifdenian (GS3159, J32/f7559). stata, but somewhat abraded on all specimens. Teleoconch of ca 6.5 whorls, spire relatively Kuroshioturris putere n. sp. (Fig. 11D, C, F) wide, rendered prominently gradate by strongly protruding peripheral keel at base of sutural Type material. Holotype TM8672, with three ramp, keel just above suture on most spire paratypes TM86735, GS1567, W19/f7516, whorls; suture descending on last whorl on some right bank Waiau River at junction with specimens, raising position of keel to mid-whorl Pakihiwi Stream (small side-stream to W), 600 height on paratype TM8673; base weakly con- m N of end of Steed forestry road and 1.3 km S vex below keel, tapering evenly and strongly of Ruapapa dam powerhouse, ca 5 km WNW into weakly concave neck and moderately long, of Cricklewood, ca 25 km straight-line distance gently tapered, incomplete but apparently un- WNW of Wairoa, N Hawke’s Bay; grid refer- notched anterior canal. Sutural ramp moder- ence W19/708424; Opoitian, with Stiracolpus ately wide, upper two-thirds steeply sloping, procellosus (Marwick, 1931), etc. almost flat; lower third deeply concave, curving out to form protruding peripheral keel. Sculp- Other material examined. Opoitian: stream in ture visually dominated by one peripheral row headwaters of Waihua River, 200 m N of Trig. O of prominent, narrow, sharply pointed, trian- no. 2, 800 m Sof junction of Putere and gular nodules formed by apex of trace of Cricklewood Roads, ca 10 km straight-line posterior sinus, otherwise predominantly of distance W of type locality (GS1567), Wairoa fine spiral cords; 20 nodules on last whorl and district, N Hawke’s Bay; grid reference W19/ 19 on penultimate whorl of holotype, 23 on last 619401; with Pelicaria parva, Zeatoma decens whorl and 22 on penultimate whorl of paratype (Marwick, 1931), Kaweka fulta Marwick, 1931, TM8673. Spiral sculpture commencing on first in a large fauna (one paratype, TM8676, teleoconch whorl as one prominent subsutural GS1576, W19/f7483). Waipipian: second-to-top row of moderately large triangular nodules and shellbed, with abundant Polinices waipipiensis one prominent peripheral row of larger trian- (Marwick, 1924), near head of E branch, Greek’s gular nodules, together occupying most of

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Creek, Sside of Arahura valley, Westland; grid whorl surface, all weakening down spire; sub- reference J33/547283; with Austrofusus pagoda sutural row gradually weakens to form 23 (Finlay, 1924), Lamprodomina, Clavatoma (one moderately prominent spiral cords (the lowest paratype, TM8677, GS12356, J33/f065). finely nodulose) over mid-spire, then weakens further to form wide zone of 34 low, narrow, Distribution. Recorded at present only from closely spaced spiral cords slightly elevated Opoitian rocks of N Hawke’s Bay and Waipipian above rest of ramp, with scarcely any nodules, rocks of Greek’s Creek, Arahura valley, Westland. on last two whorls of holotype, fading out altogether on some paratypes; 45 fine, closely Dimensions. Holotype: H 13.7, D 6.3 mm; spaced spiral threads develop on upper part of paratype TM8673: H 14.0, D 6.6 mm; paratype, wide, strongly concave sutural ramp, lower TM8676: H (incomplete) 12.7, D 5.9 mm. third almost smooth; three very narrow, closely spaced spiral cords develop around peripheral Description. Shell small (H 14 mm), tall and nodule row on last 23 teleoconch whorls, moderately slender, with tall spire and relatively uppermost very faint, central most prominent; short last whorl (siphonal canal apex and outer two prominent, narrow spiral cords on upper 78 AG Beu

part of base, upper cord margining suture and the last few whorls than in K. angustata, lower appearing from suture on holotype, upper scarcely raised at all in some specimens. The cord appearing from suture on some paratypes; base also has only two moderately prominent 13 lower, narrower, closely spaced, very fine spiral cords, narrow and quite closely spaced, spiral threads above and between two promi- high up below the peripheral angulation, and nent cords on base, 1215 further fine threads the remainder of the base and the sutural ramp below, fading out over siphonal canal. Axial bear fine, closely spaced spiral threads in sculpture of growth lines only, apart from K. putere n. sp., whereas there are three peripheral nodules. Aperture unthickened, with- prominent, rather more widely spaced cords out parietal callus, inner lip slightly excavated in K. angustata. The result is an overall into previous whorl; outer lip thin and simple, appearance that is more finely sculptured, without stromboid notch; profile of outer edge although with a slightly shorter spire, and of outer lip biarcuate, moderately convex over with a sharper and more narrowly defined sutural ramp, forming narrowly rounded poster- peripheral keel in K. putere n. sp. than in ior sinus at peripheral nodule row, curving K. angustata. Paratype TM8676 (GS1576, Wai- forwards again moderately below ramp. River, 800 m Sof junction of Putere and Cricklewood Roads, Wairoa district) has a Remarks. Kuroshioturris angustata has been complete outer lip, including the sinus apex thought of as a late Pliocene (Nukumaruan) flared outwards at the peripheral keel as is seen to Recent species only (Beu & Maxwell in many Recent specimens of K. angustata, and 1990:418) and so, potentially, Kuroshioturris is provided the lip description included above. a group that arrived in New Zealand late The similarity in size, shape, sculpture, sinus in Pliocene time. However, a similar form, shape and protoconch leave little doubt that K. putere n. sp., a little wider, more prominently K. putere n. sp. is closely related to K. and sharply nodulose at the periphery than angustata, and possibly was its direct ancestor; K. angustata and with only two prominent the change seems to have been an abrupt basal spiral cords, is now recorded from speciation event. However, the specimen of OpoitianWaipipian rocks. The specimens K. angustata listed above from Waihua River, from N Hawke’s Bay were included in the N Hawke’s Bay (GS1409, W19/f8477, Manga- faunas reviewed by Marwick (1965), but were panian or early Nukumaruan) has a slightly not described by him, and labels on some of the shorter spire and more triangular peripheral

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 specimens show that he was undecided whether nodules than in most other specimens of K. they belonged in Gemmula or ‘Micantapex’, angustata, although it agrees with K. angustata although some are clearly labelled as related to in having the subsutural fold prominent and ‘Micantapex’ cf. angustatus. K. putere n. sp. has nodulose down onto the last whorl, and in a slightly narrower and more strongly protrud- having more numerous spiral cords on the base ing peripheral keel than in K. angustata, than in K. putere n. sp. The late Miocene producing a slightly wider and more strongly (Tongaporutuan) bathyal species K. pulcherri- concave sutural ramp. The peripheral nodule ma differs from K. putere n. sp. in having three row also bears three narrow spiral cords more prominently nodulose spiral cords below the obviously in K. putere n. sp. than in K. peripheral keel and in retaining a prominently angustata, and as the central cord protrudes nodulose subsutural fold onto the last whorl, so the most, the keel is raised into a row of that its obvious sculpture consists of five rows pointed, spine-like nodules rather than the of similar, triangular nodules, whereas other lower, subquadrate nodules of K. angustata. species suggested above to be potential mem- Also, the subsutural fold of K. putere n. sp. is bers of Kuroshioturris, such as Bathytoma considerably lower and less well defined over filaris, are more similar to K. putere n. sp., Marine Mollusca of isotope stages of the last 2 million years in New Zealand 79

differing in their more even, fine spiral sculp- 1996:29; Tucker 2004:1072; Spencer et al. ture and less prominent peripheral nodules. 2009:211; Maxwell 2009:249. Aoteadrillia wanganuiensis chordata. Powell Etymology. The species name is from the type 1942:92; Boreham 1959:59; Powell 1966:86, pl. locality, near Putere-Cricklewood Road in N 13, fig. 7; Fleming 1966:73; Tucker 2004:199. Hawke’s Bay; a noun in apposition. Aoteadrillia bisecta Powell 1942:92, pl. 1, fig. 6; Fleming 1966:73. Aoteadrillia chordata. Beu & Maxwell 1990:419; Subfamily Spencer et al. 2009:211; Maxwell 2009:249.

Genus Aoteadrillia Powell, 1942 Type material. Pleurotoma wanganuiensis, two Aoteadrillia Powell 1942:87. Type species (by syntypes TM83067, from ‘Shakespeare Cliff’, original designation): Pleurotoma wanganuien- Wanganui; almost certainly from Tainui sis Hutton, 1873, (Mangapanian?) Nukumar- Shellbed (OIS 13), which formerly cropped uanRecent, New Zealand. out at Shakespeare Cliff, within Wanganui City. The slightly larger syntype (TM8306) Aoteadrillia wanganuiensis (Hutton, 1873) was labelled ‘lectotype’ when received from (Figs. 12AG, J, Fig. 13AG, J) NMNZ many years ago, but this unpublished designation has no status; the two syntypes are Pleurotoma wanganuiensis Hutton 1873b:4; conspecific and both are well preserved. Drillia Hutton 1893:50, pl. 6, fig. 28; Harris 1897:46. chordata, lectotype (designated by Boreham Drillia chordata Suter 1908:184, pl. 7, fig. 16; 1959:59, as ‘paratype’ in error) TM1109, with Suter 1913:475, pl. 21, figs. 5, 5a (new synonym). three paralectotypes, TM1110TM1112, ex Drillia wanganuiensis. Suter 1914:29, pl. 2, Suter collection, Recent, dredged off Otago fig. 13. Heads. alpha, holotype TM8308 Austrodrillia alpha King 1933:349, pl. 36, fig. 6; (Fig. 12F), from Onoke Formation (early Beu 1967:114. Nukumaruan), cliffs E of Lake Ferry, Palliser Austrodrillia beta King 1933:349, pl. 36, fig. 7. Bay, SWairarapa. Austrodrillia beta, holotype Austrodrillia gamma King 1933:350, pl. 36, fig. 8. TM8309 (Fig. 12D), from the same locality. wanganuiensis. Powell 1934a:264. Austrodrillia gamma, holotype TM8310 (Fig.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Aoteadrillia gamma. Powell 1942:90; Fleming 12B), from the same locality. Aoteadrillia 1966:73. thomsoni, holotype AIM AK70914, with one Aoteadrillia beta. Powell 1942:90; Fleming paratype AK71790, from E shore of Lake 1966:73. Grassmere, Marlborough (Nukumaruan?); Aoteadrillia alpha. Powell 1942:90; Fleming two paratypes TM8315TM8317 (Fig. 13A, 1966:73; Beu 1979:88; Beu & Maxwell C), from type locality, collected by JA Thom- 1990:418. son; four paratypes TM8318-TM8321, from Aoteadrillia thomsoni Powell 1942:90, pl. 2, fig. GS2329, U23/f6381, W-facing hillside 0.5 km 7; Fleming 1966:73; Beu & Maxwell 1990:419; N of Whetukura, 4 km SE of Ormondville, Maxwell 2009:249 (new synonym). Dannevirke district, SHawke’s Bay, probably Aoteadrillia trifida Powell 1942:91, pl. 12, fig. Mangapanian (below uppermost Te Aute lime- 13; Fleming 1966:73; Beu & Maxwell 1990:419, stone, that is, Te Onepu Limestone; Beu 1995). Maxwell 2009:249 (new synonym). Aoteadrillia trifida, holotype TM8311 (Fig. Aoteadrillia wanganuiensis. Powell 1942:91; 12G), from GS2624, S27/f8463, Makara- Fleming 1966:73; Beu & Maxwell 1990:367, Mangaopari River junction, White Rock pl. 49p; Powell 1979:231; Spencer & Willan Road, SWairarapa; Pukenui Limestone, early 80 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 12 (AG,J) Aoteadrillia wanganuiensis (Hutton), type species of Aoteadrillia Powell, 1942; SEM; A, paratype of Aoteadrillia bisecta Powell, TM8312, GS2610, S27/f8449, Makara River, S Wairarapa, early Nukumaruan; height 14.2 mm; B, holotype of Austrodrillia gamma King, TM8310, Onoke Formation (early Nukumaruan), cliffs E of Lake Ferry, Palliser Bay; height 9.8 mm; C,E,J, GS10963, R22/f6619A, 2 specimens, probable topotypes of A. wanganuiensis, Tainui Shellbed (Castlecliffian, OIS 13), ‘‘the buttress’’, Castlecliff; C, height 12.2 mm; E, height 12.2 mm; J, protoconch of C; D, holotype of Austrodrillia beta King, TM8309, Onoke Formation (early Nukumaruan), cliffs E of Lake Ferry, Palliser Bay; height 12.8 mm; F, holotype of Austrodrillia alpha King, TM8308, Onoke Formation (early Nukumaruan), cliffs E of Lake Ferry, Palliser Bay; height 12.8 mm; G, holotype of Aoteadrillia trifida Powell, TM8311, GS2624, S27/f8463, junction Mangaopari Stream and Makara River, S Wairarapa, early Nukumaruan; height 8.8 mm. (H,I) Aoteadrillia apicarinata (Murdoch & Suter), GS10858, U22/f09, Ashcott Road, NW of Pukeora hill, W of Waipukurau, Hawke’s Bay, Nukumaruan, SEM; H, whole specimen, height 11.1 mm; I, protoconch of H. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 81 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 13 (AG,J) Aoteadrillia wanganuiensis (Hutton), SEM; A, C, 2 paratypes of Aoteadrillia thomsoni Powell, GS2329, U23/f6381, Takapau Survey District, Dannevirke, early Nukumaruan; A, TM8318, height (incomplete) 8.5 mm; C, TM8319, height 11.4 mm; B,E,G, 3 specimens, GS10337, Q32/f8843, bathyal early Nukumaruan fauna, hill above Glenstrae woolshed, Oaro, Marlborough; B, height 12.3 mm; E, height 12.0 mm; G, height 8.0 mm; D,F,J, ‘‘chordata form’’, 2 specimens, Recent, RM5589, Portobello sta. Mu7045, 540490 m, Papanui Canyon, off Otago Peninsula; D, height 10.0 mm; F, height 10.2 mm; J, protoconch of F. (H) Aoteadrillia waihuaensis Powell, holotype, GS1560, W19/f8480, TM8698, Waihua River upstream from Stream, N Hawke’s Bay, Waipipian-Mangapanian; H 8.1, D 3.1 mm. (I,K) Aoteadrillia finlayi Powell, GS11465, V21/f6015, Mairau Mudstone (late Nukumaruan), ‘‘the Watchman’’ hill, Napier; I, height 18.3 mm; K, protoconch of I. 82 AG Beu

Nukumaruan. Aoteadrillia bisecta, holotype Dimensions. GS2610, S27/f8449, TM8312, AIM AK70911, from Petane, central Hawke’s paratype of Austrodrillia bisecta, Makara Riv- Bay, late Nukumaruan; three paratypes TM8 er, SWairarapa: H 14.1, D 5.1 mm; GS2329, 312TM8314 (Fig. 12A), from GS2610, S27/ U23/f6381, TM8318, paratype of Aoteadrillia f8449, Makara River, 1 km upstream from thomsoni, Whetukura, SHawke’s Bay: H 12.7, junction with Ruakokopatuna River, SWair- D (slightly incomplete) 4.9 mm; GS11465, V21/ arapa; Pukenui Limestone, early Nukumaruan. f6015, Mairau Mudstone, ‘the watchman’ hill, Ahuriri, Napier, Nukumaruan (occurring with Other material examined. Not listed; abundant A. finlayi): H 11.7, D 4.3 mm; H 11.5, D 4.3 and widespread. mm; GS10337, O32/f8843, hill above Glenstrae Station, Oaro, Marlborough, Nukumaruan: H Distribution. Aoteadrillia wanganuiensis is com- 14.8, D 4.9 mm; H 12.5, D 4.3 mm; H 11.9, D mon and widespread in Nukumaruan and 4.3 mm; GS5833, Y14/f7505A, Te Piki, near Castlecliffian rocks wherever they occur in East Cape, OIS7: H 10.8, D 3.9 mm; H 10.1, D New Zealand, in North Canterbury, Marlbor- 3.8 mm; Recent, RM5589, 540490 m, Papanui ough, Wairarapa district, Hawke’s Bay and Canyon, E Otago: H 9.7, D 3.6 mm; H 10.5, D Wanganui Basin, as well as Castlecliffian and 3.7 mm; H 11.1, D 3.7 mm; Austrodrillia alpha Haweran rocks in the Bay of Plenty (Matata, King, holotype: H 12.8, D 4.5 mm; Austrodrillia Ohope, Te Piki) and Castlecliffian rocks at beta King, holotype: H 12.9, D (incomplete) 4.5 Cape Kidnappers (Fleming in Kingma mm; Austrodrillia gamma King, holotype: H 1971:98). At most localities where faunas are 9.3, D 4.0 mm; Aoteadrillia trifida Powell, preserved that lived in shallow shelf environ- holotype: H 9.0, D 3.3 mm. ments on soft substrates it is much the most common conoidean found, although in still Remarks. My experience of biostratigraphy of shallower water Neoguraleus species are more MangapanianNukumaruan rocks in Wairara- common. Coarsely sculptured forms living in pa and North Canterbury suggests that cooler water than more finely sculptured speci- gradually changing forms of Aoteadrillia wanga- mens at present indicates that the various nuiensis have a low-level utility for biostratigra- intergrading named forms are of little biostra- phy. Extremely coarse variants are characteristic tigraphical utility, except for the dominance of of early Nukumaruan time. However, this coarsely sculptured forms resembling Drillia apparently is ecologically (probably tempera-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 chordata in early Nukumaruan rocks of Wair- ture-) controlled variation, and all the named arapa and North Canterbury. Recent speci- forms listed in the synonymy are part of the mens in NMNZ occur from the Otago fiords variation of one species. I have suggested most of (Dusky Sound, Thomson Sound, Doubtful the synonymy previously (Beu 1967:114). Sound) and the Otago shelf and canyons north The first point is to separate Aoteadrillia to New Plymouth on the west coast and to the finlayi from this complex. I wrongly included it in Bay of Plenty on the east coast. There are no the synonymy of A. wanganuiensis previously records in NMNZ from the southernmost (Beu 1967). A. finlayi is easily separated from all , Stewart Island, the Chatham other similar forms by its markedly larger size Islands, or the northern North Island north of and narrow shape, as though it added another the central Bay of Plenty. However, a single 12 whorls to its height compared with all other empty shell is present among BW Hayward’s Aoteadrillia species without increasing its dia- dredged molluscs from Port Pegasus, Stewart meter, by its consistently very fine sculpture of Island (GNSRM6061), suggesting that it is two or more closely spaced, low, narrow spiral rare near the extremities of its range rather than cords around the periphery crossing 2024 fine, completely absent. close axial ridges, by its finer spiral sculpture on Marine Mollusca of isotope stages of the last 2 million years in New Zealand 83

the base, with narrow secondary threads be- Petane, Hawke’s Bay (late Nukumaruan) and tween the primary cords, and by its slightly taller from GS1590, O32/f8039, near Leader River protoconch with a slightly more prominent keel bridge, Hawkeswood, North Canterbury (early on the second whorl. The keeled protoconch led Nukumaruan). Evidently he was unable to Marwick (1965:43, pl. 11, fig. 18) to assign an distinguish between Nukumaruan and Recent apical fragment of A. finlayi from GS2818 (W19/ coarsely sculptured variants, as I also conclude f8494, highway above Waihua rail tunnel, north- here. Powell (1979:231) later considered ern Hawke’s Bay; Nukumaruan, with rare the chordata form to be part of the variation Struthiolaria frazeri and the convexa form of of A. wanganuiensis. The variation of extremes Pelicaria vermis)toParacomitas, but the two of the chordata form led to the recognition of fine, closely spaced, peripheral spiral cords on the ‘species’ A. alpha (King, 1933) (Fig. 12F), the teleoconch spire whorl fragment demon- A. beta (King, 1933) (Fig. 12D), A. gamma strate that his specimen is A. finlayi. A. finlayi (King, 1933) (Fig. 12B), A. thomsoni Powell, apparently is limited to Nukumaruan rocks, and 1942 (Fig. 13A, C), A. trifida Powell, 1942 (Fig. I have seen specimens definitely assigned here 12G) and A. bisecta Powell, 1942 (Fig. 12A). only from late Nukumaruan (Petane Group) Specimens resembling the holotype of Drillia rocks of central and northern Hawke’s Bay. chordata, from off Otago (Fig. 13D, F), are in However, specimens from the Recent bathyal fact among the more weakly sculptured of the fauna and bathyal rocks in SWairarapa also are variants of A. wanganuiensis, and can be very similar to A. finlayi (see below). matched in most large collections of Nukumar- Powell (1942:92) discussed ‘the species uan and Castlecliffian age from Wanganui and grouped around [Aoteadrillia] wanganuiensis’, central Hawke’s Bay, particularly by specimens describing ‘the lengthening of the spire’ with of the form Powell (1942) called A. bisecta.I increasing age, and recognizing several of the have no doubt that the chordata form inter- more extreme sculptural variants as ‘divergent grades completely with A. wanganuiensis. Nukumaruan forms’. Unfortunately, he al- Forms such as that named Austrodrillia beta lowed little variation within one species. In by King (1933) (Fig. 12D) are considerably particular, the significance of the form named more extreme, almost resembling a Paracomitas Drillia chordata by Suter (1908:184, pl. 7, fig. species in their prominent peripheral row of 16) has troubled taxonomists since it was smooth, narrowly rounded, obliquely angled proposed. Although Powell (1942:92) ranked nodules, but again intergrade completely with

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 this as a subspecies of A. wanganuiensis apply- ‘typical’ A. wanganuiensis through many of the ing only to Recent specimens collected off Hawke’s BayWairarapa early Nukumaruan Otago, he noted that ‘two forms of wanga- specimens. The immediate ancestor of this nuiensis occur throughout its [time] range, the complex seems likely to have been the Waipi- typical species having long axials crossed by pian (Mangapanian?) species A. waihuaensis three to four spiral cords, and the second form Powell (1942:91, pl. 12, fig. 14; Fig. 13H), chordata, with shorter and stronger axials which is smaller and more finely sculptured forming a double peripheral series of strong than all the NukumaruanRecent forms, with rounded nodules’. Specimens from deeper narrow peripheral nodules similar to those of water off Otago (bathyal material, from the the holotype of Austrodrillia beta and some heads of the Otago canyons) also consist mostly paratypes of A. thomsoni and A. trifida. A. of the chordata form, although a few specimens consequens (Laws 1936:121, pl. 17, figs. 76, 78) of the ‘typical’ finely sculptured form occur from Kaawa Creek, SW Auckland (Opoitian, also. Despite limiting the name to Recent early Pliocene) is still narrower and more Otago specimens, Powell (1942) also recorded strongly monocarinate, resembling Kuroshio- specimens as A. wanganuiensis chordata from turris angustata, and makes a plausible link 84 AG Beu

between the PlioceneRecent species related to Pliocene sea-temperature change in New Zeal- A. wanganuiensis and the early Miocene (Alto- and (Fleming 1944; Beu et al. 1977; Beu 1985, nian) species A. callimorpha (Suter, 1917), the 1995). As well as Z. delicatula, other southern earliest species of Aoteadrillia named at present. cool-water molluscs such as Cominella nas- Powell (1942) named still further sculptural soides and the subantarctic spider crab Jacqui- variants of A. wanganuiensis as A. thomsoni notia edwardsii (Jacquinot in Jaquinot & Lucas, and A. trifida, both of which have several 1853) (Yaldwyn & Beu, appendix in Beu et al. narrow peripheral spiral cords. The form he 1977) were dispersed as larvae in the proto- named A. bisecta has a row of massive axial Southland Current to the central and southern folds around the periphery, subdivided to North Island at 2.4 Ma, in the first strongly varying degrees by a median spiral groove, marked evidence of Pleistocene cooling in New but intergrades through decreasing nodule Zealand marine faunas. The abrupt appearance strength with the chordata form. of this cool fauna in Wanganui Basin is used to Late Nukumaruan specimens in Hawke’s identify the base of the Nukumaruan Stage Bay are mostly typical, finely sculptured Aotea- (Beu in Cooper 2004). I suggested (Beu 2010: drillia wanganuiensis, although they occur with 102112) that a similar pattern was followed by the confusingly similar but markedly taller and Pelicaria vermis, in which the variation became slightly more finely sculptured species A. fin- extreme from early Nukumaruan time on, layi. At Castlecliff, Wanganui, in siltstone indicating that this species evolved direct devel- formations of the Castlecliffian coastal section, opment*and, consequently, now has a paucis- only the typical form of A. wanganuiensis piral protoconch*at about 2.4 Ma, at the occurs (Fig. 12C, E). The A. alpha-beta-gam- beginning of Nukumaruan time. A similar ma-bisecta-trifida range of variants, with mark- interpretation was also suggested (Beu 2010: edly coarser sculpture, seems to represent an 9192) for the highly variable species Stiracolpus early Nukumaruan excursion by A. wanga- symmetricus. Aoteadrillia wanganuiensis fol- nuiensis into greater sculptural variation that lowed a similar pattern. However, the proto- was later reversed. The complete intergradation conch type remained constant in Aoteadrillia between the finer and coarser sculptural ex- throughout its history, so apparently it did not tremes makes it impossible to recognize sepa- undergo any fundamental change of develop- rate species, but the recognition of coarsely mental type. A reasonable explanation seems to sculptured forms limited to early Nukumaruan be that Aoteadrillia was lecithotrophic, with a

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 rocks has definite biostratigraphical utility. relatively brief demersal larval life, and merely The most significant connotation of the shortened the period before metamorphosis Recent distribution of the ‘chordata form’ during earliest Nukumaruan time. I conclude coarse variants recognized by Powell that the coarsely sculptured forms of Aoteadril- (1942)*limited to eastern Otago, among the lia wanganuiensis are cool-water variants of no most southern of present-day records of the taxonomic significance, and I regard the many species*is that fossils of similar but even more trivial forms recognized by Powell (1942) as part coarsely sculptured forms of Aoteadrillia wan- of the variation of A. wanganuiensis. ganuiensis in early Nukumaruan rocks of Wairarapa to North Canterbury reflect cool Aoteadrillia finlayi Powell, 1942 (Fig. 13I,K) sea temperatures at the deposition site. The spread of the cool-water Zygochlamys delicatu- Aoteadrillia finlayi Powell 1942:93, pl. 1, fig. 5; la community northwards to form a major Fleming 1966:73; Beu & Maxwell 1990:419; element of the early Nukumaruan fauna of Tucker 2004:378; Maxwell 2009:249. Wairarapa and North Canterbury has long Paracomitas n. sp. Marwick 1965:43, pl. 11, been a helpful concept for understanding late fig. 18. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 85

Aoteadrillia alpha. Beu 1967:114 (in part not A. 17 km E of Alderman Islands (two); M.60222, alpha King, 1933). 660765 m, 35 km WNW Orete Point, Cape Runaway (one; large peripheral nodules, Type material. Aoteadrillia finlayi, holotype doubtfully referred); M. 61112, 127134 m, 17 AIM AK70912, with one paratype AK71786, km E of Portland Island, Mahia Peninsula from ‘Petane corner’, cliff at junction of High- (one); M.60343, 434446 m, 41 km SE of ways 3 and 5, Sof Esk River bridge, 20 km N of Cape Campbell (two). Napier, central Hawke’s Bay; Mairau Mud- stone, late Nukumaruan (not seen). Distribution. I am aware of specimens definitely referred to Aoteadrillia finlayi only from late Other material examined. Nukumaruan: high- Nukumaruan rocks of central and north-central way above Waihua rail tunnel, between Wai- Hawke’s Bay. However, the list above also hua and Wairoa valleys, N Hawke’s Bay includes a form that seems indistinguishable (GS2818, W19/f8494, one spire apex; the basis from A. finlayi (pointed out to me by BA of Marwick’s [1965:43, pl. 11, fig. 18] record of Marshall, NMNZ) occurring in bathyal Man- ‘Paracomitas n. sp.’); ‘Petane corner’, junction gapanian and Nukumaruan rocks, and in bath- Highways 3 and 5, Esk River bridge, N of yal depths in the Recent fauna from Mayor Napier, CR Laws collection (GNS, one); lo- Island, N Bay of Plenty, to Cape Campbell, cality as last (GS10937, V20/f8500A, three); Marlborough (it is not strictly bathyal; M.61112 Maharakeke Mudstone (of Thomson 1926), is from 127134 m). Some specimens seem Maharakeke Road, W foot of Pukeora Hill, indistinguishable from Hawke’s Bay Nukumar- Waipukurau (GS10857, U23/f7049, three); Ma- uan ones, and this form tentatively is referred to karetu Mudstone (of Thomson 1926), Ashcott A. finlayi. This species apparently is still living Road, N foot of Pukeora Hill, Waipukurau along eastern New Zealand, so the variation (GS10858, U22/f09, 12); Devils Elbow Mud- and time range of A. finlayi require further stone, top of Devils Elbow hill, Highway 2, N study as more material becomes available. of Napier (GS10849GS10936, V20/f8555, 46); Darkys Spur Formation, Darkys Spur Dimensions. GS10849, Devil’s Elbow: H 20.4, D road, between Waikawau and Devils Elbow 6.5 mm; H 19.3, D 6.0 mm; H 19.0, D 6.3 mm; H (GS11225, V20/f8019, one); Mairau Mudstone, 18.7, D 6.1 mm; H (slightly incomplete) 18.9, D ‘the watchman’ hill, Ahuriri lagoon, Napier 6.2 mm; H 18.3, D 6.5 mm; GS11465, Mairau

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (GS11465, V21/f6015A, 21); Mairau Mud- Mudstone, ‘the watchman’ hill, Ahuriri, Napier stone, walkway W side of Roro-o-kuri, One- (occurring with A. wanganuiensis): H 18.6, D hunga Road, N end of Ahuriri ‘lagoon’, Napier 6.0 mm; H 15.9, D 5.8 mm; H 14.6, D 5.6 mm; H (GS12386, V21/f05, 74 specimens). (slightly incomplete) 14.2, D 5.4 mm. Bathyal form referred tentatively to Aotea- drillia finlayi: Mangapanian: NMNZ M.42867, Remarks. Aoteadrillia finlayi is distinguished road cut 1 km NW of Tupurupuru-Westmere above from A. wanganuiensis. Specimens recog- Road junction, SWairarapa, Mangapanian nized by BA Marshall in bathyal faunas (S (underlying Pukenui Limestone) (14). Nuku- Wairarapa, and in the Recent bathyal fauna maruan: NMNZ M.147776, cliffs E of Lake from N Bay of Plenty to Cape Campbell) Ferry, Palliser Bay (one). Recent: NMNZ M. apparently are conspecific with A. finlayi. This 12901, 494 m, off Mayor Island (three); indicates the possibility that A. finlayi had a M.60246, 482550 m, 17 km E of Mayor Island wide bathymetric range (as A. wanganuiensis (one); M. 118497, 357312 m, SE of Alderman does), still lives around New Zealand in the Islands (one; closely similar to Hawke’s Bay bathyal zone, and possibly retreated there after Nukumaruan fossils); M.61150, 443527 m, Nukumaruan time. 86 AG Beu

Aoteadrillia apicarinata (Marshall & Murdoch, ities 13 in Nukumaruan rocks of Hawke’s Bay 1923) (Fig. 12H,I) (1: walkway W side of Roro-o-kuri, Onehunga Road, Bay View, N of Napier; Mairau Mud- Drillia apicarinata Marshall & Murdoch stone, late Nukumaruan; 2: ‘the watchman’ hill, 1923:125, pl. 17, figs. 76, 78. in the former Ahuriri Lagoon, W of Napier Aoteadrillia apicarinata. Powell 1942:90; Flem- airport, also in Mairau Mudstone; 3: Ashcott ing 1966:73; Powell 1966:87; Beu & Maxwell Road, that is, the locality where A. apicarinata 1990:418; Richardson 2002:12; Tucker 2004:80; is recorded here as moderately common). Maxwell 2009:249. Distribution. Limited to late Nukumaruan Type material. Drillia apicarinata, holotype rocks of Hawke’s Bay. TM8302, from Waikopiro, near Ormondville, southern Hawke’s Bay, late Nukumaruan, ex H Dimensions. Drillia apicarinata, holotype: H Suter collection. The illustration of Drillia (incomplete) 8.6, D 3.3 mm; GS10858, Ashcott apicarinata by Marshall & Murdoch (1923, pl. Road: H 11.1, D 4.0 mm (Fig. 32A,B). 12, fig. 5) shows an almost complete specimen, and the height is stated to be 10 mm. The Remarks. Aoteadrillia apicarinata is a very dis- specimen in GNSidentified as the holotype has tinctive small species, with even, narrow, promi- the anterior end (siphonal canal and anterior nent, closely spaced spiral cords over the entire half of the last whorl) broken away, and a note surface below the sutural ramp, and a few fine on the label by CA Fleming states ‘canal noted spiral cords on the ramp, as well as a prominent broken’ on 10 February 1947 (when the Mar- subsutural spiral ridge (as in all other Aotea- shall & Murdoch collection was incorporated in drillia species) composed of one or two spiral the NZGScollection by CA Fleming). As cords. Moderately prominent, rounded axial Marshall & Murdoch (1923) stated that they ridges begin abruptly at the border of the ramp had only one specimen, the GNSspecimen (the shoulder angle), and extend well down the labelled ‘type’ is the holotype. last whorl, without forming nodules at the intersections with the spiral cords; they are Other material examined. Nukumaruan: the much fewer and more widely spaced than those distinctive species Aoteadrillia apicarinata is of A. wanganuiensis. The protoconch is unusual, reasonably common at only one locality I am as it blends gradually into the teleoconch, and

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 aware of, the former road cutting on Ashcott the spiral sculpture begins early on the first Road (abandoned by the 1980s deviation of protoconch whorl and passes onto the teleo- Highway 2) immediately NW of Pukeora Hill, conch unchanged. The initial protoconch half- SW of Waipukurau, Hawke’s Bay (GS10858, whorl is low, superficially smooth and rounded, U22/f09, four). The type locality, ‘Waikopiro’, but actually bearing numerous fine spiral threads E of Ormondville in SHawke’s Bay, seems not (Fig. 12H) as in A. wanganuiensis (Figs. 12J, 13J), to have been re-collected during the last 80100 and the prominent, median, sharply defined keel yr. Powell (1942:90) also recorded material commences gradually over the next quarter- from ‘inner harbour, Napier’, that is, cliffs of whorl; one or two spiral threads develop gradu- the former Ahuriri Lagoon, in Mairau Mud- ally below the keel over the succeeding half- stone, high in the Petane Group (late Nuku- whorl, so after one whorl all specimens display maruan), presumably based on specimens in the some spiral sculpture, and it simply continues to Finlay collection, as I have seen no material of strengthen and to develop more cords as the A. apicarinata from this area. Richardson shell grows. It is impossible to identify the (2002:12) also listed specimens from his local- protoconch-teleoconch junction, indicating Marine Mollusca of isotope stages of the last 2 million years in New Zealand 87

that A. apicarinata had lecithotrophic develop- 1966:30, pl. 2, figs. 2, 3; Fleming 1966:73; Powell ment. Its restriction to a few localities in Hawke’s 1969:298, pl. 234, figs. 2, 3; Grant-Mackie & Bay suggests the possibility that development Chapman-Smith 1971:680; Powell 1979:231; was direct. A. apicarinata does not resemble A. Spencer & Willan 1996:29; Tucker 2004:596; wanganuiensis (type species of Aoteadrillia) very Spencer et al. 2009:211. closely but, as pointed out by Powell (1942:90), it Antimelatoma ahiparana Powell 1942:98, pl. 2, is quite similar to the early Pliocene (Opoitian) fig. 11; Powell 1966:30; Powell 1979:231, pl. 45, species A. consequens. Evidently the genus Ao- fig. 14; Spencer & Willan 1996:29; Tucker teadrillia consists of at least two discrete lineages 2004:40; Spencer et al. 2009:211 (new synonym). that evolved from the early Miocene species A. Antimelatoma benthicola Powell 1942:98, pl. 2, callimorpha, culminating respectively in A. api- fig. 10; Powell 1966:30; Powell 1979:232, pl. 45, carinata and A. wanganuiensis. fig. 12; Spencer & Willan 1996:29; Tucker 2004:120; Spencer et al. 2009:211 (new synonym). Genus Antimelatoma Powell, 1942 Antimelatoma n. sp. Fleming 1953:245. Antimelatoma maorum. Maxwell 2009:249. Antimelatoma Powell 1942:97. Type species (by original designation): Drillia maorum E. A. Smith, 1877 (Pleurotoma buchanani Hutton, Type material. Pleurotoma buchanani type 1873), late PlioceneRecent, New Zealand. material consists of two syntypes (TM86289) formerly glued to one wooden tablet, which Antimelatoma buchanani (Hutton, 1873) (Fig. is labelled ‘Drillia buchanani (Hutton) Type. 14BF, H) (Suter, 1912). (Type of Pleurotoma buchanani Hutton, 1873). Shakespeare Cliff, Wang.’. The Pleurotoma buchanani Hutton 1873b:4; Hutton specimens are labelled lectotype and paralecto- 1880:42; Hutton 1893:50, pl. 6, fig. 26; Harris type, but I am not aware of any publication of 1897:47. this designation. Hutton (1873b:4) cited speci- Pleurotoma (Drillia?) maorum EA Smith mens from ‘Shakespeare Cliff’ and ‘Awamoa’, 1877:497 (new synonym). but of course only the former two syntypes Drillia buchanani. Hutton 1878:16; Suter belong in this species. Hutton’s Awamoa speci- 1914:29. men is no longer identifiable. The Wanganui Drillia maorum. Hutton 1880:44; Hutton specimen (TM8628; Fig. 14B) that was for-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 1885b:116. merly glued by its dorsum (so the aperture does Surcula trailli. Suter 1899b:68; Suter 1902b:211; not bear glue) is here designated the lectotype, Suter in Hutton 1904:71 (not Pleurotoma trailli to be certain that this name continues to be Hutton, 1873; Suter 1913:474). applied in its accustomed sense rather than to a Drillia buchanani maorum. Suter 1905a:73; Miocene species from Oamaru. It presumably Murdoch & Suter 1906:282; Suter 1913:474, pl. came originally from Shakespeare Cliff within 46, fig. 22. Wanganui City and almost certainly from Surcula buchanani maorum. Suter 1905b:200. Tainui Shellbed (OIS 13). The paralectotype is Melatoma buchanani. Hedley 1922:250. TM8629. Pleurotoma (Drillia?) maorum, holo- Melatoma buchanani maorum. Powell 1927:296. type BMNH 1852.3.19.52 (Fig. 14F; Dell Antimelatoma buchanani. Powell 1942:97; Flem- 1963a, pl. 2, fig. 7), from ‘New Zealand, leg. ing 1966:73; Powell 1966:30; Tucker 2004:159; Lieut. Col. Bolton’. Antimelatoma ahiparana, Maxwell 2009:249. holotype AIM AK72181 (Fig. 14E), with 13 Antimelatoma buchanani maorum. Powell paratypes AK71791 (Fig. 14C, H), Recent, 1942:97; Dell 1963a:176, pl. 2, fig. 7; Powell dredged in 42 m off Ahipara, NW Northland, 88 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 14 (A,G,I,J) Antimelatoma waimea n. sp.; A,I, holotype, TM8670, GS3013, J32/f9113, upper Waimea Creek, between Taramakau and Arahura Rivers, Westland, Kapitean; height 16.5 mm; G,J, paratype, TM8671, GS3030, J32/f9130, lower Waimea Creek, Westland, Opoitian; G, incomplete shell, height 13.4 mm; J, protoconch of G. (BF,H) Antimelatoma buchanani (Hutton); B, lectotype of Pleurotoma buchanani, TM8628, Tainui Shellbed, Castlecliffian (OIS 13), ‘‘Shakespeare Cliff’’, Wanganui; H 21.0, D 6.8 mm; C,H, paratype of Antimelatoma ahiparana Powell, AIM AK71791, dredged, 42 m, off Ahipara, Northland, ex Finlay collection; C, height 12.8 mm; H, protoconch of C; D, GS4003, R22/f8002, Landguard Sand (Haweran, OIS9), Landguard Bluff, Wanganui; height 15.1 mm; E, holotype of A. ahiparana, AIM AK72181, same data as paratype; H 12.5, D 4.6 mm; F, holotype of Pleurotoma (Drillia?) maorum Smith, BMNH 1852.3.19.52, Recent, ‘‘New Zealand’’; H 20.4, D 7.0 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 89

ex HJ Finlay collection. Antimelatoma benthi- Distribution. The Antimelatoma buchanani ‘ahi- cola, holotype AIM AK70910, ex HJ Finlay parana form’ is known fossil in Landguard collection, from 160225 m, off Otago Heads Sand (OIS 9) in Wanganui Basin only, and (not seen). occurs in the Recent fauna off W North Island and in W Cook Strait (taken in 24103 m). Specimens from further south (NMNZ Other material examined, A. ahiparana form. M.59044, 91 m, off Kahurangi Shoals, NW of Haweran: Landguard Sand (OIS 9), Landguard Kahurangi Point, NW Nelson; four chalky; Bluff, E of Wanganui River (GS4003, R22/ substrate labelled as ‘subfossil shells’) are f7394, six; including re-collections); recorded normal, relatively finely sculptured Antimelato- also by Fleming (1953:245) from Landguard ma buchanani. The specimens from flats inside Sand, as ‘Antimelatoma n. sp.’; NMNZ Farewell Spit (NMNZ M.50033) are intermedi- M.15713, Landguard Bluff (one). Recent: ate in most characters between the buchanani RM2090, dredged, 42 m, off Ahipara (five; and ahiparana forms. This is therefore inter- two presented to J Marwick by W La Roche; preted as an ecophenotypic form intergrading three ex CA Fleming collection); NMNZ with A. buchanani, the typical form of which M.4229, off Reef Point, Ahipara (three); occurs throughout N and E New Zealand (and M.90601, Ahipara Bay (45, consistently A. presumably along the W South Island), and is ahiparana form); M.118479, 103 m, NW of an abundant fossil in Nukumaruan to Haweran Ahipara, 34841.90?S, 172833.50?E (eight, spiral rocks throughout New Zealand. Upwelling N cords as in A. ahiparana form, axial ridges more of the northernmost North Island possibly prominent); M.118483, 90 m, NW of Ahipara, restricts gene flow there between the buchanani 34850.00?S, 172846.10?E (four fragmentary); and ahiparana forms. A. buchanani occurs M.54961, 24 m, off Pakawau Beach, Golden widely, syntopically with Aoteadrillia wanga- Bay, Nelson, 40837.00?S, 172848.00?E (one nuiensis, at most Nukumaruan and younger small, coarse nodules); M.50033, 2426 m, off fossil localities in Wanganui Basin, the Bay of flats inside Farewell Spit, Golden Bay (three Plenty, Hawke’s Bay, Wairarapa and Marlbor- large, more prominent peripheral nodules than ough-North Canterbury, but consistently is less most A. buchanani, spiral sculpture more like abundant than A. wanganuiensis. the normal form of A. buchanani). Fleming (1953:131, 146, 151, 188, 203204, Dimensions. Lectotype of Pleurotoma buchana- Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 215, 218, 226, 230, 232, 239, 240, 245) recorded ni: H 21.0, D 6.8 mm; paralectotype: H 17.5, D 6.6 mm; holotype of Pleurotoma (Drillia?) Antimelatoma buchanani from many Nukumar- maorum: H 20.4, D 7.0 mm; holotype of uanCastlecliffian formations in Wanganui Antimelatoma ahiparana: H 12.5, D 4.8 mm; Basin (Hautawa Shellbed, Mangamako paratypes AK71791: H 13.8, D 4.9 mm; H 13.1, Shellbed, Waipuru Shellbed, Kaimatira Pumice D 4.7 mm; H 12.8, D 4.6 mm; GS4003, Land- Sand, Kupe Formation, Seafield Sand, Lower guard Bluff: H 14.8, D 5.2 mm; H (incomplete) Castlecliff Shellbed, Pinnacle Sand, Tainui 13.3, D 5.0 mm; H 11.3, D 4.2 mm; NMNZ Shellbed, Shakespeare Cliff Siltstone, Upper M.50033, 2426 m, inside Farewell Spit: H Castlecliff Shellbed, Karaka Siltstone and 19.2, D 6.3 mm. Landguard Sand). It also occurs commonly throughout mudstone formations in Hawke’s Remarks. Specimens of Antimelatoma from Bay, Wairarapa and N Canterbury-Marlbor- Landguard Sand, Landguard Bluff, Wanganui, ough, and in CastlecliffianHaweran rocks at differ from the common, widespread form of Cape Kidnappers and at Ohope Beach and Te Antimelatoma buchanani in having a longer, Piki, Bay of Plenty. more steeply sloping sutural ramp, only two 90 AG Beu

prominent spiral cords around the periphery buchanani maorum (E. A. Smith, 1877). Dell down the entire teleoconch, and prominent, (1963a:176, pl. 2, fig. 7) pointed out that this widely spaced, clearly defined spiral cords on form had never been illustrated previously, the base. It has three prominent, widely spaced apart from Suter’s (1913, pl. 46, fig. 22) spiral cords over the convex area of the base featureless little drawing, and supplied a photo- below the peripheral two, and above the con- graph of the holotype, in BMNH. Dell’s cave area of the neck. Well-preserved specimens illustration and examination of the holotype also have a glossy exterior. These characters are (Fig. 14F) show that this form differs from the shared with the Recent form named A. ahipar- common NukumaruanCastlecliffian form of ana by Powell (1942:98, pl. 2, fig. 11) and Antimelatoma buchanani, as noted by Powell comparing the Landguard specimens directly (1942:97), by the slightly more prominent with the holotype and 13 paratypes of A. uppermost two spiral cords, at the periphery, ahiparana showed that they are closely similar in A. buchanani maorum, and their more even in size, shape and sculpture. The protoconchs strength in A. buchanani buchanani, although also are identical. As Landguard Sand at Powell admitted that ‘This does not always Landguard Bluff is the horizon and locality apply, however’. Powell claimed a further slight with the greatest number of warm-water mi- but constant difference in the lengths of the grants recorded at Wanganui ( danieli, anterior canals, but I cannot see any difference Eunaticina papilla, Pupa affinis (A. Adams, between fossil and Recent specimens in this 1855), Zelippistes benhami, Philine tepikia Rud- character, despite this later (Powell 1979:231) man, 1970 [see below] and possibly Stiracolpus being maintained as the major difference be- vigilax), this seemed to be another record at tween the two forms. A. buchanani is highly Wanganui of a species now limited to northern variable in Recent material in NMNZ, some New Zealand. However, the distribution of specimens in a few lots from other areas Antimelatoma forms along the west coast of resembling the Ahipara specimens (e.g. New Zealand seems better interpreted as a NMNZ M.71413, 8994 m, Ranfurly Bank, more coarsely sculptured ecophenotypic form off East Cape; one specimen of A. ahiparana occurring from the Ahipara region southwards, form with 13 coarsely sculptured but otherwise intergrading with more finely sculptured speci- typical A. buchanani). A few specimens almost mens of the common form of A. buchanani completely lack axial ridges, nodules and pro- south of western Cook Strait. The common minent spiral cords on the last few whorls

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 finely sculptured form occurs from N of Cape (particularly NMNZ M.58826, 139 m, ENE Reinga to the Three Kings Islands and of Tolaga Bay, E North Island, 38815.20?S, throughout eastern New Zealand. The Land- 178838.60?E; two large). Most Recent speci- guard Bluff specimens then probably are merely mens from eastern New Zealand are insepar- standard western New Zealand specimens of A. able from NukumaruanCastlecliffian fossils, buchanani, possibly reflecting the cutting off and the variation is all part of the single species (possibly by closure of the Manawatu Strait) of A. buchanani. The range of variation now the dispersal of larvae from eastern New accepted for A. buchanani also includes the Zealand after about OIS11 10. Specimens form named A. benthicola by Powell (1942:98, from lower in the Castlecliff section are all pl. 2, fig. 10), based on a single only slightly typical ‘eastern New Zealand’ specimens of A. more coarsely sculptured specimen from shal- buchanani, with more even, fine sculpture, a low water off E Otago; Otago specimens in shorter sutural ramp, and the two peripheral NMNZ are typical A. buchanani. cords not distinguished from their neighbours. Powell (1942, 1966) referred only the New Landguard Sand specimens are also similar Zealand Nukumaruan to Recent species to to the form long known as Antimelatoma Antimelatoma, but Cotton (1947:11) included Marine Mollusca of isotope stages of the last 2 million years in New Zealand 91

eight Recent Australian species in Antimelato- Distribution. Known only by the type material. ma, although he later reduced this to three species (Cotton 1959:393): A. gratiosa (G. B. Dimensions. Holotype: H 16.5, D 5.3 mm. Sowerby III, 1896), A. harpularia (Desmoulins, 1842) and A. agasma Cotton, 1947. Cotton was Description. Shell small for genus, tall and followed by Macpherson & Gabriel (1962:232), gracefully slender, with tall, narrow spire and who referred two Victorian Recent species here, relatively short last whorl (siphonal canal apex A. harpularia and A. subviridis (May, 1910). and outer lip slightly incomplete). Protoconch GNSmaterial of ‘ Antimelatoma’ harpularia of 1.5 whorls, with low, wide, blunt apex (WM19103, Portland, Victoria; four) has pro- followed by one taller, evenly inflated whorl, minent, long, narrow, closely spaced axial all smooth; passing into teleoconch gradually ridges and weak spiral cords and is more nearly by development of progressively more promi- similar to species of Hedley, 1922 nent brephic axial ridges, towards end of ridges than to A. buchanani, and Pleurotoma harpular- gradually developing low, smooth subsutural ia was placed in Splendrillia by Powell cord and medial angulation defining sutural (1966:83), although the other species in Cot- ramp; brephic axial ridges curved, with apex ton’s (1959:393) list and ‘A.’ subviridis were not abapertural; similar to that illustrated by included in Powell’s (1966) monograph. Wells Powell (1942, text-fig. C10) for A. buchanani, (1990:87, pl. 4, figs. 1, 2) and Tucker (2004:965) but with more prominent brephic axial ridges also included Drillia subviridis in Splendrillia, and lacking spiral cords. Teleoconch of seven Wells (1990:77) synonymized Antimelatoma whorls, spire rendered prominently gradate by agasma with Splendrillia woodsi (Beddome, peripheral angulation at base of sutural ramp, 1883), and Wells (1991:65, pl. 1, fig. 3) included at about upper third on spire whorls; base Drillia gratiosa in Splendrillia (Tucker weakly convex below angulation, passing gra- 2004:439). Therefore, none of the Australian dually into weakly concave neck and moder- Recent species listed in Antimelatoma by Cot- ately long, gently tapered, unnotched anterior canal. Sculpture visually dominated by promi- ton (1947, 1959) was correctly assigned there. nent, narrow, long axial ridges, moderately strongly opisthocline in conformity with outer Antimelatoma waimea n. sp. (Fig. 14A, G, I, J) lip, each interspace slightly wider than one ridge, commencing abruptly and forming small

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Type material. Holotype (Fig. 14A,I) TM8670, rounded nodules at lower margin of sutural GS3013, J32/f9113, grid reference J32/557353, ramp, producing a much more deeply concave Kapitean (late Miocene), occurring with Aetho- ramp than on A. buchanani; 12 axial ridges on cola coerulescens (Finlay, 1930); main course of last whorl and 13 on penultimate whorl of Waimea Creek, 200 m N of Stafford Loop holotype. Spiral sculpture of one prominent, Road and 400 m WNW of Goldsborough, 4 km smooth, rather narrow subsutural cord, an upstream from Stafford and ca 9 km upstream almost smooth sutural ramp (crossed only by from mouth of Waimea Creek, between Tar- weak growth ridges defining posterior sinus), amakau and Arahura Rivers, Westland; one one narrow and two much more prominent incomplete paratype (Fig. 14G, J) TM8671, spiral cords around periphery, the uppermost GS3030, J32/f9130, grid reference (estimated) two closely spaced and the next two below J32/546359, Opoitian (early Pliocene), with more distantly spaced, with lower, narrower, Marama murdochi (Marwick, 1927) and Amal- more closely spaced cords below, gradually da (Baryspira) cf. tholiculus (Marwick, 1931), becoming revealed down spire as ramp des- ca 1.25 km downstream from type locality, cends; cords are weakly convex-crested and main course of Waimea Creek, Westland. sharply defined; two further cords on sides of 92 AG Beu

antepenultimate and three on penultimate Antimelatoma quemadensis (Ihering) whorl below more prominent cords; followed (1907:195, pl. 6, fig. 39; proposed in Fusus) below by ca eight similar cords on base of last (Oligoceneearly Miocene, Yegua Quemada, whorl, passing into ca 10 further weak, low, near Santa Cruz, Argentina; Monte Leon closely spaced cords on siphonal canal; small, Formation), was referred to this genus by Beu narrowly rounded nodules (much more obvious et al. (1997:95). It differs from the New Zealand than any seen on A. buchanani) formed at species in having a less prominent angulation sculptural intersections on uppermost two pro- around the base of the sutural ramp, and the minent spiral cords on last two whorls, weaker axial ridges (although also said by Ihering higher up spire; cords scarcely raised over axial [1907:196] to be 12 per whorl) are less promi- ridges lower down last whorl. Aperture un- nent than in both A. buchanani and A. waimea thickened, without parietal callus, outer lip n. sp. Comparison is still required to be certain slightly incomplete but evidently thin and that this species belongs in Antimelatoma. The simple, without stromboid notch; profile mod- age of Antimelatoma waimea n. sp. moves the erately concave over sutural ramp, forming time of Neogene immigration of Antimelatoma posterior sinus; curving forwards moderately to New Zealand from South America, as below ramp to form standard Antimelatoma postulated by Beu et al. (1997:95), to during convex outer lip profile. late Miocene time, much earlier than the Nukumaruan date envisaged previously. I am Remarks. Antimelatoma waimea n. sp. from late not aware of any other possible origin for the Mioceneearly Pliocene rocks of Westland has New Zealand species of this genus. more prominent axial and spiral sculpture than A. buchanani, and also has a slightly shorter, Etymology. The species name is that of the type more strongly nodulose teleoconch (particu- locality; a noun in apposition. larly with a shorter anterior siphonal canal, although it is slightly to highly incomplete on Family Conidae both specimens) with a more strongly concave sutural ramp and more prominent, narrow, Subfamily Pseudotominae? widely spaced, more strongly oblique axial ridges than any forms of A. buchanani. The Genus Austrotoma Finlay, 1924 protoconch (Fig. 14J) is very well preserved on

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 both holotype and paratype, and differs Austrotoma Finlay 1924:515. Type species (by significantly in its larger size and in having original designation): Bathytoma sulcata exca- more prominent brephic axial ridges (where vata Suter, 1917, Otaian (early Miocene), the sculpture gradually passes from that of the Kaipara Harbour, Northland, New Zealand protoconch to the telcoconch) and in lacking the spiral cords on the last whorl of the protoconch of A. buchanani. The differences Remarks. A position for this subfamily(?) in seem unexpectedly marked from A. buchanani, Conidae is provisional. A cladogram of Brazi- particularly in the protoconch, suggesting that lian conoideans (Simone 2007, fig. 11), based this species may not have had a direct ancestor- on Simone’s research on caenogastropod com- descendent relationship with A. buchanani. The parative anatomy, shows ‘Pleurotomella’ forms named by Powell (1942) now all being aguayoi (referred below to Austrotoma)asthe considered part of the variation of the single nearest relative of of the taxa Simone species A. buchanani, A. waimea n. sp. is only studied, so a position in Conidae seems likely. the second species of Antimelatoma recognized This group is not recognized in the prelimi- in New Zealand. nary molecular phylogeny of Conoidea by Marine Mollusca of isotope stages of the last 2 million years in New Zealand 93

Puillandre et al. (2008), but none of its possible unnecessary replacement names), Surculites members was studied by them. Recognition Conrad, 1865 and its subgenus or synonym that Austrotoma is still living in the SW Atlantic Notogenota Powell, 1942, Nekewis Stewart, provides an opportunity to examine the anato- 1926, Tahuia Maxwell, 1992, and Zeatoma mical characters and familial position of the Maxwell, 1992. Marshallena and potential subfamily Pseudotominae, recognized were included in Family Turridae, Subfamily by Beu & Maxwell (1990:417) and Maxwell Cochlespirinae by Taylor et al. (1993:165), but (1992:153; 2009:249) as a useful subfamily of any close phylogenetic relationship seems un- distinctive, large, wide taxa, most of which have likely with the small, slender genus Cochlespira an unusually short, widely open anterior sipho- Conrad, 1865, with its long, narrowly con- nal canal and an only weakly contracted last tracted siphonal canal, and Cochlespira is the whorl. Of course, this subfamily name was used furthest removed from A. aguayoi of all Con- by many earlier workers, such as Finlay oidea in Simone’s (2007, fig. 11) cladogram. (1924:516) and as ‘the Acamptogenotia group’ Pseudotominae was listed as a synonym of by Hickman (1976:47). The subfamily name Family Conidae, Subfamily Clathurellinae by usually has been attributed to L Bellardi Bouchet et al. (2005:256) and this seems a (1877:209). However, Bouchet et al. phylogenetically more likely association. How- (2005:145) attributed the subfamily name to A ever, anatomical and molecular evaluation of Bellardi (1875:19) and pointed out that Pseu- this group is still required, based expressly on dotoma Gray is an incorrect spelling of Pseu- Megasurcula carpenteriana and/or Austrotoma dostoma, leaving Pseudotoma A. Bellardi aguayoi and other living Brazilian species of available. Austrotoma. Hickman (1976) threw new light Few Recent species seem correctly referred on the characters of supposedly geographically to this group, apart from the several Indo-West restricted genera in this group when she de- Pacific Recent bathyal species of Marshallena scribed Oligocene species of both Pseudotoma Allan, 1927 (Powell 1969:365372; M. philippi- and Austrotoma from the Keasey Formation of narum (Watson, 1882)*Sysoev in Poppe NW Washington State, USA. She reconsidered 2008:784, pl. 687, fig. 8). Another is Mega- the relatively minor differences between these surcula carpenteriana (Gabb, 1865) from and Megasurcula, ranking all these groups as California, USA (Powell 1966:32, pl. 2, fig. subgenera of Pseudotoma. She also illustrated 18), type species of Megasurcula Casey, 1904 the Italian Pliocene species Pseudotoma bonellii

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (by subsequent designation, Grant & Gale Bellardi, 1877 (Hickman 1976, pl. 4, figs. 13, 1931:495). Grant & Gale (1931:497500, pl. 14), which bears the prominent subsutural ridge 25, figs. 34b) compared M. carpenteriana with that supposedly differentiates Austrotoma from Pseudotoma intorta (Brocchi, 1814) (Grant & Pseudotoma, and Pseudotoma morreni (de Ko- Gale 1931, pl. 25, fig. 1a,b; Hickman 1976, pl. ninck, 1838) (Hickman 1976, pl. 4, figs. 6, 7), 4, figs. 12, 15; Fig. 16C), type species of Rupelian (Oligocene), Germany, which closely Pseudotoma A. Bellardi, 1875. Hickman resembles Austrotoma and Megasurcula species. (1976), Beu & Maxwell (1990:417) and Maxwell Species of Nekewis and Surculites occur with (1992) included the following genera in sub- those of Pseudotoma and Austrotoma, so the family Pseudotominae: Belophos Cossmann, Keasey Formation has an unusually diverse 1901, Austrotoma Finlay, 1924, Liratomina pseudotomine fauna. Hickman (1976) also Powell, 1942, Belatomina Powell, 1942, Mar- suggested that Belophos Cossmann, 1901 might shallaria Finlay & Marwick, 1937, Marshallena be the earliest name for the genus or subgenus Finlay, 1926, Megasurcula Casey, 1904, Pseu- usually known as Austrotoma, and her evi- dotoma A. Bellardi, 1875 (Acamptogenotia dence certainly suggests that groups such as Rovereto, 1899;Pseudotomina Finlay, 1924, Pseudotoma and Austrotoma are very similar 94 AG Beu

and are possibly congeneric rather than merely [originally ca 37], D 27.8 mm; Q2186: H consubfamilial. Molecular comparison of M. [incomplete] 21.6, D 14.2 mm). This demon- carpenteriana and A. aguayoi would be an strates that Struthiolariopsis is indeed a conoi- excellent beginning to understanding relation- dean, with an obvious, moderately deep anal ships in this group; it would not be surprising if sinus occupying the concave sutural ramp. they fall into one clade, that is, constitute a However, it is more coarsely sculptured than single genus. most Austrotoma species and has a prominent Marwick (1924a:161, fig. 1a) suggested that subsutural fold and a more strongly tapered the poorly known genus Struthiolariopsis Wilck- last whorl with a narrower siphonal canal, so ens (1904:208; type species by monotypy: Fusus that it resembles a nodulose species of Marshal- ferrieri Philippi, 1887; Maastrichtian [late Cre- lena or Nekewis more closely than Austrotoma. taceous], Quiriquina Island, Chile) also was Both Philippi’s and Wilckens’ drawings based on a relatively large conoidean resembling are inaccurate. Struthiolariopsis ferrieri is not Austrotoma. Both Wilckens’ (1904, pl. 18, fig. 5) closely similar to Austrotoma, but is possibly an drawing and Marwick’s (1924a, fig. 1a) copy of early member of the ‘group’ or subfamily it resemble Austrotoma in their single row of Pseudotominae. peripheral nodules, prominent spiral cords Belophos, Austrotoma, Lirotomina and Be- around the gently tapering base, and tall, almost latomina, recognized as separate genera by smooth, weakly concave sutural ramp occupied Powell (1942, 1944, 1966), are among the by a shallow anal sinus. However, Philippi’s many groups requiring reassessment in Con- (1887:43, pl. 2, fig. 6) original drawing seems to oidea, as Hickman (1976) pointed out. Belophos show a different, more coarsely sculptured, woodsii (Tate) (1888:147, 173, pl. 4, fig. 3) (Fig. buccinoidean shell. Bandel & Stinnesbeck 15E, H, K) has usually been considered to be (2000) did not refer to Fusus ferrieri, but Klaus the valid name for the type species of Belophos Bandel (Universita¨ t Hamburg, pers. comm. Cossmann (1901:162; type species (by original June 2008) stated that it is among the species designation): Bela woodsii Tate, 1888, Fossil they referred to Pyrifusus Conrad, 1858 (Bandel Bluff, near Wynyard, N Tasmania; gender & Stinnesbeck 2000, list p. 784, ‘Pyrifusus neuter). However, there is apparently no reason several species’). Sven Nielsen and Steffen Kiel not to use the original name, B. cancellatum (Institut fu¨ r Geowissenschaften, Christian-Al- (Tenison Woods, 1877). Tate’s species name brechts-Universita¨ t Kiel pers. comm.) consider apparently is an unnecessary replacement name

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 that Quiriquina material of Struthiolariopsis for Cominella cancellata Tenison Woods ferrieri is similar to species of the conoidean (1877:107); I have been unable to find any genus Beisselia Holzapfel, 1889 (type species: earlier usage of Cominella cancellata. The Koenenia speciosa Holzapfel, 1888, late Cretac- combination Belophos cancellatum was reintro- eous, Europe; Wenz 1943:1316, fig. 3746). They duced for this species by Goudey (2006:46, figs. provided digital images of the remaining syntype 21, 22; p. 48, fig. 2) and is followed here. of Fusus ferrieri (SGO.PI.567) sent to them by Illustrations of Belophos cancellatum by Coss- Dr Daniel Frassinetti (Paleontologı´ a Inverteb- mann (1901, pl. 6, figs. 9, 10), Powell (1966, pl. rados, Museo Nacional de Historia Natural, 4, figs. 2, 3) and Goudey (2006) show a shell Santiago, Chile), showing that it is conspecific closely resembling Tahuia formosa (Allan, with the material illustrated here. They also lent 1926) (Maxwell 1992, pl. 22g,n) and Marshal- Quiriquina material of S. ferrieri (Fig. 15I, J) (in laria decipiens Maxwell (1992, pl. 22d,f,j,k), collection S. Kiel, Institut fu¨ r Geowissenschaf- that is, it has more prominent axial ridges and ten, Christian-Albrechts-Universita¨ t Kiel, five peripheral nodules than New Zealand species incomplete specimens, all calcite neomorphs, assigned to Austrotoma (although not more so including Q3020: H [very incomplete] 30.0 than in Recent Brazilian species discussed Marine Mollusca of isotope stages of the last 2 million years in New Zealand 95

below). Specimens examined (WM18996, Fossil threads between the primary spiral cords, and Bluff, near Wynyard, N Tasmania, type local- again lack the obvious, fine, close axial threads ity; Longfordian, early Miocene; three speci- present in Tahuia, Marshallaria and Belophos. mens; NMV P315217-P315224, eight specimens B. cancellatum also closely resembles the short- lent by Dr TA. Darragh; Fig. 15E, H, K) show er and more robustly sculptured of the South that, as described by Powell (1966), B. cancel- American and Antarctic Paleogene fossil spe- latum has a relatively blunt, low-conical proto- cies previously assigned to Austrotoma (Stilwell conch of about 3.54 whorls, with a small, & Zinsmeister 1992; Nielsen 2003; Griffin & blunt apex, convex outlines, and raised, flat- Nielsen 2008:297, pl. 20, figs. 2427) although, crested spiral cords on the last whorl, similar to again, these species lack the fine axial threads that of T. formosa (shown accurately in the characteristic of Belophos and Tahuia. The drawing by Cossmann 1901:163, fig. 42). It is range of sculpture and shape exhibited by also very similar to those of Marshallaria and Austrotoma species in New Zealand also in- Zeatoma species. Comparison of specimens cludes most shell shapes of species assigned by showed that Tahuia formosa is smaller, has a Powell (1942, 1944) to Belophos, Liratomina more rounded peripheral angulation, and has a and Belatomina. Few Australian or New Zeal- slightly shorter, blunter protoconch than B. and Austrotoma species have as deeply concave cancellatum. However, B. cancellatum is quite a sutural ramp or as prominent a peripheral variable in axial fold development (Fig. 15E, H, keel as Belophos cancellatum, but again the SW K) and some specimens have very weak axial Atlantic Recent species discussed below and a sculpture over the last half-whorl; the speci- few extreme New Zealand species such as A. mens illustrated by Cossmann (1901, pl. 6, figs. clifdenica Powell, 1942 (Fig. 15M) are similar to 9, 10) have a slightly more rounded periphery B. cancellatum in this respect. In view of the than those I have examined, and closely resem- range of sculpture shown by the material ble Tahuia formosa in all characters. Some illustrated by Hickman (1976) and the South undescribed species of Marshallaria (e.g. American and Antarctic species, it seems likely GS9723, NO2/f7594, Paratoetoe, opposite Te that the differences between Belophos, Tahuia, Hapua, N shore Parengarenga Harbour, N Marshallaria, Austrotoma and Lirotomina are Northland; Altonian, early Miocene; three trivial, that is, these names possibly are syno- specimens) have a sharper, more prominent nyms. All of them (along with Megasurcula) are peripheral angulation than species illustrated also possibly simply synonyms of Pseudotoma.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 by Maxwell (1992), and resemble Tahuia and Even Powell (1944:27), when describing Lira- Belophos species in all characters. Both Belo- tomina adelaidensis, commented that ‘this spe- phos cancellatum and Tahuia formosa have a cies has a strong superficial resemblance to sharply ridge-margined siphonal fasciole Austrotoma’. His new species Austrotoma in- formed by their shallowly notched anterior expectata and L. adelaidensis are very similar in canals, and bear highly distinctive, fine, closely all characters; it appears that his sole reason for spaced, axial and spiral threads over the entire placing L. adelaidensis in Liratomina was the teleoconch, rendering them particularly closely absence of the subsutural fold that he consid- similar. The sculpture of some Marshallaria ered diagnostic of Austrotoma. Large Conoidea species is little different, although wider spiral of this ‘group’ or subfamily are widespread and cords predominate on most species. In contrast, their differences are much less marked than Zeatoma species are characterized by their Powell supposed, so the occurrence of species exceedingly fine, closely spaced spiral threads of this group living in the SW Atlantic is not between the primary spiral cords, not crossed really surprising, and A. aguayoi is possibly not by any obvious fine axial sculpture. Austrotoma very closely related, phylogenetically, to New species have fewer, slightly coarser spiral Zealand Austrotoma species. 96 AG Beu

This discussion suggests that in a relatively Other material examined. Brazil: trawled by conservative, but still finely subdivided classifi- fishermen, 2535 m, off Vitoria, Espirito Santo cation, Tahuia is best ranked as a synonym of State, S Brazil (WM19004, two); trawled by Belophos, Belatomina is best ranked as a syno- shrimpers at 100120 m, off Nitero´ i, Rio de nym of Marshallaria, and Liratomina is best Janeiro State, S Brazil, August 2004 ranked as a synonym of Austrotoma. It also (WM19012, ex Museu de Zoologia da Univer- seems possible that Pseudotoma is the earliest sidade de Sa˜ o Paulo, MZSP 70943; two; Fig. name for a genus that has also been named 15C,D); dredged, 100120 m, off Arraial de Marshallaria and Belatomina. However, in my Cabo, Rio de Janeiro State, Brazil, Sept. 2008 opinion Belophos (Tahuia), Marshallaria (WM19015, four; Fig. 15A, B). Argentina: (Belatomina), Austrotoma (Liratomina), dredged, ‘old bed’, 38826?S,57840’W, Megasurcula and Pseudotoma all should be 4050 m, off SE Argentina, presented by Laura retained as genera at present, until at least Schejter, Lab. de Bentos, Inst. Nacional de DNA sequences of M. carpenteriana and A. Investgacio´ n y Desarrollo Pesquero, Mar del aguayoi can be compared. Marshallena seems Plata, Argentina, 2009 (WM19053, one). In well separated from these other genera by its January 1998 I also examined 20 lots dredged slightly longer, narrower and more constricted off the coast of northern Argentina in Museo anterior siphonal canal. The relatively fine axial Nacional de Ciencias Natural (MACN), Bue- and spiral sculpture present on A. aguayoi, only nos Aires. Most lots were dredged on the inner slightly coarser than that of Belophos cancella- continental shelf, but one is labelled ‘banco de tum, and its relatively weak subsutural spiral mejillones [mussel bank], Necochea’, ca 100 km ridge suggest that a position in Pseudotoma Sof Mar del Plata (MACN 31127, four speci- might be just as suitable as one in Austrotoma. mens), that is, it occurs as shallow as the low- However, a few New Zealand species of Aus- water spring tide line, where Trophon geversia- trotoma lack a prominent subsutural ridge (e.g. nus (Pallas, 1774) and banks of Aulacomya atra the undescribed species in Fig. 15F), it is absent (Molina, 1782) are abundant along the very from the species Powell (1944) referred to wide, gently shelving, sandy Argentinean shore. Liratomina, and it is present or absent in various Presumably Austrotoma predates Aulacomya, European Neogene species assigned to Pseudo- among other prey. toma (Hickman 1976, pl. 4, figs. 1215); its taxonomic significance seems low. Further re- Distribution. Nu´ n˜ es Corte´ s & Narosky

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 search is obviously required on this group. (1997:61) and Forcelli (2000:109, fig. 321) recorded Austrotoma aguayoi as ranging from Austrotoma aguayoi (Carcelles, 1953) (Fig. Rio de Janeiro State, Brazil, to Chubut Pro- 15AD) vince, Argentina, in 3570 m. Absala˜ o et al. (2005:34) recorded specimens from three sta- Clathurella aguayoi Carcelles 1953:12, pl. 4, fig. tions (D2, 22853?S,41809?W, 82 m; D3, 22852?S, 20ad. 41809?W, 80 m; R1#1, 21838’57ƒS,40810’47ƒW, Pleurotomella aguayoi. Rios 1994:178, pl. 59, 100180 m) off south-central Brazil, between fig. 823; Nu´ n˜ es Corte´ s & Narosky 1997:61, Salvador (Bahia State) and Cabo de Sa˜ o Tome´ illus.; Forcelli, 2000:109, fig. 321; Tucker (Rio de Janeiro State), so it occurs further 2004:39; Absala˜ o et al. 2005:34, fig. 95, 112; north and in slightly deeper water than re- Simone 2007:198, fig. 11. corded previously, although all records are in temperate waters. Type material. Carcelles (1953:12) listed the type material, in Museo de Historia Natural de Dimensions. WM19015, dredged, 100120 m, Montevideo (not seen). off Arraial de Cabo, Rio de Janeiro State: H Marine Mollusca of isotope stages of the last 2 million years in New Zealand 97 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 15 Conidae, subfamily Pseudotominae? (AD) Austrotoma aguayoi (Carcelles), Recent, dredged off Rio de Janeiro State, S Brazil; A,B, WM19015, 100120 m, off Arraial de Cabo; H 25.0, D 12.0 mm; A, unwhitened, note ; B, whitened with MgO; C,D, WM19012, 100120 m, off Nitero´ i, H 24.3, D 12.4 mm. (E,H,K) Belophos cancellatum (Tenison Woods), type species of Belophos Cossmann, 1901, Freestone Cove Sandstone (Longfordian, Early Miocene), Fossil Bluff, N Tasmania, NMV; E, P.315222, H 29.2, D 15.2 mm; H, P.315218, H 30.2, D 15.1 mm; K, P.315217, H35.1, D 15.3 mm. (F) Austrotoma n. sp., weak subsutural fold, GS8101, Y14/f7618A, Oweka Creek, between Whangaparaoa and Hicks Bay, East Cape, Opoitian (Early Pliocene); H 48.2, D 19.5 mm. (G) Austrotoma cf. nervosa Powell, GS9700, J38/f9681, Southburn Sand (Altonian, Early Miocene), 1.5 km SW of Sutherlands, Tengawai River, S Canterbury; H 37.7, D 17.4 mm. (I,J) Struthiolariopsis ferrieri (Philippi), type species of Struthiolariopsis Wilckens, 1904, Quiriquina I., Chile, Maastrichtian (Late Cretaceous), in collection S. Kiel; I, Q2186, H 21.6, D 14.2 mm; J, Q3020, H 30.0, D 27.8 mm. (L) Austrotoma echinata Powell, GS2155, D45/f8428, left bank Waiau River 400 m above bridge, Clifden, Southland, Lillburnian (Middle Miocene); H 29.0, D 15.1 mm. (M) Austrotoma clifdenica Powell, GS11213, D45/f002, Slip Point Siltstone (Clifdenian, early Middle Miocene), between Long Beach and Slip Point, Clifden, Southland; H 38.8, D 19.3 mm. 98 AG Beu

25.0, D 12.0 mm; H 23.5, D 10.2 mm; ably typical Austrotoma species resembling A. WM19012, trawled by shrimpers, 100120 m, excavata (Suter, 1917) (Beu & Maxwell 1990, off Niteroi, Rio de Janeiro State: H 24.3, D pl. 26h), A. clifdenica Powell, 1942 (Fig. 15M; 12.4 mm; H 23.2, D 11.2 mm; WM19053, Beu & Maxwell 1990, pl. 26e), A. nervosa 4050 m, dredged, ‘old mussel bed’, off SE Powell, 1942 (Fig. 15G; A. cf. nervosa) and A. Argentina: H 20.3, D 10.6 mm. lawsi Powell, 1942 (Beu & Maxwell 1990, pl. 22 l), with a slightly wider anterior end, smaller Remarks. This Recent species is included to nodules at the periphery, and a markedly point out that Austrotoma (as used tradition- weaker subsutural fold than A. lawsi. The ally, that is, ignoring the possibility that this peripheral nodules are more prominent in the genus should be known as Belophos or even photograph by Rios (1994, fig. 823) than in Pseudotoma) is not the ‘restricted Australian most New Zealand fossils, and are also very and New Zealand’ genus it has still usually been prominent in some of the Femorale web page thought to be, despite comments to the con- illustrations. However, other New Zealand trary by Hickman (1976), but is still living in fossil species such as A. echinata Powell, 1942 shallow water in the SW Atlantic Ocean. (Fig. 15L) have much more prominent periph- Several fossil species of Austrotoma occur in eral nodules than the Recent shells. The speci- OligoceneMiocene rocks of South America mens illustrated by Nu´ n˜ es Corte´ s & Narosky (Chile: Nielsen 2003:190, pl. 39, figs. 715, A. (1997:61) and Forcelli (2000:109, fig. 321) are echinulata (Hupe´ , 1854)Pleurotoma turbinel- orange-brown, with faintly paler spiral bands. loides G. B. Sowerby II, 1846, junior primary Specimens illustrated on the Femorale web homonym of P. turbinelloides Reeve, 1846; page are darker red-brown to purplish brown Fusus modestus Philippi, 1887); undescribed banded with white, whereas other illustrated ones observed near Santa Cruz in S Argenti- Atlantic species are almost plain white. nean Patagonia (M Griffin pers. comm.); Recent Atlantic specimens of Austrotoma Oligocene/Early Miocene, King George Island, aguayoi are particularly similar to Belophos Scotia Arc: Karczewski (1987, pl. 35, fig. 4; as cancellatum (Fig. 15E, H, K) in their fine, Perissodonta cf. exilis); and in Eocene rocks of closely spaced axial and spiral sculpture, but Seymour Island, Antarctic Peninsula (Stilwell have a slightly better-developed subsutural fold & Zinsmeister 1992, pl. 23ad,fi). A Recent and coarser axial ridges than B. cancellatum.In occurrence in the SW Atlantic is therefore not view of the great range of characters displayed Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 surprising, but provides an example of the by Belophos, Austrotoma, Pseudotoma, Mar- opposite theme to much of this series of papers: shallaria and Megasurcula species, the generic a supposedly ‘restricted Australian and New position of A. aguayoi is undecided at present, Zealand’ genus that is extinct in Australia and but I tentatively place it in Austrotoma until New Zealand but survives in the temperate DNA sequences can be compared with Mega- South Atlantic. Coloured photographs were surcula carpenteriana and Marshallena species. published by Nu´ n˜ es Corte´ s & Narosky Absala˜ o et al. (2005, fig. 112) illustrated the (1997:61) and Forcelli (2000:109, fig. 321) and smooth, polished, bulbous protoconch of one illustrations of Brazilian specimens of this whorl present on Austrotoma aguayoi. This Austrotoma species and two other, apparently lecithotrophic protoconch passes gradually unnamed species are published on a recent web into the teleoconch and so is significantly page among 1960 colour photos of Conoidea different from the sharply demarcated, nar- (http://www.femorale.com.br/shellphotos/thumb rowly pointed one of 45 whorls on New page.asp?familyTURRIDAE&nav102). Zealand Austrotoma species. Powell (1942: These demonstrate that A. aguayoi is a reason- 71, text-fig. B6) described the protoconch of Marine Mollusca of isotope stages of the last 2 million years in New Zealand 99

New Zealand species as ‘polygyrate of 45 Genus Mitrellatoma Powell, 1942 whorls with a minute globular tip, the first three whorls being smooth and the remainder Mitrellatoma Powell 1942:109. Type species (by bearing strong flat-topped spiral cords and thin original designation): Columbella angustata axials towards its close’ (similar to the descrip- Hutton, 1886, Nukumaruan (late Pliocene), tions and illustrations by Maxwell central Hawke’s Bay, New Zealand. [1992:154158] of the protoconchs of Zeatoma, Marshallaria and Tahuia, and to Cossmann’s Remarks. Glibert (1954:40, pl. 1, fig. 16; pl. 5, [1901:163, fig. 42] illustration of that of Belo- fig. 13) assigned Pleurotoma ligata Defrance, phos cancellatum). This again seems likely to be 1826 to ‘ (Mitrellatoma)’, and it a distinction between a direct-developing (Bra- was also placed in Mitrellatoma by Powell zilian) and either a demersal lecithotrophic or a (1966:88). However, P. ligata is certain to be planktotrophic protoconch type (New Zeal- a homologue rather than a member of clade and), of no significance in phylogeny. Two of Mitrellatoma (if there is one). P. ligata has a the specimens in WM19015 (off Arraial de taller, narrower spire than M. angustata (Hut- Cabo) have the operculum placed in the ton, 1886), the type species of Mitrellatoma, aperture (Fig. 15A); its width fills the aperture and also differs in having a weakly concave and it is long and narrowly leaf-shaped with an rather than almost flat sutural ramp and more anterior terminal nucleus, although not as long prominent spiral sculpture. Glibert (1954) and as the aperture, and similar to that illustrated Powell (1966) pointed out similarities to Asthe- by Powell (1966:14, text-fig. F, opercula, fig. notoma P. ligata 176) for Benthofascis biconica (Hedley, 1903). species, and should be referred Pleurotomella esmeralda Olsson (1964:104, to Asthenotoma provisionally. Kilburn (1986: pl. 17, figs. 44b) is a further superficially 682, figs. 97, 101) referred a third species to similar species from the Early Pliocene Onzole Mitrellatoma, the Recent species M. Formation at Punta Gorda, Esmeraldas Pro- Kilburn, 1986, from southern Mozambique, vince, Ecuador. Olsson did not provide a East Africa. This little shell (height 6.4 mm) comparison with any other species, but his has a much deeper anal sinus than M. angusta- illustrations show a species with a slightly ta, and its sculpture of even, wide, low spiral more narrowly contracted base and less pro- cords is distinctive. Again this seems highly truding peripheral nodules than in A. aguayoi. likely to be a convergent form rather than a

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Although the Ecuadorean species appears to be species of clade Mitrellatoma, particularly as I shorter than Megasurcula species and more question below whether Mitrellatoma is a similar to Austrotoma, like many of the other recognizable clade even in New Zealand. Such species discussed here it is intermediate in many small, simple, featureless shells will not be characters between these two supposed genera. classifiable until the radula is known, at least, and their DNA sequences require comparison for their relationships to be resolved finally. Subfamily Clathurellinae Remarks. Puillandre et al. (2008) recognized Mitrellatoma angustata (Hutton, 1886) (Fig. that a large number of subfamily-level clades 16H, K) (seven in the relatively small number of taxa they examined) is subsumed within ‘subfamily Columbella angustata Hutton 1886:333; Hutton Clathurellinae’. It will be many years before the 1893:45; Pace 1902:54. real relationships of New Zealand taxa are Alcira angustata. Suter 1915:29, pl. 8, fig. 9. revealed, and the present classification is re- Mitrellatoma angustata. Powell 1942:110, pl. tained in the meantime. 12, fig. 7; Powell 1966:88, pl. 13, fig. 15; 100 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 16 (A,I) Paracomitas protransenna (Marshall & Murdoch), GS10857, U23/f7049, Maharakeke Mudstone (Nukumaruan), Maharakeke Road, W of Waipukurau, Hawke’s Bay; A, height 11.2 mm; I, protoconch of A. (B,J) Paracomitas augusta (Murdoch & Suter), TM1107, paralectotype, ex Suter collection, Recent, 200 m, off Great Barrier Island, E of Auckland; B, height 6.4 mm; J, protoconch of B. (C) Pseudotoma intorta (Brocchi), type species of Pseudotoma A. Bellardi, 1875, WM7025, Castelnuovo don Bosco, near Asti, NW Italy, Piacenzian (Pliocene); H 57.5, D 21.4 mm. (D,F) Gymnobela gypsata (Watson); D, NMNZ M.59541, 17231549 m, head of Hikurangi Trench, E of South Island, H 47.9, D 21.3 mm; F, 1 of 2 syntypes, BMNH 1887.2.9.979980, HMS‘‘ Challenger’’ Sta. 169, 1280 m, off East Cape, New Zealand; H 36.8, D 15.8 mm. (E,G) Paracomitas gemmea (Murdoch), type species of Paracomitas Powell, 1942, 2 specimens; GS4061, R22/f6392, Lower Kai-Iwi Siltstone (Castlecliffian, OIS 23), Castlecliff coast; E, H 13.8, D 5.4 mm; G, H 14.9, D 5.7 mm. (H,K) Mitrellatoma angustata (Hutton), type species of Mitrellatoma Powell, 1942, specimen resembling holotype, SEM, GS2220, V20/f8462, Devils Elbow Mudstone (late Nukumaruan), top of Devils Elbow hill, Hawke’s Bay; H, height 10.5 mm; K, protoconch of H. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 101

Fleming 1966:74; Beu & Maxwell 1990:419; ellatoma angustata, Kai Iwi’, but these speci- Tucker 2004:72; Maxwell 2009:248. mens from the lower part of the Castlecliff coastal section represent a small (H 4.4 mm) Type material. Columbella angustata, holotype columbellid with even, narrow, closely spaced in CMC according to Powell (1942:110), but spiral grooves and no sinus in the outer lip, not included in the catalogue of CMC fossil closely resembling Zemitrella choava (Reeve, types by Bradshaw et al. (1992); from Petane, 1859). None of these is conspecific with the 30 km N of Napier, central Hawke’s Bay, holotype of M. angustata. precise locality unknown; Tangoio Limestone or Mairau Mudstone, Petane Group, late Dimensions. Holotype, CMC M3361: H 9.8, D Nukumaruan; collected by Augustus Hamilton. (slightly incomplete) 3.8 mm; GS2220, Norton Hiller (CMC pers. comm. 11 December TM8589: H 10.5, D 4.0 mm. 2007) reported that the holotype (M3361) is still present in CMC, and was merely missed out of Remarks. This virtually unknown species is the catalogue by Bradshaw et al. (1992), and I included to point out its apparently distinctive have now examined it. characters and the lack of knowledge of its relationships, in the hope that more specimens Other material examined. The one specimen of might come to light so its status is clarified. Mitrellatoma angustata I am aware of other Mitrellatoma angustata is one of the rarest and than the holotype is that illustrated by Powell least well understood of Hawke’s Bay Nukumar- (1942, pl. 12, fig. 7), from Devils Elbow uan molluscs, being recorded only as the holo- Mudstone at the top of Devils Elbow hill, type (as stated by Powell [1942:109], the holotype Napier-Wairoa Highway (Highway 2) ca 35 is a rather poor, severely abraded specimen) and km N of Napier, late Nukumaruan (TM8589, the single specimen illustrated by Powell (1942, GS2220, V20/f8462; Fig. 16H, K). Despite pl. 12, fig. 7) (also significantly abraded). Powell extensive re-collection at this highly diverse (1942:109) stressed its great similarity to a small locality over 50 yr, I have not collected another specimen of a Phenatoma species, and it appears specimen. feasible to me that this species is based on a A specimen in GNS(‘middle marls’, New- highly aberrant individual of a Phenatoma ton Range, E of Lake , Hawke’s Bay, species, such as the weakly sculptured Nukumar- that is, high in Petane Group in the block NE uan species P. precursor Powell (1942:109, pl. 10,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 of Arapaoanui River, N of Devil’s Elbow; fig. 8). P. precursor occurs uncommonly in among the many Conoidea presented to GNS Devils Elbow Mudstone on Devils Elbow hill, by HJ Finlay) was labelled ‘Phenatoma angu- where the illustrated specimen of M. angustata stata’ by Finlay. It is a relatively large (H 14.7 was collected. P. precursor also occurs com- mm) but poorly preserved specimen of the monly in shallower-water facies such as shore- Phenatoma species now usually known as P. face sands in central Hawke’s Bay (e.g. GS11225, precursor Powell, 1942, a species that occurs V20/f8002, Darkys Spur Formation, long NS uncommonly in Petane Group mudstone units horizontal cutting on Darkys Spur road, be- throughout Hawke’s Bay. The anal sinus of this tween Waipunga and Kaiwaka, 5 km WNW of specimen is very weak and faint, indicating that Devils Elbow, late Nukumaruan; 10 specimens). the sinus can be suppressed through injury. The However, it must be stressed that the two known one other lot of four specimens in GNS specimens of M. angustata are very similar, and assigned to this species was received from an at present it seems just as likely that this is a early collector at Wanganui (R Murdoch?) and distinct but very rare species. M. angustata bears an early label reading ‘Alcira angustata apparently differs from Phenatoma in its Hutton’, and a more recent one reading ‘Mi- much smaller size (the larger is 10.5 mm high, 102 AG Beu

compared with up to ca 25 mm in P. precursor sculpture and a much more strongly protruding and up to 35 mm in Recent specimens of P. rosea peripheral keel, but is otherwise similar to (Quoy & Gaimard, 1833)), its smoother and Paracomitas species in most characters, and a more evenly, consistently sculptured surface similar cochlespirine taxon could well have with very even, wide, low, closely spaced spiral given rise to P. protransenna. The smooth cords, with obscurely defined edges, and only peribasal cord, a prominent peripheral carina faint axial sculpture of growth lines in the spiral and a very similar shoulder sinus to that of interspaces (although both specimens assigned Paracomitas are present on all the close rela- here are abraded, and the apparently distinctive tives of Cochlepira (listed by Powell 1966:20). A sculpture might merely result from abrasion), its further helpful character of C. maora is the tall, shallower anal sinus, and its more evenly inflated clearly carinate protoconch, quite different protoconch of fewer whorls than in Phenatoma from the bulbous, paucispiral, smooth (lecitho- (M. angustata: Fig. 16H, K; Powell 1942, text- trophic) protoconch of most Cochlepira species. fig. B15, p. 38; P. rosea: Powell 1942, text-fig. This demonstrates that the keeled protoconch C21, p. 39). However, on the basis of the two of Paracomitas is not greatly discordant with known specimens, even the protoconch charac- other cochlepirine protoconchs although, of ters possibly could be based on aberrant or course, the great reliance placed on protoconch severely abraded individuals, as the protoconch characters by Powell (1942, 1966) is not fol- (Fig. 16K) is obviously abraded. In the circum- lowed here. In view of the strong teleoconch stances, assignment to Subfamily Clathurellinae similarity, Paracomitas is here referred to sub- (alongside Phenatoma) is provisional. Assign- family Cochlespirinae (sensu stricto). ment of extra-New Zealand taxa to a ‘genus’ Mitrellatoma clearly is unjustified, and further, Genus Paracomitas Powell, 1942 better-preserved specimens are required to clar- ify the status of M. angustata and the genus Paracomitas Powell 1942:61. Type species (by Mitrellatoma. original designation): Surcula castlecliffensis Marshall & Murdoch, 1919 (Pleurotoma gemmea Murdoch, 1900), Castlecliffian (Pleis- Subfamily Cochlespirinae tocene), New Zealand. Remarks. The subfamily Cochlespirinae (sensu stricto) was revealed as a distinct clade by Remarks. Species of Paracomitas are character-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Puillandre et al. (2008, fig. 1), although some ized by their small size, their prominent per- of the taxa formerly included in it on anatomi- ipheral carina, a rather weak peribasal cal grounds, such as Leucosyrinx and Comitas, angulation consisting of a smooth spiral belong in other, unnamed clades, and even the cord extending from the suture, a relatively species included in Comitas are polyphyletic. long, narrow anterior siphonal canal, a broad, Powell (1942) originally suggested a relation- relatively shallow anal sinus occupying the ship between Paracomitas and Comitas Finlay, width of the almost smooth, concave sutural 1926, although admitting that Paracomitas was ramp, a forward-extended outer lip below the ‘puzzling’. However, a relationship with Co- sinus, and a protoconch with a low, dome- chlespira Conrad, 1865 and similar genera shaped first whorl and a prominent, narrow, seems much more likely. The relatively early, median carina on the succeeding whorl. The rather generalized species Cochlepira maora earliest species in an apparently simple evolu- (Marshall & Murdoch, 1923) (Tahusyrinx tionary lineage is P. protransenna (Marshall & finlayi Powell, 1942; Maxwell 1992:152, pl. Murdoch, 1923), limited to Nukumaruan 21q,r; Kaiatan [late Eocene], McCulloch’s rocks. This species is unusual in having the Bridge, SCanterbury) has more complex spiral peripheral carina smooth, whereas later species Marine Mollusca of isotope stages of the last 2 million years in New Zealand 103

bear gemmae on the carina where it is crossed clear statement ‘Type and paratypes in Wanga- by low axial ridges. I (Beu 1970b) proposed a nui Museum’, so the illustrated specimen is subgenus Paracomitas (Macrosinus) for species considered to be the holotype. with a deeper anal sinus, more uniformly present spiral sculpture and a less protruding Other material examined. Nukumaruan: none periphery. It is conceivable that this group was was reported from Wanganui Basin by Fleming ancestral to Paracomitas, but the differences (1953); Devils Elbow Mudstone, top of Devils are great enough to suggest that the similarity is Elbow hill, Highway 2, ca 35 km N of Napier superficial, and Macrosinus is regarded here as (GS2220, V20/f8462, one; GS10849, V20/f8572, a distinct genus. The species revised here are three); above Te Onepu Limestone, tributary of those previously assigned to Paracomitas (sensu Porangahau Stream, 3 km S of Takapau, S stricto). Hawke’s Bay (GS2330, U23/f6382, one); head- waters of tributary of Waikoukou Stream, 3 km Paracomitas protransenna (Marshall & Mur- NNE of Ormondville, SHawke’s Bay (GS2369, doch, 1923) (Fig. 16A, I) U23/f6414, one); Maharakeke Road, W foot of Pukeroa Hill, Waipukurau (GS10857, U23/ Surcula protransenna Marshall & Murdoch f7049, one); Ashcott Road, NW foot of Pu- 1923:126, pl. 12, fig. 6. keora Hill, Waipukurau (GS10858, U22/f09, Parasyrinx protransenna. Finlay 1924:514. nine); Mairau Mudstone, Seafield Road, Bay- Paracomitas protransenna. Powell 1942:62; view, N of Napier (GS10875, V21/f6575, one); Fleming 1966:71; Tucker 2004:796. Mairau Mudstone, walkway W side of Roro-o- Paracomitas (Paracomitas) protransenna. Beu kuri, Onehunga Road, N end of Ahuriri & Maxwell 1990:327, pl. 42r; Maxwell ‘lagoon’, Napier (GS12386, V20/f05, six); Paul- 2009:249. sen Road, 2.5 km N of Rangitoto trig. station, SE of Takapau, S Hawke’s Bay (GS11479, Type material. Surcula protransenna, holotype U23/f6528, one). TM8287, with six paratypes TM82888293, from Waikopiro, near Ormondville, southern Distribution. Restricted to Nukumaruan rocks; Hawke’s Bay, Nukumaruan; ex Marshall & all records are from the upper siltstone forma- Murdoch Collection; stated by Marshall & tions of Petane Group in central Hawke’s Bay, Murdoch (1923:127) as ‘Waikopiro. (Seven and laterally equivalent units in SHawke’s Bay

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 specimens, Suter collection)’, that is, the types and SWairarapa, of Nukumaruan age. Speci- are from Suter’s collection from Waikopiro. mens are never abundant, but are more wide- The tube of specimens received from the spread in Hawke’s Bay Nukumaruan rocks Wanganui Museum remained in its original than above records suggest. condition when I examined it; five specimens were separated from the two in the entrance to Dimensions. Surcula protransenna, holotype: H the tube, which were identified by a small paper 12.3, D 4.6 mm; Marshall & Murdoch’s mea- label in P Marshall’s handwriting reading sured paratype, TM8288: H 9.7, D 3.5 mm; ‘type’. These are assumed to be the two speci- GS10858, Ashcott Road: H 13.0, D 4.5 mm; H mens whose dimensions were provided by 13.1, D 5.1 mm; H 13.2, D 4.6 mm; GS10849, Marshall & Murdoch (1923:127): ‘Length 13 Devils Elbow Mudstone, top of Devils Elbow mm; width, 5 mm. Another example: Length, hill: H 16.8, D 5.8 mm; GS2220, locality as last: 10 mm, width, 4 mm’ (see actual measurements H 12.6, D 4.4 mm. below). This indicates that the larger of the specimens labelled ‘type’ is the figured one. Remarks. Paracomitas protransenna is one of Marshall & Murdoch (1923:127) also made the the most distinctive of New Zealand conoi- 104 AG Beu

deans, easily recognized by its small size (H blue sandy clay’, Wanganui, Castlecliffian, also 1217 mm), its very prominent, smooth, nar- from Marshall & Murdoch collection. rowly convex peripheral carina, its obvious, smooth peribasal angulation, its elongate, Other material examined. Castlecliffian: re- graceful shape, its moderately deep, wide anal ported by Fleming (1953:192, 184, 199, 200, sinus, and its prominently keeled protoconch. 207) from Lower Kai-Iwi Siltstone and Omapu Therefore, despite its small size, it is useful as a Shellbed (OIS 23), Kaikokopu Shellbed and Nukumaruan index fossil, although it is only Upper Westmere Shellbed (OIS 19) and Upper moderately common, and seems to be recorded Kai-Iwi Siltstone (OIS 17) in Wanganui Basin; only from Hawke’s Bay and Wairarapa; most specimens in reference collection: coast between records are from siltstone and sandy siltstone Kai-Iwi & Okehu streams, Castlecliff (GS1163, beds of Petane Group, particularly the more R22/f6498, six); Lower Kai-Iwi Siltstone (OIS offshore units of highstand systems tracts. The 23), Castlecliff (GS4069, R22/f6400, seven); sutural ramp and the whorl surface between the data as last (GS4061, R22/f8392, six); Pinnacle two carinae are more strongly concave than in Sand (OIS 14), ‘the pinnacles’ gully, Castlecliff later species of Paracomitas, although it seems (GS4022 re-collection, R22/f6360, three); also likely that P. gemmea (Murdoch, 1900) is two good specimens with slightly more promi- simply its direct descendent. A few specimens nent peripheral nodules and a more prominent of P. protransenna have small nodules on the peribasal cord from 240250 m in the Mikonui- peripheral carina on early spire whorls, fading 1 well, off Hokitika (?early Castlecliffian) and out down the spire, and these more closely fragments of what is probably this species from resemble P. gemmea than completely smooth almost all other samples in the top 170520 m specimens do. of the well (see Beu 2010: 70). Paracomitas gemmea (Murdoch, 1900) (Fig. 16E, G) Distribution. Records range from OIS23-14, and it probably occurs in Tainui Shellbed (OIS Pleurotoma gemmea Murdoch 1900:217, pl. 20, 13), as well as in the Mikonui-1 well (?latest fig. 9. Nukumaruanearly Castlecliffian). However, Surcula castlecliffensis Marshall & Murdoch almost all are from siltstone units (the more 1919:255. offshore parts of highstand systems tracts), and Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Paracomitas gemmea. Powell 1942:62; Powell the offshore habitat of Paracomitas suggests 1966:29; Fleming 1966:71; Tucker 2004:412. that its occurrences are likely to be limited to Paracomitas castlecliffensis. Powell 1942:62, pl. fine-grained, offshore depositional facies. The 10, fig. 1; Powell 1966:29, pl. 2, fig. 1; Fleming real time range is likely to have been earliest 1966:71. Castlecliffian to Haweran, when it gave rise to Paracomitas (Paracomitas) gemmea. Maxwell P. augusta, and this species is of low reliability 1988:60; Beu & Maxwell 1990:368, pl. 49q; for biostratigraphy. Maxwell 2009:249. Dimensions. Pleurotoma gemmea, holotype: H Type material. Pleurotoma gemmea, holotype 12.7 (incomplete), D 5.3 mm; Surcula castle- TM6842 (Beu & Maxwell 1990, pl. 49q), from cliffensis, holotype: H 12.2 (incomplete), D 4.2 Castlecliff, Wanganui, ‘blue-clay cliffs, west of mm; GS4061, Lower Kai-Iwi Siltstone, Castle- Wanganui Heads’, Castlecliffian, from Mar- cliff: H 16.5, D 6.7 mm; H 15.0, D 5.7 mm; shall & Murdoch collection. Surcula castleclif- GS4069, as last: H 17.4, D 7.3 mm; H 17.8, D fensis, holotype TM8286, from ‘Castlecliff, in (slightly incomplete) 6.7 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 105

Remarks. Powell (1942:62) maintained Paraco- Barrier Island, E of Auckland. Suter’s speci- mitas castlecliffensis and P. gemmea as distinct mens (TM11078) are labelled ‘syntypes’ in species, claiming that P. gemmea has fewer, Suter’s handwriting, so it is not clear that the more prominent nodules on the peripheral NMNZ specimen (M.1715) was originally se- carina than P. castlecliffensis. However, Max- lected as the holotype, and it is here designated well (1988:60) and Beu & Maxwell (1990:368) as the lectotype. Paracomitas augusta powelli, pointed out that the two forms intergrade holotype NMNZ M.8840 (Marshall 1996), completely, and represent a single species. P. from 106 m, E of Stephens Island, Cook Strait, gemmea differs from P. protransenna in having 40842?S, 174810.6?E; two paratypes M.8841, a less prominent peripheral carina, which bears from the same locality. gemmae (short axial costae), in its taller and flatter interval between the two carinae, and in Other material examined. Recent, typical Para- having numerous fine, closely spaced spiral comitas augusta, 15 lots in NMNZ, all from NE cords over the entire surface below the periph- North Island: M.147932, 357447 m, E of eral carina, rather than being entirely smooth North Cape (four small); M.42261, 124 m, off over at least the last 34 whorls as in P. Poor Knights Islands, Northland (one); protransenna. P. gemmea also is larger and M.35840, 113121 m, W of Poor Knights wider and has a less prominent peripheral Islands (seven); M.35744, 80 m, off Cape Brett carina, smaller peripheral nodules, and more (four); M.11068, 207220 m, off Mayor Island prominent spiral cords below the carina than in (two); M.66590, 198273 m, NW of Mayor P. augusta. Island (11); M.61018, 293348 m, NW of Mayor Island (three); M. 11059, 494 m, off Paracomitas augusta (Murdoch & Suter, 1906) Mayor Island (one); M.60242, 482550 m, E (Fig. 16B, J) of Mayor Island (two); M. 61326, 203248 m, SE of Mayor Island (one); M.147933, 108113 Pleurotoma (Leucosyrinx) augusta Murdoch & m, N of Aldermen Islands (two); M.44432, 145 Suter 1906:286, pl. 22, figs. 1417. m, off Aldermen Islands (one); M.66723, Turris (Leucosyrinx) augusta. Suter 1913:472, 178248 m, SE of Aldermen Islands (one); pl. 21, fig. 3. M.59831, 139 m, ENE of Tolaga Bay (one); Paracomitas augusta. Powell 1942:62; Dell M.7960, 366 m, off Cape Kidnappers (one). 1956b:176; Boreham 1959:59; Powell 1966:29; P. augusta powelli form: 24 lots in NMNZ, all

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Tucker 2004:99. from W New Zealand, ranging from NW of Paracomitas augusta powelli Dell 1956b:56, fig. Ahipara, Northland (M.15803, 90 m; three) to 25; Powell 1979:229; Spencer & Willan 1996:29; Bradshaw Sound, Thompson Sound and Tucker 2004:785. Doubtful Sound in the W Southland Fiords; Paracomitas augusta augusta. Powell 1979:229; most material is from W Cook Strait. Spencer & Willan 1996:29. Paracomitas (Paracomitas) augusta augusta. Distribution. Typical Paracomitas augusta, with Spencer et al. 2009:211. a single prominent peribasal spiral cord, seems Paracomitas (Paracomitas) augusta powelli. to be limited to the E North Island, NMNZ Spencer et al. 2009:211. material ranging from off Cape Kidnappers to North Cape. The ‘powelli form’, with two Type material. Pleurotoma augusta, lectotype prominent peribasal spiral cords and several NMNZ M.1715 (Marshall 1996:28), two para- other obvious basal spiral cords, occurs along lectotypes TM11078 (Boreham 1959:59; Figs. the W coast of New Zealand from Ahipara, N 16B, J), Recent, dredged in 200 m off Great Northland, to W Otago. This is interpreted 106 AG Beu

here as one species, although the geographical augusta. Subdivision or not of the peripheral separation suggests that further study is justi- nodules by a median groove occurs along both fied. Records range from 80550 m, but the coasts, and most specimens have a weak groove deepest ones are transported chalky shells, and around the nodules. This form therefore seems the normal depth range of living specimens likely to be part of the variation of P. augusta. seems to be about 100300 m. Certainly, all Recent material in NMNZ is significantly smaller, narrower and with a Dimensions. Pleurotoma augusta lectotype: H markedly narrower peripheral spiral ridge 10.32, D 3.9 mm (Murdoch & Suter 1906:286); than in the Castlecliffian P. gemmea, and P. paralectotypes: TM11078: H 6.4, D 2.6 mm; gemmea and P. augusta clearly are distinct H 5.6, D 2.5 mm; NMNZ M.42261, off Poor species. The distinction between P. gemmea Knights Islands: H 11.6, D 4.1 mm; M.61018, (Castlecliffian) and P. augusta (Recent) offers NW of Mayor Island: H 11.0, D 4.0 mm; some possibility of biostratigraphical utility in M.59831, ENE of Tolaga Bay: H 9.8, D offshore facies (such as oil wells), but the 4.1 mm. precise age is unknown of the speciation event when P. augusta presumably evolved. Remarks. The type material of Paracomitas augusta has a single row of very prominent, laterally protruding, slightly obliquely set gem- Subfamily Raphitominae mae around a much narrower peripheral keel than in P. gemmea, and a prominent, narrow, Genus Gymnobela Verrill, 1884 smooth peribasal keel issuing from the suture. The gemmae are larger and the peribasal cord is Gymnobela Verrill 1884:157. Type species (by considerably more prominent than in P. gem- subsequent designation, Cossmann, 1896:63): mea, giving the shell a different, narrower, more Gymnobela engonia Verrill, 1896, Recent, deep finely sculptured appearance than either fossil water, North Atlantic. species. Other than a few weak spiral threads on the area between the two keels, the teleoconch is Spergo Dall 1895:680. Type species (by subse- almost smooth, apart from faint growth ridges. quent designation, Dall 1918:331): Spergo gla- The protoconch also is even more strongly diniformis Dall, 1895, Recent, off Hawaii. carinate than in either fossil species, but this

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 possibly results from better preservation. Clarke 1959:234. Type species (by origi- Dell (1956b:56, fig. 25) named a subspecies nal designation): Pleurotomella (Theta) lyronu- Paracomitas augusta powelli, from depths of Clarke, 1959, Recent, abyssal, North 106 and 137 m in Cook Strait. This form differs Atlantic. from P. augusta in having two (rather than one) prominent, well separated peribasal keels, Speoides Kuroda & Habe in Habe 1961:77; slightly more prominent peripheral nodules, appendix, p. 29 Type species (by monotypy): the peripheral nodules subdivided into two Speoides yoshidai Kuroda & Habe, 1961, Re- rows by a deep groove, and more prominent cent, deep water, Japan to New Caledonia and secondary spiral cords. Examination of the 24 N New Zealand. lots of specimens in NMNZ (see above) showed that the size, shape and sculpture are little different from those of P. augusta, apart from Remarks. Sysoev & Bouchet (2001:305) the consistent presence of two peribasal cords regarded Theta and Speoides as synonyms that are more prominent than the others, rather of Gymnobela Verrill, 1884, pointing out than one on the E North Island form of P. that Spergo Dall, 1895 is probably a further Marine Mollusca of isotope stages of the last 2 million years in New Zealand 107

synonym, as had already been suggested by 11251150 m, off Castlepoint (one); M.59718, Powell (1966:137). Bouchet & Ware´ n (1980:61, 999984 m, N Mernoo Slope, Chatham Rise figs. 128, 252) illustrated the diagonally reticu- (one); M.59541, 17231549 m, 42841.70?S, late protoconch of Gymnobela characteristic of 174828.00?E, head of Hikurangi Trench be- the Raphitominae. Pleurotoma gypsata Watson, tween Kaikoura and Mernoo bank (one, lar- 1881 from deep water around central E New gest; Fig. 16D). Zealand has been referred to Paracomitas pre- viously (Powell 1942, 1979) but is similar to Distribution. Material of Gymnobela gypsata Gymnobela yoshidai and is illustrated here to has been collected so far from off East Cape show that it is not correctly referred to Para- to the N Chatham Rise, in 7401723 m. comitas (Speoides yoshidai Kuroda & Habe in Habe 1961:77, pl. 38, fig. 17, appendix p. 29; Dimensions. Pleurotoma gypsata, syntypes: H Habe 1964:120, pl. 38, fig. 17; Spergo (Speoides) 40.8, D (incomplete) 17.7 mm; H 36.8, D yoshidai: Powell 1966:137, pl. 22, figs. 11, 12; (incomplete) 15.8 mm; large illustrated speci- Hasegawa et al. in Okutani 2000:667, pl. 332, fig. men (Fig. 36D), NMNZ M.59541, NW Mer- 237; Gymnobela yoshidai: Sysoev & Bouchet noo Slope: H 47.9, D 21.3 mm. 2001:308, figs. 134144, 173). Remarks. Pleurotoma gypsata is a large (H Gymnobela gypsata (Watson, 1881) (Fig. 16D, F) 4050 mm) Gymnobela species that has been collected in small numbers in recent years in Pleurotoma gypsata Watson 1881:413; Watson deep water (700 m) off central E New 1886:297, pl. 25, fig. 1. Zealand. This species has no bearing on rela- Surcula gypsata. Suter 1899b:69; Suter 1913:486, tionships within the lineage of small Paracomi- pl. 21, fig. 13. tas species discussed above. Paracomitas gypsata. Powell 1942:62; Dell The material of Gymnobela yoshidai illu- 1963a:176, pl. 2, figs. 10, 11; Powell 1966:29; strated by Sysoev & Bouchet (2001, figs. Powell 1979:230; Spencer et al. 1996:29; Tucker 134144, 173) is quite variable in relative tele- 2004:444. oconch width and nodule prominence, but G. Paracomitas (Paracomitas) gypsata. Spencer yoshidai differs from G. gypsata in its narrower et al. 2009:211. shape, taller spire, markedly narrower and more steeply inclined sutural ramp, and smaller and

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 less obviously oblique peripheral nodules. As Type material. Pleurotoma gypsata Watson, much of the material of G. yoshidai recorded by two syntypes BMNH 1887.2.9.979980, from Sysoev & Bouchet (2001:308) is from New HMS‘ Challenger’ Sta. 169, off East Cape, New Caledonia and the Norfolk Ridge, as far south Zealand, in 1280 m. Watson (1886:298) clearly as 24857?S, it is not surprising to find that this or a listed ‘two dead and broken specimens’ (Fig. similar species extends its range further south 16F). Therefore, although the larger syntype down the Norfolk and Kermedec Ridges to was illustrated by Dell (1963a, pl. 2, figs. 10, 11) encompass northern New Zealand (NMNZ and identified as a ‘type’, under ICZN Article five lots, from 514540 m, SNorfolk Ridge, 74.6 this usage of ‘type’ does not constitute NW of Cape Reinga, to 924712 m, Rumble V designation of a lectotype. volcano, SKermadec Ridge, off Bay of Plenty).

Other material examined. NMNZ M.151262, Genus Pleurotomella Verrill, 1873 865741 m, Calyptogena Bank, Ritchie Ridge, off Mahia Peninsula (two); M.118135, 900 m, Pleurotomella Verrill 1873:15. Type species (by SE of Cape Kidnappers (one); M.74976, monotypy): Pleurotomella packardi Verrill, 108 AG Beu

1873, Recent, bathyal, North Atlantic (Bouchet form a uniform group with strongly inflated, & Ware´ n 1980:38, figs. 96, 97, 216; Fig. 17B, C, evenly rounded whorls, apart from the concave D, G, H). sutural ramp, and with a strongly and rapidly contracted base and prominent, narrow, and in Systenope Cossmann 1889:289. Type species (by most species obviously curved axial ridges. This original designation): Raphitoma (Systenope) is the genus for which Hedley (1922) and Powell polycolpa Cossmann, 1889, Eocene, Paris Basin. (1944, 1966, 1979) used the name Asperdaphne in Australia and New Zealand. As usual, Tasmadaphne Laseron 1954:45. Type species (by Powell (1966:132) proposed a distinct subge- original designation): Tasmadaphne spicula La- nus, Anomalotomella, for species with a dome- seron, 1954, Recent, New South Wales (Powell shaped, paucispiral protoconch, as distinct 1966:124, pl. 19, fig. 19) (new synonym). from the ‘typical’ species with a conical, diagonally cancellate raphitomine one, but Fusidaphne Laseron 1954:47. Type species (by these are developmental types within one genus original designation): Fusidaphne Laser- (see Numanami [1996:214222] for discussion on, 1954, Recent, New South Wales (Powell and illustration of Antarctic species assigned to 1966:124, pl. 20, fig. 1) (new synonym). Anomalotomella). Tasmadaphne spicula Laser- on (1954:46, pl. 11, figs. 242, 243) appears to be Anomalotomella Powell 1966:132. Type species a further species of Pleurotomella with a (by original designation): Pleurotomella anom- lecithotrophic protoconch, and Fusidaphne bul- alapex Powell, 1951, Recent, (new lata Laseron (1954:46, pl. 12, figs. 246, 247; synonym). Powell 1966:124, pl. 20, fig.1) is probably another. Neopleurotomoides Shuto (1971:5; Remarks. The characters of the type species of type species [by original designation]: Clathur- Pleurotomella, P. packardi Verrill, 1873 (Fig. ella rufoapicata Schepman, 1913, Recent, bath- 17B, C, D, G, H; dimensions of illustrated yal, Indonesia) was distinguished from specimen: H 9.2, D 4.9 mm) were clarified by Pleurotomella by Bouchet & Ware´ n (1980:43) Bouchet & Ware´ n (1980, figs. 96, 97, 216), who because of its different protoconch sculpture, of commented that they included a far less diverse medially keeled whorls with vertical (axial) group of species in the genus than the large, riblets, unlike the usual raphitomine diagonally heterogeneous assemblage included by Powell reticulate, evenly convex whorls of plankto- Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (1966). Powell’s (1966) lack of recognition of trophic species of Pleurotomella. In this light, the ‘Australasian’ genus Asperdaphne in this the generic synonymy proposed here is tenta- assemblage implies that he was not aware of the tive, and the protoconch characters of type characters of P. packardi. Four Atlantic species species of genera will need to be compared included in Pleurotomella by Bouchet & Ware´ n before the synonymy is accepted. The New do not belong here (P. anceyi (Dautzenberg & Zealand Recent species referred to Asper- Fischer, 1897), P. sandersoni Verrill, 1884, P. daphne, other than A. ula (Watson, 1881) lottae Verrill, 1885 and P. megalembryon Daut- (referred below to Propebela Iredale, 1918), zenberg & Fischer, 1896; the genus Azorilla that is, A. expeditionis Dell (1956a:137, figs. Nordsieck, 1968 is available for P. lottae and P. 204, 205; Powell 1979, fig. 56.4; Fig. 17A, F) megalembryon; Bouchet & Ware´ n [1980:49]; and A. aculeata (Webster, 1906) (Suter 1913, pl. Morassi & Bonfitto [2006:231] referred Pleur- 22, fig. 10) and several unnamed species otomella anceyi to Cryptodaphne Powell, 1942). in NMNZ, have inflated, evenly rounded The other North Atlantic Pleurotomella species whorls without a peripheral carina, and with Marine Mollusca of isotope stages of the last 2 million years in New Zealand 109 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 17 (A,F) Pleurotomella expeditionis (Dell), SEM, RM5590, Portobello sta. Mu7045, 540490 m, Papanui Canyon, off Otago Peninsula, RV ‘‘Munida’’; A, height 8.2 mm; F, protoconch of A. (B,C,D,G,H) Pleurotomella packardi Verrill, type species of Pleurotomella Verrill, 1872; SEM, 2 specimens, MNHN, Biogas Sta. CP37, Golfe de Gascoigne, E Atlantic, 2175 m, 47834’N, 08839’W; B, protoconch; C,D,G,H, second specimen, height 9.2 mm; C, protoconch of D; G, granulose sculpture on teleoconch sutural ramp; H, sculpture of protoconch in C. (E) Amalda (Baryspira) olsoni Beu, GS11225, V20/f8002, Darkys Spur Formation (late Nukumaruan), Darkys Spur, W of Devils Elbow, Hawke’s Bay; largest specimen seen, height 42.8 mm. 110 AG Beu

a reversed-L-shaped raphitomine anal sinus, The one obvious difference between Austra- and are congeneric with most of the Australian lasian species and the North Atlantic species species referred to Asperdaphne by Hedley included in Pleurotomella by Bouchet & Ware´ n (1922, pl. 54, figs. 170173; pl. 55, figs. (1980:3441) is that, once again, most Atlantic 174176), Powell (1942, pl. 6, figs. 7, 8) and species have a tall, diagonally cancellate raphi- Cotton (1947:22), and with Pleurotomella pack- tomine protoconch, whereas most Australasian ardi. However, these species clearly are not species have a blunt, paucispiral, spirally ridged congeneric with Daphnella versivestita Hedley, to finely and irregularly nodulose protoconch 1912, the Australian Recent type species of (Fig. 17F). However, the type species, P. pack- Asperdaphne Hedley, 1922 (Hedley 1912:148, ardi, has a blunt, short protoconch with fine, pl. 63, fig. 12; Laseron 1954:45, pl. 11, fig. 230; rather irregular spiral threads over the upper- Powell 1966, pl. 20, fig. 16). A. versivestita most third and short, diagonally arcuate ridges closely resembles Tritonoturris robillardi (H. over the lowest two-thirds of each whorl (Fig. Adams, 1869) (Recent, Mauritius; Powell 17B, C, H) and apparently has lecithotrophic 1966, pl. 20, fig. 23), type species of Tritono- development. Greater enlargement showed that turris Dall, 1924, in all characters. The only the protoconch is quite coarsely granulose and obvious difference between them is that T. the teleoconch more finely so (Fig. 17G). Again robillardi has a typical raphitomine cancellate this protoconch difference is solely a develop- protoconch, whereas that of A. versivestita is mental difference, and Pleurotomella is adopted ‘paucispiral, blunt tipped, closely packed with here for the Australian and New Zealand species minute spiral threads and developing two spiral previously assigned to Asperdaphne. keels on the last whorl’ (Powell 1966:127), that Several genera are represented among the is, it has lecithotrophic development. A. versi- species included in Asperdaphne by Hedley vestita also resembles Tritonoturris species in its (1922), but at least some Australian species relatively large size (H 23, D 9 mm; Hedley are correctly referred to Pleurotomella (see 1912:148) compared with most other species below), although protoconchs will need to be assigned to Asperdaphne by Hedley and Powell. checked to be sure that some do not belong in Asperdaphne appears to be an earlier name for Neopleurotomoides. Some other species in- Tritonoturris. Asperdaphne versivestita resem- cluded by Hedley (1922) or Cotton (1947) in bles macandrewi (EA Smith, 1882) Asperdaphne may well belong in Pleurotomella, very closely (Li et al. 2010:198, figs. 58,11), but require further research. Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 apparently differing only in protoconch shape Fenestrodaphne pulchra Powell (1944:61, pl. (tall and multispiral in T. macandrewi, short 6, fig. 10), type and only species referred to and paucispiral in A. versivestita). Fenestrodaphne Powell, 1944, is very similar to Sysoev (in Poppe 2008:750, pl. 670, fig. 1) ‘Asperdaphne’ tasmanica (Tenison Woods, used Asperdaphne for a species from the Phi- 1877) (Hedley 1922, fig. 12, p. 343), but neither lippines, A. peradmirabilis (E. A. Smith, 1879), can be referred to Pleurotomella. Rather, they but this also seems to be a relatively weakly seem to be species of Jeffreys, 1870. sculptured species of Tritonoturris. The other ‘Asperdaphne’ vestalis Hedley (1922:344, fig. 13; Australian and New Zealand species referred to Laseron 1954:46, pl. 11, figs. 240, 241) is also a Asperdaphne are smaller than Tritonoturris species of Taranis (WM8381, 120 m, off Port versivestita and lack the prominent axial ridges, Jackson; two specimens examined). Hedley effuse outer lip, short last whorl and short, (1922:341) noted of ‘Asperdaphne’ esperanza widely open anterior siphonal canal that char- (May, 1910) that it is closely related to acterise Tritonoturris; most are typical species Bathytoma gratiosa Suter, 1908, from southern of Pleurotomella. New Zealand, and as B. gratiosa is a typical Marine Mollusca of isotope stages of the last 2 million years in New Zealand 111

species of Taranis, A. esperanza evidently probably synonyms of Pleurotomella, Neopleur- belongs in Taranis also. The late PA Maxwell otomoides or Taranis, but require reassessment. (pers. comm.) recognized a New Zealand early Fleming (1953:226) included a record in his Cenozoic species of Taranis with a tall, diag- list of molluscs from Pinnacle Sand (OIS 1514) onally reticulate, planktotrophic raphitomine of ‘New genus aff. Asperdaphne n. sp.’, but protoconch. Bouchet & Ware´ n (1980, figs. examination of the single specimen showed that 272273) also illustrated a similar, if slightly it belongs in Neoguraleus (GS4098, R22/f6425, shorter protoconch for the Atlantic species uppermost 23 cm of Pinnacle Sand in the Taranis malmii (Dall, 1889). Taranis therefore Castlecliffcoastalsection,in‘thepinnacles’gully, belongs in Subfamily Raphitominae, being the only locality where this interval is accessible). simply a raphitomine that has lost its radula Australian and New Zealand species tenta- (Taylor et al. 1993), and the Subfamily Tar- tively referred here to Pleurotomella (pending aninae of Kantor & Sysoev (1989) and Taylor assessment of a possible position in Neopleur- et al. (1993) was synonymized with Raphitomi- otomoides) are: nae by Bouchet et al. (2005:256) and other Pleurotomella aculeata (Webster, 1906), Recent, authors. Taranis therefore is closely related New Zealand. phylogenetically to Pleurotomella, which ex- Pleurotomella aculeola (Hedley, 1915) (Laseron plains Powell’s (1944) association of T. pulchra 1954:47, pl. 11, figs. 244, 245), Recent, New with ‘Asperdaphne’. Material illustrated in the South Wales. New Caledonian fauna (MNHN ‘Turridae Pleurotomella amplecta (Hedley 1922, pl. 54, group’, in preparation) also suggests that figs. 171, 172), Recent, Australia. many extra-New Zealand species assigned to Pleurotomella balcombensis (Powell 1944, pl. 6, Mioawateria Vella, 1954 are actually species of fig. 7), Fyansford Formation, Balcombe Bay, Taranis with a tall, diagonally cancellate, Melbourne, middle Miocene. planktotrophic raphitomine protoconch and a Pleurotomella brenchleyi (Angas, 1877) (Laser- wide sutural ramp, and Mioawateria is possibly on 1954:45, pl. 11, figs. 231, 232), Recent, New simply a synonym of Taranis. Bonfitto & South Wales. Morassi (2006) proposed the new genus Pleurotomella buccinoides (Shuto 1983:18, pl. 2, Acanthodaphne for A. sabellii Bonfitto & Mor- figs. 12, 13, text-fig. 4; proposed in Tritonotur- assi, 2006 (Recent, Gulf of Aden; type species), ris), Recent, Sydney Harbour. A. abbreviata (Schepman, 1913) (Recent, In- Pleurotomella bullata (Laseron 1954:47, pl. 12,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 donesia) and A. (Laws, 1947) (Early figs. 246, 247), Recent, New South Wales. Miocene, Northland, New Zealand), and com- Pleurotomella capricornea (Hedley 1922, pl. 54, mented that some species of Mioawateria are fig. 173), Recent, E Australia. similar to Acanthodaphne species, so the rela- Pleurotomella chapplei (Powell 1944:59, pl. 6, tionships of Acanthodaphne, Mioawateria and fig. 6), Muddy Creek, western Victoria, middle Taranis need reconsideration. Miocene. In summary, (1) the type species of Asper- Pleurotomella compacta (Hedley 1922, pl. 55, daphne is a lecithotrophic species of Tritono- fig. 174), Recent, Australia. turris; (2) most other Australian and New Pleurotomella contigua (Powell 1944, pl. 6, Zealand species that have been referred to fig. 8), Fyansford Formation, Altona Bay, Asperdaphne belong in Pleurotomella; (3) some Melbourne, middle Miocene. of the other species previously referred to Pleurotomella cuspidata (Chapple, 1934), Asperdaphne belong in Taranis; and (4) many Fyansford Formation, Balcombe Bay, of the other genera previously proposed for Melbourne, middle Miocene. Raphitominae similar to Pleurotomella (parti- Pleurotomella expeditionis (Dell, 1956), Recent, cularly by Powell, Laseron and Shuto) are New Zealand (Fig. 17A, F). 112 AG Beu

Pleurotomella hayesiana (Angas, 1877) (Laser- oblique growth ridges parallel to the lip. The on 1954:45, pl. 11, figs. 233, 234), Recent, New spiral sculpture consists of many narrow, South Wales. closely spaced, smooth, convex-crested cords Pleurotomella rugosa (Laseron, 1954) (Laseron over the entire teleoconch, other than a weakly 1954:46, pl. 11, figs. 235237), Recent, New sculptured sutural ramp; a rounded, smooth South Wales. subsutural fold is prominent initially, but fades Pleurotomella sepulta (Laseron, 1954) (Laseron out down the spire to be little more prominent 1954:46, pl. 11, figs. 238, 239), Recent, New than the other fine spiral cords on the last few South Wales. whorls. The axial sculpture consists of moder- Pleurotomella spicula (Laseron 1954:47, pl. 11, ately prominent, evenly rounded, strongly figs. 242, 243), Recent, New South Wales. opisthocline folds, inclined forwards at ca 458 Pleurotomella vercoi (G.B. Sowerby III, 1896) to the horizontal, with interspaces each equal in (Hedley 1922, pl. 55, figs. 175176), Recent, width to one fold, forming low, spirally elon- Australia. gate nodules on the spiral cords where they Pleurotomella n. sp. A and B of Long (1981:46, cross the axial folds. The aperture is simple, pl. 7, figs. 8, 9; as Asperdaphne), Aldingan (late unthickened, without a parietal callus or strom- Eocene), Brown’s Creek Clay and Glen Aire boid notch; the anterior end is incomplete on all Clay, SE Victoria. material, but has a weak, rounded peribasal angulation and contracts only weakly below Genus Onoketoma n. gen. that, over the neck; the siphonal canal evidently is relatively short. Type species. Insolentia solitaria King, 1933, early Nukumaruan Late Pliocene bathyal fauna Remarks. Onoketoma n. gen. seems most simi- in Onoke Formation, Palliser Bay, SWairarapa. lar to species of Pleurotomella with a lecitho- trophic protoconch, but the protoconch is still Diagnosis. A genus assigned to the Raphitomi- simpler and smoother than in any similar, nae on the basis of its deep, reversed-L-shaped previously named Raphitominae. On the tele- posterior sinus and only very short adapical oconch, its initially prominent subsutural cord, limb to the sinus, but readily distinguished from fading out down the spire, its smooth, weakly all previously named genera of Raphitominae carinate periphery, its only weakly contracted by its low, dome-shaped protoconch of 1.5 base, its flat sutural ramp, its almost straight

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 smooth, rounded whorls, passing gradually into whorl profile between the peripheral and the teleoconch by the acquisition of axial peribasal angulations, and its sculpture domi- sculpture; a few distinctive, curved axial ridges nated by even, closely spaced spiral cords are are present on the last half-whorl of the characters distinguishing Onoketoma from protoconch. The spire is moderately tall, gra- Pleurotomella. The spiral sculpture is finer date, with a relatively narrow, flat, steeply and more closely spaced, the axial threads are sloping sutural ramp equal in width to the much less obvious, the teleoconch shape is vertical whorl side above the ramp, followed taller and narrower with a taller spire and a below by a narrowly rounded, smooth periph- more steeply inclined, flatter sutural ramp, and eral angulation lacking prominent nodules. The the axial ribs are more prominent than in posterior (anal) sinus is deeply and evenly Taranis. concave, widely open, occupying the full width King (1933:350) had only the incomplete of the sutural ramp, swinging forward only a holotype when he proposed ‘Insolentia’ solitar- little adapically to the narrow subsutural ia. Powell (1942:69) reassigned it to Comitas cord, but followed below by a long, forward- ‘without prejudice’, as more appropriate than a swinging outer lip, with obvious, fine, steeply position in Insolentia on the basis of the Marine Mollusca of isotope stages of the last 2 million years in New Zealand 113

protoconch. Beu & Maxwell (1990) suggested a weaker axial sculpture, without any axial position in Austroturris Laseron, 1950 because costae, in their shorter spire, in their shallower of a general resemblance to the Australian posterior sinus, in their more prominent and Recent species Austroturris steira (Hedley, more widely spaced spiral cords, and in their 1922) (Laseron 1954, pl. 1, figs. 3, 4) and, in shorter protoconch, which is ‘closely axially particular, Austrocarina recta (Hedley, 1903) costate’ all over, with ‘two weak spiral keels’ (Laseron 1954, pl. 4, fig. 88) but I have since (Powell 1944:60), and is distinguished from the examined these species (material in GNS); they teleoconch only by the addition of further are very much smaller and have a much spiral cords. This contrasts strongly with the shallower anal sinus than Onoketoma solitaria, smooth protoconch with a few brephic axial and any resemblance is superficial. Further ridges where it joins the teleoconch in Onoke- material from Palliser Bay has enabled the toma solitaria, but it is helpful in suggesting a taxonomic position of O. solitaria to be reas- relationship between Onoketoma and Taranis. sessed, although all specimens are a little incomplete. King (1933:350) noted that the Etymology. From the name of the formation folds are inclined to the shoulder angulation and locality where the type species occurs, at a prominent angle, and it is about 458in the Onoke Formation at Lake Onoke, Palliser available material (Fig. 18B, C, G), indicating Bay; combined with ‘toma’ (abbreviation of an unusually long, forward-swinging anterior Pleurotoma), used in the names of many genera lip. The deep reversed-L-shaped sinus, with of Conoidea. Gender feminine. almost no adapical limb but a long anterior limb, seems to indicate a position in subfamily Onoketoma solitaria (King, 1933) (Fig. 18B, C, Raphitominae. The entire surface is covered G, H) with fine, low, closely spaced, convex-crested spiral cords or threads. All specimens examined Insolentia solitaria King 1933:350, pl. 36, fig. have a simple aperture with thin lips, and no 11. columellar or parietal callus, although none has ?Comitas solitaria. Powell 1942:60; Fleming a complete aperture or anterior siphonal canal. 1966:71; Powell 1966:29. One of the few other conoideans I am aware Austroturris(?) solitaria. Beu & Maxwell 1990. of that resembles Onoketoma solitaria closely is p. 419; Tucker 2004:918. Plicisyrinx binicostata Sysoev & Kantor, 1986 ‘Insolentia’ solitaria. Maxwell 2009:249.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (Kantor & Sysoev 2006:217, pl. 109C) but this has a shallower sinus and lacks obvious spiral Type material. Insolentia solitaria, holotype cords, and was placed in subfamily Cochlespir- (Fig. 18B) TM8322, from Onoke Formation, inae by its authors. The similarity is evidently cliffs east of Lake Ferry, which is on Lake superficial. It is likely that Onoketoma solitaria Onoke, Palliser Bay, SWairarapa. The age of will be found living in bathyal depths around the Palliser Bay Pliocene bathyal fauna is New Zealand. Recent specimens will allow the accepted as early Nukumaruan because of the subfamilial position to be reassessed using occurrence of occasional specimens of Zygo- radular characters. Of genera referred pre- chlamys delicatula. I am still not aware of any viously to the Raphitominae, the nearest in confirmed records of Z. delicatula older than teleoconch characters to Onoketoma solitaria is Nukumaruan. Fenestrodaphne Powell, 1944, but I suggest above that this is a synonym of Taranis Other material examined. Nukumaruan: Jeffreys, 1870. Taranis pulchra (Powell, 1944) GS15629, R28/f078, cliffs between Lake Ferry and the similar Recent species T. tasmanica and Whangaimoana, Palliser Bay (seven speci- differ from Onoketoma solitaria in having much mens). 114 AG Beu Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 18 (A,DF,I,J) Propebela abnormis (Hutton), GS10849, V20/f8572, 2 specimens, Devils Elbow Mudstone (late Nukumaruan), top of Devils Elbow hill, Hawke’s Bay; A, F, J, height 5.5 mm; F, protoconch of A; J, granules on protoconch in F; D, E, I, height 5.9 mm; E, protoconch of D; I, granules at protoconch termination. (B,C,G,H) Onoketoma solitaria (King), type species of Onoketoma n. gen., Onoke Formation (early Nukumaruan), cliffs E of Lake Ferry, Palliser Bay; B, holotype, TM8322, H 12.5, D 5.1 mm; C,G,H, GS15629, R28/f078; C, SEM, height 8.1 mm; G, H 12.4, D 4.9 mm; H, protoconch of C. Distribution. Onoketoma solitaria is known so it seems likely that it eventually will be found in far only from the bathyal Nukumaruan fauna the Recent bathyal fauna of New Zealand, in Onoke Formation at Palliser Bay. However, where a large number of conoidean taxa Marine Mollusca of isotope stages of the last 2 million years in New Zealand 115

remains unnamed (Spencer et al. 2009:211); few Belalora Powell 1951:171. Type species (by of the Onoke Formation taxa have proven to original designation): Bela pelseneeri Strebel, be distinct from living ones. 1908, Recent, Antarctica.

Dimensions. Holotype, TM8322: H 12.5, D 5.1 Paramarshallena Long 1981:27. Type species mm; GS15629, cliffs E of Lake Ferry, almost (by original designation): Paramarshallena pro- complete specimen: H 12.4, D 4.9 mm; most pebeloides Long, 1981, late Eocene (Aldingan), complete specimen: H 10.9, D 4.1 mm; H Brown’s Creek Clay, Johanna coast, SVictoria, (incomplete) 11.0, D 4.5 mm. Australia (new synonym).

Remarks. Little can be added to the comparison under the generic heading above. On a few Remarks. Beu & Maxwell (1990:328) suggested specimens the axial folds fade out before the that Antiguraleus Powell, 1942, with a New last whorl, rendering the teleoconch unusually Zealand type species, is a synonym of Oenopota narrow, but other equally large specimens have Mo¨ rch (1853:37; type species: Fusus pleuroto- axial folds continuing to the end of the last marius Couthouy, 1838 [Buccinum pyrami- whorl. The fine spiral cords vary from close and dale Stro¨ m, 1788], Recent, Norway to Iceland even to more widely spaced; those with widely and Spitzbergen) (Bogdanov 1990, fig. 447; spaced cords appear as if every second one were Kantor & Sysoev 2006, pl. 116J). This was suppressed, but they intergrade completely with based on the similarity in both teleoconch and the more common, evenly and finely sculptured protoconch characters between New Zealand shells. and Australian species referred to Antiguraleus by Powell (1942) and to Paraguraleus by Powell (1944), and the North Atlantic species Subfamily Oenopotinae referred to Oenopota by Bouchet & Ware´ n (1980:6675). In particular, the characteristic Genus Propebela Iredale, 1918 low, dome-shaped protoconch sculptured with many closely spaced microscopic granules is Propebela Iredale 1918:32. Type species (by identical in some species placed in the two original designation): Murex Monta- genera (Bouchet & Ware´ n 1980, figs. 258269). gu, 1803, PlioceneRecent, NE Atlantic. Bouchet & Ware´ n (1980:6667) listed 12 gen-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 eric names as synonyms of Oenopota, including Antiguraleus Powell 1942:146. Type species (by two of Powell’s (1951) apparently ‘restricted original designation): Antiguraleus otagoensis Antarctic’ genera: Propebela Iredale, 1918; Powell, 1942, Pleistocene(?) and Recent, New Turritomella, Nodotoma, Funitoma, Cestoma, Zealand (new synonym). Granotoma, Nematoma, Curtitoma, Venustoma and Canetoma, all of Bartsch (1941a,b), and Paraguraleus Powell 1944:49. Type species (by Belalora and Lorabela of Powell (1951). Since original designation): (Paraguraleus) then, Bogdanov (1987) has proposed Subfamily balcombensis Powell, 1944, middle Miocene, Oenopotinae, and (Bogdanov 1990) revised the Victoria, Australia. Synonymized with Antigur- boreal taxa of this group in detail. Hasegawa aleus by Powell (1966:106) (new synonym). et al. in Okutani (2000, pls. 325327) and Kantor & Sysoev (2006, pls. 112120) also Lorabela Powell 1951:171. Type species (by have illustrated many of the boreal species original designation): Bela theilei Powell, assigned to these genera, with clear colour pho- 1951, Recent, Antarctica. tographs. Bogdanov (1990) treated Oenopota, 116 AG Beu

O. (Nodotoma) Bartsch, 1941, Curtitoma previously assigned to Antiguraleus, and a Bartsch, 1941, Granotoma Bartsch, 1941, Obe- short, paucispiral, granulose protoconch, with sotoma Bartsch, 1941 and Propebela and P. a few low spiral cords in some species, but not (Canetoma) Bartsch, 1941 as distinct generic the loosely coiled, strongly spirally sculptured groups, while listing several of Bartsch’s protoconch of Oenopota. The supposedly re- (1941a) genera as synonyms of these. His stricted ‘Antarctic genera’ Lorabela and Bela- generic classification was followed by both lora of Powell (1951) also differ from Propebela Hasegawa et al. in Okutani (2000) and Kantor in only trivial protoconch characters, and seem & Sysoev (2006). to be further synonyms of Propebela, as sug- Here I adopt Propebela Iredale, 1918 as the gested previously by Bouchet & Ware´ n (1980). genus to contain ‘Asperdaphne’ ula and other The New Zealand and Australian species pre- Australian and New Zealand species previously viously assigned to Antiguraleus and Paragur- referred to Antiguraleus Powell, 1942 and aleus (which was synonymized with Paraguraleus Powell, 1944. Most of the remain- Antiguraleus by Powell 1966:106) seem to me ing possible synonyms in Bouchet and Ware´ n’s to belong in Propebela. These include Antigur- (1980) list are clearly considered by workers in aleus stellatomoides Shuto (1983:14, pl. 2, figs. the relevant regions to be distinct genera. 8, 9, text-fig. 3), from off North Reef, Queens- Bogdanov’s (1987) Subfamily Oenopotinae ten- land. Shuto (1983) mentioned several similar tatively was ranked as a separate subfamily of species (Mangilia woodwardiana Melvill, 1917, Conidae by Taylor et al. (1993:166), with Persian Gulf; Pleurotoma angicostata Reeve, genera listed as recognized by Bogdanov 1844; and Mangilia misera Thiele, 1925, Agul- (1990), and well as by Bouchet et al. has Bank, South Africa, a species apparently (2005:256), Kantor & Sysoev (2006:221), and not treated in Kilburn’s revisions of South Hasegawa (2009:338350, figs. 350412) and so African Conoidea) that may well be further is adopted here. species of the cosmopolitan genus Propebela. It is clear from Bogdanov’s (1990) mono- The distribution of species assigned to Prope- graph that the similarities between Antiguraleus bela seems bipolar at present, but the addition and some genera closely related to Oenopota of Shuto’s (1983) tropical species suggests that are not due to convergence; they are closely this is the result of the poor knowledge of small related phylogenetically, and some are prob- tropical deep-water conoideans, which are en- ably congeneric. However, the species assigned ormously diverse.

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 by Powell (1942) to Antiguraleus are much Long (1981:27) proposed Paramarshallena more closely similar to species assigned by for a single Australian Eocene species, P. prope- Bogdanov (1990) to Propebela Iredale, 1918 beloides. His description of the protoconch than to Oenopota. The type species, Propebela (‘paucispiral, dome-shaped, of 1.52.0 whorls, turricula (Montagu, 1803) (Bogdanov 1990, tip partly immersed, first whorl smooth, second figs. 368370, 404D,E; Kantor & Sysoev developing orthocline axial ribs before merging 2006, pl. 120E,F) is closely similar in all into teleoconch’) and the illustration of the characters to such New Zealand species as standard Propebela teleoconch show that this is Antiguraleus subtruncatus (Powell 1942, pl. 8, a species of Propebela, in the sense interpreted fig. 1) and A. abernethyi, A. multistriatus and A. here, assuming that the protoconch actually fusiformis of Dell (1956a, pl. 20, figs. 200, 202, bears microscopic granules. The small size (H 206). The many other boreal species assigned to 8.613.5 mm), the weak spiral sculpture, the Propebela also are very similar, with a shoulder peripheral nodules and the protoconch charac- angulation defining a narrow to moderately ters rule out a relationship with Marshallena.In wide sutural ramp, weak to moderately promi- my opinion Paramarshallena is a further syno- nent axial and spiral sculpture as in species nym of Propebela. Unnamed Japanese Propebela Marine Mollusca of isotope stages of the last 2 million years in New Zealand 117

species illustrated by Hasegawa (2009, figs. (OIS13; GS4013,R22/f6353). Antiguraleus is 406412) with prominent, sharp peripheral no- not listed in the fauna from Kupe Fm (OIS17), dules closely resemble Propebela propebeloides but a specimen is present in GS4118, R22/ (Long) in all characters. f6438, from the bivalve shellbed member. All Australian species of this group will require specimens (only one in each collection) are reassessment. New Zealand species now as- from the Castlecliff coastal section. They differ signed to Propebela Iredale, 1918 are: from Propebela abnormis (Fig. A, DF, I, J) in Propebela abernethyi (Dell, 1956), Recent. their taller and narrower shape and straighter Propebela abnormis (Hutton, 1885), Mangapa- outlines, lacking the strongly outward-inclined nianNukumaruan (late Plioceneearly Pleis- outer lip and wide sutural ramp of P. abnormis, tocene) (Fig. A, DF, I, J). and are more nearly similar to P. decepta.Itis Propebela decepta (Powell, 1942), Nukumar- clear that although P. abnormis is limited to uanCastlecliffian (Pleistocene) (see below) Nukumaruan rocks, species similar to P. de- (Fig. B, F, I). cepta are not. The common Hawke’s Bay Propebela fenestrata (Powell, 1942), Recent. Nukumaruan species P. abnormis and P. de- Propebela fusiformis (Dell, 1956), Nukumaruan cepta are illustrated here (Figs. A, DF, I, J, (Pleistocene)Recent. B, F, I), although they are shorter and Propebela infanda (Webster, 1906), Recent. more heavily sculptured than most other New Propebela makaraensis (Vella, 1954), Nukumar- Zealand species referred to Propebela. uan (Pleistocene). Propebela multistriata (Dell, 1956), Recent. Propebela ula (Watson, 1881) Propebela munda (Suter, 1909), Recent. Propebela murrhea (Webster, 1906), Recent. Pleurotoma (Drillia) ula Watson 1881:413. Propebela otagoensis (Powell, 1942), Castleclif- Pleurotoma (Bela) ula. Watson 1886:321, pl. fian (middle Pleistocene)Recent. 22, fig. 1. Propebela pedica (Powell, 1942), Recent. Mangilia ula. Suter 1899b:71. Propebela pulcherrima (Powell, 1942), Recent. Bela ula. Suter 1913:485, pl. 21, fig. 12 (with Propebela rishworthi (Vella, 1954), Tongapor- further synonymy). utuan (late Miocene). Asperdaphne ula. Powell 1942:162; Dell Propebela rossiana (Powell, 1942), Nukumar- 1956a:136, pl. 20, fig. 207; Dell 1963a:176, pl. uan (Pleistocene)Recent. 2, fig. 9; Powell 1979:244, fig. 56.5; Spencer et

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Propebela subtruncata (Powell, 1942), Nuku- al. 2009:209. maruan (Pleistocene)Recent. Mioawateria depressispira Beu 1969:494, fig. 7 Propebela taranakiensis (Marwick, 1926), Ton- (new synonym). gaporutuan (late Miocene). Antiguraleus(?) depressispirus. Maxwell Propebela ula (Watson, 1881) (depressipira 1988:68; Beu & Maxwell 1990:420; Maxwell Beu), Nukumaruan (Pleistocene)Recent (see 2009:248. below). Wanganui Basin Castlecliffian records of Type material. Pleurotoma ula, holotype Propebela (as Antiguraleus) by Fleming BMNH 1887.2.9.1004, from HMS‘ Challenger’ (1953:219, 226, 230) and in the GNSreference Sta. 169, off East Cape, New Zealand, in 1280 collection are: Antiguraleus n. sp. aff. abnormis, m (illustrated by Dell, 1963a, fig. 9). Mioawa- Lower Castlecliff Shellbed (OIS 15; GS4104, teria depressispira, holotype in Geology R22/f6518), A. cf. abnormis, Pinnacle Sand Department, Victoria University of Wellington, (OIS14; GS4023,R22/f6361), Antiguraleus n. VM349, from locality V1901, S28/f6884, sea sp., uppermost Pinnacle Sand (GS4098, R22/ cliffs E of mouth of Whangaimoana Stream, f6425), and Antiguraleus n. sp., Tainui Shellbed Palliser Bay, early Nukumaruan. 118 AG Beu

Other material examined. Opoitian: GS1567, (1990:420) recognized that the lack of a raphi- W19/f7516, Waiau River at junction with tomine protoconch and the absence of a Pakihiwi Stream, inland from Wairoa, N prominent moniliform subsutural fold rule out Hawke’s Bay (the locality from where Acirsa a position in Mioawateria for ‘M’. depressispira, cf. cookiana is recorded, above) (one incom- and assigned it to Antiguraleus. The holotype of plete, similar to P. ula but with less sharply P. ula (Dell 1963a, pl. 2, fig. 9) and other, contracted whorls below the sutural ramp). shorter specimens (Dell 1956b:228, fig. 207) Nukumaruan: the only other fossil specimen is seem to differ from A. depressispira only in from the same locality as the holotype of M. being slightly taller, and height is obviously depressipira, in the collection of D Cowe, variable in this species. Greytown (Maxwell 1988:67). Recent: NMNZ M.12832, 548 m, E of Taiaroa Head, E Otago Genus Liracraea Odhner, 1924 (three; labelled ‘compared with type’); NMNZ M.9149, ‘Alert’ stn 5417, 475640 m, ENE of Liracraea Odhner 1924:44. Type species (by Taiaroa Head, E Otago (one). original designation): Clathurella epentroma Murdoch, 1904, Recent, southern New Zealand. Distribution. Propebela ula occurs at present along the E coast of New Zealand, where it is Remarks. The type species of Oenopota, O. recorded only from the E Otago canyons to pyramidale (Stro¨ m, 1788) (Recent, Arctic East Cape, in 4751280 m. Fossils are recorded Ocean), has a loosely coiled, paucispiral proto- only from the Palliser Bay Nukumaruan bath- conch with three or four prominent, angling yal fauna, where they suggest a greater depth of spiral keels (Bogdanov 1990, figs. 397D,E, deposition (500 m) than previously thought. 399A,B) and much more closely resembles the A possible Opoitian relative suggests that this New Zealand genus Liracraea Odhner, 1924 group has occupied bathyal environments off than it does Propebela. New Zealand species New Zealand since early Pliocene time. assigned to Liracraea reach only 79mmin height, whereas the specimens of O. pyramidalis Dimensions. Pleurotoma ula, holotype: H 5.5, D illustrated by Bogdanov (1990, figs. 191208) 3.0 mm; Mioawateria depressispira, holotype: H are 10.317.7 mm high. Several of the species 4.6, D 3.7 mm (Beu 1969:495); NMNZ assigned to Liracraea also are narrower, have a M.12832, off Taiaroa Head: H 5.9, D slightly shallower anal sinus, have straighter

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 3.4 mm; H 5.9, D 3.3 mm; M.9149, off Taiaroa and more prominent axial ridges, and have a Head: H 4.8, D 2.8 mm. shorter anterior siphonal canal than most Oenopota species. However, O. pingelii (Mo¨ ller, Remarks. ‘Asperdaphne’ ula, referred to Asper- 1842) (Arctic Ocean; Bogdanov 1990, figs. daphne (that is, Pleurotomella) by Powell (1942, 209, 210; Kantor & Sysoev 2006, pl. 116F; 1966), seems better referred to Propebela. The Figs. 19A, D, 20AC) resembles Liracraea protoconch agrees with that of Propebela. species more closely in shape, sculpture and ‘Asperdaphne’ ula has a shallow but obviously protoconch sculpture, and Bogdanov’s figured U-shaped sinus with its apex in the centre of the specimens are only 10.410.7 mm high (illu- sutural ramp, and clearly does not have the strated specimen: H 7.5, D 3.0 mm). SEM deep reversed-L-shaped sinus of Pleurotomella. comparison of specimens of Liracraea odhneri The small, short, strongly keeled shell from Powell, 1942 (Fig. 19C, G) and L. subantarctica Palliser Bay named Mioawateria depressipsira Powell, 1942 (Fig. 19E, H, J) with O. pingelii by me (Beu 1969:494) belongs in Propebela and (Figs. 19A, D, 20AC) revealed minor differ- seems likely to be a synonym of Propebela ula. ences. Liracraea species have more prominent, Maxwell (1988:67) and Beu & Maxwell regular spiral cords on the protoconch than in Marine Mollusca of isotope stages of the last 2 million years in New Zealand 119 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 19 (A,D) Oenopota pingelii (Mo¨ ller), MNHN, Arctic Ocean, unlocalised, ex Petit de la Saussaye collection; SEM, whole specimen, height 7.5 mm (see also Fig. 20). (B,F,I) Propebela decepta (Powell), GS10858, U22/f9588, Ashcott Road, W of Waipukurau, Hawke’s Bay, Nukumaruan; B, whole specimen, height 6.5 mm; F, protoconch of B; I, granules on protoconch. (C,H,J) Liracraea subantarctica Powell, RM2835, Carnley Harbour, Auckland Islands; C, height 6.2 mm; H, protoconch of C; J, granules between 2 spiral cords in H. (E,G) Liracraea odhneri Powell, 2 specimens, RM5585, Portobello sta. Mu7045, 540490 m, Papanui Canyon, off Otago Peninsula, RV ‘‘Munida’’; E, height 6.6 mm; G, height 6.4 mm. 120 AG Beu

Figure 20 Oenopota pingelii (Mo¨ ller), MNHN, Arctic Ocean, unlocalised, ex Petit de la Saussaye collection, SEM; specimen in Figs. 19A,D; (A,B) protoconch; (C) granules on protoconch in B.

O. pingelii. Also, the protoconch microsculp- species. Liracraea seems likely to be a synonym ture of minute granules looks similar on the of Oenopota. two at first sight, but the granules of O. pingelii When proposing Liracraea, Odhner are arranged in regular axial lines, whereas (1924:44) did not compare it with Oenopota, those of Liracraea species are uniformly and although he must have been familiar with it. evenly distributed, or have a tendency to form Presumably this implies that he did not con- spiral rows. The protoconch is one whorl taller, sider a close phylogenetic relationship between and so narrower, in O. pingelii than in Lir- New Zealand and Arctic Ocean species to be acraea, and on the last protoconch whorl of O. possible. The apparently bipolar distribution is pingelii the lines of granules develop into still a significant obstacle to the adoption of regular, narrow, relatively widely spaced, gran- Oenopota in New Zealand, but it is possible ulous axial ridges that are not present on that Oenopota species remain to be recognized Liracraea species. Despite these differences, between the Arctic/boreal and New Zealand comparison with the protoconchs of the other species. However, it seems best not to adopt species assigned to Oenopota by Bogdanov Oenopota for New Zealand species until their Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (1990) demonstrated that Liracraea probably position can be checked by comparison of intergrades with Oenopota. Several Oenopota DNA sequences. species have few, prominent spiral cords on a loosely coiled protoconch closely similar to that Clade Heterobranchia of species assigned to Liracraea (Bogdanov 1990, figs. 395GE, O. declivis (Love´ n, 1846); Family Aplustridae fig. 396, O. elegans (Mo¨ ller, 1842); fig. 397A,B, O. (Dall, 1884); fig. 398, O. impressa Genus Hydatina Schumacher, 1817 (Mo¨ rch, 1869); fig. 399A,B, O. pyramidalis). Several of these have the minute surface Hydatina Schumacher 1817:55, 186. Type spe- granules evenly and uniformly arranged, as in cies (by subsequent designation, Gray New Zealand species assigned to Liracraea, and 1847:162): Hydatina filosa Schumacher, 1817 whether there are axial ridges on the last whorl (Bulla physis Linne´ , 1758), Pleistocene & of the protoconch also differs from species to Recent, world-wide tropics. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 121 Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011

Figure 21 (A,B) Bulla quoyii Gray; A, internal mould, GS13743, Y14/f022, farm road cutting at top of Te Piki Member (Haweran, OIS7), Te Piki, near East Cape; H 41.8, D 26.6 mm; B, RM90, Recent, Mission Bay, Auckland; H 34.5, D 22.1 mm. (C,D) Philippia lutea (Lamarck), GS1932 re-collection, U21/f9488, Okauawa Stream bridge, Kereru Road, central Hawke’s Bay, late Nukumaruan; dorsal and apertural views, H 8.3, D 12.3 mm. (E) Philine tepikia Rudman, GS14777, O06/f087, Waiparera church, NW Hokianga Harbour, Northland, Haweran, OIS7; H 14.7, D 10.8 mm. (F,H) Bulla mabillei Locard, lectotype of Bulla australis Fe´ russac, MNHN moll. 20975, ‘‘Port Jackson’’ (unlocalised; France?); H 41.1, D 29.4 mm (MNHN photos). (G) Hydatina physis (Linne´ ), GS14778, O06/f064, W side Te Rewa Point, NW shore Hokianga Harbour, Northland, Haweran (OIS7); H 22.2, D 18.8 mm.

Aplustrum Schumacher 1817:63, 108. Type Hydatoria Iredale 1936:334. Type species (by species (by subsequent designation, Gray original designation, Iredale 1936:338): Bulla 1847:162, as Amplustrum): Bulla amplustre cinctoria Perry, 1811 (Hydatina zonata Linne´ , 1758, Recent, Indo-West Pacific (syno- (Lightfoot, 1886);Bulla physis Linne´ , 1758), nym according to Rudman 1972b:135). Recent, Indo-West Pacific (synonym according 122 AG Beu

to Rudman 1972b:136) (unavailable, no dis- Hydatina physis forma fasciata. Nordsieck & tinction from related taxa; ICZN Article 13.1). Garcia-Talavera 1979:174, pl. 44, fig. 31.

Type material. Investigating the type material Remarks. Rudman (1972b) synonymized Aplus- and synonymy of this world-wide tropical trum Schumacher, 1817 with Hydatina Schu- opisthobranch is beyond the scope of this macher, 1817, using Hydatina as the senior paper. A synonymy up until 1895 was provided by Pilsbry (1895:387), and Rudman (1972b, synonym, a first reviser’s action. Bouchet et al. 2004) revised the world-wide synonymy of the (2005) showed that Aplustridae Gray, 1847 is a genus Hydatina and of H. physis (Linne´ , 1758), senior synonym of Hydatinidae Pilsbry, 1895. although he did not provide a detailed chreso- nymy, and neither he nor any other author has Hydatina physis (Linne´ , 1758) (Fig. 21G) provided information on the type material. Syntype(s) of Bulla physis apparently remain Bulla physis Linne´ 1758:727. in Linne´ ’s collection, housed by the Linnean Bulla vesicaria Lightfoot 1786:136. Society of London, as it is not in Dance’s Bulla velum Gmelin 1791:3433. (1967:2122) list of species not represented. The Bulla fasciata Bruguie` re 1792:380 (senior syno- type material of H. stroemfelti Odhner is in the nym of H. stroemfelti Odhner, 1932; Nordsieck Swedish Natural History Museum, Stockholm & Garcia-Talavera 1979:174); junior secondary (SMNHI 4195; Sandberg & Ware´ n 1993:129), homonym of Bulla fasciata (Mu¨ ller, 1776), and probably no type material exists for any of included in Bulla by Gmelin (1791:3433). the other names in the synonymy. Bulla cinctoria Perry 1811, pl. 40. Hydatina filosa Schumacher 1817:186. Other material examined. Haweran: Akera-Dia- Bulla (Hydatina) straminea Menke 1853:136. la-Anadara locality, Te Rewa Pt, N Hokianga Hydatina physis. Pilsbry 1895:387, pl. 45, figs. Harbour, Northland, OIS7 (GS14778,O06/ 1417 (with further synonymy); Powell f064, one; Fig. 21G). 1924:284; Habe 1950:17, text-fig. 1; Kira 1962:115, pl. 40, fig. 17; Powell 1964:18; Distribution. OIS7, Hokianga Harbour; Re- Kuroda et al. 1971:284, pl. 64, fig. 17; Rudman cent. An uncommon straggler to NE New 1972b:122, figs. 114, pl. 1A,B,H; Powell Zealand harbours during summer months;

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 1976a:159; Powell 1979:270, pl. 54, fig. 6; throughout the tropical Indo-West Pacific and Atlantic. Nordsieck & Garcia-Talavera 1979:174, pl. 44, fig. 30; Kay 1979:420, figs. 131A, 135H; Dimensions. GS14778, Te Rewa Point, Hokian- Bosch et al. 1995:180, fig. 830; Voskuil 1995:32, ga: H 22.2, D 18.8 mm; RM2902, Recent, pl. 1, figs. 14; Spencer & Willan 1996:33; Takou Bay, Northland: H 26.2, D 21.0 mm; Hasegawa et al. in Okutani 2000:737, pl. 367, NMNZ M.95230, Rangiputa Beach, North- fig. 1; Ardovini & Cossignani 2004:243, three land, August 1988: H 33.2, D 24.3 mm; H figs; Spencer et al. 2009:212; Maxwell 2009:251. 38.4, D 31.1 mm; H 43.1, D 34.3 mm; M.95229, Hydatina stroemfelti Odhner 1932:26, pl. 1, figs. as above, after storm, July 1988: H 56.4, D 2123; Duffus & Johnston 1969:36. 47.0 mm; H 54.7, D 46.8 mm. Hydatina vesicaria. Bartsch 1940:192; Abbott 1954:276, pl. 13q; Warmke & Abbott 1961:141, Remarks. Hydatina physis is a large, very thin- pl. 27a; Rehder 1967:23; Abbott 1974:314, pl. shelledbullomorphopisthobranchfound‘world- 10, fig. 3938; Voskuil 1995:33, pl. 1, figs. 5, 6. wide ... in warm seas’ (Rudman 1972b; Powell Marine Mollusca of isotope stages of the last 2 million years in New Zealand 123

1979:270; although there appear to be no eastern H. vesicaria). The exterior characters and Pacific records). Large adult shells reach at least internal anatomy are indistinguishable, and the 57 mm high and 47 mm wide (Powell 1979:270; patternofspiralcolourbandsontheshellishighly andabove).Living specimenshavebeencollected variable.ThesmallerspeciesH.amplustre(Linne´ , in northeastern New Zealand sporadically over 1758) differs in its slightly thicker, stronger shell many years. They were reported first by Powell and fewer colour bands, which include two wide (1924) and have since been collected in Parengar- pink bands outlined in black on most specimens enga Harbour, in the far north of the North (Rudman 1972b), but again the animal charac- Island, in some numbers (Powell 1964, 1976a, ters are very similar. The more widely black- 1979). Specimens have also been found further banded form from the Marquesas Islands named south, at Houhoura Harbour, Whangaroa H. exquisita by Voskuil (1995: 37, pl. 1, figs. 13, Harbour, Takou Bay and Rangiputa Beach, to 14) is probably a colour form of H. amplustre. as far south as Tryphena and Port Fitzroy, Great Hydatina species have an annual life-cycle Barrier Island, east of Auckland. The shell and a planktotrophic larval stage, like Akera is ovate, simple and virtually smooth, with a soluta (Gmelin, 1791) (Burn & Thompson 1998; simple, pear-shaped aperture and wide, evenly Willan 1998), which also occurs abundantly in rounded anterior end, no umbilicus, a thin outer the outcrop between Te Rewa Point and lip, and a slightly sunken, narrow spire with the Waiparera (Beu 2004). They are able to meta- posterior surface of the spire whorls revealed. It is morphose and settle in an area during favour- easily distinguished from Bulla species by the able conditions, breed, and then not necessarily much thinner and more fragile shell, by the be seen again in that area for some years. shorter and wider shape with a wider aperture, Presumably Hydatina physis arrived and and by the earlier whorls revealed in the thrived at several times in sheltered northern spire apex, with the protoconch visible, rather North Island harbours during warm intergla- than the deep, narrow, apical tube of Bulla. cial periods, as it still does today. Powell (1964, Recent specimens have a striking colour pattern 1976a, 1979) suggested that the New Zealand of narrow, dark brown spiral lines on a whitish occurrences are pseudopopulations recruited background, although Rudman (2004) empha- from the tropical southwest Pacific, but not sized the variability of the colour pattern. breeding in New Zealand. However, the large Miguel Griffin (Museu de La Plata, Argenti- numbers of large shells reported in New na) and I collected a reasonably complete fossil Zealand on some occasions suggest that tem-

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 specimen of Hydatina physis at Te Rewa Point, porary breeding populations are likely in New northern Hokianga Harbour (OIS7). The shape Zealand at present, although the two possibi- is within the range of variation of Recent speci- lities would be indistinguishable in the fossil mens. The specimen bears several very low, record (25 living specimens at Takou Bay, indistinct, widely spaced, spiral angulations. Northland; Powell [1964, 1976a]; 17 from Rudman (2004; W Rudman AMSpers. comm. Rangiputa Beach in NMNZ M.95230). 16 November 2007) reported that H. zonata (Lightfoot, 1786) (Powell 1979:270, pl. 54, fig. 7, as the synonym H. albocincta (van der Hoeven, Family Architectonicidae 1839); Spencer et al. 2009:212) (alsoH. velum (Gmelin, 1791) and H. cinctoria (Perry, 1811)), Genus Philippia Gray, 1847 with fewer, wider spiral colour bands, is probably merely a colour form of H. physis (see also Philippia Gray 1847:146. Type species (by illustrations in Voskuil 1995:34, pl. 1, figs. 110; monotypy): Solarium luteum Lamarck, 1822, figs. 7, 8 identified as H. albocincta, figs. 9, 10 Recent, Indo-West Pacific (mainly SAustralia identified as H. zonata, figs. 5, 6 identified as and N New Zealand; Bieler 1993:113, fig. 89). 124 AG Beu

Philippia lutea (Lamarck, 1822) (Fig. 21C, D) Type material. Solarium luteum, lectotype (de- signated by Bieler 1985:236; Bieler 1993, fig. 86) Solarium luteum Lamarck 1822b:5; Deshayes and five paralectotypes MHNG 1095/41, ‘Ha- 1830 (in 18301832):159; Kiener 18381839:9, bite les mers de la Nouvelle Hollande’ (Aus- pl. 4, fig. 9b (in part; not fig. 9a,Philippia tralia); not seen. Iredale & McMichael (1962:3, hybrida (Linne´ , 1758); Bieler 1993:110); Deles- 68) designated the type locality as ‘Sydney, N. sert 1841, pl. 24, fig. 2ac; Deshayes 1843:100; S. W.’. Bieler (1993) did not know the location Philippi 1853:9, 31, pl. 1, figs. 10, 11 (in part; of any type material of S. luteum var. novae- not pl. 4, fig. 11,P. hybrida; Bieler 1993:110); hollandiae Philippi, from ‘Neuholland?’ (Aus- Reeve 1864, pl. 3, fig. 14; Suter in Hutton tralia); it is likely to be in Philippi’s collection in Museu Nacional de Historia Natural, Santiago, 1904:79; Bieler & Petit 2005:48. Chile (not accessible for consultation). Philippia lutea. Gray 1847:146; H Adams & A Adams 1858:243, pl. 25, figs. 88b; Angas Other material examined. Nukumaruan: Man- 1867:201; Hutton 1878:24; Hutton 1880:70; gatahi River, on NW side of Okauawa Stream Finlay 1926b:401; Thiele 1929:184, fig. 170; junction, Kereru Road, SW of Maraekakaho, Cotton & Godfrey 1933:72, pl. 1, fig. 1; Powell central Hawke’s Bay (GS1932 re-collection, 1937:75, pl. 9, fig. 35; Wenz 1939:670, fig. 1911; U21/f9488; one; Fig. 21C, D); Devils Elbow Powell 1946a:77, pl. 9, fig. 35; Powell 1958a:97, Mudstone, cutting at top of Devils Elbow hill, pl. 9, fig. 35; Iredale & McMichael 1962:68; Napier-Wairoa Highway, N of Napier Macpherson & Gabriel 1962:100, fig. 126; (GS10849, V20/f8572; one). Recent: rather un- Powell 1962:89, pl. 9, fig. 35; Beu 1970a:128; common throughout northern New Zealand Powell 1976b:107, pl. 16, fig. 35; Beu in Suggate (44 lots examined in NMNZ and eight in et al. 1978:724, fig. 11.13.6; Powell 1979:248, pl. GNS), from the Three Kings Islands (north- 48, fig. 2; Beu & Maxwell 1990:422; Bieler ernmost: NMNZ, M.117094, Send of North 1993:110, figs. 8689; Spencer & Willan East Island, 3488.30?S, 172810?E, one specimen; 1996:31; Spencer et al. 2009:211; Maxwell M.93986, reef between Great Island and Farm- 2009:249. er Rocks, 3489.00?S, 172810?E; three) to as far Solarium luteum var. novaehollandiae Philippi south as Mount Maunganui, Bay of Plenty 1853:41. (GNSRM276, two) and Boulder Bay, Motu- Solarium (Philippia) luteum. Chenu 1859:233, hora Island, Bay of Plenty (NMNZ, M.33021,

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 fig. 1355; Hanley 1863:237, pl. 253, figs. 53, 54 M.33071, 37851.70?S, 176857.60?E; two) and (in part; not fig. 52,P. hybrida; Bieler East Cape (NMNZ, M.117625; three); NIWA 1993:111); Marshall 1887:16, pl. 5, fig. 71, 72; stn B619, 44842?S, 167833.25?E, off Four-Mile Cooper 1899:137. Point, N of entrance to Sutherland Sound, Fiordland, 19 October 1962 (seven live-col- Architectonica (Philippia) lutea. Suter lected); NMNZ M.160306, 44855.50?S, 1913:316, pl. 46 (1915), fig. 4. 167823.50?E, middle of George Sound, Fiord- Architectonica lutea. May 1921:102; May land, 37 m, 14 February 1987 (one fragment). 1923:97, pl. 46, fig. 1; Bucknill, 1924:56, pl. 7, fig. 20. Distribution. Philippia lutea occurs moderately Philippia (Philippia) lutea. Robertson 1973:37; commonly around southern Australia (Western Garrard 1977:509, 525, fig. 14 (operculum), pl. Australia, N to Vansittart Bay, 14803?S; around 5, figs. 1318; Boss & Merrill 1984:359, pl. 57, southern Australia and Tasmania, to Wool- fig. 1; Bieler 1985:236, pl. 2, fig. 8 (lectotype). goolga, northern New South Wales, 30807?S; Philippia hybrida. Hinton 1978, pl. 10, fig. 10 Garrard 1977:526) and in the NE North Island (not Trochus hybridus Linne´ , 1758). warm province of New Zealand, as far south as Marine Mollusca of isotope stages of the last 2 million years in New Zealand 125

the E Bay of Plenty. Bieler (1993:110114) 0.760.90) and very pale coloration. Specimens discussed the characters, variation, and distri- from northern New Zealand are pale yellow bution of P. lutea, and thought records from ochre to reddish tan in colour, apart from the the Philippines and Howland Island require prominent, bright, short, closely spaced, alter- confirmation. P. japonica (Pilsbry & Stearns in nating yellow and dark brown dashes around Pilsbry, 1895) (Bieler 1993:114, fig. 90) is the two peripheral spiral cords. The Fiordland extremely similar, differing only in having a specimens also all contain the dried animal, and slightly smaller protoconch (averaging 0.81 mm apparently have been preserved in ethanol for in diameter, rather than 0.94 mm in P. lutea), some time, so the shells possibly have faded. and it is possible that P. lutea is a poorly They also were collected from an unusually understood, wide-ranging, western Pacific spe- great depth for the species (92 m), as Garrard cies occurring from Japan to New Zealand, (1977:526) recorded the greatest depth as 82 m. although rare between the latitudinal extremes, They have 3545 narrow colour bands around and varying clinally in protoconch size. Such a the peripheral cords of the last whorl, the main distribution would be similar to that of the character stated by Bieler (1993:113) to distin- planktotrophic tonnoideans that occur in both guish P. lutea (with 2352 colour bands) from Japan and AustraliaNew Zealand, but are the AtlanticMediterranean species P. hybrida rare in the tropics in between ( lampas (with 1926 colour bands; Melone & Taviani (Linne´ , 1758); parthenopeus (von 1984:166, fig. 25; Ardovini & Cossignani Salis Marschlins, 1793); Monoplex exaratus 2004:232, lower right two figs). Garrard (Reeve, 1844); Beu 2010). (1977:526) described New South Wales speci- mens as ‘invariably elevated conic and coloured Dimensions. NIWA stn B619, off entrance to yellow ochre or light buff, a few specimens light Sutherland Sound, Fiordland: H 15.0, D 18.0 grey’, with H:D 0.70.8. In contrast, Victorian mm (H:D 0.83); H 15.2, D 16.8 mm (H:D 0.90); specimens are predominantly pale to dark grey, H 13.3, D 15.8 mm (H:D 0.85); H 12.0, D 15.8 with H:D 0.60.7, South Australian specimens mm (H:D 0.76); H 10.9, D 13.6 mm (H:D 0.80); have a reddish tinge and a more depressed spire, H 11.9, D 13.4 mm (H:D 0.89); H 9.9, D 13.1 mm and Western Australian specimens are still more (H:D 0.76); GNSWM15278, Collaroy Beach, depressed (H:D 0.450.55) and are predomi- Narrabeen, Sydney, NSW: H 10.5, D 12.5 mm nantly pale to deep blackish red. Recent speci- (H:D 0.84); H 7.6, D 10.8 mm (H:D 0.70); GNS mens from northern New Zealand are

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 RM3907, Ocean Beach, Whangarei Heads, intermediate in spire height (H:D 0.710.82), Northland: H 8.7, D 12.3 mm (H:D 0.71); whereas the two known New Zealand fossils RM5122, Otahei Bay, Bay of Islands: H 10.3, have rather low spires (H:D 0.670.75). In view D 12.5 mm (H:D 0.82); RM5350, north head, of the range of variation in Australian speci- Parengarenga Harbour: H 9.0, D 12.1 mm (H:D mens described by Garrard (1977), it is likely 0.74); GS1932, Kereru Road: H 8.3, D 12.3 mm that the Fiordland specimens were recruited as (H:D 0.67); GS10846, Devils Elbow: H 9.3, D planktotrophic larvae from a New South Wales (incomplete) 13.4 mm (H:D 0.75). population. The specimen recorded by me (Beu Remarks. A sample of seven specimens of 1970a:128; Beu in Suggate et al. 1978:724, Philippia lutea in NIWA stn B619, off Suther- figs. 11.13.6; Fig. C, D) as Philippia lutea land Sound, Fiordland, is unusual for the large from late Pliocene (late Nukumaruan) rocks size. Garrard (1977:525) recorded ‘fully grown’ of central Hawke’s Bay has been compared Australian specimens reaching H 10, D 15 mm, again with Recent specimens of P. lutea, and whereas the Fiordland specimens reach H 15.2, with the one other incomplete but slightly D 18.0 mm. They also have tall spires (H:D larger fossil specimen collected since, from 126 AG Beu

Devils Elbow Mudstone at the top of Devils species included; no type species was designated Elbow hill. These specimens have a rather by Gray [1847], Hermannsen [18461852], depressed spire for P. lutea and are weakly Pilsbry [1895:327] or Malaquias & Reid sculptured, with almost completely smooth [2008], and I am not aware of any other upper whorl surfaces, but display the two designation). almost equally prominent, almost smooth per- ipheral spiral cords characteristic of Philippia. Vesica Swainson 1840:360. Type species (by They can be matched among Recent Northland subsequent designation, Malaquias & Reid specimens of Philippia lutea. Their occurrence 2008:457): Bulla Linne´ , 1758. presumably resulted from the transport of planktotrophic larvae southwards from Quibulla Iredale 1929:349. Type species (by the present range in Northland during warm original designation): Bullaria botanica Hedley, Nukumaruan interglacial periods. Although 1918 (Bulla quoyii Gray, 1843), Pleistocene the geographical subdivision of colour and and Recent, southern and eastern Australia and spire height around Australia indicates some northern New Zealand. restriction of genetic exchange, Architectonici- dae are among the most wide-ranging of Bulla quoyii Gray, 1843 (Fig. 21A, B) gastropods, well-known for having long plank- totrophic larval lives allowing distributions ?Bulla nebulosa Schro¨ ter 1804:20 (nomen du- similar to those of the most wide-ranging bium; Pilsbry 1895:350; Malaquias & Reid tonnoideans (Scheltema 1968, 1971, 1979; 2008:524). Scheltema & Williams 1983; Scheltema et al. Bulla australis Gray 1825:408; Gray 1827:490; 1996). Their larvae easily could be transported Gray 1835:308; Gray 1843:243; Brazier from Australia and from northern New Zeal- 1885:89; Whitelegge 1889:275; Pilsbry and to central and southern New Zealand. 1895:346, pl. 36, figs. 17, 18; Pritchard & Gatliff 1903:214; Iredale 1929:349, pl. 33, fig. 4 (junior Family Bullidae primary homonym of Bulla australis Fe´ russac, 1822). Genus Bulla Linne´ , 1758 Bulla striata. Quoy & Gaimard 1833:354, pl. 26, figs. 8, 9 (incorrect usage of Bulla striata Bruguie` re, 1792).

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Bulla Linne´ 1758:725 (placed on the Official List of Generic Names in Zoology in ICZN Bulla australis Quoy & Gaimard 1833:357, pl. Opinion 196 1954). Type species (ICZN Opi- 26, figs. 38, 39; Deshayes 1836:673; H Adams & nion 196): Bulla ampulla Linne´ , 1758, Pleisto- A Adams 1854:16; Chenu 1859:389, fig. 2938; cene and Recent, Indo-West Pacific. Martens 1873:38; Watson 1886:638 (junior primary homonym of B. australis Fe´ russac, Bullus Montfort 1810:330. Type species (by 1822 and of B. australis Gray, 1825). monotypy): Bullus ampula (sic) ‘de Lamarck’ Bulla quoyii Gray 1843:243; EA Smith 1874:5, (Bulla ampulla Linne´ , 1758). pl. 1, fig. 11; Pilsbry 1895:348, pl. 39, fig. 71; Willan 1977:1117, fig. 1, pl. 1aj, pl. 2ad, pl. Bullaria Rafinesque 1815:142. Replacement 3ac; Willan 1978:58, figs. 1524; Wells name for Bulla Linne´ , 1758. 1985:30, pl. 1, figs. 17; Spencer & Willan 1996:33; Richardson 1997:11; Burn & Thomson Bullea Blainville 1825:477. Type species (desig- 1998:957, fig. 16.36A,B; Morley 2004:120, fig; nated here): Bulla ampulla Linne´ , 1758 (three Malaquias & Reid 2008:524, figs. 25C, 27DK, Marine Mollusca of isotope stages of the last 2 million years in New Zealand 127

28F, 29E, 31F, 32E, 34A,B, 36D,E, 37H,I, 39; Type material. Willan (1978:59) reviewed the Spencer et al. 2009:213. type material of most of the names in the above Bulla oblonga A Adams 1850:577, pl. 123, fig. synonymy. An expanded synonymy was pro- 74; H Adams & A Adams 1854:16; Angas vided by Malaquias & Reid (2008:524); these 1867:226; GB Sowerby II 1868, pl. 3, fig. are summarized here. As noted by Beu (2010: 4ac; Hutton 1873a:52; Tenison Woods 6263), EA Smith (1874:5, pl. 1, fig. 11) 1878:47; Hutton 1880:121; Pilsbry 1895:346, illustrated the ‘type’ (the largest of 10 syntypes, pl. 35, figs. 1214. BMNH 1842.11.18.135142; Willan 1978:59; Bulla castanea A Adams 1850:584, pl. 124, fig. lectotype selected by Malaquias & Reid 106a. [2008:524, fig. 27G]largest, H 22.5 mm; Bulla (Bullea) substriata Menke 1853:136 (pro- repeated by Beu [2010, fig. 1B]) of Bulla quoyii, posed partly as replacement name for Bulla in BMNH, collected in New Zealand by striata of Quoy & Gaimard, 1833, not of Frederick Strange (almost certainly from the Bruguie` re). Bay of Islands, N North Island). Bulla oblonga, Bulla tenuissima GB Sowerby II 1868, pl. 2, fig. three syntypes BMNH 197647 (one figured, 4; Pilsbry 1895:347, pl. 39, fig. 72 (synonym Malaquias & Reid 2008, fig. 27D), ‘Philippines according to Wells 1985:30, pl. 1, figs. 17). and Annaa I.’, Cuming collection (incorrect; Bulla quoyi. Martens 1873:38; Hutton 1873a:52; probably from Port Jackson, New South Hutton 1880:121; Pilsbry 1895:348, pl. 39, fig. Wales). Bulla (not Haminea) castanea, probable 71; Powell 1979:275, pl. 52, fig. 2; Beu in holotype BMNH 1881.5.20.22 (Malaquias & Yoshikawa et al. 1980:251; Beu & Maxwell Reid 2008, fig. 27F), from the Lombe-Taylor 1990:422; Maxwell 2009:251. collection, from ‘shores of New Zealand’. Bulla Bulla australis var. oblonga. Pilsbry 1895:346, australis Gray, three syntypes BMNH pl. 35, figs. 1214. 20060585, from ‘Australia’ (not seen; Mala- Haminea castanea. Pilsbry 1895:374, pl. 41, fig. 14 quias & Reid 2008:524). Bulla australis Quoy & Bulla dubiosa Mabille 1896:113 (no locality or Gaimard, 11 syntypes examined in MNHN types; synonym according to Malaquias & Reid (one illustrated, Valde´ s&He´ ros 1998:708, fig. 2008:524). 3I), from King George Sound (‘port de Roy Bullaria australis. Suter 1913:534, pl. 49, fig. 6. Georges’), Albany, southern Western Austra- Bullaria australis quoyi. Suter 1913:535. lia. When describing Bulla substriata, Menke Bullaria australis. Hedley 1916:72. (1853b:136) mentioned B. striata of Quoy &

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 Bullaria quoyi. Hedley 1916:72. Gaimard (1833), although he did not expressly Bullaria botanica Hedley 1918:M104 (replace- propose it as a replacement name. The type, ‘ad ment name for Bulla australis Gray, 1825, Novam Hollandiam’, was collected by Preiss, preoccupied); May 1921:103; Macpherson so came from southern . Part 1958:50, pl. 46, fig. 14; Macpherson & Gabriel of Menke’s collection was acquired some years 1962:242, fig. 281. ago by SMF in the Bronn collection, but R Quibulla botanica. Iredale 1929:349, pl. 38, fig. Janssen (SMF pers. comm. November 2007) 4; Iredale & McMichael 1962:88. reported that most of Menke’s collection was Quibulla quoyi. Dell 1955:31, fig. 61; Morton & dispersed before Bronn acquired it, and no type Miller 1968:534, fig. 202.5; Rudman 1971:657, material of Bulla substriata is present in SMF. figs. 1D,E, 812; Penniket & Moon 1970:68, pl. Bulla tenuissima, one syntype BMNH 196750 31, fig. 2. (Wells 1985, pl. 1, fig. 7; Malaquias & Reid Bulla (Quibulla) quoyi. Powell 1965:167, pl. 22, 2008, fig. 27E; not seen), from Swan River, fig. 10. Western Australia. Bulla botanica. Burn 1966:266; Coleman Willan (1978) included ‘Bulla striata Quoy & 1975:14, fig. 9. Gaimard, 1833:354, pl. 26, fig. 8, 9...(non Bulla 128 AG Beu

striata Bruguie` re, 1792)’ in the synonymy of & Reid 2008, fig. 40). The spiral grooves on the Bulla quoyii. However, Quoy & Gaimard base of the shell provide a character shared only (1833:354) used a name they attributed to with the Atlantic species B. striata Bruguie` re, Lamarck for specimens from ‘Baie des Iles, a` 1792 and B. occidentalis A. Adams, 1850 (Mala- la Nouvelle-Ze´ lande’. Lamarck (1822a:33) at- quias & Reid 2008:526). Therefore, B. quoyii is tributed the name Bulla striata in turn to apparently another of the Atlantic taxa that Bruguie` re, and this is merely a misidentification. arrived in Australia and New Zealand via South There is no separate type material in MNHN Africa during Pleistocene time as planktotrophic for Bulla striata of Quoy & Gaimard’s usage. larvae transported in the Antarctic Circumpolar Current. It joins a small group of taxa including Other material examined. Haweran: Te Piki, Mytilus galloprovincialis Lamarck, 1819, Mod- road cut 6 km E of Whangaparaoa, near East iolula phaseolina (Philippi, 1844) (Beu 2004), Cape, OIS7 (GS13743,Y14/f022, cutting Pecten (Beu 2006), Lutraria (Lutraria) grandis towards top of farm road at west end of (Hutton, 1873) (Beu 2006), and the tonnoideans outcrop, collected by Y. Ota; one specimen, olearium, , Monoplex limonitized internal mould; Fig. 21A). parthenopeus (von Salis Marschlins, 1793), pos- sibly other Monoplex species such as M. exaratus Distribution. The one fossil record I am aware of (Reeve, 1844), and Semicassis labiata (see in New Zealand is the specimen recorded here above). Other tonnoideans have been trans- from the Te Piki member, inland from Whan- ported in the same way from the North Pacific gaparaoa, near East Cape (OIS7). Recent via South America (Argobuccinum, ). specimens occur from Cape Maria van Diemen More such migrants probably await recognition. and Spirits Bay south to Cape Runaway, eastern Bay of Plenty, with one record from Nelson in Dimensions. Larger syntypes of Bulla australis the northern South Island (Willan 1978:59). In Quoy & Gaimard: H 58.7, D 35.3 mm; H 57.8, Australia Bulla quoyii occurs from central D 33.5 mm; H 52.6, D 32.2 mm; GS13743, farm Western Australia (Houtmans Abrolhos Is- road above Te Piki outcrop: H 41.8, D 26.6 lands) to South Australia, Victoria, Tasmania mm; RM2164, Rangaungu Harbour, North- and northern New South Wales (Malaquias & land: H 48.2, D 29.4 mm; NMNZ largest Reid 2008:526, fig. 39), but seems not to have specimens, M.94074, shell bank, Kauanga been reported as a fossil. Malaquias & Reid Channel, Parengarenga Harbour: H 65.6, D

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 (2008:526) mentioned that Australian specimens 42.7 mm; H 61.0, D 38.0 mm. reach a larger size (H 62.2 mm) than New Zealand ones (H 45.0 mm) and, while this might Remarks. Bulla quoyii is a tall, narrow species be so on average, the presence of specimens in with a weakly ‘waisted’ outline formed by a Parengarenga Harbour, Northland, up to 65.6 weakly concave outer lip, and with a few faint mm high (see below) shows that there is no spiral grooves around the base. Malaquias & consistent difference in dimensions. Reid (2008:426) stated that it is the largest The close phylogenetic relationship of Bulla species in the genus. It has been recorded fossil quoyii with E PacificAtlantic species of Bulla, previously only in the ‘Otamaroa Terrace’ and a corresponding lack of relationships with faunal list (Beu in Yoshikawa et al. 1980:251) W Pacific species, has been pointed out by M and in the Te Piki faunal list of Richardson Malaquias (BMNH pers. comm. October 2007) (1997, 1999). The single fossil specimen (an based on molecular phylogeny. B. quoyii falls internal mould) I am aware of, the basis of all within a clade that includes B. gouldiana Pilsbry, three previous records, is illustrated here (Fig. 1895 (E Pacific), B. punctulata A. Adams, 1850 21A). It was collected by Professor Yoko Ota in (E Pacific), and B. mabillei (Atlantic) (Malaquias 1978 as part of her work on the uplifted Marine Mollusca of isotope stages of the last 2 million years in New Zealand 129

terraces of the northeastern Bay of Plenty (1822:573) stated that his type material of Bulla (Yoshikawa et al. 1980). As it was collected australis came from ‘Port Jackson’, New South from a relatively concentrated, deeply weath- Wales, Australia, but the species was not ered fossiliferous horizon apparently overlying illustrated and has remained unknown. It was the Te Piki Member on the farm access track at defined only by comparison with the preceding the SW end of the Te Piki outcrop, Yoshikawa and succeeding species: ‘Un peu plus petite que et al. (1980) assumed that this fauna repre- la pre´ ce´ dente [a little smaller than the preceding sented the marine cover beds of the Otamaroa species: B. pisum Fe´ russac, no locality], se Terrace (last interglacial terrace, OIS5e). Also, rapprochant de la suivante par son forme’ the inclusion of such estuarine species as [approaching the following in form: B. or- Macomona liliana (Iredale, 1915) and Bulla bignyana Fe´ russac, from near La Rochelle, quoyii that are otherwise rare or unknown in Atlantic coast of France; that is, presumably Te Piki Member made assignment to Te Piki a specimen of either Bulla striata or B. mabillei]. Member seem unlikely. However, I have since The remaining syntype of Bulla australis Fe´ r- re-examined this site. The fauna includes abun- ussac, MNHN 20975, illustrated here (Fig. 21F, dant specimens of Pecten novaezelandiae (Re- H) and by Valde´ s&He´ ros (1998:708, fig. 3H; eve, 1852) of the tainui phenotype (Beu 2006) Fig. 41G,H; H 41.1, D 29.4 mm) resembles B. and at least one specimen of Eunaticina papilla. vernicosa Gould, 1859 (Malaquias & Reid They occur with several other species that are 2008:527;B. angasi [not of Pilsbry, 1895; a common throughout the Te Piki Member, such synonym of B. mabillei; Malaquias & Reid as Dosinia (Fallartemis) lambata, D. (Kereia) 2008:492], the name used by Willan [1978:60, greyi, Dosina zelandica and , figs. 1, 2, 2537] for the second Bulla species in but are otherwise unknown (D. greyi)or northern New Zealand and central eastern unusual constituents of uplifted terrace faunas. Australia to New Caledonia and Samoa). It is The leached, limonite-cemented nature of the definitely not a specimen of B. quoyii. Because material merely results from its position at the of this similarity and the locality ‘Port Jackson’, top of the Te Piki exposure. There is no doubt I investigated this name in case it is an earlier that the fauna reported by Yoshikawa et al. name for B. vernicosa. The remaining syntype (1980:251) merely is a shell concentration at the differs from B. vernicosa in having a more top of Te Piki Member. The fauna suggests that evenly convex outer lip, not weakly concave in the member shallowed and became more es- the centre as in B. vernicosa, and the colour

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 tuarine in nature towards the top, as would be pattern is a more reddish-purple, diffused, expected of the deposits of a single interglacial cloudy one with relatively large, variably period in a small enclosed bay. No fossiliferous blended, circular spots, without the many finer marine deposits are known associated with the dots that make up the pattern of B. vernicosa. Otamaroa Terrace, and the specimen of B. The syntype is accompanied by a hand-written quoyii is assigned to OIS7 rather than OIS label stating ‘specimenBulla mabillei Locard 5e. Bulla quoyii is a characteristic member of [1897], Atlantic. Robert Burn, January 1992’. the northern North Island warm-water fauna, Comparison of specimens and illustrations and the fossil locality lies within its living range. (Locard 1897:50, pl. 11, figs. 1, 2; Talisman stn 107, 70 m, St Vincent, Cape Verde Islands; a Status of Bulla australis Fe´rrusac. Willan (1978) large specimen, H 60, D 42 mm; Poppe & Goto did not resolve the status of Bulla australis 1991:195, pl. 37, figs. 23, 24; Ardovini & Fe´ russac, 1822, the senior homonym of B. Cossignani 2004:242, central left two figs) australis Gray, 1825. Malaquias & Reid showed that Burn’s identification is correct. (2008:524) treated it as an earlier name for This was confirmed by Malaquias & Reid Bulla quoyii, but a nomen dubium. Fe´ russac (2008:524). B. australis Fe´ russac is identifiable 130 AG Beu

only by its remaining syntype, which appar- following names are concluded to be synonyms ently (like many type specimens of other of Philine: Lobaria Mu¨ ller (1776:28, 260); molluscs) bears an incorrect locality. Because Bullaea Lamarck (1801:63); Laona A. Adams of the confusion over the possible type locality, (1865:324); Utriculopsis M. Sars (1870:177); and the consequent identity of the species, the Hermania, Johania and Ossiania of Montero- remaining syntype of B. australis Fe´ russac, sato (1884:147); Yokoyamaia, Choshiphiline and MNHN 20975 (Fig. 21F,H) is here designated Philinorbis of Habe (1950:50, 52); Globiphiline the lectotype of Bulla australis Fe´ russac, 1822, Habe (1958:120); Rhinodiaphana Lemche which is, therefore, an earlier name for the (1967:208); Retusophiline and Philingwynea of Atlantic species B. mabillei Locard, 1897. As Nordsieck (1972:20, 22); and Pseudophiline the name Bulla australis Fe´ russac has not been Habe (1976:154). used as the valid name for a species after 1899, and B. mabillei is a well-established name, it is Philine tepikia Rudman, 1970 (Fig. 21E) likely that B. australis qualifies as a nomen oblitum. However, Malaquias & Reid Philine tepikia Rudman 1970:31, fig. 2M, pl. (2008:492) pointed out that both B. cruentata 3C; Maxwell 2009:251. A. Adams, 1850 and B. angasi Pilsbry, 1895 Philine tepikiensis (sic) Beu & Maxwell also are earlier names for B. mabillei. They 1990:422. retained the name B. mabillei, pending an application to the ICZN, so B. australis Fe´ russac should be included in the same Type material. Philine tepikia, holotype application. TM4855, from GS4003, R22/f7394, Landguard Sand, Landguard Bluff, east of Wanganui City (Haweran, OIS9); one paratype AUGD Family Philinidae G5868, from Te Piki member, road cut inland from Whangaparaoa, near East Cape (Hawer- Genus Philine Ascanius, 1772 an, OIS7).

Philine Ascanius 1772:331. Type species (by Other material examined. Haweran: the only monotypy): Philine quadripartita Ascanius, specimen of Philine tepikia I am aware of other 1772 (Bulla aperta Linne´ , 1767), Recent, than the type material is from Hokianga

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 North Atlantic & Mediterranean. Harbour (GS14777, O06/f67, bank at back of tidal flats in front of Waiparera Church; Haweran, OIS7). Remarks. The first published synonymy for Philine that I am aware of is that by Valde´ s Distribution. Known only by the three speci- (2008:713714), who, however, did not include mens recorded here, from Landguard Bluff, Te some of the names concluded here to be Piki, and Waiparera (OIS9 7). The shell is synonyms. Pilsbry (1895) provided no synony- my, referring to Fischer’s (1883, in extremely thin and fragile, and fossils are 18801887:563) treatment. As for Hydatina, difficult to collect, so its range is likely to be the details of the synonymy are beyond the greater than these specimens suggest, and its scope of this paper, as I am unable to determine biostratigraphical utility is minimal. the type species of some genera, but following Fischer (1883), Rudman (1970; 1972a: Dimensions. Philine tepikia, holotype: H 29.5, 185186), Kitao & Habe (1982), Burn & D 22 mm (Rudman 1970:31); GS14777, Thomson (1998:951) and Valde´ s (2008), the Waiparera: H 14.6 (incomplete), D 10.8 mm. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 131

Remarks. Philine has a very thin, fragile, The species now recorded from Landguard translucent shell with a very large, simple Sand, only, at Wanganui and otherwise known aperture, widely open anteriorly, with the fossil or living only in the northern small, ovate spire whorls almost completely New Zealand warm-water province are enveloped by the last whorl, and with a flat or Capulus danieli, Stiracolpus vigilax (ahipa- slightly sunken spire apex. Many, but not all, ranus?;pagoda?), Eunaticina papilla, Pupa species have spiral sculpture of weak spiral affinis and Philine tepikia. The warm-water, grooves, punctate on some species, and some northeastern North Island species Zelippistes species have spines or other protrusions from benhami also is recorded from Landguard the upper (adapical) part of the outer lip. Fred Sand, and otherwise apparently is recorded at Brook collected a single specimen of the Wanganui only from Tainui Shellbed (OIS 13). relatively large species P. tepikia from Other northeastern North Island and/or tropi- GS14777, NW Hokianga Harbour (OIS 7). cal Pacific species occurring in Landguard The holotype (Rudman 1970:31, fig. 2M, pl. Sand, with longer time ranges, include Amyg- 3, fig. C) is an almost complete specimen 29.5 dalum striatum (Hutton, 1873), Limaria orien- mm high, collected from Landguard Sand (OIS talis (A. Adams & Reeve, 1850) and Leucotina 9) at Landguard Bluff by CA Fleming. Rud- casta (A. Adams, 1853). man (1970) also recorded a specimen from Te Piki, Cape Runaway (OIS7). The Waiparera specimen is 14.3 mm high, and agrees with the Acknowledgements holotype of P. tepikia in having unusually Funded by the New Zealand Foundation for Re- prominent dorsal growth ridges, and 810 quite search, Science and Technology through Backbone prominent, narrow, closely spaced, punctate Fund Contract CO5XO412 (National Paleontology spiral ridges in the apical depression. The Collection). Dedicated to the late Dick Dell and Waiparera specimen has only 34 faint, shallow Charles Fleming, who inspired my life-long interest spiral grooves across the adapical part of the in malacology and paleontology, and to the late Phil Maxwell, my friend and colleague for 40 years. exterior below the apex, rather than the numer- Many of the new records from Wanganui Basin ous, slightly more closely spaced ones across were collected and recognised by Charles Fleming, the adapical two-thirds of the shell on the who charged me with recording them. Bruce Mar- holotype. However, this difference seems likely shall (NMNZ) provided extensive, critically impor- to be part of the variation of one species; the tant data through his gathering and curation of the

Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 number of grooves presumably increases as the huge NMNZ collection, as well as many suggestions shell grows, and the holotype is twice the size of on taxonomy and biogeography over many years, the Waiparera specimen. Therefore, it appears and provided valuable comments on the manuscript. that a relatively large Philine species was Many New Zealand and Australian ‘‘Wanganui endemic to New Zealand during OIS9 7, and working group’’ colleagues have contributed locality that the similarly large species P. angasi (Crosse data and stratigraphy over the years: Bob Carter & Fischer 1865:38, pl. 2, fig. 8), which differs (James Cook University of North Queensland, from P. tepikia in lacking spiral grooves, Townsville), Tim Naish & Brent Alloway (School of Earth Sciences, Victoria University of Wellington; reached New Zealand (or possibly evolved formerly of GNS), Brad Pillans (ANU), and the late from P. tepikia) only after OIS7. The largest Tony Edwards. , Pierre Lozouet specimen of P. tepikia recorded so far is not and Virginie He´ ros (MNHN) generously provided from the warm-water northeastern North Is- access to the MNHN collections, library, and type land, where P. angasi occurs at present, but material, and loaned specimens. Tom DeVries (Bur- from Landguard Sand (OIS 9) at Wanganui. ton, Washington State, USA) and Luiz Simone Landguard Sand has produced the largest (Museu de Zoologia, Universidade de Sa˜ o Paulo) number of warm-water migrants at Wanganui. discussed American Austrotoma records and 132 AG Beu

provided references and specimens, and a specimen Absala˜ o RS, Pimenta AD, Caetano CHS 2005. was sent by Laura Schejter (Lab. de Bentos, IN- Turridae (Mollusca, Neogastropoda, Conoidea) IDEP, Mar del Plata). Norton Hiller, S. Stollmon coletados no litoral sudeste do Brasil, programa and Kate McCaughan (CMC), Todd Landers and REVISEE ‘‘Score’’ central. Biocieˆ ncias, Porto Alegre 13: 1947. Owen Lee (AIM), Jack Grant-Mackie and Neville Adams A 1850. Monograph of the Family Bullidae. Hudson (AUGD), Anita Eschner (NHMW), and in In: Sowerby GB II ed. Thesaurus Conchylior- particular, Kathie Way and Amelia McLennan um, or monographs of genera of shells, vol. (BMNH) provided many type data and loaned types. 2(11). London, GB Sowerby. Pp. 553608, pls. Other comparative material was loaned or data 117125. provided by Bill Rudman, Ian Loch, Alison Miller Adams A 1865. On some new genera of Mollusca and Janet Waterhouse (AMS), and Sven Nielsen from the seas of Japan. The Annals and and Steffen Kiel (Institut fu¨ r Geowissenschaften, Magazine of Natural History, Series 3, 15: 322324 [April 1865]. Christian-Albrechts-Universita¨ t Kiel) and Klaus Adams H, Adams A 18531858. The genera of Bandel (Universita¨ t Hamburg) discussed and loaned Recent Mollusca, arranged according to their Quiriquina fossils. Yuri Kantor (AN Severtsov organisation. London, J. van Voorst. Vol. 1, Institute of Ecology and Evolution, Russian Acad- x484 p.; vol. 2, 661 p.; vol. 3, 138 pls. [issued emy of Sciences) discussed Conoidea. Manuel Mala- in 36 parts; dates (vol. 2, p. 661): vol. 1, pp. quias (BMNH) provided information on Bulla 1256, pls. 132, 1853; pp. 257484, pls. 3360, phylogeny. The referee, TA Darragh (Museum 1854; vol. 2: pp. 192, pls. 6172, 1854; pp. Victoria, Melbourne) is thanked for information on 93284, pls. 7396, 1855; pp. 285412, pls. species of Dennantia. The late lamented Andrew 97116, 1856; p. 413540, pls. 117128, 1857; pp. 541661, pls. 129138, 1858; vol. 1, pp. ix Grebneff (Geology Department, University of Ota- 1858]. go) provided information on Epitoniidae and a Adams A, Angas GF 1864. Descriptions of new translation of Nu¨ tzel (1998). Maureen Coomer, species of shells from the Australian seas, in the Pauline Muir, Denise Garbutt, and particularly collection of George French Angas. Proceedings Kristin Garbett (GNSLibrary) found numerous of the Zoological Society of London 1863: obscure references, Marianna Terezow (GNS) took 418428 [April 1864; Trew 1992]. the light photographs, Philip Carthew (GNS) carried Allan J 1959. Australian shells, with related out extensive work on the photos and arranged the living in the sea, in freshwater and on land plates, and Scott Morgan and James Xia (Industrial [revised edition]. Melbourne, Georgian House, xxi487 p. [reprinted 1962]. Research Ltd, Lower Hutt), Marianna Terezow, and Anderson DT 1966. Further observations on the life George Scott (GNS) took the SEM micrographs. histories of littoral gastropods in New South Wales. Proceedings of the Linnean Society of Downloaded By: [Beu, A. G.] At: 20:27 16 March 2011 New South Wales 90: 242251. References Angas GF 1867. List of species of marine Mollusca Abbott RT 1954. American seashells. Princeton, D. found in Port Jackson harbour, New South van Nostrand Co. xiv541 p. Wales, and the adjacent coasts, with notes on Abbott RT 1974. American seashells. The marine their habits, etc. Part I. Proceedings of the Mollusca of the Atlantic and Pacific coasts of Zoological Society of London 1867: 185233. North America. Second edition. New York, van Angas GF 1871. A list of additional species of Nostrand Reinhold. marine Mollusca to be included in the fauna Abbott ST, Carter RM 1999. Stratigraphy of the of Port Jackson and the adjacent coasts of New Castlecliffian type section: 10 mid-Pleistocene South Wales. Proceedings of the Zoological sequences from the Wanganui coast, New Society of London 1871: 87101. Zealand. New Zealand Journal of Geology Ardovini R, Cossignani T 2004. West African sea- and Geophysics 42: 91111. shells (including Azores, Madeira and Canary Abbott ST, Naish TR, Carter RM, Pillans BJ 2005. Is.). Ancona, l’Informatore Piceno. 319 p. Sequence stratigraphy of the Nukumaruan stra- Ascanius P 1772. Philine quadripartita, et førut totype (Plio-Pleistocene, c. 2.081.63 Ma), obekant Sjø-krøk, aftecknadt och beskrifvet. Wanganui Basin, New Zealand. Journal of the Konglige Vetenskapsacadamiens Handlingar Royal Society of New Zealand 35: 123150. 33: 329331. Marine Mollusca of isotope stages of the last 2 million years in New Zealand 133

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