The malacologicalsocietymalacological society of Japan
Cenozoic History of Indo-Pacific and other Warm-Water Elements in the Marine Mollusca of New Zealand'
C. A. FLEMING
(N.Z. Geelogical Survey (D.S.I.R.), Lower Hutt, New Zealand)
== - - a.M 7 Xitiltkec ;}60L 6 rL ti 7 7 F' ?thLll2wt(tgen･tz ip 1"Jts C. A. 7 tr ay]" G'ffilgl Text-figs. 1-6) "The In 1925 Dr J. MARwicK published a paper in the (;edg.{nleboefe Vcrb(t,e,k on ". Indo-Pacific Element in the Marine Tertiary Mollusca of New Zealand At that time, about 1000 species of marine molluscs were known as Cenozoic fossils in New Zealand. During the next two decades, which I have elsewhere called the Golden Age of New Zealand Molluscan systematics (FLEMiNG 1966a), the total number of species was almost tripled through the work of MARwicK, FiNLAy, PowELL, and others, and great advances were made in the biostratigraphy and correlation of New Zealand Tertiary and Quaternery sediments. It therefore seems timely to review the subject on the basis of a new checklist that records a!most 3000 species, classed in 842 genera (FLEMING 1966b). is at New Zealand, in an isolated geographic position in the South West Pacific, Study of and present subject to biotic influence from several directions. fossil faunas floras indicates that these infiuences existed in the past and that their relative im- The most portance has fluctuated during Permian to Recent time (FLEM:NG 1962). important biotic elements (FLEMiNc; 1963) are those that came from the nearest Iand (Australian), from parts of the southern cool temperate zone (Austral), and from the tropics to the north (Malayo-Pacific). The name Malayo-Pacific, used for the elements and of apparent tropical derivation in the New Zealand biota, refers to both plants and animals that used persistent dispersal avenues southward from the Malayan Melanesian regions. The marine mollusca that came by this Malayo-Pacific route are, of course, members of the Indo-West Pacific fauna. the Biogeographic interpretation of Paleozoic and Mesozoic marine faunas uses concept of the Tethys Sea, a vast mediterranean waterway in quasi-Equatorial regions Tethyan of the 0'ld World, characterised by many distinctive tropical and sub-tropical ttt . r.. - --."- ttt t "Evolution, Migration of 1) This paper was p;Esented te symposium No. 4, Distributien, and Eleventh Pacific Science Congress, Tokyo, Plants and Animals in the Pacific Area" at the Japan, 1966. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 106 VENUS: 25 (3.4) 1967 elements generally absent from Boreal and south-temperate areas. During the Terti- ary, earth movements are said to have led to partial fragmentation of the Tethysand to somewhat independent development of the faunas in the separated basins, for in- stance in the Caribbean and Mediterranean regions, at least after the mid-Tertiary (EKMAN, 1953), but cQmmunication was maintained, for the most part, between the parts of the far-fiung Indo-West Pacific Realm, extending from the Red Sea to Hawaii. The Malayo-Pacific faunal element which plays a fiuctuating role in the Tertiary history of New Zealand marine mollusca was apparently derived from the developing Indo-West Pacific fauna, and came to New Zealand by the immigration ef planktonic larv･ae under the infiuence of favourable currents, either directly from the north or via East Australia, particularly at times when the subtropical and tropical zonal circulatory systems extended further south than at present. Dofnition of Indo-Pactiic Genera For the purpose of this analysis, approximately 842 genera and subgenera of marine Mollusca with a Cenozoic fossil record were classified as (1) endemic to the NewZealand subregion, (2) Australian (known only from New Zealand and temperate Australia), (3) Austral (south-temperate circumpolar), (4) Indo-Pacific, and (5) Cos- mopolitan (cf. FLEMING, 1963). "Indo-Pacific" The term is here used for a broader group of genera than those now restricted to the Indo-West Pacific, and corresponds perhaps more closely with the `C concept of Tethyan" used in an earlier part of the geological record and with "tropicopolitan" the of some writers. As defined here, the Indo-Pacific element in- cludes genera now eharacteristic of the Indo-West Pacific Realm, or of parts of it such as the East Indies and Southern Japan (e.g. Myadora), and in addition genera extending more widely in the Mediterranean and Caribbean (e.g. Ctenoides) and even mto temperate seas (e.g. Pecten) if there is fossil evidence that such extended dis- tributions are the result of colonisation by originally Mediterranean-Indo-Pacific genera. On the other hand, genera extending into Polar seas have been excluded, even if an origin in the Indo-Paeific is not improbable (LimqPsis, Laternula, C)'clocardia). Some extinct genera are included solely on the basis of their fossil distribution and that of living relatives in warm seas (e.g. Conilithes, Atztria). A few endemic gene'ra or subgenera with obvious afinities in the Indo-Pacific have been included (e.g. Tra- cdycardium (Ovicardium), GomPhina' (Goniphinella), Tudiclana (Tudiclidae), Eocithara (Harpidae). Many other endemic genera are probably of Indo-Pacific origin. Indo- Pacific subgenera within a single genus were plotted separately and a few subgenera were plotted separately from the genus (sensu lato) when the latter differs in range from the former. The classification of genera into biogeographic elements depends on imperfect data on their present and past distribution; the decisions are inevitably sttbjective, and will be revised by other workers. For this reason a full list of the NII-Electronic Library Service The malacologicalsocietymalacological society of Japan FLEMiNG : Cenozoic history of Indo-Pacific Elements in N.Z. mollusca 107 genera here classed as Indo-Pacific, with their ranges in New Zealand, is presented as Appendix I. Additional Indo-Pacific genera in the New Zealand Recent fauna, lacking a fossil record, have not been dealt with. IVbw Zbaland Time Scale and Mollzascan Faunas The system of local Cenozoic stages proposed by FiNLAy and MARwicK (1947), with later modifications, was used for plotting ranges. The stages are based primarily on the succession of foraminiferal faunas, supplemented by macrofossils, especially "shelf" above the Miocene. Faunas of Mollusca, mainly representative of environ- ments, are known from most of the 31 post-Cretaceous stages, the only gap being between the Wangaloan (Danian, Paleocene) and Bortonian (Middle Eocene). For this review, the faunas for this interval (Matau Fauna, FiNLAy and MARwicK, 1937; Otaio Gorge Fauna, MARwicK, 1960, etc.) are grouped as Lower Eocene (Heretaungan Stage). The molluscan assemblages known vary considerably in diversity, preservation, and eeological facies,' and some have been studied mote thoroughly than others. These factors undoubtedly influence the analysis and the conclusions derived from it. Yet reasonably rich faunas have been described from all the main divisions of the Cenozoic, and the deficiencies are considered unlikely to falsify the main trends. Method of Analysis Using published records and unpublished identifications in N.Z. GeologicaL Survey files, the writer plotted the known occurrences of each genus and subgenus in the checklist of New Zealand Cenozoic Mollusca (FLEMiNG, 1966b) on a time scale based on the 31 post-Cretaceous stages and substages. The genera classed as Indo-Pacific were then re-plotted to show their total range (inclusive of gaps in the record). In a few cases, major gaps in the record (combined with systematic criteria) were in- terpreted as evidence of renewed colonisation of a genus. For example, the mid- Tertiary record of SPondylus was treated as a different invasion from the Cretaceous record of SPondylus, and the recent record of SePtder as a different invasion from the Lower Miocene record. Doubtless many other double or multiple invasions have been overlooked for lack of suficiently discriminating taxonomy. The recorded ranges have been treated empirically, despite the fact that many records depend on chance discovery or recognition of a single specimen (e.g. Argonattta), and despite the pro- bability that many absences are due to lack of suitable facies (e.g. absence of Patelloida before Holocene). Text-figure 1 shows a section of the plot illustrating the data for 30 of the 175 genera tabulated. NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 108 VENUS: 25 (3.4) 1967 Changes in Immigration Rate The numbers of Indo-Pacific genera appearing for the first time in each stage are plotted in Text-fig. 2. A few Indo-Pacific genera had already appeared in the Upper Cretaceous and some of them (Cucullaea, Glyqymerita) persisted. In interpret- ing the numbers of iticoming Indo-Pacific genera we must not give too much signifi- ALE-CENEEOCENEL]G MIOCENE PLIO-CENEPLEISTOCENE LMUCENE L MU 6)aEa=ana2lEsABlA8af=stasC,"iasS--Jo)a)l)=)=)lj)o)dio=6=Eg8ot C'heilea Xlenopht)ra C.vpraea s. Iat. 7)'ivia s. Iat. Mltltrivia Proterato C.v-praerttto Ertttctpsis ee ProtoopriUtfa Potinices rvFeverittl e Eunattctna e Sigarutotrema Ec'hinopitoria Gaieodea Eu(lolium e lbntr[t Onis'eid[ttt Cotuhraria Mon(iplex Chctrottia Distor"'ie CPersonella)' e Ficus Prof['us e Priscqficus Ptervnotus ' Pterechetus Murek s. str. e Chicoreus T>{phis s. str, Text-fig. 1. Section of plot showing first and last records of Indo-Pacific mollusc genera in the Cenozoic of New Zealand. Recerded ranges indicated by arrows and records in a single stage by circles. 'minor cance to fluctuations. The most important trend is the maximum of Indo- Pacific colonisations in the Lower Miocene. Pre-Miocene fiuctuations are infiuenced by our relatively poor knowledge of the Lower Eocene (later Dannevirke Series) and upperrnost Eocene (Runangan) Mollusca, but there is some evidence that immigration reached a secondary peak in the Eocene. Following the prominent burst of invasions in the Lower Miocene, the abrupt fall almost to zero is an artifact due to ignorance of the Hutchinsonian Mollusca, but a more gradual decrease in immigration rate dur- ing the rest of the Miocene and Pliocene is implied by the data. During the !atest NII-Electronic LibraryMbrary Service The malacologicalsocietymalacological society of Japan FnEMiNG: Cenozoic history of lndo-Pacific Elements in N.Z. mollusca 109 Pliocene (Waitotaran) and earliest Pleistocene (Hautawan) stages no Indo-Pacific colonists are reported. In this interval, which includes New Zealand's earliest Pleisto- cene glaciation, the Subantarctic Zone of surface water seems to have moved north, favouring prominent southern colonists (FLEMiNG, 1944), but limiting, we may judge, the opportunity for successful colonisation from the north. In the warmer and pre- sumably inter-glacial stages Qf the Lower Pleistocene, however, and particu!arly in 30 < nt m z m o 20 in o 10 M pt ra E p zo Text-fig. 2. Histpgram showing numbers of first appearances of Indo-Pacific' genera in each stage of the New Zealand Cenozoic. ' - the CastlecliMan, immigration from the Indo-Pacific was renewed with the appearance of such genera as Anadara, Pecten (s.str.), Hetiacus, Acrilloscala, Ctipulus, Eunaticina, Tonna and Leucotina, some doubtless having reached New Zealand via East Australia, which was the source of other newcomers at the same time. ' Short-lived Immigrants "last When appearances" are plotted in the same way (Text-fig. 3) we find, perhaps eontrary to expectation, that the extinction of Indo-Pacific genera has not been concentrated in a few periods of crisis, but has been well distributed throughout Cenozoic time. No･ less than 39 Indo-Pacific genera (22% of the total) are recorded from single stages, and their extinction apparently followed shortly after their establishment. Twelve of the 32 Indo-Pacific genera that combine to make the Otaian Stage of the Lower Miocene the peak of Indo-Pacific colonisation were not recorded after that stage. Other genera disappeared progressively through the later Miocene, Pliecene and Pleistocene, so that the modern fauna, despite new arrivals, differs notably from the Tertiary Fauna in lacking such Indo-Pacific elements as Cucullaea, NII-Electronic Library Service The malacologicalsocietymalacological society of Japan 110 VENUS: 25 (3.4) 1967 Arca, Isagnomon, Chama, Maoricardium, Solecurtus, HiPPon,yx, Cheilca, Cypraeidae, Polinices, Ficzts, Chicoreus, Olivella, and Conidae. Lotver Miocenc Climax of Indo-Pacifc bij7"ence The greatest number of Indo-Pacific incomers is recorded for the Otaian Stage in the Lower Miocene (lewer Aquitanian), which now includes the rich faunas of Pakaurangi Point described by LAws (1939--1944). The Indo-Pacific genera present in at least northern New Zealand during the Lower Miocene include Saptifer, 1'riclzonz);a, lsognomon, Sl,ondylus, Chama, Pitar, Parmpholas, Rocellaria, Psettdastralium, Strebloceras, Oniscidea, Ptei"'notus, Murex (s. str.), Chicorezas, Cronia, ComllioPhila, Etrema, Eubela, and other Turridae. Together -(r:atmzmo 20 Lo 10 ecveasEpZ, o Text-fig. 3. Histogram showing numbers of last appearances of Indo-Pacific genera in each stage of the New Zea]and Cenozeic, with warmth loving organisms in other phyla, especially hermatypic reef corals reported by SQuiREs (1962), these genera are the basis for the conclusion of New Zealand paleontologists that the peak of Tertiary warmth in the South West Pacific came in the Lower Miocene (BEu, 1966), and not in the Eocene as in the north-east Pacific (DuRHAM, 1950). For the Mollusca, the evidence for a Lower Miocene maximum of subtropical in- fluence is illustrated in Text-fig. 4, which shows the total number of Indo-Pacific genera recorded in each stage. The data of Text-figs. 2 and 4, showing respectively the number of incoming and presumably immigrant genera per stage, and the total number of genera of Indo- "pine-tree Pacific origin in each, have been replotted as a diagram" in Text-fig. 5, using a time scale in which the intervals are proportional to those of the absolute time scale (Geological Society of London, 1964). NII-Electronic Library Service The malacologicalsocietymalacological society of Japan FLmnNa: Cenozoic history of Indo-Pacific Elements in N,Z. mollusca lll 1CX) 80o 9b a)U< ¢ ioa 40z' Lo 20 ecptcaEDOZ Text-fig. 4. Histogram showing tetal number of Indo-Pacific genera of Mollusca in each stage of the New Zealand Cenozoic. Absolute Immigration Rate Text-fig. 5 emphasizes the uneven length of the New Zealand Cenozoic stages. By accepting the current correlation of the stages with the standard divisions of the Cenozoic and the ages in years of the latter (Geological Society, 1964), it is possible to assess the approximate duration of each stage and thus to calculate a rough immi- gration rate per million years. Thus during two Oligocene stages spanning the time from 36 to 27 million years (5% million years each), 11 and 16 Indo-Pacific genera appeared for the first time, at rates of 2 and 3.9 genera per million years. Text-fig. 6, showing the immigration rates for the whole Cenozoic, emphasizes the importance of the Lower Miocene peak of invasion, and of the Lower PIeistocene immigration when Indo-Pacific colonists, few in total numbers, re-colonised at a rapid rate during quite short interglacial intervals. New Zealand Peripheral to Indo-Pacijic Even during the Lower Mio ¢ ene climax of Indo-Pacific influence, when reef corals NII-Electronic Library Service The malacologicalsocletyofmalacological society of Japan 112 VENUS:25 (3･4) 1967 o2 'l::'}'"':'/': '.:1'.]:'.'"- PLEISTOCENE r' ,':' /,: ,: WwWPWo ozpt .:.,=.-"t..n-'C,'';,} al8 7 12 ptzptooE slScSaPuPhPoLw 18 utec 26 taZutUooAo Lo 37enzo:lE tuzmoo" 4s AbDpDhDm 49 54 klj ,.l ptptUopt m n O IO 15 !O 25 6S Tgxt-fig. 5.of Diagram illustrating numbers of Indo-Pacific mollusc genera and Inde-Pacific immigrantfiuc!uatinggenera during the Cenezoic of New Zealand, NII-Electronic Library Service The malacologicalsocietymalacological society of Japan Fi[EMrNG: Cenozoic history of Indo-Pacific Elements in N.Z. mollusca 113 uaec zo:tE ecptm .cCptmzrao -z PALEO-CENEEOCENE LItcENE PLIO-CENE Text-fig.6. Histegram showing fluctuating rate of immigration of Indo-Pacific genera to New Zealand during the Cenozoic. were living in northern New Zealand (SQuiREs, 1962), the number of Indo-Pacific amounted genera to less than a quarter of the fauna, and as mentioned above many of the Indo-Pacific immigrants, throughout the Cenozoic, were short-lived, apparently unable to maintain themselves in New Zealand environments. There is a large list of characteristic Indo-Pacific genera that are not represented in New Zealand (Appendix II). These facts suggest that New Zealand always lay outside the Indo-Pacific Realm, even when Indo-Pacific elements were at their maximum. NII-Electronic Library Service The malaoologioalmalacological societysooiety of Japan ・ 114 VENUS ： 25 （3 4）1967 要 約 3 000 ニ ー ー は 3 4 合 わ せ て 842 属 （亜 属 ）， ， 種 に 上 る が ， ェ ジ ラ ソ ド の 新 生 代 海 産 軟 体 類 化 石 ，第 紀，第 紀 一 ー 3 つ 2 二 ＝ ジ プ ソ ド と 濠 洲 の 温 帯 域 に の み ， （） 南 こ れ 等 は 次 の 5 の 群 に 分 け られ る 。 （1） 固 有 属 種 ， （） 半 球 の 湿 帯 環 極 性， 印度 太 平 洋 系 （古 テ テ ／ ス 海 系 及 び 汎 熱 帯 性 を 含 む ）、 汎 世 界 的 。 い て 太 の 先 ず 各 属 の 地 質時 代 の 各 時 期 に 於 け る 出 現 状 況 を 知 り （第 1 図 ），次 に 或 る 時 期 に 於 印 度 平 洋 系 も 2 に 二 点 が り，続 い て 下 部 中 の で 出 現 の 最 初 が そ の 時 期 に 当 る も の の 数 を と る と （第 図）， 始 新 世 次 的 な頂 あ 一 の に は 新 世 に 最 高 の 頂 点 が 見 られ る が ， 鮮 新世 に 向 か い 漸 減 す る 。 方 鮮 新 世 の 最 後期 と 最新 世 最 初 期 印 度 ニ ージ ー ラ ソ ド に る の 氷 河 期 を 含 み 太 平 洋 系 の 移 住 は 全 く知 られ て い な い 。 こ の 時 期 は ュ 於 け 最 新 世 最 初 ， む し ろ 南 方 冷 水 域 か ら の 移 住 に 好 適 で ， 北 方 暖水 域 か らの 移 住 は 押 え ら れ た と 思 わ れ る 。 し か し 温 暖 な 間 氷 期 と 思 わ れ る 下 部 最 新 世 の 時 期 に は 印 度 太 平 洋 か らの 移 住 が 再 開 さ れ て い る 。 次 に 各 属 の 出現 の 最 後 の 時 点 か の で 22 の か ら見 れ ば （第 3 図 ）特 定 の 時 期 に 多 く絶 減 し て い る よ う な 事 は な い 。 ま た な り 属 （ ％ ） そ 出現 が 単 一時期 に 限 ら れ ，移 住 後 間 も な く消 滅 し て い る よ う で あ る 。 ． Aquitanian 4 最 も 多数 の 印 度 太 平 洋 系 の 属 が 見 ら れ る の は ド部 中 新 統 の Otaian 階 （下 部 ） で （第 図）， z ！1に た の と は 異 南 西 太 「 洋 地 区 で は 下 部 中 新 生 代 に 於 け る 温 暖 化 の 頂 点 が 北 東 太 平 洋 地 区 で 始 新 1こ あ た っ り， の の の 度 上 に わ し （第 5 図） 百 新 世 に あ っ た と 考 え ら れ る 。 各 時 期 に 於 け る こ れ ら の 諸 属 総 数 絶 対 年 代 尺 堤 い IC る 万 年 を 単 位 と し た 移 住 率 を 求 め れ ば （第 6 図） 下 部 中 新 世 と 下 部 最新 世 の 極 く短 か 間 氷 期 と 於 け 移 住 の 最 盛 期 が 明 ら か に 見 られ る 。 ． 一 冖 で に 於 い て さ え r一卩 太 系 の も の は ＝ ＝ ジ ラ ソ ド 印度 太 平 洋 か ら の 彫 響 が 最 高 あ っ た 下 部 中 新 itt ， 度 平 洋 ニ ・一 の の も の が ュ 貝 類 相 の 1 ！4 以 下 で あ っ た し ， ま た 移 住 者 も短 命 で あ っ た 。 ま た 印 度 太 平 洋 諸 属 中 ， 多 く ジ ー プ ソ ドに は 出現 し て い な い （第 2 表 ）。 ’ い ＝ ー ジ ー ラ ソ ド は か ら の が で っ た に 於 て さ え も， 常 に 印度 以 上 要 す る 1C ユ ， 印 度 太 平 洋 影 響 最 酉 あ 時 期 太 平 洋 海 洋生物 の 区 か らは 外 に あ っ た も の と 思 わ れ る 。 References roy ．Soc ． ．1966 ： Sea Temperatures of New Zealand during the Cenozoic Era ， Trans ． BEu ， A ，G ， − N ．Z ． （Geology ） 5 ： 177 187 ． ・ ． Bull ．Ceol．Soc．Amer ． DuRHAM ， J， WyAI v ，1950 ： Cenozoic Marine Climates of the Pacific Coast − 61 ： 1243 64． ” “ ＆ ．London ． EKMAN ， S．，1953 ： Zoogeography of the Sea ． Sidgwick Jackson and Associated Molluscan Faunas of F 】NLAY ，H ．J． and MARwTaK ， 」，，1937 ： The Wangaloan Kaitangata ．Green Is！and Sub −Division ． N ． Z ． Geot． Surv ． pa9 ． Buit．15， 一 and Tertiary ． N ．Z Sci． ，1947 ： New Divisions of the New Zea！and Upper Cretaceous ≠ − Tech ． B ，28 （4）： 228 36 ． of Pliocene Climatic Change in New Zea！and ， Trans ． FliEMIN〔｝，CA ，1944 ： Molluscan Evidence − roy ． Soc ， N ， Z ．74 （3 ）： 207 220 ． ゼ s 20 2 ； 53− 1962 ： New Zealand Biogeography ． APaleontologis Approacll ． Tuatara （） ， 108． and Trans ． ． ： The Nomenc ！ature of Biogeographic Elements in the New Zea ！ Biota． ， 1963 − rey ． Soc． N ． Z ，（Genera1）1 （2）； 13 22， Zea 】and Cenozoic Mollusca． A Historical Survey ． ，1966a ： The Description of the New ’ − 7＞．Z 、ノt Geol ． GeoPkys ． 8 （6）： 1149 74． … ’ with a Checklist of New Zealand ，1966b ： Marwick s Illustrations of New Zealand Shells Cenozoic MoUusca ， NZ ． Dept ． sci ． ind． Res ． BulL 173 ． ・ ．Soc．Lond ． 120S ： Geological Society of I．ondon ，1964 ： Phanerozoic Time Scale ． Quart． f， Geol 260 −2 ， − − Kaipara Nos ．1 3． Trans ．ア 尸． 1939 44 ： The Molluscan Fauna at Pakaurangi Point， ， の LAws ， C ．R ， 一 NII-ElectronicN 工 工 Eleotronio Library Service The malacologicalsocietymalacological society ofJapanof Japan FgreMtNa: Cenozoic histery of lndo-Pacific Elements in N.Z, mollusca 115 Soc. N.Z. 68: 466-503; 71: 134-51; 73: 297-312. MAxwicK, J., 1925: The Indo-Pacific Element in the Marine Tertiary Mo]lusca of New Zealand, Gedenkboek Verbeek, 8: 369-77. , 1960: Early Tertiary Mollusca from Otaio George, South Canterbury, N.Z. Geol. Surv. Pal. Bult, 33. SQuiR]ts, D.F., 1962: Additional Cretaceous and Tertiary Corals frum New Zealand, Trans. ray. Soc. N,l (Geology) 1 (9):133-50. Appendix I Jndo-Pacifc Genera in the New Zeatand Cenoaoic Motlusca Geelogical distribution is given in New Zealand stage symbols (FiNbAy and MARwicK, 1947), and in the Standard Tertiary divisions. Ctenoides : L. Miocene(Po)-Pliocene(Ww) BIVALVIA Crassostrea: IJ, Miocene? (Tt)-Pliocene Nucinella: U. O]igocene (Ld)-Recent (Ww) Cucuilaea (Latiarca): U, Cretaceous NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan 116 VENUS: 25(3 ･ 4) 1967 Hkeliotis (Sulculus): L. Miocene (Po)-Recent Polinices : Paleocene (Dt)-Pliocene (Ww) Scutus: U, Eocene Pterochelus: U. Eocene Miocene Heltacus : L. Pleistocene (Wc)-Recent (Ak)-L. and L. Pleistocene SPirolaxis : L. Miocene (Po) CPa) (Wn) -Recent Strebloceras: L. Miocene (Pa)-M. Miocene (sw)IJ. Mblrex (s.str.): L. Miocene (Po) Ilyrazus: Oligocene (Ld)-M. Miocene Chicoreus (Sirat"s): L. Miocene (Po-Sc) L, Acrilloscala : L. Pleistocene (Wok) CorallioPizila: Miecene (Po-Sc) Cirsotrema : M. Eocene (Ab)-Recent Zafra: L. Miocene (Pa)-I.. Pleisto- cene EPitonium (RZyaloscala) : L. Miecene (Pa)- (Wn) RecentL. Tudiclidae CTudiclana): Paleocene (Dt) -Pliocene Clathroscala : Miocene (Po) (Wo) Harpidae (Eocithara):L. Eocene (Dm)-U. Ciathrus : L. Miecene (Lw) Oligocene (Ld) Ba,zyspira L, Eocene Aciis :Korovina L, Pliocene (Wo)-Recent (s.Iat,): (Dh)-Recent L. Miocene(Po)- Pliocene(Wp) : Pliocene (Wo) (SPinasPira): -Pliocene Otivetla M. Miocene HiPPonyx : U. Eocene (Ak) (Wp) (Lamprodomina): (Sl)- Plio¢ ene Dall'iella: L, Miocene (Po) (Wp) Lvria: L. Miocene CaPulus : L, Pleistocene (Wc) (Sc) -l'liocene Cheitea : U. Eocene (Ak) (Wo) APhera: L. Miocene (Pa) XenoPhora : U. Eocene (Ak)-Recent BoneUetia: M. Eoccne (Ab)-Recent Cypraea (s.lat. including ? Bernayia and ? Merica: L. Miocene(Po)-Pliocene(Wp) EocyPraea): M. Eocene (Ab)-U. Miocene 7'rigonostoma: Oligocene (Lwh-Ld) (Tt)U, [1initas: U, Eocene (Ak)-L. Miocene Trivia (s.lat,) Eocene(Ak)-PliocenetWo) (Pa) Ellatrivia : L. Miocene (Sa)-Recent Marginella (Kogomea): L. Oligocene (Lwh)- Proterato : U, Eecene (Ak)-Pliocene CWp) Recent CYPraerato : L, Miocene (Pa-Sc) (Glabella): L. Oligocene (Lwh)-Recent Protoaypridea : L. Miocene (Po) (Volvarina): PIiocene (Wp)-Recent EratoPsis : L. Miocene (Po) (Volvarinella): M. Eocene (Ab)-Recent NII-Electronic Library Service The malacologicalsocietymalacological society ofJapanof Japan FbEMTN{x Cenozoic history of Indo-Pacific Elements in N.Z. mollusca 117 Closia : L. Miocene (Po)-Recent Leucetina : L, Miecene (Po-Pa) and L. Conus (s.lat,): M. Eocene CAb)-U, Miocene Pleistecene CWn-Ht) (Tk) PuPa;Crenilabium L. Miocene (Pa)-Recent Conilithes (==Conospirus): U. Eocene (Ak)- : L. Miocene (Lw)-Pliocene U. Miocene (Tt) (Wp) Gemmula : M. Eocene (Ab)-Pliocene(Wo) Tornatellaea : Paleocene CDt)-L. Eocene (Dh) BathNtoma: L, Miocene (Lw)-U. Miocene Ringicula: U. Oligocene (Ld)-Recent (Tk) Atys(Agiculastrum): L. Miocene (Po-Sc ?) MicantaPex: L. Miocene (Sa)-Recent Limulatys: L. Miocene (Po-Pa) CryPtoconus: L. Eocene (Dh) Ryruncutus: L. Miocene (Sc) Marshallena: M. Eocene(Ab)-Pliocene(Wn) Rhi2orus : L. Miocene(Pa)-Pliocene(Wp) inquisitor: L. Oligocene (Lwh)-M, Mio- Sinuatodostomia: Pliocene (Wo) (? Lower cene (Sw) Miocene, Pa) S)rntomodritlia:L. Miocene (Sa-Sc) Creseis: L. Miocene (Lw>-U. Miocene Tomopleura: Paleocene (Dt)-M, Miocene (Tt) (Sw) Ilyalocylis: U. Eocene (Ak-Ar) Microdriliia: L. Miocene (Po) Vaginelia: L. Miocene (Po-Sa) TurridruPa: M. Miocene (Sl) Siphonaria : Pliocene (Wp)-Recent Borsonia: M. Eocene (Ab)-L, Miocene POLYPLACOPHORA (Sc) Cordieria: U, Eocene (Ak) Craspedochiton (Notopiax): U. Oligocene (Ld) -Recent Etrema: L. Miocene (Po) Anacithara : U, Oligocene(Ld)-M. Miocene CEPHALOPODA (Sl) Eubeta: L. Miocene (Po)-U, Miocene Aturia: L, Eocene (Dt)-U. Miocene (Tt) (Tt) Clin"ra: L. Miocene (Pa ?) Hlercoglossa: M, Eocene (Ab) Thatcheria : U. Miocene(Tt)-Pliocene(Wo) EutrePhoceras: U, Cretaceous(Mp)-U, Eocene Perirhoe (Dimidacus): U. Miocene(Tk)-Plio- (Ak) cene (Ww) Argonauta: Pliocene (Wo)-Recent Appendix II Some lvdo-Pacinc Genera of Motlusca Absent from N,Z. Cenozoic ･ Trisidos Strombus Antigvna HexaPlex Pinctada LambisRostelZaria Lioconcha Marchia Vulselta CirceGafrarium DruPaMancinelta Malleus T"rbinellu Annachtamys TibiaCassisCyPraecassis AsaPhis 1)reneBabylonia Gloripallium Trochus (s.str.) Amusium Chtorostoma HdmpaOlivaVasumXLincus Placuna Natica (s.str.) TectusTurbo Tridacna Natiearius (s.str.) Hippopus SePta(lymatiu}n Phasianelia Lentiltaria (s,str,) Delphin"la (Angaria) C)'mbiotacca Codafeia Distorsio (s. str.) GenaTectarius AuticaMeloCymbium Fimbria (Corbis) Bblrsa(lyrineum Fragum Cerithium Discors BiPlexMaleaHlzusteZlum Rhinoclavis Terebra (s,str,) llbmicardium Terebralia 7'urris ArcoPagia TelescoPium NII-Electronic Library Service