ISSN 0031-0204

Transactions and Proceedings

of the

Palaeontological Society of Japan

New Series No. 165

Palaeontological Society of Japan

April 30, 1992 Co-Editors Tsunemasa Saito and Kei Mori

Language Editor Martin Janal (New York)

Editorial Board

Kunihiro lshizaki (Tohoku University), Yukio Isozaki (Yamaguchi University), Tomoki Kase (National Science Museum), Itaru Koizumi (Hokkaido University), Yasumochi Matoba (Akita University), Kei Mori (Tohoku University), Hiroshi N oda (University ofTsukuba), Tsunemasa Saito (Tohoku University), Yokichi Takayanagi (Ishinomaki Senshu University), Kazushige Tanabe (University of Tokyo), Yukimitsu Tomida (National Science Museum), Kazuhiko Uemura (National Science Museum)

Officers for 1991-1992

President: Kiyotaka Chinzei Honorary President: Teiichi Kobayashi Councillors: Kiyotaka Chinzei, Takashi Hamada, Y oshikazu Hasegawa, Itaru Hayami, Hiromichi Hirano, Hisayoshi Igo, Noriyuki Ikeya, Junji Itoigawa, Tomoki Kase, Makoto Kato, Tatsuaki Kimura, Itaru Koizumi, Kei Mori, Hiroshi Noda, Ikuo Obata, Kenshiro Ogasawara, Tomoo Ozawa, Tsunemasa Saito, Yokichi Takayanagi, Kazushige Tanabe Members of Standing Committee: Hiromichi Hirano (General Affairs), Tomoki Kase (Finance), Tsunemasa Saito (Editor in Chief, TPPSJ), Kei Mori (Co-Editor, TPPSJ), Noriyuki Ikeya (Planning), Hiroshi Noda (Membership; Co-Editor, Special Papers), Kazushige Tanabe (Foreign Affairs), Haru Hayami (Editor, "Fossils"), Juichi Yanagida (Co-Editor, Special Papers), Tatsuaki Kimura (Friends of Fossils) Secretaries: Akito Asai, Tatsuo Oji (General Affairs), Masanori Shimamoto (Editorial of l'PPSJ), Hiroshi Kitazato (Planning), Katsuo Sashida (Membership), Yukimitsu Tomida (Foreign Affairs), Katsumi Abe (Editorial of "Fossils"), Takeshi Ishibashi (Editorial of Special Papers), Tamiko Ohana (Friends of Fossils) Auditor: Kuniteru Matsumaru

The fossil on the cover is Trilophodon sendaicus Matsumoto, an extinct elephant, which was described from the Pliocene Tatsunokuchi Formation developed in the vicinity of Sendai, Northeast Honshu, Japan. (IGPS colI. cat no. 87759 (A), length about 18.5 cm)

All communication relating to this journal should be addressed to the PALAEONTOLOGICAL SOCIETY OF JAPAN c/o Business Center for Academic Societies, Honkemagome 5-16-9, Bunkyou-ku, Tokyo 113, Japan Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1009-1023, 3 Figs., April 30, 1992

932. MOLLUSCAN FAUNA FROM THE LATE MIOCENE HATS USE FORMATION IN THE MIURA PENINSULA, KANAGAWA PREFECTURE, JAPAN*

KIYOSHI OKUMURA

Department of Natural Science, Naruto University of Education, Naruto City, Tokushima Prefecture, 772

and

YUT AKA Y AMAGISHI

Kurihama Junior High School, Yokosuka City, Kanagawa Prefecture, 238

Abstract. The Hatsuse Formation distributed in the southern part of the Miura Peninsula mainly consists of well-bedded marine tuff breccia, lappili tuff, and scoriaccous sandstone and siltstone of marine origin with intercalated thin layers of pumice and conglo­ meratic sandstone. The molluscan fossils occur from a rather coarse-grained sandstonc associatcd with fragments of pyroclastic rocks. The Hatsuse Fauna named herein is com­ posed of three species of Gastropoda, one species of Scaphopoda, and 15 species of Bivalvia. The Hatsuse Fauna is characteristically an admixture of euneritic and subneritic forms, though all of them are warm-water dwellers. In consideration of specics composition and their mode of occurrence, the shallow-water dwellers arc interpreted to bc the ones which were displaced into the outcr neritic environment. The geologic age of the Hatsuse Fauna indicates Late Miocene, and is correlated with the Zushi Fauna distributed in the northern part of the Miura Peninsula, Kanagawa Prefecture and the Senhata Fauna of the Boso Peninsula, Chiba Prefecturc.

Key words. Hatsuse, Miura, Zushi, Senhata, Mollusca.

Introduction ed in the southern part of the Miura Penin­ sula have been lithologically divided into the The Hatsuse Formation was originally Misaki, Aburatsubo and Hatsuse Formations described by Suzuki (1932) for the marine in ascending order, and those formations have strata distributed around Hatsuse-Machi in tentatively been correlated with Tertiary the northern part of Miura City, southern­ strata in the northern part of the Miura most part of the Miura Peninsula, Kanagawa Peninsula (Akamine el al., 1956; Mitsuna­ Prefecture. He also reported a large number shi and Yabe, 1968). The precise geological of molluscan fossils from this formation, age of these strata has remained undeter­ though these fossils were undoubtedly col­ mined. One important reason for this is that, lected from part of the Pleistocene Miyata hitherto, no fossils have been found from the Formation. Hatsuse Formation. Subsequentely, the Tertiary strata distribut- Recently, numerous fossil molluscs were 'Received Sepember I, 1991; accepted Decembcr 20, collected by the present authors from the 1991 Hatsuse Formation. This is probably the

1009 1010 Kiyoshi Okumura and Yutaka Yamagishi first fossil collection from this formation. Formation (Pleistocene) consists mainly of This fauna i ncl udes some characteristic loose sandstone. species as shown in Table 1. It can be con­ The Hatsuse Formation shows a strike tren­ sidered that those fossil moll.uscs carry very ding generally from N 60"W to EW with the important information for age determination exception of the N agahama and Arasaki areas and correlation of the Tertiary strata in this where it trends from N 30"E to N60"E. Also, area. marked folds and faults which trend in a The purpose of the present work is to northwest-to-southeast direction are devel­ discriminate the molluscan fossils from the oped in this formation. Hatsuse Formation, to attempt to determine There are three major synclines in Mito, the geological age, and to correlate these Nagahama and Kaneda and two anticlines in strata with molluscan faunas in the northern Kurosaki-no-Hana and Kami-Miyada. part of the Miura and Boso Peninsulas. Their axes cross the line of strike of the Hatsuse Formation (Figure I). The Acknowledgments Minami-Shitaura and Hikihashi faults are reverse faults thrusting up from southwest to The authors wish to express their apprecia­ northeast and can be observed in several tion to Dr. Saburo Kanno, former Professor places. The Hikihashi Fault, one of the of the University of Tsukuba for his kind thrust faults, is the boundary between the encouragement during the present study and Hatsuse and Aburatsubo Formations. critical reading of the manuscript. The maximum thickness of the Hatsuse Formation is estimated to be about 460 m by Brief note on the geology around Akamine et al. (1956). the fossil localities Fossil localities and fossil occurrences The Miocene strata, in which the present fossil localities are incl uded, are overlain Molluscan fossils were collected from two unconformably by the Pleistocene Miyata localities near the Wakamiya Shrine, Kanda, Formation. According to Akamine et al. Hatsuse-machi, Miura City. One (Loc. A) is (1956), the Miocene strata distributed in the a small exposure on the east side of the Public southern part of the Miura Peninsula can be Hall of Kanda region, Miura City and other divided into three stratigraphic units: Misa­ one (Loc. B) is a small cutting exposed on the ki, Aburatsubo and Hatsuse Formations in mountain foot about 250 m south of the ascending order. These formations are main­ shrine, and about 450 m northeast of Loc. A ly characterized by pyroclastic rocks. (Figure I). Both fossil localities can be The Misaki Formation is overlain confor­ regarded as of equivalent stratigraphic posi­ mably by the Aburatsubo Formation, which tion. is overlain partially by the Hatsuse Forma­ Loc. A: Molluscan fossils occurred from a tion. The Hatsuse Formation consists main­ massive, conglomeratic tuffaceous sandstone. ly of brownish or yellowish andesitic tuff Most of the molluscs are sporadically scat­ breccia and lapilli tuff which are partially tered, and are mostly inner or outer molds, interbedded with scoriaceous sandstone, pyro­ except for the following species; Glycymeris clastic siltstone and thin layers of pumic and rotunda (Dunker), Chlamys miurensis (Yo­ conglomeratic sandstone. Cross-laminations koyama), Miyagipecten matsumoriensis are well developed in this formation and serve Masuda, and Amussiopecten akiyamae to distinguish it from the underlying Abura­ Masuda. tsubo Formation. The overlying Miyata The dominant species at Loc. A include 932. Molluscan Fauna from the Hatsuse Formation 1011

Glycymeris rotunda, Chlamys miurensis, rocky or gravelly bottoms at depths ranging Amllssiopecten akiyamae, Miyagipecten ma­ from about 50 to 200 m (Habe and Kosuge, tSlimoriensis, Laevicardium sp., and Clinocar­ 1967), whereas the latter inhabits rocky or dillm sp. shelly bottom from littoral depths to 180 m Loc. B: Molluscan fossils occured from a (Keen, 1963). Another rather deep-water pyroclastic, conglomeratic sandstone. F os­ dweller is Fissidentalium found at Loc. Band sils are sporadically scattered as in Loc. A, this species presently lives from about 50 to but are more dense in this locality than in 200 m depth (Habe and Kosuge, 1967). Loc. A. The dominant species from this In terms of faunal similarities between the locality are as follows; Glycymeris rotunda, A and B localities, only a few species are In docrassatella cf. tenuilirata (Shuto), common to both, namely Glycymeris rotunda, Laevicardium a ngustum (Y 0 koyama), Chlamys miurensis, Amussiopecten akiyamae Oxyperas bernardi (Pilsbry), and Anisocor­ and Azorinus abbreviatlls. Other than these bula cf. venusta (Gould). four, the two localities are characterized by The molluscan fossils from both localities different molluscs. are poorly preserved; even such thick-shelled J udgi ng from the foregoi ng, it is clear that molluscs as Cypraea, Clinocardium and the present molluscan association represents a Anisocorbula are represented by water-worn mixture of shallow and rather deep-water fragments or incomplete specimens, though a dwellers, and of rocky- or shelly-bottom few species are rather well preserved. dwellers and sandy- or muddy-bottom dwellers. These species are considered to Characteristics of the Hatsuse Fauna have been transported for some distance from the initial living site in shallow to deep The molluscan fossils discriminated from waters. the Hatsuse Formation incl ude three species Most of the species constituting the present of Gastropoda, one species of Scaphopoda molluscan association are shallow, warm­ and 15 species of Bivalvia. Among them, water dwellers with the exception of a few two species of Gastropoda, one species of deep-water dwellers. The three extant Scaphopoda, and II species of Bivalvia are species among the fauna, namely Glycymeris identified specifically. One species of rotunda, Oxyperas bernardi, and Azorinus Bivalvia is discriminated as a new species abbreviatus, are known to be distributed in (Table I). the warm Kuroshio Current region and in From a bathymetrical viewpoint, most of temperate waters off the coast of southern to the fossil molluscan species from Loc. A are central Japan (south of latitude 3YN, namely considered to be shallow-water dwellers in south of the Boso Peninsula). fine-grained sandy or silty bottoms because of the presence of such genera as Cypraea and Geological age and correlation A zorin us whose present-day distribution ranges from the intertidal zone to 50 m depth. Some species of the Hatsuse Fauna are However, other of the molluscs could pos­ restricted to strata of Miocene age. sibly live in deeper waters down to about 300 Miyagipecten matsumoriensis has been report­ m. The molluscan fossils from Loc. Bare ed from middle to late Miocene strata dis­ also represented by rather shallow-water tributed in areas north of the Kanto region. inhabitants in fine-grained sandy or silty Amussiopecten akiyamae has been reported bottoms except for Mikadotrochus and Cre­ from late Miocene formations in Inagozawa pidula. and Zushi (Masuda, 1962) and also in Sen­ The former genus dwells on deep-water hata (O'Hara and Ito, 1980; described as A. 1012 Kiyoshi Okumura and Yutaka Yamagishi

Table 1. Geologic distributions of the molluscan fossils from the Hatsuse Formation.

Miocene Local. locali ties Plio. Pleist. Recent Species E M L A B

~ h- 1 Mikadotrochus cf. yoshiwarai + 2 Crepidula cf. nidatoriensis hr-- + 3 Cypraea sp. hr-- + 4 Fissidentalium cf. yokoyamai hr-- + 5 Glycymeris rotunda + + 6 Chlamys miurensis + + 7 Amussiopecten akiyamaea r-- + + 8 Miyagipecten matsumoriensis + 9 Indocrassatella cf. tenuili rata hf-- + 10 Laevicardium angustum + 11 L. sp. hr-- + 12 Clinocardium cf. hataii hf-- + 13 C. hatsusense, n. sp. hr-- + 14 C. sp. hr-- + 15 Pitar kaniei f-- + 16 Oxyperas bernardi + 17 Azorinus abbreviatus + + 18 Ani socorbura cf. venusta hI- + 19 A. sp. hI- +

Abbreviation Plio.: Pliocene, Pleist.: Pleistocene, E: Early, M: Middle, L: Late, Local.: Fossil local­ ities, h: Known only from the Hatsuse Formation. praesignis). To the best of the present Zushi Formation (Shikama, 1973) and the authors' knowledge, this species seems to have Senhata Formation (O'Hara and Ito, 1980). a restricted stratigraphic range from the latest From these data, the molluscan fauna from middle Miocene to late Miocene. Chlamys the Hatsuse is correlated with the Zushi miurensis was originally described by Y 0- Fauna (Shikama, 1973) in the Miura Penin­ koyama (1920) from the basal part of the sula and with the Senhata Fauna (O'Hara Zushi Formation in Zushi City. Subse­ and Ito, 1980) in the 80so Peninsula. quently, Shikama (1973) reported it from the This correlation is supported by the geo­ basal part of the Zushi, O'Hara and Ito (1980) logic ranges of the respective constituent from the Late Miocene Senhata Formation of species of the Hatsuse Fauna (Table I). the 80so Peninsula, Ozaki (1954) from the Moreover, this correlation agrees well with basal conglomerate of the "Tyosi" Pliocene in that proposed by Mitsunashi and Yazaki Choshi City, and Kamada and Hayasaka (1968) and Mitsunashi et al. (1979), who (1959) from the Pliocene of Futaba-Tomioka based their correlation on the tephra distribu­ area in Fukushima Prefecture. Mikadotro­ tion in the southern Kanto region including chus yoshiwarai was originally described the 80so and Miura Peninsulas. According from the "Tyosi" Pliocene by Ozaki (1954). to their result, the typical HK and OK key It was also recorded from the late Miocene tephra beds are recognized in the Hatsuse and 932. Molluscan Fauna from the Hatsuse Formation 1013

Senhata Formations in the Miura and Boso basal part of the Zushi Formation (late Peninsulas (Omori et al., 1986). Miocene) of the Miura Peninsula (Shikama, 1973) and the Senhata Formation (late Systematic description Miocene) of the Boso Peninsula (O'Hara and Ito, 1980), besides the type locality in Choshi Family Pleurotomariidae City (probably from the basal part of the Genus Mikadotrochus Lindoholm, 1927 Naarai Formation, late Pliocene). Mikadotrochus cf. yoshiwarai (Ozaki, 1954) Registration number.-NUH [Naruto Uni­ versity palaeontological collections from the Figure 2-1 a, b Hatsuse Formation] 87001. Compared with.- Pleurotomaria yoshiwarai Ozaki, 1954, p. 9-10, pis. Family Crepidulidae 1-2. Genus Crepidula Lamarck, 1799 Mikadotrochus yoshiwarai (Ozaki). Shikama, 1973, p.92, pI. 16, figs. 3-4; O'Hara and Ito, 1980, p. Crepidula cf. nidatoriensis Otuka, 1934 138-139, pI. 7, figs. la-c. Figure 2-5a, b, c Remarks.-The specimen at hand is incom­ Compared with.- plete. The characteristic features of the pres­ Crepidula nidatoriensis Otuka, 1934, p. 626, pI. 48, ent species are as follows; shell rather thin, figs. 63a-b. depressed trochiform with more than six whorls; each whorl slightly convex; surface Shell medium for the genus, semi-ovate in of whorls ornamented with many spiral outline, with two whorls; apex strongly tur­ threads bearing granules which are formed by ned posteriorly and nearly touching body their intersection of growth lines. whorl; surface almost smooth except for fine The original description of the species by radial lines running from central part of Ozaki (1954) states "surface with fine, smooth, dorsal area to aperture. Length 21 mm, revolving striae crossed by growth lines." height 16 mm, width 20 mm, minimum diame­ However, the Hatsuse specimen differs slight­ ter of aperture 16 mm. ly from the type specimen in its whorl sculp­ Remarks.-The single specimen in hand is ture. The difference between these two speci­ somewhat abraded. The present specimen is mens is possibly due to their state of preserva­ close to C. nidatoriensis Otuka, 1934, report­ tion; actually, the type specimen appears to ed from the Kadonosawa Miocene, but the be an abraded shell. A narrow and obscure present species is distinct by having a more slit band revolves at a position somewhat conspicuous apex, which more strongly anterior to the middle of the whorl. In other curves posteriorly than that of the latter, and words, five spiral striae above (poste.rior of) the fine radial lines running from the central the slit and three spirals beneath the slit are part of the dorsal area to the aperture and counted on the fifth whorl. Base and umbili­ crossing it at a right angle. C. nidatoriensis cal portions are missing. sogai Kamada, 1962, reported from the The present species is closely related to N umanouchi and Kokozura Miocene in the Mikadotrochus yoshiwarai (Ozaki, 1954) loban district is similar to the present speci­ described from the Neogene of the Choshi men, but the strongly curved apex and fine (Tyosi) district, Chiba Prefecture, but the radial lines on the dorsal part in Crepidula cf. incomplete, small, immature specimen under nidatoriensis serve to distinguish it from discussion does not permit more precise com­ Kamada's subspecies. This specimen pos­ parison with the species. sibly represents a new form, but is insufficient M. yoshiwarai has been reported from the to permit the establishment of a new species. 1014 Kiyoshi Okumura and Yutaka Yamagishi

Registration number.-NUH 87002. Family Glycymeriidae Genus Glycymeris da Costa, 1778 Family Cypraeidae Subgenus Glycymeris s.s. Genus Cypraea Linnaeus, 1758 Glycymeris (Glycymeris) rotunda Cypraea sp. (Dunker, 1882)

Figure 2-4a, b, c Figure 2-2a, b

Remarks.-More or less depressed incom­ Glycymeris rotunda (Dunker). Makiyama, 1927, p. plete specimens are at hand. These speci­ 29, pI. 1, figs. 1-4; Makiyama, 1952, p. 131-138; Hayasaka, 1962, pI. 45, figs. 8a-9b; Habe and mens are characterized by having no visible Kosuge, 1967, p.126, pI. 46, fig. 15; Kuroda, spire, body whorl covered completely by Habe and Oyama, 1971, p.532, pI. 71, fig. 6; callus, and narrow aperture with denticulated Matsukura 1986, p.89, pI. 6, figs. 14, 15; outer lip. Cypraea is usually distributed in Okumura, 1988, p. 39, pI. 12, fig. I. warm temperate to tropical regions. Pectunculus yamakawai Yokoyama, 1922, p. 190, pI. 16, figs. 4, 5. Registration number.-NUH 87003. Remarks.-The present species is re­ Family Dentaliidae presented by the following characteristics: I) Genus Fissidentalium Fischer, 1885 thick and solid, obliquely suborbicular shell, Subgenus Fissidentalium s.s. medium in size, more or less inflated shell; 2) Fissidentalium (Fissidentalium) cf. yo­ beak pointing slightly forwards; 3) lig­ koyamai (Makiyama, 1931) amental area narrow, triangular with sub­ equilateral chevron grooves; 4) about ten Figure 2-9 strong teeth arranged obliquely in anterior Compared with.- and posterior sides; 5) surface sculptured by Dentalium (Fissidentalium) yokoyamai Makiyama, numerous low radial ribs, and inner margin 1931, p.44, pI. 1, fig. I. with about 40 crenations. Several well preserved articulated as well as Shell slender with elongate tube, gradually nonarticulated valves were collected. This attenuating toward apex; slightly curved species occurs rather frequently at both local­ dorsally; shell very thin with large numbers ities A and B. of flat-topped, slightly elevated longitudinal Geographic distribution.-N. lat. 31" to 42" riblets crossed by fine growth lines; longi­ (Kuroda and Habe, 1952). This species lives tudinal ribs bifurcated, separated by inter­ in fine-grained sandy bottoms in a depth spaces which are wider than the ribs. Lon­ range of 30-300 m. gest specimen at hand; 40 mm in length, Geologic range.-Late Miocene to Recent. diameter of aperture 7 mm, and 3 mm at apex. Registration number.-NUH 87005. Remarks.-The present species is closely allied to Makiyama's species described from the Kakegawa Pliocene, Shizuoka Prefecture, Family Pectinidae and D. (F.) watanabei Kanno from the Chi­ Subfamily Chlamydinae chibu Basin, Saitama Prefecture, but the pres­ Genus Chlamys R6ding, 1798 ent specimens differ from the latter two by Chlamys miurensis (Yokoyama, 1920) having numerous and fine longitudinal riblets Figure 2-8, 10, 11 on the external surface. However, the few Pecten miurensis Yokoyama, 1920, p. 157-158, pI. 12, rather abraded specimens at hand do not figs. 2-6. permit the establishment of a new species. Chlamys miurensis (Yokoyama). Ozaki, 1954, p. 15, Registration number.-NUH 87004. pI. 9, figs. 1-2; Kamada and Hayasaka, 1959, p. 932. Molluscan Fauna from the Hatsuse Formation 1015

N t

Tokyo Bay ~ [J]J 2 D 3

Mito I I 4 * 5 ~ 6

o 0.5 lkm I I I

Figure 1. Fossil localities in the Miura Peninsula 1 : Hatsuse Formation (Miocene) 2: Miyata Formation (Pleistocene) 3: Alluvium deposits 4: Fault 5: Syncline 6: Fossil locality 10 16 Kiyoshi Okumura and Yulaka Yamagishi 932. Molluscan Fauna from the Hatsuse Formation 1017

21, pi. 2, fig. 2; Masuda, 1962, p. 177, pi. 23, fig. 87008. I ; Shikama, 1973, p. 166, pi. 17, fig. 4; O'Hara and Ito, 1980, pi. 16, fig. 2. Subfamily Pectininae Genus Amussiopecten Sacco, 1897 Remarks.-This species was originally de­ Amussiopeclen akiyamae Masuda, 1962 scribed by Yokoyama (1920) from the Figure 3-4a, b Tagoegawa Conglomerate Member (the basal conglomerate of the Zushi Formation) dis­ Amllssiopecten planicostlilatlis (Nomura and Niino). tributed in the northern part of the Miura Akiyama, 1957 (non Nomura and Niino, 1932), Peninsula. This species is characterized by p. 32, pi. 7, figs. 2-4. Amllssiopecten akiyamae Masuda, 1962, p.224, pi. having 22-25 rather compressed radial ribs on 27, figs. 1-3; Masuda, 1986, pi. 2, fig. 4. the surface which bear several riblets on the Amllssiopecten iitomiensis (Otuka). Shikama, 1973, respective radials; the radial ribs appear to pi. 17, fig. 6. be flush with the surface owing to abrasion, A I11l1ssiopecten praesignis (Yokoyama). 0' Hara and but there are some scaly ri blets as seen in the Ito, 1980, p. 131-132, pi. 14, figs. 5-7; pi. 15, figs. 1-5. ventral margin illustrated in Figures 2-9, 10, II. Remarks.-This species was originally de­ Two or three intercalary riblets bearing scribed by Masuda (1962) based on specimens distinct scales appear in the interspaces of the collected from the Inagozawa Formation in radial ribs. This species closely resembles Chiba Prefecture. Amussiopecten planicos­ Chlamys kaneharai which has a wide distri­ tulatus described by Akiyama (1957) based bution essentially in middle Miocene strata in on specimens collected from the Zushi For­ the Kanto region, but C. miurensis is distin­ mation is also included in the present species. guished by having several riblets on each A. iitomiensis (Otuka) can be distinguished radial ribs, whereas only three riblets are from the present species by having fewer but developed on each radial of C. kaneharai. more prominent radial ribs than those of the Moreover, among the three radial riblets of C. present species and no intercalary threads kaneharai, the central one is most conspicu­ between the radials. A. praesignis (Yo­ ous. I n addition, this species possesses two koyama) also differs from the present species or three intercalary riblets in the interspaces by having squarish, rather high, flat-topped of the radials, whereas C. kaneharai is pro­ radial ribs on the right valve and rather broad vided with only one scaly riblet between and roof-shaped radials on the left valve. adjacent radial ribs. Geographic distribution.-This species have Geographic distribution.-Pacific coast, been reported only from the southern Kanto south of Fukushima Prefecture to Shizuoka region, the 80so and Miura Peninsulas. Prefecture. Geologic range.-Late Miocene. Geologic range.-Late Miocene to late Registration number.-NUH 87009. Pliocene. Registration number.-NUH 87006,87007, Genus Miyagipecten Masuda, 1952

~ Figure 2. Molluscan fossils from the Hatsuse Formation (l). la-b. Mikadotrochlls cf. yoshiwarai Ozaki, X 1.2, Loc. B, Reg. No. NUH 87001. 2a-b. Glycymeris (Glycymeris) rotllnda (Dunker), X I.l, Loc. A, Reg. No. NUH 87005. 3a-b. Clinocardillm cf. hataii Hayasaka, X 1.1, Loc. A, Reg. No. NUH 87015. 4a-c. Cypraea sp., X 1.3, Loc. A, Reg. No. NUH 87003. Sa-c. Crepidlila cf. nidatoriensis Otuta, X 1.2, Loc. B, Reg. No. NUH 78002. 6a-b. Laevicardilll11 sp., xO.8, Loc. A, Reg. No. NUH 87014. 7. Clinocar­ dilll11 sp., X I.l, Loc. A, Reg. No. NUH 87017. 8,10,11. Chlal11Ys l11illrensis (Yokoyama), xO.6, 8: Loc. A, Reg. No. NUH 87006, 10: Loc. B, Reg. No. NUH 87007, II: Loc. A, Reg. No. NUH 87006. 9. Fissidentalilll11 (s. s.) cf. yokoyamai Makiyama, X l.1, Loc. B, Reg. No. NUH 87004. 1018 Kiyoshi Okumura and Yutaka Yamagishi

Miyagipecten malsumoriensis Masuda, 1952 The anterior shell margin is rounded and the posterior one truncated. The species some­ Figure 3-6 what resembles Crassalella pauxilla (Yo­ Miyagipeclen malsumoriensis Masuda, 1952, p.252, koyama, 1925; Kanno, 1960; Kanno and pI. 24, figs. 4a-7b; Masuda, 1957, p.31, pI. 4, Chung, 1973), but it differs from the latter by figs. I a-4; Masuda, 1962, p. 226, pI. 27, fig. 13 ; having a more inflated large sheiL However, Masuda, 1977, p.5-6, pI. I, figs. 1-4; Masuda, a more precise comparison between these 1986A, pI. 2, fig. I ; Masuda, 19868, pI. 8, fig. 9 ; Masuda and Noda, 1977, pI. 2, fig. 6; Masuda allied species is prevented due to the poor and Takegawa, 1965, pI. I, fig. II ; O'Hara and condition of the materiaL Ito, 1980, pI. 16, figs. I a -c. Registration number.-NUH 87011, 87012. Remarks.-Only one left valve is at hand. This species is characterized by having a Family Cardiidae large, circular outline and low convex thin Genus Laevicardium Swainson, 1840 sheiL The shell surface is nearly smooth Laevicardillm anguslllm (Yokoyama, 1925) except for fine, concentric growth lines and Figure 3-3a, b radial striations on the umbonal area. The inner surface is smooth without inner rib lets. Cardium anguslUlll Yokoyama, 1925, p. 12, pI. 4, figs. 2a-b. The anterior auricle of left valve is rather Cardium (Laevicardium) anglislUm Yokoyama. smaiL Miyagipecten matsumoriensis is Kuroda, 1931, p. 53, pI. 5, fig. 10. known from Middle and Upper Miocene Laevicardium anguslUm Yokoyama. Kanno, 1960, strata in Northeast Honshu (Masuda, 1952, p.259, pI. 39, figs. 14a-b. 1957,1977; Masuda and Noda, 1977; Remarks.-An internal mold is in the Masuda and Takegawa, 1965; Iwasaki, 1970) collection. This species is characterized by and the Boso Peninsula (O'Hara and Ito, having an extremely high shell compared with 1980). its length (1.5 times the length), a strongly The genus Miyagipeclen has been reported convex and longitudinally ovate outline, from the lower part of the Yakataga F orma­ su bequilaterally concave anterodorsal mar­ tion of southern Alaska by Kanno (1971). gin, strongly convex posterodorsal margin, Geographic distribution.-Northeast Hon­ and prominent umbo. shu and Kanto region. Geographic distribution.-Shigarami For­ Geologic range.-Middle to Late Miocene. mation in Nagano Prefecture, Hatsuse Forma­ Registration number.-NUH 87010. tion in Kanagawa Prefecture, and Chichibu Basin in Saitama Prefecture. Family Crassatellidae Geologic range.-Early Miocene to Early Genus lndocrassatella Chavan, 1952 Pliocene. lndocrassatella cf. tenllilirata (Shuto, 1957) Registration number.-NUH 87013. Figure 3-5, 12 Laevicardillm sp. Compared with.- Crassalelliles lenuiliralUm Shuto, 1957, p. 72-73, pI. Figure 2-6a, b 22, figs. 6-8, 12; O'Hara and Ito, 1980, pI. 16, fig. 5. Remarks.-An incomplete left valve is at hand. This unnamed species has a strongly Remarks.-Two abraded specimens are at abraded shell from the central disk to the hand. This species is characterized by hav­ ventral border, and lacks shell material from ing a subtrapezoidal, moderately inflated thin the umbonal area as shown in Figure 2-6a. shell with regularly rounded concentric ribs. The characteristic features of this species are 932. Molluscan Fauna from the Hatsuse Formation 10 19

Figure 3. Molluscan fossils fro m the Hatsuse Formation (2). I. Pilar kaniei Shikama, X I.I , Loc. A, Reg. No. NUH 870 19. 2. C/inocardium halsusense n. sp. , X 1.5 , Loc. A, Reg. No. NUH 87016. 3a- b. Laevicardium anguslUm (Yokoyama), x l.l , Loc. B, Reg. No. NUH 870 13. 4a- b. Amussiopeclen akiyamae Masuda, X 0.5, Loc. A, Reg. No. N UH 87009. 5, 12. indocrassale//a cf. lenui/iala (Shuto), X 1.1 , 5: Loc. A, Reg. No. NUH 870 12, 12: Loc. A, Reg. No. NUH 870 11. 6. Miyagipeclen malsumoriensis Masuda, x O.7 , Loc. B, Reg. No. NUH 870 10. 7,10. Azorinus abbrevialus (Gould), X I, 7: Loc. B, Reg. No. NUH 8702 1, 10 : Loc. A, Reg. No. NUH 87022. 8. Anisocorbu/a cf. venUSla (Gould), X 1.7, Loc. B, Reg. No. NUH 87023. 9. Oxyperas bernardi (Pi lsbry), X I, Loc. A, Reg. No. NU H 87020. II. Anisocorbu/a sp., X 1.3 , Loc. B, Reg. No. NUH 87024. 1020 Kiyoshi Okumura and Yutaka Yamagishi its medium size and inflated-ovate and rather squarish radial ribs, separated by interspaces thick shell. Hinge apparatus is missing. which are narrower than breadth of radial Registration number.-NUH 87014. ribs; hinge inaccessible; inner margin crenulated corresponding to external radial Genus Clinocardium Keen, 1936 ribs. Clinocardium cf. hataii Hayasaka, 1956 Measurements (in mm.).-

Figure 2-3a, b Specimens Length Height Thickness Valve Type NUH 87016 14.0 10.5 3.7 right Holotypc NUH 870171 13.0 10.0 3.0 right Para type Compared with.- NUH 870172 11.0 8.0 2.5 right ClinocardiunJ hataii Hayasaka, 1956, p. 18, pI. 2, figs. NUH 870173 10.0 7.5 2.5 left 3a-b. Remark.-This new species is character­ Remarks.-The present species was origi­ ized by its quadrilateral shape, which serves nally described by Hayasaka (1956) from the to distinguish it from other allied species. Pliocene of the Futaba district, Fukushima Registration number.-NUH 87016 Prefecture. The characteristic features of this (Holotype); 870171, 870172, 870173 (Par­ species include: shell medium in size, equi­ atype). lateral, suborbicular and moderately inflated with conspicuous beak; anterodorsal margin Clinocardium sp. rounded, merging with rounded ventral bor­ der; posterior and posterodorsal margins Figure 2-7 broadly rounded; surface sculptured with Remarks.-An inner mold is at hand. more than 20 strong radial ribs, but an accu­ This specimen is medium-sized, its width as rate determination of the number of radials is long as high, moderately convex, and equilat­ difficult to obtain because of their poor state eral with prominent umbo. The surface is of preservation; hinge unknown. ornamented with many radial ribs, how many This species somewhat resembles L. is uncertain owing to the inadequate state of shiobarense described from the Kanomata­ preservation. zawa Miocene of Tochigi Prefecture, but the Registration number.-NUH 87018. lesser number of strong radials in the present species serve to distinguish it from L. shiobar­ ense. Family Veneridae Registration number.-NUH 87015. Gen us PitaI' Romer, 1857 PitaI' kaniei Shikama, 1973

Clinocardium hatsusense, n. sp. Figure 3-1

Figure 3-2a, b Pilar kaniei Shikama, 1973, p. 201, pI. 17, figs. 12-13. Description.-Shell small for the genus, Remarks.-The present species was origi­ rather flattened, thin, quadrilateral shape; nally described by Shikama (1973) from the posterodorsal margin nearly straight meeting late Miocene Zushi Formation of the Miura with narrowly rounded posteroventral bor­ Peninsula. The following characters of the der; ventral margin broadly and regularly present species correspond well with Shi­ arched, ascending to anterior margin; antero­ kama's species; shell medium in size and dorsal margin short, making a rather distinct moderately inflated; beak prominent and corner with anterior margin; umbo promi­ turned forward; ventral margin broadly nent; apical angle about lOY; surface or­ arched; posteroventral corner slightly an­ namented with about 50 flat-topped and gulated, anterior margin narrowly rounded; 932. Molluscan Fauna from the Hatsuse Formation 1021 length about 28 mm., and thickness about Geologic range.-Late Miocene to Recent. 6.5- 7.0 mm. Registration number.-NUH 87021, 87022. Geographic distribution.-Miura Peninsula, Kanagawa Prefecture. Family Corbulidae Geologic range.-Late Miocene. Gen us A nisocorblila Iredale, 1930 Registration number.-NUH 87019. Anisocorblila cf. venllsta (Gould, 1861)

Figure 3-8 Family Mactridae Genus Oxyperas March, 1853 Compared with.- Oxyperas bernardi (Pilsbry, 1904) Anisocorblila venslisia (Gould). Hayasaka, 1961, p. 63, pI. 5, figs. 7a-b; pI. 6, figs. 8a-b; Matsu­ Figure 3-9 shima, 1969, pI. II, fig. 14; Kuroda, Habc and Oyama, 1971, p.707, pI. 102, fig. 15; Itoigawa Oxyperas bernardi (Pilsbry). Kuroda, Habe and (in ltoigawa, Shibata and Nishimoto, 1974), p. Oyama, 1971, p.670, pI. 95, figs. 1-2; Matsuura 103, pI. 32, figs. 7a-8b. 1977 pI. 9, fig. 19; O'Hara and Ito, 1980, pI. 16, fig. 6. Remarks.-Two specimens are at hand. Oxyperas aff. bernardi (Pilsbry). Hayasaka, 1973, The present species is characterized by having pI. 6, fig. 4. a solid, slightly inflated, subtrigonal shell Remarks.-An incomplete, strongly abrad­ with a small, weakly pointed beak situated ed right valve missing the anterior quarter of more or less anteriorly, and a blunt ridge the shell is at hand. running from the beak to posteroventral cor­ Geographic distribution of living species: ner. The present species closely resembles South of the 80so Peninsula (about N. lat. Anisocorbula venusta, but these much abrad­ 35') to N. lat. 25' (Kuroda and Habe, 1952), ed specimens do not permit definite and living from littoral depths to 100 m (Ku­ identification with typical A. venllsta. roda, Habe and Oyama). Registration nllmber.-NUH 87023. Geologic range.-Late Miocene to Recent. Registration number.-NUH 87020. A nisocorbula sp. Figure 3-1 I Family Solecurtidae Remarks.-Two poorly preserved speci­ Genus Azorinus Recluz, 1869 mens are at hand. Shells are small in size, Azorinus abbreviatus (Gould, 1861) ovate-trigonal in shape with a blunt ridge Figure 3-7, 10 running from beak to posteroventral corner. Concentric surface sculpture is slightly visible Azorinlls abbreviailis (Gould). Habe and Kosugc, 1967, p. 160, pI. 60, fig. 18; Hayasaka and Ha­ near the ventral margin. The hinge appara­ shimoto, 1970, p. 20, pI. I, figs. 10-11 ; Kuroda, tus is inaccessible. Habe and Oyama, 1971, p. 681, pI. 99, fig. 5. Various species of this genus are distributed in the west Pacific regions south of southern Remarks.-Two abraded specimens are at Hokkaido in littoral to inner neritic waters. hand. The shell is thin, weakly swollen, and Registration number.-NUH 87022. characterized by a slight depression running from the umbo to ventral margin. Living specimens of this species inhabit muddy bot­ References cited toms in waters from 10 to 80 m deep, and are Akamine, H., Iwai, S., Koike, K., Naruse, S., Ogose, distributed south of the 80so Peninsula to the T., Omori, M., Seki, Y., Suzuki, K. and East China Sea, Southeast Asia and Indian Watanabe, K., 1956: Geology of the Miura Ocean. Peninsula. Earlh Sci., no. 30, p.I-8. (in 1022 Kiyoshi Okumura and Yutaka Yamagishi

Japanese with English abstract) the so-called Paleogene Formation in northern Akiyama, M., 1957: Amussiopecten iitomiensis Taiwan. Ibid., vol. 13, p. 91-127, pis. ID-l2. (Otuka) and its allies from Japan. Trans. Proc. Kcen, M., 1963: Marine molluscan genera of west­ Palaeont. Soc. Japan, N.S., no. 25, p. 31-39, pis. ern North America. 126 p. Stanford Univ. 6-7. Press, Stanford, Calif. Habe, T. and Kosuge, S., 1967: Shells. 223 p., 64 pis. Kuroda, T., 1931: Fossil Mollusca. In Honma, Hoikusha, Osaka. (in Japanese) Shinano Chubu Chishitsu-Shi (Geology of Cen­ Hayasaka, S., 1956: Pliocene Mollusaca from the tral Shinano), Part 4, p. 1-90, pis. 1-13. (in Futaba district, Fukushima Prefecture, Japan. Japanese) Saito Ho-on Kai Mus., Res., Bull., no. 25, p. 13- -- and Hahe, H., 1952: Check list and bihliogra­ 20, pI. 2. phy of the Recent marine mollusca of Japan. --, 1961: The Geology and paleontology of the Hosokawa Print. Co., Tokyo, 21D p. Atsumi Peninsula, Aichi Prefecture, Japan. Sci. --, Habe T. and Oyama, K., 1971 : The sea shells of Rep., Tohoku Unil'., 2nd Ser. (Ceol.), vol. 33, no. Sagami Bay. BioI. Lab. Imp. Household. 741 p. I, p. I-IOJ, pis. 1-12. (in Japanese), 489 p. (in English), 121 pis. Maru­ --, 1962: Chevrons of glycymcrid shell. Trans. zcn, Tokyo. Proc. Palaeont. Soc. Japan, N.S., no. 47, p. 291- Makiyama, J., 1927: Molluscan fauna of the lower 297, pI. 45. part of the Kakegawa Series in the province of --, 1973: Pliocene marine fauna from the Tanc­ Totomi, Japan. Mem. Coli. Sci., Kyoto Imp. ga-Shima, South Kyushu. Sci. Rep., Tohoku Univ., Ser. B., vol. 3, no. I, art. I, p. 1-1479, pis. Univ., 2nd Ser. (Ceol.), Spec. Vol., no. 6, p.97- 1-6. 108, pis. 6-7. --, 1931 : Stratigraphy of the Kakegawa Pliocene -- and Hashimoto, S., 1970: Discovery of marine in Totomi. Ibid., vol. 7, no. I, art. I, p. I-53, pis. fossil from immediately bclow thc Ata pyroclas­ 1-3. tic flow deposits, Onejime-Cho, Osumi Peninsula, --, 1952: On the mutation of the fossil C(l'cymeris Kagoshima Prefecture. Rep., Fac. Sci., K ago­ rOll/nda. Trans. Proc. Palaeonl. Soc. Japan, N. shima Univ., no. 3, p. 17-22, pI. l. S., no. 5, p. 131-138. Itoigawa, J., Shibata, H. and Nishimoto, H., 1974: Masuda, K., 1952: On some Miocene Pectinidae from Molluscan fossils of the Miocene Mizunami environs of Sendai. Ibid., no.8, p.249-254, Group, Central Japan. Bull., Mizunami Fossil pI. l. Mus., no. 2, p.43-203, pis. 1-63. (in Japanese --, 1957: A note on Miyagipecten matsumoriensis with English abstracl) Masuda. Saito Ho-on Kai Mus., Res. Bull., no. Iwasaki. Y., 1970: The Shiobara-type Molluscan 26, p. 31-39, pI. 4. Fauna. An ecological analysis of fossil Mollus­ --,1962: Tertiary Pectinidae of Japan. Sci. Rep., cas. Jour. Fac. Sci., Univ. Tokyo, Sec. 2, vol. Tohoku Univ., 2nd Ser. (Ceol.), vol. 33, no. 2, p. 17, pt. 3, p. 351-444, pis. 1-7, 25 figs. 117-238, pis. 18-27. Kamada, Y., 1962: Tertiary marine mollusca from --, 1977: Miocene molluscs from the Shimokuro­ the Joban coalficld, Japan. Palaeont. Soc. sawa Formation, Ichinoseki City, Iwate Prefec­ Japan, Spec. Papers, no. 8, p. 1-187, pis. 1-2l. ture, Northeast Honshu, Japan. Saito Ho-on -- and Hayasaka, S., 1959: Remarks on a fossil K ai Mus., Res. Bull., no. 45, p. 3-10, pI. l. marine fauna from Tateishi, Futaba district, in --, 1986A: Historical change of the Neogene the Joban coalfield, Fukushima Prefecture. mollusca fauna of Japan with special refercnce to Saito Ho-on Kai Mus., Res. Bull., no. 28, p. 17- pectinids. Monogr. Mizunami Foss. Mus., no. 6, 28, pI. 2. p. 1-21, pis. 1-3. (in Japanese with English Kanno, S., 1960: The TertiGlJI System of the Chi­ abstract) chibu basin, Saitama Prefecture, central Japan. --, 1986B: Notes on origin and migration of Part II. Paleontology. p. 123-396, pls.31-5l. Cenozoic pectinids in the North Pacific. Japan Soc. Promot. Sci., Tokyo. Palaeonl. Soc. Japan, Spec. Papers, no. 29, p. --,1971: Tertiary molluscan fauna from the Yaka­ 95-110, pis. 7-10. taga district and adjacent area of southern Alas­ -- and Noda, H., 1977: Remarks on the Miocene ka. Palaeont. Soc. Japan, Spec. Papers, no. 16, marine fauna from Watarimachi. Miyagi Prefcc­ p. 1-154, pis. 1-18. ture, Japan. Saito Ho-on Kai Mus., Res. Bull., -- and Chang, L. S., 1971: Amussiopecten and its no. 45, pI. 13-16, pI. I. associate fauna from northern Taiwan. Ceol. -- and Takegawa, 1965: Remarks on thc Miocene Palaeol1l. Southeast Asia, vol. 12, p. 189-206, mollusca from thc Scnnan district, Miyagi Prefec­ pis. 30-3l. ture, Northeast Honsh u, Japan. Ibid., no. 35, p. -- and Chung, C. T., 1973: Molluscan fauna from 1-14, pis. 1-2, I text-fig. 932. Molluscan Fauna from the Halsuse Formation 1023

Matsukawa, A., 1986: Cenozoic Glycymerid bivalves Vnir. Tokyo, vol. 12, p. 566-638. of Japan. Palaeont. Soc. Japan, Spec. Papers, --, 1934: On some fossils from the northern foot of no. 29, p. 77-94, pis. 5-6. Mt. Minobu and the Hayakawa tutlite of Hakone. Matsushima, Y., 1969: On the molluscan fossils from Jour. Geol. Soc. Japan, vol. 41, no. 492, p.562- the Alluvial deposits in Yokohama City. Bull. 568. (in Japanese) Kanagawa Pref Mus. no. 2, p. 79-96, pis. 10-13. Ozaki, H., 1954: On the palaeontology of the basal (in Japanese with English abstract) conglomerate of Pliocene, in Tyosi City, Kanto Matsuura, N., 1977: Molluscan fossils from the late Region. Bull. Natn. Sci. Mus .. Tokyo. N.S., Pleistocene marine terrace deposits of Hokuriku vol. I, no. I (no. 34), p. 9-21, pI. 1-10. region, Japan Sea side of central Japan. Sci. Shikama, T., 1973: Moll uscan assemblages of the Rep.. Kanazawa Vniv., vol. 22, no. I, p. 117-162, basal part of the Zushi Formation in Miura pis. 1-20. Peninsula. Sci. Rep.• Tohoku Vnil' .• 2nd Ser. Mitsunashi, T. and Yazaki,K., 1968: Geologic map (Geol.), Spec. Vol., no. 6, p. 179-204, pis. 16-17. of the Miura Peninsula, Scale I: 25,000. Shuto, T., 1957: Crasatellites and Venericardia from Geological Survey of Japan. the Miyazaki Group (Paleontological study of -- et al., 1976: Geological map of Tokyo Bay and the Miyazaki Group). Mem. Fac. Sci.• Kyushu adjacent area, Scale I: 100,000. Geological Vnil' .. Ser. D, vol. 6, no. 2, p. 69-90, pI. 22. Survey of Japan. Suzuki, K., 1932: Pliocene formation north of O'Hara, S. and Ito, M., 1980: Molluscan fossils from Atsugi-Machi, Kanagawa Prefecture (II). Jour. the Senhata Formation in Boso Peninsula. Geol. Soc. Japan, vol. 39, no. 462, p. 97-13 I. (in Prof S. Kanno Memorial Vol., p. 121-136, pis. Japanese) 14-17. Yokoyama, M., 1920: Fossils from the Miura Penin­ Okumura, K., 1988: Molluscan f()ssils from the Pleis­ sula and its immediate north. Jour. Coli. Sci.. tocene Ninomiya Group in the Oiso Hills, Imp. Vnir. Tokyo., vol. 39, art. 6, p. 1-193, pis. 1- Kanagawa Prefecture, central Japan. Cullllral 20. Properties Rep.. Ninomiya Town Educ. Com­ --,1922: Fossils from the Upper Musashino of the mitee, no. 16, p. 1-99. (in Japanese with English Kazusa and Shimosa. Ibid., vol. 59, art. I, p. 1- abstract) 200, pis. 1-17. Omori, M., Hayama, K. and Horiguchi, M., 1986: --, 1925: Mollusca from the Tertiary Basin of Kanto region (Geology of Japan. no. 3). 335 p. Chichibu. Jour. Fac. Sci.• Imp. Vnil'. Tokyo. Kyoritsu Publ. Co., Tokyo. (in Japanese) Sec. 2, vol. I, pt. 3, p. 111-126, pis. 14-15. Otuka, Y., 1934: Tertiary structures of the northwest­ --, 1926: Tertiary mollusca from Shiobara in ern end of the Kitakami Mountainland, Iwate Shimotsuke. Ibid, vol. I, pt. 4, p. 127-136, pis. Prefecture, Japan. Bull. Earthquake Res. Inst .. 16-20.

Boso m1$., Kami-miyada J·.'8EE, Kaneda ~EE. Kurihama ;7.JI.U~, Kurosaki-no-hana ~ ~ /14, Misakiguchi -=~ I I, Mito -=P. Miura .=:00. Miurakaigan -:':iflji'liH¥, Nagahama ffi:i%, Sagami t§t!i'!. Wakamiya %'8.

-=rffi~!,:!1'b~tmt}]F~il€l/i,kff.:iM?o{r.Em:: Ij~J~il€l\iJ\1*iJJ¥J.l{r.EO) 19 t~~~5J1j L. :?CO) 5 ~. g~.3g, ~~.IM~~a~.15~(5~I~g~~~)0)~.~ff0kom~ O)~!~(;, fiJj:JJJ(:glt.""t"'"' -c ll£i'lJ.El=.g""(-;I; Q -=- C, neritic 1:,~,M C outer neritic 1:,~,g 0)i1t.g­ Ie::. ~ 0 -c -=- 0) {r.ElWt;;fiJj:IJJ(: ~.h -c l' Q -=- C, t t.:, 4'!fu>, 0 -=- O)lW~ft.!=r~tJl:f~MO) {, 0)""(-;1; Q -=- !.: ~§Jj 07')'1'::' Lt.:o lJJi!:H il'f· IUi¥ f1Z Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1024-1039, 8 Figs., April 30,1992

933. INTRASPECIFIC VARIATION AND HETEROCHRONY OF PHANEROLEPIDA PSEUDOTRANSENNA OZAKI (GASTROPODA: TURBINIDAE) FROM THE PLIOCENE NOBORI FORMATION, PACIFIC SIDE OF SOUTHWESTERN JAPAN*

RYUICHI MAJIMA

Geological Institute, Yokohama National University, Yokohama, 240

and

AKIHIRO MURATA

Institute of Geosciences, Shizuoka University, Shizuoka, 422

Abstract. The recent discovery of 79 specimens of Phanerolepida pseudotransenna Ozaki from the Pliocene Nobori Formation indicates that the species shows a very wide range of individual variation in its surface ornamentation. This variation is interpreted to be caused by differences in timing of the onset of ornamental development among individuals in the population. An evolutionary lineage, Nehalemia hieroglyphica --> Phanerolepida oregonensis --> P. pseudotransenna --> P. transenna, is established on the basis of their stratigraphic occurrences, and by comparison of ontogenetic developments of shell ornamen­ tation. At least the first two speciation events are considered to be the results of hetero­ chrony. Phanerolepida rehderi MacNeil is designated as a junior synonym of P. pseudotransen­ na. Four opercula are reported and illustrated in association with their own shells, near to their life position.

Key words. Phanerolepida, intraspecific vanatlOn, heterochrony, Gastropoda, Pliocene, Nobori Formation, southwestern Japan.

Introduction North America (Hickman, 1972) to its later occurrences which have been restricted to the Phanerolepida is a very interesting genus Japanese Neogene and Recent faunas, are of from the malacological, conchological and special interest. paleontological viewpoints. Ecologically it In Japan, the following five species of is a homalopomatine, exceptionally living in Phaneroleida have previously been known: the very deep sea, and morphologically it Phaneroleida transenna (Watson, 1879), bears a peculiar rhombohedral network pat­ Pliocene (Otuka, 1949; Okutani, 1968)­ tern of ornamentation. The evolution and Recent (Watson, 1879 and 1886; Dall, 1907; migration of Phanerolepida, from its first Ok utani, 1964, 1966, 1968, 1969; N oda and appearance in the early Oligocene of western Ogasawara, 1976); P. expansilabrum Kuroda, 1931, Miocene 'Received September 17, 1991; revised manuscript (Kuroda, 1931; Watanabe et al., 1950; accepted December 20, 1991 Shibata, 1974 [as P. cf. expansilabrumJ;

1024 933. Intraspecific variation and heterochrony of Phanerolepida 1025

Noda and Kikuchi, 1980; Itoigawa el al., Abbrevi a tions 198 I, 1982 [as P. cf. expansilabrllmJ) ; P. pselldotransenna Ozaki, 1956, ? Miocene The following abbreviations are used in (Hatai and Masuda, 1962) - Pliocene (Ozaki, this paper: 1956; N oda and Ogasawara, 1976; Mimoto, IGSU, Institute of Geosciences, Shizuoka 1986). University, Shizuoka, 422 Japan; and P. rehderi MacNeil, 1961, Pliocene (Mac­ NSM, National Science Museum, Shinzyuku­ Neil, 196 I ; N oda and Ogasawara, 1976; ku, Tokyo, 169 Japan. Noda, 1980, 1988); and P. okinawana Noda, 1988, Pliocene (Noda, Collecting localities 1988). Their morphological comparison and tax­ Specimens of Phanerolepida pseudotran­ onomic validity have not been well senna examined for this study were collected documented because some of them were from the massive mudstone of the Pliocene proposed on the basis of poorly preserved Nobori Formation of the Tonohama Group, material in which the type material of P. exposed in a quarry at Nobori, Hanemachi, pselldotransenna is included. From summer Muroto City, Kochi Prefecture, on the Pacific of 1987 to April of 1990, we collected 79 side of southwestern Japan (Figure I). Our specimens of P. pselldotransenna, including three localities in the quarry from which the many that are well preserved. The aim of species was collected are as follows: this study is to describe P. pseudotransenna IGSU loc. no. 62-35- I : a cliff located in in detail, to show the mode of individual southern end of the quarry, consisting of ontogenetic variation, to explain the origin of massive, pale gray mudstone intercalated with this variation, and to evaluate the role of lenticular molluscan shell beds about IO cm heterochrony in the evolution of the genus. in maximum thickness and over 6 m in maxi-

SEA OF JAPAN

PACIFIC OCEAN

Figure 1. Index map showing the fossil localities of Phanerolepida pseudotransenna Ozaki from the upper Pliocene Nobori Formation. The fossil localities are plotted on a I : 25,OOO-scale topographic map of Japan, Quadrangle "Hane," Geographical Survey of Japan. 1026 Ry uichi Majima and Akihiro Murala

Figure 2. A view of the quarry cliff ( I) a nd the mode of occurrence of Phanerolepida pseudotransenna Ozaki (2 - 4) at IGSU loc. no. 62 - 35 - 3. mum width. The matrix of the lenticul ar their life position (Figure 8). shell beds is composed of mudstone. The massive mudstone of the three local­ IGSU loc. no. 62- 35 - 2: a cliff, si tuated iti es sporadicall y yields the following deep­ near the center of the quarry, consistin g of water species: Tenuileda ikebei (Suzuki and massive, pale brownish gray, weak ly weather­ Kanehara, 1936), Portlandia sp., Limopsis ed mudstone. tajimae Sowerby, 1914, L. chilaniana Yoko­ IGSU loc. no. 62- 35- 3: a large cliff (Fig­ yama, 1926, Glycymeris nipponica (Yoko­ ure 2- 1) exposed in northern end of the yama, 1920), Acesla golialh (Sowerby, 1883), quarry consisting of massive, pale gray mud­ Deleclopeclen peckhami (Gabb, 1869), stone. Nemocardium sp., Cuspidaria hirasei K uro­ Except fo r the lenticul ar shell beds at IGSU da, 1948, Fissidentalium yokoyamai (Maki­ loc. no. 62- 35- 1, the moll uscan fo ssi Is from yama, 193 1), Balhybembix aeola (Watson, these localities are considered to have been 1878), Ginebis argenteonitens (Lischke, 1872), preserved in SiIU, because: ( I) they occur Phanero/epida pseudolransenna Ozaki , 1956, sporadicall y in massive mudstone (Figure 2- Aula cofusus cerulescens Kuroda a nd Habe, 2 - 4) ; (2) many bi valves are preserved in 1966, Neplunea noboriensis Ozaki, 1956, articulated condition; and (3) four speci­ Turrancilla sua vis (Yokoyama, 1926), Ful­ mens of Phanerolepida pseudolransenna bear goraria (Musashia) cancellala Kuroda a nd o percula within their own apertures near to Habe, 1950, F. (Saolomea) delicata (Fulton, 933. Intraspecific variation and heterochrony of Phanerolepida 1027

1940), and Makiyamaia coreanica (Adams hyugensis (Shuto, 1964), G. hagenoshitensis and Reeve, 1849). (Shuto, 1964), Mammilla sp., Oyptonatica The lenticular shell beds at IGSU loc. no. andoi (Nomura, 1935), C. adamsiana (Dun­ 62- 35 - 1 bear the species A nadara (Tosarca) ker, 1859), Babylonia ela/a (Yokoyama, tosaensis (N oda, 1965), Glycyme ris cf. yesso­ 1923) , Siphonalia tonohamaensis N omura, ensis (Sowerby, 1886), Chlamys satoi (Yoko­ 1937, S. y abei Nomura, 1937, Baryspira al­ yama, 1928), Chlamys sp., Cryptopecten bocallosa (Lischke, 1873), Oliva mustellina vesiculosus (Dunker, 1877), Amussiopecten (Yokoyama, 19 23), Granulifusus dualis (Yo­ praesignis (Yokoyama, 1922), Venericardia koyama, 1928), Conus sp., Orthosurcula per­ panda (Yokoyama, 1926), Dimya sp., Cras­ virgo (Yokoyama, 1928), Spirotropis subde­ satellites sp., Paphia sp., Placamen tiara (Dil­ c!ivis (Yokoyama, 1926), and A rchitectonica lwyn, 18 17), Dentalium sp., Umbonium (Su­ sp. Because these molluscs are composed of chium) suchiense Yokoyama, 192 3, Phanero­ both sha ll ow a nd deep water species, they are lepida pseudotransenna Ozaki, 1956, Turri/el­ considered to have been more or less rewor­ la perterebra Yokoyama, 1923, Glossaulax ked .

..y 1a (SRFsRFsFRFsF) (SRF) (SRF)

Figure 3. Phanero/epic/a pseuc/olransenna Ozaki. la - 3b, IGS U loco no. 62- 35 - 1, X 1.7 : la - lb, IGSU M37 ; 2a-2b, IGSU M3 8; 3a - 3b, IGSU M39. Letters around shel l indicate successive ornament patterns on body whorl. 4, 5, IGSU loc. no. 62- 35 - 3, X 2.6: 4, IGSU M60; 5, IGSU M64. Arrows R, F, and s indicate the extent of the respective patterns on body whorl. 1028 Ryuichi Majima and Akihiro Murata

Table 1. Comparison for letter formula between Shinzato and Nobori specimens. See text for details.

SHINZA TO FORMATION NOBORI FORMATION OKINAWA (NODA AND OGASAWARA, 1976) (IGSU LOC, NO. 62-35-3) 43 SPECIMENS 25 SPECIMENS LETTER FORMULA NUMBER OF LETTER FORMULA NUMBER OF SPECIMENS SPECIMENS

SRF 10 SRF 11 SRFsF 5 SRFsF 2 SsF SRFs 21 SRFs SRs 2 SsFs 1 Ss 3 ?Ss SR SRFRF 2 SRFsRs SRFRFRF SRFRFRFs SRFRFRFRF SRFsRFsRF SRFsFRFRF (A) SRFRFRFsRFs LETTER FORMULA LETTER FORMULA SRF SRF SRF(s)F -+ SRFRF SRF(s)F -+ SRFRF S(s)F-+SRF SRF(s)-+SRFR SRF(s)-+SRFR SR(s)-+SR S(s)F(s)-+SRFR S(s)-+SR ?S(s)-+ ?SR SR SRFRF SRF(s)R(s)-+SRFR SRFRFRF SRFRFRF(s)-+ SRFRFRFR SRFRFRFRF SRF(s)RF(s)RF -+SRFRFRF SRF(s)FRFRF -+ SRFRFRFRF (B) SRFRFRF(s)RF(s)-+SRFRFRFRFR LETTER FORMULA NUMBER OF LETTER FORMULA NUMBERS OF SPECIMENS SPECIMENS SR 5 SR (1 +?1) SRF 11 SRF 11 SRFR 22 SRFR 2 SRFRF 5 SRFRF 4 SRFRFRF 2 SRFRFRFR SRFRFRFRF 2 (C) SRFRFRFRFR 933. Intraspecific variation and heterochrony of Phanerolepida 1029

Intraspecific variation of Phanerolepida A): SRF (10 specimens), SRFsF (5 speci­ pseudotransenna Ozaki mens), SsF (I specimen), SRFs (21 speci­ mens), SRs (2 specimens), SsFs (I specimen), Ornamentation of the teleoconch of and Ss (3 specimens). Phanerolepida pseudotransenna is divided We observed 79 specimens (see Appendix) into four patterns: three or four spiral cords from the Nobori Formation, of which we on the smooth surface (S in abbreviation; analyzed 25 well preserved specimens from Figure 3-4-5 [arrow SJ), a roughly devel­ the IGSU loc. no. 62-35-3, where P. oped rhombohedral network (R; Figure 3- pseudotransenna occurs autochthonously and 4-5 [arrow RJ), a finely developed rhom­ most abundantly. Among the 25 specimens bohedral network (F; Figure 3-4 - 5 [arrow (Table I-A, Figures 4,5), 11 specimens are FJ), and a smooth surface occasionally classifiable into Type SRF (Figure 5-9a­ bearing very fine spiral striae (s; Figure 3-5 lOb), 2 specimens into SRFsF (Figure 5-8a- [arrow sJ). The pattern boundaries are com­ 8b), I specimen into SRFs, and I specimen monly sharply separated from each other, but into a questionable Ss (Figure 5-12a-12b). the pattern R, when R immediately follows The remaining 10 specimens bear ornamenta­ the pattern S, is sometimes superposed by tion that is not classifiable into any of the weak spiral cords that continue from the types of Noda and Ogasawara (1976) (Table spiral cords of the pattern S (Figure 3-la) I-A). They consist of the Type SR (I speci­ and the R is sometimes gradually changed men; Figure 5-11 a - II b), SRFRF (2 speci­ into the s (see boundary between the Rand s mens; Figure 5-6a - 6b), SRFsRs (I speci­ in Figure 3-5). In the following discussion, men; Figure 5-7a - 7b), SRFRFRF (I speci­ we use a letter formula for the description of men; Figure 5-5a-5b), SRFRFRFs (I speci­ the shell ornamentation for respective indi­ men; Figures 3-4,5-4a-4b), SRFRFRFRF viduals. The above four abbreviations of the (I specimen; Figure 5-1 a - I b), SRFsRFsRF pattern are arranged ontogenetically. For (I specimen), SRFsFRFRF (I specimen; example, when the teleoconch is sculptured Figure5-3a-3b), and SRFRFRFsRFs (1 from earlier to later with S -> R -> F -> s, specimen; Figures 3-5, 5-2a-2b). The then we use the letter formula "SRFs." ranges of individual variation of the shell­ Noda and Ogasawara (1976) showed the surface ornamentation of the Shinzato and individual variety of the shell ornamentation Nobori specimens are quite different (Table of Phanerolepida rehderi MacNeil, 1961, here I-A). considered to be a junior synonym of P. We propose herein a sorting scheme for the pseudotransenna Ozaki, 1956, by using mate­ surface ornamentation. As noted above, the rial from the upper Pliocene Shinzato Forma­ pattern R sometimes becomes coarser and tion, Okinawa Prefecture, southern Japan. changes gradually into the pattern s (Figure Noda and Ogasawara's (1976) work is sum­ 3-5). Thus, the pattern s is considered to be marized as follows. the coarsest part of the pattern R. In the Surface ornamentation of teleoconch following discussion of pattern arrangement changes ontogenetically. The first, second, the s is replaced by R (Table 1-8). and rarely third teleoconchs have the pattern In this sorting scheme (Table 1-8), the S, and its later teleoconchs are sculptured three patterns S, R, and F are regularly arran­ with R, F, and/or s. The degree of develop­ ged in both the Shinzato and Nobori speci­ ment of these four patterns (S, R, F, and s) is mens: that is, the teleoconch ornamentation greatly variable in individuals. The 43 Shin­ always begins with the pattern S; the S is zato specimens were classified into the follow­ always followed by the pattern R; the R is ing 7 different development types (Table 1- always succeeded by the pattern F; and the R 1030 Ryuichi Majima and Akihiro Murata

SPECIMEN VOLUTION LETTER FORMULA LATE (IGSU·M) EARLY 2 2 I I n SRFRFRFRF 64 SRFRFRFsRFs 69 SRFsRFsRF 72 SRFsFRFRF 6~ SRFRFRFs m SRFRFRF 65 SRFRF 74 SRFRF 75 SRFsRs 97 SRF 77 SRF H SRFsF LEGEND 91 SRF 71 SRF D1 63 SRF 02 59 SRF 76 SRFsF rn3 94 SRF ~4 66 SRF 70 SRF U5 6l SRF ~6 67 SRF 83 I7 SRFs -,- 68 SR ~8 933. Intraspecific variation and heterochrony of Phanerolepida 1031

and F appear repeatedly in some specimens M60, MIOO and M65 in Figure 4) are very (Table I-C). closely arranged in comparison with the These differences among individual orna­ others. We consider these close arrange­ mental patterns are not considered to be ments to be caused by slow growth rate due controlled by age differences because there is either to the later adult stage, worse trophic no relationship between number of whorls conditions, or reproductive activity. and ornament type (Figure 4; Noda and Ogasawara, 1976, fig. 3). Evolution of Phanerolepida We propose a model to explain the different ranges of individual variation We hypothesize that the Recent Phanero­ between the Shinzato and Nobori specimens. lepida transenna evolved from Eocene Ne­ This model maintains that the differences halemia hieroglyphica through Oligocene P. among these individual ornamental types are oregonensis and Pliocene P. pseudotransenna. controlled by the differences in timing of the In the speciation events represented, at least onset of ornamental development in individ­ two events are considered to have been the uals and that the onset timing of respective results of heterochrony, as discussed below. individuals differs successively (Figure 6-1). Phanerolepida is considered to have In Figure 6-1, if the onset timing is extremely evolved from the western North American accelerated, then an SRFRFRFRF-type orna­ species Nehalemia hieroglyphica Hickman, mentation results, while if the onset timing is 1974. Nehalemia hieroglyphica Hickman greatly delayed, then an SR-type ornamenta­ shares characteristics of both Phanerolepida tion appears. In this model, the range of and Homalopoma and occurs near the top of individual variation on shell ornamentation the Eocene Cowlitz Formation, approximate­ is primarily controlled by the range of varia­ ly 450 m stratigraphically beneath the oldest tion in timing of the onset of the ornament occurrence of Phanerolepida, P. oregonensis development (Figure 6-1). The Nobori Hickman, in the lower Oligocene Keasey specimens are, therefore, considered to have a Formation (Hickman, 1974). On the wider range in timing of the onset of orna­ teleoconch surface, N. hieroglyphica is or­ ment development than the Shinzato speci­ namented with strong spiral ribs separated by mens. This mode of individual variation approximately equal interspaces (Hickman, controlled by such a mechanism has been 1974). This ornamentation is considered to reported by Majima (1985, 1988), who studied be comparable to the pattern S of P. the heterochronic evolution of three naticid pseudotransenna (Figure 6-3). gastropods. Phanerolepida oregonensis is considered to The patterns Rand F in the later adult have evolved from N. hieroglyphica, and has a stage of some specimens (lGSU M69, M72, net-like, coarse rhombohedral pattern of sur-

+- Figure 4. Individual variation of ornamentation on shells of Phanerolepida pseudotransenna Ozaki at IGSU loc. no. 62-35-3. Respective lateral columns illustrate successive ornamental patterns. The "0" position of the volution coincides with the boundary between the patterns Rand S but not with protoconch/ teleoconch boundary. In many specimens, the protoconch/teleoconch boundary is indistinct due to weathering. Legend: I, protoconch or pattern S (three or four spiral cords on the smooth surface) indetermi­ nable due to weathering; 2, protoconch; 3, pattern of three or four spiral cords; 4, roughly developed rhombohedral network pattern; 5, finely developed rhombohedral network pattern; 6, smooth surface occasionally bearing very fine spiral striae; 7, sculptural boundary preserved; 8, sculptural boundary or early conchs indeterminable due to weathering. IGSU M86 specimen (Figure 5-12a-b; Table I-A, ? Ss) is not illustrated in this figure because of its imperfect preservation of ornamentation. So the total number of specimens in this figure is 24, but not 25 mentioned in the text. 103 2 Ryuichi Majima and Akihiro Murata

F

~ " 2a / (SRFRFRFs) 4a (SRFRFRFRF) (SRFRFRFsRFs) (SRFsFRFRF)

2b " S

(SRFRF) (SRFsRs) (SRFsF) /'

7b

8b

11a (SRF) (SRF) (SR) (5?s?s) 933. Intraspecific variation and heterochrony of Phanerolepida 1033 face ornamentation, and its early teleoconchs can be stated. The faint spiral sculpture of have a faint spiral sculpture superposed on P. transenna may be indistinct in some speci­ the rhombohedral network (Hickman, 1972) mens, and the fine rhombohedral network (Figure 6-2). The coarse rhombohedral net­ pattern is commonly widely developed, finer work pattern is considered to be comparable in grain size than the pattern F of P. to the pattern R of P. pseudotransenna, and pseudotransenna, and may be interrupted by the superposed faint spirals are probably the coarse rhombohedral network pattern. comparable with the spiral ribs of N. hierogly­ The evolutionary lineage Nehalemia hiero­ phica, and also with the pattern S of P. glyphica (ornament pattern: S) --> P. pseudotransenna. oregonensis (S + R --> R) --> P. pseudotransen­ Phanerolepida pseudotransenna is inferred na (S --> R <- variation --> S --> R --> F --> R to have evolved from P. oregonensis in a --> F --> R --> F --> R --> F --> R) is considered migration process from the eastern Pacific to to be an example of peramorphosis (Figure the western Pacific during the late Oligocene 6). Peramorphosis is the occurrence of the or early Miocene. The former species shows ancestral adult morphology in a descendant a very wide range of individual variation in juvenile stage of development (McNamara, ornamentation and has a variant bearing 1986). A highly variable onset timing of SR-type ornamentation which is interpreted ornament development in individuals occurs as a relic of P. oregonensis. In some speci­ in the evolutionary process from P. oregonen­ mens of P. pseudotransenna, the pattern R sis to P. pseudotransenna. Although the that immediately follows the pattern S is wide range of individual ontogenetic varia­ superposed by weak spiral cords that con­ tion in P. pseudotransenna makes interpreta­ tin ue from the spiral cords of the pattern S tion of the heterochrony complicated, this (Figure 3-la), just as with the early teleocon­ evolution is evidently a peramorphosis chs of P. oregonensis. This similarity sup­ because many specimens of P. pseudotransen­ ports the above interpretation for the lineage. na have the ancestral patterns Sand R in Phanerolepida transenna is considered to their younger stages. have evolved from P. pseudotransenna, and commonly shows an ontogenetic ornamental sequence on the teleoconch, i.e., faint spiral sculpture --> coarse rhombohedral network Systematic Paleontology pattern --> fine rhombohedral network pat­ Class Gastropoda Cuvier, 1797 tern, which are considered to be comparable Subclass Prosobranchia Milne-Edwards, 1848 with the patterns S, R, and F of P. Order Archaeogastropoda Thiele, 1925 pseudotransenna, respectively. The deriva­ Family Turbinidae Rafinesque, 1815 tion of P. transenna from P. pseudotransenna Subfamily Homalopomatinae Keen, 1960 is supported by evidence that P. pseudotran­ senna has a variant bearing SRF-type orna­ mentation that is considered to be a fore­ Gen us Phanerolepida Dall, 1907 runner of P. transenna. The precise individ­ Type species. - Turbo transenna Watson, ual variation of the ornamentation of P. 1879, by monotypy. Holocene, Pacific side transenna is not well known because of the of central and southwestern Japan (Noda and lack of available specimens, but the following Ogasawara, 1976).

~ Figure S. Phanero/epida pseudotransenna Ozaki. IGSU loco no. 62-35-3. X 1.7. la-lb, IGSU M73; 2a-2b, IGSU M64; 3a-3b, IGSU M72; 4a-4b, IGSU M60; 5a-5b, IGSU MIOO; 6a-6b, IGSU M74; 7a-7b, IGSU M75; 8a-8b, IGSU M61; 9a-9b, IGSU M94; lOa-lOb, IGSU M67; lla-llb, IGSU M68; 12a - 12b, IGSU M86. Letters around the shell indicate the successive patterns on body whorl. 1034 Ryuichi Majima and Akihiro Murata

------Ontogeny------Early whorl ~.. ------late whorl t

::0 :::>'" "'"'" ~ :::> i :c.=- -='" g: '" ~ !!! i' §: I gO

-'"~ o

Phanerolepida pseudotransenna PERAMORPHOSIS..

i? XXXXXXXXXXXXXJ• 2 (S+R) ... (R) PhanerolepidC! PERAMORPHOSIS• oregonensis ? 3 (S) Nehalemia hieroglyphica Figure 6. Schematic illustration of a model for the explanation of individual variation of Phanerolepida pseudotransenna Ozaki (I), and a heterochrony of Nehalemia hieroglyphica Hickman (3) ---> Phanerolepida oregonensis Hickman (2) ---> P. pseudotransenna Ozaki (I) lineage. Note that the difference of individual varia­ tion between Shinzato and Nobori specimens is interpreted to be caused by variation in the onset timing of ornament development for each individual.

Phanerolepida pseudotransenna Mimoto, 1986, p. 36, pI. 15, figs.5a-6b. Ozaki, 1956 ? Phanerolepida pseudotransenna Ozaki. Hatai and Masuda, 1962, pI. 40, figs. 26, 27. Figures 3,5, 7, 8 Phanerolepida rehderi MacNeil, 1960, p. 30, pI. 7, figs. 6-8, 11-13; Hickman, 1972, figs. 3,7, II, 13; Phanerolepida pseudotransenna Ozaki, 1956, p. 1-2, Noda and Ogasawara, 1976, pI. I, figs. la-lOb; pI. I, figs. 1-2; Noda and Ogasawara, 1976, pI. I, Masuda and Noda, 1976, p. 246; Noda, 1980, p. fig. 13; Masuda and Noda, 1976, p. 246 ; II, pI. 6, fig. I ; Noda and Kikuchi, 1980, pI. I, 933. Intraspecific variation and heterochrony of Phanerolepida 1035

Figure 7. Four views of the ho lotype of Phanero/epida pseudolransenna Oza ki. NSM PI 4368. x 1. 8. Nisinotani, Nobori, Ha ne- mach i, Mu­ roto C ity, Kochi Prefecture, Pacifi c side of so uthwest­ ern Ja pa n. Upper Pli ocene No bori Formation. Figure 8. Opercula of Phan ero/epida pseudOiransenna Ozaki. IGSU loc. no. 62 - 35 - 3. X 1.7. 1, 3 - 5, opercul a preserved in the aperture near to thei r life positi o n. 2, operculum occurring iso­ fi gs. 2a, 2b, 13, ?4 [ figs . 3, 4, showing opercul a lated from its own shell . I, IGSU M73; 2, IGSU isolated from the shell ] ; Noda, 1988, p. 34, pI. 5, M117 ; 3, IGSU M112 ; 4, IGSU M69 ; 5, IGSU figs. I a- 3c. Mill. Phanero/epida Iransenna (Wa tson). Aoki and Baba, 198 4, p. 73 , figs . I a, I b [ not Ph anero/epida lran­ Pliocene species of Okinawa, so uthern Japan, senna (Watson, 1879)]. is here designated as a junior syno nym of the present species for the first time. The two Type malerial.- NSM PI 4368 (ho lotype: species are identica l to each other in shell Figure 7) from Nisino ta ni , N o bo ri , Ha ne­ form. Although the two species have machi (Ozaki, 1956), Muro to City, Kochi different ra nges of individua l o rn amenta l Prefecture, Pacific side of southwestern Japa n. variations (Ta ble I- A), these differences are Pliocene Nobori Formation of the Tono­ interpreted as intraspecific variatio n resulting ham a Group. fr o m the variational ra nge of onset timing o f Stratigraphic occurrence. - ? Miocene: T o­ o rnament develo pment (Figure 6- 1) . kigawa Formatio n, Saitama Prefecture (H atai Phanerolepida expansilabrum Kuroda, a nd Masuda, 1962 ). Pliocene: N o bo ri For­ 1931 from the Miocene o f J a pa n is intermedi­ mation, Kochi Prefecture ; Shinzato Fo rma­ ate in age between P. oregonensis a nd P. tion, Okinawa Prefecture (MacNeil , 1960; pseudolransenna. Because P. expansilabrum N o d a and Ogasawara, 1976 ; N oda, 1980, has previously been know n o nl y by poorly 1988 ). preserved specimens, specific comparisons Remarks. - We collected 5 opercula of P. among the three species must await availabil­ pseudolransenna from IGSU loco no. 62 - 35 - ity of numero us well preserved specimens o f 3, of which 4 are preserved in their own the Mi ocene species. apertures near to their life position (Figure 8- 1, 8- 3 - 5). These o percula are indistinguish­ Acknowledgments able from those of the Recent species, P. transenna. We would like to tha nk Prof. H . N oda, Phanerolepida rehderi MacNeil, 1960, a University o f T sukuba, for his kind di sc us- 1036 Ryuichi Majima and Akihiro Murata sion. We are grateful to Dr. J. Mark Erick­ Mus., No. 3~B, p. 1~330. (in Japanese). son, St. Lawrence University, Dr. T. Koike, Kuroda, T., 1931: Fossil Mollusca. In Homma, F., Shinano Chabu Chisitsu-Shi (Geology of Cen­ Yokohama National University, and an tral Shinano), part 4, p. I ~90, pis. I ~ 13. (in anonymous reviewer for their review of the Japanese). early or final version of the manuscript and MacNeil, F.S., 1960: Tertiary and Quaternary Gas­ providing helpful comments. Dr. T. Kase, tropoda of Okinawa. u.s. Geol. Surv., Prof Paper 339, p. I ~ 148, pis. I ~ 19. (published in National Science Museum, kindly sent us 1961). photographs of the holotype of Phanerole­ McNamara, K.J., 1986: A guide to the nomenclature pida pseudotransenna. Dr. Warren C. Blow, of heterochrony. Jour. Paleont., vol. 60, no. I, Smithsonian Institution, kindly allowed p. 4~13. access to the type specimens of Nehalemia Majima, R., 1985: Intraspecific variation in three species of Glossaulax (Gastropoda: Naticidae) hieroglyphica and Phanerolepida oregonensis from the late Cenozoic strata in central and in his care. Messrs. R. Uchimura and H. southwest Japan. Trans. Proc. Palaeont. Soc. Ishimoto, Shizuoka University, kindly assist­ Japan, N.S., no. 138, p. III ~ 137, pis. 17~ 19. ed in collecting fossil specimens. This study 1988: Evolution of Japanese Glossaulax was funded by Grants-in-Aids for Science species (Gastropoda: Naticidae). II: G. hyugensis, G. hagenoshitensis and G.nodai. Fos­ Research from the Ministry of Education, sils (Palaeont. Soc. Japan), no. 45, p. 1~7. (in Science and Culture, the Government of Japanese with English abstract). Japan (Nos. 02740394 and 03640654). Masuda, K. and Noda, H., 1976: Checklist and Bibliography of the Tertiary and Quaternary Mollusca of Japan, 1950~ 1974. 494 p. Saito References cited Ho-on Kai, Sendai. M imoto, K., 1986: Fossils in Kochi Prefecture. Aoki, N. and Baba, K., 1984: Additions to the Illustrations of the specimens kept by members molluscan fossils from the Nobori Formation, of the Kochi Fossil Society. 54 p. Kochi Fos­ Shikoku. Ann. Rep., Ins!. Geosci., Univ. Tsu­ sil Society, Kochi. (in Japanese). kuba, no. 10, p. 73~79. Noda, H., 1980: Molluscan fossils from the Ryukyu Dall, W.H., 1907: Descriptions of new species of Islands, southwestern Japan. Part I. Gastro­ shells, chiefly Buccinidae, from the dredgings of poda and Pelecypoda from the Shinzato Forma­ the U.S. "Albatross" during 1906, in the north­ tion in southeastern part of Okinawa-jima. Sci. western Pacific, Bering, and Japanese Sea. Rep., lnst. Geosci., Univ. Tsukuba, Sec. B, vol. I, Smithsonian Misc. Coli., vol. 50, pt. 2, p. p. 1~95, pis. 1~12. 139~ 173. --, 1988: Molluscan fossils from the Ryukyu Hatai, K. and Masuda, K., 1962: Megafossils from Islands, southwest Japan. Part 2. Gastropoda near Higashi-Matsuyama City. Saitama Prefec­ and Pelecypoda from the Shinzato Formation in ture, Japan. Trans. Proc. Palaeont. Soc. Japan, the middle part of Okinawa-jima. Sci. Rep., N.S., no. 46, p. 254~262, pI. 40. Ins!. Geosci., Univ. Tsukuba, Sec. B, vol. 9, p. Hickman, C. S., 1972: Review of the bathyal gas­ 29~85, pis. 5~ 19. tropod Phanerolepida (Homalopomatinae) and -- and Kikuchi, Y., 1980: Phanerolepida expan­ description of a new species from the Oregon silabrum (Kuroda) (Gastropoda) from the Oligocene. Veliger, vol. 15, no. 2, p. 107~112. Miocene N antaisan Volcanic Breccia, Ibaraki --, 1974: Nehalemia hieroglyphica, a new genus Prefecture, northeastern Japan. Venus, vol. 39, and species of archaeogastropod (Turbinidae: no. 1, p. 69~ 73, pI. 1. (in Japanese with English Homalopomatinae) from the Eocene of Oregon. abstract). Veliger, vol. 17, no. 2, p. 89~91. -- and Ogasawara, K., 1976: On the variation of Itoigawa, J., Shibata, H., Nishimoto, H., and Oku­ Phanerolepida rehderi MacNeil. Contr., Dept. mura, Y., 1981 : Miocene fossils of the Mizunami Geol. Miner., Niigata Univ., no. 4, p. 431 ~436, pI. Group, central Japan. 2. Molluscs. Monogr. 1. (in Japanese with English abstract). Mizunami Fossil Mus., No. 3~A, p. 1~53, pis. I~ Okutani, T., 1964: Report of the archibenthal and 52. (in Japanese). abyssal Mollusca mainly collected from Sagami --, --, --, and --, 1982: Miocene fossils of Bay and adjacent waters by the R.V. Soyo-Maru the Mizunami Group, central Japan. 2. Mol­ during the year 1955~ 1963. Jour. Fac. Sci., luscs (continued). Monogr. Mizunami Fossil Univ. Tokyo, Sec. 2, vol. 15, p. 371~447. 933. Intraspecific variation and heterochrony of Phanerolepida 1037

--, 1966: Archibenthal and abyssal Mollusca col­ Koti Prefecture, Japan. Bull. Natn. Sci. Mus.. lected by the R.V. Soyo-Maru from Japanese Tokyo, vol. 3, no. I (no. 38), p. 1-6, pI. 1. waters during 1964. Bull. Tokai Reg. Fish. Shibata, H., 1974: Phanerolepida cf. expansilabrum Res. Lab., vol. 46, p. 1-32. Kuroda. In, J. Itoigawa, H. Shibata, and H. --, 1968: Systematics, ecological distribution and Nishimoto, Geology and Paleontology of palaeoecological implication of archibenthal and Mizunami City. Bull. Mizunami Fossil Mus., abyssal Mollusca from Sagami Bay and adjacent no. I, p.43-203, pis. 1-63. (in Japanese). areas. Jour. Fae. Sci.. Unil'. Tokyo. Sec. 2, vol. Watanabe, K., Arai, J. and Hayashi, T., 1950: 17, pt. I, p. 1-98. Tertiary geology of the Chichibu basin. Bull. --, 1969: Synopsis of bathyal and abyssal megalo­ Chiehibu Mus. Nat. Hist., no. I, p. 29-92, pis. 1- invertebrates from Sagami Bay and the south off 6. (in Japanese). Boso Peninsula trawled by the RjV Soyo-Maru. Watson, R.B., 1879: Mollusca of H.M.S. 'Challenger' Bull. Tokai Reg. Fish. Res. Lab., vol. 57, p. 1-61. expedition. IV. Trochidae continued, viz. the Otuka, Y., 1949: Fossil Mollusca and rocks of the genera Basilissa and Troehus, and the Tur­ Kiyosumi Group exposed at Minato-machi, binidae, viz. the genus Turbo. Jour. Linn. Soc. Chiba Prefecture, and its environs (1st paper). London, vol. 14, p.692-716. [not seen]. Japan. Jour. Geol. Geogr., vol. 21, nos. 1-4, p. --, 1886: Report of the Scaphopoda and Gas­ 295-309, pI. 13. tropoda collected by H.M.S. Challenger during Ozaki, H., 1956: Some new and interesting molluscs the year 1873-1876. Res. Voy. Challenger. Rep. from Miocene Nobori Formation in the eastern Sci.• Zoology, vol. 15, p. 1-722.

Hane 3fJfN. Kochi ~:ttl!, Muroto ~p, Nishinotani g§ /1:)-, Nobori E;, Okinawa rq,~g, Saitama ~~. Shinzato ~1[, Tokigawa m~JII. Tonohama Jl!trA.

~~iJj!,O)!'f~*lCE;~.i£: Phanerolepida pseudotransenna Ozaki (.im:[email protected]: 1) "'" ? T""/-ij--tf' =fil-) O)fiP'l~~c~lF.fli: Phanerolepida pseudotransenna Ozaki O)JI:ilf:96~ ~ht.::. 75 j~lt*f:I:.~~rnH\uflit-:'~F~t-:.* ~ t.e~~>a:-~i"o .: o)~~li{fEI-(;$:r,,'O)~~96~O)Im~IF.fWlO) ~~t-:.~IZSIi"."6 c m~ ~ h."6 o ~ff.:1¥J.i£:lli c ~O)~~O)OO-(;$:961:.J:O)~~O).Itti:iJ' I? Ne­ halemia hieroglyphiea .... Phanerolepida oregonensis .... P. pseudotransenna .... P. tansenna c \" ? im1~*7IJiJ~Jt5E ~ ht.::.o -l" 0) ? t;" 1't.e < c HJJtlbO) 2 """:)O)fi53'1~li~1F.f1tO)t.t* -Ctl;."6 c1f;;tl?h."6 o Phanerolepida rehderi MacNeil >a:-OO-(;$:~~O).l:~~t-:.i1i\;":)I,,-C P. pseudotransenna O)~* :/ / .::. A C L- t.::.o ~ c f*"'? t.::. 4 """:)o)~iJ~1:..f,.!,IF.fO){ftlit-:.:ilf:\' ,tkfrn"C'.i£:lli L- t.::.o r,,'~~Ili- . HEHIIB~ 1038 Ryuichi Majima and Akihiro Murata

Appendix. Measurements, letter formula and remarks for all examined specimens of Phanerolepida pseudotransenna Ozaki from the Nobori Formation. Number Letter formula of Specimen Locality Shell Shell Aperture of shell-surface Remarks IGSU IGSU height width height whorl ornamentation M37 62-35-1 17.0 20.0 12.4 4 1/2+ SRFsRFsFRFsF Fig. 3-1 M38 62-35-1 17.8 20.3 13.1 4+ SRF Fig. 3-2 M39 62-35-1 17.7 19.8 12.4 4 1/2+ SRF Fig. 3-3 M40 62-35-1 16.9 20.2 12.3 4+ SRF M41 62-35-1 16.4 19.0+ 10.8 3 1/2+ SRF M42 62-35-1 15.6 17.2+ 11.4 4 1/2+ SsRF M43 62-35-1 17.8+ 17.7+ 12.5+ 3 1/2+ SRF?F M44 62-35-1 10.9+ 16.5+ 3+ SRF M45 62-35-1 12.3+ 16.9 11.8 1/2+ ?F Deformed M46 62-35-1 11.3+ 15.3+ 9.8+ 1/2+ ?F M47 62-35-1 11.0+ 18.5 13.5 1/2+ ?F M48 62-35-2 14.0+ 20.0+ 9.5+ 3+ ?F Deformed M49 62-35-2 9.9 15.0 8.0 3 1/2+ ?RF Deformed M50 62-35-2 14.7 17.1+ 12.2 3+ ?R?R Deformed M51 62-35-2 15.1 19.5+ 10.1 4 1/2+ ?F Deformed M52 62-35-2 15.8 18.4 11.9 4 1/4 SRF?F Deformed M53 62-35-2 12.7+ 17.0 11.5 2+ ?F Deformed M54 62-35-2 11.3+ 15.8+ 11.0+ 2+ ?F?F? Deformed M55 62-35-2 8.4+ 15.2+ 8.0+ 2 1/2+ ?FR Deformed M56 62-35-2 11.0+ 19.1+ 1+ ?F? Deformed M57 62-35-2 8.8+ 13.8+ 1/2+ ?F Deformed M58 62-35-2 8.8+ 11.2+ 3+ ?F? Deformed M59 62-35-3 16.3 18.8 10.7 5+ SRF Fig.4 M60 62-35-3 13.5 16.7 9.8 4 1/2+ SRFRFRF Figs. 3-4, 4, 5-4 M61 62-35-3 15.8 18.1 11.6 4 1/2+ SRFsF Figs. 4, 5-8 M62 62-35-3 14.2 17.0 10.6 3 1/2+ SRF Fig. 4 M63 62-35-3 15.9 17.4 11.7 4+ SRF Fig. 4 M64 62-35-3 17.2 19.0 11.9 4 1/2+ SRFRFRFsRFs Fig. 3-5,4, 5-2 M65 62-35-3 14.8 18.2 10.5 3+ SRFRF Fig. 4 M66 62-35-3 17.2 20.3 13.0 4+ SRF Fig. 4 M67 62-35-3 17.0 19.5+ 12.5 3 1/2+ SRF Figs. 4, 5-10 M68 62-35-3 14.1 18.0 11.2 5+ SR Figs. 4, 5-11 M69 62-35-3 16.3 17.8 10.9 3 1/2+ SRFsRFsRF Figs. 4, 8-4, with operculum M70 62-35-3 16.8 19.1+ 12.4 4+ SRF Fig. 4 M71 62-35-3 16.3 19.6+ 12.4 4+ SRF Fig. 4 M72 62-35-3 15.6+ 16.7+ 11.1+ 3 1/2+ SRFsFRFRF Figs. 4, 5-3 M73 62-35-3 15.0+ 17.1+ 12.0+ 5 SRFRFRFRF Figs. 4, 5-1, 8-1, with operculum M74 62-35-3 16.3+ 16.4+ 12.1+ 6 1/2 SRFRF Figs. 4, 5-6 M75 62-35-3 15.0 17.6+ 12.4 3 1/2+ SRFsRs Figs. 4, 5-7 M76 62-35-3 14.0 19.5 9.3+ 4+ SRFsF Fig. 4 933. Intraspecific variation and heterochrony of Phanerolepida 1039

Number Letter formula of Specimen Locality Shell Shell Aperture of shell-surface Remarks rGSU rGSU height width height whorl ornamentation M77 62-35-3 15.4 17.3 11.7 3 1/2+ SRF Fig. 4 M78 62-35-3 14.5 18.4 11.9 3+ ?RF M79 62-35-3 16.2 19.3 12.8 3 1/2+ ?FR?s M80 62-35-3 16.6 19.5 13.1 4 1/2+ ?RF Deformed M81 62-35-3 14.9+ 17.3+ 11.6+ 3+ ?RFsR?F Deformed M82 62-35-3 14.3+ 19.7+ 10.8+ 3+ ?F Deformed M83 62-35-3 14.0 17.8 11.3 3 1/2+ SRFs Fig. 4 M84 62-35-3 14.2 18.8+ 10.2+ 3+ SRF?F Deformed M85 62-35-3 14.7+ 18.4+ 13.0 2+ ?FRF?F M86 62-35-3 14.5 16.5+ 11.3 4+ S?s?s Fig. 5-12 M87 62-35-3 16.4 18.6 13.0 3 1/2+ SRF?F?F Deformed M88 62-35-3 13.5 18.3+ 12.8+ 3 1/2+ ?FR?F Deformed M89 62-35-3 16.7 18.1+ 13.8 3 1/2+ SRF?F M90 62-35-3 13.3+ 17.3+ 10.6+ 3+ ?F Deformed M91 62-35-3 14.2+ 15.1+ 10.8+ 5 1/2 SRF Fig. 4 M92 62-35-3 12.2+ 15.3+ 10.0+ 4+ SR? M93 62-35-3 13.1+ 17.5+ 10.9+ 4+ S?F? Deformed M94 62-35-3 11.6+ 14.1+ 8.9+ 6 SRF Figs. 4,5-9 M95 62-35-3 11.4+ 13.4+ 9.3+ 4 1/2+ SR? M96 62-35-3 14.1+ 14.8+ 10.2+ 5+ SRF?F M97 62-35-3 12.7+ 18.4+ 3+ SRF Fig. 4 M98 62-35-3 12.7+ 18.0+ 11.0 ?R? Deformed M99 62-35-3 21.4 12.6 ?RF MIOO 62-35-3 13.3+ 15.4+ 5 1/2+ SRFRFRF Figs. 4, 5-5 MIOI 62-35-3 13.6+ 19.0+ 11.0+ 3+ ?RF?F MI02 62-35-3 12.7+ 11.2+ ?F Deformed MI03 62-35-3 11.9+ 15.3+ 9.8+ 2 1/2+ ?F?F Deformed MI04 62-35-3 9.3+ 13.4+ 8.3+ ?F Deformed MI05 62-35-3 14.8+ 3+ ?RF MI06 62-35-3 15.0+ 11.2+ 10.3+ 3 1/2+ SR?F MI07 62-35-3 14.1+ 11.4+ 4+ SR?FRF?F MI08 62-35-3 12.5+ 15.0+ 8.9+ 4+ SRF?Fs MI09 62-35-3 10.6+ 14.5+ 2 1/2+ SRF?FsRF? MIlO 62-35-3 13.9+ 13.3+ 2 1/2+ ?F? Mill 62-35-3 13.2+ 14.9+ 10.3+ 4 1/2+ ?F?F? Fig. 8-5, with operculum MI12 62-35-3 13.3+ 9.5+ ?F? Fig. 8-3, with operculum MI13 62-35-3 11.6+ 11.9+ 8.7+ 3 1/2+ SRF? Deformed MI14 62-35-3 19.2+ 3+ S?F?F MI15 62-35-3 12.6+ 15.3+ 9.5+ 4+ SRF?F MI16 62-35-3 7.1+ 10.5+ 7.0+ 3+ ?RF? Trans. Proc. Palaeont. Soc. Japan, N.S., No. 165, pp. 1040-1069, 12 Figs., April 30, 1992

934. VERBEEKINID AND NEOSCHW AGERINID FUSULINACEANS FROM THE AKIYOSHI LIMESTONE GROUP ABOVE THE PARAFUSULINA KAERIMIZENSIS ZONE, SOUTHWEST JAPAN*

KATSUMI UENO

Institute of Geoscience, University of Tsukuba, Ibaraki, 305

Abstract. The Akiyoshi Limestone Group above the Paraflislilina kaerimizensis Zone in the Kaerimizu area can be divided into the Paraflislilina kaerimizensis, Afghanella ozawai, Neoschwagerina craticlilifera robusta, Verbeekina verbeeki-Afghanella schencki, Neoschwagerina fusiformis, Verbeek ina verbeeki and Colania dOllvillei Zones in ascending order, mainly on the basis of the stratigraphic distribution of verbeekinid and neoschwager­ inid fusulinaceans. In this paper, 10 species of verbeekinid and neoschwagerinid fusulinaceans including one new subspecies, Neoschwagerina craticlilifera robusta, are described. Also, the fusulinacean biostratigraphy of the Akiyoshi Limestone Group above the Parafllslllina kaerimizensis Zone in the Kaerimizu area is briefly noted.

Key words. Akiyoshi Limestone Group, fusulinacean biostratigraphy, Middle Per­ mian, Parafllslllina kaerimizensis Zone, Neoschwagerinidae, Verbeekinidae.

Introduction Concerning the biostratigraphic zonation by fusulinaceans in the Permian sequence of the The Akiyoshi Limestone Group, covering Akiyoshi Limestone Group, Ota (1977) sub­ almost the whole area of the Akiyoshi Pla­ divided it into II zones. Recently, Ozawa teau situated in the central part of Yamaguchi and Kobayashi (1990) established 17 Prefecture, southwest Japan, is considered to fusulinacean zones in the same part. have formed as a large organic reef complex The fusulinacean family Verbeekinidae during Late Paleozoic time (Ota, 1968). arose from the family Ozawainellidae at the This limestone contains rich shallow marine beginning of the Middle Permian and the fossil remains such as foraminifers, algae, family Neoschwagerinidae was derived from sponges, corals, brachiopods, mollusks and the former in the middle of that period. others. Since Ozawa's fundamental work in These two families flourished through Middle 1923, fusulinacean foraminifers have often Permian time. It is known that they have been studied in connection with the biostrati­ significant value for biostratigraphic zonation graphic subdivision of the Akiyoshi Lime­ in the Middle Permian of the Tethyan realm. stone Group, except for its lowermost part About 30 species of the families Verbee­ (e.g. Ozawa, 1925b; Toriyama, 1954,1958; kinidae and Neoschwagerinidae have been Hasegawa, 1958, 1963; Murata, 1961; Sa­ recorded from the Akiyoshi Limestone kaguchi et al., 1966; Yanagida et al., 197 I ; Group. Ota et al., 1973; Ota, 1977; Matsusue, 1986 ; Very recently, I have discussed the origin Ueno, 1989; Ozawa and Kobayashi, 1990). and early evolution of verbeekinid and neo­ 'Received September 17, 1991; accepted December schwagerinid fusulinaceans, based on bios­ 20, 1991 tratigraphic evidence obtained from the

1040 934. Verbeekinids and neoschwagerinids from Akiyoshi 1041

Parafusulina kaerimizensis, Afghanella schencki, Neoschwagerina craticulifera, Ver­ beekina verbeeki and Colania douvillei Zones. I recognized seven fusulinacean biohor­ izons characterized by the first and last occur­ Kdma (;) rence of some marker species in the stratigra­ aerimizu t ?< phic interval between Ota's Parafusulina kaerimizensis and Colania douvillei Zones (Figure 2). Each biohorizon is well recog­ nized in the examined sections of the Kaerim­ izu area and defines the base of each fusulinacean zone established herein. The fusulinacean zones are distributed almost concentrically in the Kaerimizu Doline, being younger toward the bottom of o 2km the doline, except for the southern part of the Akiyoshi I ...... surveyed area where the Afghanella ozawai Figure 1. Map showing the Kaerimizu area in and Neosch wagerina craticulifera robusta the Akiyoshi Plateau. Zones are shown as the inlier in the Para­ fusulina kaerimizensis Zone (Figure 3). Consequently, the Akiyoshi Limestone Akiyoshi Limestone Group below the Misel­ Group in this area lies almost horizontally lina (M.) claudiae Zone in the Kaerimizu and is in reverse order. area (Ueno, 1991 a, 1991 b). A total of II species of verbeekinid and This paper deals with the biostratigraphy neoschwagerinid fusulinaceans were identi­ of verbeekinid and neoschwagerinid fied from the Akiyoshi Limestone Group fusulinaceans from the Akiyoshi Limestone above the Parafusulina kaerimizensis Zone in Group above the Parafusulina kaerimizensis the Kaerimizu area. Their stratigraphic dis­ Zone in the Kaerimizu area, and 10 species of tribution is shown in Figure 4. The them including one new subspecies, Neo­ fusulinacean biostratigraphy of the Para­ schwagerina craticulifera robusta, are de­ fusulina kaerimizensis Zone through the scribed. In addition, the fusulinacean bio­ Colania douvillei Zone is summarized as fol­ stratigraphy of the Akiyoshi Limestone lows. Group above the Parafusulina kaerimizensis Parafusulina kaerimizensis Zone: This Zone is briefly noted. zone is defined as the stratigraphic interval between the first occurrence of Parafusulina Fusulinacean biostratigraphy kaerimizensis and that of Afghanella ozawai, and is about 15 m in thickness. Stratigra­ The Kaerimizu area is a large doline locat­ phic relationship with the underlying Misel­ ed in the northeastern part of the Akiyoshi lina (M.) claudiae Zone is conformable. Plateau (Figure I). Ota (l977) divided the Well sorted foraminiferal sparitic limestone Akiyoshi Limestone Group in this area into and algal foraminiferal sparitic limestone are the following 10 fusulinacean zones in predominant. The following fusulinaceans ascending order: the Fusulinella biconica, are discriminated: Rauserella aff. breviscula Triticites simplex, Pseudofusulina vulgaris, Sosnina, Rauserella sp., Minojapanella (Rus­ Pseudofusulina ambigua, Misellina claudiae, siella) pulchra Miklukho-Maklay, Neo- 1042 Katsumi Veno

FU SULINACEAN Z 0 N E BIOHORIZON Coiania douvillei .... F. Colania douvillei Verbeekina verbeeki .... L. Neoschwagerina fusiform is & Pseudodoliolina pseudolepida Neoschwag. fusiformis .... F. Neoschwagerina fusiform is Verbeekina verbeeki- Afghanella schenck.i .... F. Verbeek ina verbeeki & Neoschwagerina Afghanella schencki craticulifera robusta .... F. Neoschwagerina craticulifera robusta Afghanella ozawai .... F. Afghanella ozawai Paraf. kaerimizensis .... F. Parafusulina kaerimizensis Mise/lina (1.4.) c/audiae .... F. Neoschwagerina simplex (PART) .... F. "'aklaya pamirica

F.: First occurrence L.: Last occurrence Figure 2. Fusulinaeean zones and biohorizons above the Parafusulina kaerimizensis Zone. fusulinella giraudi Deprat, N. pseudogiraudi giraudi Deprat, N. pseudogiraudi (Sheng), (Sheng), Neofusulinella sp., Pseudoreichelina Pseudoendothyra spp., Parafusulina kaerim­ sp., Pseudoendothyra spp., Parafusulina izensis (Ozawa), Pseudodoliolina cf. ozawai kaerimizesis (Ozawa), P. lutugini (Schel­ Yabe and Hanzawa, Afghanella ozawai Han­ Iwien) and Chusenella alpina (Kochansky­ zawa and Afghanella sp. Devide and RamovS). Neoschwagerina craticulifera robusta Afghanella ozawai Zone: Afghanella oz­ Zone: This zone is in conformable contact awai Zone, about IO m thick, conformably with the subjacent Afghanella ozawai Zone overlies the Parafusulina kaerimizensis Zone. and is about 20 m in thickness. The lower Limestone facies of the present zone is similar limit of the present zone is defined by the first to that of the Parafusulina kaerimizensis occurrence of Neoschwagerina craticulifera Zone. The fusulinacean fauna incl udes robusta, subsp. nov. Algal foraminiferal Rauserella aff. breviscula Sosnina, Rauserella sparitic limestone and foraminiferal micritic sp., Schubertella rara Sheng, Neofusulinella limestone are predominant. The identified

~ Figure 3. Distribution offusulinacean zones and fossil localities in the Kaerimizu area. I: Misellina (M.) claudiae Zone (Me), 2: Parafusulina kaerimizensis Zone (Pk), 3: Afghanella ozawai Zone (Ao), 4: Neoschwagerina craticutijera robusta Zone (Nc), 5: Verbeekina verbeeki-Afghanella schencki Zone (V-A), 6: Neoschwagerina fusijormis Zone (Nf), 7: Verbeekina verbeeki Zone (Vv), 8: Cotania douvillei Zone (Cd), 9: fossil locality studied herein. 934. Verbeekinids and neoschwagerinids from Akiyoshi 1043

100m I

:::::::::::::::(nnnnH~ 2 ...... 3 •...... !TITIillJs

6 7 8 • 9 1044 Katsumi Ueno fusulinaceans are as follows: Rauserella alf. with a subordinate amount of algal micritic breviscula Sosnina, Rauserella spp., Dunbar­ limestone and muddy limestone. The ula schubertellaeformis Sheng, Neofusulinella fusulinacean fauna includes Rauserella cf. pseudogiraudi (Sheng), Pselldoreichelina sp., sphaeroidea Sosnina, Rauserella sp., Dunbar­ Pselldoendothyra spp., Parafuslilina kaerim­ ula schubertellaeformis Sheng, Dunbarula izensis (Ozawa), Chusenella referta Skinner spp., Schubertella sp., Neofusulinella sp., and Wilde, C. alpina (Kochansky-Devide and Staffella sp., Pseudoendothyra spp., Para­ Ramovs), C. schwagerinaeformis Sheng, fusulina edoensis (Ozawa), P. gigantea (De­ Pselldodoliolina cf. ozawai Yabe and Hanza­ prat), Parafusulina spp., Chusenella? cf. wa, and Neosch wagerina craticulifera roblls­ pindingensis (Sheng), Chusenella sp., Verbee­ ta, subsp. nov. kina verbeeki (Geinitz), Pselldodoliolina Verbeekina verbeeki-Afghanella schencki pseudolepida (Deprat), Afghanella schencki Zone: This zone, about 10 m in maximum Thompson and A. cf. sumatrinaeformis (Gub­ thickness, is defined as the stratigraphic inter­ ler). val between the first occurrence of both Neoschwagerina fusiform is Zone: Neo­ Verbeekina verbeeki and Afghanella schwagerina fusiformis Zone, about 10 m in schencki and that of Neoschwagerina fusifor­ maximum thickness, is in conformable con­ mis, and conformably overlies the Neo­ tact with the underlying Verbeekina schwagerina cra tic lllifera robusta Zone. verbeeki-Afghanella schencki Zone. In this Foraminiferal sparitic limestone is dominant zone, foraminiferal sparitic limestone is pre-

N < (") 1> 2 I < o (') < c.. :J o 1> CD 444 • Maklaya pamirica • Neoschwagerina simplex Parafusulina kaer imizensis ." c::: Afghanella ozawai en Afghanella sp. c::: Pseudodoliolina ct. ozawai Neoschwag. craticulifera robusta Verbeekina verbeeki ---...... --- Afghanella schencki Afghanella ct. sumatrinaeformis Pseudodoliolina pseudolepida Neoschwagerina fusiform is Neoschwagerina haydeni Colania douvillei 4 : Biohorizon

Figure 4. Stratigraphic distribution of verbeekinid, neoschwagerinid and some related fusulinaceans. (abbreviations of zone names: see legend of Figure 3) 934. Verbeekinids and neoschwagerinids from Akiyoshi 1045

dominant with a subordinate amount of for­ imizensis Zone through the Colania douvillei aminiferal micritic limestone and muddy Zone is considered to be the Murgabian of limestone. The following fusulinaceans are the standard stratigraphic scheme in the Teth­ identified: Rauserella sp., Dunbarula yan province proposed by Leven (1980). schubertellaeformis Sheng, Dunbarula spp., Schubertella sp., Neofusulinella sp., Systematic paleontology Pseudoendothyra spp., Parafusulina sp., Sch wagerina sp., Pseudodoliolina pseudole­ All specimens identified in this paper are pida (Deprat), Neoschwagerina fusiformis deposited in the paleontological collections Skinner and Wilde, and N. haydeni Dut­ of the Akiyoshi-dai Museum of Natural kevich and Khabakov. History. The abbreviation ASM is the prefix Verbeekina verbeeki Zone: The lower of the specimen repository number. limit of the Verbeekina verbeeki Zone is defined by the last occurrence of Neo­ Order Foraminiferida Eichwald, 1830 sch wagerina fusiformis and Pseudodoliolina Suborder Fusulinina Wedekind, 1937 pseudolepida. In the same horizon, Verbee­ Superfamily Fusulinacea von kina verbeeki appears again, which is diag­ Moller, 1878 nostic in the Verbeekina verbeeki-Afghanella Family Verbeekinidae Staff schencki Zone but barren in the Neo­ and Wedekind, 1910 schwagerina fusiformis Zone. The thickness Subfamily Verbeekininae Staff of the present zone is about 10 m. A strati­ and Wedekind, 1910 graphic hiatus may exist between the Verbee­ kina verbeeki Zone and subjacent ones, judg­ Genus Verbeekina Staff, 1909 ing from the distribution of fusulinacean Type species. - Fusulina verbeeki Geinitz, zones. The following fusulinaceans are di­ 1876. scriminated: Rauserella sp., Dunbarula schubertellaeformis Sheng, Dunbarula spp., Verbeekina verbeeki (Geinitz, 1876) Neofusulinella sp., Pseudoreichelina sp., Pseudoendothyra spp., Chusenella referta Figures 5-1-6,6-1-5 Skinner and Wilde, and Verbeekina verbeeki Fusulina I'erbeeki Geinitz, 1876, p.399-400. (Geinitz). Schwagerina l'erbeeki (Geinitz). Staff, 1909, p.506, Colania dOllvillei Zone: The Colania pI. 7, figs. 5, 7; Deprat, 1912, p. 40-41, pI. I, figs. douvillei Zone occupies the highest stratigra­ 7-11; Colani, 1924, p. 108-113, pI. 15, fig. 14, pI. phic position in the Kaerimizu area and only 18, figs. 2,5,16-21,25-27, pI. 19, figs. 1-7, II, 13. crops out at the bottom of the doline. It is at Verbeekina verbeeki (Geinitz). Ozawa, 1925b, p. 48- 51, pI. 10, figs. 6-7; Lange, 1925, p. 260-261, pI. least 5 m in thickness and in conformable 3, fig. 66a, pI. 5, fig. 66b; Ozawa and Tobler, contact with the Verbeekina verbeeki Zone. 1929, p. 48, pI. 5, fig. 5; Chen, 1934, p. 101-102, The lower limit of the present zone is defined pI. 16, fig. I ; Gubler, 1935, pI. I, figs. 10-16; by the first occurrence of Colania douvillei. Thompson, 1936, p. 197-200, pI. 24, figs. 1-8 ; The upper limit, however, can not be H uzimoto, 1936, p. 10 I-I 04, pI. 9, fig. 8, pI. 21, figs. 1-3; Thompson, 1948, pI. 16, figs. 7-12 confirmed in the Kaerimizu area. Algal (same as pI. 24, figs. 1-2,4,6-8 of Thompson, sparitic limestone and muddy limestone are 1936, p. 197-200); Kochansky-Dcvide and dominant. The present zone is rather poor Ramovs, 1955, p. 389,416, pI. 4, figs. 1-3; Chen, in foraminifers and only two species of 1956, p.9,47-48, pI. 9, figs. 5-6, pI. 13, figs. 1-2; Miklukho-Maklay, 1957, p. 113-114, pI. 3, fig. 2; fusulinaceans, Pseudoendolhyra sp. and Kobayashi, 1957, p.301-302, pI. 9, figs. 5-7 ; Colania dOllvillei (Ozawa), are identified. Toriyama, 1958, p.205-208, pI. 37, figs. 1-6, pI. The geologic age of the Parafusulina kaer- 38, figs. 1-6; Rauser-Chcrnoussova and Fursen- 1046 Katsul17i Vena

Figure S. 1- 6. VerbeekinG verbeeki (Geinitz), la, 2-4 : axia l sections, ASM25156, ASM25162, ASM25161 , ASM25155, 5,6 : sagittal secti o ns, ASM25164, ASM25165, x 10, lb : enlarged part of la, x 40. 934. Verbeekinids and neoschwagerinids from Akiyoshi 1047

ko, 1959, pI. 10, fig. 5; Nogami, 1961, p.167- section of polyvalent individual; ASM25159 169, pI. 2, figs. 1-4; Sheng, 1963, p. 85-86, 215- from Loc. T A256. 216, pI. 26, figs. 1-5; Hanzawa and Murata, 1963, pI. 4, fig. 8; Igo, 1964, p. 62-63, pI. 2, fig. Description. - Shell large and almost spheri­ I ; Ishii and Nogami, 1964, p. 24, pI. 8, figs. 4-6; cal with very shallow umbilicus. Mature Pasini, 1965, pI. 16, fig. I ; Leven, 1965, p. 140, specimens having 13 to 15 volutions, 5.40 to pI. 4, fig. 1 ; Ishii, 1966, pI. 6, fig. 9; Sheng, 1966, 7.20 mm in length and 5.15 to 7.00 mm in p. 138-139, pI. 23, fig. I (same as pI. 13, fig. 1 of Chen, 1956, p. 9,47-48); Skinner and Wilde, width, with a form ratio of about 1.00. 1966, pI. 17, fig. 6; Igo, 1967, pI. 6, figs. 1-2 ; Inner 3 to 5 vol utions eostaffelloid and Leven, 1967, p. 205, pI. 38, fig. I ; Choi, 1970, p. tightly coiled with rotated axis of coiling in 348-349, pI. 15, figs. 1-3; Ozawa, 1970a, pI. 2, some case. Beyond the fifth volution, shell fig. I; Tien, 1970, p.35-37, pI. 6, figs. 1-10; becomes spherical and loose rather rapidly. Toriyama and Pitakpaivan, 1973, p. 50-53, pI. 6, figs. 1-5; Zhang and Wang, 1974, p. 293, pI. 152, Chamber height almost constant throughout fig. 3; Toriyama, 1975, p.66-68, pI. 14, fig. 12; length of shell. Radius vectors of the first to Rozovskaya, 1975, pI. 27, figs. 6-7; Sheng and twentieth volutions of well oriented axial Sun, 1975, p.47, pI. 2, fig. 13; Ozawa, 1975, pI. section (Figure 5-1) 0.05, 0.08, 0.12, 0.20, 0.34, 10, figs. 10-11 ; Toriyama and Kanmera, 1977, p. 0.60,0.88, 1.19, 1.56, 1.91, 2.23 and 2.51 mm, 8-9, pI. I, figs. 8-10; Lin et at., 1977, p.83-84, pI. 25, fig. 9; Ota, 1977, pI. I, figs. 5-6; Liu et and form ratios 0.50,0.53,0.67,0.92,0.97, at., 1978, p. 81, pI. 18, fig. 17; Ding, 1978, p. 288, 1.05, 1.06, 1.09, 1.05, 1.03, 1.03 and 1.04, pI. 100, fig. 2; Chen and Yang, 1978, p. 105, pI. respectively. 27, fig. 3; Kobayashi, 1979, pI. 4, figs. 9-10; Proloculus spherical and exceedingly min­ Hasegawa et at., 1979, pI. 85, figs. 1-5,7; Tien, ute for shell size. Its outside diameter varies 1979, p. 130-132, pI. 21, figs. 3-9, pI. 22, fig. I B, pI. 23, fig. 3B; Kahler and Kahler, 1979, p. 215- from 0.030 to 0.060 mm, averaging 0.045 mm 217, pI. 2, fig. 4, pI. 10, figs. 1-2, 4-5; Wang et for 8 specimens. at., 1981, p.58-59, pI. 19, fig. 6; Wang et at., Spirotheca thin and composed of a single 1982, p. 99, pI. 27, figs. 1-2; Xie, 1982, p. 66, pI. structureless layer in inner 3 volutions, but of at., 1977, 32, fig. 3(same as pI. 25, fig. 9 of Lin et a tectum and lower fibrous alveolar kerioth­ p. 83-84); Sun et aI., 1983, p. 33, pI. 9, fig. 5 ; Chen, 1984, p.64, pI. 17, figs. 1-2; Yang, 1985, eca beyond the fourth volution. Alveolar pI. 2, fig. 34; Tien, 1986a, pI. 7, fig. 13; Tien, keriotheca finer than that observed in neo­ 1986b, pI. 12, fig. II ; Ishibashi, 1986, p. 105-107, schwagerinids. Thickness of spirotheca of pI. 15, figs. 2-8, pI. 16, figs. 1-3; Wang and Tang, the first to thirteenth volutions of above­ 1986, pI. 2, fig. 16; Kobayashi, 1986, p. 145, pI. 4, fig. 8, pI. 9, fig. 6; Wang and Zhou, 1986, pI. 2, mentioned specimen 0.010, 0.010, 0.010, 0.020, figs. 1-2; Han el at., 1987, pI. 2, figs. 1-2,6; 0.015,0.025,0.030,0.040,0.055,0.065,0.090, Loeblich and Tappan, 1988, pI. 298, figs. 1-5 0.065 and 0.050 mm. (same as pI. 24, figs. 1-2,6-8 of Thompson, 1936, Septa thin, unfluted throughout shell p. 197-200); Kobayashi, 1988a, p.8, pI. 11, figs. length and bending anteriorly. They are 10-14; Kobayashi, 1988b, p.443, figs. 5-3 -4; Tien, 1989, pI. 33, figs. 7-8; Ozawa and Kobaya­ composed of the same elements as the spiroth­ shi, 1990, pI. 10, fig. 11. eca. Average septal counts of the first to Verbeekina verbeeki forma A, Pitakpaivan, 1965, p. thirteenth vol utions for 3 specimens 6, 9, 11, 53-54, pI. 6, fig. 5. 11, 8, 8, 10, 12, 18, 19, 25, 30 and 36, respec­ tively. Material.-Axial sections; ASM25157 Parachomata numerous, small and low, from Loc. SF77, ASM25158 from Loc. SF79, and semicircular in transverse section. They ASM25155, ASM25156 from Loc. SF80, are well developed in inner juvenile volu­ ASM25160 from Loc. SA 120, ASM25161, tions, but sporadically and inconstantly ob­ ASM25162, ASM25163 from Loc. SA 152. served in some outer ones, because they Sagittal sections; ASM25164 from Loc. develop only on both sides adjacent to septa SA96, ASM25165 from Loc. SA152. Axial and decrease in height rather rapidly toward 1048 Katsumi Vena the center of the cham bers. The Akiyoshi specimens are quite similar to Remarks. - Verbeekina verbeeki (Geinitz) the topotypes restudied and redescri bed by is one of the most fami liar verbeekinids in the Thompson (1936). Middle Permian of the Tethyan province. In the Kaerimizu area, the distribution of

Sb Figure 6. 1- 5. Verbeekina verbeeki (Geinitz), 1- 4: axial sections, ASM25l57, ASM25l58, ASM25l60, ASM25163, ASM25159, Sa: axial section of polyva lent individua l, X 10, 5b: enlarged part of Sa, X 40. 934. Verbeekinids and neoschwagerinids from Akiyoshi 1049

Verbeekina verbeeki (Geinitz) is stratigra­ mcra, 1977, p. 12-14, pI. I, figs. 13-18, pI. 2, figs. phically separated into two intervals, namely 1-6; Lin el al., 1977, p.89, pI. 27, fig. II; Liu the Verbeekina verbeeki-Afghanella sclzencki el al., 1978, p.83, pI. 20, fig. 5; Ding, 1978, p. 288, pI. 98, fig. 6; Toriyama and Kanmcra, 1979, Zone and the Verbeekina verbeeki Zone. p. 54-56, pI. 8, figs. 7-13; Hasegawa el al., 1979, Most of the individuals found in the former pI. 83, figs. I, 2,4; Ticn, 1979, p. 132-134, pI. 20, zone are generally smaller in size and have figs. 4-6; Kahler and Kahler, 1979, p. 224-225, fewer volutions than those in the latter one, pI. 3, fig. 3; Sun et al., 1983, p.37, pI. 9, fig. 3; although other important morphological Chen, 1984, p. 68, pI. 19, fig. 4; Yang, 1985, pI. 2, fig. 28; Tien, 1986a, pI. 7, fig. II (same as pI. 20, characters are in close agreement. fig. 5 ofTien, 1979, p. 132-134); Tien, 1986b, pI. Occurrence.-Abundant in the Verbeekina 11, fig. 10, pI. 12, figs. 1-3, 5D, liB; Han et al., verbeeki-Afghanella schencki and Verbee­ 1987, pI. 2, fig. 4; Kobayashi, 1988a, p. 10, pI. 5, kina verbeeki Zones. figs. 18-21, pI. 6, fig. 18; Kobayashi, 1988b, p. 443-445, figs. 6-1 - 8; Tien, 1989, pI. 32, figs. 6- 7; Ozawa and Kobayashi, 1990, pI. 10, fig. 8. Subfamily Pseudodoliolininae Leven, 1963 Pseudodoliolina cf. pseudolepida (Deprat). Kawano, 1961, p. 105-107, pI. 10, figs. 8-10. Pselldodoliolina aIr. pselldolepida pseudolepida (Dc­ Genus Pseudodoliolina Yabe prall. Kanmera, 1963, p. 108-109, pI. 14, figs. and Hanzawa, 1932 15-18. Type species. - Pseudodoliolina ozawai Pselldodoliolina ozawai Yabe and Hanzawa. Chen Yabe and Hanzawa, 1932. and Yang, 1978, p. 112, pI. 30, figs. 4-5. Pseudodoliolina sp., Hasegawa et al., 1979, pI. 83, figs. 3,5 -7; Tien, 1989, pI. 32, figs. 8-10 (9: same as Pseudodoliolina pseudolepida pI. 11, fig. 1 of Tien, 1986b). (Deprat, 1912) Material. - Axial sections; ASM25167 from Figures 7-1- 10 Loc. AL169, ASM25166 from Loc. ALl80.5, ASM25168 from Loc. KN4, ASM25169 from Doliolina pseudolepida Deprat, 1912, p. 46, pI. 5, figs. Loc. T A26l, ASM25170 from Loc. SA 119, 6-9, pI. 6, fig. 4. Doliolina lepida mut. pseudolepida Deprat, Deprat, ASM25171 from Loc. EB98, ASM25172 from 1913, p. 50. Loc. MT253. Sagittal sections; ASM25l73, Doliolina lepida (Schwager). Ozawa, 1927, p. 152- ASM25174 from Loc. T A263. Tangential sec­ 153, pI. 45, figs. 1-2 (not pI. 37, figs. Ie, 7b, 8b, tion; ASM25175 from Loc. KS201. 9b); Gubler, 1935, p. 95-99, pI. 4, figs. 3-10, II?, Description. - Shell medium to large for 12?, pI. 5, figs. 14-15; Ciry, 1941, pI. I, fig. 2. Pseudodoliolina pseudolepida (Deprat). Thompson genus, cylindrical with broadly rounded and Foster, 1937, p. 141-142, pI. 25, figs. 2-4; polar regions and convex lateral slopes. Thompson, 1948, pI. 17, figs. 3-4 (same as pI. 25, Mature specimens of II to 14 volutions, 4.55 figs. 2,4 of Thompson and Foster, 1937, p. 141- to 6.23 mm in length and 1.90 to 3.10 mm in 142); Kochansky-Devide and Ramov~, 1955, p. 416-417, pI. 4, figs. 7-10, pI. 8, fig. 6; Toriyama, width. Form ratio ranges from 1.97 to 2.51, 1958, p.211-212, pI. 39, figs. 20-25; Nogami, averaging 2.20 for 16 specimens. 1961, p.163-165, pI. I, figs. 7-11; Ishii and Shell expands gradually through growth Nogami, 1961, pI. 25, fig. 7 (same as pI. I, fig. 7 of with an almost straight axis of coiling. The Nogami, 1961, p. 163-165); Suyari, 1962, p.32, first volution spherical, but later ones become pI. 12, fig. 10; Hanzawa and Murata, 1963, p. 23, pI. 20, figs. 1-5; Sheng, 1963, p. 97-98,229-230, cylindrical rather rapidly. Radius vectors of pI. 29, figs. 1-9; Sheng, 1966, p. 143, pI. 26, fig. the first to thirteenth volutions of illustrated 4; Ozawa, 1970a, pI. 3, figs. 7-8; Tien, 1970, p. axial section (Figure 7-1) 0.11,0.17,0.21, 39-40, pI. 7, figs. 6-9, pI. 8, figs. 1-4; Toriyama 0.27,0.32,0.41,0.49,0.59,0.71,0.85,1.06,1.25 and Pitakpaivan, 1973, p. 53-57, pI. 6, figs. 6-13 ; and 1.41 mm, and form ratios 1.00,1.97,2.10, Zhang and Wang, 1974, p. 294, pI. 152, fig. 8 ; Sheng and Sun, 1975, p.50, pI. 14, figs. 1-2; 2.30,2.63,2.71,2.80,2.83,2.83,2.74,2.50,2.31 Ozawa, 1975, pI. 10, fig. 13; Toriyama and Kan- and 2.17, respectively. 1050 Katsumi Vena

13

15 Figure 7. I- tO. Pseudodoliolina pseudolepida "'4(Oeprat), 1-3,5- 8: axial sections,• ASM25170 , ASM25169, ASM25167, ASM25171 , ASM25166, ASM25168, ASM25172, 4,9 : sagittal sections, ASM25173, ASM25174, 10 : tangential section, ASM25175. 11 , 12. Colania douvillei (Ozawa), 11: diagonal section, ASM25213, 12 : axial section, ASM25212. 13- 15. Pseudodoliolina cf. ozawai Yabe and Hanzawa, 13: tangential section, ASM25177, 14 : oblique section, ASM25178, 15 : sagitta l section, ASM25176. All X 10. 934. Verbeekinids and neoschwagerinids from Akiyoshi 1051

Proloculus commonly spherical but rarely Subfamily Neoschwagerininae Dunbar irregular. Its outside diameter varies from and Condra, 1927 0.150 to 0.300 mm, averaging 0.211 mm for 22 specimens. Gen us Neosch wagerina Yabe, 1903 Spirotheca thin and composed of a single Type species. - Schwagerina craticulifera structureless layer in inner few vol utions, but Schwager, 1883. consists of a tectum and a lower, less dense layer in outer ones. Alveolar structure not Neosch wagerina craticulifera robusta observable even in outermost volution. Ueno, subsp. nov. Thickness of spirotheca of the first to thir­ teenth volutions of above-mentioned speci­ Figures 8-1-10, 10-6-11 men 0.010,0.005,0.010,0.010,0.010,0.015, Material. - Axial section of the holotype; 0.015,0.015,0.010,0.020,0.020,0.020 and ASM25179 from Loc. T A246. Axial sections 0.025 mm. of paratypes; ASM25180 from Loc. T A246, Septa numerous and slightly bending anter­ ASM25181 from Loc. AL98, ASM25182 from iorly. They consist of the same elements as Loc. ALIOO, ASM25183 from Loc. ALIOI, the spirotheca. ASM25184 from Loc. ALl08, ASM25185 Parachomata narrow and high, well devel­ from Loc. KS-AD7, ASM25186 from Loc. oped in all volutions, and attain about Yz to KS-AD9, ASM25187 from Loc. KS158, % of chamber height. ASM25188 from Loc. NS254. Sagittal sec­ Remarks. - Pseudodoliolina pseudolepida tions of paratypes; ASM25189 from Loc. (Deprat) is one of the most common species TA246, ASM25190 from Loc. TA247, of the genus and widespread in the Middle ASM25191 from Loc. T A250, ASM25192 Permian of the Tethyan province. The Aki­ from Loc. EB68, ASM25193 from Loc. yoshi specimens are quite identical with KSI55.5. Slightly oblique axial section of Deprat's original ones. paratype; ASM25194 from Loc. ALl03. This species is easily distinguished from Diagnosis. - Large Neoschwagerina having Pseudodoliolina ozawai Yabe and Hanzawa an inflated fusiform shell with bluntly by a larger shell and greater number of volu­ pointed polar ends, almost straight or slightly tions. convex lateral slopes, very small proloculus, Hasegawa et al. (1979) reported Pseudo­ and low, broad and fan-shaped primary trans­ doliolina sp. from the Verbeekina heimi Sub­ verse septula. Inner few volutions skew­ zone (equivalent to the Verbeekina verbeeki­ coiled, outer ones planispiral. Parachomata Afghanella schencki Zone in this study) in narrow and high. the Kaerimizu area. Judging from their Description. - Shell large for genus and illustrations, Pseudodoliolina sp. has a slight­ inflated fusiform with bluntly pointed polar ly smaller proloculus than P. pseudolepida ends. Lateral slopes almost straight or (Deprat). However, this difference is hardly slightly convex. Mature specimens having sufficient to warrant the specific separation of 16 to 19 volutions, 4.50 to 5.95 mm in length these forms. and 3.55 to 4.85 mm in width. Form ratio Occurrence. - Abundant in the Verbeekina ranges from 1.06 to 1.44, averaging 1.27 for 15 verbeeki-Afghanella schencki and Neo­ specimens. sch wagerina fusiformis Zones. Inner 2 or 3 volutions tightly and skew­ coiled. Beyond the third or fourth volution, shell expands gradually with a straight axis of Family Neoschwagerinidae Dunbar coiling. Radi us vectors of the first to seven­ and Condra, 1927 teenth volutions of the holotype 0.05,0.08, 1052 K alsumi Vena 934. Verbeekinids and neoschwagerinids from Akiyoshi 1053

0.12,0.17,0.22,0.29,0.39,0.51,0.67,0.77,0.91, morphological characters of Neoschwagerina 1.05,1.25,1.43,1.64, 1.83 and 1.97 mm, and craticulifera robust a, subsp. nov. suggest that form ratios 0.90, 1.03, 1.25, 1.24, 1.45, 1.62, the present new subspecies is phylogenetically 1.62, 1.57, 1.5 I, 1.58, 1.52, 1.49, 1.43, 1.43, 1.38, intermediate between N. craticulifera (s.s.) 1.36 and 1.38, respectively. and N. margaritae Deprat. Prolocul us very small and spherical. Its Measurements. - See Table I. outside diameter ranges from 0.040 to 0.090 Occurrence.-Abundant In the Neo­ mm, averaging 0.054 mm for 17 specimens. schwagerina craticulifera robusta Zone. Spirotheca thin to medium, composed of a tectum and alveolar keriotheca. Thickness Neoschwagerina fusiformis of spirotheca of the first to seventeenth vol u­ Skinner and Wilde, 1967 tions of the holotype 0.010,0.010,0.015, 0.015,0.015,0.030,0.040,0.065,0.055,0.030, Figures 9-1- 14 0.020, 0.030, 0.050, 0.020, 0.030, 0.020 and Neoschwagerina fusiformis Skinner and Wilde, 1967, 0.025 mm. Primary transverse septula low, p. 16-17, pI. 19, figs. 1-8; Ding, 1978, p. 288- broad and fan-shaped, and well developed 289, pI. 98, fig. 3; Wang et al., 1981, p. 64-65, pI. beyond the fourth volution. They are obser­ 18, figs. 1-2, II ; Chen, 1984, p.69, pI. 19, fig. 3 (same as pI. 98, fig. 3 of Ding, 1978, p. 288-289). ved immediately above each parachomata and abut tops of parachomata adjacent to Material. - Axial sections; ASM25195, septa. No secondary transverse septula ASM25196, ASM25197 from Loc. KN2, developed even in outermost volution. ASM25198, ASM25199A,B from Loc. KN4, Septa composed of downward deflections ASM25200 from Loc. KN21, ASM25201, of tectum and alveolar keriotheca. Septal ASM25202 from Loc. T A265, ASM25203A,B counts of the second to seventeenth vol utions from Loc. AL168. Sagittal sections; of typical sagittal section of para type (Figure ASM25204 from Loc. KN2, ASM25205 from 8-8) 3?,9, 14, 17,20, 19,22,21,26,26,25,26, Loc. KN5, ASM25206 from Loc. ALI80. 29, 30, 34 and 33, respectively. Axial septula Description. - Shell medium to large for present beyond the sixth volution. Two genus and fusiform to elongate fusiform with axial septula occasionally inserted between 2 bluntly pointed poles and almost straight to adjacent septa in outer volutions. slightly convex lateral slopes. Mature speci­ Parachomata high, rather narrow, and mens of 14 to 18 volutions 4.65 to 7.90 mm in begin to occur in the third volution. They length and 2.70 to 4.70 mm in width. Form attain a height about % that of chambers. ratio ranges from 1.43 to 1.90, averaging 1.65 Remarks. - Neosch wagerina cra ticulifera for 17 specimens. robusta, su bsp. nov. is closely similar to N. Axis of coiling straight through growth craticulifera (s.s.), especially in the nature of except for inner few volutions of some speci­ the primary transverse septula and para­ mens. Radius vectors of the first to fifteenth chomata, but differs from the latter in having vol utions of typical axial section (Figure 9- a larger shell, more vol utions and a smaller 12) 0.08, 0.11,0.13,0.18,0.24,0.29,0.35,0.42, form ratio. The present new subspecies also 0.51, 0.69, 0.79,0.97, 1.13, 1.29 and 1.46 mm, resembles Neoschwagerina margaritae De­ and form ratios 0.53,0.76, 1.15, 1.66, 1.49, prat, but differs from the latter in having a \.69,1.89, 2.02, 1.98, 1.71, 1.99, 1.84, 1.77, 1.78 smaller shell and a different shell shape. The and 1.75, respectively.

+---- Figure 8. 1-10. Neoschwagerina craticulifera robusta. subsp. nov., 6a: axial section of the holotype, ASM25179, 1,2,4, 5, 9, 10: axial sections of para types, ASM25184, ASM25180, ASM25181, ASM25188, ASM25186, ASM25182, 3,7,8: sagittal sections of paratypes, ASM25189, ASM25191, ASM25190, X 10, 6b : enlarged part of 6a, X 40. o V> +>-

Table 1. Measurements of Neoschwagel'ina craticulifera I'obusla, subsp. nov. F.R.: From ratio, D.P.: Diameter of proloculus (in mm). Radius vector Reg. no. Figure Length Width F.R. D.P. I 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 I ASM25179 8-6 5.69 4.20 1.35 0.060 0.05 0.08 0.12 0.17 0.22 0.29 0.39 0.51 0.67 0.77 0.91 1.05 1.25 1.43 1.64 1.83 1.97 - 2 ASM25180 8-2 5.53 3.85 1.44 0.065 0.06 0.09 0.12 0.16 0.21 0.28 0.37 0.49 0.61 0.77 0.92 1.10 1.29 1.47 1.65 1.86 - 3 ASM25181 8-4 5.50 4.05 1.36 - 0.05 0.07 0.12 0.17 0.22 CUI 0.40 0.52 0.65 0.81 0.95 1.11 1.27 1.49 1.68 1.88 - 4 ASM25183 10-9 5.60 4.35 1.29 0.050 0.04 0.07 0.10 0.13 0.19 0.25 0.34 0.44 0.57 0.72 0.89 1.04 1.22 1.40 1.55 1.78 1.99 2.19 5 ASM25184 8-1 5.10 4.05 1.26 0.050 0.05 0.08 0.11 0.15 0.21 0.30 0.40 0.52 0.64 0.75 0.88 1.08 1.27 1.46 1.64 1.82 2.03 6 ASM25187 10-10 5.18 4.00 1.30 0.045 0.05 0.06 0.10 0.13 0.17 0.25 0.34 0.46 0.57 0.70 0.83 1.00 1.15 1.31 1.46 1.71 1.92 - 7 ASM25186 8-9 5.15 4.85 1.06 0.045 0.05 0.08 0.11 0.15 0.20 0.29 0.42 0.53 0.67 0.81 0.95 1.12 1.29 1.46 1.67 1.86 2.18 2.39 2.52 - -- - "-----.- --

From ratio >:: I 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 s:::, c;:; I 0.90 1.13 1.25 1.24 1.45 1.62 1.62 1.57 1.51 1.58 1.52 1.49 1.43 1.43 1.38 1.36 1.38 - s::: 2 0.73 0.56 0.71 0.94 1.09 1.28 1.45 1.43 1.44 1.42 1.48 1.44 1.41 1.38 1.39 1.40 - ~ 3 - 1.28 1.25 1.47 1.52 1.56 1.63 1.52 1.57 1.60 1.56 1.58 1.52 1.47 1.48 1.44 - -. 4 0.60 0.64 0.85 1.15 1.33 1.58 1.66 1.78 1.66 1.58 1.46 1.40 1.34 1.36 1.40 1.36 1.21 - ~:::: 5 0.70 0.76 0.87 1.00 1.27 1.16 1.37 1.36 1.32 1.44 1.42 1.37 1.39 1.30 1.29 1.26 - Cl 6 0.60 0.77 0.75 1.00 1.28 1.31 1.31 1.28 1.34 1.33 1.30 1.28 1.25 1.24 1.27 1.24 1.23 - 7 0.70 0.75 0.87 1.13 1.28 1.50 1.35 1.40 1.37 1.32 1.31 1.23 1.21 1.20 1.18 1.17 1.10 1.09 - - ---

Thickness of spirothcca I 2 3 4 5 6 7 8 9 10 II 12 13 14 15 16 17 18 19 I 0.010 0.010 0.015 0.015 0.015 0.030 0.040 0.065 0.055 0.030 0.020 0.030 0.050 0.020 0.030 0.020 0.025 - 2 0.005 0.005 0.010 0.010 0.015 0.020 0.025 0.020 0.055 0.045 0.060 0.030 0.045 0.040 0.040 -- 3 0.005 0.005 0.010 0.015 0.015 0.020 0.020 0.030 0.030 0.030 0.080 0.040 0.030 0.070 0.045 0.040 - 4 0.005 0.005 0.010 0.010 0.015 0.015 0.025 0.040 0.050 0.040 0.040 0.040 0.040 0.030 0.030 0.075 0.035 0.035 5 0.005 0.005 0.010 0.010 0.020 0.025 0.030 0.025 0.060 0.055 0.060 0.025 0.030 0.025 0.030 0.020 CUl25 6 0.005 0.010 0.010 0.015 0.015 0.020 0.020 0.025 0.050 0.030 0.030 0.025 0.030 0.025 0.040 0.040 0.030 - 7 0.010 0.010 0.015 0.020 0.020 0.045 0.030 0.035 0.040 0.035 0.025 0.030 0.035 0.040 0.030 0.050 0.030 0.040 - 934. Verbeekinids and neoschwagerinids from Akiyoshi 1055

Proloculus small and spherical. Its outside schwagerina fusiformis Zone. diameter varies from 0.030 to 0.130 mm, aver­ aging 0.081 mm for 21 specimens. Neoschwagerina haydeni Dutkevich Spirotheca rather thin for genus and com­ and Khabakov, 1934 posed of a tectum and alveolar keriotheca. Thickness of spirotheca of the first to fifteenth Figures 10-1 - 5 volutions of specimen mentioned above 0.005, 0.005,0.005,0.010,0.025,0.030,0.030,0.025, Neoschwagerina craticulifera (Schwager). Hayden, 0.030,0.030,0.025,0.025,0.025,0.035 and 1909, p. 248-249, pI. 21, figs. 1-7. Neoschwagerina craticulifera haydeni Dutkeyich and 0.040 mm. Primary transverse septula thin Khabakoy, 1934, p.94-99, pI. 2, figs. 6-8, pI. 3, and well developed beyond the third volu­ figs. 1-2; Thompson, 1946, p.155-156, pI. 23, tion. They occur immediately above each figs. 12-13; Thompson, 1948, pI. 20, figs. 3-4 parachomata and abut tops of parachomata (same as pI. 23, figs. 12-13 of Thompson, 1946); Kanmera, 1957, pI. 19, fig. 24 (same as pI. 23, fig. adjacent to septa. One secondary transverse 12 of Thompson, 1946); Toriyama, 1958, p. 220- septulum rarely and rather inconstantly inser­ 223, pI. 41, figs. 7-8 (not pI. 41, fig. 6); Suyari, ted between 2 adjacent primary transverse 1962, p. 35-36, pI. 11, fig. 5; Ishizaki, 1962, p. septula. 173-175, pI. 12, figs. 4-6. Neoschwagerina haydeni Dutkeyich and Khabakoy. Septa thin, bending anteriorly and com­ Honjo, 1959, p. 147-149, pI. 3, fig. 10; Minato posed of downward deflections of spirothecal and Honjo, 1959, pI. 3, fig. 2; Sheng, 1963, p. element. Septal counts of the third to six­ 103-104,236-237, pI. 32, figs. 1-8; Zhang and teenth volutions of typical sagittal section Wang, 1974, p. 292, pI. 151, fig. 7; Chen and (Figure 9-2) 12, 14, 15, 15, 16, 17, 24, 23, 27, Yang, 1978, p. 116, pI. 32, figs. 3-5; Liu et al., 1978, p. 89, pI. 21, fig. 6; Toriyama and 27, 30, 28, 30? and 28? Axial septula begin Kanmera, 1979, p.77-80, pI. 12, figs. 19-22, pI. to occur in the fifth vol ution. One axial 13, figs. 1-8; Hasegawa et al., 1979, pI. 84, figs. septulum inserted between 2 adjacent septa in 1-8; Wang et al., 1981, p. 63, pI. 15, figs. 1-2,5, inner volutions, but usually 2, rarely 3 in 7; Wang et al., 1982, p. 106, pI. 30, fig. 7; Sun et al., 1983, p. 39, pI. 10, fig. 13; Lin, 1984, p. outer ones. 171-172, pI. 14, fig. 12; Chen, 1984, p. 69, pI. 21, Parachomata narrow and high, reaching fig. 2; Yang, 1985, pI. 2, fig. 12; Kobayashi, about % the height of chambers. 1988b, p.445-447, figs.7 - 2-11; Ozawa and Remarks. - Neoschwagerina fusiformis was Kobayashi, 1990, pI. 10, figs. 12-14. originally described by Skinner and Wilde Neoschwagerina cf. haydeni Dutkeyich and Khaba­ koy. Ding, 1978, p. 289, pI. 101, fig. 2. (1967) from the Permian of Tunisia. The holotype specimen (Skinner and Wilde, 1967, Material. - Axial sections; ASM25207 pI. 19, fig. I) has a minute proloculus and from Loc. ALl7S, ASM2520S from Loc. skew-coiled inner volutions, and is consid­ ALISO, ASM25209 from Loc. KN21, ered to be a microspheric form of the species. ASM25210 from Loc. SAI4S. Sagittal sec­ Most of the Akiyoshi specimens are quite tion; ASM25211 from Loc. KN 19. similar to one of the paratypes (Skinner and Description. - Shell medium to large for Wilde, 1967, pI. 19, fig. 2). genus and thickly fusiform with rounded This species somewhat resembles Neo­ polar ends and almost straight lateral slopes. schwagerina haydeni Dutkevich and Khaba­ Mature specimens having 15 to 17 volutions, kov in its essential morphological characters, 5.10 to 6.00 mm in length and 3.S0 to 4.15 mm especially in the nature of septula and para­ in width, giving form ratios of 1.31 to 1.54. chomata. However, the former can be distin­ Axis of coiling straight except for inner few guished from the latter in having a more volutions of some specimens. Shell expands elongate shell and larger form ratio. gradually through growth. Radius vectors of Occurrence. - Abundant In the Neo- the first to sixteenth vol utions of illustrated 1056 Katsumi Vena 934. Verbeekinids and neoschwagerinids from Akiyoshi 1057 axial section (Figure 10-1) 0.06,0.10,0.14, Subfamily Lepidolininae 0.20,0.26,0.35,0.43,0.52,0.63,0.76,0.88, 1.03, Miklukho-Maklay, 1958 1.16, 1.31, 1.46 and 1.66 mm, and form ratios 0.58,0.74,0.81, 1.10, 1.29, 1.31, 1.53, 1.63, 1.66, Genus Colania Lee, 1934 emend. 1.61, 1.59, 1.53, 1.54, 1.58, 1.53 and 1.50, Ozawa, 1970 respectively. Type species. - Colania kwangsiana Lee, Proloculus small and spherical, being 0.060 1934. to 0.070 mm in outside diameter averaging Colania douvillei (Ozawa, 1925) 0.063 mm for 3 specimens. Spirotheca thin for genus and composed of Figures 7-11-12 a tectum and alveolar keriotheca. Thickness Neoschwagerina globosa (Yabe). Douville, 1906, p. of spirotheca of the first to sixteenth volutions 182, pI. 17, pI. 18, figs. 1-2; Deprat, 1912, p.51, of above-mentioned specimen 0.005, 0.005, pI. 4, figs. 1-2 (not pI. 4, figs. 3-4); Deprat, 1913, p. 55; Deprat, 1914, p. 29-30; Ozawa, 1922, p. 0.010,0.015,0.015,0.020,0.015,0.020,0.020, 368-372; Colani, 1924, p. 152-153, pI. 23, figs. 0.035,0.035,0.030,0.025,0.025,0.025 and 1-2,4-14,22-34,36-38, pI. 25, figs. 9, 13, pI. 26, 0.030 mm. Thin primary transverse septula figs. 3-5. first appear in the third or fourth vol uti on. Neosch wagerina douvillei Ozawa, 1925a, pI. 3, fig. 6, pI. 4, fig. 5; Ozawa, 1925b, p.55-57, pI. II, They develop immediately above each para­ figs. 5-7; Huzimoto, 1936, p.114-115, pI. 23, chomata and abut tops of parachomata adja­ figs. 1-5; Toriyama, 1947, p. 78-79, pI. 17, fig. 8 ; cent to septa. One secondary transverse se­ Chen, 1956, p. 12,58-59, pI. 13, figs. 3-4, 6-7 (not ptulum sometimes inserted between 2 adja­ pI. 13, fig. 5, pI. 14, fig. 7); Toriyama, 1958, p. 223-227, pI. 41, figs. 9-13, pI. 42, figs. 1-6; Sa­ cent primary ones. kagami, 1958, p.92-93, pI. 14, figs. 7-10; Septa thin, bending anteriorly and com­ Kanuma, 1960, p. 70-71, pI. 13, figs. 5-7 ; posed of downward deflections of tectum and Nogami, 1961, p. 180-183, pI. 5, figs. 1-5; Sada, alveolar keriotheca. Axial septula begin to 1961, p. 123-125, pl. 12,figL 1-8; Sheng, 1963, p. occur in the fifth volution. Beyond the sev­ 102-103,235-236, pI. 33, figs. 1-7; Hanzawa and Murata, 1963, pI. 4, figs. 3-4, pI. 17, figs. 1-10; enth volution, 2 axial septula commonly Sheng, 1966, p. 146, pI. 25, fig. I ; Sheng and Sun, observed. Three axial septula rarely inserted 1975, p.51-52, pI. 13, figs. 11-14; Lin el al., between 2 adjacent septa in outer few volu­ 1977, p. 92, pI. 28, fig. 12; Liu el al., 1978, p. 89, tions. pI. 24, fig. I ; Han, 1980, p. 90, pI. 34, figs. 13-15 ; Xie, 1982, p. 69, pI. 34, fig. I ; Wang et al., 1982, Parachomata narrow and high adjacent to p. 106, pI. 30, fig. 5. septa, but low and semicircular in central part Neosch wagerina cf. douvillei Ozawa. Kawano, 1960, of chamber. pI. 25, figs. 12-13; Kawano, 1961, p. 107-108, pI. Remarks. - Neoschwagerina haydeni Out­ II, figs. 2-5; Yoshimura, 1961, pI. 3, fig. 4 ; kevich and Khabakov somewhat resembles N. Suyari, 1962, p. 36, pI. 10, fig. 14. Colania douvillei (Ozawa). Ozawa, 1970a, pI. 7, figs. craticulifera robusta, subsp. nov., described 8-10; Choi, 1972, p.371-372, pI. 46, figs. 1-6; above, in shell shape. However, the former Toriyama, 1975, p. 103, pI. 20, figs. 22-26; Tor­ can be easily distinguished from the latter in iyama and Kanmera, 1977, p. 17-24, pI. 2, figs. 7- having a thinner spirotheca, more slender 17, pI. 3, figs. 1-16; Ota, 1977, pI. I, figs. 3-4; Sakagami and Miyama, 1988, fig. 1-15; Kobaya­ transverse septula and parachomata. shi, 1988a, p. 13, pI. 7, figs. 7-9, pI. 8, figs. 1-8 ; Occurrence. - Common in the Neo­ Ozawa and Kobayashi, 1990, pI. II, figs. 1-2. schwagerina fusiformis Zone. Gifuella douvillei (Ozawa). Hasegawa el al., 1979,

+-- Figure 9. 1-14. Neoschwagerina fusiformis Skinner and Wilde, 1,3-7,9-11, 12a, 13: axial sections, ASM25203A, ASM25198, ASM25196, ASM25195, ASM25197, ASM25200, ASM25203B, ASM25201, ASM25199A, ASM25199B, ASM25202, 2,8,14: sagittal sections, ASM25204, ASM25205, ASM25206, X 10, 12b: enlarged part of 12a, X 40. 1058 Katsumi VenD

Figure 10. 1- 5. Neoschwagerina haydeni Dutkevich and Khabakov, 1,2,4,5: ax ia l sectio ns, ASM25209, ASM25208, ASM2521O, ASM25207, 3 : sagitta l section, ASM252 ll. 6- 11. Neoschwagerina craticulifera robusta, subsp. nov., 7,9, 10 : axial sections of para types, ASM25l85, ASM25l83, ASM25 l87 , 6, 11: sagittal sections of paratypes, ASM25 l93, ASM25 l92, 8: sli ghtly oblique axial secti o n of paratype, ASM25l94. A ll X 10. 934. Verbeekinids and neoschwagerinids from Akiyoshi 1059

pI. 86, figs. 1-6. Taishaku, Atetsu and Omi. The present not Neoschwagerina dOllvillei Ozawa. Gubler, 1935, p. I I I-I 13, pI. 6, fig. 2, pI. 7, figs. 7-8,10-1 I, pI. specimens agree closely with ones reported 8, fig. 6; Ishii, 1966, pI. 5, fig. 4; Igo, 1967, pI. 7, from these limestone plateaus. figs. 6-8, pI. 8, fig. 7; Tien, 1970, p. 48-49, pI. I I, Occurrence. - Rare in the Colania douvillei figs. I - I 5, pI. 12, figs. 1-13 ; Wang et aI., 198 I, p. Zone. 62-63, pI. 18, fig. 5; Yang, 1985, pI. 2, fig. 14. not Colania dOllvillei (Ozawa). Ozawa, 1970b, p. 35-38, pI. 3, figs. 1-10, pI. 4, figs. 1-10, pI. 5, figs. Subfamily Sumatrininae Silvestri, 1933 1-5, pI. 6, figs. 1-9; Tien, 1979, p. 137- 139, pI. 22, figs. 1- 10, pI. 23, figs. 1-2; Tien, 1986a, pI. 8, Genus Afghanella Thompson, 1946 fig. 2 (same as pI. 22, fig. 2 of Tien, 1979); Tien, 1986b, pI. I I, figs. 1-6, 9A; Tien, 1989, pI. 33, Type species. - Afghanella schencki figs. 1-6(2-3: same as pI. I I, figs. 5, I of Tien, Thompson, 1946. 1986b). not? Neoschwagerina dOllvillei Ozawa. Chen and Hanzawa, 1954 Yang, 1978, p. 117, pI. 32, figs. 7-9. Afghanella ozawai Figures I I-I - 17 Material. - Axial section; ASM25212. Diagonal section; ASM25213. Both from Neoschwagerina (Yabeina) schellwieni (Deprat). Ozawa, 1925a, pI. 2, figs. 8- 10. Loc. EBIII. Yabeina schellwieni (Deprat). Ozawa, 1925b, p.60- Description. - The following description is 61, pI. 10, figs. 3a, 4 (4: same as pI. 2, fig. 9 of based on one axial section (Figure 7-12). Ozawa, 1925a). Shell large and inflated fusiform with Cancellina schellwieni (Deprat). Ozawa, 1927, p. broadly rounded polar ends and straight axis 161, pI. 34, fig. 18 (same as pI. 2, fig. 9 of Ozawa, 1925a), pI. 44, fig. I b, pI. 45, fig. 3. of coiling. Length and width of shell having Afghanella ozawai Hanzawa, 1954, p. 3-7, pI. I, figs. 13 volutions, 6.80 mm and 4.55 mm, respec­ 1-6, pI. 2, figs. 1-4; Kanmera, 1957, pI. 19, figs. tively. Form ratio about 1.50. Shell 12-13; Toriyama, 1958, p.252-254, pI. 48, figs. expands gradually through growth. Radius I 1,13-14 (not pI. 48, figs. 12, 15-17), Kawano, 196 I, p. I 17- I 19, pI. 15, figs. 3-11 ; Hanzawa and vectors of the first to twelfth volutions 0.22, Murata, 1963, pI. 2, figs. 1-2; Ozawa, I 970a, pI. 0.33,0.44,0.54,0.67,0.79,0.96,1.16,1.36, 1.56, 6, figs.6-7 (same as pI. 19, figs.12-13 of 1. 74 and 1.95 mm, and form ratios 0.91, 1.15, Kanmera, 1957); Lin et al., 1977, p. 95, pI. 30, 1.22,1.28,1.37,1.48,1.52,1.48,1.57, 1.49, 1.53 fig. 3; Hasegawa et al., 1979, pI. 82, figs. I I - I 6 ; and 1.51, respectively. Prolocul us somewhat Xie, 1982, p. 72, pI. 35, fig. 6 (same as pI. 30, fig. 3 of Lin et al., 1977, p. 95); Kobayashi, 1988a, irregular, being 0.210 mm in outside diameter. p. 14, pI. 5, figs. 22-25; Ozawa and Kobayashi, Spirotheca thin and composed of a tectum 1990, pI. 10, figs. 1-2. and alveolar keriotheca. Thickness of spiro­ Afghanella schencki Thompson. Toriyama, 1958, p. theca of the first to twelfth volutions 0.015, 250-25 I, pI. I?, 2,4-5, 10 (not pI. 48, figs. 3,6-9) ; Ota, 1977, pI. 2, figs. 1-2. 0.030, 0.020, 0.025, 0.020, 0.025, 0.040, 0.030, 0.025,0.035,0.035 and 0.040 mm. Primary Material. - Axial sections of megalospheric transverse septula thin and begin to appear in form; ASM25214 from Loc. EB49, the second volution. No secondary trans­ ASM25215, ASM25216 from Loc. EB50, verse septula present. Parachomata narrow ASM25217, ASM25218 from Loc. EB56, and high, meeting the tops of primary trans­ ASM25219, ASM25220 from Loc. T A231, verse septula adjacent to septa. ASM25221 from Loc. MT210, ASM25222 Remarks. - Colania douvillei (Ozawa) is from Loc. NS220, ASM25223 from Loc. widespread in the Middle Permian rocks in SF49, ASM25224A from Loc. KS-AD24. the eastern part of the Tethyan province, and Sagittal sections of megalospheric form; also in the huge limestone masses in the Inner ASM25224B from Loc. KS-AD24, ASM25225 Zone of southwest Japan, such as Akiyoshi, from Loc. SF33, ASM25226 from Loc. AL92, 1060 Katsumi Vena 934. Verbeekinids and neoschwagerinids from Akiyoshi 1061

ASM25227 from Loc. EB49. Axial sections of septa. One secondary transverse septulum microspheric form; ASM25228 from Loc. inserted between 2 adj acent primary trans­ SA56, ASM25229 from Loc. AL90. verse septula in outer volutions. Description. - Shell small to medi urn for Septa consist of downward deflections of genus and inflated fusiform to fusiform with tectum and anterior and posterior extensions bl untly pointed poles. Dimorphism well of keriotheca. Lower half of septa usually observed. Megalospheric specimens having solidified and pendant-shaped in cross sec­ 7 to 9 volutions, 2.55 to 4.20 mm in length tion. Septal counts of the first to eighth and 1.53 to 2.53 mm in width. Form ratio volutions of one illustrated sagittal section ranges from 1.38 to 1.99, averagi ng 1.67 for 20 (Figure 11-4) 8, 18,23,23,24,26, 28? and specimens. Microspheric form of 9 to 10 31? Axial septula begin to appear in the volutions 2.95 to 4.55 mm in length and 1.40 second or third volution. Two axial septula to 2.30 mm in width, giving form ratios of usually inserted between 2 adjacent septa. 1.98 to 2.11. Parachomata massive and triangular in trans­ Axis of coiling straight throughout in verse section, rarely narrow and high, and megalospheric form. Inner 2 or 3 volutions well developed in all volutions. tightly and skew coiled in microspheric form. Remarks. - Small shell size and the nature Radius vectors of the first to seventh volutions of axial and transverse septula in Afghanella of well oriented axial section of megalospher­ ozawai Hanzawa indicate that it is one of the ic form (Figure 11-2) 0.15,0.22,0.32,0.44, most primitive species in this genus. The 0.58, 0.73 and 0.92 mm, and form ratios 1.40, stratigraphic position of the present species 1.64, 1.62, 1.66, 1.72, 1.67 and 1.65, respective­ also supports this conclusion. ly. Afghanella ozawai Hanzawa somewhat Proloculus of megalospheric form large for resembles A. robbinsae and A. african a, both shell size, spherical to somewhat irregular, described by Skinner and Wilde (1967) from being 0.165 to 0.320 mm in outside diameter, the Permian of Tunisia. However, the for­ averaging 0.239 mm for 49 specimens. mer differs from the latter two in having a Proloculus of microspheric form minute and thicker spirotheca and fewer axial septula spherical, being 0.030 to 0.040 mm in outside between 2 adjacent septa. diameter, averaging 0.036 mm for 5 speci­ Occurrence. - Abundant in the Afghanella mens. ozawai Zone. Spirotheca thin to medium, composed of a tectum and alveolar keriotheca. Thickness of Afghanella schencki Thompson, 1946 spirotheca of the first to seventh vol utions of Figures 12-1 - 11 above-mentioned specimen 0.010,0.025, 0.040,0.025, 0.070?, 0.050 and 0.030 mm. Afghanella schencki Thompson, 1946, p. 153-155, pI. Primary transverse septula developed in all 25, figs. 1-12; Thompson, 1948, pI. 18, figs. 1-5 (same as pI. 25, figs. 1-3,7,10 of Thompson, volutions. Their tips slightly swollen. 1946, p. 153-155); Chen, 1956, p. 14,67-68, pI. Lower edges of primary transverse septula in 12, figs. 4-9; Toriyama, 1958, p. 250-251, pI. 48, contact with tops of parachomata adjacent to figs. 3, 6-9 (not I?, 2, 4-5,10); Sheng, 1958, p.

+-- Figure 11. 1-17. Afghanella ozawai Hanzawa, 1-3,5,7,9,10,12-15: axial sections of megalo­ spheric form, ASM25217, ASM25215, ASM25224A, ASM25218, ASM25220, ASM25214, ASM25219, ASM25216, ASM25221, ASM25222, ASM25223, 4,6,8,11: sagittal sections of megalospheric form, ASM25226, ASM25225, ASM25227, ASM25224B, X 10, 16a, 17: axial sections of microspheric form, ASM25229, ASM25228, X 20, 16b: enlarged part of 16a showing minute proloculus and skew coiled juvenile volutions, X 100. 18-20: Afghanella sp., tangential sections, ASM25230, ASM25231, ASM25232, X 10. 1062 Katsumi Vena

Figure 12. 1- 11. Afghanella schencki Thompson, 1,2,4- 8,10,11 : axial sections, ASM25238, ASM25235A, ASM25235B, ASM25234, ASM25240, ASM25236, ASM25237, ASM25239, ASM25233, 3,9: sagittal sections, ASM25242, ASM25241. 12- 16. Afghanella cf. sumatrinaeformis (Gubler), 12- 15: axial sections, ASM25244, ASM25245, ASM25246, ASM25243, 16: sagittal section, ASM25247. All X 10.

277, 289- 290, pI. 4, figs. 9- 10 ; Sheng, 1963 , p. 7, 9- 10 ; Yang, 1985, pI. 3, fig. 21 ; Loeblich and 109-110, 244-245, pI. 36, fig. 20 ; Pasini, 1965 , pI. Tappan, 1988, pI. 306, figs. 3- 6 (same as pI. 25, 17, fig. l a, Ib (same as pI. 25, fig. 12, 7, ofThomp­ figs. 2- 3, 6- 7 of Thompson, 1946, p. 153- 155) ; son, 1946, p. 153-155); Sheng, 1966, p. 151 , pI. Ozawa and Kobayashi, 1990, pI. 10, figs. 5- 7. 26, fig. 6; Leven, 1967, p. 198, pI. 36, figs. 4, 7- 8, Afghanella ozawai Hanzawa. Toriyama, 195 8, p. pI. 37, fig . 3; Zhang and Wang, 1974, p. 291 , pI. 252-254, pI. 48 , figs. 12, 15 - 17 (not pI. 48, figs. II , 151 , fig. I ; Rozovskaya, 1975, pI. 33, figs. 5- 7 13- 14). (same as pI. 25, figs. 2- 3, 7 of Thompson, 1946, p. Afghanella schencki schencki Thompson. Tor­ 153- 155); Lin et al., 1977, p.95, pI. 30, fig. 4 ; iyama and Kanmera, 1979, p. 58 - 61 , pI. 9, figs. 8- Chen and Yang, 1978, p. 121 , pI. 34, figs. 3- 4 ; 24. Liu el al., 1978, p. 96, pI. 23, fig. 4; Kahler and Kahler, 1979, p. 251 - 252, pI. 8, fig. 5, pI. 9, fig. 2 ; Material. - Axial sections; ASM25233 Hasegawa et al., 1979, pI. 82, figs. 5- 10; Da and Sun, 1983, p. 113, pI. 30, fig. 3 ; Sun et al., 1983, from Loc. MT240, ASM25234 from Loc. p. 39- 40, pI. 10, fig. 5 ; Chen, 1984, p. 71 - 72, pI. MT243, ASM25235A,B from Loc. SA 119, 22, fig. I ; Sun and Zhang, 1985, p. 506, pI. I, figs. ASM25236 from Loc. SAI20, ASM25237 934. Verbeekinids and neoschwagerinids from Akiyoshi 1063

from Lac. SA 130, ASM25238 from Lac. the Bamian Limestone of Afghanistan. The SF69, ASM25239 from Lac. T A254, proloculus in the Akiyoshi specimens is ASM25240 from Lac. EB97. Sagittal sections; slightly smaller than that in the original ones. ASM25241 from Lac. SA 130, ASM25242 Afghanella schencki Thompson can be from Loc. MT239. easily distinguished from A. ozawai Hanzawa Description. - Shell small to medi urn for in having a more elongate shell, more volu­ genus and elongate fusiform with bluntly tions, smaller prolocul us, thinner spiro theca pointed polar regions and almost straight to and more developed primary transverse and slightly convex lateral slopes. Mature shell axial septula. The stratigraphic distribution of 9 to 11 volutions 3.33 to 4.55 mm in length of these two species is also different. and 1.50 to 2.35 mm in width. Form ratio Occurrence. - Abundant in the Verbeekina ranges from 1.74 to 2.37, averaging 1.96 for 18 verbeeki-Afghanella schencki Zone. specimens. Shell expands gradually through growth. Afghanella cf. sumatrinaeformis Axis of coiling straight throughout. Radius (Gubler, 1935) vectors of the first to ninth vol utions of typi­ cal axial section (Figure 12-1) 0.12, 0.16, 0.25, Figures 12-12-16 0.33, 0.43, 0.55, 0.68, 0.83 and 0.99 mm, and Compare- form ratios 1.48,1.81,1.80, 1.85,2.00, 1.96, Neaschwagerina annae (Volz). Hayden, 1909, p. 1.96, 1.94 and 1.86, respectively. 250-251, pI. 22, figs. 8-14. Prolocul us small and spherical. Its out­ Neascll\vagerina suma/rinae/armis Gubler, 1935, p. side diameter varies from 0.070 to 0.195 mm, 123-127, pI. 5, figs. 3-4,10,17, pI. 7, fig. 2. averaging 0.121 mm for 19 specimens. Suma/rina suma/rinae/armis (Gubler). Ciry, 1941, Spirotheca thin and consists of a tectum pI. I, figs. 8-9. A/ghanella sumalrinae/armis (Gubler). Kochans­ and probably very thin alveolar keriotheca. ky-Devide and Ramov5, 1955, p.391-392,417, Thickness of spirotheca of the first to ninth pI. 5, figs. 1-10; Chen, 1956, p.14,68-69, pI. 7, vol utions of specimen mentioned above 0.0 15, figs. 9-11 ; Kanmera, 1957, pI. 19, figs. 16-17 0.025, 0.020, 0.015, 0.025, 0.015, 0.020, 0.025 (same as pI. 5, figs. 3-4 of Gubler, 1935, p. 123- 127); Sheng, 1966, p. 152, pI. 22, fig. II (same as and 0.025 mm. pI. 7, fig. 9 of Chen, 1956, p. 14, 68-69); Leven, Primary transverse septula very thin and 1967, p. 198-199, pI. 37, figs. 1-2; Ozawa, 1970a, developed in all volutions. Tips of primary pI. 6, figs. 10-11 (same as pI. 5, figs. 3-4 of Gub­ transverse septula slightly swollen and in ler, 1935, p. 123-127); Tien, 1970, p. 55, pI. 13, contact with tops of parachomata adjacent to figs. 9-10, pI. 14, figs. 5-9; Rozovskaya, 1975, pI. 33, figs. 8-9 (same as pI. 5, figs. 3-4 of Gubler, septa. One secondary transverse septul urn 1935, p. 123-127); Toriyama and Kanmera, inserted between 2 adjacent primary trans­ 1979, p.63-68, pI. 10, figs. 14-24; Yang, 1985, verse septula in outer 3 or 4 volutions. pI. 3, fig. 22; Loeblich and Tappan, 1988, pI. Septa thin and bending anteriorly. Tips of 306, figs. 7-9 (same as pI. 5, figs. 3-4, 10 of Gub­ ler, 1935, p. 123-127). septa solidified and pendant-shaped. Septal A/ghanella cf. schencki Thompson. Kanmera, 1957, counts of the first to tenth volutions of illust­ pI. 19, figs. 14-15. rated sagittal section (Figure 12-3) 9,14,16, 18,16,18,21,19,21 and 26, respectively. Material. - Axial sections; ASM25243 Axial septula first appear in the fifth volution. from Lac. T A256, ASM25244 from Lac. Two, rarely 3 axial septula observed in outer SA 122, ASM25245 from Loc. SA 124, 3 or 4 volutions. Parachomata narrow and ASM25246 from Loc. MT238. Sagittal sec­ high. tion; ASM25247 from Loc. MT239. Remarks. - Afghanella schencki was origi­ Description. - Shell medi urn to large for nally described by Thompson (1946) from genus and thickly fusiform with bluntly 1064 Katsumi Ueno pointed to rounded axial regions and slightly specific identification is postponed until bet­ convex lateral slopes. Mature specimens hav­ ter material is accumulated. ing 11 to 13 volutions, 4.05 to 4.98 mm in Occurrence. - Rare in the lower part of the length and 2.58 to 3.50 mm in width, giving Afghanella ozawai Zone. form ratios of 1.26 to 1.53. Shell expands gradually through growth. Proloculus small Acknowledgments to medium, being 0.090 to 0.175 mm in out­ side diameter, averaging 0.124 mm for 4 speci­ I would like to express my deep apprecia­ mens. Spirotheca thin and composed of a tion to Prof. Hisayoshi Igo of the University tectum and thin alveolar keriotheca. Pri­ of Tsukuba for critical reading of the manu­ mary transverse septula observed in all volu­ script. Thanks are also extended to Prof. tions. Tips of primary transverse septula Sumio Sakagami ofChiba University, Messrs. solidified and pendant-shaped, joining with Akihiro Sugimura and Takehiko Haikawa of tops of parachomata. One, rarely 2 secon­ the Akiyoshi-dai Museum of Natural History dary transverse septula inserted between 2 for their contin uous advice and encourage­ adjacent primary transverse septula. Septa ment, and Mr. Yasuyuki Miyama of thin but usually coated with dense calcareous Sumitomo Cement Co., Ltd. for fruitful dis­ material. Tips of septa slightly swollen. One cussion on fusulinacean taxonomy. axial septulum usually occurs between 2 adja­ cent septa beyond the fifth volution, but 2 axial septula rarely inserted in outer ones. References cited Parachomata prominent and well developed Chen, G.X., 1984: Protozoa. In, Regional Geol. in all volutions. Surv. Team, Hubei ed., The Palaeontological Remarks. - The present species quite agrees Atlas of Hubei province, p.12-73, pis. 1-22. with Afghanella sumatrinaeformis (Gubler) Hubei Sci. Tech. Press, Wuhan. (in Chinese) in the essential shell characters except for Chen, J.R. and Yang, Z.R., 1978: Fusulinida. In, Xinan Inst., Geol. Sci. chiefly ed., having a smaller proloculus. Paleontological Atlas of Southwest China, Si­ Afghanella cf. sumatrinaeformis (Gubler) chuan vol., 2, p. 17-123, pis. 3-34. Geol. Publ. differs from A. schencki Thompson in having House, Beijing. (in Chinese) a more thickly fusiform shell, smaller form Chen, S., 1934: Fusulinidae of South China, Part I. ratio and more volutions. Palaeont. Sinica, ser. B, vol. 4, fasc. 2, p. 1-185, pis. 1-16. Occurrence. - Common in the Verbeekina --, 1956: Fusulinidae of South China, Part II. verbeeki - Afghanella schencki Zone. Palaeont. Sinica, N.S., ser. B, no. 6, p. 1-71, pis. 1-14. Afghanella sp. Choi, D.R., 1970: Permian fusulinids from Imo, southern Kitakami Mountains, N.E. Japan. Figures 11-18-20 Jour. Fac. Sci., Hokkaido Univ., ser. 4, vol. 14, no. 3, p. 327-354, pis. 9-15. Material. -Tangential sections --, 1972: Colania douvillei (Ozawa), a fusulinid ASM25230 from Loc. KS 175, ASM25231 foraminifera from the northern Kitakami Moun­ from Loc. AL89, ASM25232 from Loc. EB56. tains, N.E. Japan. Trans. Proc. Palaeont. Soc. Japan, N.S., no. 86, p. 369-374, pI. 46. Remarks. - Three tangential sections are Ciry, R., 1941: Les fusulinides de Turquie. Ann. referable to Afghanella sp. This uniden­ Paleont., vol. 29, p. 53-78, pI. I. tified species resembles Afghanella ozawai Colani, M., 1924: Nouvelle contribution a l'etude Hanzawa, which coexists with it, in the nature des fusulinides de I'Extreme-Orient. Mem. Servo of the transverse septula. However, the for­ Geol. l'lndochine, vol. II, fasc.l, p. 1-191, pis. 1- 29. mer has a larger and more elongate shell than Da, Y.T. and Sun, Q.L., 1983: Fusulinida. In, those of the latter. At any rate, the exact Regional Geol. Surveying Team of Xinjiang et 934. Verbeekinids and neoschwagerinids from Akiyoshi 1065

al. chiefly eds., Paleontological A tlas of North­ of some fusulinid foraminifera from the Kitakami west China, Xinjiang Weiwuer Aut. Region, 2, massif, Japan. Sci. Rep., Tohoku Univ., 2nd (Late Paleozoic), p. 7-113, pis. 1-30. Geol. Publ. ser. (Ceol.), vol. 35, no. I, p. 1-31, pis. 1-20. House, Beijing. (in Chinese) Hasegawa, Y., 1958: Note on the geological structure Deprat, J., 1912: Etude des fusulinides de Chine et of the Akiyoshi Plateau, S.W. Honshu. Earth d'Indochine et classification des calcaires a Science (Chikyu Kagaku), no. 39, p. 15-18. (in fusulines. Mem. SeT!'. Ceol. I 'In dochine, vol. I, Japanese with English abstract) fasc. 3, p. 1-76, pis. 1-9. --, 1963: New find of fossils in the reddish --, 1913: Etude des fusulinides de Chine et tuffaceous shale in the Akiyoshi province. d'Indochine et classification des calcaires a Ibid., no. 64, p.32-36. fusulines (lie Memoir). Les fusulinides des cal­ --, Kato, M., Nakamura, K. and Minato, M., 1979: caires carboniferiens et permiens du Tonkin, du Permian, Appendix. In, Minato, M., Hunaha­ Laos et du Nord-Annam. Ibid., vol. 2,. fasc. I, shi, M., Watanabe, J. and Kato, M. eds., The p. 1-74, pis. 1-10. Abean Orogeny, p. 184-191, pIs. 73-90. Tokai --, 1914: Etude des fusulinides de Chine et Univ. Press. d'Indochine et classification des calcaires a Hayden, H.H., 1969: Fusulinidae from Afghanistan. fusulines (Ille Memoir). Etude comparative des Rec. Ceol. SUT!'. India, vol. 38, p.230-257, pIs. fusulinides d'Akasaka (Japon) et des fusulinides 17-22. de Chine et d'Indochine. Ibid., vol. 3, fasc. I, p. Honjo, S., 1959: Neoschwagerinids from the Aka­ 1-45, pis. 1-8. saka Limestone (A paleontological study of the Ding, Q.X., 1978: Permian Fusulinaceans. In, Akasaka Limestone, I st report). Jour. Fac. Sci., Hubei Inst. Geol., Working Group of Stratigra­ Hokkaido Univ., ser. 4, vol. 10, no. 1, p. 111-161, phy of Sanxia ed., Stratigraphy and Paleon­ pIs. 1-12. tology of the Sinian to the Permian in Xiadong Huzimoto, H. (Fujimoto, H.), 1936: Stratigraphical district, p.277-290, pls.96-101. Geol. Publ. and palaeontological studies of the Titibu system House, Beijing. (in Chinese) of the Kwanto-mountainland, Part 2. Douville, M.H., 1906: Les calcaires a fusulinides de Palaeontology. Sci. Rep., Tokyo Bunrika l'Indochine. Bull. Soc. Ceol. France, ser. 4, vol. Daigaku, sect. C, vol. I, no. 2, p. 29-125, pIs. 1- 6, p. 576-587, pis. 17-18. 26. Dutkevich, G.A. and Khabakov, A.V., 1934: Perms­ Igo, H., 1964: Permian fossils from northern Pahang, kie otlozheniya vostochnogo Pamira i paleo­ Malaya. Japan. Jour. Ceo!. Ceogr., vol. 35, no. geografiya verkhnego Paleozoya tsentral'noy I, p. 55-71, pI. 2. Azii. Trudy Tadzhikskogo Kompleksnogo --, 1967: Some Permian fusulinids from Pahang, Ekspeditsiya, 1932, Ceologiya Pamira, 8, ONTI, Malaya. Ceol. Palaeont. Southeast Asia, vol. 3, p. 1-112, pis. 1-3. (in Russian) p. 30-38, pis. 4-9. Geinitz, H.B., 1876: Zur Geologic von Sumatra, I. Ishibashi, T., 1986: Fusulines from the Ryukyu Zur Geologie von Sumatra's Westkuste. Islands, Part 3. Iheya-jima 2, Tonaki-jima and Palaeontogr., Bd. 22, p. 399-404. Okinawa-jima (Paleontological study of the Gubler, J., 1935: Les fusulinides du Permien de Ryukyu Islands-X). Mem. Fac. Sci., KYlishu l'Indochine, leur structure et leur classification. Univ., ser. D, vol. 26, no. I, p. 95-123, pis. 13-22. Mem. Soc. Ceol. France, N.S., vol. II, fasc.4, Ishii, K., 1966: On some fusulinids and other for­ no. 26, p. 1-173, pis. 17-24. aminifera from the Permian of Pahang, Malaya Han, J.X., 1976: Protozoa. In, Shenyang Inst. Geol. (Notes on the geology and palaeontology of Mineral Resour. ed., Paleontological A tlas of Malaya-V). Jour. Ceosci., Osaka City Unil'., Northeast China, no. 1 (Paleozoic vol.), p. 18-95, vol. 9, art. 4-V, p. 131-142, pIs. 5-6. pis. 1-35. Geol. Publ. House, Beijing. (in -- and Nogami, Y., 1961: On the new genus Chinese) Metadoliolina. Trans. Proc. Palaeont. Soc. Han, N.R., He, L.Y. and Zhou, Y.M., 1987: Shell Japan, N.S., no. 44, p. 161-166, pI. 25. structures of Verbeekina. Acta Micropalaeont. -- and --, 1964: Contributions to the geology Sinica, vol. 4, no. 4, p. 423-426, pis. 1-2. (in and paleontology of Cambodia, Part I. Per­ Chinese with English abstract) mian fusulinids. JOllr. Ceosci., Osaka City Hanzawa, S., 1954: Notes on Afghanella and Univ., vol. 8, art. 2, p. 9-68, pIs. 1-8. Sumatrina from Japan. Japan. Jour. Ceol. Ishizaki, K., 1962: Stratigraphical and paleonto­ Ceogr., vol. 24, p. 1-14, pis. 1-3. logical studies of the Onogahara and its neigh­ -- and Murata, M., 1963: The paleontologic and bouring area, Kochi and Ehime Prefectures, stratigraphic considerations on the Neoschwager­ southwest Japan. Sci. Rep., Tohoku Univ., 2nd. ininae and Verbeekininae, with the descriptions ser. (Ceol.), vol. 34, no. 2, p. 95-185, pIs. 7-12. 1066 Katsumi Vena

Kahler, F. and Kahler, G., 1979: Fusuliniden (For­ Guguk Bulat (Padanger Oberland, Sumatra). aminiferen) aus dem Karbon und Perm von Verh., Geol.-mijnb. Genootsch. Nederland en Westanatolien und dem Iran. Mitt. OSterr. geol. Kolonien., geol. ser., Deel 7, p. 157-239, pis. 1-5. Ges., vol. 70 (1977), p. 187-269, pis. 1-10. Leven, E Ya., 1965: 0 stratigraficheskom znachenii Kanmera, K., 1957: Revised classification of Cancel­ roda Polydiexodina Dunbar et Skinner, 1931. lina, Neoschwagerina, and evolution of Voprosy Mikropaleont., vol. 9, p. 129-146, pis. 1- Sumatrininae and Neoschwagerininae. Mem. 6. (in Russian) Fac. Sci., Kyushu Univ., ser. D, vol. 6, no. I, p. --, 1967: Stratigrafiya i fuzulinidy permskikh otl­ 47-64, pis. 19-20. ozheniy Pamira. Akad. Nauk SSSR, Trudy --, 1963: Fusulines of the Middle Permian Kozaki Geol. Inst., vol. 167, p. 1-224, pis. 1-39. (in Rus­ Formation. Ibid., vol. 14, no. 2, p. 79-141, pis. sian) 11-19. --, 1980: Ob'yasniitel'naya zapiska k stratig­ Kanuma, M., 1960: Stratigraphical and paleonto­ raficheskoy shkale permskikh otlozheniy oblasti logical studies of the southern part of the Hida Tetis. 51 p., Minist. Geol. SSSR, Vses. Orden a plateau and north-eastern part of the Mino Lenina Nauchno-Issled. Geol. Inst. (VSEGEI), mountainland, central Japan, Part II. Leningrad. (in Russian) Paleontology, no. 4. Ozawainellinae, Schubertel­ Lin, J.X., 1984: Protozoa. In, Yichan Inst., Geol. linae, Verbeekininae and Neoschwagerininae. Mineral Resour. chiefly ed., Biostratigraphy of Bull. Tokyo Gakugei Univ., vol. II, p. 55-83, pis. the Yangtze gorge area, (3), Late Paleozoic Era, 10-13. p. 110-177, pis. 1-16. Geol. Publ. House, Beijing. Kawano, M., 1960: Some fusulinids from the Aralani (in Chinese with English abstract and descrip­ Conglomerate in the northeastern part of tion of new species) Yamaguchi City. Sci. Rep., Tohoku Un iv. , 2nd --, Li, J.X., Chen, G.X., Zhou, Z.R. and Zhang, B. ser. (Geol.), Spec. Vol., no. 4, p. 223-230, pis. 24- F., 1977: Fusulinida. In, Hubei. Inst., Geol. 25. Sci. et al. eds., Paleontological Atlas of Central --, 1961: Stratigraphical and paleontological South China, no. 2 (Late Paleozoic), p. 4-96, pis. studies of the Paleozoic formations in the western 1-30. Geol. Publ. House, Beijing. (in Chinese) part of the Chugoku Massif. Bull. Fac. Educ., Liu, Z.A., Xiao, X.M. and Dong, W.L., 1978: Prot­ Yamaguchi Univ., Mathematics & Science, vol. ozoa. In, Working Group of Stratigraphy and II, spec. no., p. 1-133, pis. 1-15. Paleontology of Guizhou ed., Paleontological Kobayashi, F., 1979: Petrography and sedimentary A tlas of South west China, Guizhou vol., no. 2 environment of the Permian Nabeyama limestone (Carboniferous to Quaternary), p. 12-98, pis. 1- in the Kuzuu area, Tochigi Prefecture, central 24. Geol. Publ. House, Beijing. (in Chinese) Japan. Jour. Geol. Soc. Japan, vol. 85, no. 10, Loeblich, A.R., Jr. and Tappan, H., 1988: For­ p.627-642, pis. 1-4. (in Japanese with English aminiferal genera and their classification. 970 abstract) p., 847 pis. Van Nostrand Reinhold Comp., --, 1986: Middle Permian foraminifers of the New York. Gozenyama Formation, southern Kwanto Moun­ Matsusue, K., 1986: Foraminiferal biostratigraphy of tains, Japan. Bull., Natn. Sci. Mus., Tokyo, ser. the lower part of the Akiyoshi Limestone Group. C, vol. 12, no. 4, p. 131-163, pis. 1-10. Sci. Rep., Dept. Geol., Kyushu Univ., vol. 14, no. --, 1988a: Middle Permian foraminifers of the Omi 4, p. 163-185, pis. 2-7. (in Japanese with English Limestone, central Japan. Ibid., vol. 14, no. I, abstract) p. 1-35, pis. 1-13. Miklukho-Maklay, A.D., 1957: Nekotorye --, 1988b: Late Paleozoic foraminifers of the fuzulinidy permi Kryma. Uchenye Zapiski Ogawadani Formation, southern Kwanto moun­ Leningradskogo Gosudarstvennogo Universiteita, tains, Japan. Trans. Proc. Palaeont. Soc. no. 225, ser. geol. Nauk 9, p. 93-159, pis. 1-14. Japan, N.S., no. 150, p.435-452. (in Russian) Kobayashi, M., 1957: Paleontological study of the Minato, M. and Honjo, S., 1959: The axial septula of Ibukiyama Limestone, Shiga Prefecture, central some Japanese Neoschwagerininae with special Japan. Sci. Rep., Tokyo Kyoiku Daigaku, sect. remarks of the phylogeny of the Subfamily Neos­ e, vol. 5, no. 48, p.247-331, pis. 1-10. chwagerininae Dunbar and Condra, 1928. Jour. Kochansky-Devide, V. and Ramovs, A., 1955: Die Fac. Sci., Hokkaido Univ., ser. 4, vol. 10, no. 2, Neoschwagerinenschichten und ihre Fusuliniden­ p. 305-336, pis. 1-6. fauna bei Bohinjska Bela und Bled (Julische Murata, M., 1961 : On the geological structure of the Alpen, Siowenien, NW-Jugoslawien). Siov. Akiyoshi Plateau. Contr., Inst. Geol. Paleont., A kad., 4, p. 359-424, pis. 1-8. Tohoku Univ., no. 53, p. 1-46. (in Japanese with Lange, E., 1925: Eine mittelpermische Fauna von English abstract) 934. Verbeekinids and neoschwagerinids from Akiyoshi 1067

Nogami, Y., 1961: Permische Fusuliniden aus dem limestone of Akasaka, province of Mino. Jour. Atetsu-Plateau SUdwestjapans, Teil2. Verbee­ Fac. Sci., Imp. Univ. Tokyo., scct. 2, vol. 2, no. 3, kininae, Neoschwagerininae u. a. Mem. Coli. p. 121-164, pIs. 34-46. Sci., Univ. Kyoto, ser. B, vol. 28, no. 2, p. 159- -- and Tobler, A., 1929: Permian Fusulinidae 228, pis. 1-7. found in Greece. Eclog. Geol. Helv., vol. 22, no. Ota, M., 1968: The Akiyoshi Limestone Group: A I, p. 45-49, pI. 5. . geosynclinal organic reef complex. Bull., Pasini, M., 1965: Fusulinidi, una chiave analitica per Akiyoshi-dai Sci. Mus., no. 5, p. 1-44, pis. 1-31. la determinazione del generi. 107 p., 18 pIs. (in Japanese with English abstract) Palaeontgr. Italica, Pisa. --, 1977: Geological studies of Akiyoshi, Part l. Pitakpaivan, K., 1965: Fusulines of the Rat Buri General geology of the Akiyoshi Limestone limestone of Thailand (Fusuline fossils from Group. Bull., Akiyoshi-dai Mus. Nat. His!., no. Thailand, Part I). Mem. Fac. Sci., Kyushu 12, p. 1-33, pis. 1-3. Unil'., ser. D, vol. 17, no. I, p. 3-69, pIs. 1-6. --, Toriyama, R., Sugimura, A. and Haikawa, T., Rauser-Chernoussova, O.M. and Fursenko, A. V., 1973: Restudy on the geologic structure of the 1959: Osnovy paleontologii, Obschaya chast; Akiyoshi Limestone Group, southwest Japan. Prosteyshie. 482 p., 13 pis. Izdatel. Akad. N auk Japan. Jour. Geol. Geogr., vol. 82, no. 3, p. 115- SSSR, Moskva. (in Russian) 135. (in Japanese with English abstract) Rozovskaya, S.E., 1975: Sostav, sistema i filogeniya Ozawa, T., 1970a: Notes on the phylogeny and otryada fuzulinida. Akad. Nauk SSSR, Trudy classification of the superfamily Verbeekinoidea Paleont. Inst., vol. 149, p. 1-227, pis. 1-35. (in (Studies of the Permian verbeekinoidean Russian) foraminifera-I). Mem. Fac. Sci., Kyushu Univ., Sada, K., 1961: Neoschwagerines from the Yukawa ser. D, vol. 20, no. I, p. 17-58, pis. 1-9. Group in the Atetsu Limestone plateau. Jour. --, 1970b: Variation and relative growth of Sci., Hiroshima Univ., ser. C, vol. 4, no. I, p. Colania douvillei (Ozawa) from the Rat Buri 117-129, pis. 11-14. Limestone (Studies of the Permian verbee­ Sakagami, S., 1958: Fusulinids from the Upper Per­ kinoidean foraminifera-II). Geol. Palaeont. mian conglomerates of the northern part of Itsu­ Southeast Asia, vol. 8, p. 19-41, pis. 3-6. kaichi, Tokyo-to, Japan. Jour. Hokkaido Ga­ --, 1975: Stratigraphy of the Paleozoic and Mes­ kllgei Univ., ser. 2, vol. 9, no. 2, p. 72-97, pis. 1- ozoic strata in the Tamagawa area, southeastern 4. part of the K wanto mountains. Sci. Rep., Dept. -- and Miyama, Y., 1988: On the limestone­ Geol., Kyushu Univ., vol. 12, no. 2, p. 57-76, pis. bearing conglomerates in the Maki and Terauchi 7-10. (in Japanese with English abstract) Formations in the Atetsu Limestone Plateau, -- and Kobayashi, F., 1990: Carboniferous to southwest Japan. Proc. Japan Acad., vol. 64, Permian Akiyoshi Limestone Group. In, Or­ ser. B, no. 4, p. 92-97. ganizing Committee Benthos '90 ed., Fossil and Sakaguchi, S., Sugano, K., Nakamura, T. and recent benthic foraminifera in some selected Kunugi, K., 1966: A study on the biostratigra­ regions of Japan. Guidebook for field trips, phy of the Kaerimizu doline, Akiyoshi lime­ 4th International Symposium on Benthic For­ stone Plateau, Yamaguchi Prefecture, Japan (Pre­ aminifera, Sendai, 1990, p. E 1-E31, pIs. 1-13. liminary). Mem. Osaka Univ., Liberal Arts Ozawa, Y., 1922: Preliminary note on the and EdIlC., B, no. 14, p.220-229. (in Japanese classification of the family Fusulinidae. Jour. with English abstract) Geol. Soc. Japan, vol. 29, no. 348, p. 352-365. (in Sheng, J.Z. (Sheng, J.c.), 1958: Some fusulinids from Japanese) the Maokou Limestone of Chinghai Province, --, 1923: Stratigraphical study on the so-called northwestern China. Acta Palaeon!. Sinica, Upper Chichibu Paleozoic system including the vol. 6, no. 3, p. 268-291, pis. 1-4. --, 1963: Permian fusulinids of Kwangsi, Kuei­ Akiyoshi-dai Limestone. Ibid., vol. 30, no. 357, chow and Szechuan. Palaeont. Sinica, N.S., ser. p. 222-243. (in Japanese) B, no. 10, p. 1-247, pIs. 1-36. --, 1925a: On the classification of Fusulinidae. -- ed., 1966: Fusulinids of China. 177 p., 27 pis. Jour. Coli. Sci., Imp. Univ. Tokyo, vol. 45, art. 4, Sci. Publ. House, Beijing. (in Chinese) p. 1-26, pis. 1-4. -- and Sun, ~.O., 1975: Fusulinids of Qinghai. --, 1925b: Paleontological and stratigraphical 97 p., 14 pis. Geol. Publ. House, Beijing. (in studies on the Permo-Carboniferous limestone of Chinese) Nagato, Part II, Paleontology. Ibid., vol. 45, Skinner, J.W. and Wilde, G.L., 1966: Permian art. 6, p. 1-90, pis. 1-14. fusulinids from Sicily. Univ. Kansas, Paleont. --, 1927: Stratigraphical studies of the fusulina Contr., Papers., no. 8, p. 1-16, pis. 1-20. 1068 Katsumi Uena

-- and --, 1967: Permian foraminifera from 1958: Geology of Akiyoshi, Part III. Tunisia. Ibid., no. 30, p. 1-22, pis. 1-32. Fusulinids of Akiyoshi. Ibid., vol. 7, p. 1-264, Staff, H. von, 1909: Beitrage wr Kenntnis der pis. 1-48. Fusuliniden. Neues Jahrb. Min. Geol. --, 1975: Permian fusulines from the Rat Buri Palaeont., Bd. 27, p. 461-508, pis. 7-8. Limestone in the Khao Phlong Phrab area, Sara Sun, Q. L. and Zhang, L.X., 1985: Early Permian Buri, central Thailand (Fusuline fossils from fusulinids from Alge mountain of Xinjiang. Thailand, Part IX). Ibid., vol. 23, no. I, p. 1- Acta. Palaeont. Sinica, vol. 24, no. 5, p. 503-510, 116, pis. 1-21. pis. 1-2. (in Chinese with English abstract) -- and Kanmera, K., 1977: The Permian fusulines Sun, X.F., Gao, C.X. and Zhang, X.L., 1983: from the limestone conglomerate formation in the Fusulinida. In, Xi'an Inst., Geol. Mineral Khao Phlong Phrab area, Sara BUfi, central Resour. ed., Paleontological Atlas of Northwest Thailand (Fusuline fossils from Thailand, Part China, Shaanxi, Gansu and Ningxia vol., Part 2 X). Geol. Palaeont. Southeast Asia, vol. 18, p. (Upper Palaeozoic), p.3-40, pis. I-10. Geol. 1-27, pis. 1-3. Publ. House, Beijing. (in Chinese) -- and --, 1979: Permian fusulines from the Suyari, K., 1962: Geological and paleontological Ratburi Limstone in the Khao Khao area, Sara studies in central and eastern Shikoku, Japan, Buri, central Thailand (Fusuline fossils from Part II. Paleontology. Jour. Gakugei, Toku­ Thailand, Part XII). Ibid., vol. 20, p. 23-93, pis. shima Univ., vol. 12, p. 1-64, pis. 1-12. 4-14. Thompson, M.L., 1936: The fusulinid genus Verbee­ -- and Pitakpaivan, K., 1973: Middle Permian kina. Jour. Paleont., vol. 10, no. 3, p. 193-201, pI. fusulines from Wait Kirinakratanaram, central 24. Thailand (Fusulinacean fossils from Thailand, --, 1946: Permian fusulinids from Afghanistan. Part VII). Ibid., vol. 12, p. 43-61, pis. 5-6. Ibid., vol. 20, no. 2, p. 140-157, pis. 23-26. Ueno, K., 1989: Carboniferous and Lower Permian --, 1948: Studies of American fusulinids. Univ. foraminiferal biostratigraphy in the Akiyoshi Kansas, Paleont. Contr., Protozoa, art. I, p. 1- Limestone Group (Studies of the Upper Palae­ 184, pis. 1-38. ozoic foraminifers in the Akiyoshi Limestone -- and Foster, c.L., 1937: Middle Permian Group, southwest Japan, Part I). Bull., fusulinids from Szechuan, China. Jour. Akiyoshi-dai Mus. Nat. Hisl., no. 24, p. 1-39, pis. Paleont., vol. II, no. 2, p. 126-144, pis. 23-25. 1-8. (in Japanese with English abstract) Tien, N.D., 1970: Quelques fusulinides de NGi Com, --, 1991a: Pamirina (Permian Fusulinacea) from sud Viet-Nam (Zone a Neoschwagerina). Ar­ the Akiyoshi Limestone Group, southwest Japan. chives Geol. Viet-Nam, no. 13, fasc. I, p. I-70, Trans. Proc. Palaeont. Soc. Japan, N.S., no. 161, pis. 1-15. p.739-750. --, 1979: Etude micropaleontologique (For­ --, 1991 b: Early evolu'tion of the families Verbee­ aminiferes) de materiaux du Permien du Cam­ kinidae and Neoschwagerinidae (Permian bodge. These Centre d'Orsay, Univ. Paris Sud, Fusulinacea) in the Akiyoshi Limestone Group, p. 1-166, pis. 1-32. southwest Japan. Ibid., no. 164, p.973-1002. --, 1986a: Foraminifera and algae from the Per­ Wang, J.H. and Tang, Y., 1986: Fusulinids from mian of Kampuchea. In, Fontaine, H. ed., The Chihsia Formation of Lengwu district in Tonglu Permian of Southeast Asia, CCOP Tech. Bull., County, Zhejiang. Acta Micropalaeont. Sinica, vol. 18, p.116-137, pis. I-10. vol. 3, no. I, p. 3-11, pis. 1-3. (in Chinese with --, 1986b: Foraminifera and algae from the Per­ English abstract) mian of Guguk Bulat and Silungkang, Sumatra. Wang, Y.H., Wang, L.L., Wang, J.H., Zhu, Z.G., Lin, Ibid., p. 138-147, pis. 11-15. G.W., Zhang, L.X. and Quian, Q., 1982: --, 1989: Middle Permian foraminifera. In, Fusulinida. In, Nanjing Inst. Geol. Mineral Fontaine, H. and Gafoer, S. eds., The Pre­ Resour. chiefly ed., Paleontological A tlas of Tertiary fossils of Sumatra and their environ­ East China, no.2 (Late Palaeozoic), p.5-108, ments. CCOP/TP 19, p. 113-148, pis. 20-35. pis. 1-30. Geo!. Pub!. House, Beijing. (in Toriyama, R., 1947: On some fusulinids from Chinese) Tosayama, Koti-ken, shikoku, with a note on the Wang, Y.J., Sheng, J.Z. and Zhang, L.X., 1981: stratigraphical range of Neosch wagerina. Fusulinids from Xizang of China. In, Nanjing Japan. Jour. Geol. Geogr., vol. 20, nos. 2-4, p. Inst. Geol. Palaeont. ed., Palaeontology of 64-82, pis. 26-27. Xizang, Book 3, p. I-80, pis. 1-21. Sci. Pub!. --, 1954: Geology of Akiyoshi, Part I. Study of House, Beijing. (in Chinese with English the Akiyoshi Limestone Group. Mem. Fac. abstract) Sci., Kyushu Univ., ser. D, vol. 4, no. I, p. 39-97. -- and Zhou, J.P., 1986: New material of 934. Verbeekinids and neoschwagerinids from Akiyoshi 1069

fusulinids from Xainza, Xizang. Bull.. Nanjing "Maokou Limestone" (Permian) at Oatieguan, Insl. Ceol. Palaeont .• Acad. Sinica. no. 10, p. Langdai, Guizhou. Acta Micropalaeonl. Sinica. 141-156, pis. 1-4. (in Chinese with English vol. 2, no. 4, p. 307-335, pis. 1-3. (in Chinese abstract and description of new species) with English abstract and description of new Xie, S.G., 1982: Protozoa. In, Geol. Bureau, Hunan genus) ed.. The Palaeontological A tlas of Hunan. Ceol. Yoshimura, N., 1961 : Geological studies of the Pale­ Mem .. ser. 2, no. I, p.2-73, pis. 1-35. Geol. ozoic groups in the Oga Plateau, central Chu­ Publ. House, Beijing. (in Chinese) goku, Japan. Ceol. Rep.. Hiroshima Unil'., no. Yanagida, J., Ota, M., Sugimura, A. and Haikawa, T., 10, p. 1-36, pis. 1-4. (in Japanese with English 1971 : On the geology of the northeastern part of abstract) the Akiyoshi limestone plateau. Sci. Repts.. Zhang, L.X. and Wang, Y.l., 1974: Permian Dept. Ceol.. Kyushu Univ., vol. II, no. I, p. 105- Fusulinida. In, Nanjing Inst., Geol. Palaeonil. 114, pis. 7-10. (in Japanese with English ed.. Stratigraphical and Paleontological Atlas abstract) of West South China. p.289-296, pis. 150-153. Yang, Z.O., 1985: Restudy of fusulinids from the Sci. Pub I. House, Beijing. (in Chinese)

Akiyoshi f.:k6', Akiyoshi-dai f.:k6'iJ. Atetsu ~iiJ1Y. Kaerimizu jffi I) 7.K. Omi WfliJ. Taishaku 7iffR. Yamaguchi ill p.

f.:k 6' E I!k.;fi ~ ff. Parafusulina kaerimizensis '* tJ Ili'f: 0) Verbeekinidae:f4 to J: If Neo­ schwagerinidae :f4f,jj~£E.~: f.:k6'iJ;lt*$O)jffi I) 7Jc~~I=-5t;fjj-t Q Parafusulina kaerim­ izensis '* J: I) 1::. {lL 0) fKi5 E I!k.;fi ~ ff t.J' G II flO) Verbeekinidae :f4 to J: U' Neoschwager­ inidae fJj.iiiO~£E.~~~5J1j L t-.:o ~ G v=- c..h Gf,jj~£E.~O) ~ J1':5t;fjj v=-~--j\;, -C. Parafusulina kaerimizensis '* tJlli'f: 0) f.:k6'E I!k.E ~ff ~ T {lL t.J' G Parafusulina kaerimizensis m. Afgh­ anella ozawai '*. Neoschwagerina craticulifera robusta m. Verbeekina verbeeki-Afghanella schencki '*. Neoschwagerina fusiformis '*. Verbeekina verbeeki m. Colania douvillei m 0) 71tE'*v=-5tm L t-.:o *~tfli-efi I ~BEfl (Neoschwagerina craticulifera robsta, subsp, nov.) ~%tsc.O) ') 'f:,0) 10 fl~~G$t-t Q c. c.

935. A NEW CRYPTIC SPECIES OF PYCNODONTE FROM RYUKYU ISLANDS: A LIVING FOSSIL OYSTER *

ITARU HAYAMI

Geological Institute, University of Tokyo, Hongo 7-3-1, Bunkyo-ku, Tokyo, 113

and

TOMOKI KASE

National Science Museum, Hyakunin-cho 3-23-1, Shinjuku-ku, Tokyo, 169

Abstract. A new enigmatic oyster was found alive from several poorly lighted subma­ rine caves of Miyako and Okinawa Islands, southern Japan. It belongs to the genus Pycnodonte, which occurs commonly in the Cretaceous-Miocene of the Tethyan realm but had been believed extinct. Its shell shape is variable, but the right valve is paper thin and has an extremely wide depositional surface of flexible outer prismatic layer. Nevertheless, every diagnostic character of Pycnodonte (s.s.) is well recognized (e.g. long dorsal margin, dorsally located and subcircular adductor muscle insertion, wide commissural shelf, ver­ micular chomata, vesicular shell structure and remarkable radial gashes on the right valve). It is suggested that this oyster is a significant example of a living fossil, and that Pycnodonte has survived for many millions of years by transforming itself into a cryptic organism.

Key words. cryptic, Pycnodonte, submarine cave, living fossil.

Introduction High levels of endemism in cryptic organ­ isms have attracted the attention of many The technical development and populariza­ biologists. Riedl (1966) made a comprehen­ tion of scuba diving in recent years enabled us sive review on the biota of coastal caves and to examine the diversity and ecology of vari­ illustrated the spectral distribution of various ous organisms indigenous to sublittoral organisms (including some bivalves) in some sheltered environments. Submarine cave wave-excavated intertidal grottoes of the faunas, in particular, have emerged as an Mediterranean region. Since Hartman and interesting subject of study because they seem Goreau (1970) discovered sclerosponges to offer substantial evidence to test various resembling stromatoporoids (problematic theories of systematics, evolutionary ecology Paleozoic and Mesozoic fossil organisms) in and marine biogeography. Cryptic faunas some sublittoral crevices of Caribbean coral also may contain paleontologically interest­ reefs, several species of articulate bra­ ing species because such sheltered places serve chiopods, bryozoans, polychaetes and as suitable refuges to archaic and anachronis­ decapod crustaceans have been recognized tic organisms. also as unique constituents of cryptic commu­ 'Received January 10, 1992; accepted February 3, nities, and their origin and evolutionary 1992 significance have been discussed by many

1070 935. A living fossil oyster 1071

investigators (Jackson, Goreau and Hartman, larly sheltered environments of Okinawa (Ie­ 1971; Jackson and Winston, 1982; Iliffe, jima and Seragaki beach). In spite of its Hart and Manning, 1983; Kobluk, 1988; enigmatic shell morphology, many essential etc.). However, the unique features of mol­ characters indicate that this oyster belongs to luscs in such sheltered environments have the genus Pycnodonte, which flourished in been little studied. Cretaceous and Paleogene periods in the On the fore-reef slopes of Shimoji-shima Tethyan realm and had been believed to be an and Ie-jima of the Ryukyu Islands, southern extinct taxon. Its cryptic habitat is also Japan, there are numerous limestone caves meaningful, because such a sheltered place which seem to have been formed by under­ may offer a refuge for archaic organisms. In ground water during some low sea-level this article we describe its morphology and stages of the Pleistocene and drowned during life habit and discuss its evolutionary the post-glacial sea-level rise. Since a few significance as a "living fossil". years ago, we have been studying the cryptic faunas (especially molluscs) in these caves On the genus Pycnodonte with the vigorous assistance of several skilled divers. As reported preliminarily (Hayami Before going into the description, previous and Kase, 1991 ; Kase and Hayami, in press.), systematic studies of pycnodonteine oysters several characteristic moll uscs incl uding Ner­ are briefly reviewed because the newly dis­ itopsis radula (a living fossil snail) have been covered cryptic species seems to belong to found alive in some of these caves, and the Pycnodonte. calcareous muddy sediments therefrom bear In spite of great intraspecific morphologic numerous dead shells of bivalves and gas­ variability and frequent convergence, natural tropods which are different from the shallow­ classification of fossil and extant oysters has water faunas of ordinary exposed environ­ been attempted by many investigators. ments. The diversity, ecology, origin and Among others, Stenzel's (1959, 1971) scheme evolutionary significance of these cryptic may be the most elaborate and influential. faunas may be an interesting subject to study. He classified oysters into two families, In spite of the apparent great species diver­ Gryphaeidae and Ostreidae, and further sub­ sity, most of these molluscs are very small in divided the former into three subfamilies, size (commonly less than 5 mm in maximum Gryphaeinae, Pycnodonteinae and Exogyr­ length). Some sclerosponges, corals, soft inae. Stenzel's scheme has been substantially sponges, articulate brachiopods, bryozoans, followed by many subsequent authors, ostracods and decapod crustaceans were also though Torigoe (1981) raised the Pycno­ found alive in some of these caves, but the donteinae as a distinct family, Harry (1985) biomass per unit space is generally much subdivided the Pycnodonteinae into three smaller than that of the exposed environment tribes (Pycnodontini, Neopycnodontini and around the caves. Hyotissini), and Carter (\ 991) regarded the The new oyster described here is an excep­ Gryphaeidae and the Ostreidae as constitut­ tionally large-sized species among the cave ing distinct superfamilies. bivalves. It was first discovered by M. Important criteria for these family- or Taniguchi, a skilled diver of Miyako Island, subfamily-level classifications are 1) shell at the poorly lighted walls and ceilings of characters (particularly, presence or absence several caves (about 20 meters below sea of chalky deposit and vesicular shell struc­ level) along the western coast of Shimoji­ ture, commissural shelf, chomata, and shape shima of Miyako Islands (Figure 1). The of muscle scar) and 2) anatomical characters same species has been found also from simi- of soft parts (particularly, the status of the 1072 Itaru Hayami and Tomoki Kase

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Miyako o- 6 km Figure 1. Map showing the localities (indicated by asterisks) of Pycnodonte (Pycnodonte) taniguchii n. sp. in Ryukyu Islands of Japan. A: Index map, B: Central part of Okinawa Island, C: Miyako Islands. promyal passage, heart-kidney-rectum com­ accordance with Stenzel's system (with some plex, digestive tract, labial palp fusion and modifications), Torigoe (1981) described the anal appendage) and 3) reproductive strat­ shell and soft parts of all the extant oysters in egy (incubatory or non-incubatory). In Japanese waters, and Harry (1985) gave a 935. A living fossil oyster 1073 synopsis of all the supraspecific taxa of extant seem to support Stenzel's (1971) scheme. oysters in the world. Discrimination of the Pycnodonteinae Stenzel (1971) distinguished the Pycno­ from the Ostreidae is justified also by compar­ donteinae from the Gryphaeinae by the pres­ ative anatomy of extant species. As illus­ ence of a well defined commissural shelf, trated by Torigoe (1981), the Pycnodontinae vermiculate chomata and vesicular shell struc­ shares a Z-shaped digestive tract with the ture, but treated the former as derived from Lophinae, but the rectum penetrates (or the latter, regarding Gryphaea (Bilobissa) comes into contact with) the pericardium of from the Middle-Upper Jurassic as transi­ the heart in Neopycnodonte and Hyotissa, tional between the two subfamilies. Subse­ while it passes the dorsum of the heart in all quent studies of shell microstructure, how­ the genera of the Ostreidae (including the ever, do not necessarily support Stenzel's Lophinae). Adductor muscle insertion is scheme. According to Siewert (1972) and subcircular in the Pycnodonteinae (like the Hudson and Palmer (1976), Pycnodonte and Gryphaeinae) but reniform in the Ostreidae. Hyotissa considerably differ from the The status of the promyal passage and labial Gryphaeinae but resemble the Ostreidae in palp fusion may be also important to discrim­ the structural chambering and in lacking well inate the Pycnodonteinae from the Ostreidae. defined crossed foliated structure. Siewert Habe (1977) regarded Pretostrea as a pycno­ (1972) supposed that Pycnodonte together donteine genus and referred several Japanese with many ostreid genera was derived from species to it, but Torigoe (1981) clarified on Lopha which constituted a stock independent the basis of anatomy that some of them from Gryphaea throughout the Jurassic. belong to Hyotissa and other species to Carter (1991) critically reviewed all the Dendostrea (of the Lophinae). These ana­ available data about the shell microstructure tomical characters are useful to see through of fossil and extant oysters, and admitted the shell convergence. structural resemblance between the Pycno­ Pycnodonte Fischer de Waldheim, 1835, as donteinae and the Ostreidae, although he did defined by Stenzel (1971) and many recent not change Stenzel's (1971) system in which workers, had been regarded as an extinct the Pycnodonteinae is regarded as a sub­ genus, though its descendant(?) genus family of the Gryphaeidae. Contrary to Sten­ Neopycnodonte Stenzel, 1971, is represented zel's description, "No prismatic shell layer by a single extant circumglobal species, N. except in Neopycnodonte': an outer layer of cochlear (Poli). Hyotissa Stenzel, 1971, calcitic simple prisms is commonly observed which comprises several tropical-subtropical in RV of many fossil and extant species and corresponds to the tribe Hyotis­ pycnodonteine species. Vesicular shell struc­ sini Harry, 1985, is another extant pycno­ ture, which was regarded by Stenzel (1971) as donteine genus, but the Lopha-like plicate diagnostic of the Pycnodonteinae, exists also valves, subequivalve shell, internal surface in a lophine oyster, Dendostrea folium. with a moire luster and huge adductor muscle Therefore, the origin and the familial posi­ insertion are dissimilar from those of tion of the Pycnodonteinae is still controver­ Pycnodonte and Neopycnodonte. sial. If much weight is given to the shell A large number of fossil species of structure and structural chambering, the Pycnodonte (including Gigantostrea, Pycnodonteinae may be considered to be Phygraea and Costeina as subgenera) have more closely related to the Ostreidae (espe­ been known mainly in the Tethyan realm cially the Lophinae) than to the Gryphaeinae. from late Early Cretaceous to early Miocene. Nevertheless, as to the content and limit of Probably owing to biased fossil preservation, the Pycnodonteinae, most recent investigators only left valves were described in many fossil 1074 Itarll Hayami and Tomoki Kase species. Though the intraspecific variation conspicuous radial gashes of RV like and detailed shell characters of the type Pycnodonte (s.s.), but its subgeneric distinc­ species are poorly known, the gen us tion can be based on the Gryphaea-like out­ Pycnodonte, as defined by Stenzel (1971 : p. line (though the resemblance is due to conver­ N 1106), seems to be characterized by strongly gence), much thicker LV, prominent umbo of convex LV, irregular but generally subcir­ L V, short dorsal margin, and undeveloped cular to semicircular outline, wide commis­ auricles. This subgenus possibly includes sural shelf, vermicular chomata, gently un­ Pycnodonte vesicularis (Lamarck, 1806) from dulated LV rarely with well defined radial the Upper Cretaceous of western Europe, west costae and flat to concave RV. Radial and north Africa, south India and New gashes of R V is a very conspicuous feature of Caledonia. Woods (1913) and Freneix Pycnodonte, though they sometimes occur (1960,1972) described the wide morphologic also in other genera of the Pycnodonteinae (e. variation of P. vesicularis, and many Eur­ g. Harry, 1985: p. 136, fig. 16) and some opean authors regarded P. radiata, the type species of the Gryphaeinae (e.g. Duff, 1978: species of Pycnodonte, as synonymous with P. pI. 8, fig 7b). vesicularis. This view was rejected and P. Subgeneric division of Pycnodonte is con­ vesicularis was referred to Phygraea by Sten­ troversial and somewhat confusing. zel (1971) and some subsequent authors, Pycnodonte (s.s.) [type species: Pycnodonte chiefly because its subcircular shell form radiala from the Upper Cretaceous of the looks different from the figured original speci­ Crimea] is generally defined by the im­ men of P. radiata. The shape difference prominent umbo, long dorsal margin, devel­ between Pycnodonte (s.s.) and Phygraea, as oped auricular parts and absence of radial discussed later, may be related to di fferent ribs on LV. Gigantostrea Sacco, 1897 [type modes of life. species: Ostrea gigantica Solander in Bran­ In addition, Costeina Vyalov, 1965 [type der, 1766, from the Eocene of England], was species: Pycnodonte (Costeina) costei Co­ regarded as synonymous with Pycnodonte quand, 1869, from the Upper Cretaceous of (s.s.) by Stenzel (1971 : p. NI107), but Palmer north Africa] may be characterized by the and Brann (1965: p. 149) and Wilson (1987 : presence of dichotomous radial ribs on the p. 13) applied this subgeneric name to several surface of LV, and Crenostrea Marwick, 1931 species from the Eocene-Miocene of the At­ [type species: Ostrea (Crenostrea) wueller­ lantic Coastal Plain of USA, and Fleming storfi Zittel, 1864, from the Oligocene of New (1966: p. 23) also to two species from the Zealand] seems to be subgenerically distin­ Paleogene of New Zealand. Gigantostrea guished by the pointed umbo of LV and very seems to differ from Pycnodonte (s.s.) in the strong chomata, though R V of their type planoconvex to biconvex outline, less ine­ species are insufficiently known. Squires quivalve shell and undeveloped auricles. and Demetrion (1990) proposed Pegma [type Phygraea Vyalov, 1936 [type species: species: Pycnodonte (Pegma) bajaensis Phygraea frauscheri Vyalov, 1936 (= Sq uires and Demetrion, 1990, from the Gryphaea pseudovesicularis GUmbel, 1861), Eocene of Baja California] as a subgenus of from the Paleocene of eastern Europe] shows Pycnodonte, but we think that it is, if not planoconvex to concavoconvex outline and synonymous with, closely related to Hyotissa.

~ Figure 2. Pycnodonte (Pycnodonte) tanigllchii n. sp., Holotype (UMUT RM 18908), brownish indi­ vidual. la: Exterior of LV with many dead shells of juvenile individuals, lb: Interior of LV, the depositional surface of middle layer showing pepper-and-saJt appearance, Ie: Exterior of RV, showing remarkable radial gashes. Id: Interior of RV, showing wide depositional surface of prismatic outer layer. All figures xO.7. 935. A living fossil oyster 1075 1076 Itaru Hayami and Tomoki Kase

Table 1. Observed specimens of Pycnodonte (Pycnodonte) taniguchi/, n. sp. No. Reg. no. Length Height Thickness Phenotype Locality Type Remarks I RM18908 I03mm 138mm 71 mm brownish W-Arch Holotype dried 2 RM18909 87 mm 124mm 71 mm whity Black Hole Paratype dried 3 RMI8910 I09mm I05mm 68 mm whity Devil's Palace Paratype in alcohol 4 RM18911 96mm 84mm 45mm brownish W-Arch Paratype in alcohol 5 RM18912 97mm 76mm 39mm brownish Seragaki Beach Paratype in alcohol 6 RMI8913 97mm 101 mm 53mm brownish W-Arch Paratype sectioned 7 123 mm 125mm 66mm brownish W-Arch anatomized 8 81 mm 84mm 28mm brownish W-Arch anatomized Registered specimens are in the University Museum, University of Tokyo (UMUT). Speciment Nos. 7 and 8 were anatomized by Dr. Torigoe of the Hiroshima University.

The geographic distribution of Pycnodonte Pycnodonte (Pycnodonte) taniguchii was said to be worldwide (Stenzel, 1971 : p. Hayami and Kase, n. sp. N 1107), but this genus has been known Figures 2-7 almost exclusively from low-middle latitudi­ nal regions. Cretaceous species are common Material. - Several living specimens from in western Europe, Crimea, north and west the submarine caves on the western coast of Africa, south India and Gulf Coast regions. Shimoji-shima, Miyako Islands, Ryukyu Paleogene species show almost the same geo­ (Table I). The holotype (UMUT RM 18908) graphic distribution as Cretaceous ones. In from one of the caves, a diving point called Japan large species of Pycnodonte seems to be "W-Arch" (Lat. 24°51'43"N, Long. 12Y09'41" rare, but Tashiro (1978) described a small E, ca. 20 m below sea level). species from the Santonian of west Kyushu, Diagnosis. - A large and concavo-convex and Hanai and Oji (1981) noticed numerous extant species of Pycnodonte (s.s.), the adult individuals (mostly LV) of an undescribed shell of which is characterized by an unusu­ species adhering to an Aptian fossil bea­ ally wide commissural shelf, an attachment chrock in north Honshu. As revised by area extending over the long geniculated Oyama, Mizuno and Sakamoto (1960: p. dorsal margin of LV, distinct vermicular 129), Ostrea crassis Nagao, 1928, from the chomata, dorsally situated subcircular ad­ Oligocene of west Kyushu seems to belong to ductor insertion, variably developed vesicular Pycnodonte, although Nagao (1928) wrongly structure, moderately thick LV, and more or described the right and left valves in reverse. less hollowed RV with a paper-thin and All the extant representatives of the flexible distal area and remarkable radial Pycnodonteinae (Neopycnodonte and Hyotis­ gashes on the external surface. sa) are also tropical to warm-temperate General shell features. - Shell large for the inhabitants (Harry, 1985,1986; Nicol, 1991). genus, often exceeding 100 mm in length and height, very inequivalve, more or less proso­ cline, nonplicated, presumably dimorphic in Systematic description coloration (commonly purplish brown but sometimes ivory white). Outline variable, usually flat and subcircular in young stage ? Family Gryphaeidae Vyalov, 1936 but becoming linguiform or fan-shaped in Subfamily Pycnodonteinae Stenzel, 1959 adult. LV strongly convex, solid, relatively Genus Pycnodonte Fischer thin except for middle-ventral area in adult de Waldheim, 1835 stage, characterized by improminent umbo, Subgenus Pycnodonte s.s. broadly undulated and puckery surface and 935. A living fossil oyster 10 77

10 1b Figure 3. Pycnodonte (Pycnodonte) wniguchii n. sp. , Holotype (UMUT RM 18908). la: Umbonal area of LV, showing the liga ment area a nd muscle insertions, Ib: Umbonal area of RV, showing the ligament area a nd muscle insertions. All figures X 1.5. generally well developed auricular parts, 1/ 10 of shell height), subcircular, without sometimes having a posterior radial sulcus elevated rim, slightly longer than high in LV but lacking any well-defined radial sculpture but nearly as long as high in RV, with its and hyote spines. Attachment area, which center placed at one-fifth to one-fourth of occupies the whole external surface of LV in shell height from the dorsal margin in adult early stage (up to 20- 30 mm in height), (unusually dorsally situated) . Gill protrac­ extends along the entire dorsal margin in tor muscle scar (Quenstedt muscle insertion adult, geniculated at the umbonal area in by Stenzel, 1971) distinct in both valves, dorsal view. R V broadly and deeply con­ situated a nteriorly in LV but subcentrally cave except flattened or feebly convex um­ nearly below the resilifer in RV. Chomata bonal area, sticking to the inner surface of LV relatively weak but distinct, moderate in with flexible distal area, extremely thin and length, vermicular, situated on the outer light (in the holotype, L V is 221 gram and RV slopes of a pair of small ridges. Commissur­ is 26 gram in weight), characterized by al shelf unusually broad, occupying almost remarkable radial gashes on the external sur­ the distal half of shell; its proximal margin face. Ligament area relatively narrow, markedly angulated on the internal surface of subtriangular with an apical angle of about RV. 60' in LV, though its apical part was com­ Shell microstructure. - The outer layer of monly corroded in RV ; resilifer small, sub­ L V often ragged but consists of accumulated central, slightly opisthocline. Subumbonal regularly foliated calcite which dips slightly cavity of LV moderately deep. Inner surface relative to depositional surface. The middle of both valves shiny but does not show a layer of LV composed of thick accumulation moire luster. Adductor muscle insertion of composite crossed foliated calcite. In comparatively small (with a diameter about purplish-brown individuals this layer catches 1078 flaru Hayami and Tomoki Kase 935. A living fossil oyster 1079

____ umbo (corroded) ~ resilifer "-j --~~Quenstedt muscle insertion If ~~ Q"'---;-~ " It t ..~ ,ermicular chomala 0.,.,.--\ \ 'I / U)'-) --'----- adductor muscle insertion ./ \ .. J proximal margin of / / _ \ commissural shelf b ' // j/ ~ distal margin of

radial gash

/ ~ \ ... -,\,~----',-- de~~~:~~i~ :~~:~cl~y~r ~ \\

---- ventral margin (incomplete)

Figure 5. Sketch of the internal surface of RV (based on the holotype). many yellowish lenses which bear numerous face. Its depositional surface extremely ellipsoidal vacuoles of about 40 pm in maxi­ wide, occupying almost the entire distal half mum diameter (Figure 6-6). The brownish of shell. Owing to this structure, the distal and yellowish parts irregularly intermingled, part of RV without backing of middle layer so that the depositional surface shows a very flexible when alive. Radial gashes pepper-and-sait appearance (Figure 2-1 b). occur only in this outer layer. A number of Development of vesicular structure quite vari­ flat cavities exist in and below the prismatic able in LV; in some individuals there are outer layer, in which vesicular structure is large cavities with very coarse vesicular struc­ commonly developed (Figure 6-3, 4). Mid­ ture between the outer and middle layers, but dle layer of R V cross foliated, solid and rarely in many other individuals this structure is not vacuolated, easily separable from outer layer. observed at all. The outer layer of RV com­ The inner layer of both valves composed of paratively thick in the middle-ventral area complex crossed foliated calcite but generally (attaining almost I mm in total), composed of very thin. several sublayers of simple prismatic calcite Soft part. - The soft part relatively small, which are interrupted by thin conchiolin seats as recognized from the unusually wide or cavities (Figure 6-2). Calcite prisms vari­ commissural shelf. Promyal passage extends able in size, but commonly 20-50 pm in between RV and visceral mass, reaching the diameter and reclined about 50· to the sur- umbonal part. Visceral mass relatively

+- Figure 4. Pycnodonte (Pycnodonte) taniguchii n. sp. 1: Holotype (UMUT RM 18908), dorsal view of articulated valves, 2a-c: Paratype (UMUT RMI891O), left, right and dorsal views of a whity articulated individual. 3a-c: Paratype (UMUT RMI8911), left, right and dorsal views of a brownish articulated individual. All figures X 0.7. 1080 flaru Hay ami and Tomoki Kase 935. A living fossil oyster 1081

small, orange in color. Labial palp broadly from those of N. cochlear, although the rec­ fused, resembling that of Neopycnodonte co­ tum does not pierce the pericardium in the chlear. Digestive tract relatively thick, Z­ present species. N. cochlear is regarded as a shaped, never looped; rectum does not pene­ circumglobal species, having been recorded trate the pericardium of the heart but comes from various regions of the Indo- West in contact with its posterodorsal part; anal Pacific, eastern and western Atlantic, and papilla very long. In summary, every ana­ Mediterranean realms. It extends to greater tomical character resembles that of Neopyc­ depths (27 to 2,100 m) than any other extant nodonte and Hyotissa and differs significantly oyster (Stenzel, 1971; Harry, 1985). In from that of the Ostreidae. (The description Japan, though it often was treated under the of anatomical characters is mainly based on name of Ostrea musashiana Yokoyama, Torigoe's personal communication.) 1920, the species occurs commonly on the Comparisons. - The present new oyster lower sublittoral to upper bathyal gravelly shows every essential character of the bottom (Kuroda, 1931; Habe, 1977; etc.). Pycnodonteinae called for by Stenzel (1971), Although the soft parts are relatively small, Torigoe (1980) , Harry (1985) and Carter the present species grows much larger than N. (1991). In the shell morphology as well as cochlear in shell size. The shape of the adult the cryptic habitat, however, the present shell is also quite different because the attach­ species is so unique that its distinction from ment area of the present species extends other extant oysters can be based on various entirely over the long dorsal margin of LV. characters. The extreme inequivalveness, Moreover, unlike N. cochlear, the simple broad commissural shelf, dorsally situated prismatic outer layer of RV is well developed, adductor muscle, and thin and deeply con­ and its depositional surface is much wider. cave RV with broad and flexible distal area The shell of N. cochlear is fragile, while the (owing to the shell consisting of simple pris­ L V and the middle-inner layer of R V are matic outer layer) in the adult stage are the generally solid in the present species. most striking diagnostic characters of this The present species resembles the type species. species of Pycnodonte (s.s.) more closely than Among extant pycnodonteine oysters, that of Neopycnodonte in various shell char­ Neopycnodonte cochlear (Poli, 1795) may be acters, e.g. concavo-convex valves, long dor­ most closely related to the present species sal margin and linguiform to fan-shaped because the two species share very dorsally outline in the adult stage and well developed situated adductor muscle, vermicular radial gashes on the external surface of RV. chomata, radial gashes and limitedly dis­ Some specimens of the present species are tributed vesicular structure in RV. Accord­ surprisingly similar in shell shape to the ing to Torigoe (pers. comm.), who anatom­ figured original specimen of its type species, ized an individual at our request, many ana­ P. radiata Fischer de Waldheim, 1835 (re­ tomical characters are not much different produced by Stenzel, 1971 : fig. J80), from the

~ Figure 6. SEM photomicrographs showing shell microstructure of Pycnodonte (Pycnodonte) laniguchii, n. sp. Unit of scale in microns. 1: Depositional surface of simple prismatic outer layer near the ventral margin of RV (UMUT RM 18913); the venter is toward the lower, X 250. 2: Oblique internal view of fractured edge of multilayered simple prismatic outer layer near the ventral margin of RV (the same individual); the venter is toward the right upper, X 500. 3: Fractured external surface of middle part of RV (UMUT RMI8909); vesiculate cavity below simple prismatic outer layer, X 100. 4: Vesiculate struc­ ture in a cavity between outer and middle layers of RV (the same individual), X 60. 5: Depositional surface of crossed foliated middle layer of LV (UMUT RM 18913), X 1,000. 6: Slightly etched surface of subver­ tical section of LV (the same individual), vacuolated lenticular part embedded in crossed foliated middle layer; the venter is toward the right and the external surface toward the lower, X 100. 1082 ftaru Hayami and Tomoki Kase

Upper Cretaceous of the Crimea. In a num­ extend over the long dorsal margin as seen in ber of fossil species of Pycnodonte RV looks the present species. All fossil Pycnodonte much smaller than LV, as described and (s.s. and Phygraea) share with the present figured by many authors. Judging from the species conspicuous radial gashes on R V, concordant ventral margins of two valves in which indicate the presence of a well devel­ all the living specimens of the present species, oped simple prismatic outer layer. the "size discordance" in those fossil species Vesicular shell structure is widespread in may be only superficial, because the marginal the fossi I and extant species of the part of R V was physically weak and may have Pycnodonteinae and has been regarded by been selectively lost before fossilization. Stenzel (1971) as diagnostic of this subfamily. Even in living specimens of the present This structure is actually observed in the species, this portion, if dried and not strength­ present species (Figure 6-3,4). In RV it is ened, is apt to be broken into pieces, just like commonly seen in small cavities beneath the the marginal apron of RV in living outer layer, although in LV its development is propeamussiids (see Hayami, 1988). Flex­ quite variable among individuals. As Carter ible distal area of RV, therefore, may be a (1991: p.355, fig.2E) ascertained the pres­ ubiquitous feature in the genus Pycnodonte. ence of locally developed vesicular structure The apparently wide range of variation of in the middle layer of a lophine oyster, shell form of the present species is partly Dendostrea folium, taxonomic evaluation of attributable to ontogenetic change. Some this structure should be reconsidered. Chel' immature specimens of the present species tsova (1969) mentioned the presence of (e.g. UMUT RM18911) show shorter and vacuolated and "pinnate" structure 1TI arcuate dorsal margin and subcircular out­ Pycnodonte radiata. The type species is line, resembling some species of the subgenus possibly similar to the present species in this Phygraea, especially Pycnodonte (Phygraea) feature, though a more detailed comparison mutabilis (Morton, 1828) from the of shell microstructure has yet to be worked Campanian-Maastrichtian of the Gulf and out. Atlantic Coastal Plain, which was described Distribution.-In addition to the type local­ by Gardner (1916) and Wade (1926) as ity, the present species was found alive in Gryphaea vesicularis and by Stephenson submarine caves on fore-reef slope ("Black (1941) as Gryphaea mutabilis. In this Hole" and "Devil's Palace" as called by respect, some specimens of Pycnodonte divers) along the western coast of Shimoji­ (Ph ygraea) vesicularis (Lamarck, 1806), a shima of Miyako Islands, and similarly widely distributed Late Cretaceous species in sheltered places at "Daidokutsu" of Ie-jima Europe, north and west Africa, India and and at Seragaki near Manza Beach of Okin­ New Caledonia (Stoliczka, 1870-1871; awa Island (all 20-30 m in depth) (Figure 1). Woods, 1913; Dartevelle and Freneix, 1957 ; Freneix, 1960, 1972), also exhibit a similar Observations and discussions outline. Those Cretaceous species of Phygraea, however, have much thicker shells, Cryptic habitat. - The habitat of the pres­ and, as interpreted by Jablonski and Bottjer ent oyster is hardly accessible to us, but its life (1983), probably were cup-shaped recliners habit can be recognized through divers' talk on soft substrata. and underwater videotapes. Various cave On the other hand, fossil species of organisms seem to show spectral distribution Pycnodonte (s.s.) were mostly lifelong sessile from the entrance to the innermost part in animals, because the shell is not very thick accordance with the change of physical and and because the attachment area seems to biological factors. Individuals of this oyster 935. A living fossil oyster 1083

umbo cavity with vesicular structure

simple prismatic outer layer epibiont (scleractinian)

1/..-::I c: -~ E .r. o ftI -ftI

" , abandoned venter

Cave waif ,:'<: ' , geniculation vacuolated lens adductor muscle epibiont (sclerosponge) 10mm

Figure 7. Subvertical section of Pycnodonle (Pycnodonle) lanigllchii, n. sp. in living position (based on a sectioned articulated individual, UMUT RM 18913).

always grow on gloomy walls or ceilings of Various epibionts are observed on the sur­ submarine caves and other sheltered places face of both valves of this oyster. Bryozoans open to fore-reef slopes under normal salinity and serpulids are most common, and certain and temperature. Such a large-sized suspen­ species of scleractinian coral, barnacle and sion feeder as this oyster probably cannot live sclerosponge are occasionally found (Figure unless a considerable amount of phytoplan­ 7). A characteristic community consisting of kton is supplied from the exposed environ­ sclerosponges, articulated brachiopods, ment. It does not live in the totally dark bryozoans, annelids and scleractinian corals innermost part of such caves, where Neritopsis (commonly noncolonial) are found at or near radula and several characteristic minute mol­ the habitat of the present oyster. Such luscs are still found alive. sclerosponge-bearing communities have been The present oyster, on the other hand, does discovered at similarly sheltered places (main­ not dwell in the ordinary exposed environ­ ly crevices and caves) in a number of Carib­ ment. This is probably the main reason why bean and Indo-West Pacific coral reefs, and such a large-sized conspicuous bivalve had their ecological and paleontological signi­ not been discovered by scientists and collec­ ficance as well as their origin have been tors until quite recently. Moreover, owing to discussed by many authors (Hartman and the wide attachment area the LV seems to Goreau, 1970; Jackson, Goreau and Hart­ adhere to the cave wall for a long time after man, 1971; Jackson, 1977,1979; Kobluk, death, and the paper-thin RV, if disarticulat­ 1988; etc.). The present oyster is possibly ed, is easily broken. Dead shells of N. also a constituent of such a cryptic commu­ radula are found on exposed bottoms and nity at least in this region. beaches, but they were certainly transported Color dimorphism. -Of eight large speci­ from sheltered environments by hermit crabs. mens examined, two are ivory white and six 1084 Itaru Hayami and Tomoki Kase are purplish brown in shell coloration (Table inside of the shell. The formation and func­ 1). There is no intermediate individual, and tion of the strongly puckered venter are prob­ the color type of infant individuals is difficult ably similar to those of hyote spines in to determine. The two phena are not Hyotissa (Stenzel, 1971: p. N 1026). different in various characters other than shell Repeated reconstruction of ventral margins coloration. This fact suggests the presence of result in the stronger convexity of LV and dimorphism, but the sample size at each local­ shell thickening of the middle-ventral por­ ity is too small to ascertain their strictly tion. Judging from the occasionally im­ sympatric relation and to clarify the relative bricated simple prismatic outer layer, recon­ frequency. Because the habitat of this oyster struction of venter may occur also in RV, is very dark, selection can scarcely act on the though the abandoned venter must be readily variation of shell coloration. broken. Shell growth. - The shell shape of the pres­ The RV of this oyster is extremely thin and ent oyster changes significantly with growth. has almost no physical strength, but its flexi­ In the initial stage (up to 20-30 mm in shell ble distal part sticks so closely to the inner height) every individual attaches itself to the surface of LV that we can hardly examine the cave wall or some other hard object (includ­ soft parts without breaking the shell. The ing the shell surface of larger individuals) strongly puckered venter of LV and unusually with the whole surface of LV. In this stage broad commissural shelf may be advanta­ both valves are nearly flat or only feebly geous, because, as interpreted by Stenzel convex, the commissural shelf is still narrow, (1971 : p. N 1025), they make it more difficult and the test thickness is almost the same for shell-breaking predators to reach the soft between the two valves. Although the part. In fact, individuals with inj ured-and­ attachment area extends along the entire dor­ repaired shell are often met with in this sal margin of LV until the latest stage, the species. main part of the generated shell margin of LV Radial gashes of RV, which were called sits up suddenly apart from the wall surface, "tuck grooves" by Harry (1985), are a very and then the shell becomes very inequivalve characteristic feature in many pycnodonteine and concavoconvex, as shown in Figure 7. oysters. As observed in the present oyster, In some cases numerous infant shells of the they occur exclusively in the simple prismatic same oyster species adhere to the shell surface outer layer (Figure 2-lc, d, 5). Because this (Figure 2-la, c), but such individuals prob­ layer is commonly multilayered, the gashes ably cannot grow larger because we have rarely penetrate the shell except near the never seen any cluster of large individuals. ventral margin. They may also contribute to Functional morphology. - As seen in the the increase of flexibility of the distal part. dorsal view of many specimens, the attach­ In many fossil species of Pycnodonte, RV ment area is commonly geniculated at the looks much smaller than LV. The presence umbonal area (Figure 4-1). The chevron­ of radial gashes, however, strongly suggests like attachment area seems to strengthen the that those species had originally the flexible adherence. Divers could not collect any distal part of R V and concordant valve mar­ specimen without using a hammer. In many gins like the present species. grown individuals of this oyster the ventral Evolutionary ecology. - In view of the very surface of LV is strongly puckered, so that conservative morphology, the present oyster this portion often looks branched in cross can be regarded as directly descended from section (Figure 7). This feature indicates some Cretaceous and Paleogene species of that a preexisting ventral margin was aban­ Pycnodonte (s.s.). This, however, does not doned, and that a new margin grew from the necessarily mean that fossil species of 935. A living fossil oyster 1085

Pycnodonte were also cryptic animals. In crusting organisms on hardground and in fact, many small individuals of a pycno­ small crevices at the base of the Bradford Clay donteine oyster, as recognized by Hanai and (Bathonian) in England. As mentioned by Oji (1981), attach themselves to the upper Boucot (1981), this seems to represent an surface of a fossil beachrock in the Aptian of exceptional preservation of ancient sheltered north Japan. Bottjer, Roberts and Hattin biota. Palmer and Ftirsich recorded that the (1978) ill ustrated many individuals of upper (exposed) surface and the lower Pycnodonte kansasense [sic] encrusting the (sheltered) surface of the crevices were inhab­ surface of an inoceramid valve in the Tur­ ited by more or less different organisms. An onian Greenhorn Limestone of Kansas. unnamed species of Plicatula, a brachiopod, Kauffman (1967) and Tashiro (1978) also some serpulids and bryozoans could be cryp­ recorded similar occurrences respectively for tic organisms because they predominantly Pycnodonte? conjesta from the Colorado occur on the lower surface of crevices. Some Group in the Western Interior and oysters belonging to Liostrea, Exogyra, Pycnodonte amakusaensis from the San­ ? Nanogyra and Lopha are occasionally tonian muddy sediments of west Japan. found adhering to the lower surface, but they Kauffman and Sohl (1973) mentioned that occur more abundantly on the exposed sur­ small individuals of Pycnodonte often at­ face of the hardground. These crevices are tached themselves to the surface of lower only about 25 cm deep and measure up to 5 valves of rudists in the latest Cretaceous cm from roof to floor; their scale must be by rudist reefs of the West Indies. Jablonski far smaller than that of the submarine caves in and Bottjer (1983), on the other hand, regard­ which the present oyster dwells. ed Pycnodonte (Phygraea) mutabilis from the Why is the present oyster found only in Late Cretaceous offshore chalky sediments of sheltered places? If Cretaceous-Paleogene the Gulf and Atlantic Coastal region as an species of Pycnodonte were noncryptic, what iceberg strategist (cup-shaped recliner by drove the descendant to such a cryptic envi­ Seilacher, 1984). This is probably true ronment? There are two possible answers. about many species of the subgenus One is an explanation regarding the change of Phygraea, because of their gryphaeate mor­ habitat as a result of overgrowth competition phology. These Cretaceous species of among reef-building organisms; that is, in Pycnodonte, both sedentarists and recliners, modern coral reefs, as Jackson (1977) inter­ are thus not considered to have been cryptic preted for the restricted distribution of organisms. sclerosponge-fauna, rapidly encrusting colo­ Typical Cretaceous and Paleogene species nial organisms predominantly occupy the of Pycnodonte (s.s.) are generally character­ surface of exposed hard bottom, excluding ized by a longer dorsal margin, better devel­ therefrom slowly growing sessile organisms. oped auricular part and commonly thinner The other explanation is based on the shell than those of Phygraea. If the attach­ assumption that the predation pressure on ment area extends over the entire dorsal bivalves is relatively low in such a sheltered margin with growth, those fossil species environment. should be regarded as lifelong sessile organ­ Many sedentary bivalves of similar size are isms like the present species. Cryptic life of known in and around Indo-West Pacific coral these fossil species is still unlikely, because reefs. Spondylus varius, Hyotissa hyotis and they occur abundantly together with many Chama lazarus are often found alive on the other epifaunal and infaunal bivalves. exposed fore-reef bottom around the caves. Palmer and Ftirsich (1974) examined the Although little is known about the growth microgeographic distribution of various en- rates of these bivalves, it is unlikely that these 1086 ftaru Hayami and Tomoki Kase noncryptic bivalves grow much more rapidly glance, but many important characters of this than the present oyster. On the other hand, subgenus are well retained. In addition to they are characterized by much thicker shells the archaic morphology and evolutionary and more strongly armoured upper valves. stasis, its cryptic habitat strongly suggests that The strongly puckered venter and broad the present species is a significant example of commissural shelf may be effective to some "living fossils". Its morphology, physiology extent against predation, but the shell of this and ecology must be informative for under­ oyster appears to be more delicate and standing the paleobiology of fossil defenseless in comparison with such massive Pycnodonte and related oysters. sedentary bivalves in the exposed environ­ Many Cretaceous and Paleogene species of ment. Though nothing is known about the Pycnodonte, however, were probably non­ actual predators of this oyster, these lines of cryptic organisms, because they occur very evidence seem to suggest that the second abundantly together with other bivalves in explanation is more plausible in this case. various kinds of sediments. Two different Submarine cave faunas often contain life habits are assumed for fossil species of deeper-water elements. Kobluk (1988) Pycnodonte. One is iceberg strategists (sec­ attributed the cause of resemblance between ondary cup-shaped recliners), and the other is cryptic and deeper-water faunas mainly to lifetime sedentarists like the present species. analogous physical factors, especially to simi­ Iceberg strategy is assumed in various families larly low illumination level. Some subma­ of Mesozoic bivalves (Jablonski and Bottjer, rine caves of Ryukyu Islands yield many 1983; Seilacher, 1984), but is believed to incredibly extraneous bivalve genera and fam­ have become much rarer in the Cenozoic, ilies, the bathymetric distributions of which probably as the result of the increase of extend to or are restricted to lower neritic, powerful predators (Hayami and Hosoda, bathyal and even abyssal bottoms (Hayami 1988). and Kase, 1991 ; Kase and Hayami, in press). As advocated by Vermeij (1977,1987) and Deep-sea origin of the present oyster, how­ others, the increase of predation pressure after ever, is rather unlikely, because there is no the Mesozoic seems to have had a serious evidence for Pycnodonte to have migrated to impact on the evolution of adaptive strategy deep seas after the Paleogene. It may have of molluscs. Bivalve evolution seems to have survived for a long geologic period by wan­ followed several trends not only in morphol­ dering sublittoral cryptic environments. ogy but also in behavior and habitat. In many free-living bivalves escaping ability Conclusion: significance as a living fossil from predators (e.g., by burrowing and by swimming) has increased. Some sedentary Although there are several extant represen­ groups seem to have developed armored tatives of the Pycnodonteinae (Neopyc­ sculptures against the attack of durophogous nodonte and Hyotissa), this subfamily seems predators. Another effective way of surviv­ to have attained its acme in Late Cretaceous ing must be to change habitat to a place of and Paleogene times. The genus Pycnodonte lower predation pressure. Neopycnodonte, (including several subgenera) was a leading which is probably also a descendant of group of this subfamily during these periods, Pycnodonte, now survives in deeper water but there is no fossil record after the Middle than other oysters. It is generally supposed Miocene. As described above, the newly that submarine caves and other cavities as discovered archaic oyster is considered to be a well as exposed deep-sea bottoms often pro­ direct descendant of Pycnodonte (s.s.). Its vide suitable refuges for archaic and relatively shell morphology may look enigmatic at a defenseless organisms. 935. A living fossil oyster 1087

The inferred habitat change of Pycnodonte resembles those of the present species, but its LV is is noticeably similar in many respects to that very thick like Phygraea. It is unlikely that the Cenomanian oyster was a cryptic organism, as Dhon­ of Neritopsis, which is a famous "living fossil" dt regarded the species as having been an iceberg gastropod (Batten, 1984). Neritopsis radula, strategist on a soft substrate. She treated the gender which is often found alive together with the of the generic name Pycnodonte as neuter, though we present oyster in the same caves, shows very are ignorant about the reason. conservative morphology. Fossil records of Neritopsis are common in the Mesozoic and References Paleogene (especially in the Tethyan realm), but N. radula from the Indo-West Pacific and Batten, R.L., 1984: Neopilina, Neomphalus, and Neritopsis, living fossil molluscs. p.218-224. a closely related Caribbean extant species are In, Eldredge, N. and Stanley, S.M. eds., Living the only known representatives after the fossils. Springer-Verlag, New York. Miocene. To conclude, it is strongly suggest­ Bottjer, D.J., Roberts, C. and Hattin, D.E., 1978: ed that Pycnodonte and Neritopsis have been Stratigraphic and ecologic significance of surviving in shallow and warm seas for many Pycnodonte kansasense, a new lower Turonian oyster from the Greenhorn Limestone of Kansas. millions of years by transforming themselves Jour. Paleont., vol. 52, p. 1208-1218. into cryptic animals. We may be able to Boucot, A.J., 1981: Principles of benthic marine have a glimpse of Mesozoic-type biota in such paleoecology. 463 p. Academic Press, New York. a cryptic environment. Further studies of Carter, J.G., 1991: Chapter 10. Evol utionary organisms in submarine caves would provide significance of shell microstructure in the Palaeotaxodonta, Pteriomorphia and Isofilibran­ concrete and significant evidence to test vari­ chia (Bivalvia: Mollusca), Chapter 11-16. ous theories in evolutionary biology. Shell microstructural data for the Bivalvia. p. 136-412. In, Carter, J.G. ed., Skeletal biomineralization: patterns, process and evolu­ Acknowledgements tionary trends, vol. 1. Van Nostrand Reinhold, New York. We thank sincerely Dr. Kenji Torigoe of Chel'tsova, N.A., 1969: [A study of the micro­ Hiroshima University for his kind informa­ structure of the shell of some Cretaceous oys­ tion about the anatomical characters of the ters for their systematization.] 87 p. Academia present oyster. Our cordial thanks are also Nauk, SSSR, Moscow. Dartevelle, E. and Freneix, S., 1957: Moll usq ues due to skillful divers, Messrs. Mitsutoshi fossiles du Cretace de la c<>te occidentale d' Afri­ Taniguchi (Miyako-jima), Shuichi Ohashi, que du Cameroun ill'Angola. II. Lamellibran­ Shigemitsu Kinjo and Hiroyuki Kinjo (Oki­ ches. Ann. Musee royal du Congo Beige, sci. nawa), and Miss Miyoko Uchima (Okinawa) geol., vol. 20, p. 1-271, pIs. 1-35. for their self-devoting cooperation in collect­ Duff, K.L., 1978: Bivalvia from the English Lower Oxford Clay (Middle Jurassic). 137 p., 13 pIs. ing the material. Palaeontogr. Soc. London. Fleming, C.A., 1966: Marwick's illustrations of New POSTSCRIPT. At the preparation of this manuscript we Zealand shells. N.Z. Depart. Sci. In dust. Res., overlooked an important work (Dhondt, 1984: The Bull. 173, p. 1-455. unusual Cenomanian oyster Pycnodonte biauri­ Freneix, S., 1960: Etude complementaire des lamelli­ culatum. Geobios, Mem. special, no. 8, p. 53-61), in branches du Cretace de Nouvelle-Caledonie. which the morphology, distribution and paleoecology Sci. de la Terre, vol. 4, no. 1-2, p. 5-56, pIs. 1-3. of a Cenomanian oyster Pycnodonte (Pycnodonte) --, 1972: Les mollusques bivalves cretaces du biauriculatum (Lamarck, 1819) were described and bassin cOlier de Tarfaya (Maroc meridional). discussed in detail. It was said that the species Notes & Mem. Servo geol. Maroc, no. 228, p. 49- distributes widely (though episodically) along the 255. northern margin of the Tethys from the Iberian Penin­ Gardner, J.A., 1916: Systematic paleontology; Mol­ sula to Central Asia, sometimes forming crowded lusca. p. 371-733, pis. 12-45. In, Clark, W.B. fossil banks as a monospecific community. The shell and others eds., Upper Cretaceous [Maryland]. shape and long dorsal margin of P. (P.) biauriculatum Maryland Geol. Surv. 1088 Itaru Hayami and Tomoki Kase

Habe, T., 1977: Systematics of Mollusca in Japan. 135-147. Bivalvia and Scaphopoda. 372 p. Hokuryu­ Kase, T. and Hayami, I., in press: Unique submarine kan, Tokyo [in Japanese]. cave mollusk fauna: composition, origin and Hanai, T. and Oji, T., 1981: Early Cretaceous bea­ adaptation. Jour. Moll. Studies. chrock from the Miyako Group, Northeast Kauffman, E.G., 1967: Coloradoan macroinverte­ Japan. Proc. Japan Acad., 57, ser. B, vol. 57, p. brate assemblages, central Western Interior, 362-367. United States. Paleoenvironments of the Harry, H.W., 1985: Synopsis of the supraspecific Cretaceous seaway - a symposium, Colorado classification of living oysters (Bivalvia: School of Mines, Golden, May 1967. p.67-143. Gryphaeidae and Ostreidae). Veliger, vol. 28, -- and Sohl, N.F., 1973: Structure and evolution no. 2, p. 121-158. of Antillean Cretaceous rudist frameworks. --, 1986: Sententia: the relevancy of the generic Verhandl. NaturJ Ges. Basel, vol. 84, no. I, p. concept to the geographic distribution of living 399-467. oysters (Gryphaeidae and Ostreidae). A mer. Kobluk, D.R., 1988: Cryptic faunas in reefs: ecol­ Malacol. Bull., vol. 4, p. 157-162. ogy and geologic importance. Palaios, vol. 3, p. Hartman, W.D. and Goreau, T.F., 1970: Jamaican 379-390. coralline sponges: their morphology, ecology Kuroda, T., 1931: An illustrated catalogue of the and fossil relatives. In, Fry, W.G. ed., The Japanese shells (7). Venus, vol. 2, no. 5, Appen­ biology of the Porifera: Symposium of the Zoo­ dix p. 55-68. logical Society of London, vol. 25, p. 205-243. Nagao, T., 1928: Paleogene fossils of the island of Hayami, 1.,1988: Functional and taxonomic implica­ Kyushu, Japan, Part 2. Sci. Rep. Tohoku Imp. tions of internal ribs of Propeamussium. Trans. Univ., ser. 2, vol. 12, no. I, p. 11-140, pis. 1-17. Proc. Palaeont. Soc. Japan, N.S., no. ISO, p. Nicol, D., 1991: Location of continents and oceans 476-490. and the distribution of living oysters -- and Hosoda, I., 1988: Fortipecten takahashii, a (Gryphaeidae and Ostreidae). Papers in Florida reclining pectinid from the Pliocene of north Paleont., no. 5, p. 1-3. Japan. Palaeontology, vol. 31, p.419-444. Oyama, K., Mizuno, A. and Sakamoto, T., 1960: -- and Kase, T., 1991: Cryptic propeamussiids Illustrated handbook of Japanese Paleogene and other bivalves from a submarine cave of molluscs. 244 p., 7 I pis. Geological Survey of Okinawa, south Japan. Program and Abstracts, Japan. 1991 International Marine Bivalve Symposium, Palmer, K.V.W. and Brann, D.C., 1965: Catalogue of p.34. the Paleocene and Eocene Mollusca of the south­ Hudson, J.D. and Palmer, T.J., 1976: A euryhaline ern and eastern United States. Part I. oyster from the Middle Jurassic and the origin of Pelecypoda, Amphineura, Pteropoda, Sca­ the true oysters. Palaeontology, vol. 19, phopoda, and Cephalopoda. Bull. A mer. p.79-93. Paleont., vol. 48, no. 218, p. 1-466, pis. 1-3. lliffe, T.M., Hart, C.W. Jr. and Manning, R. B., 1983: Palmer, T.J. and Fiirsich, F.T., 1974: The ecology of Biogeography and the caves of Bermuda. a Middle Jurassic hardground and crevice fauna. Nature, vol. 302, p. 141-142. Palaeontology, vol. 17, p. 507-524. Jablonski, D. and Bottjer, D.l., 1983: Soft-bottom Riedl, R., 1966: Biologie der Meereshijh/en. 636 p. epifaunal suspension-feeding assemblages in the Verlag Paul Parey, Hamburg u. Berlin. Late Cretaceous. p.747-812. In, Tevesz, M.J. Seilacher, A., 1984: Constructional morphology of S. and McCall, P.L. eds., Biotic interactions in bivalves: evolutionary pathway in primary ver­ recent and fossil benthic communities. Plenum sus secondary soft-bottom dwellers. Palaeon­ Press, New York. tology, vol. 27, p. 207-237. Jackson, J.B.c., 1977: Competition on marine hard Siewelt, W., 1972: Schalenbau und Stammesgeschich­ substrata: the adaptive significance of solitary te von Austern. Stuttgarter Beitr. Naturk, ser. and colonial strategies. American Naturalist, B, no. I, p. I-57. vol. I I I, p. 743-767. Squires, R.L. and Demetrion, R., 1990: New Eocene --, Goreau, T.F. and Hartman, W.D., 1971: marine bivalves from Baja California Sur, Mex­ Recent brachiopod-coralline sponge communities ico. Jour. Paleont., vol. 64, p. 382-391. and their paleoecological significance. Science, Stenzel, H.B., 1959: Cretaceous oysters of southwest­ vol. 173, p. 623-625. ern North America. Congo Geol. Intern., XXa -- and Winston, J.E., 1982: Ecology of cryptic sesion, Ciudad de Mexico, 1956, EI sistema coral reef communities. I. Distribution and Cretacico, vol. I, p. 15-37. abundance of major groups of encrusting organ­ Stenzel, H.B., 1971: Treatise on invertebrate isms. Jour. Exper. Mar. Bioi. Ecol., vol. 57, p. paleontology. Part N, Part 3, p. N954-NI224. 935. A living fossil oyster 1089

Geol. Soc. America and Univ. Kansas. tion: evidence from snails, predators and Stephenson, L.W., 1941 : The larger invertebrate fos­ grazers. Paleobiology, vol. 3, p. 245-258. sils of the Navarro Group of Texas. Univ. --, 1987: Evolution and escalation. 527 p. Prin­ Texas Publ., no. 4101, p. 1-641. pIs. 1-95. ceton Univ. Press. Stoliczka, P., 1870-1871 : Cretaceous fauna of south­ Wade, B., 1926: The fauna of the Ripley Formation ern India. Vol. 3. The Pelecypoda. Palaeont. on Coon Creek, Tennessee. U.S. Geol. Surv., Indica, part 1-4, p. 1-222, pIs. 1-13 (1870), part Prof Paper 137, p. 1-272, pIs. 1-72. 5-13, p. 223-537, pIs. 14-50 (1871). Wilson, D., 1987: A new pycnodont oyster from the Tashiro, M., 1978: On some interesting bivalves from Pungo River Formation, and an annotated list of the Cretaceous Himenoura Group in Kyushu. the Cenozoic pycnodonts of the Atlantic and Trans. Proc. Palaeont. Soc. Japan, N.S., no. 110, Gulf coastal plain. Smithsonian Contr. p. 319-329, pis. 43,44. Paleobiol., no. 61, p. 13-20. Torigoe, K., 1981: Oysters in Japan. Jour. Sci. Woods, H., 1913: A monograph of the Cretaceous Hiroshima Univ., ser. B, div. I (Zool.), vol. 29, Lamellibranchia of England. vol. 2, pt. 9. no. 2, p. 291-419 (36 pIs. incl.). Palaeontogr. Soc. London, vol. 66, p. 341-473, Vermeij, G.J., 1977: The Mesozoic marine revolu- pIs. 55-62.

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l'frE:E:j(~~tJ'G9£JiI.~nt-:'t:·;7 / r"/T~O)Iffi.1:.f! (1:.§"·"Cl'91~:O;l;""'): -gtl7IJ~0) Ti1!!.~:t-o J: lfi'rpi':llli~ ® 1*:0) iWl.iti1PJilIl (i*1tfJ 20 m) 0)7W1fili l '~iii v.::. tJ, t.>: I) :kml 0) JiI,,\§}1I n t.>: l,;I; "'" tJ~9£JiI, ~ nt-:.o IDtMjffi· ~l'UIfl:l'tI\:~ • ~ff{ffl>a:~W L tda:llt .: O)f! ft ~ 'Y ::I '7 ;iJ'" "'" v.::.~ {, iii l ,tJ~, :m1:.O)Jlt~f! f.::.~~i- 9 {, 0) ftt.>: <, B llUc -tl5t=*cv'::' T 7- A iW~-c'iJ\?i * L, ,*,ffit!fx~ll.jI( L L!: 'j!j xJ:d't -C §" t-:. Pycnodonte ~O)JEtff!-c' t5 9': C tJ~1ljj G tJ'v'::' t.>:--:;t-:. o Pycnodonte (Pycnodonte) laniguchii [TP~: :;;J-:;;J-~ 'Y ::I '7 ;1;'''''' (ffif;i];)] c~fHt

-C~cax: L, {-0)*1~tl1:.!\'kJ"f:l¥J~~>a:~~-t" 9 0 {-O)Mtk· IDt13ftllt~v.::.@;tstJ~, ~tt.m ~O)7i-~t-': vttJ, G t.>: 9 :tiIDto) mH~mlfHl\;o) J: ') v.::.fi <, 1:.Bifv'::'Ii*~-c·, "jiif L a:~-t"o LtJ'L Pycnodonte(s.s.) 0)-t""'-C0)!f!fW: (17UxJi, *l''W~V.::.i''d--:;-ClAtJ~9~ ~iii, 'W{J!U v.::. {:Ii: !i:-t" 9 F9MO) F,fJIDi:~njJt $.l' commissural shelf, ~tko) chomata, i[!!O)*tk

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Benthic foraminiferal assemblages of two piston cores 7 ~.;; l' :1 ~1\:lI:.E G')ili'<# '7 ;/ ~ t tlffiJ:l[fi'i~ taken from south of the Ishigaki Island ...... il'Ii~ im

·································Xu Xuedong· Ujiie Hiroshi l!ID~iJ' I?.lr-t.::. ~ ~ -. :li:/ '7 G')ff:* U~~·········Lll Q §'f ~.reEKJ:;.,e.'7~~t~.L~.~e~MB*W ~.~ilf.=&ftG')~ME.1*Ktovt;., •••~G')R# G')wi7i'.Il£ilJG')MIJ.Ij····································'HR ~ ~~·······································!EI33EIDl· ~*m- *~ •• lI:.E UE~.HI:.E~m:m Lt.::.tl3l 'L ·······································f!!!:frlli* • Allmon, W...... i')Jj(I!j!f~ • Lll Q jlfz .~:ttfJ.Ktovt;.,*.~G')ft~.!RreEilf !lI,~:;t "'" ~ = !::"" A :ll1.G').lJHm%:t~············*~,Wt1.ilt ·············· .. ··············EMrrOO~~ • AJJ ~1J.Ij • 1tE -'< *Jl B*wlllffv-1-G')~.lr-~.lr-Kt~~5tlt~.~G'):/p *.~I::.tovt;., Dosinia japonica t D. tomikawensis G') '1 ~ :1]'1 ilfJf;························WkIlilJJt!!*J',(~ • tj"Lll*~ '5l:.If. tJ:j~JH::. ":> l 'L ...... ·· .. ·····~ttj" 5itjlf .tfitltt~tfitltKtovt;.,~~*G')~ • !:: -If '7 :Ii 1 t$"EG')~jjg t ~!ill····· .. ··m ~ • llii!lm& ...... !jfftj"f!::J<: • tmp~= . tfi~llilJ.lj '1 ~ ;t-!l IvG')~~gif.!ill UriIJ - <'f { T:;ti!l.~iJ'l? e~e/.~eJ:l[~Ktovt;.,'*'~*E~~R.!RG')~~...... ;; 3;/ '1' 7 :=. '" • i!iiJli9lime $ ...... ····wili'<'ffl::J<:

;tJJF,,'*ffj!l)::.~tJ'7·;/ f ~ '1 :=. G') Btl ,t!. u::.xt-t;":[email protected] i.&J1s:. I mtif.t::.tovt;., ~JL.!RG')*:;:~.G') 2l'ii "'wili'<'ffl::J<: ~Hil?r ...... ·:lil:lR.illft- Cenomanian/Turonian J:l[~I::'tovt ;.,l!7t* ~ &1*lt A /~ Soft bottom l::.tovt;.,1jt1*~MIIlIIl\\JlG') growth form 1 ~ t~~JJ:tG')xtr.t ...... ··~l33bl*c'f - ~twJ!i:~?i'i~ilf,*,G')~~iJ;f.f":>.'* - ···*1'tJIIlj[

Palaeontological Society of Japan (PSJ) Council Actions

During its meeting on January 24, 1992, the PSJ Council enacted the following changes to its membership.

New mem bers elected : Yuuji Abe, Toshiaki Futagawa, Kouzi Hasegawa, Tomoteru Hiyoriyama, Yoshiyuki Hayashi, Yoshinori Hikida, Haruki Horikawa, Masumi Ichikawa, Shigeru lnaba, Seiji Kakuta, Takafumi Katoh, Sanao Kashikura, Hakuichi Koike, Minoru Koitabashi, Ken-ichi Kouko, Katsue Koya, Yukihide Matsumoto, Hiroshi Matsuo, Takehiro Morishige, Masafumi Murayama, Takehiro Nakamura, [093

Takashi Nakashima, Ricardo Franco-Nieto, Hiroyuki Oishi, Itsuro Oshiro, Johann G. Rigor, Kazuyoshi Ryuzaki, Takeshi Saito, Takeyoshi Saito, Tak uj i Sasaki, Takehiro Sato, Chieko Shimada, Hiroko Suzuki, Harumi Sugawara, Ken Takagi, Michio Takahashi, Isamu Terui, Yutaka Tsubaki, Shigeki Togashi, Hideo Yabe, Koichi Yoshiba

New Fellows elected; Kazumi Akimoto, Ren Hirayama, Ryuichi Majima, Tomoko Matsuda, Atsushi Matsuoka, Keij i Matsuoka, Tomio Nakagawa, Hideo Nakaya, Harufumi Nishida, Hiroshi Nokariya, Masayuki Oishi, Kiyoshi Okumura, Osamu Sakamoto, Tokiyuki Sato, Toshio Takagi, Susumu Tomida, Akifumi Tomizawa, Y oshitaka Yabumoto

Resigned members; (Fellow) Kazumi Suyari (Ordinary members) Tsutomu Kinoshita, Hidekazu Yoshida

Deceased members; (Fellow) Toru Makino (Foreign members) Li-Sho Chang 1094

Bibliography 1986-1990 (7) flJ ff ,: ""'J \, '-r

B*<5i£4&.J't~-C'f:l:., 1986-1990~0) 5~rs'IC0~~ht.:~Jt (mt'!AAliJtuJ: (J'T ~:;/7 ~ 71 O)~o'l'$ ~~IfffJt'l¥) ~tlHJIHcJ: I) flJrr-f1.>':' C Ict! I) i Lt.:o ':'0) Bibliography ICf:l:.ltt*illi I) B*<5i£4&.J't~~Jl.O) RIiti)(~cp.c}lc~J(~3& L i -ftJ;, B *0)~f4HlI1"? t.:OOF9 • 007l-0)?F~ jbj;0~ Lt.:ilIitiJt Jb -2;'00 1.> .:. Uc L i -fo Jt~O)~~f:l:.%~Jl. 0) $15~£lif! c V(rrt! 0 ,t.: 0' c ,i!\o' i -fO)-c', J:.ile 5 ~5tO)~IitiJt ~ A r ~r O){JUIC ltt"?--C{'PIlltL. 1992~7 )131 B~lcu.iZ';l)r~o'o Jt~ ~ A r{'PIlltIC~ L --C l:l:.r~eO)l±/i!~jP1l:~~w& l--Cr ~ 0'0 t!U, /F1JJl t!.:. c iI;lll) i Lt.: i?, ~ Jl.~ ~IH.:.':itJt-r~o'o

[ l±/i!~J1!l] L AAliJtf:l:.0~0)~f(;II~IC3'1 7' ('7 -7' P~~-2;'tr) L--Cr~ 0'0 7r:O)A 3'1 /vU J: lJ';t}1\l!ii:fIllil!JH:l:.ltt* 0) Bibliography Icltt 0' i -f 0 (AAliJtflh' c L --ct}1\l!ii:lcFn,MjICt! 1.> c ~ f:l:.~ ~~-c'~W1 ~-lt--Crn < .:. c tJ;;t; I) i-fo 2, ~M,!~-c't! 0 ,:Jl<'!O)AAliJt-c' Jb:@:--C f'Plllt L --Cr ~ 0'0 3. *i~t.i5, ~, ~, Jl:, ~I11ii, uJ:U,!~i5f:l:.1!iimt-U!o,-c'r~o,o 4. AAliJt3'1 r /J...f:l:.gxJtO)~fif:l:.gxJt-C', :fIlJtO)~fia:fllJt"eil~, :1<::k:fllJt.gxJto) 3' 1 r /v;t; 1.> o'f:l:.7r: O)~, ~§'O)ll1W<, {fffl~~{JUlcltto':'5ht!o'J: 5IcL--Cr~0'o 5. AAliJtcp-c'1lI1"? --C 0' 1.> F9'GO)5t~llf • 1liff(;1:I:. Index O)m~~~~, l, &;,-nIfffJt.:.' c IcileA L --C r ~ 0 '0 6. 7r:hi'hO):w:r'5tfF-c'?F~~ (#lc7I-OOA) O)liIf~~lcJ: 1.> B*0)~*4~1lI1"?t.:gIitiJtlcu3K#~0)~ fiICI:l:.JjrJ CIDJ;l::-c' ~ A r ~f'Plllt l--Cr~ 1.> ii" 7r:O)§'u:mi?-ltr~o'o

AAliJt~Ar.iZ';#7t =r 305 "':)

[ fJU]

WA TANABE Kozo (1973): Proflislilinella assemblage in the Omi Limestone, Niigata Prefecture, central Japan (Studies of Carboniferous fusulinacean of Omi, part I), Transactions and Proceedings of the Palaeontological Society of Japan, New Series, no. 95, pp. 371- 394, Plates 51-53, Figure I, Tables I-I L Carboniferous Fusuline

BANDO Yuji and KATTO Jiro (1980): On the Upper Triassic ammonoids from the Sampozan Group at Hitsuzan, Kochi City in Shikoku. In A. Taira and M. Tashiro (eds.): Selected papers in honor of Prof. Jiro Katto. Geology and Paleontology of the Shimanto Belt. Rinyakosaikai Press, Kochi, pp. 95-100. (~:mm~LIJ 0) :=:::i:llJ!illf J: I) Il€tfj Lt.::=::~*e1&JtIlo) 7 /'.f: 7 1 r IC"':) 0 '--C) (J,E,) Triassic Ammonoidea 1095

HANZA W A Shoshiro (1961): Cretaceous and Tertiary three-layered larger Foraminifera and their allied forms: their classification and geographical and stratigraphical distributions. Fossils (Palaeontological Society of Japan), no. 2, pp. 1-24, Figures 1-29, Tables 1-2. (f&JtIlB.ilUc· ~=:*c=:Jl'Il*~1l1L!R&V' -i"O)ilIfbl:fif!0)5H]lt.;: IS> V'~.:.Jm:F1f1,¥i¥J • Jl'IlilL'¥i¥J5j';fp) (J.) Cretaceous to Tertiary Larger Foraminifera [~~t.,*, O)*)(~JHj# ~ ;fO)(~Iiii)( O){JU]

KOBA Y ASH 1 lwao and KAKIZAKI Takeo (1978): [Preservation of vertebrate bones in nodules from the Tsurushi Formation]' Saito Ryojiro Sensei Taishoku Kinenshi, pp.43-49, Plates 1-2. (~TJW'@: / :: ~ -/H.:.§J:!!!~nt.::1'tftEO)i*m (J.) Miocene Mammalia

Taxa index for the bibliography

ANIMALIA ANNELIDA General MOLLUSCA Problematica, Trace fossils Cephalopoda and others Coleoidea CHORDATA Ammonoidea Mammalia Nautiloidea Proboscidea Mollusca excl. Cephalopoda Artiodactyla Bivalvia Aves Scaphopoda Reptilia Gastropoda Amphibia Polyplacophora and Monoplacophora Pisces COELENTERATA Osteichtyes Anthozoa Chondrichtyes Zoantharia PROTOCHORDATA Tabulata Graptolithia Scleractina Conodontochordata Tetracorallia ECHINODERMATA Octocorallia Holothuroidea Stromatoporata Echinoidea Hydrozoa Crinoidea Scyphozoa BRACHIOPODA PORIFERA BRYOZOA PROTOZOA excl. MASTIGOPHORA ARTHROZOA Radiolaria Insecta Rhizopoda Crustacea Foraminifera Malacostraca Fusuline Cirripedia Larger Foraminifera Ostracoda Benthic Smaller Foraminifera Branchiopoda Planktonic Foraminifera Trilobita 1096

VEGETABILIA PTERIDOPHYTA General Pteropsida Problematica and others Articulatae Palynology NON TRACHEOPHYTA ANGIOSPERME Charophyta Dicotyledoneae Calcareous Algae GYMNOSPERME Coccolithophoridae Coniferopsida Dinoflagellata Ginkgopsida Diatomae Cycadopsida Silicoflagellata Pteridospermopsida Others

Geologic age index for the bibliography

GENERAL OR INDEPENDENT TO AGE LATE CRETACEOUS PRE-CAMBRIAN MESOZOIC TO CENOZOIC PALEOZOIC LATE CRETACEOUS TO TERTIARY OLD PALEOZOIC CENOZOIC CAMBRIAN TERTIARY CAMBRIAN TO ORDOVICIAN PALEOGENE ORDOVICIAN PALEOCENE SILURIAN PALEOCENE TO EOCENE SILURIAN TO DEVONIAN EOCENE NEW PALEOZOIC EOCENE TO OLIGOCENE DEVONIAN OLIGOCENE CARBONIFEROUS OLIGOCENE TO MIOCENE CARBONIFEROUS TO PERMIAN NEOGENE PERMIAN MIOCENE PALEOZOIC TO MESOZOIC MIOCENE TO PLIOCENE PERMIAN TO TRIASSIC PLIOCENE MESOZOIC PLIOCENE TO PLEISTOCENE TRIASSIC NEOGENE TO QUATERNARY JURASSIC QUATERNARY JURASSIC TO CRETACEOUS PLEISTOCENE CRETACEOUS PLEISTOCENE TO HOLOCENE EARLY CRETACEOUS HOLOCENE

------0 ------

Erratum

MA TSUMOTO, T A TSURO On some acanthoceratid ammonites from the Turonian of Hokkaido. Trans. Proc. Palaeont. Soc. Japan, N.s., no. 164, p. 910-927, Decem­ ber, 1991. Figure 6 caption, printed on a gummed-label and inserted in this number, should be substituted for the erroneous figure caption appearing on page 920 of No. 164.

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~ IT ~ El * ~ ':t ~ ~ ~ 1992if. 4 Jl 27 B ~p jjliJ T 113 }1:[Ji':jIfl))CglR::,~:i65-16-9 1992if. 4 Jl 30 B ge IT B::~~~tlJ-t /'?-P'.J 'I[ ~ 03-5814-5801 ISSN 0031-0204 $I ~ ~ *11i~.iI . ~ rg. B ::E1:.1o/.J~~¥IH!j··;j1C~ $i~:ji;f$ Jiij, :: § 'lii. ¥r 1 6 5 % • ~IJ MiiJ ~ ftll 13 m~ it IR /7, T (J) El l1§ IBJ 8-45 2.500 F9 ttE~:±lJiHPjjlIJ1*J'\~t± ttE~*~ ::t± 022-288-5555 }1:[Ji': 3455-4415 Transactions and Proceedings of the Palaeontological Society of Japan

New Series No. 165 April 30, 1992

CONTE:NTS

TRANSACTIONS 932. Kiyoshi Okumura and Yutaka Yamagishi: Molluscan fauna from the Late Miocene Hatsuse Formation in the Miura Peninsula, Kanagawa Prefecture, Japan ...... 1009 933. Ryuichi Majima and Akihiro Murata: Intraspecific variation and hetero­ chrony in Phanerolepida pseudotransenna Ozaki (Gastropoda: Turbinidae) from the Pliocene Nobori Formation, Pacific side of southwestern Japan ... 1024 934. Katsumi Ueno: Verbeekinid and Neoschwagerinid fusulinaceans from the Akiyoshi Limestone Group above the Parafusulina kaerimizensis Zone, southwest Japan...... 1040 935. Itaru Hayami and Tomoki Kase: A new cryptic species of Pycnodonte from Ryukyu Islands: A living fossil oyster...... 1070

PROCEEDINGS ...... 1090