ISSN 0031-0204

Transactions and Proceedings

of the Palaeontological Society of Japan

New Series No. 148

Palaeontological Society of Japan September 30, 1986 Co-Editors Hisayoshi 100 and Takashi HAMADA

Officers for 1985-1986

President: Toshimasa TANAI Honorary President: Teiichi KOBAYASHI Councillors; Kiyotaka GHINZEI, Takashi HAMADA, Tetsuro HANAI, Yoshikazu HASEGAWA, Itaru HAYAMI, Hisayoshi IGO, Junji ITOIGAWA, Tadao KAMEl, Tatsuaki KIMURA, Tamio KOTAKA, Kei MORI, Ikuwo OBATA, Tsunemasa SAITO, Yokichi TAKAYANAGI, Toshimasa TAN AI Members of Standing Committee: Kiyotaka CHINZEI (Membership), Takashi HAMADA (Co­ Editor of Transactions), Yoshikazu HASEGAWA (Foreign Affairs), Itaru HAY AMI (Planning), Hisayoshi IGO (Co-Editor of Transactions), Tatsuaki KIMURA (Finance), Ikuwo OBATA (General Affairs), Yokichi TAKAYANAGI (Editor of "Fossils"), Juichi YANAGIDA (Editor of Special Papers) Secretary: Hiroshi NODA (Editor of Transactions) Auditor: Yoshiaki MA TSUSHIMA

The fossil on the cover is Unuma (Spinunuma) echinatus ICHIKAWA and YAO, a Middle multisegmented radiolaria from Unuma, Gifu Prefecture, central Japan (photo by A. YAO, x 260).

All communication relating to this journal should be addressed to the PALAEONTOLOGICAL SOCIETY OF JAPAN c/o Business Center for Academic Societies, Yayoi 2-4-16, Bunkyo-ku, Tokyo 113, Japan Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 409-421, pis. 81-85, September 30,1986

816. SOME INOCERAMIDS (BIV AL VIA) FROM THE CENOMANIAN () OF JAPAN-I NEW OR LITTLE KNOWN FOUR SPECIES FROM HOKKAIDO AND KYUSHU*

TATSURO MATSUMOTO

c/o Department of Geology, Kyushu University 33, Fukuoka 812

and

MASAYUKINODA

Minami·Oita Junior High School, Oita 870

Abstract. In a series of papers in 3 parts some new and little known ino· ceramid species from the Cenomanian of Japan are to be described. In this part we describe the following four species including three new ones. (1) Birostrina sp. nov., which belongs to the group of B. concentrica but has a radial groove on the posterior part and ribs of moderate intensity in the late growth-stage. It is repre­ sented by a number of specimens from the Middle Cenomanian of the Mifune (central Kyushu), Mikasa (central Hokkaido) and Kotanbetsu (northwestern Hok­ kaido) areas. (2) Inocermus sp. nov., which is allied to 1. comancheanus but has a subrectangular outline with less convex valves and weaker subcostae. Its holotype came from the Lower Cenomanian of the Mikasa area. (3) 1. heinzi Sornay, which is represented by several specimens from the Upper Cenomanian and/or lowest part of the Lower Turonian of the Saku area (northwestern Hokkaido). They are very similar to the holotype from Madagascar. (4) Mytiloides sp. nov., which has a larger beak angle and more convex anterior margin than M. submytiloides and is char­ acterized by regular concentric ribs combined with fine concentric lirae. Its holo­ type came from the Zone of Euomphaloceras septemseriatum (Upper Cenomanian) of the Mikasa area.

Introduction commISSiOn on Cretaceous Stratigraphy of the lUGS (1976-1984), we have obtained a great Through the field works in various areas of amount of mega-fossils which may be useful Japan to cooperate in the IGCP Project No. 58 for biostratigraphic and other aims. The major Mid-Cretaceous Events (MCE) and (1975-1982) parts of the collections are those of ammonites also in the Regional Working Group for the Sub- and inoceramid bivalves and some of them have been and would be evaluated in improving the *Received March 20, 1985; Read June 16, 1985 and biostratigraphic zonation and at Osaka City University. correlation.

409 410 Tatsuro MATSUMOTO and Masayuki NODA

In a series of papers entitled at the heading, primarily on external characters should be we are going to describe some species of the revised on the ground of more important interior Inoceramidae from the Cenomanian of Japan, features, but has not yet given fully his scheme which have been recorded only preliminarily of classification nor his description of generic or left untouched for full description. As the diagnosis, except for a few genera, e.g. Myti­ persons concerned with the cooperative study [oides. At present, we have no competent idea vary case by case, the results are to be published of our own scheme of overall taxonomy and successively in three parts by different author­ have to depend on the existing concepts of ships. Dobrov and Pavlova (1959), Cox (1969), Seitz For some reasons, some species from the (1961, 1965, 1967, 1970), Sornay (1966), Cenomanian of Japan are represented by a small Dhondt (1983) etc., as well as Kauffman (op. number of specimens. This is probably due to cit; 1977), with some modification. our collection failure and indeed unfavourable For the terms to describe morphological for getting an adequate conception of a species. characters we follow primarily those defined It would be, however, better to describe them by Nagao and Matsumoto (1939), which have at this moment than to leave them unrecorded been used by Noda (1983,1984 etc.) and Matsu­ for a long time, so that someone may obtain a moto and Noda (1985) with some modification. satisfactory sample of population in future to Readers may refer to Efremova (1978) to com­ revise the present definition. In Part I we de­ pare them with other authors'. scribe four species of which three belong to such The symbols (or letters) to abbreviate the category. The material of this study is the dimensions and angles in the tables of measure­ specimens from the Mikasa, Saku and Kotan­ ments in this paper are as follows: betsu areas of Hokkaido, but one of the species h=height, l=length, H=maximum linear dimen­ depends also on those from the Mifune area sion from beak to posteroventral extremity, of Kyushu which were once treated by Tamura L=ditto perpendicular to H, s=length of and Matsumura (1974). hinge line (previously indicated as HL), b= The repositories of the described or men­ breadth (previously indicated as th or T), tioned specimens are as follows with abbrevia­ a=anterior hinge angle (angle between hinge tion in parentheses: line and anterodorsal margin), ~=angle of Geological Collections, Kyushu University, umbonal inflation, 'Y=posterior hinge angle Fukuoka (GK) (angle between hinge line and posterior Geological Collections, Faculty of Education, margin), r=obliquity (angle between axis of Kumamoto University, Kumamoto (KE) growth and hinge line); L=left valve, R=right Institute of Geology and Palaeontology, valve Tohoku University, Sendai (IGPS) The terms to describe the concentric orna­ University Museum, University of Tokyo, mentation vary between authors and also in Tokyo (UMUT) accordance with the characters. In this paper we use the following terms: Systematic descriptions concentric lines or lirae: fine markings on the Family Inoceramidae Zittel, 1881 surface of shell, which are not impressed on the internal mould, except in the case of the As to the authorship of this family we follow composite internal mould Vokes' (1980) indication. concentric rings: fine concentric elevations Kauffman (in Kauffman et al., 1977 and of the third order, which may correspond to elsewhere) commented that the current tax­ the concentric lines or lirae but are somewhat onomy of this family at generic level based more elevated than them and may be dis­ cernible even on the internal mould 816. Cenomanian Inoceramids from Japan-I 411

concentric subcostae or riblets (minor ribs): et Matsumoto; Tamura and Matsumura, concentric elevations of the second order, p. 49, pI. 1, figs. 1-5. well recognized even on the internal mould, 1976. Inoceramus concentricus nipponicus being larger (i.e. thicker and higher) than Nagao et Matsumoto; Noda and Matsu­ the concentric rings but somewhat smaller moto, pI. Cr-31, fig. 2. (narrower and lower) than and often branched 1976. Inoceramus concentricus costatus Nagao from or intercalated between the concentric et Matsumoto; Noda and Matsumoto, pI. elevation of the first order Cr-31, fig. 3. concentric ribs or ridges (major ribs): con­ 1977. Birostrina n. sp., Kauffman, p. 175. centric elevations of the first order. When they are broad but low or weak the term Material:-Holotype, KE 1976 (PI. 81, Fig. 1), concentric undulations may be employed. composite internal mould of left valve from loco concentric depressions: concentric furrow 4 (Yaseto, Mifune area, Kyushu) of Tamura and along the constricted or geniculated part of Matsumura (1974). Paratypes, KE 1977 from the shell in the case when the shell shows an loco 1 (Omine), KE 1978 from loco 5 (Okadake abrupt change in convexity. Playground), KE 1979 from loco 2 (Asanoyabu) Radial or divergent ornaments do not occur and KE 1706 from loco 3 (Subayashi) of the Mifune area; also GK. H 10164 and H 10165 in the described species, except for a radial both valves, GK. H 10166, small left valve, and sulcus or groove in a species. Radial markings which reflect the interior others (GK. H 10167-H 10169) from loco Ik shell morphology may appear on some well 2021a, Pombets gorge, Mikasa district (colI. M. Noda and T. Matsumoto); GK. H 10082, H preserved specimens, but they should not be 10157-H 10159 from loco R 100 and GK. H treated as ornamentation. 10153-H 10156, H 10160-H 10162 from loco R 101 of the Kotanbetsu area (colI. T. Matsu­ Genus Birostrina J. Sowerby, 1821 moto and H. Okada). Type species:-Inoceramus sulcatus Parkinson, Diagnosis:-Rather small, inequivalve, asym­ 1819. metrically ovoid shell of the group of Birostrina Diagnosis:-Shell small, inequivalve, with concentrica, characterized by the presence of a narrow and prominent left umbo and prosogy­ radial sulcus on the posterior part and the rous beak; hinge line short, without or with very development of rather irregular concentric ribs narrow posterodorsal auricle; surface ornamented and riblets in late growth stage in addition to with fine concentric lirae and with or without finer concentric lirae or rings. concentric ribs, riblets and/or radial folds. Description:-The holotype (PI. 81, Fig. 1) Remarks:-Presence of the radial folds is is small and probably not a full-grown indi­ not a generic criterion and the genus is best vidual, but it does show diagnostic characters. represented by Inoceramus concentricus Parkin­ Tamura and Matsumura (1974, pI. 1, figs. 1a- son, 1919. Although the group of 1. concentricus 1b) have carefully given two figures in somewhat was taken too much comprehensively by Nagao different views, which clearly show the posterior and Matsumoto (1939), Birostrina evidently radial sulcus and the details of the posterodorsal ranges up to the Cenomanian in the North part as well as other features, although the Pacific region. authors did not mention these two characters in the text. KE 1979 (Tamura and Matsumura, Birostrina tamurai sp. nov. 1974, pI. 1, fig. 5; this paper PI. 81, Fig. 3) is somewhat larger than the holotype but deficient PI. 81, Figs. 1-6; PI. 82, Figs. 1-3; in the anteroventral and posteroventral portions. PI. 83, Figs. 1-3; PI. 85, Figs. 4-5 KE 1977 (Tamura and Matsumura, 1974, pI. 1, 1974. Inoceramus concentricus costatus Nagao fig. 2; this paper PI. 85, Figs. 4-5) is also in- 412 Tatsuro MATSUMOTO and Masayuki NODA complete but the largest of the Mifune speci­ is gently concave below the umbo and broadly mens, attaining to about 75 mm in H in a re­ convex in the lower part towards the antero­ stored outline. GK. H 10164 (PI. 81, Fig. 6; ventral edge, where it is bent to the asym­ PI. 82, Fig. 1) from Mikasa and GK. H 10153 metrically rounded ventral margin, which passes (PI. 82, Fig. 2) from Kotanbetsu may represent to the gently arcuate posterior margin. The the normal size of this species. The right valve narrow posterodorsal edge is somewhat flat­ of the former is badly eroded, but this deficiency tened. is supplemented by another specimen, GK. H A radial sulcus runs on the posterior part 10165 (PI. 81, Fig. 5; PI. 82, Fig. 3) from Mikasa, of the disk near the margin of the umbonal which however, is somewhat distorted. GK. inflation in the left valve, extending towards 10154 from Kotanbetsu has both valves and the posteroventral part, where the ribs may more right valves are preserved in the Kotan­ show slight sinuosity on crossing the shallow betsu sample (e.g. GK. H 10155-10157, H sulcus. A faint, corresponding sulcus is dis­ 10159-10161). Shell layer is partly preserved cernible on some right valve specimens, e.g. in some Kotanbetsu specimens. GK. H 10154 and H 10155. On the basis of a considerable number of The shell is ornamented with numerous these specimens the specific characters are well concentric lirae or rings and rather irregularly recognized. Evidently the left valve is more developed concentric ribs and riblets. The convex than the right, having a fairly inflated, lirae or rings are distinct on the young part prominent but narrow umbonal part whose beak (e.g. up to 25 mm in H in the holotype) as fine is considerably curved inward and forward, and dense markings on the shell surface and exceeding the hinge line. sometimes even on the internal mould, some The hinge line is nearly as long as or slightly of which may be somewhat stronger than others. longer than a half of shell length, forming an Major ribs and branched or intercalated minor angle (a) of about 80° with the anterior margin ribs are developed in late growth stages with and that (r) of about 140° with the posterior rather irregular intensity and distance, being margin. The axis of growth extends towards combined with the finer rings on the inter­ the posteroventral extremity nearly straightly spaces. Major ribs extend to the posterior part, or obliquely with a forward gently concave whereas minor ribs or riblets and rings on curvature, forming an angle m of about 40° the main part of the disk fade away as they to 50° with the hinge-line. The anterior margin approach the zone of radial sulcus. Some of the

Explanation of Plate 81

Figs. 1---U. Birostrina tamurai sp. nov...... Page 411 1. Holotype, KE 1976 (LV), from Mifune; lateral (a), anterior (b), posterior (c) an d dorsal (d) views, X 1. 2. 2. Replica of holotype in the lighting somewhat different from 1a to show the posterior sulcus, x 1. 3. KE 1979 (LV) from Mifune, lateral view, x 1. 4. GK. H 10166 (LV), from Mikasa; lateral (a), anterior (b) and posterior (c) views, xl. 5. GK. H 10165 (BV) from Mikasa; left (a) and right (b) lateral, anterior (c) and posterior (d) views, x 1. 6. GK. H 10164, left lateral view in the disposition a little different form PI. 82, Fig. la, x 1.1. BV: both valves, LV: left valve, RV: right valve. Photos. (Pis. 81-85) by M. Noda. MATSUMOTO and NODA: Cenomanian Inoceramids from Japan-I Plate 81

ld 816. Cenomanian Inoceramids from Japan-I 413

Table Measurements of selected specimens.

Specimens h l/h H L L/H b b/h s/l ex i3 r Ii KE 1976 (L) 34.5 26.0 .75 37.0 26.7 .72 14.0 .41 1B.0 .61 BOo 60° 140° 50° GK. H 10164 (L) 59.6 36.7 .62 62.5 3B.0 .61 27.0 .45 19.0 .52 BO° 55° 1300 56° GK. H 10164 (R) 52.B 36.7 .70 56.B 3B.0 .67 19.0 .52 130° 560 GK. H 10153 (L) 4B.4 34.2 .71 1B.5 55° GK. H 10157 (R) 40.0 30.B .77 42.0 31.6 .75 12.0 .30 ~15.4 ~.50 100° 1370 50°-65° GK. H 100B3 (.L) 50.0 42.0 .B4 5B.6 37.B .65 21.4 .42 21.4 .51 90° 55° 145° 500 ribs extended from the main part of the disk from the Albian of England. This is interesting, reach the narrow, flat, posterodorsal edge with but B. tamurai has concentric ribs of moderate gradual weakening, as seen in the well preserved intensity in the late growth-stage. The young specimens (e.g. holotype and GK. H 10082). shell of B. tamurai ornamented with fine con­ Etymology:-This species is dedicated to centric lirae and rings may be similar to that Professor Minoru Tamura of Kumamoto Uni­ Albian species, which has not yet been clearly versity who has contributed much to the palae­ described nor illustrated. The sulcus may be ontology of the Mesozoic Bivalvia. the same kind of radial plication as those of Comparison and discussion:-We owe much B. sulcata (Parkinson, 1819) and B. subsulcata to ErIe G. Kauffman (1977, p. 175) who sug­ (Wiltshire, 1879) (see Woods, 1911). gested that the presence of the posterior radial Occurrence:-The type locality is Yaseto, sulcus should be evaluated as a specific char­ Toyono-mura, Shimomashiki-gun, Kumamoto acter in Birostrina. In fact, the sulcus is observ­ Prefecture (central Kyushu). This and other able in a number of examined specimens from nearby localities are recorded by Tamura and particular beds of the three areas. In some Matsumura (1974). They are all in the lithic specimens (e.g. GK. H 10082 figured by Noda sandstone of the Lower Formation of the Mifune and Matsumoto, 1976; this paper PI. 83, Fig. Group. Eucalycoceras sp. cf. E. spathi (Col­ 3) the sulcus is faint. lignon) is recorded to occur in the same forma­ The lectotype of Inoceramus concentricus tion (Matsumoto in Tamura and Matsumura, var. costatus Nagao et Matsumoto, 1939 was 1974). On the evidence of this ammonite and designated by Tamura and Matsumura (1974, p. other associated fossils (Tamura, 1979) the 49) to the one illustrated by Nagao and Matsu­ Lower Mifune is assigned to the Middle Ceno­ moto, 1939, pI. 24, figs. 1a---"1!, i.e. UMUT. MM manian. 6480 [=1-690]. It has no posterior sulcus and is Other described specimens came from two distinguished from B. tamurai. It resembles places. One is loco Ik 2021a at Pombets gorge closely Inoceramus concentricus var. porrectus in a tributary of the River Ikushumbets, Mikasa Woods (1917, pI. 4, figs. la-b), as Nagao and district (central Hokkaido), in the Mikasa Sand­ Matsumoto (1939, p. 271) have already pointed stone between the layers rich in trigonians. out. Kauffman considered it as identical with Calycoceras cf. C. asiaticum (Jimbo) occurs B. porrecta (Woods) from New Zealand. We fragmentarily at loco Ik 2021b in a trigonian would defer the conclusion about this point bed. For the route map and stratigraphic section until we could examine more carefully the see Matsumoto (1965, figs. 3 and 5). The other specimens from New Zealand and Japan. is locs_ R 100 and R 101, on road side of High­ Kauffman (op. cit.) compared the species way 239, about 800 m linearly west of Kiri­ represented by Tamura and Matsumura's speci­ tachi Pass, Kotanbetsu area (northwestern mens (that is now called B. tamurai) with an Hokkaido), from calcareous nodules in the undescribed species, "Birostrina n. sp. (sulcate)", mudstone of the Middle Yezo Group. Caly- 414 Tatsuro MATSUMOTO and Masayuki NODA

coceras sp. of newboldi group was found from than long later. Hinge plate fairly thick; hinge loco R 100. For the map and stratigraphic sec· line moderately long, somewhat more than half tion of this route see Matsumoto and Okada of shell-length. Anterior hinge angle (O!) about

0 0 (1973, figs. 7-8). 105 and angle of umbonal inflation «(3) 85 • To sum up, the known age of Birostrina Anterior margin nearly straight, with very gently tamurai is Middle Cenomanian. The fossils arcuate curvature towards rounded anteroventral occur separately in the Mifune area but in groups edge; ventral margin rather broadly and asym­ often with both valves conjugated in the Mikasa metrically rounded, passing gradually to gently and Kotanbetsu areas. arcuate posterior margin. Posterior hinge angle (1) from 1200 (in young stage) to 1400 (later). Inoceramus takahashii sp. nov. Axis of growth runs along the central line of PI. 84, Figs. 1a-c flank, forming angle (0) of about 65 0 with the hinge line. Holotype:-GK. H. 10145, both valves, col­ Shell in the main growth-stage ornamented lected by Takemi Takahashi from the upper with numerous, fairly dense, rather weak con­ reaches of the 8th branch of the Kami-ichi-no­ centric subcostae, which become very weak in sawa, a tributary to the River Ikushumbetsu, the last stage. Major ribs of the first order scarce­ Mikasa district, central Hokkaido. Outer and ly developed, except for the indistinct ones in inner shell layers preserved on the umbonal the early middle grwoth-stage, but the occur­ part; inner shell layer attached on some contigu­ rence of comparatively more distinct subcostae ous part; the rest main part of left valve and at periodic intervals associated with somewhat some portions of right valve internal mould. deeper interspaces may imply incipient or Specific characters:-Shell subequivalve; the hidden major ribs. Fine concentric lirae of the umbo of left valve slightly more projecting and third order discernible on the surface of the incurved than the right. Valve gently convex outer shell layer which is preserved on the in the main part, but for somewhat more inflated umbonal part of the holotype. umbonal part. Posterior part flattened, passing Etymology:-This species is dedicated to Mr. gradually from the main part of the disk, except Takemi Takahashi of Mikasa, who is one of the the umbonal part, where there is a faint auricular most eminent naturalists in collecting scientifi­ depression. Anterior side steep, with concave cally valuable fossils from Hokkaido. anterodorsal part. Remarks:-The holotype itself is fairly good, Shell subrounded rectangular in rough outline, but no more specimen has been obtained from nearly as high as broad in early stage and higher the same bed and the extent of variation is un- Measurements of holotype (left valve).

Specimen h l/h H L L/H b b/h sll r GK. H 10145 92.0 71.0 .77 102.0 72.0 .71 24.0 .26 41.0 .58 1050 850 1400 650

Explanation of Plate 82

Figs. 1-3. Birostrina tamurai sp. nov...... Page 411 1. GK. H 10164 (BV) from Mikasa; left lateral (a), right lateral (b), anterior (c), posterior (d) and dorsal (e) views, X 1. 2. GK. H 10153 (LV) from Kotanbetsu; lateral (a), anterior (b), posterior (c) and dorsal (d) views, x 1. 3. GK. H 10165 (BV) from Mikasa; anterior view in the lighting somewhat different from PI. 81, Fig. 5d, x 1. MATSUMOTO and NODA : Cenomanian Inoceramids from Japan-I Plate 82 816. Cenomanian Inoceramids from Japan-I 415 known at present. There are, however, some Queensland (Australia) under I. multiplicatus specimens from other areas within Hokkaido, Stoliczka var. elongatus Etheridge, I. pernoides which are comparable but not quite identical Etheridge and I. sp. aff. I. pro blema ticus with the holotype. They will be described in d'Orbigny seem to be allied to I. comanchean us. Part 3. The third one is especially similar to I. taka­ Comparison:-This species is somewhat similar hashii in the shell-form but different in having to Inoceramus comancheanus Cragin, 1895 from more distinct concentric subcostae on its young­ the Albian of Texas. By courtesy of Dr. W. A. er half and stronger concentric ribs on its later Cobban, there are plaster casts (GK. H 9194 and half. The age of the Marathon Formation was H 9195) of the two syntypes (USNM. 32686 A, considered as Cenomanian by Etheridge but B) (for their and other illustrations see Hill, must be Albian in the light of recent knowledge 1901 and Reeside, 1923) from the Duck Creek about the Cretaceous stratigraphy of Australia Limestone. Furthermore, Pergament (1964) (Ludbrook, 1978). described an example from the Albian of Kam­ To sum up, the described specimen found by chatka under Inoceramus cf. I. comancheanus T. Takahashi represents a new species belonging and then Eigenheer and Sornay (1974) another to the group of I. comanchean us. example from the basal Cenomanian of Montlaux Occurrence :-In a calcareous nodule derived (Alpes-de-Provence, France) under Inoceramus from the sandstone referable to the lower part aff. I. comancheanus. Our species differs from of the Mikasa Formation in the upper reaches them in having more subrectangular outline, of the 8th branch (Hachi-no-sawa in Japanese) with larger umbonal angle (~) and more broadly of the Kami-ichi-no-sawa, a tributary of the rounded ventral margin and weaker and denser River Ikushumbets. As Sharpeiceras sp. was concentric su bcostae. Especially, the concentric obtained by T. Takahashi form a nearby locality, ribs are distinctly stronger in the late growth­ the sandstone is most probably referred to the stage of I. comancheanus. In the two syntypes Lower Cenomanian. (left and right valves) of that species the main part of the flank is more convex and slopes Inoceramus heinzi Sornay, 1965 down less gradually to the posterior flat part, PI. 83, Figs. 4a-t\; PI. 85, Figs. 2-3b with a very faint, incipient, radial sulcus between the two parts. The specimen from the basal 1933. Inaequiceramus inaequiualuis (SchlUter); Cenomanian of Montlaux, mentioned above, Heinz p. 246, pI. 18, fig. 1. is more compressed than the American types 1965. Inoceramus heinzi Sornay, p. 7, pI. B, and is similar to ours in that respect, but is ex­ fig. 4. tended more obliquely and has more irregular Material:-GK. H 10148, internal (PI. 83, Fig. and stronger ribs than the holotype of our 4) and external moulds of left valve from loco T species. Anyhow, that French species is not 5046 B; GK. H 10149 (PI. 85, Fig. 2), left valve, either identical with ours. with shell attached on the main part, from loco I. takahashii is somewhat similar to I. pre­ T 5046 A; GK. H 10150, internal (PI. 85, Fig. 3) fragilis Stephenson, 1953 (see also Kauffman and external moulds of right valve from loco T et al., 1977), from the Middle Cenomanian of 5045 B; poorly preserved GK. H 10151, from Texas and Oklahoma, in the general outline loco T 5046 B (I. cf. 1. heinzi); all collected by and ornamentation, but the latter has a more T. Matsumoto. convex main part of the flank, more distinct Description :-Shell medium-sized, inequivalve auricular sulcus or break, and finer but more and inequilaterai. Beak of left valve projected distinct concentric subcostae. considerably beyond hinge and curved remark­ The specimens described long ago by Etheridge ably inward and forward, as seen in GK. H (1872) from the "Marathon Formation" of 416 Tatsuro MATSUMOTO and Masayuki NODA

10148. Valve moderately convex in the umbonal somewhat larger anterior hinge angle, longer part, inclined abrupty forward to the almost hinge line and larger ratio of L/H than our vertical and somewhat concave anterior side and described specimens and also than the holotype. gently towards the flattened posterodorsal wing The specimens IGPS. 22709 and UMUT. MM like part. Convexity along the growth·axis 6492 [=GT. 1-710] from pebbles of the River gradually decreased towards ventral margin. Obirashibe, described under Inoceramus sp. Outline suboval, extending obliquely towards indet. (sp. nov.?) by Nagao and Matsumoto the posteroventral extremity, with axis of growth (1939, p. 280, pI. 27, fig. 1 and pI. 28, fig. 1), forming angle (8) of about 500 with hinge line; are somewhat similar to this Sornay's form, as beak anterior, hinge line slightly shorter than a Kauffman (1977, p. 176) has pointed out, but half of shell length, forming angle (a) of 75- they are larger, "broader" and flatter. Probably 800 with anterior margin and that (')') of 1450 they represent a distinct species, but we have to 1500 with the posterior margin; anterior not yet been successful to decide the strati­ margin broadly arcuate with gentle an tero­ graphic position of this peculiar form. dorsal concavity; ventral margin asymmetrically Occurrence :--Loc. T 5046 A, B, Saku-gakko­ rounded, passing gradually to gently convex, no-sawa, lower part of Member II c (clayey mud­ arcuate posterior margin. stone); also loco T 5045 B, close to T 5046, Shell ornamented with rather low concentric uppermost part of Member II b (fine-grained major ribs or undulations and also concentric sandy siltstone), Saku area, Teshio Mountains rings, the latter of which may not be impressed (see route map of Matsumoto and Okada, 1973). on internal mould but well discernible on shell The main part of II b, lower than loco T 5045, is surface or external mould. Posterior muscle referred to the Middle Cenomanian on the evi­ impression is observable slightly behind the dence of ammonites, Desmoceras (Pseudo­ midline in GK. H 10150. uhligella) japonicum Yabe, Calycoceras spino­ Comparison and discussion :-The described sum (Kossmat), C. cf. C. bathyomphalum (Kos­ specimens are very similar to the holotype smat) and Euomphaloceras asura Matsumoto et illustrated by Heinz (1933, pI. 18, fig. 1) and Muramoto [= ?E. multicostatum (Basse)], and also by Sornay (1965, pI. B, fig. 4) of 1. heinzi the main part of II c, higher than loco T 5046, Sornay from Madagascar. is referred to the Lower Turonian on the evi­ The specimens (no. 534 A, B) from Anko­ dence of Mytiloides mytiloides (Mantell), Fagesia maka, drawn by Sornay (1965, fig. 3), have cf. F. rudra (Stoliczka) and F. thevestensis

Measurements of described specimens.

Specimens h l/h H L LiH b b/h sll a !3 r 0 GK. H 10148 70 65 .93 78 52 .67 19 .27 32.0 .49 85° 65° 145° 50° GK. H 10150 47 42 .89 55 42 .76 12 .26 19.5 .46 90° 150° 50°

Explanation of Plate 83

Figs. 1-3. Birostrina tamurai sp. nov...... Page 411 1. GK. H 10155 (RV) from Kotanbetsu; lateral view, x 1. 2. GK. H 10158 (LV) from Kotanbetsu; lateral (a) and posterior (b) views, x 1. 3. GK. H 10082 (LV) from Kotanbetsu; lateral (a), anterior (b), posterior (c) and dorsal (d) views, x 1. Fig. 4. Inoceramus heinzi Sornay ...... Page 415 GK. H 10148 (LV) from Saku; lateral (a), dorsal (b), posterior (c) and anterior (d) views, x 1. MATSUMOTO and NODA: Cenomanian Inoceramids from Japan-I Plate 83 816. Cenomanian Inoceramids from Japan-I 417

Pervinquiere. Therefore, the beds exposed at ribs, which are of moderate intensity in the locs. T 5045 and T 5046 are probably referable middle growth-stage and gradually weakened to the Upper Cenomanian and/or lowest part later; surface with fine and regular concentric of Lower Turonian. Iirae, which may be somewhat prominent on the summit of concentric ribs. Genus Mytiloides Brongniart, 1822 Test very thin. Faint radial striae, which may be tracks of mantle retractor muscles, discernible Type species:-Ostracites labiatus Schlotheim, on the main part of internal mould, but for a 1813. narrow zone along the growth axis, which may Remarks:-For generic diagnosis and distinc­ be posterior adductor muscle insertion area. tion from Inoceramus Sowerby, 1814 readers Remarks:-Fragmentary shells with regular may refer to Kauffman et al., 1977 and Keller, concentric ribs are contained in the calcareous 1982, whom we follow. nodule together with the holotype, but they are too incomplete to show the entire outline. Mytiloides mikasaensis sp. nov. Comparison :-This species is distinguished PI. 85, Figs. 1a-c from Mytiloides submytiloides (Seitz, 1935) (Holotype; Seitz, 1935, p. 444, pI. 37, fig. 1, Holotype:-GK. H 10045, composite internal text-fig. 8a), from the Upper Cenomanian and mould of right valve, with test attached partly, lowest Turonian of Europe and North America collected by Tatsuo Muramoto from loco Ik (Wiedmann and Kauffman, 1985), in its larger 1038 on the right bank of the River Ikushum­ anterior hinge angle (a) and larger umbonal bets, Mikasa district, central Hokkaido. angle (~) more convex and forward projecting Specific characters:-Valve gently convex anterior margin, and more regular concentric from beak to ventral margin and also from front ribbing combined with finer and more regular to rear, inclined fairly steeply on anterior side concentric lirae. and gently to posteior, passing to rather narrow M. mikasaensis is similar in shell-form to M. flat wing. Hinge plate thin. Hinge line rather goppelnensis (BadiIIet et Sornay, 1980) (Keller, short, a little more than a third of shell length. 1982), from the Lower Turonian of Europe and Beak nearly but not exactly at the anterior end North America, especially to the form described of hinge line, suberect, projecting slightly. under M. goppelnensis tourtenayensis (Badillet Shell extended obliquely towards the postero­ et Somay, 1980) from France, but the former ventral extremity, with H fairly longer than L has less arcuate growth-axis and more regular and with axis of growth conyexly curved for­ and less crowded concentric ribs combined more ward, showing angle (0) of 45° to 35° with hinge regularly with finer lirae. line. Anterior hinge angle (a) and angle of umbo­ For so~e reasons, Seitz (1935), Kauffman nal inflation (~) large. Anterior margin convex, (in Kauffman et al., 1977 and other papers) and with forward arcuate anteroventral margin, pass­ Troger (1981) seem to have misunderstood ing gradually to broadly and asymmetrically Inoceramus opalensis Bose, 1923 in the original curved ventral margin, and then curved rather sense, which is represented by Bose (1923, pI. abruptly at the posteroventral extremity to 13, figs. 1-3) and never so oblique as the so­ gently arcuate, long, posterior margin. Posterior called M. opalensis by them. This has been hinge angle (r) very large in late growth-stage. revised by BadiIIet and Somay (1980), to whose Shell ornamented with regular concentric opinion we are quite agreeable. W. A. Cobban Measurements of holotype.

Specimen h I/h H L L/H b b/h 5/1 a ~ 'Y 5 GK. H 10045 41.0 51.0 1.24 51.8 37.6 .73 11.0 .27 19.0 .37 1400 1300 1550 450 350 418 Tatsuro MATSUMOTO and Masayuki NODA

(1983, personal communication) told us that Acknowledgements examples of correctly identified M. opalensis occur near the boundary of Lower and Middle We wish to thank Professors Minoru Tamura Turonian, i.e. near the boundary of Mammites and Hakuyu Okada and Messrs. Takemi Taka­ nodosoides Zone and Collignoniceras woollgari hashi, Tatsuo Muramoto and Kikuwo Muramoto Zone. for their kind help to this study, including the Mytiloides modeliaensis (Sornay, 1981), from supply of valuable specimens. Thanks are ex­ the Lower Turonian of Colombia and North tended to Professors Itaru Hayami of the Uni­ America, is similar to M. mikasaensis with respect versity of Tokyo and Tamio Kotaka of Tohoku to the shell-form, especially to the convex an­ University, who gave us facilities to study there terior margin, but the former has on the average some type specimens in their charge, and also to less oblique growth-axis and more crowded Drs. W. A. Cobban, E. G. Kauffman and Jacques and sharper concentric ribs. Sornay who gave us good suggestions. Occurrence:-Loc. Ik 1038, right bank of the main stream of the River Ikushumbets, Zone of References cited Euomphaloceras septemseriatum, Upper Ceno­ manian of the Mikasa district, central Hokkaido Badillet, G. and Sornay, J. (1980): Sur quelques (see Matsumoto, 1965, figs. 2 and 4 for the loca­ formes du groupe d'Inoceramus labiatus tion ofIk 1038). decrites par O. Seitz. ImpossibiIite d'utiliser ce groupe pour une datation stratigraphique du inferieur du Saumurois (France). C.R. Summary Acad. Sci. Paris, vol. 290 D, p. 323-325. As a summary the species described in this Bose, E. (1923): Algunas faunas cretacicas de Zacutecas, Durango y Guerrero. Inst. Gesl. paper are listed below with indication of their Mexico, Bol., vol. 42, p. 1-219, pIs. 1-19. known age and area. Cox, L. R. (1969): Inoceramidae. In Moore, R. Birostrina tamurai sp. nov., Middle Ceno­ C. (ed.), Treatise on invertebrate paleon­ manian of the Mifune (central Kyushu), tology, Part N, Mollusca 6, p. N 314-N 321, Mikasa (central Hokkaido) and Kotanbetsu Geol. Soc. Amer. & Univ. Kansas Press. (northwestern Hokkaido) areas Cragin, F. W. (1892): A contribution to the Inoceramus takahashii sp. nov., Lower Ceno­ invertebrate paleontology of the Texas manian of the Mikasa area Cretaceous. Geol. Surv. Texas, 4th Ann. Inoceramus heinzi Sornay, Upper Cenomanian Rep., no. 2 (inaccessible). and (?) Lower Turonian of the Saku area Dhondt, A. V. (1983): Campanian and Maast­ (northwestern Hokkaido) richtian inoceramids: a review. Zittliana, Mytiloides mikasaensis sp. nov., Upper Ceno­ vol. 10, p. 689-701. Dobrov, S. A. and Pavlova, M. M. (1959): Ino­ manian of the Mikasa area. ceramidae. In Moskvin, M. M. (ed.), Atlas We hope that more examples of these species of the Upper Cretaceous fauna of Northern would be found from other localities in the Caucasus and Crimea. Trudy vses. nauchno­ future, so that their biostratigraphic and biogeo­ issled. Inst. prir. Gazov, p. 130-165, pIs. 1- graphic implications may become clearer. 23 (in Russian).

Explanation of Plate 84

Fig. 1. Inoceramus takahashii sp. nov...... Page 414 Holotype, GK. H 10145 (BV) from Mikasa; right (a) and left (b) lateral and anterior (c) views, x 1. MATSUMOTO and NODA: Cenomanian Inoceramids tram Japan-I Plate 84 816. Cenomanian Inoceramids from Japan-I 419

Efremova, v. I. (1978): On methods and unifica­ 16, no. 1, p. 1-80, pIs. 1-18. tion of measurements of morphological --and Noda, M. (1985): A new inoceramid elements of inoceram tests. In Pergament, (Bivalvia) species from the Upper Campanian M. A. (ed.), Jurassic and Cretaceoous Ino­ (Cretaceous) of Hokkaido. Proc. Japan cerams and their Stratigraphic Importance, Acad., vol. 61, ser. B, p. 1-3. p. 99-104, tables, Acad. Sci. USSR (in --and Okada, H. (1973): Saku Formation of Russian). the Yezo geosyncline. Sci. Repts. Dept. Eigenheer, R. and Sornay, J. (1974): Sur une Geol., Kyushu Univ., vol. 11, no. 2, p. 275- forme apparenteee a Inoceramus comanche­ 309 (in Japanese with English abstract). anus Cragin dans Ie Cenomanien basal de Nagao, T. and Matsumoto, T. (1939): A mono­ Montlaux (Alpes-de-Provence). Bull. Mus. graph of the Cretaceous Inoceramus of Natn. d'Rist. Nat., 3 ser., no. 229, p. 141- Japan. Part I. Jour. Fac. Sci., Rokkaido 144. Imp. Univ., Ser. 4, vol. 4, p. 241-299, Etheridge, R. (1872). Description of the Paleo­ pIs. 23-34. zoic and Mesozoic fossils of Queensland. Noda, M. (1983): Notes on the so-called Ino­ Quart. Jour. Geol. Soc. London, vol. 28, p. ceramus japonicus (Bivalvia) from the 317-350, pIs. 14-25. Upper Cretaceous of Japan. Trans. Proc. Heinz, R. (1933): Inoceramen von Madagascar Palaeont. Soc. Japan, N.S., no. 132, p. 191- und ihre Bedeutung fiir die Kreide-Strati­ 219, pIs. 41-46. graphie. Zeits. Deut. Geol. Ges., vol. 85, p. --(1984): Notes on My tiloides incertus (Creta­ 241-259, pIs. 16-22. ceous Bivalvia) from the Upper Turonian of Hill, R. T. (1901): Geography and geology of the the Pombets area, central Hokkaido. Ibid., Black and Grand Prairies, Texas, with de­ no. 136, p. 455-473, pIs. 84-86. tailed description of the Cretaceous forma­ --and Matsumoto, T. (1976): Mesozoic mol­ tions and special reference to artesian luscan fossils of Japan 4 (Certaceous Ino­ waters. Rep. U.S. Geol. Surv., vol. 21, no. 7, ceramus). In Matsumoto, T. (ed.), Atlas 666 p., 71 pIs. (inaccessible). of Japanese Fossils, no. 45, Cr 31-Gr 36, Kauffman, E. G. (1977): Systematic, biostrati­ Tsukiji Shokan, Tokyo. graphic, and biogeographic relationships be­ Pergament, M. A. (1966): Zonal stratigraphy tween Middle Cretaceous Euramerican and and inocerams of the lowermost Upper north Pacific Inoceramidae. Palaeont. Soc. Cretaceous of the Pacific coast of the Japan Spec. Pap., no. 21, p. 169-212. USSR. Trans. Acad. Sci. USSR, Geol. Inst., --, Hattin, D. E. and Powell, J. D. (1977): vol. 146, 83 p., 36 pIs. (in Russian). Stratigraphic, paleontologic and paeoenviron­ Reeside, J. B. (1923): The fauna of the so-called mental analysis of the Upper Cretaceous Dakota Formation of northern central rocks of Cimarron County, northwestern Colorado and its equivalent in southeastern Oklahoma. Mem. Geol. Soc. Amer., vol. 149, Wyoming. U.S. Geol. Surv. Prof. Pap., no. 150 p., 12 pIs. 131 H, p. 199-207, pIs. 45-50. Keller, S. (1982): Die Oberkreide der Sack-Mulde Seitz, O. (1935): Die Variabilitiit des Inoceramus bei Alfeld (Cenoman-Unter-Coniac). Litho­ labiatus v. Schloth. Jahrb. Preuss. Geol. logie, Biostratigraphie und Inoceramen. Landesan., vol. 55 (for 1934), p. 429-474, Geol. Jahrb., Reihe A, Heft 64, 154 p., 8 pIs. 36-40. pIs. --(1961): Die Inoceramen des Santon von Ludbrook, N. H. (1978): Australia. In Moullade, Nordwestdeutschland. I Teil. Beih. Geol. M. and Nairn, A. E. M. (eds.), The Phanero­ Jahrb., Heft 47,186 p., 15 pIs. zoic Geology of the World II The Mesozoic, --(1965): Die Inoceramen des Santon und A, Chapt. 7, p. 209-249, Elsevier, Amster­ Unter-Campan von Nordwestdutschland. II. dam. Ibid., Heft 69, 194 p., 26 pIs. Matsumoto, T. (1965): A monograph of the --(1967): Die Inoceramen des Santon und Collignoniceratidae from Hokkaido. Part I. Unter-Campan von Nordwestdeutschland. Mem. Fac. Sci., Kyushu Univ., Ser. D, vol. III. Ibid., Heft 75, 171 p., 27 pIs. 420 Tatsuro MATSUMOTO and Masayuki NODA

--(1970): tiber einige Inoceramen aus der group by T. Matsumoto. Ibid .• vol. 23, no. Oberen Kreide. Ibid., Heft 86, 171 p., 28 1, p. 47-56, pI. 1. pis. Trager, K.-A. (1967): Zur Paliiontologie, Bio­ Sornay, J. (1965): La faune d'inocerames du stratigraphie und faziellen Ausbildung der Cenomanien et du Turonien inferieur du unteren Oberkreide (Cenoman bis Turon). sud-oest de Madagascar.. Ann. PaIeont., Teil I Paliiontologie und Biostratigraphie Invertebres, vol. 51, p. 1-18, pis. A-G. der Inoceramen des Cenoman bis Turons --(1966): Idees actuelles sur les inocerames Mitteleuropas. Abh. Staatl. Mus. Min. Geol.• d 'apres divers travaux recents. Ibid., Inverte­ vol. 12, p. 13-207 (incl. 14 pis.). bres, vol. 52, p. 59-92. Vokes, H. E. (1980): Genera of the Bivalvia: --(1981): Inocerames (Bivalvia) du Turonien A systematic and biostratigraphic catalogue inferieur de Colombie (Amerique du Sud). (Revised and updated), 307 p., Palaeonto. Ibid., Invertebres, vol. 67, p. 135-148, pis. Res. Inst., Itahaca, N.Y. 1-2. Wiedmann, J. and Kauffman, E. G. (1978): Mid­ Stephenson, L. W. (1953): Larger invertebrate Cretaceous biostratigraphy of northern fossils of the Woodbine Formation (Ceno­ Spain. Ann. Mus. d'Hist. Nat. Nice, vol. 4 manian) of Texas. U.S. Geol. Surv. Prof. (for 1976), III, p. 1-84 (incl. 12 pIs.). Pap .• 242 (for 1952), p. 1-226, pIs. 1-59. Woods, H. (1911): A monograph of the Cretace­ Tamura, M. (1979): Cenomanian bivalves from ous Lamellibranchia of England. Vol. 2, the Mifune Group, Japan. Part 3. Mem. Fac. pt. 7. Palaeontogr. Soc .• 1910, p. 261-284, Educ. Kumamoto Univ .• no. 28, Nat. Sci., pis. 45-50. p. 59-74, pIs. 1-3. --(1917): Certaceous faunas from the north­ --and Matsumura, M. (1974): On the age of east of South Island, New Zealand. N.Z. the Mifune Group, central Kyushu, Japan, Geol. Suv. Paleont. Bull., no. 4, p. 1-41, with a description of ammonite from the pis. 1-20.

Asanoyabu T:\1;O)ii. Hachi-no-sawa i\O)iR. Ikushumbets [IkushunbetsuJ ~lF5.m. Kami-ichi­ no-sawa J::-O)iR. Kiritachi ~:ll:. Kotanbetsu i1:f'HJI]. Kumamoto Jm*. Mifune f,l.jJ~H. Mikasa :=:~. Obirashibe 'P¥Ii. Okadake I8iJffi. Omine **. Pombets [PonbetsuJ 9"f-}]JJ. Saku f.£~. Saku-gakko-no-sawa fti:~'¥=t.lCO)iR. Shimomashiki Tt.d:l1iX;, Subayashi !Itt"" Teshio ;'klji, Toyono-mura ~!ilfN, Yaseto i\ifjip.

Explanation of Plate 85

Fig.1. Mytiloides mikasaensis sp. nov...... Page 412 Hoiotype, GK. H 10045 (RV), from Mikasa; lateral (a), anterior (b) and postero- dorsal views, x 1. Fragmentary left valve is attached on the opposite side. Figs. 2-3. Inoceramus heinzi Sornay ...... Page 415 2. GK. H 10149 (LV) from Saku; lateral view, x 1. 3. GK. H 10150 (RV) from Saku; lateral (a) and anterior (b) views, xL Figs. 4-5. Birostrina cf. B. tamurai sp. nov...... Page 411 4. KE 1977 (LV), incomplete, secondarily compressed, large example (internal mould), x 1. 5. Rubber-cast from the external mould (posteroventral portion) of KE 1977, x 1. MA TSUlvIOTO and NODA: Cenomanian Inoceramids tram Japan-I Plate 85 816. Cenomanian Inoceramids from Japan-I 421

*fBl~r~*-l! / ""7 =- 7 :/1\M'E[1' / -l! '7 A,A- I. ~tifij.ii! C fL1'/iE[0) J: <:m I?.h -c ~ ,t.eli' -:,f-: Hi: *:f!S0) -l! / ""7 =- 7 :/li' I?ll, t. tf. J: <~~~.h -c ~,t.eli' "? t-:1'jlitli' t.e IJ il!> GO) ,"(", k-.h I? ~ ;i¥/xIl~JliJn... -0' <• ;: 0) iffiljx'"("Il/X 0) 4 fi~IlG$lt L t-:. (1) Birostrina tamurai sp. nov. ; tJlitrl':' Inoceramus concentricus costatus Nagao et Matsumoto c Iif-'i.h -C ~ ,t-: ~ 0)0) -$'"(", B. concentrica ~I.:.~-t G lit, Wl:0)~$1.:.1tdlt~O)Mlitil!> IJ, hX:~f~MI.:.l!1Jlit;e~ -t Go fL1+1 O)~ti1\1lrff$~ Utifij)1HIH~~5]IJN.O'itft5]JjO) -l! / ""7 =- 7 :/ cp$E[o (2) Inocera- mus takahashii sp. nov.; I. comanchean us ~I.:.~-r G li~, Wl:O)~!1l\1.J~l(t~1.i}f;'"(", IJ~I? .7,..li~~~<' !lli1!1J~~'!I~'o ~Ui~5]IJo)-l!/""7=-7:/T$E[, (3) I. heinzi Sornay; ""7!J"-j/,A'j] J"1Ji(E[O)~O)I':' i'1J:

817. LEPIDOLINA COLUMBIANA ( FUSULINID) FROM BRITISH COLUMBIA, CANADA*

HIROYA GOTO

Institute of Geosciences, College of Liberal Arts, Kobe University; Kobe 657

KUNITAKA MARUOKA

Higashinada High School, Kobe 658

and

KEN·ICHI ISHII

Institute of Geosciences, College of Liberal Arts, Kobe University, Kobe 657

Abstract. Nine species of Yabeina have been reported from British Columbia. On the basis of statistical study of specimens collected from Marble Canyon and Upper Hat Creek, it is concluded that our specimens as well as specimens of the nine species of "Yabeina" (Y. columbiana, Y. minuta, Y. obesa, Y. dawsoni, Y. gracilis, Y. dunbari, Y. cylindrica. Y. ampla and Y. paruula) belong to one species which should be referred to the genus Lepidolina. They are described under the species L. columbiana (Dawson).

Introduction Recently Ishii and others (1985) reexamined It has been suggested that Middle Permian distribution of neoschwagerinids in the Tethys. fusulinids of the Cordilleran region including According to them, Neoschwagerina and Yabeina British Columbia, Oregon and California resem· are distributed allover the Tethys, while distri· ble Tethyan fusulinids unlike those of the Mid· bution of Colania and Lepidolina is restricted Continent, North America (Dunbar, 1932, to East Asia including Southeast Asia, China Thompson and Wheeler, 1942, Ross, 1967 and and Japan, which forms the Paleopacific bio· others). With this suggestion in mind, one of geographic province in Middle Permian time. us (Goto) visited the Marble Canyon region Our paleontological restudy of "Yabeina" during his stay in British Columbia in 1980 columbiana from British Columbia has revealed and collected limestone samples containing that this species does not belong to the genus many specimens of "Yabeina" columbiana. Ya be ina, but is assignable to the genus Lepi· dolina. In this paper we intend to present a result *Received April 15, 1985; Revised manuscript of our study on species of the genus Yabeina received June 19, 1986: Read January 22, 1984 described from Marble Canyon, British Columbia at Kyoto University. on the basis of specimens collected by Goto.

422 817. Lepidolina columbiana from B.C. Canada 423

Historical reviews:-According to Duffell and Geological setting McTaggart (1951), the study of fusulinids of the Marble Canyon limestone, British Columbia Geological mapping of the area including the was started in 1872 by Selwyn. In 1877 Dawson Marble Canyon region, British Columbia was revisited Selwyn's original locality in Marble made by Duffell and McTaggart (1951). Trettin Canyon and on the basis of the fusulinids (1980) described Permian Cache Creek Group assigned a age to the Marbl~ of the Marble Range. According to them the Canyon limestone. Dawson (1879) described Marble Canyon Formation is a constituent of Loftusia columbiana from Marble Canyon. Cache Creek Group. The Marble Canyon For­ Although Loftusia Brady is originally based on mation is composed of the massive and mostly a Tertiary species, Dawson concluded that L. recrystallized limestone with intercalations of columbiana was a Carboniferous form rather volcanic rocks, argillites and cherty quartzites, than Cretaceous or Eocene because of the typically exposed in the Marble Canyon and accompanying fusulinids. Pavilion Mountains (Text-fig. 1). In the western Dunbar (1932) restudied Dawson's thin sec­ limit of the distribution of the limestone in tions of specimens of Loftusia columbiana and Pavilion Mountain, strata are dipping mainly described the species under the genus Neo­ to the west and southwest at about 60 degrees, schwagerina. He pointed out that the limestone but near the eastern boundary, dips are to the at Marble Canyon is Permian in age. Based on a west at about only 20 degrees. Though the study of well oriented section of Loftusia colum­ repetition of strata resulting from minor fold­ biana Dawson, Thompson and Wheeler (1942) ing and faulting is conspicuous, it is probable pointed out that this species should be referred that the limestone has a thickness of about to the genus Yabeina Deprat. They also described 1800 m. The Marble Canyon Formation is Yabeina minuta, Schwagerina pavilioensis var. flanked on both sides by chert, argillite, green­ acris, Nankinella? and Staffella in addition to stone, minor limestone, quartzite and their Yabeina columbiana from samples of talus in metamorphic equivalents, which are constitu­ ents of the Cache Creek Group. The Marble Marble Canyon. Thompson, Wheeler and Danner Canyon Formation yields Permian foraminifers, (1950) redescribed Y. columbiana and Y. minuta crinoids, blastoids, corals and brachiopods, associated with Yabeina ? n. sp. Schwagerina and this formation has been assigned to the acris, S. andersoni ? and Codonofusiella duffeli Guadalupian Formations of the other regions from the Marble Canyon region. Skinner and by many investigators (Duffel and McTaggart, Wilde (1966) described seven new species of the 1951 and Trettin, 1980). genus Yabeina listed below in addition to Y. Our samples were collected from the follow­ columbiana, Y. minuta, Schwagerina canadensis ing two localities (Text-fig. 1). and Chusenella spicata from the same !ocality Loc. 1, Marble Canyon: Talus on the north of Marble Canyon material. The seven species shore of the Pavilion Lake. Massive recrystallized of the genus Yabeina are Yabeina obesa Y limestone. Type locality of "Yabeina" colum­ dawsoni, Y. gracilis, Y. dunbari, Y. cylind'rica: biana (Dawson). Sample obtained from Loc. 1 Y. ampla and Y. parvula. are grayish limestone, containing fossils of Acknowledgement:-We wish to express our fusulinids, smaller foraminifers and algae. The thanks to Dr. Raymond C. Douglass of U.S. preservation of these fossils is generally poor, Geological Survey, Prof. Wilbert R. Danner of because of recrystallization of limestone owing University of British Columbia, Canada and to regional metamorphism. Prof. Koichiro Ichikawa of Osaka City University Loc. 2, Upper Hat Creek: About 30 km south for their kind advice. of Loc. 1. Bedded limestone, partly recrystal­ lized. Samples obtained from Loc. 2 are dark 424 Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

[[[]]

[[I]]] 2

500 45'N

~1-__51...! -_19 km q i !

Text-fig.!. Geological sketch map of the Marble Canyon region. Loc. 1: Marble Canyon, Loc. 2: Upper Hat Creek, 1: Pavilion beds, 2: western belt of Cache Creek Group, 3: central belt of Cache Creek Group (Marble Canyon Formation), 4: eastern belt of Cache Creek Group. (after Trettin, 1980)

gray limestone showing micritic texture. Only First, the comparison of individuals treated fusulinids are found as fossils. This locality is this time with those of species of Yabeina and newly found by Goto. Lepidolina described previously is made with reference to the form ratio and the diameter Morphologic variation of proloculus (Text-fig. 2). The individuals treated this time and those of nine species of Before going over the systematic paleon­ Yabeina described by Skinner and Wilde from tology, some results of morphological analysis the same region are recognized to be situated of yabeinid specimens from two localities are on the intermediate position between Lepi­ presented in order to examine the scope of dolina multiseptata from East Asia (cf. Ozawa, specific variation. 1975), and the Yabeina group including Y. Morphometry of the specimens from Marble globosa, Y. packardi, Y. kaizensis, Y. igoi, Y. Canyon and Upper Hat Creek is carried out katoi, etc. In Text-fig. 2, measured values of with respect to the following shell characters proloculus of the present specimens and those such as form ratio, diameter of proloculus, of nine species of Yabeina mentioned already thickness of spirotheca and first appearance are overlapping with that of Lepidolina asiatica of secondary transverse septula. but are not overlapping with that of most of 817. Lepidolina columbiana from B.C. Canada 425

19. Lepldalino mulliseptata multiseptata Text-fig. 2. Diam eter of proloculus / against form ratio for present specimens 400 6 from Upper Hat Creek (open circle) and 20. L. m. shiraiwensis Marble Canyon (solid circle). Variation ranges of species of L epidolina and Yabeina are shown as vertical bars and squares. 1: E 300 "Yabeina" columbiana, 2: "Y." minuta, 3 : o o "Y." obesa, 4: "Y." dawsoni, 5: "Y." gracilis, 6: "Y." dunbari, 7: " Y . " cylindrica, 8: "Y." ampla, 9: "Y. " parvula, 10: Y. cascadensis, 11: Y. decora, 12: Y. fu si­ formis, 13 : Y. paclwrdi, 14: Y. kaizensis, being indicated by symbol of black triangle, 15: Y. igoi, 16 : Y. inouye i, 17: Y. haloi, 18: Y. globosa, 19: L epidolina multiseptala muitiseptata, 20: L. m. shiraiwensis, 21: L . asiatica, 22: L . japonica. Variation range

o 10 2.0 30 4.0 of the species of Yabeina (s. str.) falls within the part below the broken line. form rotio

Text-fig. 3. Comparison of spirotheca, primary and secondary transverse septula among L epidolina multiseptala multiseplala, L. columbiana and Yabeinaglobosa. a : L. 111 . multiseptata, topotype, loco Phnom Bak, Sisophon, Cambodia (coll. Ishii), b: L. columbiana, KU 3002, enlarged part of pI. fig. 2, loco Marble Canyon, British Columbia (colI. Goto), c: Y. globosa, topotype, loco Mameishi limestone, Akasaka, Gifu Prefec­ ture, Japan (colI. Ishii). 426 Hiroya GOTO, Kunitaka MAR UOKA and Ken-Ichi ISHII the Yabeina group. This text-figure shows that we described that present specimens and those some specimens, which were referred to "Yabe­ of nine species by Skinner and Wilde belong to ina" columbiana and "Y." ampla by Skinner the genus Lepidolina. and Wilde, are below 100 microns in diameter N ext, the frequency distribution curves for of proloculus. However, there is a possibility magnitude of form ratio of the specimens from that thin sections of these specimens were not Marble Canyon and Upper Hat Creek show a cut through the center of proloculus. Since bell shape (Text-fig. 4). The bell shape in the diameter of proloculus of representative species frequency distribution curves of specimens of Yabeina such as Yabeina globosa, Y. cascad­ from Marble Canyon and Upper Hat Creek may ensis, Y. decora, Y. fusiformis, Y. packardi, Y. be attributed to that the specimens come from kaizensis, Y. igoi, Y. inouyei and Y. katoi is a population. Most of the specimens have a form mostly below 100 microns, we regard the genus ratio of 1.5 to 2.0. The specimens of "Yabeina" Yabeina as fusulinids having the prolocular columbiana described by Skinner and Wilde diameter of less than 100 microns. (1966) are within this range of form ratio (Text­ Secondarily, the present specimens and fig. 4). those of nine species of "Yabeina" from Marble Incidentally several morphologic characters Canyon have the generic character of Lepidolina including form ratio, diameter of proloculus, that secondary transverse septula are thin bar thickness of spirotheca, first appearance of and occasionally pendant club-shape in cross secondary transverse septula are said to change section (Text·fig. 3). In view of these differences, phylogenetically in the subfamily Neoschwageri­ ninae (Ozawa, 1970). For example, the more advanced species of the genus Lepidolina which Marble Canyon 8 Upper Hat Creek 20 is included in the Neoschwagerininae have n = 43 larger form ratio, larger proloculus, thinner m = 1.81 spirotheca and the earlier appearance in onto­ en 15 0 geny of secondary transverse septula. In addition, :::J -0 the magnitude of form ratio also changes accord­ > n = 29 ing to change of environment such as the change -0 m= 1.78 C in speed of water current and in wave action. 10 't-' It seems that species of Eopolydiexodina and 0 Monodiexodina having larger form ratio are "- Q) n = 14 found in limestone of higher energy environ­ .0 E 5 m= 1.86 ment such as sparite, while the species of Yabe­ :::J C ina and Verbeekina having smaller form ratio are observed in limestone of low energy environ­ ment such as micrite. The larger value of form o ratio in the Marble Canyon specimens is larger than that of Upper Hat Creek. In view of re­ for mral' i sembling lithology of limestone in the two l'r";;;:;::~;:::::; localities, the above mentioned feature may l "Y."columbiana, "Y. "ampla reflect the phyletic increase in form ratio. I/y' ilobesa, "yN dunbari, I'y/lparvu/a The specimens from both localities are com­ Text-fig. 4. Frequency distribution pared with one another by plotting thickness curves of form ratio for the present speci­ of spirotheca and volution number on which mens from Marble Canyon and Upper Hat secondary transverse septula first appears (Text­ Creek. n: measured individual number, m: figs. 5, 6). On these two text-figures, present mean value. specimens and those of nine species of "Yabe- 817. Lepidolina columbiana from B.c. Canada 427

thickness of spirotheca. in microns first appearance of secondary transverse septu Ia Ip 30 40 50 volution number I I I -1 5, 10 -2 01 -3 02 -4 5- 04 , -6 ~5 ~7 06 -8 07 08 I -:9 I , 9 0 ---10 ~10 ~11 I 11 0 --:12 012 ----13: ?13 014 ""',--15 I 015 , !------16 I I , I , --~--17 : 170 I 18 I 718 19 --::i 190--0 : I I 20 21 20d--o ,I ,r- 21 ; I ?22

en Marble Canyon Ci :::J ~ II !II II :.a> ~I .~ '0 10 Upper Hat Creek 0 Upper Hat Creek '­ ... Q)

Marble Canyon Table 1. Relative growth coefficients (a) form ratio for the present specimens, Lepi­ 5.0. A 0..1-1.0. dolina multiseptata multiseptata, B 1.0.-1.5 L. m. shiraiwensis, L. asiatica 1.5-2.0. 3.0. C D 2.0.- 2.5 A (after Ozawa, 1975) and Yabeina E 2.0. E 2.5- 3.0 D globosa (after Hanzawa and Mura­ E ta,1963). c E (; 1.0. Form ratio a u -Q) > 1.0-1.5 1.02 II'> 0..5 ~ 1.5-2.0 0.82 '0 Marble Canyon 2.0-2.5 0.76 ~ 0..3 2.5-3.0 0.79 0..2 3.0-3.5 0.69 0.5-1.0 0.99 1.0-1.5 0.76 0.1 0.2 0.3 0..5 1.0. 2.0. 3.0 5.0 Upper Hat Creek 1.5-2.0 0.85 half length, in mm 2.0-2.5 0.85 Text-fig. 7. Growth curves of the 2.5-3.0 0.79 present specimens from Marble Canyon. Lepidolina m. multiseptata 0.63 Line A is a mean curve of specimens whose L. m. shiraiwensis 0.77 form ratio are within 1.0 to 1.5, and so L. asiatica 0.82 forth. Yabeina globosa 0.98 Upper Hat Creek farm ratia 5.0. A 0.1- 1.0. Table 1 demonstrates the relative growth co­ B 1.0.- 1.5 efficients - that is inclination of a - of the 3.0. C 1.5- 2.0. respectively classified groups, which is led by E D 2.0.- 2.5 2.5- 3.0. A E 2.0. E applying method of least squares. As seen from c C Table 1, all. the individuals from Marble Canyon ... D and Upper Hat Creek are within the range of 0 1.0. u E -Q) about 0.7 to 1.0. It suggests that these two > II'> groups would belong to the same species. More­ ~ 0..5 over, they differ from those of m. '0 Lepidolina ...0 multiseptata, while they are difficult to be dis­ 0.3 tinguished from L. asiatica, L. m. shiraiwensis 0..2 and Yabeina globosa by the relative growth coefficient only. However, they can be distin­ guished from specimens of the latter two species 0.1 0.2 0.3 0..5 1.0. 2.0. 3.0. 5.0. on basis of prolocular diameter. half length, in mm In conclusion, all specimens from Marble Text-fig. 8. Growth curves of the Canyon and Upper Hat Creek treated this time present specimens from Upper Hat Creek. as well as those of nine species of "Yabeina" For line A to E see Tex t-fig. 7. described by Skinner and Wilde are regarded to belong to the same species. They belong undoubtedly to the genus Lepidolina and not values which were calculated for all individuals to Yabeina. They are described under the species classified by range of form ratio in each locality. Lepidolina columbiana (Dawson). 817. Lepidolina columbiana from B.C. Canada 429

Table 2. Measurement of Lepidolina columbiana (Dawson)

KU 3001, from Marble Canyon, PI. 86, Fig. 1. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.18 0.30 0.40 0.50 0.65 0.85 1.00 1.20 1.45 1.68 1.91 2.18 2.53 rv 0.25 0.35 0.41 0.50 0.64 0.79 1.00 1.15 1.45 1.65 1.88 2.13 fr 0.72 0.85 0.97 1.00 1.00 1.08 1.00 1.04 1.00 1.02 1.02 1.02

Length: 5.08 mm, width: 4.45 mm Thickness of spirotheca: 30 microns Diameter of proloculus: 250 microns First appearance of sts: 5th volution

vn: volution number, hI: half length (mm), rv: radius vector (mm), fr: form ratio, sts: secondary transverse septula. Note: Thickness of spirotheca is measured at 10th volu tion on all specimens.

KU 3002, from Marble Canyon, PI. 86, Fig. 2. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.28 0.34 0.45 0.66 0.91 1.20 1.65 2.00 2.40 2.79 rv 0.20 0.30 0.40 0.50 0.65 0.85 1.05 1.20 1.43 1.65 fr 1.38 1.13 1.11 1.31 1.39 1.41 1.57 1.67 1.68 1.69

Length: 5.70 mm, width: 3.30 mm Thickness of spirotheca: 12 microns Diameter of proloculus: 302 microns First appearance of sts: 4th volu tion

KU 3003, from Marble Canyon, PI. 86, Fig. 3. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.29 0.48 0.65 0.83 1.20 1.50 1.85 2.40 2.83 3.15 3.50 3.80 rv 0.24 0.33 0.40 0.45 0.53 0.60 0.69 0.80 0.98 1.23 1.43 fr 1.19 1.46 1.63 1.83 2.29 2.50 2.68 3.00 2.90 2.57 2.45

Length:' 7.40 mm, width: 3.10 mm Thickness of spirotheca: 28 microns Diameter of proloculus: 320 microns First appearance of sts: 6th volution 430 Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

KU 3004, from Marble Canyon, PI. 86, Fig. 4. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.26 0.42 0.65 0.95 1.20 1.58 1.95 2.53 2.95 3.60 4.37 4.93 rv 0.15 0.29 0.33 0.43 0.50 0.65 0.81 0.96 1.13 1.30 1.50 1.66

fr 1.79 1.45 ~.OO 2.24 2.40 2.42 2.41 2.63 2.62 2.77 2.91 2.97

Length: 10.5mm, width: 3.60 mm Thickness of spirotheca: 25 microns Diameter of proloculus: 250 microns First appearance of sts: 6th volution

KU 3005, from Marble Canyon, PI. 86, Fig. 5. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.12 0.29 0.49 0.83 1.10 1.35 1.63 1.80 2.36 2.65 3.30 3.60 3.93 rv 0.10 0.19 0.23 0.28 0.35 0.44 0.49 0.58 0.65 0.77 0.91 1.02 1.20 fr 1.20 1.53 2.16 3.00 3.14 3.10 3.36 3.13 3.62 3.44 3.65 3.53 3.28

Length: 7.30 mm, width: 2.57 mm Thickness of spiro theca : 15 microns Diameter of proloculus: 123 microns First appearance of sts: 6th volution

KU 3006, from Upper Hat Creek, PI. 86, Fig. 6. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.10 0.19 0.28 0.35 0.52 0.62 0.80 0.97 1.09 1.20 1.30 1.43 1.53 rv 0.13 0.20 0.35 0.45 0.59 0.82 0.95 1.15 1.25 1.45 1.58 1.75 1.90 fr 0.77 0.93 0.79 0.78 0.88 0.75 0.84 0.84 0.87 0.83 0.83 0.81 0.80

Length: 3.10 mm, width: 4.00 mm Thickness of spirotheca: 28 microns Diameter of proloculus: 121 microns First appearance of sts: 6th volution

KU 3007, from Upper Hat Creek, PI. 86, Fig. 7. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hI 0.07 0.19 0.25 0.34 0.43 0.58 0.74 0.94 1.20 1.45 1.70 1.98 2.25 2.53 2.80 rv 0.09 0.16 0.25 0.29 0.45 0.57 0.70 0.90 1.03 1.35 1.66 1.90 2.15 2.41 2.80 fr 0.78 1.23 1.00 1.16 0.96 1.02 1.05 1.04 1.17 1.07 1.02 1.04 1.05 1.05 1.04

Length: 5.60? mm, width: 5.35 mm Thickness of spirotheca: 26 microns Diameter of proloculus: 144 microns First appearance of sts: 8th volution 817. Lepidolina columbiana from B.C. Canada 431

KU 3008, from Upper Hat Creek, PI. 86, Fig. 8. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hi 0.25 0.35 0.54 0.85 1.15 1.35 1.60 1.85 2.15 2.45 2.75 3.28 rv 0.16 0.27 0.38 0.48 0.60 0.70 0.85 101 1.16 1.32 1.47 1.66 fr 1.56 1.30 1.43 1.79 1.92 1.93 1.88 1.84 1.85 1.86 1.86 1.98

Length: 6.67 mm, width: 3.10 mm Thickness of spirotheca: 25 microns Diameter of proloculus: 260 microns First appearance of sts: 8th volution

KU 3009, from Upper Hat Creek, PI. 86, Fig. 9. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hi 0.17 0.30 0.43 0.60 0.85 1.10 1.39 1.98 2.32 2.55 3.00 rv 0.15 0.19 0.23 0.30 0.40 0.51 0.66 0.80 0.95 1.10 1.23 1.45 fr 1.13 1.58 1.87 2.00 2.13 2.17 2.10 2.47 2.44 2.32 2.44

Length: 6.20 mm, width: 2.60 mm Thickness of spirotheca: 20 microns Diameter of proloculus: 200 microns First appearance of sts: 8th volution

KU 3010, from Upper Hat Creek, PI. 86, Fig. 10. vn 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 hi 0.28 0.40 0.61 0.95 1.17 1.61 1.94 2.50 3.00 3.55 3.95 4.40 5.00 rv 0.15 0.23 0.31 0.43 0.55 0.60 0.83 0.98 1.10 1.25 1.45 1.60 1.76 fr 1.83 1.78 1.97 2.21 2.12 2.68 2.35 2.65 2.73 2.84 2.72 2.75 2.85

Length: 10.00? mm, width: 3.65 mm Thickness of spirotheca: 21 microns Diameter of proloculus: 214 microns First appearance of sts: 7th volution

Systematic paleontology Loftusia colum biana Dawson, 1879, p. 69-75, pI. 6, figs. 1-7; Silvestri, 1932, p. 87, pI. 2, Family Verbeekinidae Staff and Wedekind, 1910 figs. 4-7. Subfamily Neoschwagerininae Neoschwagerina columbiana, Dunbar, 1932, p. Dunbar and Condra, 1928 46-48, pI. 1, figs. 1-4. Yabeina minuta Thompson and Wheeler, 1942, Genus Lepidolina Lee, 1933 p. 707-708, pI. 106, figs. 6-10; Thompson, Lepidolina columbiana (Dawson) Wheeler and Danner, 1950, p. 62, pI. 8, figs. 4-7; Ishii and Nogami, 1964, p. 22, pI. 4, PI. 86, Figs. 1-10. figs. 4-7; Skinner and Wilde, 1966, p. 309- 432 Hiroya GOTO, Kunitaka MARUOKA and Ken-Ichi ISHII

311, pI. 34, figs. 3-7; Yang, 1978, p. 120, Measurement:-See Table 2. pI. 34, figs. 1-2. Remarks:-Morphologic variation within the Yabeina columbiana, Thompson and Wheeler, species is given in detail in the preceding chapter. 1942, p. 708-710, pI. 106, fig. 5, pI. 107, The present species differs from Lepidolina fig. 5, pI. 108, fig. 1, pI. 109, figs. 1-4; multiseptata multiseptata and L. m. shiraiwensis Thompson, Wheeler and Danner, 1950, p. in having rather smaller proloculus. Moreover 61, pI. 8, figs. 1-3; Kanmera, 1954, p. 16- the secondary transverse septula appear in 18, pI. 3, figs. 1-5, 7, questionable specimens, pI. 3, fig. 6; Nogami, 1958, p. 101-102, pI. 1, slightly later stage and the thickness of spiro­ figs. 9-10; Chisaka, 1960, p. 249-250, pI. theca is larger in the former than in the latters. 7, fig. 3, p. 54, fig. 18c; Skinner and Wilde, The secondary transverse septula of the 1966, p. 47-48, pI. 38, figs. 1-6; Yamagiwa present species appear in earlier stage than those and Saka, 1972, p. 268-269, p. 31, fig. 4; of L. asiatica. Although measured values of size Zang, 1982, questionable specimens, p. 209, of proloculus and form ratio of the present pI. 34, figs. 3, 7. species overlap those of L. asiatica, these values Yabeina obesa Skinner and Wilde, 1966, p. 50, of L. asiatica are concentrated on the end of the pI. 40, figs. 1---7. morphometrical variation of L. columbiana Yabeina dawsoni Skinner and Wilde, 1966, p. (Text-fig. 2). 50, pI. 41, figs. 1-7. Yabeina gracilis Skinner and Wilde, 1966, p. 50- The present species resembles L. japonica 51, pI. 42, figs. 1-9. (Huzimoto, 1936) in size of proloculus, but the Yabeina dunbari Skinner and Wilde, 1966, p. former has generally a smaller form ratio than 51-52, pI. 43, figs. 1,. the latter (Text-fig. 2), so far as Huzimoto's Yabeina cylindrica Skinner and Wilde, 1966, p. original description of L. japonica is concerned. 52, pI. 44, figs. 1-7. Occurrence:-The specimens at hand have Yabeina ampla Skinner and Wilde, 1966, p. 52- been collected from the type locality, Marble 53, pI. 45, figs. 1-4, pI. 46, fig. l. Canyon, and from Upper Hat Creek, British Yabeina parvula Skinner and Wilde, 1966, p. Columbia, Canada (Marble Canyon Formation 53-54, pI. 46, figs. 2-11, pI. 47, figs. 1-2. of the Cache Creek Group). They occur together Description:-Shell large, inflated fusiform to with many specimens of Schwagerina in the two elongate fusiform in. shape, with straight to localities, and also with specimens of Colania slightly curving axis of coiling, gently convex in Upper Hat Creek. to concave lateral slopes. Mature specimens of Materials:-Reg. no. KU 3001, KU 3002, KU 11 to 13 volutions or more. 5.0 to 10.0 mm 3003, KU 3004 and KU 3005 from Marble long, and 2.6 to 5.4 mm wide, form ratios of Canyon. KU 3006, KU 3007, KU 3008, KU 0.8? to 3.3 3009 and KU 3010 form Upper Hat Creek. These Outside diameter of proloculus 100 to 330 materials are collected by H. Goto, and are microns, mostly 150 to 250 microns. Spiro­ deposited in Institute of Geosciences, College of theca composed of a tectum and a keriotheca Liberal Arts, Kobe University. with very fine alveoli. Thickness of spirotheca 10 to 30 microns in outer volutions. Septa and primary transverse septula present through­ References out shell and essentially slender. Axial septula Chisaka, T. (1960): On some Permian Fusuli­ thin, first appear in 4th to 5th volution. Second­ nids from the Takagami Conglomerate, ary transverse septula first appear in 4th to 8th Choshi Peninsula, Chiba Prefecture, Japan. volution, mostly in 5th to 7th volution, usually Jour. Call. Arts and Sci. Chiba Univ., vol. 3, one septulum between adjacent primary trans­ no. 2, p. 235-254, pIs. 1-9. verse septula throughout shell, but two septula Dawson, G. M. (1879): On the new species of in most outer volutions. Loftusia from British Columbia. Geol. Soc. 817. Lepidolina columbiana from B.C. Canada 433

London Quart. Jour., vol. 35, p. 69-75, pI. vol. 25, no. 2, p. 97-114, pis. 1-2. 6. Ozawa, T. (1970): Notes on the phylogeny and Duffel, S. and McTaggart, K. C. (1951): Ashcroft classification of the Superfamily Verbeek­ Map - Area, British Columbia. Ceol. Suru. inidea. (Studies of the Permian Verbeek­ Canada, Mem. 262. inoidean foraminifera-I). Mem. Fac. Sci. Dunbar, C. O. (1932): Neoschwagerina in the Kyushu Uniu., Ser. D, Ceol., vol. 20, no. 1, Permian faunas of British Columbia. Royal p. 17-58, pis. 1-9. Soc. Canada, Proc. and Trans., 3rd ser., sec. -- (1975): Evolution of Lepidolina multi­ 4,vol. 26, p. 45-49, pI. 1. septata (Permain foraminifer) in East Asia. Geological Institute of Southwest China (ed.). Ditto, vol. 23, no. 2, p. 117-164, pis. 22- Yang (1978): Paleontological monograph of 26. Southwest China. Part 2 (Sichuan). Ceol. Ross, C. A. (1976): Development of fusulinid [nst. Southwest China, Pub., p. 1-684, pis. (Foraminiferida) faunal realms. Jour. Pale­ 1-191 (in Chinese). ont., vol. 41, no. 6, p. 1341-1354. Hanzawa, S. and Murata, M. (1963): The pale­ Sada, K. and Yokoyama, T. (1966): Upper ontologic and stratigraphic considerations on Permian fusulinids from the Taishaku the Neoschwagerininae and Verbeekininae, Limestone in West Japan. Trans. Proc. Palae­ with the descriptions of some fusulinid onto Soc. Japan, N.S., no. 63, p. 303-315, foraminifera from the Kitakami Massif, pis. 33-34. Japan. Sci. Rep. Tohoku Uniu. (Ceology), Selwyn, A. R. C. (1872): Journal and report of vol. 35, no. I, p. 1-31, pis. 1-20. preliminary explorations in British Colum­ Huzimoto, H. (1936): Stratigraphical and palae­ bia. Ceol. Suru. Con., Pept. Prog., 1871- ontological studies of the Titibu System of 72, p. 16-72. the Kwanto-mountainland, Part 2. Palae­ Skinner, J. W. and Wilde, G. L. (1966): Permian ontology. Sci. Rep. Tokyo Bunrika Daigaku, fusulinids from Pacific Northwest and sec. C, vol. I, no. 2, p. 29-125, pis. 1-26. Alaska. Uniu. Kansas Paleont. Contr., Paper Ishii, K. and Nogami, Y. (1964): Contributions 4, p. 1-64, pis. 1-49. to the geology and paleontology of Cam­ Thompson, M. L. and Wheeler, H. E. (1942): bodia. Part 1. Permian fusulinids. Jour. Permian fusulinids from British Columbia, Ceosci., Osaka City Uniu., vol. 8, Art. 2, Washington and Oregon. Jour. Paleont., p. 9-70, pis. 1-8. vol. 16, p. 700-711, pis. 105,-109. -, Okimura, Y. and Ichikawa, K. (1985): Notes --, -- and Danner, W. R. (1950): Middle on Tethys biogeography with reference to and Upper Permian fusulinids of Washing­ Middle Permian fusulinaceans. In Nakazawa, ton and British Columbia. Contr. Cushman K. and Dickins, J. M., (eds.), The Tethys, Foundation Foram. Res., vol. 1, pts. 3-4, her paleogeography and paleobiogeography p. 40-63, pis. 7-8. from Paleozoic to Mesozoic, p. 139-155, Trettin, H. P. (1980): Permian rocks of the Tokai Uniu. Press. Cache Creek Group in the Marble Range Kanmera, K. (1954): Fusulinids from the Upper Clinton Area, British Columbia. Canada Permian Kuma Formation, Southern Kyu­ Ceol. Suru. Paper, 79-17, p. 1-17. shu, Japan, with special reference to the Yamagiwa, N. and Saka, Y. (1972): On the fusulinid zone in the Upper Permian of Lepidolina zone discovered from the Shima Japan. Mem. Fac. Sci. Kyushu Uniu., Ser. D, Peninsula, Southwest Japan. Trans. Proc. Ceol., vol. 4, no. 1, p. 1-38, pis. 1-6. Palaeont. Soc. Japan, N.S., no. 85, p. 260- Morikawa, R. and Suzuki, Y. (1961): Fusulinids 274, pIs. 31-32. form the Akasaka Limestone (Part 2). Sci. Zang, L. (1982): Fusulinids of eastern Qinghai­ Rep. Saitama Uniu., ser. B, vol. B, no. 1, p. Xizang plateau. In Nanjing Institute of 43-74, pis. 4-22. Geology and palaeontology (eds.), Strati­ Nogami, Y. (1958): Fusulinids from the Maizuru graphy and Palaeontology in W. Sichuan Zone, Southwest Japan. Part 1. Ozawainel­ and E. Xizang. Acad. Sinica, Sichuan linae, Schu bertellinae and N eosch wageri­ People's Pub., p. 119-244, pIs. 1-36 (in ninae. Mem. Coil. Sci. Kyoto Uniu., ser. B, Chinese with English abstract). 434 Hiroya GOTO, Kunitaka MAR UOKA and Ken-Ichi ISHII

7' ~ T.{ 'Y -/ "" :I P :.; i::." 7 piC Lepidolina columbiana (~/t.- L, ~~*JJw.!:I:~): 7' ~ T ~ 'Y -/ "" :I P:'; i::."7j1!!lJijtl.J'~~HE. 9,fiJ!1l) Yabeina mtl.J~¥Ilfi~n-C~'Qo <';1" 1. l:iicJ-lillJijtll)<"-7' It.- 4'1'.::.*:.;c7'Y/~- "'Y r !J~-!JI.J'~ Yabeina ~c~nt-:WjW.!1;{I::{:i~M!tH... ~ni~¥Il~~nk~ll)cM~-cm~MIl)~.~AIl).N~fi~ko ~Il)M~. ~*~~.I.J' ~i1GlI\t~nt-: Yabeina ~11 Lepidolina Hii~<6IJ, 1.J'-?i'ij-f1!I::::.~-tQc*IJil'JiLt-:o ~ll)fIi ~ Lepidolina columbiana (DAWSON) c l.. -C ~G1liX Lt-:o 1~~!tf:5!r. • JLii7d~;f. • 1j;JI:{~-

Explanation of Plate 86

Lepidolina columbiana (Dawson) All figures x10 Figs. 1, 2, 6, 7. Specimens of inflated fusiform. 1. KU 3001, 2. KU 3002 from Marble Canyon. 6. KU 3006,7. KU 3007 from Upper Hat Creek. Figs. 3, 8. Specimens of fusiform. 3. KU 3003 from Marble Canyon. 8. KU 3008 from Upper Hat Creek. Figs. 4, 5, 9, 10. Specimens of elongate fusiform. 4. KU 3004, 5. KU 3005 from Marble Canyon. 9. KU 3009, 10. KU 3010 from Upper Hat Creek. GOTO et al: Lepidolina columbiana lrom B.C. Canada Plate 86

6 Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 435-446, pis. 87-89, September 30,1986

818. ON A NEW TEMPSKYA STEM FROM JAP AN*

HAR UFUMI NISHIDA

The International Budo University, 841 Monomizuka, Shinkan, Katsuura, Chiba 299-52

Abstract. Tempskya iwatensis sp. nov. was described from the Taneichi dis­ trict, Iwate Prefecture, Japan. The age of the specimen was dated as Santonian based on marine mollusca. Stratigraphically, this is the youngest Tempskya species known with certainty. The past distribution of Tempskya is plotted on paleogeo­ graphic map in relation to the Cretaceous paleoclimate.

Introduction North America, chiefly in the Rocky Moun­ tains. One species from Atlantic Coastal Plain Tempskya is a genus of leptosporangiate (T. whitei Berry) has also been excluded by ferns flourished during the Cretaceous and earlier workers for anatomical comparison be­ probably becoming extinct by the end of the cause this species is also based on poorly pre­ period. It had a tree habit sometimes attain­ served material. Endo (1926) reported a Temp­ ing a height of up to six meters. Some species skya stem from the Cretaceous of Kii (Waka­ are thought to have had a prostrate or vine­ yama Pref.). His specimen was based on a root like habit with dorsiventral stems. The stem of mass with no embedded rhizomes. Ogura (1927) Tempskya is generally described as a "false reidentified that specimen as part of a fern stem stem" consisting mainly of a mass of adventi­ with Cyatheaceous affinity, highly possibly tious roots in which are embedded slender Cyathocaulis Ogura. rhizomes and petioles of the plant. About 12 years ago Biichi Fukkiri, an amateur A comprehensive reference to the history rock collector, found a petrified trunk at the and taxonomy of Tempskya was prepared by seashore of Oh-hama, Yagi, Taneichi Town, Ash and Read (1976). According to them ten Sannohe-Gun, Iwate Prefecture (locality, 40°21' species of Temp$kya have been described up to 12"N, 141°45'30"E; Text-fig. 1). The area 1976. This excluded most of the European around the fossil locality, north of Kuji City is species described prior to T. rossica Kidston et called here the Taneichi district. Two thirds Gwynne-Vaughan (1911), which were too of the trunk was embedded in sandstone which poorly preserved to allow anatomical compari­ is widely distributed along the coast. Fukkiri son with later described tempskyas. Nine species, used the specimen to decorate a corner of his other than T. rossica, have been described in garden. Ryozo Sugiyama of the Ichinohe Senior High School (Taneichi Senior High School at *Received July 10, 1985; Revised manuscript that time) noticed the peculiar shape of the received July 15, 1986; Read June 24, 1984 specimen and advised Fukkiri to donate it to at Akita University; Contributions from the the Iwate Prefectural Museum. The specimen Laboratory of Phylogenetic Botany, Faculty is now deposited in that museum. I was asked of Science, Chiba University, No. 106. to identify the specimen by the curator of the

435 436 Harufumi NISHIDA

ceramus (Sphenoceramus) naumanii Yokoyama and Polyptycoceras cf. subundulatum (Yoko­ yama) that clearly indicate Santonian age (Matsu­ moto, 1977). This major sandstone member is exposed along the coast at Yagi where Fukkiri found the stem embedded in the sandstone of the intertidal zone. It is highly probable that the age of the stem is Santonian. The upper member consists of alternating layers of coarse­ grained sandstone and conglomerate. In the Kuji district, about 20 km south of the Taneichi district, the Upper Cretaceous shallow marine to terrestrial sediments called the Kuji Group are exposed. The Kuji Group is divided into three formations, the Tamagawa, the Kunitan and the Sawayama in ascending order. Three members of the Taneichi Formation can be stratigraphically correlated with three formations of the Kuji Group, respectively (Matsumoto et al., 1982). Tanai (1979) described a fossil flora from the Tamagawa and Sawayama Foramations of the Kuji Group. The former is dated as Late Text-fig. 1. Map showing fossil locality. Coniacian and the latter as Early Campanian based on both marine faunas and comparisons museum. Through anatomical studies the speci­ of floral composition with that of other locali­ men has been identified as a new species of ties, especially that of North Saghalien. The Tempskya and this paper deals with its descrip­ middle member of the Taneichi Formation is, tion and taxonomical treatment. therefore, correlated with the Kunitan Forma­ Geological setting and the age of the fossil:­ tion, which is Santonian in age. In the Taneichi district the Taneichi Formation of Off the Pacific coast of the Taneichi district late Cretaceous period is exposed along the the Paleogene sediments of the Noda Formation Pacific coast between Kuji City and Taneichi are exposed. Petrified woods are also known to Town. The Taneichi Formation was formerly occur from the Noda Formation, although they thought to be Miocene. It consists mainly of have not yet been studied taxonomically. It shallow-marine sediments and can be subdivided seems unlikely that the Tempskya specimen into three members. could have been derived from the Paleogene The lowest member unconformably overlies sediments. the granite basement. It consists of basal con­ Ikeya (1981) examined ESR (Electron Spin glomerate and overlying sandstone in which Resonance) of the petrified woods that had coal seams and tuff beds are intercalated. The been collected as rolled stones at the coast of middle member, which is a major part of the Taneichi district. He reported that other bio­ Taneichi Formation, consists of greenish gray logical materials of the same age show char­ sandstone. Oyster beds partly intercalate in acteristic ESR patterns. Ikeya compared the the lower part of this member. From this mem­ ESR of the rolled stones of Taneichi district ber Terui et al. (1975) reported the occurrence with that of in situ petrified woods collected of a Cretaceous marine fauna including Ino- from both the Taneichi and the Noda Forma- 818. New Tempskya from Japan 437

tions. The result indicated that the ESR of the by roots. The anatomical description which rolled stones is very similar to that of the Creta­ will be given below is derived from observa­ ceous in situ material. To ascertain the age of tions of these incompletely preserved compo­ the Tempskya specimen the examination of its nents. Description of certain diagnostic features ESR pattern will provide useful evidence. that are thought to be taxonomically important, for example, the relative length of internodes Material and method and amount of xylem parenchyma, are in con­ formity with the characteristics used in the The specimen before cutting was a petrified artificial key to the species prepared by Ash columnar trunk, yellowish white on the surface, and Read (1976). about 50 cm long, with diameters ranging from The rhizomes are very slender, 2.7 - 3.6 mm 7 to 11.5 cm. One end is almost smooth but is in diameter, with long to medium internodes, slightly knobby. The trunk weakly tapers to with slight overlapping leaf bases (usually two one end. It was cut into five pieces as is shown or three). Leaf traces depart from only one in Text-fig. 2. side of a rhizome alternately in two rows (PI. Peels prepared by etching the material about 87, Fig. 1). The epidermis is not clearly defined. four minutes in 20 to 25 percent hydrofluolic No dermal hairs nor their remnants are present. acid solution were made into slides for study. A hypodermal layer that is comparable to the outer cortex as described by Ash and Read Systematic description (1976) was not clearly defined. The only evi­ dence of a hypodermal layer is one or two ex­ Order Filicales ternal layers of the outer cortex that consist Family Tempskyaceae Read and Brown, 1937 of less sclerotized cells (PI. 87, Fig. 3). The cortex consists of two layers. The outer cortex, Genus Tempskya Corda, 1845 that is comparable to the middle cortex cells Tempskya iwatensis H. Nishida, sp. nov. in other species, is sclerotic, about six or seven PIs. 87-89; Text-fig. 2. cells wide, with cells that are simple-pitted and contain reddish brown contents. The inner Description :-Other than some longitudinal cortex is 15 - 17 cells wide, and composed of grooves the specimen has a rather smooth ex­ thick-walled parenchyma cells that become ternal surface. It constitutes a so-called false smaller near periphery of the layer. stem that is characteristic of Tempskya, consist­ The stele is solenostelic with a well-developed ing of numerous adventitious roots and a small pith (PI. 87, Fig. 4). The endodermis is a single number of solenostelic rhizomes and petioles. layer of tangentially flattened cells. The pericycle In the false stem the rhizomes give rise to peti­ is two to three cells thick. Internal to the peri­ oles in all directions. Therefore the false stem cycle is a layer of very small cells. This layer is does not exhibit dorsiventrality (Text-fig. 2). not always detected but when present probably In section 1 (Text-fig. 2) of the false stem no represents protophloem. Metaphloem is devel­ rhizome was present. Rhizomes appear and oped internally to the protophloem and is a increase in number toward the other end of layer three to five cells thick. The xylem plate section 1 indicating that it is nearer the base of the concentric bundle is slightly thicker on of the false stem. Throughout the length of the ventral side and the tracheids of which it the false stem the exact number of rhizomes is composed are scalariform. The position of the could not be determined because most of the proto xylem is not clear because of nearly equal rhizomes had been decayed before fossilization. size of tracheids. However, smaller elements The rhizomes and petioles are poorly pre­ tend to be located at the outer edge of the served because of decay and some disruption xylem cylinder. Xylem parenchyma is relatively 438 Harufumi NISHIDA

b

1 em C-U; ~ d-( \ ,: '' : : A I '

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"- eo " "5l E ~ u .,., ... 0 @ '"

'0\ '!J ~ ,. C • , ~ " "~1. @, ..~ .~I" It~-

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.., 4 "" 818. New Tempskya from Japan 439 abundant (PI. 88, Fig. 1). Parenchyma cells are The epidermis is decayed and thus lost. The often alined in groups of five to more than outer cortex is sclerenchymatous and consists eight cells. The cells of endodermis, pericycle of cells similar in structure and contents to the and xylem parenchyma contain reddish brown cells of the corresponding layer in the rhizome. contents similar to those occur in outer cortex. It is 15 - 17 cells thick at petiole base. The The periphery of pith, which is about three inner cortex consists of thick-walled parenchyma cells thick, consists of thick-walled parenchyma cells. The stele is C-shaped and consists of an cells (PI. 87, Fig. 4). Toward the center of the amphiphloic concentric bundle with a continuo pith cells become smaller and thick·walled. The ous C·shaped xylem plate. Both adaxial margins preservation of the pith is quite poor and the of xylem plate curve inward. The abaxial arc pith area is often invaded by adventitious roots. of the xylem strand is rather flat. Near the However, at places the pith consists of sclerotic point of insertion of the petiole its xylem strand cells as in other species of Tempskya (PI. 87, is thicker and lacks apparent proto xylem groups Fig. 2). (PI. 88, Fig. 3). Distally in the petiole the xylem Roots depart endogenously from the sole­ strand diminishes in thickness until it becomes nostele of the rhizome. The size, shape, state one or two cells thick except for its adaxial of development and state of preservation of margins where xylem is three or more cells roots vary considerably (PI. 89, Fig. 2). They thick. As the xylem strand thins three proto­ are equal to the roots of other species of Temp­ xylem groups become apparent (PI. 88, Fig. 2). skya in general structure. Root hairs are present At places endodermis, pericycle, protophloem (PI. 89, Fig. 4). The cortex consists of outer and metaphloem are preserved (PI. 89, Fig. 1). and inner layers. The outer cortex is parenchy­ The center of pith consists only of thick-walled matous, with cells that are more thick-walled parenchyma cells (PI. 89, Fig. 1). near the periphery. The inner cortex is generally Diagnosis of the species:-False stem radially sclerenchymatous, although the degree of sclero­ symmetrical, 7 to 12 cm in diameter at base. tization varies. In some roots the inner cortex Rhizome slender, 2.7 to 3.6 mm in diameter. is completely thick-walled parenchymatous, or Internodes long to medium usually two or three in rare cases only a layer adjacent to the endo­ overlapping petiole bases in one section. Cortex dermis is parenchymtous. two-layered; outer layer sclerotic, inner layer In mature roots the xylem is diarch, whereas thick-walled parenchymatous. Much xylem pa­ in juvenile roots such as shown in PI. 89, Fig. 3, renchyma in xylem strand. Proto xylem in­ the central xylem mass is too small to be clearly definite in xylem of rhizome. Pith of the petiole defined. In such young roots the inner cortex consists of thick-walled parenchyma. is not typically sclerotized, although the degree Locality:-Oh-hama, Yagi, Taneichi Town, of sclerotization is not always correlated with Sannohe-Gun, Iwate Prefecture; 40021'12''N, the age of roots. 141 °45'30"E. Petioles are very slender, 1.2 x 1.6 mm in Horizon and age:-Taneichi Formation; Upper diameter, and do not persist in the tissue of the Cretaceous (Santonian). false stem after departure from the rhizome. Specimen:-Holotype (IPMM 31003) is de­ However, the petiole is better preserved than posited in the Iwate Prefectural Museum. Both the rhizome and persists for a greater distance ends of the holotype specimen and their micro­ in the false stem than the parent rhizome. preparations are deposited in the Laboratory of

~ Text-fig. 2. External features of specimen IPMM 31003 (E) and serial cross sec­ tions of upper part of false trunk (A-D) showing its radially symmetrical constitution. A to D correspond to the surfaces indicated as a to c. Arrowhead indicates the level where the section 1 mentioned in the text was made. 440 Harufumi NISHIDA

Phylogenetic Botany, Faculty of Science, Chiba false stem indicates that the specimen represents University. a basal part of the false stem. Based on careful examination of numerous false stems, Andrews and Kern (1947) estimated that T. wesselii Affinity and discussions attained a height of 3.6 m if the basal diameter Affinity:-Nine species of Tempskya deserve is 25 cm. The thickest false stem of T. wesselii anatomical comparisons with the specimen rep­ attains a diameter of 40 cm or more. With this resented here (Table 1). According to the separa­ diameter as a basis for calculation the plant tion of species by Ash and Read (1976) the would have been nearly 5.8 m high. If it is pos­ specimen belongs to the intragenetic group that tulated that the height of a plant is in propor­ has radially symmetrical false stems. Among tion to the basal diameter of its stem, the height species with radially symmetrical false stem T. of T. iwatensis with a false stem 10 - 12 cm in rossica is compared favorably with the Iwate diameter at base, is estimated as 1.45 - 1.74 m. specimen in the histology of the cortex, amount Andrews and Kern also mentioned that the of xylem parenchyma and the length of inter­ lower one sixth of the false stem consists of a nodes, which are characters regarded by earlier root mass alone. The lower 25 to 30 cm of the workers to be taxonomically valuable. The new Iwate specimen appears to consist of a root form is distinguished from T. rossica by the mass without rhizomes. smaller size of the rhizome and the absence of Age of Tempskya:-Most Tempskya species a central depression in the abaxial arc of a leaf known to date are derived from the Lower trace. Although T. zelleri also has a histology Cretaceous strata. Ash and Read (1976) revised similar to the Iwate specimen, it is highly char­ the age of Tempskya-bearing beds in North acteristic in having angular stelar strands that America including the Patapusco Formation never occur in the species described here, as well bearing T. whitei. Most beds are now believed as other early described species. to be Late Albian in age (Table 1). Some species Apart from the radially symmetrical com­ extend from Late Albian into Early Cenomanian. position of the false stem the specimen ana­ According to Read and Brown (1937), European tomically resembles T. knowltoni and T. minor, tempskyas occur mainly in Wealden (Berriasian both belonging to the dorsiventral group. Smaller to Barremian), but in part in Aptian (Lower size of rhizome, simple disposition of cortical Greensand). Perucer beds in Czechoslovakia, tissues and long to medium internodes are the which yield T. varians Velenovsky, are con­ common features. The specimen is distinguished sidered to be Cenomanian in age (Read and from T. knowltoni in having many free petioles Brown, 1936). Tempskya rossica from the Kara­ in the false stem and in having a larger quantity ganda River Basin is the only specimen that is of parenchyma in xylem strand. T. minor has derived from unconfirmed Upper Cretaceous many free petioles in the false stem but lacks strata. With the possible exception of T. rossica, xylem parenchyma. The petiolar pith consisting T. iwatensis is presentely the only Tempskya of thick-walled parenchyma is characteristic known to occur in the Upper Cretaceous. of the specimen and is a feature not found in Ferns associated with Tempskya:-Fern leaves other species. Although the Iwate specimen is are lacking in the Tempskya beds of the Taneichi partially characterized by poorly delineated Formation. However, from the Kuji Group rich characters such as the variable amount of xylem fern flora has been recorded by Tanai (1979). parenchyma, the combination of features de­ Although the Kunitan Formation, which is cor­ scribed here lead me to designate the specimen related with the Tempskya-bearing bed of the as a new species. Taneichi Formation does not contain plant Size of the plant:-Absence of well-preserved fossils, both the Tamagawa and Sawayama rhizomes and petioles in whole length of the Formations are rich in ferns. The Gyliakian Table 1. Anatomical comparisons of Tempskya iwatensis, sp. nov., with nine species of anatomically preserved Tempskya.

Size of Internode Outer Inner Size of Xylem Species Age False stem* rhizome length** eorlex*** cortex false slem paren· (mm) (em in diam.) ehyma

T. rossiea Senonian? R 6.0-7.0 0 se. pa. 5.5-9.5 2**** Kidsloll et Gwynne·Vaughan

T. Imowltolli Seward L. Albian- D 2.5-3.5 0 se. pa. 1.5-6.5 2 E. Cenomanian

T. grallciis Read et Brown L. Albian R 4.6-6.0 - se. pa. + irr. se. 10.4-]2.2 3 00 T. millor Read et Brown L. Aptian- D 2.0-3.5 0 se...... pa. 9.0-14.4 1 90 Albian ~ T. wesselii Arnold L. Albian R 4.0-5.0 + se. pa. + 7.0-40.0 3 E; doub. se. or more ~ T. wyomillgellsis Arnold L. Albian R 6.0-8.0 0 se. pa. + ? 3 ~ doub. se. i'" T. superba Arnold L. Albian? R 10.0-15.0 - sc. pa. + irr. se. more than 3 ~c 12.0 ~ ~ T. reesiciei Ash et Read L. Albian R 3.0-5.0 0 pa. pa. island 10.0-30.0 2 'g in se. ;:s

T. zelleri Ash el Read L. Albian R 3.0-8.0 + se. pa. 17.0 2

T. iwaiellsis, sp. nov. Santonian R 2.7-3.6 0 se. pa. 7.0-12.0 3

*R: radially symmetrical, D: dorsiventral. **+: very long, 0: long to medium, -: very short. ***In some species this layer is originally described as middle cortex. ****3: much, 2: little, 1: absent.

>I>­ .....>I>­ 442 Harufumi NISHIDA

flora of North Saghalien of the same age as T. for much wider and detailed investigations on iwatensis contains 21 species of ferns, of which these Cretaceous ferns in an effort to clarify eight are common to the Tamagawa and Sawa­ the taxonomic position of Tempskya. yama floras. Because the affinities of Tempskya Other plants associated with Tempskya:­ are still uncertain it is impossible to relating Seward (1924) reported the occurrence of an Tempskya stems to the isolated remains of araucarian root associated with T. knowltoni. fern leaves. When Arnold (1945) described T. wyomingensis In looking for possible affinities of Tempskya he recorded Cyacadeoidea sp. from the same to fern families several earlier workers have locality. According to Andrews and Kern (1947) emphasized the occurrence of Anemia fremonti Cupressinoxylon sp., Cycadeoidea sp. and di­ Knowlton foliages (Schizaeaceae) in a horizon cotyledonous woods occur with T. wesselii in the same as or close to that bearing Tempskya Idaho. (Seward, 1924; Read and Brown, 1937). The From the Taneichi Formation four species Sawayama flora contains A. elongata (Newb.) of gymnospermous woods have been recognized Knowlton and the Gyliakian flora contains A. by M. Nishida (pers. com.). Although the de­ cf. elongata (Kryshtofovich and Bykovskaya, tailed taxonomic accounts will be published 1960). elsewhere, he tentatively recognizes Araucario­ Many anatomically preserved fragments of xylon kiiense Ogura, Cupressinoxylon vectense ferns embedded in Late Cretaceous calcareous Barber, Taxodioxylon albertense Shimakura and nodules have been described from Hokkaido a new species of Piceoxylon. Cycadeoidea was (Stopes and Fujii, 1910; Ogura, 1927, 1930; described from the Late Cretaceous Upper Yezo Hashimoto, 1971; Nishida and Nishida, 1979; Group of Hokkaido (Kryshtofovich, 1920; Nishida, 1981a, b). Most of these belong to Endo, 1925). The floral components associated Cyatheaceae. The age of the strata bearing with Tempskya in Japan and those in North such nodules extends from Cenomanian to America are very similar and indicate similar Campanian, and is partly comparable with the paleoecological conditions at these localities. age of the Taneichi Formation. Only one spore­ A possible habitat for Tempskya, suggested by bearing fragment, Schizaeopteris mesozoica Andrews and Kern (1947) is a tropical montane Stopes et Fujii, however, suggests the presence cloud-forest, where most species of the tree of Schizaeaceae. There is no record of Tempskya ferns occur today. from either Hokkaido or Saghalien. To avoid Distribution of Tempskya and its relation to further speculation I emphasize here the need paleoclimate:-Parrish et al. (1982) predict rain-

Explanation of Plate 87

Fig. 1. C.s. of dorsal half of a half-decayed rhizome, showing leaf traces and free petioles (arrow­ heads) departing in two rows. X 15.6. Bar 1 mm. Fig. 2. C.s. of a best preserved rhizome. Sclerotic pith is partly preserved (arrowhead). Outer cortex is decayed but inner cortex consisting of thick-walled parenchyma is well preserved. X 29.0. Bar 0.5 mm. Fig. 3. C.s. of a rhizome showing construction of tissues. Outer cortex is preserved only at its periphery. Cells of outermost layer of outer cortex are larger and thinner-walled than inner cells, indicating the presence of a hypodermal layer. ic; inner cortex, oc; outer cortex, vb; vascular bundle, p; pith. X 30.7. Bar 0.5 mm. . Fig. 4. Vascular tissue in fig. 3 enlarged. Protoxylem indefinite. Note abundant xylem paren­ chyma. Phloem is badly preserved. Periphery of pith consists of thick-walled parenchyma. e; endodermis, pe; pericycIe, ph; phloem, xy; xylem. X 61.4. Bar 0.1 mm. NISHIDA : New Tempskya from Japan Plate 87 818. New Tempskya from Japan 443 fall patterns, based on atmospheric circulations reviewed by Read and Brown (1937). North of the past, to explain the distribution of coals American localities are mapped by Ash and and evaporites in the Mesozoic and Cenozoic. Read (1976). In North America, Tempsllya Paleobotanical data are useful in testing such a occurs in the Rocky Mountain Geosyncline, hypothesis. Text-fig. 3 shows a distribution of Mexican Geosyncline and Atlantic Coastal Tempskya plotted on a paleogeographic map. Plain, extending from 3Io N to 4SoN and 103°W The base map (Owen, 1983) shows the distri­ to 118°W. In Europe Tempsllya-bearing beds bution of the continents in the Turonian stage. are located from 500 N to 53°N and 2°W to Contours of rainfall patterns (relative values 65°E. The locality of T. iwatensis is approxi­ o only) are based on Cenomanian data by Parrish mately 40 N, and 141° E. et al. (1982) who also present Maestrichtian data. The Rocky Mountains and Mexican Geosyn­ Although the age of Tempskya extends from clines, which are major sources of North Ameri­ Berriasian to Santonian, I preferred here to use can Tempsllya, are developed in the eastern Cenomanian data as being representative of the interior of the Rocky Mountains that provided paleoclimate of Tempskya age. Because judging a rain shadow against the westerlies. Another from the paleoclimate maps by Parrish et al. North American locality faces the Atlantic coast. (1982), the paleogeography and paleoclimate of In Europe, England and Ireland localities and Tempskya-bearing localities had been subject to French to Bohemian localities are all on islands. minor change through the Cretaceous period. The Karaganda River Basin near the Ural Moun­ The worldwide occurrence of Tempskya is tains is situated at the margin of the Asian conti-

o· - .,. - -1

Text-fig. 3. Distribution of Tempshya on paleogeographic map showing rainfall patterns in Cenomanian. Paleogeography and rainfall patterns are redrawn after Parrish et al. (1982) on base map by Owen (1983). Numbers show relative values of rainfall. 200; high, 200-100; moderately high, 100-50; modera tely low, 50; low, shaded; approximate contour of land masses. 444 Harufumi NISHIDA

nental lowland on the north coast of Tethys. by wet oceanic winds. The possible habitat of The Taneichi district is located at the east Tempskya could be a humid low to high mon­ Pacific margin of Asia continent. tane forest in subtropical or even a warm tem­ Tree ferns need high humidity for their perate climate. The fact that evaporites are continuous growth and thus the postulated never associated with these localities supports habitat for Tempskya is a tropical or subtropical the idea of a lower evaporation rate resulting humid forest developed at high elevations (1,200 in high humidity (Parrish et al., 1982). - 2,100 m) on undulating hills or mountains Tempskya has not yet been described from (Andrews and Kern, 1947). In tropical to sub­ Asian inlands at latitudes comparable to those tropical South America, cloud-forests rich in of other localities. This may be the product of ferns are well developed on the Andean slopes a dry climate. Much more information must at higher elevations (1,800 - 2,600 m). be obtained, however. Wide latitudinal distri­ Species of Tempskya are distributed in low­ bution of the American Tempskya is in marked lands at rather high latitudes except in North contrast to the restricted distributional pattern America where several localities occur at nearly of the European members. In North America 31°N. In these localities rainfalls are predicted a moist habitat occurs along the northward to be moderately high as well as moderately extension of the north Atlantic, whereas in low without affecting the distribution of Temp­ Europe land masses are confined to latitudes skya. These paleogeographical and paleoclima­ around 50o N. This may account for restricted tological conditions differ somewhat from the distribution of Tempskya in Europe. postulated habitat of Tempskya. On the naturalness of intragenetic groups:­ From my observations of living species of Whether the false stem of Tempskya is radially Cyatheaceae in South America, Australia and symmetrical or dorsiventral has been used as south Japan, continuous atmospheric moisture one of important criteria in classifying species seems to be a most important factor limiting in~o two subgeneric groups (Read and Brown, their growth rather than total amount of rain­ 1937; Andrews and Kern, 1947; Arnold, 1958; fall, if the temperature is moderate. Under Ash and Read, 1976). However, Andrews and moderate temperatures coastal cliffs or moun­ Kern (1947) questioned this grouping and tains of no more than 400 - 500 m seems to be suggested that dorsiventrality reflects either a high enough to produce sufficient humidity for juvenile stage or only a part of an entire false tree-fern growth. The Juan Fernandez Islands stem of a plant. No fossil evidence supporting and Pacific coast of South Chile are examples these suggestions has been discovered as yet. of such localities. According to Parrish et al. The dorsiventral group (T. knowltoni and T. (1982) the climate in a given area is regarded minor) has slender rhizomes less than 3.5 mm in to be wet when precipitation exceeds evapora­ diameter. The radially symmetrical group has tion, even though the total amount of pre­ thicker rhizomes usually more than 4. 0 mm in cipitation is low. All localities are placed either diameter, except for T. reesidei and T. zelleri near the coast or on islands that are influenced where diameters decrease to 3.0 mm in the lower

Explanation of Plate 88

Fig. 1. C.s. of rhizome. Part of a xylem strand. Abundance of xylem parenchyma is better dis­ played. X 170. Bar 0.1 mm. Fig. 2. Enlargement of a free petiole shown in Plate 87, Fig. 1. Thin xylem strand with three protoxylem groups (arrowheads) is shown. Outer cortex is lost. X 140. Bar 0.1 mm. Fig. 3. C.s. of a petiole base showing a vascular strand without central depression on its abaxial arc. Protoxylem is not clear in xylem plate. Upper side is abaxial. X 43. Bar 0.5 mm. NISHIDA: New Tempskya from Japan Plate 88 818. New Tempskya from Japan 445 extremities of the rhizome. The simple structure men that is now deposited in the Iwate Prefec­ of cortical tissues is another feature common tural Museum to which I also acknowledge for to the dorsiventral group. The consistent rela· the loan of the specimen. This study was partly tionship between internal structure and the supported by a Grant-in-Aid for Encouragement dorsiventrality appears to support the natural· of Young Scientists from the Ministry of Educa­ ness of the grouping. tion, Science and Culture, No. 60740385. T. iwatensis, however, displays characteristics that are intermediate between the two groups. References This species belongs to the radially symmetrical group, but at the same time displays a slender Andrews, H. N. and Kern, E. M. (1947): The Idaho tempskyas and associated fossil rhizomes and a simple disposition of cortical plants. Ann. Missouri Bot. Gard., vol. 34, tissues as in the dorsiventral group. This combi· no. 2, p. 119-183, app. p. 185-186, pis. nation of characteristics in T. iwatensis suggests 15-27. that there is no strict boundary between the Arnold, C. A. (1945): Silicified plant remains two groups. The construction of the false stem from the Mesozoic and Tertiary of western is still, however, a good feature in identifying North America. 1. Ferns. Papers Michigan Tempskya species. Acad. Sci. Arts and Letters, vol. 30, p. 3- The rhizomes of Tempskya are dorsiventral. 34, pis. 1-11. Therefore, the fern that is hypothetically an­ --(1958): A new Tempskya. Contr. Michigan cestral to Tempskya could have had prostrate, Univ. Mus. Paleontology, vol. 14, no. 8, p. 133-142, pis. 1-3. dorsiventral rhizomes like those of the living Ash, S. R. and Read, C. B. (1976): North Ameri­ schizaeaceous ferns. The false stems first formed can species of Tempskya and their strati­ by such an ancestor probably was a dorsiventral graphic significance. U.S.G.S. Prof. Paper, type, so that the primitive form of Tempskya 874, p. 1-42, pis. 1-13. would have structure similar to T. minor with Endo, S. (1925): Nilssonia-bed of Hokkaido its dorsiventral stem and that exhibits the sim­ and its flora. Sci. Rep. Tohoku Imp. Univ., plest structure of all. The slightly younger age of 2nd Ser., vol. 7, no. 3, p. 57-72, pis. 11-17. T. minor supports this idea. However, it remains --(1926): On the genus Tempskya from Kii. uncertain whether or not T. minor and another Chikyu, vol. 6, p. 5-6 (in Japanese). dorsiventral species, T. knowltoni, are direct Hashimoto, W. (1971): On a new Cretaceous tree descendants of the ancestral form. Tempskya fern from Nakagawa-machi, Teshio Province, Hokkaido. Sci. Rep. Tokyo Kyoiku Daigaku, is a highly variable group that diversified from Sec. C, vol. 11, no. 103, p. 1-10, 3 pis. a simple ancestral form in the Cretaceous. T. Ikeya, M. and Miki, T. (1981): ESR (Electron iwatensis is a species, that in the course of its Spin Resonance) of fossil bones, woods evolution, has retained the simple internal and natural minerals. Earth, vol. 3, no. 8, structures of some other tempskyas. p. 477-483 (in Japanese). Kidston, R. and GWynne-Vaughan, D. T. (1911): On a new species of Tempskya from Russia. Acknowledgement Verh. Russ. K. min. Gesell., vol. 48, p. 1- I am grateful to Prof. Makoto Nishida of 20, pis. 1-3. Kryshtofovich, A. N. (1920): A cycadean trunk Chiba University for his continuous encourage­ from Hokkaido. Jour. Geol. Soc. Tokyo, ment and suggestions. Dr. Wilson N. Stewart, vol. 27, no. 325, p. 1-8, pI. 19. Prof. Emeritus of the University of Alberta, --and Baykovskaya, T. N. (1960): Mesozoic Canada, kindly reviewed the manuscript and gave flora of Saghalien, 122 p., 21 pis., Academia me valuable suggestions. Thanks are due to Mr. Nauka, Moscow (in Russian). Ryozo Sugiyama of Ichinohe High School for Matsumoto, T. (1977): Zonal correlation of the giving me an opportunity to examine the speci- Upper Cretaceous of Japan. Pal. Soc. Japan, 446 Harufumi NISHIDA

Owen, H. G. (1983): Atlas of continental dis­ Spec. Paps .• no. 21, p. 63-74. placement, 200 million years to the Present. --, Obata, I., Tashiro, M., Ohta, Y., Tamura, 159 p., Cambridge Univ. Press, London. M., Matsukawa, M. and Tanaka, H. (1982): Parrish, J. T., Ziegler, A. M. and Scotese, C. R. Correlation of marine and non-marine for­ (1982): Rainfall patterns and the distribu­ mations in the Cretaceous of Japan. Fossils, tion of coals and evaporites in the Meso­ no. 31, p. 1-26 (in Japanese): zoic and Cenozoic. Palaeogeogr.. Palaeo­ Nishida, H. (1981a): Anatomical studies of a new climatol.• Palaeoecol .• vol. 40, p. 67-101. specimens of Yezopteris polycycloides Read, C. B. and Brown, R. W. (1937): American Ogura from the Upper Cretaceous of Hok­ Cretaceous ferns of the genus Tempskya. kaido, Japan. Jour. Jap. Bot., vol. 56, p. U.S.G.S. Prof Paper. 186-F. p. 105-131, 169-180, pI. 8. pis. 16-17. --(1981b): A revision of the genus Cya­ Stopes, M. C. and Fujii, K. (1910): Studies on thorachis in Japan. Bot. Mag. Tokyo, vol. the structure and affinities of Cretaceous 94, p. 249-259. plants. Phil. Trans. Roy. Soc. London, B. --and Nishida, M. (1979): Thyrsopterorachis. vol. 201, p. 1-90, pIs. 1-9. gen. nov., a tree fern stipe from the Upper Tanai, T. (1979): Late Cretaceous floras from Cretaceous of Hokkaido, Japan. Ibid .• vol. the Kuji district, northeastern Honshu, 92, p. 187-195. Japan. Jour. Fac. Sci. Hokkaido Uniu .• Ser. Ogura, Y. (1927): On the structure and affinities 4. vol. 19, no. 1-2, p. 75-136, pIs. 1-14. of some fossil tree-ferns from Japan. Jour. Terui, K., Terui, K., Yanagisawa, H. and Koba­ Fac. Sci. Imp. Uniu. Tokyo. Bot.. vol. 1, p. yashi, T. (1975): Cretaceous fossils from the 351-380, pIs. 2-8. Taneichi Formation in the north coast of -(1930): On the structure and affinities of Rikuchu, Iwate Prefecture. Jour. Geol. Soc. some Cretaceous plants from Hokkaido. Japan, vol. 81, no. 12, p. 783-785. Ibid .• vol. 2, p. 381-412, pI. 18-21.

Taneichi :fl1!m. Kuji 7-.~. Oh-harna *~. Vagi /\*. Tarnagawa :li)II. Sawaya rna /Rlli. Kunitan 1'!!Ift. Noda !!fm. Hokkaido ~tila~.

B *£7;/ 7 7-=\'- 70) ~:fl1!t.:."?~'-C: !ft=F~~PlillllmlBJ /\**~O) ~JIr.. '-KHNIO) fijl!ft.t ~ ••~ •••~U.Lk.~*~. *~*~.O)*~~~ 7;/77-=\,-7 (Thm~~a). O)ifi.:il:tIl~ c nn' -? t.:o t~*li:fl1!mJ\1. 11jilJW~t.:.7};ffl Lt.::;: c n;? n'n;nhoo .1*J007}1tillFt.:. "? ~'-C\-dNFa-ffifUr:t.:o l§1Ri'int

Explanation of Plate 89

Fig. 1. Enlargement of a petiolar vascular bundle at its adaxial margin. Pith consists of thick­ walled parenchyma (arrowhead). e; endoderm is. X 227. Bar 0.1 mm. Fig. 2. Adventitious roots displaying various degrees of preservation and construction of tissues. X 31. Bar 0.5 mm. Fig. 3. Adventitious root with inner cortex composed of cells having less thickened walls. Xylem mass is poorly developed. X 81. Bar 0.1 mm. Fig. 4. Long intermingled root hairs. X 163. Bar 0.1 mm. NISHIDA : New Tempskya from Japan Plate 89 Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 447-462, pIs. 90-92, September 30, 1986

819. THE SECOND ADDITION TO THE FAUNA OF YOKOKURA-Y AMA, SHIKOKU, JAP AN*

TEIlCHI KOBAYASHI

Japan Academy, Ueno Park, Taito-ku, Tokyo

and

TAKASHI HAMADA

Department of Earth Science and Astronomy, College of Arts and Sciences, University of Tokyo, Komaba, Meguro-ku, Tokyo

Abstract. Nine species of including six new species and two new subspecies from Mt. Yokokura are described in this article. A new genus, Ichi­ yamella, is instituted on Ichiyamella subglobula, sp. nov. Besides this genus Japono­ scutellum and Apolichas are indigenous to Japan or Eastern Asia. Some notes are given on the Scutelluidae, Meroperixinae, Illaenidae, Goldillaeninae, Sphaerexo­ chinae, Dindymeninae and the Lichidae in addition to Sphaerexochus, Dindymene and Dicranopeltis and Dindymene (?) megacranidia.

In the monograph on Silurian Trilobites and Hamada. of Japan etc. (1974), 26 species of trilobites Japonoscutellum japonicum puteatum, subsp. were described from Mt. Yokokura. Recently Japonoscutellum japonicum laticephalum, subsp. 11 species were added to them (1984, 1985). nov. A further addition is made here with 9 species Subfamily Scutelluinae R. and E. Richter (?) and subspecies as listed below. The faunal analy­ Scutelluid, gen. et sp. indet. Subfamily.Meroperixinae Kobayashi and sis is postponed to another occasion. Hamada+ Palaeontological notes are given on the families Opoa (?) trinodosa, sp. nov. and subfamilies marked with plus in the list Family Illaenidae Hawle and Corda+ below beside a few genera. Ichiyamella is a new Subfamily Goldillaeninae Balashova+ genus proposed on this occasion. Illaenoides (?) magnisulcatus, sp. nov. Illaenoides (?) abnormis, sp. nov. List of additional trilobites Family Proetidae Salter Subfamily Cyphoproetinae Pillet Family Scutelluidae R. and E. Richter+ Cyphoproetus latiaxis, sp. nov. Subfamily Planiscutelluinae Ko bayashi Family Cheiruridae Salter Subfamily Sphaerexochinae Opik+ Sphaerexochus hiratai Kobayashi and Hamada *Received September 17, 1985; Studies on Family Encrinuridae Angelin Japanese Trilobites and Associated Fossils, Subfamily Dindymeninae Kielan+ No. XXXIX.

447 448 Teiichi KOBA YASHI and Takashi HAMADA

Dindymene (?) sp. indet. and produced into genal spines. Pygidium is Ichiyamella subglobula, sp. nov. bell-shaped; axial lobe nearly as long as broad, a Ichiyamella megacranidia (Kobayashi and Hama­ little shorter than one-third of pygidium; median da) rib bifurcate in posterior; 7 pairs of pledural Family Lichidae Hawle and Corda+ ribs present on its two sides. Lamproscutellum Subfamily Lichinae Hawle and Corda shaanxiense Zhou, 1982 and L. latilimbatum Dicranopeltis tricornis, sp. nov. Zhou, 1982, occur in Middle , Sha­ Subfamily Homolichinae Phleger an xi. Apolichas truncatus Kobayashi and Hamada Leioscutellum Wu, 1977 founded on L. tenuicaudatum Wu, 1977 from Lower Silurian Ack:lOwledgments in Guizhou, resembles Paralejurus, but its glabella is less expanded in anterior, occipital furrow de­ Here the authors record their best thanks to scribing two arcs, eye large and genal angle Messrs. S. Asami, I. Fujiyama, A. Kaneko, F. rounded on cephalon and 8 pairs of pleural ribs Kojima, Y. Mizuno, S. Nakajima, K. Noda and on two sides of the simple somewhat broad T. Uchida for collecting the trilobites dealt with median rib on the bell-shaped pygidium. here, and to the Japan Academy for the grant to Australoscutellum Chatterton and Campbell, this study. 1980 was instituted on Bronteus longispinifer Mitchell, 1887 from the Ludlovian in New Description of trilobites South Wales, Australia. According to the authors Family Scutelluidae Richter and Richter, 1956 of the genus it is allied to Microscutellum in the glabellar pattern, but it has only 5 or 6 pairs Leioscutellum and Lamposcutellum were of pleural ribs, besides a median rib which seems described from China respectively by Wu (1977) to be trisected. This genus indicates probably and Yin (1980). The latter is one of the oldest an early isolate branch of the Thysanopeltinae. scutelluids that bear some characteristics of the Un icapeltis Maximova, 1977, whose type­ Eobronteinae as well as the Planiscutelluinae. species is Unicapeltis unica Maximova, 1977 The former may be the oldest genus of the from Middle of Salair, is quite dis­ Paralejurinae. tinctive in its glabellar configuration, particularly Lamposcutellum Yin, 1980, whose type is in the connection of the posterior pair of pro­ L. guizhouense Yin, 1980, from Middle Ordo· found lateral furrows in describing a backwardly vician in Guizhou, is allied to. Eobronteus, convex arc in the axial part. A pair of shallow Protobronteus and Planiscutellum in one or paraxial furrows extend forward from both another characteristic. Its glabella is constricted sides of this arc delimiting middle lateral lobes. at about one-third of the cephalon from the Nevertheless, this genus resembles Thysano­ posterior margin, more expanded in anterior, peltella, Arctopeltis and Altaepeltis. It would reaching the frontal margin and provided with be another aberrant genus of the Thysano­ three pairs of lateral furrows; occipital furrow peltinae. transversal; neck ring uniform in sagittal length; axial furrow well pronounced in middle and pos­ Subfamily Planiscutelluinae Kobayashi terior; small paraxial nodes occasionally present and Hamada, 1974 at the constriction; eyes of medium size opposed Genus Japonoscutellum Pftbyl and Vanek, 1971 at posterior furrows and lobes; eye-ridge extend­ ing from middle lateral furrow to eye; fixed Japonoscutellum japonicum (Kobayashi cheek anterior to this ridge divided into two and Hamada) puteatum, subsp. nov. subequal parts by another ridge; lateral marginal PI. 90, Figs. la-c. borders extending from two sides of glabella 819. Silurian Trilobites from Shikoku, 2nd Addition 449

This cephal on agrees with J. japonicum in furrow to palpebral lobe; ala small, suboval, major aspects, but it disagrees with the typical located on lateral side of posterior lateral lobe form of the species in the middle and anterior of glabella. Facial suture diagonal posterior to lateral furrows which are disconnected from eye; its anterior branch extending from eye to the axial furrow and strongly pitted, somewhat lateral end of frontal glabellar lobe. Fine striae smaller eyes and the distinct eye-ridge and ala. like terrace-lines on doublure seen most of dorsal The cheek is narrow and depressed, but test which are subparallel anterior margin on provided with a narrow convex lateral border. glabella, particularly dense and distinct on A narrow ridge runs along the lateral margin of frontal lobe, more or less longitudinal on neck the eye. The anterior branch of the facial suture ring and subparalltl to axial furrow on fixed extends from the eye and then diagonally crosses cheek. the cheek to the marginal border, interrupting Comparison :-This cranidium resembles those the above ridge. Fine linear ornaments are seen of Planiscu tellum and Kosouopeltis, but the on the glabella, particularly well in its anterior essential distinction from them is the absence part. of the preglabellar depression. Planiscutellum Occurrence:-Gomi quarry, Yokokura-yama; has the large concave preglabellar depression collected by Kojima. in front of the glabella. The depression is very narrow in Kosouope/tis, but it separates the Japonoscutellum japonicum (Kobayashi glabella from the subvertical doublure clearly. and Hamada) laticephalum, subsp. nov. This cranidium reveals the characteristics of Japonoscutellum in the glabellar pattern, but it PI. 90, Figs. 2a-b. disagrees with J. japonicum s. str. in the lateral Description :-Cranidium broader than long, expansion of the cranidium in anterior, eyes most expanded laterally in anterior, gently located more posteriorly and possession of inflated and becoming subvertical along frontal distinct eye-ridges and smaller alae. The linear margin with forward convexity; sagittal profile ornamentation is well developed in this sub­ nearly straight except for occipital furrow and species. anterior bent of frontal lobe. Glabella fungiform, Occurrence :-The Yokokura-yama limestone; most expanded at frontal lobe, strongly con­ collected by Noda. tracted at posterior lateral furrow and as wide as occipital ring at middle lateral furrow; pos­ (?) Subfamily Scutelluinae R. and terior furrow expanded adaxially, separating E. Richter, 1956 posterior and middle lateral lobes of subequal Scutelloid, gen. et sp. indet. length; middle furrow longer and anterior furrow PI. 90, Fig. 3. longest; both linear; anterior lateral lobe shorter than middle or posterior lobe; these three pairs A small dorsal shield, about 14 mm long. of lateral furrows all interrupted by axial zone Thorax is composed of ten segments, most which is a little narrower than posterior furrow; expanded laterally at about posterior one-third, un furrowed axial zone becoming a little wider axial lobe broader than pleural lobe in anterior, between anterior pair of furrows; occipital but regularly narrows back as far as the axial furrow deeper than these lateral furrows; occipi­ and pleural lobes become subequal in breadth. tal ring somewhat shorter than posterior lateral Pleural furrows are absent. lobe on lateral side; frontal lobe arching down Pygidium is suboval, wider than long and in anterior and becoming subvertical; axial moderately convex, convexity being emphasized furrow deep. Fixed cheek narrow and depressed; in one-third near anterior margin; axial lobe eyes of medium size opposed at posterior lateral sm~l subtriangular and well inflated; its median lobes; eye-ridge extending from middle lateral part smooth, parallel-sided and slightly elevated 450 Teiichi KOBA YASHI and Takashi HAMADA above lateral parts. Pleural ribs seven on each wavy backward; many tubercles spread in whole side; median rib a little broader than others; glabella, mostly in interspace between ridges, all of them flat·topped and separated from one but ridges are absent in posterior part; median another by narrow furrows; doublure about tubercle at two-thirds from front very large; a half as wide as pleural lobe. pair of tubercles on its postero-Iateral sides, The pygidium disagrees with other pygidia second largest; axial furrow deep; basal lateral from the Yokokura-yama limestone in the lobes subtriangular; eyes of medium size opposed outline and convexity of the pygidium and the in posterior. flat ribs and narrow furrows. The median rib ObselVation and comparison:-This species is may be simple. Because the cephalon is ill­ represented by an imperfect cranidium whose preserved, its generic reference cannot be made. fixed cheeks and neck ring are largely gone, Occurrence:-Gomi quarry, Yokokura·yama; but a semicircular scar of the right eye-band collected by Kojima. suggests that the fixed cheek is very narrow. It is certain that this cranidium belongs to Subfamily Meroperixinae Kobayashi the Scutelluidae, but it does not fit in any and Hamada, 1974 generic diagnosis so far known. The nearest to it is Opoa whose glabella is, however, simply Ligiscus Lane and Owen, 1982, whose type­ parallel-sided in posterior, but it is truncate­ species is L. arcanus Lane and Owen, 1982 from conical and provided with a pair of the basal the Silurian near the Llandovery-Wenlock bound­ lateral lobes in this species. The test is granulate ary, in western North Greenland, is an effaced as in Opoa, but the interspace reveals a honey­ scutelluid probably referable to the Mero­ comb structure in Opoa, while it is ridged in the perixinae. In this genus the occipital ring is anterior side and three large tubercles are present ill-defined, but a median tubercle is present in in the posterior part of the glabella in this the palce. species. Therefore, it is probable that this species Genus Opoa Lane, 1972 will be found to be a distinct genus in future. Among the pygidia referred to Microscutel­ Opoa (?) trinodosa Kobayashi and Hamada, lum primigenium there are two kinds. With this sp. nov. find it becomes probable that the strongly PI. 90, Figs. 4a--c. granulate pygidium having geniculate pleural area like Opoa damesi Lane to which the authors 1974. Microscutellum primigenium Kobayashi have compared belongs probably to this species. and Hamada (pars), p. 74, pI. 5, figs. 9a--c Occurrence :-Yokokura-yama limestone, at only Gomi quarry, collected by Asami. Descrip tion :-Glabella three-fourths as long as broad, semi elliptical transversally expanded in Family Illaenidae Hawle and Corda, 1874 anterior one-third of sagittal length and conical and tapering back as far as its half breadth in Raymond's bipartition of the Illaenidae into the remainder, gently ascending to one-third of the Illaeninae and Bumastinae (1916) has been sagittal profile from posterior margin where a widely accepted by Hupe (1955), Balashova large median tubercle exists; lateral slope steep (1975) and others. In Moore's Treatise (1959) in conical part, whereas it is gentle in expanded Jaanusson discriminated the Ectillaeninae in the anterior; frontal rim narrow with terrace lines family and provisionally added the Theamata­ separated from glabella by narrow marginal spidae Hupe, 1953 to it as a subfamily. Later furrow; irregular linear ridges distributed in Bruton (1970) erected the Panderininae for anterior and middle parts of glabella which are Panderia. In additing the Goldillaeninae Bala­ subparallel to frontal margin, but they become shova, the authors (1974) classified the family 819. Silurian Trilobites from Shikoku, 2nd Addition 451 into the Illaeninae, Panderininae, Ectillaeninae, the contrary, the occipital furrow is completely Buamastinae, Goldillaeninae and (?) Theama­ effaced, at least in the grown stage, although it spidinae beside three genera whose subfamily may remain in the immature stage as seen in reference is uncertain. Faillaena clava Chatters ton and Ludvigsen, Balashova (1960) instituted the Goldillaeninae 1976, in fig. 34, pI. 6. Among the Illaenidae in the Scutelluidae to include Goldillaenus and Bumastus (Middle Ordovician-Silurian) reveals Goldillaenoides the latter of which had the particularly high specialization in the very occipital ring. While Jaanusson (1979) accepted broad glabella and almost complete oblitera­ Goldlllaenus as a subgenus of Illaenoides Weller, tion of the trilobation in the thorax and pyg­ 1908, Lane (1972) brought back the Goldil­ idium before the appearance of the Scutelluidae. laeninae to the Scutelluidae. Distribution :-In the Ordovician fauna of Chatterton and Ludvigsen (1976) instituted China this family is well represented by eight a new genus, Failleana with Failleana clava as or nine genera in four or five subfamilies as its type species and suggested that the Bumasti­ follows: nae be placed in synonymy with the Illaeninae , Stenopareia, Nanillaenus, Cekovia, as they found the rostral flange in Bumastus Trigocekovia and Spinillaenus in Illaeninae (Bumastoides) aplatus. Lane and Thomas (1978) Zdicella and Zbirovia in Ectillaeninae on the other hand referred Bumastus to the Bumastus and Dysplanus in Bumastinae Scutelluidae and added three new Bumastus­ Meitanillaenus in Goldillaeninae like Silurian genera, i.e. Cybantyx, Litotix and (?) Theamatopsis in Theamatopsinae Rhax the last of which is based on Rhax pol­ Trigocekovia Xia, 1978 whose monotypic lincitrix Lane and Thomas from the Llandovery­ species is T. fonicatus Xia, 1978, resembles Wenlock of Queensland, Australia. Thus the Cekovia, but its cephalon is triangular in dorsal taxonomy of the family Illaenidae and its rela­ view and hem~sphaeric in frontal view. Spinil­ tion to the Scutelluidae is now a moot question. laenus Xia, 1978, of which S. sinensis Xia, 1978 The Typhlokorynetidae Shaw, 1966 and the is monotypic is similar to Illaenus and OcW­ Portaginidae Lesperance, 1968 are represented laenus, particularly to the latter, but it has the respectively by Typhlokorynetes Shaw, 1966 concave marginal border which is produced and Portagius Lesperance, 1968. If they are postero-Iaterally into a triangular genal spine as included in the Illaenacea, the superfamily is far as the third thoracic segment. The thorax certainly comprehensive and much remains to consists of 10 segments, instead of 8 in OcW­ be solved particularly of the effaced illaenoids. laenus. Meitanillaenus which appeared first in The Illaenidae appeared already in the early Hunan in the late Ordovician age survived in Ordovician and died out before the Devonian the Silurian period when Ptilillaenus and period. The Scutelluidae on the contrary first Wuchuanella also occur. appear in the Middle Ordovician rocks in China In Japan the family is represented by many and the wide divergence has taken place from Silurian species in two subgenera, Bumastus the Planiscutelluinae during the late Silurian and Bumastella of Bumastus beside a species and Devonian periods. The trend of evolution of Goldillaenus and two species of "Illaenoides" was quite different between the two families. which belong probably to a new genus. It is improbable that the Scutelluidae were derived from the effaced Illaenidae or vice Subfamily Goldillaeninae Balashova, 1959 versa. In the Scutelluidae the effacement of the cephalic segmentation for example is most In the Silurian fauna of China the family advanced in the Meroperixinae (Upper and Illaenidae is represented by the following three Middle Silurian), but still the occipital lobe is endemic genera whose type-species are cited in generally well defined. In the Illaenidae, on brackets. 452 Teiichi KOBA YASHI and Takashi HAMADA

Ptilillaenus Lu, 1962 (Ptilillaenus lojoping­ Ordovician of Northeast USSR has the pygidium ensis Lu, 1962) with axial lobe composed of tre trapezoidal Meitanillaenus Chang, 1974 (Meitanillaenus short anterior part and the longer posterior binodosus Chang, 1974) part but ill-defined near the terminal part. Wuchuanella Wu, 1978 (Wuchuanella quad­ rata Wu, 1978) Genus Illaenoides Weller, 1907 Ptilillaenus lojopingensis was primarily found­ Illaenoides (?) magnisulcatus Kobayashi and ed on cranidium and pygidium. A free cheek Hamada, sp. nov. from Lojoping- which Chang (1974, pI. 18, fig. 8) PI. 90, Figs. 5a-b. referred to this species has the lateral border which is rounded at the genal angle and becomes Description:-Illaenoid pygidium gently con­ broader near the angle. A shallow marginal vex except for marginal inclination, semielliptical furrow runs along the border. The axis of the in outline, widest a little behind anterior margin associate pygidium is long, conical and composed where axis is a little projected beyond pleural of 7-8 rings and a round terminal lobe. The lobes; anterior margin forming a broad obtuse marginal border is nearly flat and moderate in angle near median point of pleural part; con­ breadth. vexity of pygidium reaching maximum at about Meitanillaenus binodosus from Meitan, Gui­ one-third from posterior end. Axial furrow com­ zhou has the cranidium nearly as long as wide pletely effaced; axial projection of anterior mar­ and strongly contracted at about one-fourth gin one-third as broad as pygidium. First pleural from posterior, much more expanded in anterior furrow extending postero-laterally from lateral than posterior where it is about half as broad as end of axial projection and confluent with mar­ the cranidium; a pair of semicircular nodes ginal furrow. Lateral border relatively thick for present on fixed cheeks at glabellar constriction; illaenid. Facetted area defined by obtuse ridge eyes there opposed, fairly large; free cheek with extending postero-laterally from median angula­ narrow lateral border and short genal spine. tion of anterior pleural margin; lateral marginal Like Bumastus hypostoma is provided with a furrow tending to shallow backward. Sagittal pair of large triangular wings; central body sub­ profile of pygidium in exfoliated part simply circular and divided into two parts by middle arching down to posterior margin from highest furrow and maculae. Thorax distinctly trilobed point. Test smooth. by axial furrow; axial ring slightly broader than Observation:-The pygidium is 8.5 mm long pleuron. Pygidium semicircular; axis short and and 11 mm wide. The axial projection of the trigonal; articulating facet distinctly limited by anterior margin measures 5.8 mm in breadth pleural furrow. and the lateral border exclusive of marginal Meitanillaenus zhitangensis Wu, 1983 from furrow 1.8 mm. the Upper Ordovician of Xijiang which has a Comparison:-The convex marginal border distinct occipital furrow and lobe may be ex­ and well developed marginal furrow are most cluded from this genus. Wuchuanella quadrata distinctive of this pygidium, suggesting that Wu is represented by two cranidia resembling this pygidium belongs to an undescribed genus. Meitanillaenus, but lacking alae. Among the Japanese trilobites it is similar to Finally, the Brontocephalidae Kolobova, Bumastus (Bu mas tella) bipuncatus, but the 1973, including Brontocephalus Tschugaeva, lateral margins converge more rapidly and the 1975, and Brontocephalina Tschugaeva, 1975, marginal border is depressed. It resembles also but excluding Bulanaspis Tschugaeva, 1956 is the pygidium of Ptilillaenus lojopingensis Lu in allied to the Goldillaeninae. Its glabella is strong­ the presence of marginal border which is, how­ ly expanded in anterior. Brontocephalina mar­ ever, not convex and the marginal furrow is ginatula Tschugaeva, 1975, from the Upper absent in that species. 819. Silurian Trilobites from Shikoku, 2nd Addition 453

Occurrence:-Yokokura-yama limestone; col­ composed of large median lobe and a pair of very lected by Fujiyama. small lateral lobes; occipital and axial furrows profound; eye medium sized, semicircular close­ Illaenoides (?) abnormis Kobayashi and set glabella behind anterior lateral furrow and Hamada, sp. nov. provided with distinct eye-socle; palpebral lobe PI. 90, Figs. 6a-c. flat and simply slanting to axial furrow; pre­ glabellar field short, arching down to concave Description :-Pygidium parabolic in outline, frontal furrow which is in turn bent up toward broader than long in ratio of 1:0.7, regularly frontal edge; facial sutures widely divergent and gently convex with narrow marginal border diagonally from eyes with postero·lateral con­ and narrow marginal furrow; posterior one-third vexity, incurved through margina,l furrow and of pygidium distinctly arching down below the rim a little beyond eyes' limit; posterior branch general base level. running backward along axial furrow, then be­ Observation and comparison:-The margin of coming diagonal to cut the cheek border on this pygidium is not perfect, but if perfect, it is parallels through eyes; narrow ridge running probably 19.3 mm long and 27 mm broad. The along posterior facial suture till it becomes axial part is about 13 mm wide on the articulat­ confluent with posterior cheek border; test ing margin. The lateral border is 3 mm across. densely granulose. Like the preceding pygidium the axial furrow Pygidium two·thirds as long as broad; an­ is completely effaced in this pygidium, but the terior pleural margin obtusely angulated at marginal border is evidently narrower and not median point; postero-lateral margin broadly so convex as in that pygidium. This pygidium arcuate; posterior margin nearly transversal. is more convex and distinctly bent down in Axial lobe one-third as wide as pygidium, well posterior one-third. Because it looks the original convex, rapidly tapering back to rounded pos­ curvature, it is quite an unique aspect among terior end whence a short postaxial ridge issues; illaenids. six axial rings separated from one another by Occurrence:-Yokokura-yama limestone; col­ deep ring furrrows. Pleural lobe divided into lected by Mizuno. five flat-topped ribs, all geniculate, less slanting in inner part than outer part; first rib distinctly Family Poetidae Salter, 1864 facetted; succeeding ribs forked by narrow Subfamily Cyphoproetinae Pillet, 1969 interpleural furrows which are deepened in distal part. Marginal border flat and smooth; Genus Cyphoproetus Kegel, 1927 its inner side depressed where pleural and inter­ Cyphoproetus latiaxis Kobayashi and pleural furrows die out. Test granulose. Hamada, sp. nov. Observation:-Because the occipital lobe is PI. 91, Figs. 1a-g, 2a-c. mesially exfoliated, it is indeterminable whether or not a median tubercle is present. The test is Description:-Cephalon semicircular in out­ broken also near the genal angles. Therefore the line, well convex except for distinctly concave weak incurvature of the cephalic margin, com­ and smooth marginal border; glabella half as wide monly seen near the angle in Cyphoproetus as cephalon, subconical, but much more ex­ cannot be ascertained in this specimen. Note· panded in posterior and broadly arcuate in front; worthy is that a round-topped narrow ridge anterior lateral furrow (2p) weak and short; pos­ resembling the so-called eye·socle by Owens, terior lateral lobe isolated by strong lateral 1963 is running in this cephal on along the pos­ furrow (lp), which is long and straight, but terior branch of the facial suture as far as the postero-laterally bent near its posterior end and posterior cheek border. On the left cheek a disconnected from occipital furrow; neck ring narrow ridge is clearly seen behind the eye. 454 Teiichi KOBA YASHI and Takashi HAMADA

The associate pygidium has a flat marginal cently Sphaerexochus fimbrisulcatus Lu, 1975, border of moderate breadth, but no distinct and S. granosus Ju, 1983, were described from marginal furrow on its inner side where pleural the Middle Ordovician in East China, S. dong­ ribs and furrows terminate. The pleural ribs are zhuangensis Zhou, 1982, from Middle Ordovician distinctly flat-topped in the inner part. in Shaan-Xi, Northwest China, S. luoheensis Comparison:-This cranidium belongs to the Nan, 1980, from Upper Ordovician, Heilongjiang, group of Cyphoproetus with a narrow pre­ Northeast China, and S. guizhouensis Wu, 1977, glabellar area and granulose test like Wenlockian from Lower Silurian, Guizhou, Southwest China. C. strabismus Owen, 1963, but the granula­ Sphaerexochus tisulcatus Tschugaeva, 1973 tion is coarser and the cranidium and glabella occurs in Lower Ordovician of Northeast USSR. are broader and the eyes smaller in this species. Further additions were made from Canada Occurrence:-Yokokura-yama limestone at and Australia as follows: Gomi quarry; collected by Uchida. Sphaerexochus arenosus Chatterton & Lud­ vigsen, 1976, from Middle Ordovician, Family Cheiruridae Angelin, 1854 Canada Subfamily Sphaerexochinae tJpik, 1937 Sphaerexochus dimorphus Perry & Chatter­ ton, 1977, from Middle Silurian, Canada Beside Sphaerexochus and Pompeckia Whit­ Sphaerexochus molongoensis Chatterton & tington (1963, 1965), this subfamily included Campbell, 1980, from Middle Silurian, Heliomera Raymond, 1905, Kawina Barton, Australia 1915, Cydonocephalus Whittington, 1963, and Sphaerexochus lorum Chatterton & Campbell, Xystocrania Whittington, 1965. Kolymella and 1980, from Upper Silurian, Australia Parasphaerexochus are two additional genera Sphaerexochus glaber Holloway, 1980, from which were instituted by Tschugaeva, 1973, Middle Silurian, U.S.A. Kawina plana Tschugaeva, 1964, and Para­ Sphaerexochus brundlyi Chatterton, 1984, from Middle Silurian, Canada sphaerexochus galeatus Tschugaeva, 1973, being Sphaerexochus johstoni Chatterton & Perry, the respective type-species. The former occurs 1984, from Middle Silurian, Canada in Middle Ordovician and the latter in late Lower Ordovician, both in Eastern Siberia, These species are represented by many ex­ Northeast USSR. cellent specimens, some of which are silicified and show their ontogeny. Thus our concept of Genus Sphaerexochus Beyrich, 1845 the genus was greatly clarified. Finally, supplemented with Sphaerexochus This is an Ordovician-Silurian genus of Trilo­ seto Lane and Owens, 1982, from the Silurian bita whose type-species is Sphaerexochus mirus (up. Llandovery-low. Wenlock) from Greenland Beyrich, 1845. According to Lane (1971) it it is nearly cosmopolitan, although none is re­ comprises 23 other species including one doubt­ ported from South America, so far as the authors ful reference. Most of them are European species, are aware. but a quarter is American and one, namely, Sphaerexochus taimyricus Balashova, 1960, is Sphaerexochus hiratai Kobayashi and an Asian species. S. mirus is reported from Hamada, 1974 Turkestan by Weber (pygidium, 1932, cranidium and pygidium, 1951) and also from Australia PI. 90, Figs. 7a-c. and North America. The authors added one 1974. Sphaerexochus hiratai Kobayashi and Silurian (up. Llandoverian-low. Wenlockian) Hamada, p. 88, pI. 6, figs. 6-10, pI. 7, species from Malaysia (1971) and two Ludo­ figs. 1-8, pI. 8, figs. 1-7, text-fig. 6F. lovian species from Japan (1974, 1976). Re- 819. Silurian Trilobites from Shikoku, 2nd Addition 455

This species is well represented by many segments in its thorax and pygidium than Din­ cranidia and pygidia, but its free cheek, hy· dymene. Furthermore, the difference in the postoma and thorax were unknown. In a small ventral facial suture between Dindymene and complete rolled shield from the same locality Plasiaspis is emphasized by Kielan. the pygidium is seen to fit nicely the broad Incidentally, Dindymenella Lu, 1976, insti­ sinuate frontal margin of the cranidium. tuted on Dindymenella sulcata Lu, 1976, from Its thorax is tapering back slowly and com­ the early Upper Ordovician in Yunnan differs posed of ten segments. The axial ring is nearly from Dindymene as well as Eodindymene in the as wide as the pleuron which is unfurrowed possession of 12 segments in thorax, instead and abruptly rounded near lateral extremity. of 10 segments in those two genera and the The anterior segments whose pleurae are more axial lobe of the pygidium is comparatively or less geniculate are more convex than the small and its spines are fairly stout and long. posterior ones. Two small round bosses are The distinct lateral furrows in three pairs on its seen on the left side of the sixth and seventh glabella are quite distinctive from those two axial rings at their antero-lateral ends, instead genera. of the postern-lateral ends. The free cheek has a narrow rim and a broad Genus Dindymene Hawle and Corda, 1847 concave border outside the relatively small cheek-roll. The anterior branch of the facial Recently Dindymene orientalis Zhou and suture runs forward and a little inward from D. deflecta Ju were described from the Upper the eye. Its posterior branch in form of a suture Ordovician, respectively in Jiangxi and Xijiang, ridge extends from the eye to the genal angle both Central China (Chou et al., 1983). Dindy­ taking a broad arcuate course and meets the mene brevicaudatus Kolobova, 1972, is known lateral margin just in front of the angle where from the Upper Ordovician in Kazakhstan. a very short rudimentary spine issues. Occurrence:-Yokokura-yama limestone; col­ Dindymene (?) sp. indet. lected by Kojima. PI. 91, Fig. 4.

Subfamily Dindymeninae Kielan, 1959 A blind cephal on composed of large vaulted non Henningsmoen, 1959 glabella and less convex cheeks of moderate size. Glabella suboval, broader in anterior, grad­ Family Dindymeninae was proposed in 1959 ually narrowing in middle and posterior parts; by Kielan to include Dindymene Hawle and posterior margin transversal and nearly straight; Corda, 1847 and a new genus, Eodindymene lateral furrows absent; a large median tubercle whose type-species is Dindymene pulchra Olin, present at a point slightly anterior to center of 1906, but she excluded Plasiaspis Prantl and glabella. Cheeks separated from glabella by pHbyl, 1948. In the same year Henningsmoen narrow straight axial furrows and opposed at proposed also the Dindymeninae as a subfamily posterior half of glabella. Marginal furrow and of the Encrinuridae, but he included Plasiaspis rim very narrow. Small tubercles densely dis­ and possibly Prosopiscus Salter, 1865 beside tributed on whole surface except for axial and Dindymene in his subfamily. Because only the marginal furrows where they are absent or year, 1959, is cited for their dates of publica­ sparce; medium sized tubercles scattered irreg­ tion, it is a question which of the family or ularly among small ones. subfamily possesses the priority. The present The genal and posterior parts of the cephalon authors, however, consider that Plasiaspis and are obscure, but the antero-lateral outline can be Prosophiscus are to be eliminated out of the figured out by tracing the tuberculate marginal subfamily, because these two genera have well rim and smooth marginal furrow. Little is known segmented glabellae and Plasiaspis has more of facial suture. 456 Teiichi KOBA YASHI and Takashi HAMADA

This cephalon belongs most probably to A Dindymene or the Dindymeninae s. str., but it is quite distinct from most species of the genus in its proportionally large glabella to the cheeks and accordingly the much greater forward projection of the glabella. The antero· lateral outline of the cheek is also different from typical Dindymene. These differences suggest its subgeneric or generic distinction from Upper 2 Ordovician Dindymene in Europe. Occurrence:-Gomi quarry of the Yokokura· Text-figs. A-C. Restoration of Three yama limestone; collected by Fujiyama. Silurian Trilobites from Mt. Yokokura. (vide p. 457) Genus IchiyameZla Kobayashi and Hamada, Al-4: Ichiyamella subg/obu/a. gen. nov. toward genal angle where it is remarkably hang­ Diagnosis:-Cephalon strongly vaulted, more ing down below general level of cephalon; eyes or less hemisphaeric and divided into a large and facial suture absent; test densely granulate. glabella and two small cheeks by gently arcuate Observation :-The holotype cephalon is 14.7 or nearly straight and forwardly divergent axial mm long, 16 mm broad and 7.5 mm high; its furrows; no lateral furrows on glabella, but a glabella 11.5 mm long, 13 mm at the widest pair of short linear furrows cutting into glabella and 4.7 mm at the base; occipital ring 3 mm from frontal margin; occipital ring relatively long and 7.5 mm wide; and its cheek 4.6 mm large; cheek lacking eyes and facial stures. wide, 5 mm high and 7.3 mm in diagonal length. Type-species:-Ichiyamella subglobula Koba· Because the diagonal furrows are confluent yashi and Hamada, sp. nov. with the occipital furrow far inside of the lateral Remarks:-The authors are of opinion that extremities of the neck ring, they look similar Dyndymene (?) megacranidia is referable to this to the preoccipital furrows of Sphaerexochus. genus. No median tubercle is present on the In this cephalon, however, there are neither glabella in these two species. axial furrows nor cheeks, if they be really the Distribution :-Silurian in Japan. posterior lateral glabellar furrows. Therefore these diagonal furrows are evidently the axial Ichiyamella subglobula Kobayashi and furrows. Hamada, sp. nov. Occurrence :-Ichiyama, near Ochi-town; col­ lected by Nakajima. PI. 91, Figs. 3a-f; Text-figs. AI-4.

Description :-Cephalon strongly vaulted, a Ichiyamella megacranidia Kobayashi and little broader than long. Glabella very large, Hamada, 1985 strongly expanded forward from narrow base 1985. Dindymene (?) megacranidia Kobayashi and well rounded in anterior; lateral furrows and Hamada, p. 213, pI. 28, figs. 3a--c. absent, but a pair of weak short linear furrows inserted from frontal margin in two lateral This species agrees with the preceding in the parts of glabella; occipital ring much wider trilobation of the strongly convex cephalon, a than glabellar base and abruptly narrowing pair of short linear frontal furrows on the glabel­ near lateral ends; axial furrow diagonal and la, small cheeks and granulate test, but it is dif­ gently arcuate. Cheek ovoid, very small in com­ ferent from that species in the roundly quadrate parison with glabella; marginal border thickened glabella limited laterally by nearly straight axial 819. Silurian Trilobites from Shikoku, 2nd Addition 457

furrows. A very narrow marginal rim and a Tsungilichas Chang, 1974 from the Lower little broader and deep intramarginai furrow Silurian of Guizhou, China was synonymized are also present in this species. The posterior with Dicranopeltis by Thomas (1978). Recently mrgin of this cephalon is unknown. Dicranopeltis cf. polytoma (Angelin) was re­ Occurrence :-Gomi quarry of the Yokokura­ ported by Zhou and Zhou (1982) to occur in yama limestone. the Upper Ordovician of Inner Mongolia.

Family Lichidae Hawle and Corda, 1874

In the Ordovician fauna of Asia this family is represented by Metopolichas, Amphilichas, Acro­ lichas (?) and Lyralichas in China, Indochina, Burma and Turkestan. Acanthopyge, Lobopyge, Graspedarges and Lichas s.l. are reported from the Devonian of Western, Central and Eastern Asia where in the last Acanthopyge is known from the Amur region and Radiolichas (?) from Mongolia, Lichas, s.l. from Viet-Nam. Gras­ pedarges and Acanthopyge (Lobopyge ?) were found in Japan. Compared to them, the occur­ rences of Silurian lichids are scarce in Asia. Lichas (Gorydocephalus) in two species are described from the Middle Silurian of Western Mongolia by Tschernysheva (1937) and Acantho­ pyge (3 spp.), Acanth?ma, Tetralichas and 2 Discrnopeltis (?) from the Kuznetsk region and BI-3: Apolichas trucatus. Kazakhstan, Turkestan by Weber (1932, 1951), Tchernysheva (1951), Maximova (1960) and Balashova (1968). Gorydocephalus is now synonymized with Trochurus.

Subfamily Lichinae Hawle and Corda, 1874 Genus Dicran op Ie tis Hawle and Corda, 1974

Phleger (1936) erected two new genera, Dicranopletoides and Makromuktis on Lichas decipiens Winchell and Marcey, 1865 and Dicranopeltis nasutus Weller, 1907, respectively, but they were synonymized with Dicranopeltis by Warburg (1939) and this opinion was upheld by Tripp (1957 and in Moore's Treatise, 1959). Makromuktis is Dicranopeltis having a median anterior projection on the glabella. It is note­ worthy that such forms occur not only in the Niagaran in North America but in the Wen­ lockian in Britain and also in the Yokokura­ yama limestone in Japan. CI-3: Dicranopeltis tricornis. 458 Teiichi KOBA Y ASH! and Takashi HAMADA

Dicranopeltis tricornis Kobayashi and toward mid-length of lateral glabellar lobes; Hamada, sp. nov. front-median lobe composed of laterally ex­ PI. 92, Figs. la-g; Text-figs. Cl-3. panded frontal part, long conical middle part narrowing back, and small posterior part ex­ Description :-Cephalon large, broad, pentag­ panding toward neck ring; lateral lobes quite onal, well vaulted, most elevated at a little an­ different in size; anterior lateral lobe long and terior to glabellar center whence a stout median occupying major part; posterior lateral lobe spine issues, subvertical anterior to spine, but small but a little larger than basal lobe; dorsal gently sloping down backward; its length about and occipital furrows stronger than lateral ones. five-eights of cephalic breadth exclusive of Fixed and free cheeks well fused; eyes small, genal spines; anterior margin nearly straight, located a little anterior to cheek-center; no eye­ diagonal and forming a broad obtuse blunt ridge discernible; suture ridge extending laterally angle of about 110 degrees, slightly more pro­ from eye, describing an arc on cheek, then truded forward in median part than antero­ facial suture cutting diagonally through lateral lateral parts; posterior margin almost as broad border as far as near intergenal angle; anterior as anterior one, transversal, but more extended branch of facial suture apparently running back at neck ring and forming obtuse angles at forward and inward from eye and crossing lateral ends; antero-lateral and postero-lateral doublure between frontal and antero-lateral margins meeting at genal angle whence a stout parts of marginal border. Genal spine stout, long lateral spine issues. protruding in same line with antero-lateral Glabella large, occupying more than a half of border and then slightly bent backward; pos­ cephalon, broadest through a point a little pos­ terior cheek border divided into two parts by terior to median spine and broadly sinuate intergenal angle at its median point.

Explanation of Plate 90

Figs. la-c. Japonoscutellum japonicum (Kobayashi and Hamada) puteatum, subsp. nov ...... Page 448 a: dorsal, b: frontal and c: left lateral views of the holotype cranidium. PA 18071, x2.0, colI. Kojima. Figs. 2a--c. Japonoscutellum japonicum (Kobayashi and Hamada) laticephalum, subsp. nov ...... Page 449 a: dorsal, b: frontal and c: right lateral views of the holotype cranidium. PA 18072, x2.8, colI. Noda. Fig. 3. Scutelloid, gen. et sp. indet...... Page 449 A dorsal view of an almost complete shield with 10 thoracic segments, though they were exfoliated. PA 18073, x3.4, colI. Kojima. Figs. 4a-c. Opoa (?) trinodosa Kobayashi and Hamada, sp. nov...... Page 450 a: dorsal, b: frontal and c: right lateral views of the holotype cranisium. PA 18074, x1.8, colI. Fujiyama. Figs. 5a, b Illaenoides (?) magnisulcatus Kobayashi and Hamada, sp. nov...... Page 452 a: dorsal and b: left lateral views of the holotype pygidium, PA 18075, x3.2, colI. Fujiyama. Figs. 6a--c. Illaenoids (?) abnormis Kobayashi and Hamada, sp. nov...... Page 453 a: dorsal, b: posterior and c: left lateral views of the holotype pygidium showing an abnormal slant of the margin, PA 18076, x1.4, coli. Mizuno. Figs. 7a-c. Sphaerexochus hiratai Kobayashi and Hamada ...... Page 454 a: left lateral, b: pygidial and c: cephalic dorsal views of an enrolled specimen of small size, PA 18077, x4.1, colI. Kojima. KOBA YASHI and HAMADA: Silurian Trilobites from Shilwku Plate 90 819. Silurian Trilobites from Shikoku, 2nd Addition 459

Fine granules distributed on test from glabella 1907), although it is not a proboscis-like projec­ to convex marginal border and three spines, tion in this species. Dicranopeltis woodwardi excluding various smooth furrows; smooth (Reed) and Dicranopeltis sp. I by Thomas (1981) interspace, however, wider than granules in has an upwardly directed spine on the median total space. glabellar lobe which is bifurcated at a distance Observation :-This species is represented by a from its root. Lichas (Metopolichas) celorhin half of a cephalon which is, however, scarcely Angelin var. coniceps Herz von Leuchtenberg by deformed. If complete, the cephalon exclusive Reed 1917 form the Ordovician of Shihtien, of three spines is about 26 mm long, 40 mm Yunnan also has a median spine on the frontal broad and 16 mm high. lobe of the glabella. These spines are all broken, but they are Occurrence:-Yokokura-yama limestone; col­ presumably fairly long. The left eye is indicated lected by Asami. by an eye-scar whose outline is obscure on the axial side. Its lateral and posterior margins are, Subfamily Homolichinae Phleger, 1936 however, traceable along the peripheral narrow Genus Apolichas Kobayashi and Hamada, 1974 furrow of the eye-socle and also with reference Apolichas truncatus Kobayashi and Hamada, to the suture ridge behind the eye. The posterior 1974 branch of the facial suture is seen in the further extension, while its anterior branch is recog­ PI. 92, Figs. 2a-c; Text-figs. Bl-3. nizable only on the doublure and its vicinity. 1974. Apolichas truncatus Kobayashi and Comparison:-This species agrees with Lichas Hamada, p. 80, pI. 8, figs. 9-12; text-fig. scabra scabra (Beyrich, 1845), the type-species 6A. of Dicranopletis Hawle and Corda, 1947 in the broad pentagonal outline of the cephalon with a Little has been known on the cheek of this pair of lateral spines and the essential configura­ species. A new cephalon shows that a left cheek tion of the cranidium, although the glabella of is separated from the tricomposite lateral lobe this species disagrees with the lectotype of of the glabella by a deep axial furrow and over­ Dicranopeltis scabra in fig. 1, pI. 4 and Dicrano­ hanging the narrow marginal border. It is reni­ pettis scabra propiqua (Barrande, 1846) restored form and strongly vaulted, but its top is lower in text-fig. 13, both in Vanek (1959) in the than the lateral lobe. Its eye is very large for outline of the median lobe, the proportional the Lichidae. length of bicomposite lobe with the posterior Compared to the type specimen this cephalon lobe and other minor details. The most impor­ is somewhat different in the anterior outline. tant distinction of this species from Dicranopeltis The main lobe of its glabella is protruded for­ scabra is the possession of the median glabellar ward, forming weak concavity on each side of spine. the frontal margin of the cephalon. Further· Dicranopeltis polytomus (Angelin, 1854) with more the convexity of the glabella is a little which Lichas elegans Tornquist, 1884 was syno­ stronger, attaining the maximum shortly behind nymized by Warburg (1939) has also a similar its center. The posterior margin of the cephalon cranidium which, however, lacks the median is not well preserved and the genal spine which spine. Tsungilichas pustulosus Chang, 1974, existed on the left cheek of the type specimen from the Lower Silurian of Guizhou is repre­ is lost in this cephalon. On the basis of these sented by a strongly vaulted cranidium resembl­ two cephala combined the dorsal aspect of the ing this species, but it has no spine on the some­ cephalon can now be restored completely. what longer glabella. Occurrence :--Yokokura-yama limestone. KAC­ In regard of the median projection this species St-0032(a) ; collected by Kaneko. is comparable to Makromuktis nasuta (Weller, 460 Teiichi KOBA YASHI and Takashi HAMADA

References from Arkansas and Oklahoma U.S.A. -Part 1. Palaeontogr. (A), Vol. 170, p. 1-85, 20 pIs. Bergstrom, J. 1973, Organization, life and sys­ Kielan, Z. 1959, Upper Ordovician trilobites tematics of trilobites. Fossils & Strata, no. 2, from Poland and some related forms from 69 pp., pIs. 1-5. Bohemia and Scandinavia. Pal. Polonica, no. Chang, W. T. 1974, Silurian trilobites, p. 173- 11,198 pp., 36 pIs. 187. In Handbook of stratigraphy and Kobayashi, T. and Hamada, T. 1965, An occur­ palaeontology in Southwest China. Nanjing rence of a new Scutellum in the Silurian of Inst. Geol. Palaeont., Acad. Sinica, 454 pp., Shikoku, .Japan. Trans. Proc. Pal. Soc. Japan, 202 pIs. N. S., no. 58, p. 74-81, pI. 7. Chatterton, B. D. E. 1980, Ontogenetic studies -- and -- , 1971, Silurian trilobites from the of Middle Ordovician trilobites from the Langkawi Islands, Northwest Malay, with Eabataot-Tine Formation, Mackenzie Moun­ notes on the Dalmanitidae and Raphiopho­ tains, Canada, Palaeotogr. (A), Vol. 171, ridae. Geol. Pal. SE Asia, Vol. 9, p. 87-134, p. 75-103, 10 pIs. pIs. 18-23. --and Campbell, K. S. W. 1980, Silurian trilo­ -- and --, 1974, Silurian trilobites of Japan bites from near Canberra and some related in comparison with Asian, Pacific and other forms from the Yasa basin. Ibid., Vol. 167, faunas. Palaeont. Soc. Japan, Spec. Pap., no. p. 77-119, 16 pIs. 18,155 pp., 12 pIs. --and Ludvigsen, R. 1976, Silicified Middle - and -- , 1976, A new Silurian Trilobite Ordovician trilobites from the south from Ofuna to, North Japan, Proc. Japan Hananni River area, district of Mackenzie, Acad., Vol. 52, no. 7, p. 367-370. Canada. Ibid., (A), Vol. 154, 1-106, pIs. --and --,1985, Additional Silurian trilobites 1-22. to the Yokokurayama fauna from Shikoku, --and Perry, D. G. 1984, Silurian cheirurid Japan. Trans. Proc. Pal. Soc. Japan, N.S., no. trilobites from the Mackenzie mountains, 139, p. 206-217, pIs. 28-30. northwestern Canada. Ibid., Vol. 184, --, Katto, J. and Hamada, T. 1984, New obser­ p.1-78, 35 pIs. vations of Shiroishi-ohsawa, Mt. Yokokura, Chou Hong-An et al. 1983, Trilobita, p. 28-254. Kochi Prefecture with palaeontological Palaeontological Atlas of East China (1),657 studies on the cephalopods and trilobites pp., 176 pIs. from the locality. Research Rep. Kochi Univ., Dean, W. T. 1978, The trilobites of the Chair of Vol. 32, Nat. Sci. p. 235-258, pIs. 1-5. Kildare Limestone (Upper Ordovician) of Kolobova, N. M. 1972, Cheiruroidae, Trilobita, eastern Ireland. Part 3, Palaeontogr. Soc. p. 242-245. pIs. 55-56. New species of Monogr., p. 99-129, pIs. 45-52. Prehistoric plants and invertebrates of the Holloway, D. J. 1980, Middle Silurian trilobites USSR, 383 pp., 75 pIs.

Explanation of Plate 91

Figs. la-g. Cyphoproetus latiaxis Kobayashi and Hamada, sp. nov...... Page 453 a, e: dorsal, b, f: frontal, c: right lateral and d, g: left lateral views of the holotype cephalon, PA 18078. a, b, c, d x4.5, e, f, g x3.0, coli. Uchida. Figs. 2a-c. Cyphoproetus latiaxis Kobayashi and Hamada, sp. nov...... Page 453 a: dorsal, b: right lateral and c: posterior views of a pygidium, PA 18079, x7.0, coll. Uchida. Figs. 3 a-f. Ichiyamella subglobula Kobayashi and Hamada, gen. et sp. nov...... Page 456 a: dorsal, b: posterior, c: left oblique, d: frontal, e: right lateral and f: left lateral views of the holotype cephalon, PA 18080, x2.0, coIl. Nakajima. Fig. 4. Dindymene (?) sp. indet...... Page 455 A fragmentary cranidium showing the granulation of various sizes, PA 18081, x3.3, coIl. Fujiyama. KOBA YASHI and HAMADA: Silurian Trilobites from Shilwku Plate 91 819. Silurian Trilobites from Shikoku, 2nd Addition 461

Lane, P. D. 1971, British Cheiruridae (Trilobita). p. 175-210, pIs. 19-28. Palaeontogr. Soc. Monogr., p. 1-95, pIs. Reed, F. R. C. 1906, The lower Palaeozoic 1-16. fossils of the northern Shan States, Burma. --1972, New trilobites from the Silurian of Palaeont. Indica. N.S., Vol. 2, no. 3, 154 pp., north-east Greenland, with a note on trilobite 8 pIs. faunas in pure limestones. Palaeontology, Shaw, A. B. 1966, Paleontology of Northwestern Vol. 15, pt. 2, p. 336-364, pIs. 59-64. Vermont, XII. Fossils from the Ordovician --and Owen, R. M. 1982, Silurian trilobites Highgate Formation. Jour. Pal., Vol. 40, no. from. Kap Schuchert, Washington Land, 6, p. 1312-1330, pIs. 161-163. western North Greenland. Rapp. Gr,pnlands Stratigraphic Research Party of the Gorge area, Geol. Unders., no. 108, p. 41-69, 5 pIs. 1978, Sinian to Permian stratigraphy and -- and Thomas, A. T. 1978, Silurian trilobites palaeontology of the eastern Gorge area. 381 from NE Queensland and the classification pp., 113 pIs. of effaced trilobites. Geol. Mag., Vol. 115, Tchernycheva, N. 1937, Silurian and Devonian no. 5, p. 351-358, 2 pIs. trilobites of Mongolia and Tuva. Trud. Lesperance, P. J. 1968, Ordovician and Silurian Mongol. Kom., no. 28, Akad. Nauk SSSR, trilobite faunas of the White Head Forma­ p. 5-32, pIs. 1-2. tion, Perce region, Quebec. Jour. Pal., Vol. Thomas, A. T. 1978, British Wenlock trilobites, 42, no. 3, p. 811-826, pI. 106. Part 1. Paeontogr. Soc. Monogr., p. 1-56, Lu Yen-hao, 1975, Ordovician trilobite faunas of pIs. 1-14 (with P. D. Lane, in part). central and south western China. Palaeontol. Tripp, R. P. 1957, The classification and evolu­ Sinica, N. S. B, no. 11, 463 pp., 50 pIs. tion of the superfamily Lichacea (Trilobita). --,1976, Trilobita typical of North and South Geol. Mag., Vol. 94, p. 104-122. China. In Lu Yen-hao et al. 1976, Ordovi­ Tschugaeva, M. N. 1973, Trilobita, p. 44-122, cian biostratigraphy and palaeogeography of pIs. 1-19. In Biostratigraphy of the lower China. Mem. Nanjing Inst. Geol. Pal., no. 7, part of the Ordovician in the Northeast of 83 pp. 14 pIs. the USSR and biogeography of the upper­ Maximova, Z. A. 1977, Devonian trilobites from most Lower Ordovician. 285 pp., 32 pIs. Novaya Zemlya and neighbouring area of --, 1975, Late Ordovician trilobites of the Soviet Arctic, p. 140-185, 5 pIs. Northeast of the USSR. Acad. Sci. USSR, --, 1977, Trilobita, p. 68-72. New Species of Order, Red Banner, Labour Geol. Inst. Palaeozoic plants and invertebrates, SSSR., Trans. Vol. 272, 76 pp., 11 pIs. Vol. 4, 214 pp., 32 pIs. Vanek, J. 1959, Celed Lichaidae Hawle et Corda, Nan Run·shan, 1980, Trilobita, p. 484-519. 1984, ze stredoceskeho starsiho paleozoika Paleontological Atlas of Northeast China (1) (Trilobita). Bohemia Centrals, Vol. 1, fasc. Paleozoic volume, 686 pp. 261 pIs. 3, p. 82-168, pIs. 1-12. (A-Sci. nat.) Owen, A. W. 1981, The Ashgill trilobites of the Warburg, E. 1925, The trilobites of the Leptaena Oslo region, Norway. Palaeontogr. Abt. A, Limestone in Dalarne. Bull. Geol. Inst. Vol. 175, p. 1-88, pIs. 1-17. Upsala, Vol. 42, p. 1-446, pIs. I-II. Owens, R. M. 1973, British Ordovician and --, 1939, The Swedish Ordovician and Lower Silurian Proetidae (Trilobita). Palaeontogr. Silurian Lichidae. Kungl. Svenska Vetens. Soc. Monograph., 98 pp., 15 pIs. -Akad. Handl. Tred., Ser. Vol. 17, no. 4, Perry, D. G. and Chatterton, B. D. F. 1977, p. 3 -162, pIs. 1-13. Silurian (Wenlockian) trilobites from Baillie­ Weber, V. N. 1932, Trilobites of the Turkestan. Hamilton Island, Canadian Arctic Archipe­ Mem. Comm. Geol. N.S. 178, 157 pp., 4 lago. Canad. Jour. Earth Sci., Vol. 14, no. 2, pIs. p.285-317. --, 1951, Upper Silurian trilobites of USSR. Phleger, F. B. Jr. 1936, Lichadian trilobites. Jour. Trudi VSEGEI, Ministry of Geology, 71 pp., Pal., Vol. 10, p. 593-615. 6 pIs. Ramskold, L. 1983, Silurian cheirurid trilobites Whittington, H. B. 1963, Middle Ordovician trilo­ from Gotland. Palaeontology, Vol. 26, pt. 1, bites from Lower Head, Western Newfound- 462 Teiichi KOBA YASHI and Takashi HAMADA

land. Bull. Mus. Compo Zool. Harvard Coil., cian trilobites from Northern Guizhou. Acta. Vol. 129, no. 1, 118 pp., 36 pIs. Pal. Sinica, Vol. 19, no. 1, p. 23-28, pI. 1. --, 1965, Trilobites of the Ordovician Table Zhang Tai-rong (1981) Trilobita, p. 134-213. In Head Formation, Western Newfoundland. Palaeontological Atlas of Northwest China Ibid., Vol. 132, no. 4, p. 275-442, pIs. Xingjiang-Weiwuer Autonomic Province Wu Honh-ji, 1977, Comments on new genera and Fascicle (1),332 pp., 92 pIs. species of Silurian-Devonian trilobites in Zhou Zhi-yi and Zhou Zhi-qiang, 1982, An Southwest China and their significance. Ashgill (Rawtheyan) trilobite faunule from Acta Pal. Sinica, Vol. 16, no. 1, p. 95-117, Ejin Qi, Nei Monggol (Inner Mongolia). 3 pIs. Acta Pal. Sinica, Vol. 21, no. 6, p. 659-671, Xian Institute of Geology and Mineral Resources, 1 pI. 1982, Palaeontological Atlas of Northwest China. Shaan-Gan-i\Iing Fascicule (1), 480 See also Kobayashi and Hamada (1974 and pp., 106 pIs. 1984) for other references here omitted. Yin Gong-zheng, 1980, New material of Ordovi-

Ertaogon ='~i'fl', Gomi 1\.1*, Guizhou yUH, Heilongjiang ,~1lI!,iL lchiyama mill, jiangxi iIlI.§, Kweichou (=Guizhou) iVH, Shaan·Xi MlI.§, Shihtien 1i(!i1J], Tienshan xLII, Yunnan ~l¥l, Xijiang WriI.

pLlmt'lt~UlO) c/)v )vK:c='lt!Rltr.A.,O);~1 2 !fl1JQ : * ~(,ij x: I':' Ii 5 HI':' !'~-t 0 9 ~TlO)= ~ 1(1); ,ie 1I'~~n-n'0o .:to)5'b1':'1:l:6:lfrfl!t2:lfrilli:fillil;fl;;l:n-n'L,:lfr~ Ichiyamella {';l;0o

.:t 0) {un.:. f*ff:'f:¥c~O)t':6br::'lEliYllt.d!:7EO) [113*t.( 1" 2 {[iii;;/; 0 0 .:-. n I? (1) ==::i;t;b. t I*l l'f-O);/; 0 flL I/.l.it.:.JiltI, 'L {, ET~frn& L L 1" 00 Ichiyamella, Japonoscutellum, Apolichas I:l:m{f.l:pj,: *ltO)~-C';/;00 'N:.j;fo-· ~Ill~.&±

Explanation of Plate 92

Figs. la-g. Dicranopeltis tricornis Kobayashi and Hamada, sp. nov...... Page 458 a, d: dorsal, b, e: left lateral and c, f: frontal views of the holotype cranidium show- ing an inflated shield with long and stout spines, PA 18082, a, b, c x2.0, d, e, f x1.0, coll. Asami. Figs. 2a-c. Apolichas truncatus Kobayashi and Hamada ...... Page 459 a: dorsal, b: frontal and c: left lateral views of a cranidium to show the trilobation and the large eye, PA 18083, x3.5, coIl. Kaneko. KOBA YASHI and HAMADA: Silurian Trilobites tram Shikoku Plate 92 Trans. Proc. Palaeont. Soc. Japan, N.S., No. 143, pp. 463-474, pIs. 93-94, September 30,1986

820. ON NEOCRIOCERAS SPINIGERUM (JIMBO), A SPECIES OF CRETACEOUS HETEROMORPH AMMONOIDS (STUDIES OF THE CRETACEOUS AMMONITES FROM HOKKAIDO-LV)*

TATSURO MATSUMOTO

c/o Department of Geology, Kyushu University, Fukuoka 812

KIKUWO MURAMOTO, TAKEMI TAKAHASHI, MINORU YAMASHITA and YOSHITARO KAWASHITA

c/o Museum of Mikasa City, Hokkaido 068-21

Abstract. Neocrioceras spinigerum (Jimbo), the type species of Neocrioceras, is redefined on the basis of the specimens in our collections as well as the hitherto described ones. Its adult body-chamber is spirally coiled in the same way as the phragmocone. It is quadrituberculate at intervals and the disposition of the ventro­ lateral tubercles is variable (opposite or alternate or irregular) among individuals and also with growth-stages. The quadrilobate suture is florid; I trifid and as deep as U; other elements bifid and of dove-tail form. Reliable records show that this species occurs in the Santonian of Hokkaido and Sakhalin. Comparison with other species is described at length, with a result that certain species previously referred to Neocrioceras should be transferred to other genera. In an appendix the genus Neocrioceras is redefined and its relationships with other genera are discussed.

In troduction to describe this species in detail with more illustrations. For some reasons, the genus Neocrioceras In this study the senior author is primarily Spath, 1921 has been understood inadequately responsible for the paleontological description by some authors. To remove this misunder­ and discussion; all the authors were engaged standing, the first step is to redescribe its type in field work of collecting fossils in respective species, Crioceras spinigerum Jimbo. This has areas concerned and in developing specimens been done concisely in a recently published from calcareous nodules. paper (Matsumoto, 1985), in which only two syntypes of Jimbo have been photographed Description of a species because of the page limitation. This paper is Neocrioceras spinigerum (JimbQ) PI. 93, Figs. 1-8; PI. 94, Figs. 1-16; *Received July 17, 1985; Read February 2, 1986 at Sendai. Text-fig. 1

463 464 Tatsuro MATSUMOTO et ai.

1894. Crioceras spinigerum Jimbo, p. 184, pI. text-fig. 5) from somewhere in Sakhalin (?). 24, figs. 1, la, 1 b. Repositories are as follows with abbreviation 1921. Neocrioceras spinigerum (Jimbo); Spath, in parentheses: p.51. Geological Collections, Kyushu University, 1933. Neocrioceras spinigerum (Jimbo); Shimi­ Fukuoka (GK). zu, p. 13, pI. 2, figs. 4-9. Institute of Geology and Palaeontology, 1963. Neocrioceras spinigerum (Jimbo); Matsu­ Tohoku University, Sendai (IGPS) moto, p. 45, pI. 67, figs. 1, la, lb. 1981. Neocrioceras spinigerum (Jimbo); Tanabe National Science Museum, Tokyo (NSM) et al., p. 220, pI. 37, figs. 2a, b; text-fig. University Museum, University of Tokyo 5. (UMUT) 1985. Neocriocnas spinigerum (Jimbo); Matsu­ The Muramotos' Collection (on display at the moto, p. 56, text-figs. 1, 2. Museum of Mikasa City), Mikasa (MC) Takemi Takahashi's Collection, Mikasa (TTC) Lectotype:-UMUT. MM7521 (Jimbo, 1894, Minoru Yamashita's Collection, Mikasa (MYC) pI. 24, fig. 1, 1a; Matsumoto, 1985, figs. lA, 1B; Yoshitaro Kawashita's Collection, Mikasa this paper, PI. 93, Figs. 1-2) from Ikandai of the (YKC) Urakawa area (southern central Hokkaido) , The specimens collected by T. Matsumoto designated by Matsumoto, 1985 (p. 56). (TMC) are kept at GK and UMUT. Other materiai:-Paralectotype, UMUT. Diagnosis:-Shell medium-sized, consisting of MM7522 (Jimbo, 1894, pI. 24, fig. 1b; Matsu­ slightly open crioceratoid planispiral whorls, moto, 1985, figs. 2A, 2B; this paper PI. 93, Figs. which are depressed in cross-section_ Body­ 3-5) from Ikandai. TTC_ 400819 (this paper, chamber about three fifths of the last whorl. PI. 94, Figs. 1-2) from Ikdandai; UMUT. Ribs fine and numerous, gently projected MM7640 from the Sosushi, a branch of the on the dorsum, sinuous on the dorsolateral part, Sanushibe, Hobetsu area (H. Yabe CoIl.); TTC. somewhat prorsiradiate on the flank, crossing 360806 (PI. 94, Figs. 6-9) from a branch stream the venter nearly vertically. Spiny tubercles in called the Y-sawa of the Kikume-zawa, a tribu­ four rows at intervals, with looping of minor tary of the Ikushumbets, Mikasa district; UMUT. ribs. The ventrolateral tubercles normally oppo­ 1-3531 (TMC_) from loc_ T 55 p and UMUT. 1- site, resting on low major ribs, but sometimes 3582 (TMC) from loco T 277 e of the Saku-Abe­ deviate slightly or even considerably from this shinai area (see Matsumoto, 1942, for the num­ disposition to give rise to irregular or alternate bered localities of TMC); TTC. 360815 (PI. 94, configuration, thus variable among individuals Figs. 14-16) from the Saku-gakko-no-sawa, two and/or with growth-stages_ The ventrolateral other unnumbered specimens of MYC. and tubercles often more numerous than the lateral YKC., all the above six from the Saku-Abeshinai tubercles by addition of intercalated ones. De­ area in the Teshio Nakagawa district; MC. Hbo- pending on the frequency of tuberculation, the 3695 (PI. 93, Figs. 6-8) from the Haboro area. intervening nodeless ribs vary from one to four, YKC. (without number) from the Obira area; occasionally showing outward branching_ H. Maeda's collection from the Onkono-sawa, Sutural formula E, L., U, I; lateral saddles Kotanbetsu area_ Also UMUT. MM7708 (M. and lobes (L, U) bifid and of dove-tail shape; Kawada ColI.) and TMC from the Naibuchi area; I nearly as deep as U and trifid; all the elements IGPS_ 36836 from Namikawa (Shimizu, 1933, finely and deeply incised. pI. 2, figs_ 4-9; this paper Text-fig. 1); GK. Description:-A completely preserved adult H5204, GK. H5205 (PI. 94, Figs. 3-5) and shell is very rare_ MC Hbo-3695, about 75 mm GK. H5218 from Kawakami (S. Nagaoka CoIl.), in diameter, is probably an adult shell, in which all the above six from South Sakhalin; NSM. body-chamber, although partly destroyed, occu­ PM9491 (Tanabe et ai., 1981, pI. 37, figs. 2a, b; pies about 2100 and its last septum is at 48 mm 820. Neocrioceras spinigerum (Jimbo) 465

Measurements (in mm):

Specimens Diameter Umbilicus Height Breadth B/H Lectotype 34.4 18.3 (.53) 9.5(.28) 14.5 (.42) 1.53 Paralectotype* 39.5 19.7 (.50) 11.6 (.29) 15.3 (.39) 1.32 Paralectotype -180° 31.0 13.8 (.45) 8.5 (.27) 11.0 (.35) 1.29 TTC.400819 24.4 11. 7 (.48) 7.8 (.32) 11.6 (.48) 1.49 TTC. 360806* 26.6 12.8 (.48) 9.0 (.34) 14.4 (.54) 1.60 TTC.360815 42.5 18.0 (.42) 13.8 (.32) (distorted) TTC.360815* 36.0 16.8 (.47) 12.4 (.34) 18.2 (.51) 1.47 MC. Hbo-3695 75.0 37.0 (.49) ~21 (.28) 34.5 (.46) ~1.6 MC. Hbo-3695* 48.0 22.5 (.47) 16.3 (.34) 24.2 (.50) 1.48 TTC.370817* ~43 ~22.5 (.52) ~15 (.35) ~23 (.53 ) ~1.5 UMUT. MM7640 20.4 11.4 (.54) 5.6 (.27) 8.0 (.39) 1.43 GK. H5205 33.2 15.6 (.47) 10.8 (.33) 15.2 (.46) 1.41 GK. H5204 27.7 13.2 (.48) 8.4 (.30) 11.7 (.42) 1.40 GK. H5118 34.0 16.0 (.47) 10.0 (.29) 14.4 (.42) 1.44 IGPS. 36836* 50.0 22.8 (.45) 16.5 (.33) 24.5 (.49) 1.48 * measured at the end of the phragmocone ~ approximate in shell diameter. It should be noted that the The observable part of the ammonitella is sub­ body-chamber is coiled in the same form as the helical and soon followed by a nearly planispiral phragmocone and never extended to a straight crioceratoid whorl. shaft as in Anisoceras. This denies the presump­ Shimizu's (1933) specimen was actually re­ tion of Spath (1921, p. 51). constructed with artificial connection between IGPS. 36836, described by Shimizu (1933, the outer and inner whorls. Therefore, the p. 13, pI. 2, figs. 4-9), is also an adult shell, slightly helicoidal form in his diagram is doubt­ but its body-chamber is incompletely preserved ful. and secondarily squashed. The whorl is characteristically depressed, but The lectotype is wholly septate. Some other the ratio of B/H is considerably variable from an specimens, e.g. paralectotype, TTC. 360815, individual to another and also with growth-stage. TTC. 370817, TTC. 360806, GK. H5118 and The measurements show that B/H ranges from GK. H5204, show the last septum at smaller 1.3 to 1.5 in many cases but occasionally exceeds diameters, from 40 mm to 25 mm, and only a beyond that range. Anyhow, the ventral part is portion of the body-chamber is preserved. They broad and its central zone between the tubercles may be more or less immature but some of is nearly flat, whereas the flanks are convex, them may be small adult shells. On the probably with the maximum breadth between the rows of adult body-chamber the ribs and tubercles are lateral tubercles. somewhat coarser than those on immature shells, In the diagram of whorl-section by Jimbo but the difference is by no means great and it (1894, pI. 8, fig. 1a) the four tubercles are is difficult to know the variation in size. illustrated as equidistant, but in the actual The ammonitella and early immature shell specimen the distance between the left and are not preserved in mot cases. NSM. PM 9491, right ventrolateral tubercles is wider than that reported by Tanabe et al. (1981), may be a com­ between the lateral and ventrolateral ones. The paratively better preserved example, but is yet same feature is observable in almost all the incomplete in lacking the very initial conch. specimens examined. In this respect the diagrams 466 Tatsuro MATSUMOTO et al. by Shimizu (1933, pI. 2, figs. 7, 8; disordered than the normal or average ones. In addition to in the explanation of plate) as well as Text-fig. the ventrolateral tubercles which correspond to 1C in this paper are correct. the lateral ones, more ventrolateral tubercles The disposition of tubercles is considerably occur in an intercalated disposition variably variable in this species. Some authors regarded in some specimens or at some growth-stages. the alternate disposition of the ventral (i.e. The tubercles in the late growth-stage have a ventrolateral) tubercles as a diagnosis of Neo­ long spine with a septate node at the base. The crioceras. This came probably from Shimizu's lateral nodes are somewhat larger than the (1933) specimen whose illustration was cited ventrolateral ones. The ventrolateral spines in the first edition of the Treatise (Wright in stretch upward and forward; the lateral ones Moore (ed.), 1957). In this specimen, IGPS. sideward. 36836, the ventrolateral tubercles are alternate The ribs are mostly simple and some of them on the figured part (Shimizu, 1933, pI. 2, fig. are looped at the tubercles. Occasionally the 5), i.e. on the last half of the phragmocone ribs are bifurcated outward. and the first 30° of the body-chamber, but The curvature of the rib, as described in the they are opposite or nearly so on other observ­ diagnosis, is indeed characteristic of this species, able parts. At the changing point from the but the degree of curvature varies to some alternate to opposite disposition on the body­ extent. The feature of the curvature, such as chamber a few ribs cross the venter obliquely dorsal projection, dorsolateral sinuosity and for adjustment (Text-fig. 1B). Incidentally the forward inclination on the flank, are generally highly squashed last part of the body-chamber weak on younger whorls, becoming more distinct of this specimen was excluded from the illustra­ later. The ribs cross the siphonal zone of the tion by Shimizu (1933, pI. 2, fig. 4). venter nearly vertically between the rows of In MC. Hbo-3695, the ventrolateral tubercles ventrolateral tubercles and some of them, looped are opposite on the body-chamber but irregular, by the tubercles, show a faint sinus (i.e. back­ if not quite alternate, on the late part of the ward arc). There are also indistinct constric­ phragmocone (see PI. 93, Figs. 8, 7). In the tions behind some of the major ribs which have lectotype and many other specimens (e.g. TTC. opposite tubercles. 360815, TTC. 370817, TTC. 360806, UMUT. The sutural pattern described in the diagnosis MM7640, UMUT. 1-3581, GK. H5118, etc.) is kept constantly. Jimbo (1894, pI. 24, fig. 1b) the ventrolateral tubercles are opposite or illustrated correctly the general pattern but nearly so. The paralectotype has also the op­ ignored minor incisions, probably because the posite ventrolateral tubercles in its young stages specimen (paralectotype) was somewhat eroded. but on the preserved last half whorl there is no Shimizu's (1933, pI. 2, fig. 9) illustration shows left ventrolateral tubercle and the ribs run minor details as well as the general pattern. irregularly. This is unusual and interpreted as Some more figures (Text-figs. lA, D) are added probably pathologic. In GK. H5205, which is in this paper. Even in small, immature specimens wholly septate, the ventrolateral tubercles are the suture is fairly deeply and finely incised. disposed irregularly or rather alternate in the In mature shells the suture is florid, with deep later part, but in the younger part they are and fine incisions and narrowed stems and opposite and rest on the low major rib. In GK. branches of the elements. Spath (1921) mis­ H5204 they are opposite on the phragmocone understood that suture of this species was simple. and tend to deviate from that disposition on the Occurrence :-Some of the above described succeeding part of the body-chamber. specimens are not precisely located in the strati­ The lateral tubercles number 7 or 8 in a half graphic sequence; others from the Upper Ura­ whorl, occurring as a rule at regular intervals kawan (K6b), approximately Santonian, and but occasionally at a longer or shorter interval often associated with Sphenoceramus naumanni 820. Neocrioceras spinigerum (Jimbo) 467

0':': :._.

c

B

L

,, , I ,I , I , I

I I , I ", I,,'

I , PI I I I o

Text-fig. 1. Some features on IGPS. 26836, an example of Neocrioceras spinigerum (Jimbo). A. External lobe of the last third suture. B. Disposition of ribs and tubercles on posterior half of the body-chamber. C. Diagrammatic whorl-section at D=50 mm. (Spines restored in which left ones (dotted lines) projected on the same plane). D. Suture at a young stage (H=5.4mm, B=7.5 mm). Scale bar=10 mm (A-C); 1 mm (D). These figures are to supple­ ment Shimizu's (1933, pI. 2, figs. 4-9). (T.M. de lin. )

(Yokoyama) and S. nagaoi (Matsumoto et Ueda). have compressed or nearly circular whorls. Besides the described specimens, fossils of They seem to be closer to N. (?) undulosum this species may be found occasionally from the Matsumoto, 1977, but are too incompletely Upper Urakawan (Santonian) of various areas preserved for a precise identification. in Hokkaido and also Sakhalin, although they Neocrioceras (Schlueterella) compressum are often hardly taken out in a complete shape Klinger (1976, p. 74, pI. 33, fig. 5; text-figs. 8j, from calcareous nodules. 109), from the Lower Santonian of Zululand, The true stratigraphic range of this species was established on a fragmentary whorl, curved beyond the Santonian and the geographic dis­ gently on a plane. The holotype clearly shows tribution outside Hokkaido and Sakhalin are a change in the curvature which suggests a sub­ yet uncertain. elliptical coiling; its whorl is laterally compressed Comparison:-The specimens from Pondoland and its suture has a very small I. These are described by Spath (1921, p. 52, pI. 7, figs. 6a­ fundamentally different from the characters c) under Neocrioceras cf. spinigerum are not of N. spinigerum. As in N. spinigerum the holo­ identified with the named species, because they type has ventrolateral and inner lateral tubercles 468 Tatsuro MATSUMOTO et al.

connected by looped ribs and intervening two and ventral rows). Its ribs do not show such a to three nontuberculate ribs. As only a single dorsal projection and dorsolateral sinuosity as fragmentary specimen was available, no remark those of N. spinigerum. Again its suture is not was given on the variation. This Zululand species known. This Gosau species is somewhat similar is provisionally referable to Schlueterella Wied­ to Kawashitaceras obiraense Matsumoto (1984b, mann, 1962 (emend. Matsumoto and Miyauchi, p. 342, figs. 2, 3A, 3B, 4A, 4B), but whether 1984). the summit of the ventral tubercle is minutely The specimens from the Lower Santonian of subdivided or not should be examined on better the Gosau Group (Austria) described by Immel preserved specimens. Anyhow, it is doubtful to et al. (1982) under Neocrioceras (Schlueterella) refer this species (maderi) to Neocrioceras. compressum seem to show an open subelliptical Ancyloceras kossmati Simionescu, 1899 (p. coiling (op. cit., pI. 10, fig. 1a), but this might 257 [21], pI. 1, figs. 6-8), from the Upper Creta­ have been affected by secondary deformation ceous of Transilvania (Roumania) was referred as suggested by some of the normally coiled to Neocrioceras (Schlueterella) by Wiedmann ammonites from the Gosau. A fragmentary (1962, p. 205). It has widely open whorls coiled phragmocone M 2 illustrated in the same paper subelliptically on a plane and the whorl seems (op. cit., pI. 9, fig. 3) indeed resembles the holo­ to be higher than broad, although the specimens type from Zululand, but the body-chamber may be secondarily deformed. Quadrituberculate and also the late part of the phragmocone (op. major ribs and intervening finer ribs without cit., pI. 10, figs. 2-4; pI. 11, fig. 3) show a zigzag tubercles encircle the whorl radially without arrangement of tubercles in the lateral and sinuosity and the ribs are looped only occasion­ probably also ventrolateral rows. This may be ally at a few tubercles. The ventral tubercles another characteristic of S. compressa. The same are opposite on the figured specimen (Simio­ character occurs more distinctly in S. kawadai nescu, 1899, pI. 1, figs. 8A, B). Again the suture Matsumoto et Miyauchi, 1984, from the Cam­ is not illustrated. This species is similar to S. panian of Hokkaido and Sakhalin, which differs compressa in shell-form but differs in its infre­ from contemporary European species S. pseudo­ quent occurrence of looped ribs at a few tuber­ armata (SchlUter, 1872) at this point. Regretta­ cles. It is provisionally referred to Schlueterella, bly the suture is not well exposed on the Gosau unless enough evidence be obtained for the specimens, but it is essential for a precise identifi­ establishement of a new genus or subgenus. cation. In this paper, however, the Gosau speci­ There are a few specimens (e.g. Hokkaido mens are provisionally identified with Schlueter­ University Collection 9467 with uncertain ella compressa. Anyhow, it is evidently distinct record of locality) which may represent an from Neocrioceras spinigerum. unnamed new species of Neocrioceras. This is N. (Schlueterella) mutinodosum (SchlUter, distinguished from N. spinigerum by a larger 1872) (see Wright, 1979), from the Turonian of size (about 100 mm in diameter of the phragmo­ Europe, should be referred to Pseudoxybeloceras cone), more open spiral form, subcircular whorl­ or P. (Christophoceras), because four rows of section and outward shifting tendency of lateral tubercles occur not only on major ribs, with tubercles in a late growth-stage. More specimens looped riblets, but also on intervening minor ribs. with reliable locality records are wanted to Neocrioceras (Neocrioceras) maderi Immel, define clearly this species. Klinger et Wiedmann, 1982 (p. 24, pI. 9, fig. 2; N. (?) undulosum Matsumoto, 1977, from pI. 11, figs. 1-2), from the Lower Santonian the Turonian of Hokkaido, is atypical in that of the Gosau Group, is distinguished from N. its immature shell shows a low helicoid shape spinigerum in its more gently arcuate and com­ and is followed by an open spiral shell in an pressed instead of depressed whorls and out­ undulating plane, but its body-chamber is gently ward shifted rows of tubercles (i.e. ventrolateral curved on that plane without a retroversal hook. 820. Neocrioceras spinigerum (Jimbo) 469

Its ornamentation is fundamentally similar to is a working hypothesis suggested by N. (?) that of N. spinigerum. undulosum Matsumoto. Neocrioceras has no affinity with Anisoceras Hyatt, because its Appendix body-chamber is spirally coiled in the same way as its phragmocone without straightly Remarks on the genus Neocrioceras extended arm and because the suture is differ­ TATSURO MATSUMOTO ent in its well developed I and also because there is no species which could suggest the con­ The genus Neocrioceras was proposed by nection. Spath (1921, p. 51), but the legal designation Schlueterella Wiedmann, 1962 is based on of Crioceras spinigerum Jimbo, 1894 as its type the incompletely known type species, Aniso­ species was settled by Diener (1925, p. 192). ceras pseudoarmatum SchWter, 1872 from the Since then this genus has been comprehended Campanian of Germany, and supplemented by widely and was even misunderstood by some a closely allied species S. kawadai Matsumoto authors. et Miyauchi, 1984, from the Campanian of the In the preceding pages the type species has northern Pacific region. Its revised generic been defined clearly and compared with other diagnosis has recently been given by Matsumoto species. In this appendix I attempt to give a and Miyauchi (1984, p. 59). It is similar to Neo­ generic diagnosis of Neocrioceras and remarks crioceras in the general mode of ornamentation on the relationships of this genus with other (four rows of tubercles at intervals with inter­ genera. The latter problem is still difficult, be­ vening untuberclate ribs and with looping of cause heteromorph ammonite species of this ribs at the tubercles) but different in its open, kind are often based on incomplete material. subelliptically (in stead of spirally) coiled whorls, Therefore, the following remarks, which are which consist of nearly straigt or gently arcuate in part cited from the recently issued papers long shafts and U-curved parts, and also in (e.g. Matsumoto, 1977; 1984a, b; Matsumoto having very small I in the suture. The mode of and Miyauchi, 1984) with some modification, disposition of ventral tubercles does not work may still be tentative. as a criterion. The shell-form of the type species Generic diagnosis:-Whorls in the main and S. kawadai in early immature stage is not growth-stages more or less open crioceratoid, known precisely and the phylogenetic origin coiled in a plane or nearly so, ornamented with of this genus is uncertain. Anyhow, on the numerous ribs and four rows of spiny tubercles, ground of the distinction mentioned above, i.e. the lateral and ventrolateral ones, at inter­ it is better to treat Schlueterella as an inde­ vals, with intervening nodeless ribs; typically pendent genus rather than to affiliate it to ribs looped at the tubercles. Suture deeply and Neocrioceras as a subgenus. finely incised, consisting of E, L, U, I; I trifid Kawashitaceras Matsumoto, 1984 was pro­ and as deep as U; other elements bifid and of posed for the type sepcies, Neocrioceras denta­ dove-tail pattern. tum Matsumoto et Obata, 1981, represented Distribution:-From Turonian to Santonian by a peculiar body-chamber from the Upper (possibly to Lower Campanian) in Japan and Turonian of Hokkaido, and another species Sakhalin, so far as reliable records are con­ K. obiraense Matsumoto, 1984, from the Upper cerned. Turonian (or possibly Coniacian) of Hokkaido. Relationships of Neocrioceras with other It has also quadrituberculate looped ribs and genera:-The phylogenetic origin of Neocrio­ intervening nodeless ribs, but the tubercles ceras is uncertain. It was tentatively sought in are distributed restrictedly on the ventral part the plastic and long-ranging genus Hyphanto­ and of dental form with minute crenulations ceras Hyatt, 1900 (Matsumoto, 1977), but this on their backward sloping summit. The tuber- 470 Ta tsuro MA TS UM OTO e tal.

cles have long spines in Neocrioceras and type species P. splendens Collignon, 1969, from Schlueterella, which would work as protection the Campanian of Madagascar, has ventrolateral against enemies. The body-chamber of Kawa­ tubercles alone on every rib but no lateral tuber­ shitaceras is long and broadly arcuate. The cle at any growth-stage. In its suture I is as deep shape of the preceding phragmocone is not as U. With respect to the shell-form and suture completely known but may be loosely open Parasolenoceras is similar to Pseudoxybeloceras. and subelliptically coiled as judged from some Periodic major ribs with stronger tubercles preserved pieces of curved parts. The peculiar are not recognized in the holotype of P. splend­ tubercles of a dental form distributed on the ens, but they appear more or less clearly in other ventral part may be favourable for this species, e.g. P. periodicum Matsumoto et Miya­ living in a shell of very unstable shape to repose uchi, 1984, P. compressum (Henderson, 1970) himself (or herself) on the bottom sediments and P. nanaimoense (Ward et Mallory, 1977) and also for him to start for occasional loco­ and fine ribs mayor may not be looped at the motion back upward in the sea water (Matsu­ tubercle. As in the case of Pseudoxybeloceras, moto, 1984b). The phylogenetic origin of this character alone would not be sufficient to Kawashitaceras is yet uncertain. Anyhow, separate a genus from Parasolenoceras. Kawashitaceras is distinct enough to be treated Lewyites Matsumoto et Miyauchi, 1984 (p. as a genus independent of Schlueterella or 64), established for Idiohamites (?) oronensis Neocrioceras. Lewy, 1969 (type species), I. (?) circularis Pseudoxybeloceras Wright et Matsumoto, Lewy, 1969, and Hamites (?) taylorensis Adkins, 1954, established on the type species, Hamites 1929, is distinct in that periodic major ribs quadrinodosus Jimbo, 1895 (emended by Matsu­ have ventrolateral tubercles where minor ribs moto, 1977), had nearly straight parallel shafts are looped and that intervening minor ribs have connected by U-curved parts. Numerous simple no tubercles. As in the suture of Schlueterella, ribs encircle the shell, of which some are peri­ I is much smaller and shallower than U. odically stronger than others. Every rib on the Hitherto reported species of Lewyites and shell of late grwoth-stages have tubercles in Parasolenoceras are all Campanian in age, where· four rows, ventrolateral and lateral ones. The as Schlueterella and Pseudoxybeloceras appeared major or stronger ribs have stronger tubercles earlier and ranged up to the Campanian. The where riblets are looped in some case but not lateral tubercles do not appear in young shells always so. Anyhow, the presence of tubercles of Ps. quadrinodosum and occur infrequently on every rib is one of the criteria to distinguish in Ps. linea tum. These facts suggest that Para­ Pseudoxybeloceras from Neocrioceras and solenoceras may have derived from Pseudoxy­ Schlueterella. The suture of P. quadrinodosum beloceras. Likewise, Lewyites may have branched figured by Jimbo shows shallow I, but that of from Schlueterella. P. lineatum (Gabb) (Matsumoto, 1959, fig. 79) The degree of differentiation into major clearly shows that I is as deep as U, being fairly and minor ribs varies from a species to another. similar to that of Neocrioceras spinigerum. I is Therefore, there is little reason to maintain somewhat smaller than other lobes as Matsumoto Christophoceras Collignon 1969 (type species (1959, p. 162) described previously but not so C. ramboulai Collignon, 1969) as a genus inde­ small as that of Schlueterella kawadai. With pendent of Pseudoxybeloceras. Some authors respect to shell-form, Pseudoxybeloceras may be (e.g. Klinger, 1976) may keep Christophoceras closer to Schlueterella than to Neocrioceras, as a subgenus of Pseudoxybeloceras, but in the but the shell-form in the early immature stage type species of Pseudoxybeloceras itself some is not precisely known in both Pseudoxybelo­ ribs are somewhat (but not much) stronger than ceras and Schlueterella. others and can be regarded as a kind of major Parasolenoceras Collignon, 1969, based on the ribs. 820. Neoerioeeras spinigerum (Jimbo) 471

As suggested by Ward and Mallory (1977), Axonoeeras Stephenson, 1941, established for Solenoeeras Conrad, 1860 (type species Hamites A. eompressum Stephenson, 1941 (type species) annulifer Morton, 1842) may be related to Para­ and other species from the Neylandville Marl solenoeeras. It is characterized by a smaller shell (possibly uppermost Campanian) of Texas, looks with greater appression of shafts. Lewy (1967) similar to Exiteloeeras in the mode of coiling, has shown a gyro conic spiral form of a young ribbing and tuberculation, but its whorls are shell in a species of Solenoeeras from Israel. irregularly in contact or partly open. In both Oxybeloeeras Hyatt, 1900 (type species genera the initial part is hooped. The suture has Ptyehoeeras erassum Whitfield, 1880) was not been illustrated for any species of Axono­ suppressed by some authors as a synonym of eeras. In the type species almost all the ribs have Solenoeeras but is kept by Matsumoto and ventral tubercles on either side of the siphonal Morozumi (1980, p. 20) as a subgenus of Soleno­ zone, but occasionally a few nodeless ribs may eeras for its larger size and absence of constric­ be intercalated. On the last part (probably adult tions. Solenoeeras is world-wide in the Cam­ body-chamber) of A. pingue Stephenson, 1941, panian and Maastrichtian. the tuberculate ribs are coarsen and alternated Exiteloeeras Hyatt, 1894 (type species An­ with one or two nodeless finer ribs. This char­ eyloeeras jenneyi Whitfield, 1880) has also acter reminds us of the ornament of Nostoeeras ventrolateral tubercles on every rib and seems Hyatt, 1894. The available evidence, however, to be loosely coiled earlier in a subelliptical form is by no means sufficient to connect Axonoeeras and later in a crioceratoid spiral (according to phylogenetically with Nostoeeras. the restroration by Scott and Cobban, 1966; Anaklinoeeras Stephenson, 1941 (type species Gill and Cobban, 1973, p. 10, fig. 6). The suture A. reflexum Stephenson, 1941, from the Ney­ of E. jenneyi (cited by Wright, 1967, fig. 251-7c landville Marl of Texas) and Planostoceras from Whitfield, 1880) is similar to that of Neo­ Lewy, 1967 (type species P. rehavami Lewy, erioeeras spinigerum in being deeply and finely 1967, from the Upper Campanian of Israel) incised and having bifid, dove-tail major ele­ are evidently offshoots of N ostoeeras, taking ments and pointed I as deep as U. Exiteloeeras the posture of body-chamber in a plane, as may have an affinity with Parasolenoeeras, but Pravitoeeras Yabe, 1902 (type species P. sig­ its true phylogenetic origin is uncertain. moidale Yabe, 1902 from the Upper Campanian The very initial stage of E. jenneyi has not of Japan) is a probable offshoot of Didymoeeras been known precisely, but Gill and Cobban taking a plane shell-form in the main growth­ (1973, p. 10) observed nearly straight (or broad­ stages (see Matsumoto et al., 1980; Morozumi, ly arcuate) limbs connected by a semicircular 1985). Some authors affiliate Anaklinoeeras bend in a juvenile (but not initial) stage. This and Planostoeeras to Nostoeeras as subgenera, suggests an affinity with such species as Para­ but the changes in shell-form seem to have solenoeeras tomitai Matsumoto, 1984a, which, occurred abruptly in these three examples. however, is based on incomplete specimens. To sum up, the tuberculate heteromorphs The two species are nearly contemporary (Upper whose main or adult shells posture nearly or Campanian), but known in much separate prov­ strictly in one plane are probably polyphyletic. inces, one from Wyoming and the other from Neocrioeeras and certain other possibly related Japan. E. uneiforme Lewy, 1969, from the Cam­ genera with four rows of tubercles and their panian of Israel, shows a crioceratoid phragmo­ presumed descendants with two rows of tuber­ cone and an unciforme body-chamber. E. ete­ cles could be grouped into a subfamily, for quenee Lewy, 1969 is peculiar in showing looped which Spath (1953) suggested the Neocriocera­ ribs at the tubercles and inervening nodeless ribs tinae. I would evaluate this suggestion, if the as in Lewyites, to which genus it could be trans­ intimate relationships among these genera were ferred. made clear with sufficient evidence. With regard 472 Tatsuro MATSUMOTO et ai. to many of them, however, the phylogenetic U.S. Geol. Surv. Prof. Paper 776, p. 1-37. origin is uncertain in our present knowledge. Henderson, R. A. (1970): Ammonoidea from the Mata Series (Santonian-Maastrichtian) of New Zealand. Special Papers in Palaeont., Acknowledgements no. 6, p. 1-82, pis. 1-15. Hyatt, A. (1894): Phylogeny of an acquired We wish to thank Professors Itaru Hayami, characteristic. Proc. Amer. Phil. Soc., Tamio Kotaka and Kametoshi Kanmera and Philadelphia, vol. 32, p. 349-647, pis. 1- Dr. Ikuwo Obta for their offer of facilities to 14. one of us (T.M.) for the study of the specimens --(1900): Cephalopoda. In Zittel, K. A. von. kept at UMUT, IGPS, GK and NSM respectively, 1896-1900: Textbook of Palaeontology, and also Dr. Kenshiro Ogasawara and Mr. Haru­ translated by Eastman, C. R., p. 502-604, yoshi Maeda for their kind help at IGPS and London. UMUT. We are also indebted to Dr. Masayuki Immel, H., Klinger, H. C. and Wiedmann, J. Noda for his kindness in taking photographs, (1982): Die Cephalopoden des Untereren to the authorities of the Japan Academy for Santon der Gosau von Brandenberg/Tirol, permission to reproduce the photographs of Osterreich. Zitteliana, vol. 8, p. 3-32, pls.1-11. Jimbo's type specimens printed in p. 57 (figs. Jimbo, K. (1894): Beitrage zur Kenntnis der 1-2) of the Proceedings of the Japan Academy, Fauna der Kreideformation von Hokkaido. vol. 61, ser. B, no. 2 (1985), and to Miss Yoshimi Paliiont. Abh., N. F., vol. 2, p. 149-194 Tanigawa in preparing the typescript. [3-48], pis. 17-25 [1-9]. Klinger, H. C. (1976): Cretaceous heteromorph References Cited ammonites from Zululand. Dept. Min. Geol. Surv. Mem., no. 69, 142 p., 43 pis., Pretoria. Adkins, W. S. (1929): Some Upper Cretaceous Lewy, Z. (1967): Some late Campanian nosto­ Taylor ammonites from Texas. Univ. Texas ceratid ammonites from southern Israel. Bull., 2901, p. 203-242, pis. 5, 6. Israel Jour. Earth-Sci., vol. 16, p. 165-173. Collignon, M. (1969): Atlas des Fossiles Carac· --(1969): Late Campanian heteromorph am­ teristiques de Madagascar (Ammonites), fasc. monites from southern Israel. Ibid., vol. 18, 15 (Campanien infedeur), p. 1-216, pis. p.109-135. 514-606, Servo Geol., Tananarive. Matsumoto, T. (1942): Fundamentals in the Diener, C. (1925): Ammonoidea neocretacea. Cretacenous stratigraphy of Japan. Part 1. Fossilium Catalogus, 1 Animalia, part 29, Mem. Fac. Sci. Kyushu Imp. Univ., ser. D, p. 1-244, Junk, Berlin. vol. 1, p. 129-210, pis. 5-20. Gill, J. R. and Cobban, W. A. (1973): Stratig­ --(ed.) (1963): A Survey of Fossils from Japan raphy and geologic history of the Montana Illustrated in Classical Monographs, 57 p., Group and equivalent rocks, Montana, 68 pis., Palaeont. Soc. Japan, Tokyo. Wyoming, and North and South Dakota. --(1959): Upper Cretaceous ammonites of

Explanation of Plate 93

Neocrioceras spinigerum (Jimbo) Figs. 1, 2. Lectotype, UMUT. MM7521, from Ikandai, Urakawa. Lateral (1) and ventral (2) views, xl. 5. Figs. 3-5. Paralectotype, UMUT. MM7522, from Ikandai, Urakawa. Lateral (3) and ventral views of pathologic body-chamber (4) and normal phragmocone (5), x 1.6. Figs. 6-8. MC. Hbo-3695, from Haboro. Lateral view (6) with Sphenoceramus nagaoi (Matsu­ moto et Ueda) and an ammonite, ventral views of a portion of phragmocone (7) and body­ chamber (8), xl. Photos by courtesy of Mr. H. Maeda (1-5) and Dr. M. Noda (6-8). MA TSUMOTO et al: Neocrioceras spinigerum (Jimbo) Plate 93

7 820. Neocrioceras spinigerum (Jimbo) 473

California. Part 2. Mem. Fac. Sci. Kyushu between Jarre Creek and Loveland, Colorado. Univ., ser. D, Spec. vol. 1, p. 1-172, pIs. U.S. Geol. Surv. Miscell. Geol. Invest. Map, 1-41. 1-439,1 map with text p. 1-4. --(1977): Some heteromorph ammonites from Shimizu, S. (1933): Note on two interesting the Cretaceous of Hokkaido. Ibid., ser. D, Senonian ammonites from Hokkaido and vol. 23, no. 3, p. 303-366, pIs. 43-61. South Saghalin. Jour. Shanghai Sci. Inst., --(1984a): Some ammonites from the Cam· sec. 2, vol. 1, p. 11-15, pI. 2. panian (Upper Cretaceous) of northern Simionescu, 1. (1899): Fauna cretacica superiora Hokkaidq. Part 1. Ammonites from the de la Urmos (Transilvania). Acad. Romana, Upper Campanian of the Teshio Mountains. vol. 4, p. 237-274 [1-38], pIs. 1-3. Palaeont. Soc. Japan, Spec. Pap., no. 27, p. Spath, L. F. (1921): On the Upper Cretaceous 5-32, pIs. 1-9. Ammonoidea from Pondoland. Ann. Durban --(1984b): An aberrant ammonite genus from Mus., vol. 3, p. 39--57, pIs. 6-7. the Cretaceous of Hokkaido. Proc. Japan --(1953): The Upper Cretaceous cephalopod Acad., vol. 60, ser. B, p. 341-344. fauna from Graham Land. Falkland lsi. --(1985): Restudy of Crioceras spinigerum Depend. Surv. Sci. Rep., no. 3, p. 1-60, Jimbo, a Cretaceous ammonite species. Ibid., pIs. 1-13. vol. 61, ser. B, p. 56-59. Stephenson, L, W. (1941): The larger inverte­ --and Miyauchi, T. (1984): Some Campanian brate fossils of the Navarro Group of Texas. ammonites from the Soya area. In Matsu· Univ. Texas Pub., no. 4101, p. 1-641 moto, T. (1984a): Some ammonites from (incl. pIs. 1-95), folded tables 1-6. the Campanian (Upper Cretaceous) of Tanabe, K., Obata, 1. and Futakami, M. (1981): northern Hokkaido. Part 2. Palaeont. Soc. Early shell morphology in some Upper Japan, Spec. Pap., no. 27, p. 33-76, pIs. Cretaceous heteromorph ammonites. Trans. 10-31. Proc. Palaeont. Soc. Japan, N.S., no. 124, --and Morozumi, Y. (1980): Late Cretaceous p. 215-234, pIs. 35-38. ammonites from the Izumi Mountains, Ward, P. D. and Mallory, V. S. (1977): Taxonomy Southwest Japan. Bull. Osaka Mus. Nat. and evolution of the lytoceratid genus Rist. no. 33, p. 1-31, pIs. 1-16. Pseudoxybeloceras and relationship to the --, --, Bando, Y., Hashimoto, H. and genus Solenoceras. Jour. Paleont., vol. 51, Matsuoka, A. (1980): Note on Pravitoceras p. 606-618 (incl. 3 pIs.) sigmoidale Yabe (Cretaceous heteromorph Whitfield, R. P. (1880): Paleontology of the ammonite). Trans. Proc. Palaeont. Soc. Black Hills of Dakota. In Newton, H. and Japan, N.S., no. 123, p. 168-178, pIs. 22- Jenney, W. P.: Report on the geology and 26. resources of the Black Hills of Dakota. --and Obata, 1. (1981): A new heteromorph U.S. Geog. Geol. Surv. Rocky Mtn. Region, ammonite from Hokkaido in the collection p. 325-468, 16 pIs. (inaccessible in Japan). of Yoshitaro Kawashita. Bull. Natn. Sci. Wiedmann, J. (1962): Ammoniten aus der Mus., Tokyo, ser. C, vol. 7, no. 3, p. 115- vascogotischen Kreide (Nordspanien). 1. 118, pIs. 1, 2. Phylloceratina, Lytoceratina. Palaeonto· Morozumi, Y. (1985): Late Cretaceous (Cam· graphica, vol. 118, A, p. 119--237, pIs. panian and Maastrichtian) ammonites from 8-14. Awaji Island, Southwest Japan. Bull. Osaka Wright, C. W. (1957): In Moore, R. C. (ed.): Mus. Nat. Rist., no. 39, p. 1-58, pIs. 1-18. Treatise on Invertebrate Paleontology. Part Schliiter, C. (1871-76): Cephalopoden der L, Mollusca, Cephalopoda, Ammonoidea, L oberen deutschen Kreide. Palaeontographica, l-L 490, Geol. Soc. Amer. & Univ. Kansas vol. 21, p. 21-24, pIs. 1-18 (1871); p. 25- Press. 120, pIs. 19--35 (1872); vol. 24, p. 121- --(1979): The ammonites of the English Chalk 263, pIs. 36-55 (1876). Rock (Upper Turonian). Bull. Brit. Mus. Scott, G. R. and Cobban, W. A. (1965): Geologic (Nat. Hist.) [Geol.], vol. 31, no. 4, p. 281- and biostratigraphic map of the Pierre Shale 332 (incl. 7 pIs.). 474 Tatsuro MATSUMOTO et al.

Wright, C. W. and Matsumoto, T. (1954): Some Univ., ser. D, vol. 4, no. 2, p. 107-134, doubtful Cretaceous ammonite genera from pIs. 7-8. Japan and Saghalien. Mem. Fac. Sci. Kyushu

Abesbinai 'it.:fZ:J*J=7"'c/71, Haboro +Jrm, Hobetsu 1l1l71I], lkandai ;tf~h lkushum· bets ~~:tf71I]. Kawakami JIII.. Kikume-zawa 1M lHi iR. Kotanbetsu [KotambetsJ 1~"fl71lJ. Mikasa =~, Naibuchi r"JiJ:iI. Nakagawa 9")11. Namikawa ~JII. Obira 'J'f. Onkono-sawa * /' ::z / iR, Saku &7-.., Saku-gakko-no-sawa itI:7-..$t:cO)iR. Sanushibe -if":>( c/ "'. Sosushi '/ ;<. c/o Teshio ::Rlll. Urakawa iJlif

13ill!*c~~~i;~7 /,-1':-71 r 0) 1 fill Neocrioceras spinigerum (JIMBO) I:''':)I,'L: Neocrio­ ceras ml,O)ti4:a::fifl"c';!; Q N. spinigerum (JI\\BO) O)i"J,;JRil;.iI G < IIJ!!f(, '2!'-:h L 1" t.n ' lW.{til; ;!;~kO)~ • • :a:~*K~~. ~O).m~.il-GMm'2!'-:hk~*~t.MG, ~&O)~m~~ i~ U':o $5'\-0) ~~rT*il;~O) iR71IH:.i~~ Q C GL 1" Q ,~, (5'HfliJ 0) ~ O)~~il;MlHiil-x~!iI'iCyIJil-) Ii, ill\lftq:.J: I), it.:liiJ-il!\l{-4:J*JO)!EftJl:IH:'J: 1)Il!:~il;;!;Q;: c~J!:{*(191:'~U,:o it.:lj\(;if WjO)tl:J.1H *flii:~JV10)r& C rnlt!(H:. ~ ~ * -t '7 ;<.~O)-t ~ i O);!; QflHi G-tt Iv~ ~~WML 1" Q ;: C ~ ~ Gil-I:. U':o la-iIii'i U~Qi&.ttEITc£jHe:;Mi0 < C, *:f!THiiJlil6J~/LI:.$ (:to t Ie: -if" /' I· =. 7 /') I:.i& G. Sphenoceramus naumanni (Yokoyama) ~ S. nagaoi (Matsumoto et Ueda) ~I-'¥?;: ct);~I,'o fIt* Neocrioceras ~1:'A:hG:hL\,'t.:[!tW.%~O)iJj\f&cO)tt"t~~tA.;r,... *& c il-tJ: I) "'tJ: Q {, 0) Ii ftl!o)~f:'f$ U.:1Jil; J:\, -;: c ~~i1Ui!tt U':o t~::4':~J'l~· N*tt7-../!$ '1(1;j~~~· UIT ~. )IITLIJ:;tJ'l~

fJ~ Neocrioceras ~f9TJiI,: ;:O)~O)%m~;~cL ftll.~cO)mlJ;J.,s~UIH~<.~~I}fj~G. fit *0)~J5L U':Jjf!;~~tlt~1j • ~.iI U':o ;!;Q ~I:' ,,:)1,'LIHfl1LO)m~~t%O)i.iJfjgtt~~~ U':o U- G*tfCl¥Jmf$~ ~ Gil-I:' -t Q 1:'li~m-t Q iJj\f;j\JH:. ":)\, 'L O)~JiI,I:'/Gviilil;;!; Q ;: !: ~1I1ji/ii G, ~~~.gI¥JM.~~

Explanation of Plate 94

Neocrioceras spinigerum (Jimbo) Figs. 1-2. TTC. 400819, from Ikandai, Urakawa. Lateral (1) and ventral (2) views of an im­ mature shell, x 1.5. Figs. 3-5. GK. H5205, S. Nagaoka's ColI. from South Sakhalin. Ventral (3), lateral (4) and fron tal (5) views, x 1. Figs. 6-9. TTC. 360806, from the Kikume-zawa, Mikasa. Two lateral (6, 8) and ventral (7, 9) . views, x 1. 2. Figs. 10-13. TTC. 360815, from Nio-no-sawa, Toyosato. Two lateral (10, 12) and ventral (11, 13) views, xL Figs. 14-16. TTC. 370817, from the Saku-gakko-no-sawa, Saku. Lateral (14) and ventral views of phragmocone (15) and a portion of body-chamber (16). Photos by courtesy of Dr. M. Noda. MATSUMOTO et al: Neocrioceras spinigerum (Jimbo) Plate 94

1 3 4 5

8 9

15 475

PROCEEDINGS OF THE PALAEONTOLOGICAL SOCIETY OF JAPAN

a*i5~4~¥~~ 135 @]19~~ n~JEm.=. '/ of- ? ;< :J11 f]~ (Trochidae) O)lJli:n<.n::. E *i11:!ffrr¥:~m 1351"1WIJ~I.I;1986ff 6 Ji 14' 15 -::J\"L ...... :;!CJ1iRlifitr.l£ E ~tJLHlnl:il: L3 ~;ttlf.¥/.JMi7:l-*-e~~llt ~htc. ($JJn ft~~~7/'~~1bO)~~~~O)W~ .. iIIrl* ~ :;lj103~)o A record of Menabites (Ammonoidea) from the Cretaceous of Hokkaido ...... @ A ~ ;1l · ...... T. Matsumoto and T. Takahashi ~tJL1Hp~J\l'iI:r (iifiif1TtrJO 0) Squalodon 'IJI1:'-::J Little known ammonite Grandidiericeras I,'L ...... 1l7;l~l!.f~& from the Cretaceous of Hokkaido ...... Recognition of the genus Enterodon from · ...... T. Matsumoto and R. Saito Iwaki Formation of Joban Coalfield and Note on an ammonite species of Pachydiscus obliteration of "Anthracothema tsuchiyai" from Awajj Island, Southwest Japan ...... (nom. nud.) ...... , ... Y. Tomida T. Matsumoto, Y. Morozumi and T. Ozawa tl*:1 l/:!l '/ 3 /' '1'0) "):n\.!lfJ~\J3nlllrpH" 11i'U{', y /v ~ l/ /,y (/{of-~ ~ /') 1:.:t,;vtZ,~/vA$C-lj- ~1tEI:'-::J\"L ...... lI1li;j,;~* /' :1"1tE ..... , ...... , .. iII~t.j-{'f- • 1JI1IW IDP. On the genus Palaeoloxodon from the East ~,1iMrjHi'i~EIK:E l I) Fusulinella. Fusulina China Sea ...... H. Otsuka 1tE[il)~llf~O)feJ~I:'-::J\"L ...... 1'l1j)j iH..f (,/$l;/iR i:%) 0) Yl Jf; !E. fR ...... '5ll:.i ~ ~~1 ...... riIIljJJ!t· f!i:1Il~~:r iH..f~fl']f3!E. 0) f'Utl.l , Gj'iiJI.I;:hI.l, Z, 1.1' ? ...... ~.W.It!l~~mlK~~~z,&/.~Mm~MO)~ ...... , Ironmn#E' fillfill£fr:b; 1!i!E.:¥fl:: ...... ji'fIUl!#lJ1 • :E111~ff~91!. ~lHjlll.7..~ 77~.0)*~~~:1-¥ ~0)3-00~W~~0) "fmlELTITI*7-,\,- ~O)ji(lIx£t\~11~ti:l'lr,~xHt .... f~ ff ...... ' ...... f!~:fr ~ffr~; ...... , ..... , ...... Il7;JH J'~ 1lt~1jl7HJc'IT;;fMff)0)""lv·qc~9iiJEffl7 ot -T, t < i:mlfti1%f19\:- "fml'I' %Jrl'flO:.:t-ovt z, I¥jQfj~HrfL!'u 1t I:'~ 0) r~-~J:.¥.!rJIl!..f!lli'!'~:,®,:;m ...... , .. III iRMj- E;lIH:.-::J\"L ...... illi 'iL,Mir.! 1:IiHtEr::"ic~J ~ htc.IL\~t1:4~0) JJJi:*~cf,R t bJi::R: fi~A.~~Ji!(~m~M~~O)~1:~R!E.Wm(~ I:.{f? 1JifjiJt;(t"t;;0)~1t ...... , ... ,1H/r{~'~; m) .. , ...... , ... " ...... , ·f:i\~til~ f!!lm E *0) Nipponitrigonia kikuchiana I::' -::J\" ILl p l)jl.~iY'J;~liy:ilftf;H·iffif'0 (3) ..... ·I)'lIWil*:kAA L ...... tlHl~'T· III ml it.lt!l J:Jf'il 0) ;:fk!Jly !(E'i ;fl3.~l J!i1l!!! fu nt ...... ~ ••D·~M7~~7/'0)iH..f1:=ttYl~E ...... *HjfflJl·*{ERT·~1Jl( ~ ...... rnf\;.iIz· i'lIi~ 1m YHtf"fmlS!lli*l I) i1f!±l Ltc. 3 ft]0)l!!!f'01tE .... /Il:i..!I1frrmljj[~llfO)J:jUf(lJj'!.tt ...... EB{I::.iIZ • · ...... *tt8~1j)j· :;!c{EJ3.';'f ~~.~·~H X'~EB~ft'MEBma cp. ~tmlJL1'IH&WJ%f~{1::0)1tEifHfullf ~O) 3 )( Restudy on "Inoceramus yubarensis NAGAO W.lfutdZ , .. , ...... " .. *~~/l£k' !5~Ij)jT et MATSUMOTO" from the Upper Cretaceous of Hokkaido ...... M. Noda ~:,...;f- ~ rJ.4 ltl.:s1:iEi{:Effli~Hi 1!R;f,C{!::1:i:f J p~1tEilJ~llfl:'-::J\"L ...... , .... §~lX5:j tlt~,5A : tliPlBjI;y-' :k1B..iIJ11 ~tiH..f.ilfi:l1t~-¥,,*,~rlrj.;J.It!lJi!(O)i'fJil/lllJi'l 0) Yl~Htfi W fl'fL!E.:t-o Hh / ~. /' HtE'ltH:. -::J\, 'L ...... ~(~0)1) ...... ~*1j)j& ... ,., ...... , .. ,., .. r~*tilZ 'l'icJllft\:;jft ;T ~ 7 " n~=txYl ~ 7J" ,/;j; ~ TO)[email protected]}f;n.~ ~1v'::1tE!i'H:'-::JI,'L ...... f~UiJ~~ 'l'icJllif1::rt ...... iill7K m ij!!;ft[fbfultl~!iH:'-::J\"L .... illilBJ3.';m· .7..IImtFJ'- ~~~Yl~ill ••~O)~~.~~.MK~~<~ IllilJEfUtEllff::'-::Jl'L ...... ·jljPH3/Y- iIIrlm'lt%ftl!:f!.!ffl1tE!IWIO)f¥HJT ...... "flll.iI­ ::. 11' .'E.1tE llfl:' -::Jl' L ...... m:J::i~1( • f~HllBiJL ~~:!l~:17~-KlZ,**'/~~Tff1 "fml!!O)Jttfi'lIDI:'-::J\"L '" .:kIB.iI.ilfi:· ***,- Limopsis tajimae 0)1:.Wi.* ...... JlIjlilifi1: 476

New members approved by the Council Meeting held on June 13,1986 Yen-Nien, Cheng; Satoshi, Chiba; Chikara, Hiramatsu; Toshiaki, Irizuki; Megumi, Kobayashi; Fumio, Kubota; Hiroshi, Kurita; Takashi, Murota; Jun, Nakano; Yukihiro, Numabe; Yoshi­ taka, Ohta; Ikuo, Sunouchi; Kiyoshi, Takatsuka; Yuichiro, Tanaka; Yasuhiro, Taniuchi; Yutaka, Toda; Akira, Tsukagoshi; Michio, Yoshida. Members resignated Juzo, Arai (Fellow); Yoshio, Hojo; Tadashi, Horiuchi.

Bibliography U)flj:(j(;:?\. \-C

B *~1:.!J&.J~~"('f:!:, 1981-1985%0) 5 ~r,~I':::'~~ ~.hf-:NiftrxO) Bibliography >£ffifJIH.:::. J: IJ flJrri" 7.l .: c re_ts: IJ i Lt..:o .: 0) Bibliography t':::'11V't*JlH IJ J-1J1:!J&.J~~~~O)lVf7C~iftrX>£'I:J,L.'I':::', II ;{:O)'tiH>£;iS"J

iJ' ",:,f..:;jl'~ .wO)~x~ i3"&I)7.l .: eeL i i" 0 xiiikO)~l(ji;11~@IAO) $ttr~£~ c L -C frl ,t..: l' c ftl,l' i i"0)"(', J:,1~ 5 -:f7J'0)~iftr:)( ~ 7, ~ YfO) (lIH.:::.ftt -:> -c f'Fl.ix L, 1986~12~ 3113 i -r::t-oiZS IJ T ~ l '0

W AT ANABE Kozo (1973) Pro!usulinella assemblage in the Omi Limestone, Niigata Prefecture, central Japan (Studies of Carboniferous fusulinacean of Omi, part 1). Transactions and Proceedings of the Palaeontological Society of Japan, New Series, no. 95, pp. 371-394, Plates 51-53, Figure 1, Tables 1-11. Carboniferous Fusuline [,*~cp O)*Jt~XO){7~]

BANDO Yuji and KATTO Jiro (1980) On the Upper 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. Rinya­ kosaikai Press, Kochi, pp. 95-100. (j','Ij~-(Jm*WO):=::

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. U:f(H EI ~e·~:=::e~*~~R~&~~O)~~.O)~.~b~K~~~~·~~~~~~) ~) Cretaceous to Tertiary Larger Foraminifera [,*~cpO)*XJH~ff ~5fIlX~iftrXO){7Ij]

KOBA YASHI [wao and KAKIZAKI Takeo (1978): [Preservation of veretebrate bones in nodules from the Tsurushi Formation] Saito Ryojiro Sensei Taishoku Kinenshi, pp. 43-49, Plates 1-2. (i!lT~~/;;:L -JH.:::.'e2jjj1~.ht..:tj·1t:OO)f*ff-) (J.) Miocene Mammalia [t.6Ux~

i'± 1. aiftrxI1~~0)~f(;II~I':::'.?I17·L-CT~l'0 ~0)7,.?I1 J!.-11ftt*O)i=I~,JH.:::.V'tl'ii"0 2. '*ltt.:g,111tl'm~ Lt.r.l '"('ill '-C

3. ~xcp"(';iS"JiJ' -:> t..:I:I'1~O)~mln • HH~11 Index ~*.I@ L, ~,f,1~ A L -C t..:~ffiJX>£ t-3j\\0 ~ 0)~frI1:t-o~ b-l±

Hrilix ~ :A I- O)jzsN:'t 'T980 {Iiltnnl'f #;111 JldtA:':r:Jlj!"f:ffllJt1l1!{~f'r~/J:.47/r'1:r~-:r;( I}~ 1:1 :4':J'i~I'-:'1&.}"¥=0)(mk 0 nmtls~.l't 0 ::(jilIi.fl'mr:~ • R1~ ~

TAXA INDEX

ANIMALIA COELENTERATA General Anthozoa Problematica, Trace fossils Zoantharia and Others Tabulata CHORDATA Scleractina Mammalia Tetracorallia Proboscidea Octocorallia Artiodactyla Stromatoporata Aves Hydrozoa Reptilia Scyphozoa Amphibia POLYFERA Pisces PROTOZOA Excl. MASTIGOPHORA Osteichtyes Radiolaria Chondrichtyes Rhizopoda PROTO CHORDATA Foraminifera Graptolithia Fusuline Conodontochordata Larger Foraminifera ECHINODERMATA Benthic Smaller Foraminifera Holothuroidea Planktonic Foraminifera Echinoidea VEGET ABILIA Crinoidea General BRACHIOPODA Problematica and Others BRYOZOA Palynology ARTHROZOA ANGIOSPERME Insecta Dicotyledoneae Crustacea GYMNOSPERME Malacostraca Coniferopsida Cirripedia Ginkgopsida Ostracoda Cycadopsida Branchiopoda Pteridospermopsida Trilobita PTERIDOPHYTA ANNELIDA Pteropsida MOLLUSCA Articulatae Cephalopoda NON TRACHEOPHYTA Coleoidea Charophyta Ammonoidea Calcareous Algae Nautiloidea Coccolithophoridae Mollusca excl. Cephalopoda Dinoflagellata Bivalvia Diatomae Scaphopoda Silicoflagellata Gastropoda Others Polyplacophora and Monoplacophora 478

GEOLOGIC AGE INDEX

GENERAL OR INDEPENDENT TO AGE LATE CRETACEOUS PRE- 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 ~~ fiJi :11!! 00 Wi 8 ilt4'iji($jz,*'li'j;I]8 1987if. if.~.t{€.~ Mt IlliJ "* '* 1987if.l}j 30 8-2}j 1 8 1986if.ll}j 15 8 1987Jt. 1361ill 1JIJf< fl,jJHIjl&f'l4toiril 1987if. 6 }j 20 8-218 1987if. 4 }j 5 8

;to ~ 6 'It

01987if.Jj[if.~ • t{€.~"elt r 8;;$: C {- 0) 1m j?1 O)fn'::::~-c1lW1l!!~;f§J i!tg-;5 A± ~-tJ: G Lfv:::. ri¥li'lJ) 7 :t 'Y -iT .... ?'Tv:::'$V1 QiJBj~'in tt~~:I1!!:I!I!.J t!t:lliAfn~fi3~-c'~;/;f-~ '" - A tJ~5t@j~n -C\"i-ro oaB'¥!l61if. 9 }j 208 ffl.tE-c'¥IH1:f • *,i:1ji:0)·H#i?0)1*~'tI2~-c'-r 0 t-.: t.:\" i rii:~~ n i -r c, 1 if.tJF9v:::.1±l ~~nQT5E-c'-rO)-c'b Q -? -C :::'rii:~

;to c. C ~ I)

O*,,"jj~m5Ev:::'lt rjt$~'-C'il0 -? -C

i Lt-.: (~~~ Jt:!'~J 0

B ;;$: r!l 1:. ¥JJ '+- ~ 1 9 8 6 if. 9 }j 25 8 X E( !K ifF ~ 2-4-16 1986 if. 9}j 308 Q ;;$: '* ~ lO}J -t! ;/ ~ - r9 (~ ~ j:l !¥: * E( 84780 '/11=) ISSN 0031-0204 ('il[ ~;5 0 3 - 8 1 7 - 5 8 0 1 ) 8 ;;$:tt~~~~¥Il~·*'i:* ~ ~ ~ ~ • ~ ffi i!:i ::i: !ff EH liS ~ JT 1111 143~ r:p .lP1J :ti * E( 'IlIl ** .~ lK ~ ~ ~t 2 / 13 2,5 0 0 fI3 ,*~Hi~ iF t:iJ.IJl1J ·t~tt:~ t± '& rn ~ ('lit ~;5 03 - 9 9 1 - 3 7 54) Transactions and Proceedings of the Palaeontological Society of Japan

New Series No. 143 September 30, 1986

CONTENTS

TRANSACTIONS 816. MATSUMOTO, Tatsuro and NODA Masayuki: Some inoceramids (Bivalvia) from the Cenomanian (Cretaceous) of Japan-I. New or little known four species from Hok- kaido and Kyushu ...... •...... •...... 409 817. GOTO, Hiroya, MARUOKA, Kunitaka and ISHD, Ken-Ichi: Lepidolina columbiana (Permian fusulinid) from British Columbia, Canada ...... •.....•....• 422 818. NISHIDA, Harufumi: On a new Tempskya from Japan .....••.....•...... 435 819. KOBAYASHI, Teiichi and HAMADA, Takashi: The second addition to the Silurian trilobite fauna of Yokokura-yama, Shikoku, Japan ...•...... •... 447 820. MATSUMOTO, Tatsuro, MURAMOTO, Kikuo, TAKAHASHI, Takemi, YAMASHITA, Minoru and KAWASHITA, Yoshitaro: On Neocrioceras spinigerum (Jimbo), a species of Cretaceous heteromorph ammonids (Studies of the Cretaceous ammonites from Hokkaido-LV) ....•..•...... •..•...... 463 PROCEEDINGS ...... •...... 475