BulL Kitakyushu Mus. Nat. Hist. Hum. Hist., Ser. A. 3: 135-143, March 31, 2005

Early Cretaceous freshwater fishes from the Tetori Group, central Japan

Yoshitaka Yabumoto Kitakyushu Museum of Natural History and Human History, Yahatahigashiku, Kitakyushu, Fukuoka, 805-0071, Japan e-mail: yabumoto @kmnh.jp

(Received September 10, 2004; accepted December I, 2004)

ABSTRACT—Early Cretaceous freshwater fish fossils from the Kuwajima Formation of the Itoshiro Subgroup, Tetori Group described in detail for the first time are assigned to Lepidotes sp. (isolated scales), Sinamiidae (many isolated bones and scales), Pachycormidae (a dentary), Osteoglossiformes (a disarticulated skull with vertebrae and scales) and Teleostei (isolated scales).

detail by Isaji et at. (2005). INTRODUCTION

Two Early Cretaceous freshwater fish assemblages SYSTEMATIC DESCRIPTION have been found in Japan. These are the fossils from the Wakino Subgroup of the Kanmon Group of Kitakyushu Class Actinopterygii City in the northern part of Kyushu, and the Itoshiro Subclass Neopterygii Subgroup of the Tetori Group in Shiramine, Ishikawa Order Semionotiformes Prefectureand Shokawa, Gifu Prefecture in central Japan Family Semionotidae (Uyeno, 1979; Yabumoto, 1994; 2000). Genus Lepidotes Agassiz, 1839 Shokawa, Gifu Prefecture and Shiramine, Ishikawa Lepidotes sp. Prefecture (Figure 1) have yielded many fish fossil speci mens. The first fish fossil, a left lower jaw, was found in Figure 2 Kuwajima of Shiramine village in 1987, and was reported together with a few ganoid scales belonging to the Two scales (SBEI-818 and SBEI-1268) have been Semionotiformes, Holostei, Osteichthyes (Azuma and found. These scales have a thick cover layer of ganoin Hasegawa, 1989). Two sinamiid fishes recognized from that makes the exposed area of the scale smooth. The Shokawa were reported at the meeting of the exposed area is rectangular, and slightly deeper than long. Palaeontological Society of Japan (Yabumoto, 1997). The dorsal overlapped area exhibits a pointed process that Since 1997, many isolated fish scales and bones have been fit into the depression of similar shape at the ventral found in Kuwajima, Shiramine. In the present paper, the margin of the inner side of the scale dorsal to it, creating specimens from this locality are described in detail for the peg-and-socket articulation. The anterior overlapped first time. area shows two processes; the upper one extending antero- dorsally with a rounded end, and the lower one extending

LOCALITY AND HORIZON anteriorly with an acute tip. There is a ridge on the inner surface from a point slightly below the middle of the All fish specimens described here have been found in anterior margin to the postero-ventral corner of the scale. Kuwajima, Shiramine, Ishikawa Prefecture, central Japan The thick ganoin covering and the dorsal and ante (Fig. I). The horizon is the Kuwajima Formation of the rior processes in the overlapped area of the scales are very Itoshiro Subgroup in the Tetori Group, and is from the similar to those of the genus Lepidotes, Semionotidae early Neocomian age. Depositional environments and (Woodward 1916-1919). Thus, the scalesare assignedto taphonomy ofthe Kuwajima Formation were described in the genus Lepidotes.

This paper is one of those given at the Symposium of Early Cretaceous Terrestrial Biota held in the Kitakyushu Museum of Natural History and Human History on the 1st and 2nd of September in 2003. 136 Yoshitaka Yabumoto

•igure 2. Scales of the genus Lepidotes from the Kuwajima Forma tion in Kuwajima. Ishikawa Prefecture. Japan, a. SBIZI-SIS: b. SBEI-1268. Scales indicate 5mm.

anterior margin. This specimen is almost the same as the anterior part of SBEI-295. SBEI-014 is a left dentary. and the first fish fossil from the Tetori Group found in 1987. It has a deep coronoid process at the posterior portion, and its low. thick anterior part bears canine-like teeth curved inward. A series of pores opening from the mandibular canal run along the ventral margin of the dentary (Figure 3b). SBEI-1951 is a right dentary. Its anterior tip is missing. Posteriorly, the coronoid process is broken at the base, and bent inward. Anteriorly, there are canine like teeth curving inward. The estimated total number of Figure I. The locality of the Early Cretaceous freshwater fish fossils. teeth is 22 (14 preserved teeth and 18 sockets for missing teeth). The anterior tooth-bearing portion is moderately Order Amiiformes curved towards the symphysis (Figure 3c. d). Family Sinamiidae Two gular plates (SBEI-300 and SBEI-1208) have Genus Sinamia Stensio, 1935 been found. SBEI-1208 is almost complete only with its Sinamia sp. anterior tip missing. Its ventral surface is exposed, the anterior portion being narrower than its posterior portion. Figures 3 and 4 There is a very narrow groove in the middle of its anterior portion. The posterior portion is triangular with convex Most fish remains from Kuwajima can be referred to lateral margins, and a slightly concave, almost straight this taxon. There are all together 18 specimens of skull posterior margin (Figure 31*). SBEI-300 is less complete bones and 277 of detached scales. than SBEI-1208. lacking part of the posterolateral portion. Three specimens of maxilla (SBEI-295. SBEI-1860 It differs from SBEI-1208 in its wider anterior portion and and SBEI-2017) have been found. SBEI-295 is an almost shorter groove on the anterior portion, not reaching the complete left maxilla with only its posterior end missing; middle of the bone. There are many pits on the inner the bone gradually deepening posteriorly. The external surface of the bone of the latter (Figure 3e). surface is exposed. It has a relatively long and stout SBEI-817 is a right hyomandibular (Figure 3g) anterior articular process for the premaxilla. The large exposed in its lateral side. It is a hatchet-shaped bone canine-like teeth along the oral margin are strongly curved with a posterior process for articulation with the opcrcle. inward. Ornamentations on its lateral surface are in the The dorsal arm is slightly longer than the ventral arm. form of pits in the anterior and grooves in the posterior The foramen for the hyomandibular trunk (containing portions (Figure 3a). fibers from the facial and lateral line nerves) opens at the In SBEI-2017 only the middle portion of a left middle of the anterior margin of the dorsal arm. and the maxilla without teeth is preserved and looks the same as groove behind the foramen is located at the middle of the the middle part of SBEI-295. anterior margin of the bone. There is a nearly right angle SBEI-1860 is the anterior portion of a right maxilla. between the posterior margin of the ventral arm and the There are two pores on its inner face along the dorso- ventral margin of the process for articulation with the Cretaceous Fishes from Tetori Group. Japan 137

Figure 3. Fossils of the genus Sinamia from the Kuwajima Formation in Kuwajima. Ishikawa Prefecture. Japan, a. left maxilla. SBEI-295: b. left dentary. SBEI-014: c. dorsal view of left dentary. SBEI-295: d. outer view ofdentary. SBEI-I95I: e. gular plate. SBEI-1208: f. gular plate. SBEI-300; g. right hyomandibular. SBEl-817: h, right frontal, SBEI-823; i. cast of the left infraorbital, SBEI-1207; j. lea infraorbital, SBEI-1257. Scales of i and j indicate 2 mm. Other scales indicate 5 mm. 138 Yoshitaka Yabumoto opercle. The angle between the posterior margin of the margins of the ventral part are convex (Figure 4e). dorsal arm and the dorsal margin of the process for SBEI-317 is a left supracleithrum with only a small articulation with the opercle is slightly acute, but the axes dorsal portion of the bone preserved; most of the ventral of the dorsal arm and the process form an obtuse angle. portion being preserved as the impression of its inner face. The dorsal margin of the dorsal arm is convex. The proportion and shape are almost the same as those of SBEI-823 is a right frontal. Its dorsal surface is SBEI-315. A lateral line canal runs through the bone covered by an enamel layer. A series of pores opening from the dorsoanterior process to the pore opening on the from the supraorbital sensory canal are found on the inner surface at the point slightly above the middle of the dorsal surface of the bone. The suture for the dermo- posterior portion (Figure 4h). sphenotic is located just behind the shallow concavity of SBEI-301 is a left supracleithrum with its outer sur the orbit. The posterior margin of the frontal indicates face exposed. The enamel layer remains on the posterior that this species has a single parietal. The width of the and ventral portions of the outer surface. Its proportions anterior and posterior portions are almost the same. The are quite different from those of SBEI-315, -317. The suture with the left frontal is slightly undulate (Figure 3h). posterior margin is almost straight, and the anterior Two infraorbital (SBEI-I207 and SBEI-1257) bones margin, rather convex. The ventral margin is thin, and have been found. Specimen SBEI-1207 is an impression almost straight (Figure 4i). of a left infraorbital (probably the last one because of the SBEI-1192 is a left supracleithrum missing the ventral short length) (Figure 3i). Its dorsal portion bends slight part. The enamel layer remains on the postero-ventral ly inward. There are pits and grooves radiating from the part of the bone. Its posterior margin is almost straight point which is slightly above the middle of the anterior (Figure 4d). This specimen is similar to SBEI-301. margin of the bone. SBEI-1257 is a left infraorbital with a Many rhombic scales with a ganoin layer have been small posterior part missing (Figure 3j). found; some are isolated, others articulated (SBE1-297, The preopercle (SBEI-312) is a narrow crescent- or FI-3-271) or assembled (SBEI-297, FI-3-277). These bow-shaped bone; the ventral three-fourths of the bone scales have a thin ganoin layer and a large overlapped being almost straight and slightly wider than the dorsal area not covered by ganoin along the dorsal and anterior part, which bends slightly anteriorly. Pores of the margins. No processes have been observed in the over preopercular sensory canal open along the posterior lapped area. The outer surface is smooth without any margin of the ventral part and the dorsal tip. The dorsal ornamentation, and the posterior margin, also smooth tip is missing, but the impression of the bone and canal is with no serrations. There is a vertical ridge in the middle preserved. The external surface of the bone has pitted of the inner surface (Figure 4j, 1). and grooved ornamentation (Figure 4a). Although the material described above consists only SBEI-316 is a right cleithrum; a crescent-shaped bone of isolated bones and scales, there is no doubt that they with a dorsal and a ventral arm. The dorsal arm is belong to the genus Sinamia based on the following covered with ridged ornament, and has a slender dorsal characters: (l)The maxilla has a relatively long, stout tip, whereas the ventral arm has a reversed V-shaped anterior articular process for the premaxilla, with large unornamented area. The two arms form an obtuse angle canine-like teeth strongly curved inward along the oral (Figure 4b). margin; (2)The shape of the frontal indicates a single SBEI-1259 is a right cleithrum missing the ventral parietal; (3)The hyomandibular is hatchet-shaped; (4)The arm, posterior margin and dorsal tip (Figure 4c). cleithrum has ridged ornamentation on its outer surface; Four supracleithra (SBEI-301, SBEI-315, SBEI-317, (5)The gular plate exists; (6)The scales are covered by a and SBEI-1192) have been found. The supracleithra are thin ganoin layer, and have large overlapped areas, but no shoehorn-shaped. Two types of supracleithra (SBEI-315 processes (Schulitze, 1996). and-317; -301 and -1192) are recognized. SBEI-315 is a complete right supracleithrum with Order Pachycormiformes three processes at the dorsal end (Figure 4e, 0- The Family Pachycormidae anterior process contains a tube for the main lateral line Gen. et sp. indet. canal traversing the bone in a posterior and slightly ventral direction (Figure 4g) with two pores opening at Figure 5 the posterodorsal corner of the bone (Figure 4f). An enamel layer is seen on the part along the posterior Only an anterior part of a dentary (SBEI-826) has margin, and some small enamel spots on the dorsal por been collected from Kuwajima, and this can be placed in tion of the bone as well. The anterior and posterior the Pachycormidae. This bone is narrow, with well- Cretaceous Fishes from Tetori Group. Japan 139

Figure 4. Fossils ofthe genus Sinamia from the Kuwajima Formation in Kuwajima. Ishikawa Prefecture. Japan, a. right preoperclc. SBEI-3I2; b. right cleithrum. SBEI-316: c, right cleithrum. SBEI-I259; d, left supracleithrum, SBEI-1 129: e. right supracleithrum. SBEI-315: f. dorsal end ofright supracleithrum. SBEI-315; g, anterior view of the dorsal end of right supracleithrum. SBEI-315; h. right supracleithrum. SBEI-317: i. left supracleithrum. SBEI- 301: j. ganoid scales, SBEI-297; k. ganoin scales, SBFI-297. Scales of b and c indicate 5 mm. Other scales indicate 2 mm. 140 Yoshitaka Yabumoto

Figure 5. Dentary of the family Pachycormidae from the Kuwajima Formation in Kuwajima. Ishikawa Prefecture. Japan. Enlarged mid-part of teeth band, below. SBEI-S26. Scale of a indicates 5 mm. Scale of b indicates 2 mm. preserved dentition. The external surface of the dentary scales probably belonging to one individual have been is exposed (Figure 5a). It has large, irregular-sized teeth found in one piece of gray siltstone (SBEI-1489a) (Figure (the anterior teeth are smaller than the posterior ones) 6a). The bones include preopercles. a left opercle. cleith- located inside of a villiform tooth band covered on the ra. infraorbitals, a left scapula, and a neurocranium outer surface of the bone along the oral margin. The shown in its left view, lacking the ethmoid region. In surfaces of the large teeth are smooth (Figure 5b). addition, many scales, the posterior part of a para- This fossil appears to belong to the family Pachycor sphenoid, and some unidentified bones have been found midae. because the dentition resembles that of the Jurassic in another piece of gray siltstone (SBEI-1489c) (Figure gen us Hypsocomuis. 6b). On SBEI-1489a. the sensory canal on the horizontal arm of the preopercle runs quite straight, with the sensory Division Teleostei canal pores open in a shallow groove. The anterior Order Osteoglossiformes portion of the frontal is wider than the posterior portion. Family, gen., ct sp. indet. Its infraorbitals behind the orbit are large. Long teeth are found on the parasphenoid. The scales are thick and Figure 6 large with radial grooves. Some of the characters observed on the specimens Disarticulated skull bones, vertebral centra, and described in this section indicate that this fish can be Cretaceous fishes from Tetori Group. Japan Ml

Figure (>. The osteoglossiform fish from the Kuwajima Formation in Kuwajima, Ishikawa Prefecture. Japan, a. SBEI-I489a: b. SBEI-I489c. Abbreviations: Bab = basibranchial; Cle = cleithrum; Fro prefrontal: lfo = infraorbital; Op = operculum: Preo-preoperculum: Pte= pterygoid; Sc = scales; Sca = scapula: Supe—supracleithrum. Scales indicate 10 mm

placed in the Osteoglossiformes. Among them, the most important are: the long teeth on the parasphenoid. and the large infraorbitals covering the cheek region.

Order, family, gen. et sp. indct.

Figure 7

Discovered from Kuwajima were many round or oval small cycloid scales. It is certain that they belong to teleosts. possibly primitive ones. Most scales are very thin, with six to eight ridges without any grooves.

CONCLUDING REMARKS

Identification of fossil fishes Although most of the fish fossils are isolated bones Figure 7. Cycloid scales belonging to the division Teleostei from the and scales, the following taxa can be recognized. They are: Kuwajima Formation in Kuwajima. Ishikawa Prefecture. Japan. SBEI-298. Scales indicate 2 mm.

Semionotidae. Lepidotes sp. Sinamiidae, Sinamia sp. In this Early Cretaceous locality, fossils of the genus Pachycormidae. gen. et sp. indeterminable Sinamia are the most abundant probably due to the Osteoglossiformes. family, gen. et sp. indeterminable depositional environment (Isa.ii et a/.. 2005) favorable for Teleostei. order, family, gen. et sp. indeterminable the preservation of robust bones and scales. Probably these specimens represent two species as two different 142 Yoshitaka Yabumoto

types of gular plates and supracleithra have been found. record (Laeliichthys ancestralis Silva Santos, 1985 from However, it is a bit difficult to compare the material from Brazil; Aptian) of the family, this fossil is significant for Kuwajima to other species of the genus because of the understanding the origin of osteoglossids. As the frontal incompleteness of the specimens. For the purpose of has wide anterior, and narrow posterior portions similar comparison the author examined the holotypes of to that of the osteoglossid subfamily Phareodontinae Sinamia poyangica Su and Li, 1990, S. chinhuaensis Wei, which consists of Phareodus, Phareoides (probably a 1976, S. huananensis Su, 1973 and S. luozigouensis Li, synonym of the Phaleodus), Brychaetus, Cretophareodus, 1984. A few differences between these previously estab it maybe further assigned as a member of the phare- lished species and the Kuwajima ones have been noticed. odontins which would make it 60 million years older than Hyomandibular SBEl-817 differs from that of S. the previous oldest record (Cretophareodus alberticus Li, huananensis in the convex outline ofthe dorsal margin of 1996 from Canada; middle Campanian) of the sub the dorsal arm (it is almost straight in S. huananensis); S. family. chinhuaensis in its smaller process for articulation with The other Early Cretaceous freshwater fish fauna the opercle and lacks ridges on its lateral surface (S. found in Japan is the Wakino fish fauna (Yabumoto, chinhuaensis has the strong ridges); 5". poyangica in the 1994). Twenty one species have been described on the strongly convex outline of the dorsal margin of the dorsal basis of the articulated specimens, and three different arm (the dorsal margin is almost straight in S. poyangica), assemblages have been recognized: these are Nipponamia- and the posterior margin of the dorsal arm forming an Aokiichthys assemblage from the Dobaru Formation acute angle with the dorsal margin; S. zdanskyi with the (lowest), Paraleptolepis-Wakinoichthys assemblage from acute angle between the posterior margin of the dorsal the Gamo Formation (the third formation from the bot arm and the dorsal margin of the process for articulation tom) and Diplomystus-Wakinoichthys assemblage from with the opercle (5". zdanskyi has an obtuse angle), the the Kumagai Formation (top). No fish remains have relatively short ventral arm, the more concave anterior been reported from the Takatsuo Formation (the second margin, and the convex dorsal margin of the dorsal arm formation from the bottom). The Wakino fish fauna was (slightly convex, almost straight in S. zdanskyi). characterized by the abundance of "Diplomystus" and The author also examined the holotype and paratype Aokiichthys (Yabumoto, 1994). of S. luozigouensis, but the hyomandibular is not pre The age of both sediments from Kuwajima and served in the types. There is a chronological difference Kitakyushu is almost the same, but except for the between the species from the Kuwajima Formation, early semionotid the faunae are different. Probably Lepidotes Early Cretaceous and S. luozigouensis, late Early is shared by both faunae. Various species of Lepidotes Cretaceous, approximately Aptian or Albian (Chang have been found from China and Korea in East Asia Mee-mann personal communication, 2003). (Chang and Chou, 1977) and all over the world (Nel The dentary SBEI-295 which was reported to belong son, 1994). However, Sinamia, which is the most abun to a semionotiform fish by Azuma and Hasegawa dant form in the Kuwajima fish fauna, has not been found (1989), is instead placed in the amiiform genus Sinamia in the Wakino Subgroup. on the basis of the deep coronoid process and canine-like The Wakino Subgroup in Japan and the Nagdong teeth curved inward by the present author. Subgroup of the Gyeongsang Group in Korea are consid The pachycormid dentary SBEI-826 is similar to that ered of the same age because both have yielded the same of the Late Jurassic Hypsocormus macrodon (Wagner, species, Wakinoichthys aoki Yabumoto, 1994 1858) from Solnhofen, Germany, but the surfaces of the (Yabumoto and Yang, 2000). Sinamiid fossils have large teeth are smooth in the specimen from Kuwajima been found in the Nagdong Subgroup, but not in the (H. macrodon has fine ridges). Pachycormids had been Wakino Subgroup. The composition of the Wakino fish reported only from marine sediments, therefore the speci fauna more resembles that of the fish fauna from the men from the Kuwajima Formation represents the first southeastern part of China than that of the Jehol Group; report of the group from non-marine strata. "Diplomystus" and Aokiichthys from the Wakino Sub group are close to Paraclupeaand Paralycoptera from the Significance of the fish fauna southeastern part of China. The osteoglossiform fish (SEBI-1489a, c) from the The difference of the fish fauna of the Kuwajima Kuwajima Formation is the best preserved fish fossil from Formation from other areas in East Asia is probably the Tetori Group collected so far. If the author's deduc caused by the difference ofthe depositional environments. tion that it belongs to the family Osteoglossidae is correct, Further research and excavation of the Kuwajima Forma then at 25 million years older than the previous oldest tion are necessary and important for the understanding of Cretaceous Fishes from Tetori Group, Japan 143 the Early Cretaceous of this area. (eds.), Mesozoic Fishes —Systematics and Paleoecology, pp. 285-298, figs. 1-7. Liu, T.-S., Liu, H.-T., and Su, T.-T., 1963. The discovery of Sinamia zdanskyi from the Ordos region and its stratigraphical signifi ACKNOWLEDGMENTS cance. Vertebrata PalAsiatica, 7: 1-30, pis. 1-8. The author expresses his sincere gratitude to Dr. Nelson, J. S. 1994. Fishes of the world, 3rd edition. John Wiley & Son, Inc., New York, xvii+600 pp. Teruya Uyeno of the National Science Museum, Tokyo Schulitze, H.-P. 1996. The scales of Mesozoic actinopterygians. and Dr. Chang Mee-mann of the Institute of Vertebrate Mesozoic Fishes —Systematics and Paleoecology, G. Arratia & Paleontology and Paleoanthropology for their critical G. Viohl (eds.): pp. 83-93, figs. 1-6. Silva Santos, R. Da. 1985. Laeiiichthys ancestralis, novo genero et reading of the manuscript and their valuable advice. He especie de Osteoglossiformes do Aptiano da Formacao Areado, thanks Dr. Makoto Manabe of the National Science Estado de Minas Gerais, Brasil. Colet. Trab. PaleontoL, Ser. Museum, Tokyo and Dr. Shinji Isaji of the Natural Ceol. 27, sec. PaleontoL e Estr. 2: 161-167. History Museum and Institute, Chiba for their advice and Stensio, E. 1935. Sinamia zdanskyi, a new amiid from the Lower Cretaceous of Shantung, China. Palaeontologia Sinica, Series information of the fossils and the fossil locality. He also C, 3(1): 1-48, figs. 1-20, pis. 1-17. thanks Dr. Jin Fan of the Institute of Vertebrate Su, D. and Li, H. 1990. Note on new Sinamia from Jiangzi, South Paleontology and Paleoanthropology for his help to east China. Vertebrata PalAsiatica, 28: 140-149, pis. 1-2. Su, T.-T. 1973. A new Sinamia (5. huananensis, sp. nov) from the examine the type specimens ofthe Sinamia in China. He Upper Jurassic of southern Anhui. Vertebrata PalAsiatica, 11: thanks Mr. Ichio Yamaguchi, Mrs. Mikiko Yamaguchi, 149-152, pi. I. and Dr. Tatsuya Sakumoto ofShiramine Village for their Uyeno, T. 1979. Early Cretaceous freshwater fishes from northern Kyushu, Japan. I. Description of two new species of the clupeid help for this study. This study was supported by the genus Diplomystus. Bull. Kitakyushu Mus. Nat. Hist., (I): 11-24, Japanese Society for the Promotion of Science Grant-in- figs. 1-3, pis. 3-4. aid for scientific research and Fujiwara Natural History Wagner, A. 1858. Geschichte der Urwelt, mit besonderer BerQcksich- tigung der Menschenrassen und des mosalschen SchSpfungsber- Foundation. ichtes. 2e Theil, vi +528pp. Wei, F. (Zhejiang Museum). 1976. New discovery of Early Cretaceous fossil fishes from Jinhua, Zhejiang. Vertebrata REFERENCES PalAsiatica, 14: 154-159, pi. 2. Woodward, A. S. 1916-1919. The fossil fishes of the English Weal- Agassiz, L. 1839. Untersuchungen iiber die fossilen Fische der den and Purbeck Formations, Part I. London Monogr. Lias-Formation. Jahrbuch f. Mineralogie, pp. 139-149. Palaeont. Soc, 1916: 1-48, pis. i-x; Part II, 1918: 49-104, pis. xi-xx; Azuma, Y. and Y. Hasegawa. 1989. Vertebrate fossils. The Part III, 1919: 105-148, pis. xxi-xxvi. report for the conservation ofthe fossil localities along the Tetori Yabumoto, Y. 1994. Early Cretaceous freshwater fish fauna in River, pp. 26-32, Shiramine Village Board of Education. Kyushu, Japan. Bull. Kitakyushu Mus. Nat. Hist., (13): 107-254, Chang, M. -M. and C.-C. Chou. 1977. On late Mesozoic fossil fishes figs. 1-96, pis. 36-59. from Zhejiang Province, China. Acad. Sin., Instil. Palaeont. Yabumoto, Y. 1997. Early Cretaceous freshwater fishes from Sho Palaeoanthro., Mem., (12): 1-59, figs. 1-28, pis. 1-25. kawa, Gifu Prefecture, Japan. Abstracts with Programs. The Isaji, S., H. Okazaki, R. Hirayama, H. Matsuoka, P. M. Barrett, T. 1997 Annual Meeting. The Palaeontological Society of Japan, Tsubamoto, M. Yamaguchi, I. Yamaguchi, and T. Sakamoto. p. 26. 2005. Depositional environments and taphonomy of the bone- Yabumoto, Y. 2000. Fossil fishes. In Fossils from the Kuwajima bearing beds of the Lower Cretaceous Kuwajima Formation, "Kaseki-kabe" (Fossil-bluff). —Scientific report on a Neocomian Tetori Group, Japan. Bull Kitakyushu Mus. Nat. Hist. Hum. (Early Cretaceous) fossil assemblage from Kuwajima Formation, Hist., Ser. A, (3): 123-133. Tetori Group, Shiramine, Ishikawa, Japan, pp. 46-49, pis. 14-15. Li, G. -Q. 1984. Discovery of Sinamia from East Jilin. Vertebrata Shiramine Village Board of Education, Ishikawa Prefecture, PalAsiatica, 22(2): 143-150, pi. I. Japan. Li, G. -Q. 1996. A new species of Late Cretaceous osteoglossid Yabumoto, Y. and S.-Y. Yang. 2000. The first record of the Early (Teleostei) from the Oldman Formation of Alberta, Canada, and Cretaceous freshwater fish, Wakinoichthys aokii, from Korea. its phylogenetic relationships. In, Arratia, G. and Violhl, G. Bull. Kitakyushu Mus. Nat. Hist., (19): 105-110, figs. 1-4.