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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 ItoshiroSubgroup, Tetori Group described in detail for the first time are assigned to Lepidotes sp. (isolatedscales), Sinamiidae (many isolated bones and scales), Pachycormidae (a dentary), Osteoglossiformes (adisarticulated skull with vertebrae and scales) and Teleostei (isolated scales).
INTRODUCTION
Two Early Cretaceous freshwater fish assemblageshave been found in Japan. These are the fossils from theWakino Subgroup of the Kanmon Group of KitakyushuCity in the northern part of Kyushu, and the ItoshiroSubgroup of the Tetori Group in Shiramine, IshikawaPrefectureand Shokawa, Gifu Prefecture in central Japan(Uyeno, 1979; Yabumoto, 1994; 2000).
Shokawa, Gifu Prefecture and Shiramine, Ishikawa
Prefecture (Figure 1) have yielded many fish fossil specimens. The first fish fossil, a left lower jaw, was found inKuwajima of Shiramine village in 1987, and was reportedtogether with a few ganoid scales belonging to theSemionotiformes, Holostei, Osteichthyes (Azuma andHasegawa, 1989). Two sinamiid fishes recognized fromShokawa were reported at the meeting of thePalaeontological Society of Japan (Yabumoto, 1997).Since 1997, many isolated fish scales and bones have beenfound in Kuwajima, Shiramine. In the present paper, thespecimens from this locality are described in detail for thefirst time.
LOCALITY AND HORIZON
All fish specimens described here have been found inKuwajima, Shiramine, Ishikawa Prefecture, central Japan(Fig. I). The horizon is the Kuwajima Formation of theItoshiro Subgroup in the Tetori Group, and is from theearly Neocomian age. Depositional environments andtaphonomy of the Kuwajima Formation were described in
detail by Isaji et at. (2005).
SYSTEMATIC DESCRIPTION
Class Actinopterygii
Subclass NeopterygiiOrder Semionotiformes
Family Semionotidae
Genus Lepidotes Agassiz, 1839
Lepidotes sp.
Figure 2
Two scales (SBEI-818 and SBEI-1268) have beenfound. These scales have a thick cover layer of ganointhat makes the exposed area of the scale smooth. Theexposed area is rectangular, and slightly deeper than long.The dorsal overlapped area exhibits a pointed process thatfit into the depression of similar shape at the ventralmargin of the inner side of the scale dorsal to it, creatingpeg-and-socket articulation. The anterior overlappedarea shows two processes; the upper one extending antero-dorsally with a rounded end, and the lower one extendinganteriorly with an acute tip. There is a ridge on the innersurface from a point slightly below the middle of theanterior margin to the postero-ventral corner of the scale.
The thick ganoin covering and the dorsal and anterior processes in the overlapped area of the scales are verysimilar to those of the genus Lepidotes, Semionotidae(Woodward 1916-1919). Thus, the scalesare assignedtothe genus Lepidotes.
This paper is one of those given at the Symposium of Early Cretaceous Terrestrial Biota held in theKitakyushu Museum of Natural History and Human History on the 1st and 2nd of September in 2003.
136 Yoshitaka Yabumoto
Figure I. The locality of the Early Cretaceous freshwater fish fossils.
Order Amiiformes
Family Sinamiidae
Genus Sinamia Stensio, 1935
Sinamia sp.
Figures 3 and 4
Most fish remains from Kuwajima can be referred to
this taxon. There are all together 18 specimens of skullbones and 277 of detached scales.
Three specimens of maxilla (SBEI-295. SBEI-1860and SBEI-2017) have been found. SBEI-295 is an almost
complete left maxilla with only its posterior end missing;the bone gradually deepening posteriorly. The externalsurface is exposed. It has a relatively long and stoutanterior articular process for the premaxilla. The largecanine-like teeth along the oral margin are strongly curvedinward. Ornamentations on its lateral surface are in the
form of pits in the anterior and grooves in the posteriorportions (Figure 3a).
In SBEI-2017 only the middle portion of a leftmaxilla without teeth is preserved and looks the same asthe middle part of SBEI-295.
SBEI-1860 is the anterior portion of a right maxilla.There are two pores on its inner face along the dorso-
•igure 2. Scales of the genus Lepidotes from the Kuwajima Formation 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 atthe base, and bent inward. Anteriorly, there are caninelike teeth curving inward. The estimated total number ofteeth is 22 (14 preserved teeth and 18 sockets for missingteeth). The anterior tooth-bearing portion is moderately
curved towards the symphysis (Figure 3c. d).
Two gular plates (SBEI-300 and SBEI-1208) havebeen found. SBEI-1208 is almost complete only with its
anterior tip missing. Its ventral surface is exposed, theanterior portion being narrower than its posterior portion.There is a very narrow groove in the middle of its anteriorportion. The posterior portion is triangular with convexlateral margins, and a slightly concave, almost straightposterior margin (Figure 31*). SBEI-300 is less completethan SBEI-1208. lacking part of the posterolateral portion.It differs from SBEI-1208 in its wider anterior portion and
shorter groove on the anterior portion, not reaching themiddle of the bone. There are many pits on the inner
surface of the bone of the latter (Figure 3e).
SBEI-817 is a right hyomandibular (Figure 3g)exposed in its lateral side. It is a hatchet-shaped bonewith a posterior process for articulation with the opcrcle.The dorsal arm is slightly longer than the ventral arm.The foramen for the hyomandibular trunk (containingfibers from the facial and lateral line nerves) opens at the
middle of the anterior margin of the dorsal arm. and thegroove behind the foramen is located at the middle of theanterior margin of the bone. There is a nearly right anglebetween the posterior margin of the ventral arm and theventral 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 thedorsal arm and the dorsal margin of the process for
articulation with the opercle is slightly acute, but the axes
of the dorsal arm and the process form an obtuse angle.
The dorsal margin of the dorsal arm is convex.
SBEI-823 is a right frontal. Its dorsal surface is
covered by an enamel layer. A series of pores opening
from the supraorbital sensory canal are found on the
dorsal surface of the bone. The suture for the dermo-
sphenotic is located just behind the shallow concavity of
the orbit. The posterior margin of the frontal indicates
that this species has a single parietal. The width of the
anterior and posterior portions are almost the same. The
suture with the left frontal is slightly undulate (Figure 3h).
Two infraorbital (SBEI-I207 and SBEI-1257) bones
have been found. Specimen SBEI-1207 is an impressionof a left infraorbital (probably the last one because of theshort length) (Figure 3i). Its dorsal portion bends slightly inward. There are pits and grooves radiating from thepoint which is slightly above the middle of the anteriormargin of the bone. SBEI-1257 is a left infraorbital with asmall posterior part missing (Figure 3j).
The preopercle (SBEI-312) is a narrow crescent- or
bow-shaped bone; the ventral three-fourths of the bonebeing almost straight and slightly wider than the dorsalpart, which bends slightly anteriorly. Pores of thepreopercular sensory canal open along the posteriormargin of the ventral part and the dorsal tip. The dorsaltip is missing, but the impression of the bone and canal ispreserved. The external surface of the bone has pittedand grooved ornamentation (Figure 4a).
SBEI-316 is a right cleithrum; a crescent-shaped bone
with a dorsal and a ventral arm. The dorsal arm is
covered with ridged ornament, and has a slender dorsaltip, whereas the ventral arm has a reversed V-shapedunornamented area. The two arms form an obtuse angle
(Figure 4b).
SBEI-1259 is a right cleithrum missing the ventralarm, posterior margin and dorsal tip (Figure 4c).
Four supracleithra (SBEI-301, SBEI-315, SBEI-317,and SBEI-1192) have been found. The supracleithra areshoehorn-shaped. Two types of supracleithra (SBEI-315and-317; -301 and -1192) are recognized.
SBEI-315 is a complete right supracleithrum withthree processes at the dorsal end (Figure 4e, 0- Theanterior process contains a tube for the main lateral linecanal traversing the bone in a posterior and slightlyventral direction (Figure 4g) with two pores opening atthe posterodorsal corner of the bone (Figure 4f). Anenamel layer is seen on the part along the posteriormargin, and some small enamel spots on the dorsal portion of the bone as well. The anterior and posterior
margins of the ventral part are convex (Figure 4e).SBEI-317 is a left supracleithrum with only a small
dorsal portion of the bone preserved; most of the ventral
portion being preserved as the impression of its inner face.
The proportion and shape are almost the same as those of
SBEI-315. A lateral line canal runs through the bone
from the dorsoanterior process to the pore opening on theinner surface at the point slightly above the middle of the
posterior portion (Figure 4h).
SBEI-301 is a left supracleithrum with its outer sur
face exposed. The enamel layer remains on the posterior
and ventral portions of the outer surface. Its proportions
are quite different from those of SBEI-315, -317. Theposterior margin is almost straight, and the anteriormargin, rather convex. The ventral margin is thin, andalmost straight (Figure 4i).
SBEI-1192 is a left supracleithrum missing the ventral
part. The enamel layer remains on the postero-ventralpart of the bone. Its posterior margin is almost straight(Figure 4d). This specimen is similar to SBEI-301.
Many rhombic scales with a ganoin layer have been
found; some are isolated, others articulated (SBE1-297,
FI-3-271) or assembled (SBEI-297, FI-3-277). These
scales have a thin ganoin layer and a large overlappedarea not covered by ganoin along the dorsal and anteriormargins. No processes have been observed in the overlapped area. The outer surface is smooth without anyornamentation, and the posterior margin, also smooth
with no serrations. There is a vertical ridge in the middle
of the inner surface (Figure 4j, 1).Although the material described above consists only
of isolated bones and scales, there is no doubt that they
belong to the genus Sinamia based on the followingcharacters: (l)The maxilla has a relatively long, stoutanterior articular process for the premaxilla, with largecanine-like teeth strongly curved inward along the oralmargin; (2)The shape of the frontal indicates a singleparietal; (3)The hyomandibular is hatchet-shaped; (4)Thecleithrum has ridged ornamentation on its outer surface;(5)The gular plate exists; (6)The scales are covered by athin ganoin layer, and have large overlapped areas, but noprocesses (Schulitze, 1996).
Order Pachycormiformes
Family Pachycormidae
Gen. et sp. indet.
Figure 5
Only an anterior part of a dentary (SBEI-826) hasbeen collected from Kuwajima, and this can be placed inthe 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 endof right 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 scalesindicate 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 dentaryis exposed (Figure 5a). It has large, irregular-sized teeth(the anterior teeth are smaller than the posterior ones)
located inside of a villiform tooth band covered on the
outer surface of the bone along the oral margin. The
surfaces of the large teeth are smooth (Figure 5b).This fossil appears to belong to the family Pachycor
midae. because the dentition resembles that of the Jurassic
gen us Hypsocomuis.
Division Teleostei
Order Osteoglossiformes
Family, gen., ct sp. indet.
Figure 6
Disarticulated skull bones, vertebral centra, and
scales probably belonging to one individual have beenfound in one piece of gray siltstone (SBEI-1489a) (Figure6a). The bones include preopercles. a left opercle. cleith-ra. infraorbitals, a left scapula, and a neurocraniumshown in its left view, lacking the ethmoid region. In
addition, many scales, the posterior part of a para-
sphenoid, and some unidentified bones have been foundin another piece of gray siltstone (SBEI-1489c) (Figure6b). On SBEI-1489a. the sensory canal on the horizontal
arm of the preopercle runs quite straight, with the sensorycanal pores open in a shallow groove. The anteriorportion of the frontal is wider than the posterior portion.Its infraorbitals behind the orbit are large. Long teeth
are found on the parasphenoid. The scales are thick andlarge with radial grooves.
Some of the characters observed on the specimens
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 thelarge infraorbitals covering the cheek region.
Order, family, gen. et sp. indct.
Figure 7
Discovered from Kuwajima were many round or ovalsmall cycloid scales. It is certain that they belong toteleosts. possibly primitive ones. Most scales are verythin, with six to eight ridges without any grooves.
CONCLUDING REMARKS
Identification of fossil fishes
Although most of the fish fossils are isolated bones
and scales, the following taxa can be recognized. They are:
Semionotidae. Lepidotes sp.Sinamiidae, Sinamia sp.Pachycormidae. gen. et sp. indeterminable
Osteoglossiformes. family, gen. et sp. indeterminableTeleostei. order, family, gen. et sp. indeterminable
Figure 7. Cycloid scales belonging to the division Teleostei from theKuwajima Formation in Kuwajima. Ishikawa Prefecture. Japan.SBEI-298. Scales indicate 2 mm.
In this Early Cretaceous locality, fossils of the genusSinamia are the most abundant probably due to thedepositional environment (Isa.ii et a/.. 2005) favorable for
the preservation of robust bones and scales. Probablythese specimens represent two species as two different
142 Yoshitaka Yabumoto
types of gular plates and supracleithra have been found.However, it is a bit difficult to compare the material fromKuwajima to other species of the genus because of theincompleteness of the specimens. For the purpose ofcomparison the author examined the holotypes ofSinamia poyangica Su and Li, 1990, S. chinhuaensis Wei,
1976, S. huananensis Su, 1973 and S. luozigouensis Li,
1984. A few differences between these previously established species and the Kuwajima ones have been noticed.
Hyomandibular SBEl-817 differs from that of S.huananensis in the convex outline of the dorsal margin ofthe dorsal arm (it is almost straight in S. huananensis); S.chinhuaensis in its smaller process for articulation withthe opercle and lacks ridges on its lateral surface (S.chinhuaensis has the strong ridges); 5". poyangica in the
strongly convex outline of the dorsal margin of the dorsal
arm (the dorsal margin is almost straight in S. poyangica),and the posterior margin of the dorsal arm forming an
acute angle with the dorsal margin; S. zdanskyi with theacute angle between the posterior margin of the dorsal
arm and the dorsal margin of the process for articulation
with the opercle (5". zdanskyi has an obtuse angle), therelatively short ventral arm, the more concave anterior
margin, and the convex dorsal margin of the dorsal arm
(slightly convex, almost straight in S. zdanskyi).The author also examined the holotype and paratype
of S. luozigouensis, but the hyomandibular is not preserved in the types. There is a chronological difference
between the species from the Kuwajima Formation, earlyEarly Cretaceous and S. luozigouensis, late EarlyCretaceous, approximately Aptian or Albian (ChangMee-mann personal communication, 2003).
The dentary SBEI-295 which was reported to belongto a semionotiform fish by Azuma and Hasegawa
(1989), is instead placed in the amiiform genus Sinamiaon the basis of the deep coronoid process and canine-liketeeth curved inward by the present author.
The pachycormid dentary SBEI-826 is similar to thatof the Late Jurassic Hypsocormus macrodon (Wagner,1858) from Solnhofen, Germany, but the surfaces of thelarge teeth are smooth in the specimen from Kuwajima(H. macrodon has fine ridges). Pachycormids had beenreported only from marine sediments, therefore the specimen from the Kuwajima Formation represents the firstreport of the group from non-marine strata.
Significance of the fish faunaThe osteoglossi form fish (SEBI-1489a, c) from the
Kuwajima Formation is the best preserved fish fossil fromthe Tetori Group collected so far. If the author's deduc
tion that it belongs to the family Osteoglossidae is correct,then at 25 million years older than the previous oldest
record (Laeliichthys ancestralis Silva Santos, 1985 from
Brazil; Aptian) of the family, this fossil is significant forunderstanding the origin of osteoglossids. As the frontalhas wide anterior, and narrow posterior portions similarto that of the osteoglossid subfamily Phareodontinae
which consists of Phareodus, Phareoides (probably asynonym of the Phaleodus), Brychaetus, Cretophareodus,it maybe further assigned as a member of the phare-
odontins which would make it 60 million years older than
the previous oldest record (Cretophareodus alberticus Li,1996 from Canada; middle Campanian) of the subfamily.
The other Early Cretaceous freshwater fish faunafound in Japan is the Wakino fish fauna (Yabumoto,
1994). Twenty one species have been described on the
basis of the articulated specimens, and three different
assemblages have been recognized: these are Nipponamia-Aokiichthys assemblage from the Dobaru Formation
(lowest), Paraleptolepis-Wakinoichthys assemblage from
the Gamo Formation (the third formation from the bot
tom) and Diplomystus-Wakinoichthys assemblage fromthe Kumagai Formation (top). No fish remains havebeen reported from the Takatsuo Formation (the secondformation from the bottom). The Wakino fish fauna was
characterized by the abundance of "Diplomystus" andAokiichthys (Yabumoto, 1994).
The age of both sediments from Kuwajima andKitakyushu is almost the same, but except for thesemionotid the faunae are different. Probably Lepidotesis shared by both faunae. Various species of Lepidoteshave been found from China and Korea in East Asia
(Chang and Chou, 1977) and all over the world (Nel
son, 1994). However, Sinamia, which is the most abun
dant form in the Kuwajima fish fauna, has not been found
in the Wakino Subgroup.The Wakino Subgroup in Japan and the Nagdong
Subgroup of the Gyeongsang Group in Korea are considered of the same age because both have yielded the samespecies, Wakinoichthys aoki Yabumoto, 1994(Yabumoto and Yang, 2000). Sinamiid fossils havebeen found in the Nagdong Subgroup, but not in theWakino Subgroup. The composition of the Wakino fishfauna more resembles that of the fish fauna from the
southeastern part of China than that of the Jehol Group;"Diplomystus" and Aokiichthys from the Wakino Subgroup are close to Paraclupeaand Paralycoptera from thesoutheastern part of China.
The difference of the fish fauna of the Kuwajima
Formation from other areas in East Asia is probably
caused by the difference of the depositional environments.Further research and excavation of the Kuwajima Forma
tion are necessary and important for the understanding of
Cretaceous Fishes from Tetori Group, Japan 143
the Early Cretaceous of this area.
ACKNOWLEDGMENTS
The author expresses his sincere gratitude to Dr.Teruya Uyeno of the National Science Museum, Tokyo
and Dr. Chang Mee-mann of the Institute of Vertebrate
Paleontology and Paleoanthropology for their critical
reading of the manuscript and their valuable advice. He
thanks Dr. Makoto Manabe of the National Science
Museum, Tokyo and Dr. Shinji Isaji of the NaturalHistory Museum and Institute, Chiba for their advice and
information of the fossils and the fossil locality. He alsothanks Dr. Jin Fan of the Institute of Vertebrate
Paleontology and Paleoanthropology for his help toexamine the type specimens of the Sinamia in China. Hethanks Mr. Ichio Yamaguchi, Mrs. Mikiko Yamaguchi,
and Dr. Tatsuya Sakumoto of Shiramine Village for theirhelp for this study. This study was supported by theJapanese Society for the Promotion of Science Grant-in-
aid for scientific research and Fujiwara Natural HistoryFoundation.
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