pleistocene clupeid and engraulidid fishes from the

Bull. Kitakyushu Mus. Nat. Hist., 8: 55-74. December 27, 1988

Pleistocene Clupeid and Engraulidid Fishes from theKokubu Group in Kagoshima Prefecture, Japan.

Yoshitaka Yabumoto

Kitakyushu Museum of Natural History, Nishihonmachi

Kitakyushu, 805 Japan

Abstract Fish fossils were found in the Pleistocene Kokubu Group in KagoshimaPrefecture, Japan. These are identified as the extant species Sardinops melanostictus in thefamily Clupeidae and Engraulis japonicus in the family Engraulididae. Figures of thecomplete skeletons of those extant species are presented for the comparison with thefossil specimens.

Several specimens of fossil fishes yielded from the Kokubu Group at Aira Townin Kagoshima Prefecture were sent to the author by Mr. Shigeru Takaki of theKagoshima Prefectural Museum for identification in March, 1987. In September,1987, I collected two more fossils. Two species, Sardinops melanostictus and Engraulis

japonicus, are recognized among these specimens. The Kokubu Group was firstdescribed by Ida, et al. (1950) and studied in detail by Otsuka and Nishiinoue(1980) and Nishiinoue and Otsuka (1982). Otsuka and Nishiinoue (1980)recognized five stratigraphic units: the Kajiki Formation, the Nabekura pyroclasticflow deposits, the Kamo Formation, the Oda pyroclastic flow deposits and theHayato Formation in ascending order and reported the fossil fishes, Sardinopsmelanostictus from the Hayato Formation at Kuwanomaru, Yoshida Town and anunknown species of the Gobiidae from the Kajiki Formation at Ooyama, Aira Town,without description. I examined their specimens in the National Science Museumand recognized Engraulis japonicus and Sardinops melanostictus. The fossils of E.japonicus and S. melanostictus are described here, compared with bones of the extantspecies. The only fossil records of S. melanostictus and E. japonicus in Japan areisolated otoliths from the Pleistocene deposits of Boso and Miura Peninsulas (Aoki,1968). This is the first fossil record of partial and almost complete skeletons of fossilS. melanostictus and E. japonicus in Japan.

Previously there were only partial osteological descriptions of S. melanostictus andE. japonicus: craniums and vertebral columns of both species (Hotta, 1961), dorsaland anal fin rays and their pterygiophores (Kinoshita, 1984) and hypobranchialapparatus (Balart, 1985) of E.japonicus. Figures of the complete skeletons of E.japonicus and S. melanostictus are presented here for the first time for comparison withfossils. The names of the bones and anatomical elements are based on Harrington

56 Yoshiiaka Vabumoto

(1955), Ridewood (1905), Chapman (1944a, 1944b), Uyeno (1975) and Pattersonand Rosen (1977).

Acknowledgments

I am very grateful to Dr. Teruya Uyeno of the National Science Museum for hisinvaluable adviceand critical reading of the manuscript, and Dr. Hiroyuki Otsuka ofKagoshima University and Mr. Shigeru Takaki of the Kagoshima PrcfecturalMuseum for the loan of the specimens and the locality information. I am alsograteful to Dr. Takakazu Ozawa of Kagoshima University for his cooperation incollecting specimens, and to Dr. Ryuzo Toriyama and Dr. Masamichi Ota of theKitakyushu Museum of Natural History for their constant encouragement.

Localities and Horizon

The locality of fish fossils at Ooyama (longitude 130°37'38"E. and latitude31°45'77"N.) is in Aira Town (Fig. 1). This locality belongs to the Kajiki Formationwhich is the lowest unit in the Kokubu Group. The locality of fish fossils atKuwanomaru (130°33'58"E. and 31°44'l9"N.) is in Yoshida Town (Fig. 2). Thislocality belongs to the Hayato Formation which is the uppermost unit in the KokubuGroup. Both towns are located at the northern part of Kagoshima Prefecture, about

t.o Kajiki Kagoshima

Fig. 1 A map showing the locality (X) for the fossil fishes from the Kajiki Formation atOoyama, Aira Town in Kagoshima Prefecture.

Pleistocene Clupeid and Engraulidid Fishes from Kagoshima 57

to Kagoshima

Fig.2. A map showing the locality (X) for the fossil fishes from the Hayato Formation atKuwanomaru, Yoshida Town in Kagoshima Prefecture.

20 km north of Kagoshima City. The age of the Kokubu Group is Early Pleistocene(Otsuka and Nishiinoue, 1980).

Class Osteichthyes

Order ClupeiformesFamily Clupeidae

Sardinops melanostictus (Temminck et Schlegel, 1846)

Material: KMNH (Kitakyushu Museum of Natural History) VP100,138,almost complete specimen with lateral side exposed; KMNH VP100,139, almostcomplete specimen with rightsideexposed. Both specimens are bent at the centerofthe body, collected from the Kajiki Formation at Ooyama, Aira Town by the author.ESK(Institute of Earth Sciences, Faculty of Science, Kagoshima University) 6053,scales, collected from the Hayato Fqrmation at Kuwanomaru, Yoshida Town by Dr.Hiroyuki Otsuka.

Description: In the specimen of KMNH VP 100,138, there are 19 dorsal finrays, located near the middle of the body (Plate 1). Three abdominal scutes in frontof the pelvic, one under' the pelvic and 7 behind the pelvic are observable. Thenumber of vertebrae is 48. Three ridges are present on the opercle. The quadrate,angular, preopercle and hyomandibuler are partially preserved and these bones are

58 Yoshitaka Yabumoto

ang

den

max

supmlsupm2 ret <*»

Fig. 3. Sardinops melanostictus: A, head region of the fossil specimen of KMNH VP 100,138;B, head bones of the Recent specimen, KMNH VR100, 094, 92mm in standard length,collected at Wakamatsu, Kitakyushu, Fukuoka Prefecture, 18 October, 1980. ang,angular; den, dentary; ecp ectopterygoid; enp, endopterygoid; hyo, hyomandibular; ino,interopercle; max, maxillary; met, metapterygoid; ope, opercle; pal, palatine; prem,premaxillary; preo, preopercle; qua, quadrate; ret, retroarticular; subop, subopercle ;supml, first supramaxillary; supm2, second supramaxillary; sym, symplectic. Cartilageis indicated by hatching.

Pleistocene Clupeid and Engraulidid Fishes from Kagoshima

WW

Fig. 4. Pleistocene Sardinops melanostictus: A, middle part of the body, KMNH VP 100,138;B, scales, ESK 6053.

hyu5

mcfr

2mm

59

Fig. 5. Caudal bones of a Pleistocene Sardinops melanostictus, KMNH VP100.139. hyu,hypural; mcfr, middle caudal fin rays; parh, parhypural bone.


similar to those of the extant species (Fig. 3). Some scales are preserved, which haveseveral grooves from the ventral and dorsal margin. These grooves are notcontinuous in the middle part of the scales (Fig. 4). Two middle caudal fin raysextend over the hypurals.

In the specimen of KMNH VP100,139, the number of dorsal rays is 17. Thenumber of vertebrae is 47. The posteroventral corner of the ceratohyal is extendedposteroventrally. The epihyal is large and the length is about three quarters of theceratohyal. Two middle caudal rays extend over the hypurals. A short ridge ispresent at the anterior part of the first hypural (Fig. 5).

In the specimen of ESK 6053, thirteen scales are preserved on a mudstone.The posterior parts are missing in some scales. Five or six grooves extend from theventral and dorsal margin to the middle of the scales and only the most posteriorgroove is continuous (Plate 1 and Fig. 4).

Family Englaulididae

Engraulis japonicus (Houttuyn, 1872)

Material: KM(Kagoshima Prefectural Museum)-F-87-l, almost complete specimen with left side exposed, but a part of the dorsal fin, opercular bones andsuspensorium are missing, collected from the Kajiki Formation at Ooyama, Aira

Town. ESK 6054, anterior part of the body, with ventral side exposed, collected

from the Hayato Formation at Kuwanomaru, Yoshida Town by Dr. HiroyukiOtsuka.

prev

Fig.6. Head region of a Pleistocene Engraulis japonicus, KM-F-87-1. bra, branchiostegal;chy, ceratohyal; cle, cleithrum; era, coracoid ; eth, ethmoid; gra, gill raker; hyph,hypohyal; prev, prevomer; P^ pectoral fin.


prem

Fig. 7. Head region of a Pleistocene Engraulisjaponicus, ESK 6054. Abbreviations see Fig. 3.

Description: In the specimen of KM-F-87-1, the mesethmoid bone projectsbeyond the prevomer. The ceratohyal is long with many short branchiostegal rays(Fig. 6). The dorsal fin is situated near the middle of the body. The anal base islong with 14 fin rays. A part of the gill arch, long gill rakers, shoulder girdle andpectoral fins are observable. These are similar to those of the extant species (Figs. 6,15, 16 and Plate 2). In the specimen of ESK 6054, the maxillaries and dentary arelong and bear small teeth. The preopercle is narrow. The suspensorium inclinetoward the rear (Fig. 7). The preserved body length is 50.9 mm. Thirteen dorsalrays, pterygiophores, and 31 vertebrae are observable. The abdominal scute isabsent. The dorsal fin is situated near the middle of the body.


Fig. 8. Scales of an extant Sardinops melanostictus, KMNH VR100, 095, 158 mm in standardlength, collected from Kokura, Kitakyushu, 10 April, 1988. Roman numerals indicatethe rows of scales and Arabic numerals are numbered from top downword.


prev para ors sph pro inca

pto

10mm

Fig. 9. Cranium of an extant Sardinops melanostictus, KMNH VRIOO 096, 159 mm instandard length, collected at Kokura, Kitakyushu, 10 April, 1988. A, dorsal view; B,lateral view; C, ventral view, bao, basioccipital; bsp, basisphenoid; epo, epiotic; eth,ethmoid; exo, exocciptial; fro, frontal; inca, intercalar; ors, orbitosphenoid; para,parasphenoid; pari, parietal; pref, prefrontal; prev, prevomer; pro, prootic; pte,pterosphenoid; pto, pterotic; supo, supraoccipital; sph, autosphenotic. Cartilage isindicated by hatching.


supml supm2

Fig. 10. Extant Sardinops melanostictus, KMNH VR100, 096. A, upper jaw; B, lower jaw;C, shoulder girdle; D,ventral view of theshoulder girdle, act, actinost; ang, angular;cle, cleithrum; era, coracoid; den, dentary; max, maxillary; mes, mesocoracoid; poc lo,lower postcleithrum; poc up, upper postcleithrum; pot, posttemporal; prem, premaxil-lary; ret, retroarticular; scap, scapula; supc, supracleithrum; supml, first supramaxillary; supm2, second supramaxillary.

Ahyph up

Pleistocene Clupeid and Engraulidid Fishes from Kagoshima

ghav ephinh

Fig. 11. Extant Sardinops melanostictus, KMNH VRIOO, 096 A, hyoid arch; B, urohyal;C, gill arches, bab, basibranchial; bah, basihyal; bra, branchiostegal; cbr, ceratobran-chial; chy, ceratohyal; epb, epibranchial; eph, epihyal; gha, groove for hyoidean artery;hyp, hypobranchial; hyph lo, lower hypohyal; hyph up, upper hypohyal; inh, interhyal;pha lo, lower pharyngeal; pha up, upper pharyngeal, suprp, suprapharyngobranchial.Cartilage is indicated by hatching.

65


Fig. 12. Extant Sardinops melanostictus, KMNH VRIOO, 096 A, dorsal fin; B, anal fin; C,pelvic girdle; D, caudal skeleton, bap, basipterygium; epu, epural; hes, haemal spine;hpap, hypurapophysis; hyu, hypural; lep, lepidotrichia; nes, neural spine; parh,parhypural bone; preu, preural centrum; ptr dis, distal pterygiophore; ptr med, medianpterygiophore; ptr pro, proximal pterygiophore; urn, uroneural bone; v ura, uralcentrum. Cartilage is indicated by hatching.


Fig. 14. Cranium of an extant Engraulis japonicus, KMNH VRIOO, 098, 59 mm in standardlength, collected at Kaie, Kaimon, Kagoshima Prefecture, 28 October, 1987. A, dorsalview; B, lateral view; C, ventral view, supt, supratemporal. Other abbreviations see

Fig. 9.


1mm 5mm

Fig. 15. Extant Engraulis japonicus, KMNH VRIOO, 098 A, suspensorium and opercularbones; B, upper jaw; C lower jaw; D, shoulder girdle; E, ventral view of theshoulder girdle. Abbreviations see Figs. 3 and 10.

Remarks

In the extant species of Sardinops melanostictus and Engraulis japonicus, the first ribsare attached to the third vertebra. The number of the dorsal pterygiophores is twofewer than that of dorsal fin rays. This information is useful in determining thenumbers of vertebrae and dorsal fin rays in fossil specimens.

The following characters indicate that the fossil specimens are members of theteleostean fish order Clupeiformes; 1) the dorsal fin is situated near the middle of the


Fig. 16. Extant Engraulis japonicus, KMNH VR100, 098.

gill arches. Abbreviations see Fig. 11.

suprp

suprp

A, hyoid arch; B, urohyal; C,

body; 2) the pelvic fin is situated near the middle of the body and below the dorsalfin; 3) two middle caudal rays extend over the hypurals.

The age of the deposits from which the fossil specimens are yielded is EarlyPleistocene (Otsuka and Nishiinoue, 1980). Twenty-one speices in 3 families of theorder Clupeiformes have been recorded in Japan (Masuda etal. 1988). Characteristic and meristic features of the fossil specimens agree well with those of Sardinopsmelanostictus and Engraulis japonicus.

The specimens of KMNH VP100,138, KMNH VP100,139 and ESK 6053 are


ptr pro

v ura2

Fig. 17. Extant Engraulis japonicus, KMNH VRIOO, 098. A, dorsal fin; B, anal fin; C,pelvic girdle; D, caudal skeleton. Abbreviations see Figs. 5 and 12.

identified as Sardinops melanostictus, on the basis of following characters: 1) theabdominal scutes are present (KMNH VP100,138); 2) three ridges are presenton theopercle (KMNH VP100,138); 3) the grooves on each scale are curved and extendfrom the dorsal and ventral margin to the middle, and are not continuous at themiddle (KMNH VP100,138 and ESK 6053); 4) the numbers ofvertebrae (47 and 48)

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Fig

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.E

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tE

ngra

ulis

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s,K

MN

HV

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0,09

8.


and dorsal rays (17 and 19) are in the range of those of the extant species (KMNHVP100,138 and 100,139); 5) the posteroventral corner of the ceratohyal is extendedposteroventrally and the length of the epihyal is about three quarters of theceratohyal (KMNH VP100,139).

The diagnostic characters of engraulidid fish are the oblique (posterior)inclination of the suspensorium and the projected mesethmoid bone beyond thevomer supporting a paired rostral organ (Nelson, 1984b; Grande and Nelson,1985). The specimens of KM-F-87-1 and ESK 6054 are identified as Engraulis

japonicus, on the basis of the following characters: 1) abdominal scute is absent; 2) theceratohyal is long and stout with many short branchiostegal rays (KM-F-87-1); 3)the mesethmoid bone projects beyond the vomer (KM-F-87-1); 4) the long maxillaryand dentary bear small teeth (ESK 6054); 5) the narrow preopercle and the obliquelyinclined suspensorium (ESK 6054); 6) the number of dorsal rays (15) is in the rangeof the extant species E.japonicus (Nelson, 1984a). Both species are now distributedin the waters of Kagoshima Prefecture.

Literature Cited

Aokj, N. 1968. Some Pleistocene fish-otoliths from the Boso and Miura Peninsulas. Trans. Proc.Palaeont. Soc. Japan. 71 (N. S.): 296-307. figs. 1-14.

Balart, E. F. 1985. Osteological development of the hyobranchial apparatus in Engraulis japonicus.Bull.Japan. Soc. Scient. Fish., 51 (4): 515-519, figs. 1-3.

Chapman, W. M. 1944a. The osteology of the Pacific deep bodied anchovy, Anchoa compressa. J.MorphoL, 74 (2): 311-329, figs. 1-15.

Chapman, W. M. .1944b. The comparative osteology of the herring-like fishes (Clupeidae) ofCalifornia. California fish and Game, 30 (1): 6-21, figs. 1-19.

Grande, L. and G. Nelson. 1985. Interrelationships of fossil and recent anchovies (Teleostei:Engrauloides) and description of a new species from the Miocene ofCyprus. Amer. Mus. Novitates,(2826): 1-16, figs. 1-12.

Harrington, R. W. Jr. 1955. The osteocranium of the American cyprinid fish, Notropis bifrenatus,with an annotated synonymy of teleost skull bones. Copeia, 1955 (4): 267-290, figs. 1-8.

Hotta, H. 1961. Comparative study of the axial of Japanese teleostei. Nippon GyogakuShinkokai, Tokyo, 155 pp., 70 pis.

Houttuyn, M. 1872. Beschrijving van eenige Japanische visschen en andere zee-schepzelen. Verh.Holl. Maatsch. Wet. Haarlem, 20 (2): 1-18.

Ida, K., S. Shinoyama, K. Saitou, and K. Kato. 1950. Natural gas in Shikine gas field,

Kagoshima Prefecture. Bull. Gtol. Surv. Japan, 1(2): 9-14, figs. 1-7. (in Japanese with Englishabstract)

Kinoshita, T. 1984. Dorsal and anal fin rays of the Japanese anchovy, Engraulis japonica, and theirpterygiophores. Bull. Fax. Fish. Hokkaido Univ., 35 (2): 66-82, figs. I-10.

Masuda, H., K. Amaoka, C. Araca, T. Uyeno, and T. Yoshino (ed.). 1988. The fishes of theJapanese Archipelago, Text. Tokai Univ. Press, xxii+456 pp., 247 figs.

Nelson, G. 1984a. Identity of the anchovy Hildebrandichthys setiger with note on relationships and

biogeography of the genera Engraulis and Cetengraulis. Copeia, 1984 (2): 422-427, figs. 1-3.Nelson, G. 1984b. Notes on the rostral organ of anchovies (family Engraulidae). Japan. J. Ichthyol.


31 (1): 86-87, fig. 1.

Nishiinoue, T. and H. Otsuka. 1982. Pollen stratigraphy of the Kokubu Group in South Kyushu,Japan. Rep. Fac. Sci. Kagoshima Univ. {Earth Sci. & Biol.), (15): 89-100, figs. 1-4, pis. 1-2. (inJapanese with English abstract)

Otsuka, H. and T. Nishiinoue. 1980. Quaternary geology of the coastal area north of Kagoshimabay, south Kyushu, Japan. Rep. Fac. Set., Kagoshima Univ. (Earth Sci. & Biol.), (13): 35-76, figs.1-7, pis. 1-3. (in Japanese with English abstract)

Patterson, C. and D. E. Rosen. 1977. Review of Ichthyodectiform and other Mesozoic teleostfishes and the theory and practice of classifying fossils. Bull. Amer. Mus. Nat. Hist., 158 (2): 81-172, figs. 1-54.

Ridewood, W. G. 1905. On the cranial osteology of theclupeoid fishes. Proc. Zool. Soc. London, 1905(1): 448-493, figs. 1-25.

Temminck, C. and H. Schlegel. 1842-1850. Pisces. Siebold's Fauna Japonica, 323pp., 144pls.,Leiden.

Uyeno, T. 1975. Pisces, pp. 181-242. in Shikama, T., (ed.), Paleontology, III, Asakurashoten,Tokyo, iv+7+527pp. (in Japanese)

Pleistocene Clupeid and Engraulidid Fishes from theKokubu Group in Kagoshima Prefecture, Japan.

Yoshitaka Yabumoto

Explanation of Plates 1-2.

Explanation of Plate 1.

The fossil specimens of Sardinops melanostictus from Kagoshima Prefecture."

A. KMNH VP100,138, from Ooyama, Aira Town.B. KMNH VP100.139, from Ooyama, Aira Town.

C. ESK 6053 from Kuwanomaru, Yoshida Town.

Scale bars indicate 10 mm.

Yabumoto, Y. Pleistocene Clupeid and Engraulidid Fishes from Kagoshima Plate 1

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Plate 2

Explanation of Plate 2.

The fossil specimens of Engraulis japonicus from Kagoshima Prefecture.

A. KM-F-87-1, from Ooyama, Aira Town.

B. head region, KM-F-87-1.C. ESK 6054, from Kuwanomaru, Yoshida Town.

Scales of A and C indicate 10 mm. Scale of B indicates 5 mm.

Yabumoto, Y. Pleistocene Clupeid and Engraulidid Fishes from Kagoshima Plate 2

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pleistocene clupeid and engraulidid fishes from the

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