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First Cretaceous Fish Fauna from Malaysia Journal of Vertebrate Paleontology ISSN: 0272-4634 (Print) 1937-2809 (Online) Journal homepage: https://www.tandfonline.com/loi/ujvp20 First Cretaceous fish fauna from Malaysia Yu He Teng, Masatoshi Sone, Ren Hirayama, Masataka Yoshida, Toshifumi Komatsu, Suchada Khamha & Gilles Cuny To cite this article: Yu He Teng, Masatoshi Sone, Ren Hirayama, Masataka Yoshida, Toshifumi Komatsu, Suchada Khamha & Gilles Cuny (2019): First Cretaceous fish fauna from Malaysia, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2019.1573735 To link to this article: https://doi.org/10.1080/02724634.2019.1573735 Published online: 01 May 2019. Submit your article to this journal View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=ujvp20 Journal of Vertebrate Paleontology e1573735 (14 pages) © by the Society of Vertebrate Paleontology DOI: 10.1080/02724634.2019.1573735 ARTICLE FIRST CRETACEOUS FISH FAUNA FROM MALAYSIA YU HE TENG,1 MASATOSHI SONE, *,1 REN HIRAYAMA, 2 MASATAKA YOSHIDA,2,3 TOSHIFUMI KOMATSU,4 SUCHADA KHAMHA,5 and GILLES CUNY 6 1Department of Geology, University of Malaya, Kuala Lumpur 50603, Malaysia, [email protected]; 2School of International Liberal Studies, Waseda University, Nishiwaseda 1-6-1, Shinjuku, Tokyo 169-8050, Japan; 3Graduate School of Science, the University of Tokyo, 7-3-1 Hongo, Bunkyo, Tokyo 113-8654, Japan; 4Faculty of Advanced Science and Technology, Kumamoto University, 2-39-1 Kurokami, Chuo-ku, Kumamoto 860-8555, Japan; 5Palaeontological Research and Education Centre, Mahasarakham University, Khamrieng, Kantharawichai, Maha Sarakham 44150, Thailand; 6Université Claude Bernard Lyon 1, ENS de Lyon, CNRS, UMR 5276 LGL-TPE, F-69622, Villeurbanne, France ABSTRACT—Cretaceous fish fossils are reported from Malaysia for the first time. They were found with dinosaur and turtle remains in nonmarine sediments in the interior of Pahang State, Peninsular Malaysia. This fish assemblage consists mostly of isolated teeth, with minor amounts of noncranial remains, including vertebral centra, scales, coprolites, and hybodont fin spines. Over 100 fish teeth were examined. Nine taxa were confirmed from this fish assemblage: six (identified and unidentified) species of hybodont sharks, Heteroptychodus kokutensis, Isanodus paladeji, Lonchidion aff. khoratensis, Mukdahanodus aff. trisivakulii, Egertonodus sp., and Hybodontidae indet., and three species of actinopterygians (ray-finned fishes): ‘Lepidotes’ sp., Halecomorphi indet., and Ginglymodi indet. This fish assemblage has strong affinities with Early Cretaceous (Barremian–early Aptian) faunas of Thailand, because the four hybodont species, H. kokutensis, I. paladeji, L. khoratensis, and M. trisivakulii, were previously known only from the Sao Khua Formation (Khorat Group) and equivalent strata of Ko Kut (Kut Island). Egertonodus has been confirmed in Asia for the first time. Overall, this fish assemblage shows a close linkage to Early Cretaceous (Barremian–early Aptian) faunas of Thailand. Based on faunal composition and biostratigraphic correlation, we suggest a Barremian–early Aptian age for this new fauna from Malaysia. Citation for this article: Teng, Y. H., M. Sone, R. Hirayama, M. Yoshida, T. Komatsu, S. Khamha, and G. Cuny. 2019. First Cretaceous fish fauna from Malaysia. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2019.1573735. INTRODUCTION GEOLOGICAL SETTING For the past 20 years, our knowledge of Cretaceous freshwater The fossil-bearing unit is informally referred to as ‘the Pahang fishes in Southeast Asia has been based mainly on finds from the vertebrate bed’ in this report (Fig. 1). The unit consists of thick Khorat Basin in northeast Thailand. Abundant isolated fish teeth fine-grained sandstones at the base, fining upward into medium- and scales have been reported from the Khorat Basin (e.g., grained sandstone-siltstone laminations. Small-scale tabular Cappetta et al., 1990, 2006; Cuny et al., 2003, 2006, 2008, 2009, cross-bedding is common in some sandstone beds. The presence 2014; Khamha et al., 2016). On the other hand, no information of dinosaurs, terrestrial turtles, and some plant fossils in the unit has been available from the other Cretaceous nonmarine basins suggests that the depositional site might have been near to a ter- in the region. restrial environment where these organisms lived. The absence of In 2014, the University of Malaya’s paleontological team marine fossils suggests that the depositional environment had reported the first discovery of Cretaceous nonmarine vertebrate little to no marine influence. Based on the Early Cretaceous fossils from the state of Pahang, Peninsular Malaysia. This age of the Pahang vertebrate bed, we consider that it is correlated included the first dinosaurs (spinosaurid and ornithischian) to the Early Cretaceous part (Temus Shale) of the Tembeling from Malaysia (Sone et al., 2015). Together with the dinosaurs, Group. The depositional environment of the Tembeling Group abundant fossils of freshwater fish and terrestrial turtle remains is generally accepted to be fluvial-lacustrine (Koopmans, 1968; were recovered, including hybodont sharks. This paper aims to Khoo, 1977; Ahmad et al., 1991; Shamsudin and Morley, 1994; systematically study this fish assemblage and to interpret its Ainul et al., 2005). likely age based on biostratigraphic correlation. Traditionally, most Mesozoic nonmarine sediments in Malaysia Institutional Abbreviations—PRC, Palaeontological Research were dated using plant macrofossils, palynomorphs, and/or fresh- and Education Centre, Mahasarakham University, Maha Sarakham, water bivalves, for example, the plant Gleichenoides gagauensis, Thailand; TF, Thai Fossil specimens, Sirindhorn Museum, Kalasin, the bivalve Trigonioides sp. (Koopmans, 1968), and spores and Thailand; UM, Department of Geology, University of Malaya, pollen of Clavatipollenites sp., Classopollis sp., and Circulina sp. Kuala Lumpur, Malaysia. (Khoo, 1977; Shamsudin and Morley, 1994; Ainul et al., 2005; Uyop et al., 2007). These earlier studies suggested an Early Cre- taceous age for those nonmarine Mesozoic sediments in Malaysia. Due to our agreement with Pahang State Government to *Corresponding author. protect the fossiliferous site, the exact location of the sampling Color versions of one or more of the figures in the article can be found outcrop cannot be provided in this report. The Global Positioning online at www.tandfonline.com/ujvp. System (GPS) information of the location is kept at the Published online 01 May 2019 Teng et al.—First Cretaceous fish fauna from Malaysia (e1573735-2) FIGURE 1. A, map showing the distribution of Jurassic–Cretaceous nonmarine sediments in Southeast Asia. Information combined from Lovatt Smith et al. (1996), Yunus (2014), and Hinthong (1999). B, sedimentary log of the fossil locality with the position of the reported materials. University of Malaya. Any scientific inquiry concerning this fossil magnification of ×1–2 aided by a teleconverter. The lighting deposit can be referred to the Department of Geology, University sources used were a Nikon fiber optic illuminator and a Nikon of Malaya, Kuala Lumpur, or to the Pahang State Government, ring light. To take images at different depths of field, step size Kuantan, or directly to any of the authors. was determined by focusing the top and bottom of the specimens. The gaps were then divided into an average of eight steps. The raw images were aligned and stacked using graphic design soft- MATERIALS AND METHODS ware program CombineZP. The fossil fish assemblage from the Pahang vertebrate bed con- Another method was used for imaging smaller teeth with sists mostly of isolated teeth, many of which show signs of lengths less than 5 mm. A field emission scanning electron micro- abrasion. This implies that they were likely transported for scope (FESEM) was used for taking photomicrographs of the some distance prior to deposition. Rare fish scales, coprolites, specimens. The acceleration voltage used was 15.0 kV, and the hybodont fin spines, and osteichthyan (bony fish) vertebral working distance used ranged from 12 to 27.7 mm. Specimens centra also occur in this deposit. were pre-coated with gold by using a Leica EM SCD005 sputter All fossils were obtained from the field by surface collecting coater. and bulk sampling. Fossil preparation was performed chiefly using a mechanical engraver, knives, and sharp-pointed tools under a stereomicroscope. Wherever necessary, both liquid and SYSTEMATIC PALEONTOLOGY gel Paraloid B-72 thermoplastic resin and cyanoacrylate glue were applied to prevent fragmentation. Screen-sieving was Phylum CHORDATA Haeckel, 1874 attempted, but it was not an effective method to free the fossils Class CHONDRICHTHYES Huxley, 1880 from the strongly cemented sandstone. More than 100 isolated Subclass ELASMOBRANCHII Bonaparte, 1838 fish teeth from both hybodont sharks and actinopterygians (ray- Order HYBODONTIFORMES Patterson, 1966 finned fishes) were examined for this study. Family HYBODONTIDAE Owen, 1846 For imaging fish teeth, we used two different methods depend- HYBODONTIDAE indet. ing on the specimen size. For specimens larger than 0.5 mm in (Fig. 3A, B) length, a focus stacking technique was applied. Images at differ- ent depths of field were taken and then were combined to produce a focus-stacked image in which the whole specimen is Hybodus sp. Cuny, Laojumpon, Cheychiw, and Lauprasert, in focus (Brecko et al., 2014). We used a Nikon D300 digital 2010:416,
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