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An Early Pleistocene Giant Tortoise (Reptilia ; Testudines ; Testudinidae) from the Bumiayu Area, Central Java, Indonesia

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An Early Pleistocene Giant Tortoise (Reptilia ; Testudines ; Testudinidae) from the Bumiayu Area, Central Java, Indonesia ｖｅｒ／化石研究会会誌 ＰＤＦ化／０９０７０５６６ 化石研究会誌４２巻１号／本文／１ 欧文 ０１‐１１ 2009.08.19 14.24 化石研究会会誌 Journal of Fossil Research, Vol.４２（１）,１-１１（２００９） ［Original report］ An Early Pleistocene giant tortoise (Reptilia ; Testudines ; Testudinidae) from the Bumiayu area, Central Java, Indonesia SETIYABUDI, Erick* Abstract A well-preserved specimen of a gigantic testudinid tortoise from the Early Pleistocene Kali Glagah Formation of the Bumiayu area, Central Java, Indonesia is describedindetail.TheBumiayu specimen is referable to the largest testudinid, Megalochelys sivalensis, typically known from the Plio- Pleistocene of Siwalik, Punjab, India, based on its huge size and prominent Y-shaped anterior projection of the epiplastron. Although Bumiayu specimen is identified as Megalochelys cf. sivalensis here, a few distinct characters of the former suggest the possibility of independent evolution resulting from the biogeographical isolation of the insular form from the continental population. Key words : Pleistocene, Systematics, Megalochelys cf. sivalensis, Testudinidae, Testudines, Java, Indonesia. Introduction 1935 ; Hooijer, 1954, 1971a, b, 1982 ; Sondaar, 1981). The Early Pleistocene non-marine deposits on Nevertheless taxonomic status of the Bumiayu the Bumiayu area, Central Java, Indonesia, have specimen should be reexamined, because this fossil yielded extinct testudinid tortoises (van der Maarel, has never been described in detail. Recently, I have 1932 ; ter Haar, 1934 ; Koenigswald, 1935 : Fig. 1). A an opportunity to compare the Bumiayu specimen large skeleton, from one individual (K1587-1589, GSI) with M. sivalensis and the other gigantic extinct including large part of the shell, is here referred to testudinids from different localities of Indonesia, Megalochelys sivalensis Falconer and Cautley, 1837. such as Sulawesi, Timor, and Flores Islands. In this Megalochelys sivalensis was originally described from paper, I describe the Bumiayu specimen with and the Plio-Pleistocene of Siwalik, India, and is considered discuss its taxonomic identity. Although M. sivalensis the largest land tortoise (roughly estimated as 2 m has been famous as the largest known terrestrial in straight shell length) (Lapparent de Broin, 2002). turtle since the 19th Century, its skeletal anatomy Megalochelys sivalensis is characterized by a prominent was largely obscured by its fragmentary nature of Y-shaped anterior projection of the epiplastron Indian materials. Thus, this report will throw new (Loveridge and Williams, 1957). The Bumiayu light on the morphology of this gigantic tortoise specimen has been tentatively assumed to be species. conspecific to M. sivalensis (ter Haar, 1934; Koenigswald, Received : 10th April, 2009, Accepted : 4th June, 2009 *Graduate School of Science and Engineering, Kagoshima University, Kagoshima, 890-0065, Japan E-mail : [email protected] ―1― ｖｅｒ／化石研究会会誌 ＰＤＦ化／０９０７０５６６ 化石研究会誌４２巻１号／本文／１ 欧文 ０１‐１１ 2009.08.19 14.24 120 130 140 JAKARTA J A V A S E A o 10o 10 N Bandung BUMIAYU SANGIRAN Purwokerto 0o I N D I A N S E A Java 0 53 106 km o 10 S 0 1 2 cm o 110 o 120 To Prupuk Legend L I M B AN G A N giant tortoise fossil locality River Cisaat river Road Cipamali river Kali Glagah (Glagah River) Cijajawal river Biuk river Cisaat river Cibenda river Cipelem river Slatri river ek river Gepr Ciluncat river Jurang river Lembu river Panujah river B A N T A R K A W U N G Panujah bening river Jambe river 0 2.5km cipati river Bodas river B U M I AY U Fig. 1. Map showing the locality of fossil turtle in Kali Glagah (Glagah River) of the Bumiayu area, Central Java, Indonesia. Materials and methods (AAJ) and the Rijksmuseum van Natuurlijke Historie, I examined the Plio-Pleistocene fossil testudinids Leiden, the Netherlands (RMNH). GSI gives the from Java, Sulawesi, Flores and Timor Islands of registered specimens from Java as K (Koenigswald). Indonesia. The Geological Museum, Geology Survey Bumiayu specimen consists of large part of carapace Institute, Bandung, Indonesia (GSI) keeps the fossil and plastron (K1857a), detached anterior portion of materials from Java, Sulawesi, and Flores Islands, plastron (K1857b), femora (right side : K1588a, and while other Sulawesi and Timor specimens are left side : K1588b), and pelvic girdle (K1589) of one stored in the Archaeological Agency in Jakarta individual. K1857a is so fragile that it was difficult to ―2― ｖｅｒ／化石研究会会誌 ＰＤＦ化／０９０７０５６６ 化石研究会誌４２巻１号／本文／１ 欧文 ０１‐１１ 2009.08.19 14.24 remove this large shell material from a protective Hirayama (RH) and Akio Takahashi (AT) as comparative girdle in order to take a photo from the ventral side. materials. Skeletal terminology used in this study Therefore, ventral view of the main part of plastron follows Walker (1973) and Zangerl (1969). The most was shown as a drawn illustration. The Indian of comparative characters follows Loveridge and materials of Megalochelys sivalensis were examined in Williams (1957), Crumly (1984), Gaffney and Meylan the Natural History Museum, London (BMNH as (1988), Meylan and Sterrer (2000), Takahashi et al . catalogued by Lydekker, 1889b). We observed extant (2003), and Claude and Tong (2004). testudinid skeletons in the private collections of Ren FORMATION LITHOLOGY HOLO- CENE VOLCANICS / ALLUVIAL / RIVER DEPOSITS . /. /. //. /. LINGGOPODO Fm . Andesitic conglomerate, blue grey clays. > 200 m /. /. //. /. .. /.. / ... / .. / .... / . GINTUNG Fm . / . Andesitic conglomerate, tuffaceous sandstone, 450- 800 m .. .. / .. argilaceous, sandstone, mudstone. / . / .. / . / ... ... ... OCENE / . / . /. / . / . ../ .. .. .. / / Tuff, tuffaceous sandstone intercalated with clays MENGGER Fm 150- 200 m . .. / . / . / . / / . and conglomerates. .. / . / . / / . / . / . Tuffaceous sandstone and conglomerate coarser PLEIST KALIGLAGAH Fm . .. .. in the upper part. .. 500 m . ./ ./ /. /. / . / / /. .. / . / ./. / . /. /. / KALIBIUK Fm . .. .. .... ... ... ......... ... .. .. Mudstones, intercalated with sandstone, calcare- 350 - 500 m ous sandstone and marls, molluscs. Reef locally in upper part, tuffaceous sandstone, . / . /. / . conglomerate alternation with sandy marls and TAPAK Fm . Late /.. /.. clays, molluscs. 500 m . /. / . / . /. / . / . .. .. .... .. / / / / / . / / . Andesitic breccias alternatively very coarse and / / / / / / / / / / / / / / / / / / / / coarse, intercalations of tuffs, lava, ashes, tuffs. / / / / / / / / KUMBANG Fm / / / / / / / / // / / / // / / / / / / / / / / / up to 2000 m / / / / / / / / / / / PLIOCENE / / / / / / / / / / / / / / / / / / / Early / / / / / / / / / . / . / . / . / Alternation of calcareous tuffaceous sandstone, . HALANG Fm / / / sandy marls, intercalations of micro breccia, cong- / . / . 800 m . / . lomerate and calcarenite. / . / . / . / / / / . / .. / . / ../ . "SERIES"(ter Haar,1934) 0.10-250 m Marls intercalation with calcareous tuffaceous / / ./. /. / sandstone, and alternation of marls with calcare- RAMBATAN Mb ous sandstone, tuff, tuffaceous sandstone, inter 800 m . .. .. .. .. calation of conglomerates, breccia, calcarenite, Late fibrous calcite beds. ... PEMALI Fm. Argilaceous marls intercalation with calcareous MIOCENE >800 m sandstone, tuffaceous sandstone. / / / . / / . / . / . Legend / / / . .. / / // / .. .. //. /. // ............ Sandstone / / . .. Andesite conglomerate Breccia // Tuff, lava . .. .. .. /. Clay . Calcareous sandstone Reef / / Sandy Marl . / . / . / / / Marl . / . Tuffaceous sandstone Ash ... Conglomerate . Fig. 2. Stratigraphy of the Kali Glagah Formation in the Bumiayu area (modified from Sumarso and Suparyono, 1974). X indicates the horizon of fossil turtle. ―3― ｖｅｒ／化石研究会会誌 ＰＤＦ化／０９０７０５６６ 化石研究会誌４２巻１号／本文／１ 欧文 ０１‐１１ 2009.08.19 14.24 Geological setting Megalochelys to Testudo gigantea Schweigger 1812 The Kali Glagah Formation mainly consists of from the Aldabra Islands (see Williams, 1952), which tuff and tuffaceous sandstone with intercalation of is now placed in genus Dipsochelys (Bour, 1982). clays and conglomerates in the upper part, and Megalochelys was also used as a subgenus of Testudo contains shells of Unio, Melania, Corbicula,and Linnaeus 1758 (Falconer and Cautley, 1844; Lydekker, Viviparus, plant remains, and vertebrate fossils in 1889a). The taxonomic history of Megalochelys is lower part (Sondaar, 1984; Sumarsono and Suparyono, often conflated with that of Colossochelys atlas 1974 ; Kastowo and Suwarna, 1996; Fig. 2). Mammals, Falconer and Cautley, 1844 also from the Plio- including carnivores, rodents, artiodactyls, and Plesitocene of India and Pakistan. Maack (1869) proboscideans, and reptiles such as testudinids were mentioned C. atlas as Colossochelys (Megalochelys) found from its sandstone beds (ter Haar, 1934). The atlas . Lydekker (1880) retained Colossochelys as the Kali Glagah Formation is considered to be continental subgenus of genus Testudo, and Lydekker (1885) first to shallow marine deposits (coastal or fluvio-deltaic gave a clear diagnosis and illustrations for C. atlas.A near the base, becoming increasingly terrestrial large portion of carapace of M. sivalensis from towards the top) (Kastowo and Suwarna, 1996). Siwalik, was reported as the first mounted skeleton Leinders et al . (1985) dated it at about 1.5 Ma based of Testudo atlas without proper description (Brown, on its vertebrate fauna from the base of the Kali 1931; Ren Hirayama, personal observation). The Glagah Formation. The vertebrates of this horizon generic name of Testudo, however, was once used as are classified as the Satir Fauna, the oldest mammalian a wastebasket taxon applied to most testudinids
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