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The oldest record of Alligator sinensis from the Late Pliocene of Western Japan, and its biogeographic implication

https://doi.org/10.1016/j.jseaes.2016.04.017Get rights and content

Highlights

  • The oldest record of Alligator sinensis is reported from the Pliocene of Japan.

  • Japanese A. sinensis is large in size (201 cm).

  • They might be wiped out due to tectonic and climatic changes in the Plio-Pleistocene.

Abstract

The late Cenozoic fossil record of alligators in East Asia is crucial in understanding the origin and past distribution of Asian alligators that are now represented by a single species, Alligator sinensis. This study reports a partial skeleton of A. sinensis from the Late Pliocene (approximately 3.0 Ma) of western Japan. This Japanese A. sinensis is large in size (>200 cm total length), comparable to the maximum size of extant individuals. It demonstrates the oldest record of A. sinensis and wider distribution of this species in the past. Tectonic and geographic history of East Asia suggests that alligators presumably dispersed into Japan before 25 Ma or after 10 Ma, yet finally were wiped out from Japan due to the semi-isolated condition of the Japanese island arc and the deteriorated climate during the Plio-Pleistocene.

Introduction

Alligator sinensis is the only extant alligatorid species found outside the Americas. It is one of the critically endangered species of the world’s crocodylians and is now restricted to a small area in Anhui Province, China (Jiang, 2010, Thorbjarnarson and Wang, 1999). Fossil and archaeological evidence (Fig. 1; Table 1) indicates that its past distribution extended north to Wangyin, Shandong Province, China (Fig. 1: #10), and south to Penghu Channel, west of Taiwan (Fig. 1: #4). If the potential A. sinensis materials are included, the distribution is extended further south to Ban Si Liam, Nakhon Ratchasima Province, Thailand (Fig. 1: #1). Alligator bones and skins also have been collected in other Neolithic and Bronze Age sites of China (Dawenkou cultural site in Tai’an, Shandong Province; Longshan cultural sites in Linqu and Sishui, Shandong Province, and Linfen, Shanxi Province; Shang Dynasty royal tombs in Anyang, Henan Province), although those remains were related to burials of high-ranked people and could have been brought from other regions (Liu, 1994, Thorbjarnarson and Wang, 2010). However, compared with the solid archaeological evidence, fossil records of A. sinensis hinting the origin and past distribution of the species are still limited; the sole definitive A. sinensis is a skull from the Late Pleistocene of Taiwan (Shan et al., 2013).

In late 1990s, a series of excavations of the Pliocene Tsubusagawa Formation at Ajimu-machi, Usa City, Oita Prefecture, Japan, yielded numerous vertebrate fossils including fishes (Cyprinidae and Bagridae), a salamander (Cryptobranchidae), crocodylians (Alligatoridae and Crocodylidae), turtles (Platysternidae, Trionychidae, and Bataguridae), birds (Pelecaniformes, Falconiformes, Anseriformes, and Gruiformes), and mammals (Stegodontidae, Cervidae, Ursidae, and Rhinocerotidae) (Aoki, 2001, Hirayama, 2001, Kato, 2001, Matsui et al., 2001, Matsuoka, 2001, Nakajima and Kitabayashi, 2001, Takahashi and Kitabayashi, 2001, Watanabe and Kitabayashi, 2001). This fossil assemblage was named “Ajimu fauna” (Takahashi, 2001). Aoki (2001) suggested the presence of A. sinensis among the Ajimu fauna based on fragmentary cranial and postcranial elements but did not provide a thorough description and a taxonomic assessment. Because any fossil A. sinensis or its potential kin from the Cenozoic of East Asia is crucial in the study of Alligator biogeography, we here provide a detailed description and a firm basis for the taxonomic identification of the Ajimu Alligator and discuss its biogeographic significance.

Section snippets

Geological setting

The Tsubusagawa Formation is distributed over Ajimu-machi and Innai-machi, Usa City, Oita Prefecture in Kyushu Region, and consists of lacustrine (Iwauchi and Hase, 1986, Matsumoto et al., 1984), fluvial, and marsh deposits (Satoguchi, 2001) intercalated by numerous volcanic ash beds. Iwauchi and Hase (1986) divided the formation into three parts: the lower part characterized by peaty mudstone with lignite beds and tuffaceous sand–siltstone; the middle part characterized by massive mudstone and

Systematic paleontology

  • Crocodylia Gmelin, 1789 (sensu Clark in Benton and Clark, 1988).

  • Alligatoridae Cuvier, 1807 (sensu Brochu, 2003).

  • Alligatorinae Gray, 1844 (sensu Brochu, 2003).

  • Alligator sinensis Fauvel, 1879.

Material: left quadrate (LBM 0142000027), left dentary (LBM 0142000753), left angular (LBM 0142000754), left articular (LBM 0142000756). The left mandible elements belong to a single individual.

  • Crocodylia indet. Gmelin, 1789 (sensu Clark in Benton and Clark, 1988).

  • cf. Alligator sinensis Fauvel, 1879.

Alligator sinensis Fauvel, 1879

The left quadrate (Fig. 3A–E) is nearly complete. Dorsally near the quadrate condyles, a scarf-joint suture for the quadratojugal is transversely broad and tapers anteriorly and becomes a butt-joint suture. In lateral view, a ventral half of the otic aperture and lateral wall of the cranio-quadrate passage are preserved. Anterior to the otic aperture, a large preotic foramen is present within a shallow depression. Near the caudal end of the cranio-quadrate passage, quadrate foramen aëreum is set

Taxonomic status of the Ajimu specimen

Aoki’s (2001) identification of A. sinensis from Ajimu is based on the following characters in the fragmentary skull and postcranial elements: non-significant “twist” of medial and lateral quadrate hemicondyles, referring to the relatively shallower medial hemicondyle; laterally displaced quadrate foramen aëreum; jugal width widest at the level of the anterior end of orbit; absence of longitudinal ridge on the lateral surface of jugal; dorsal margin of jugal downturned like “cutting edge” at

Acknowledgements

We express our sincere gratitude to E. Kitabayashi and all the people who engaged in the laborious excavation work and the management of the Ajimu fauna collection. We are grateful to E. Kitabayashi, M. Kitada, M. Tanimoto, K. Watanabe, and K. Yamamoto for discussion, H.-y. Shan and M.-l. Yang for their help during the visit to NMNS, Taichung, Taiwan. K. Chiba helped locate some of the literature. Comments from two anonymous reviewers significantly improved the manuscript. This work was

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