Original articleA review of theropod dinosaurs from the Late Jurassic to mid-Cretaceous of Southeast AsiaUne revue des dinosaures théropodes du Jurassique supérieur au Crétacé moyen en Asie du Sud-Est
Introduction
The most abundant and diverse non-avian theropod fossils found in Southeast Asia are from Thailand. Most of them are from the Sao Khua Formation, but also can be found in the Khok Kruat and Phu Kradung formations. Some of these finds were studied almost twenty years ago, whereas others have been discovered recently and need careful study. In this paper, an overview of theropod dinosaurs during the Mesozoic in Southeast Asia is provided, with a specific emphasis on new specimens and information on theropods found in Thailand.
The discovery of non-avian theropod dinosaurs from Thailand started in the late 1980s. Until now, eleven non-avian theropods and a Mesozoic bird have been reported. The theropods pertain to various groups and date from the Late Jurassic to the Early Cretaceous of northeastern Thailand (ca. 160–113 million years ago). There are two metriacanthosaurids (= sinraptorids) from the Phu Kradung Formation, which may be Late Jurassic or Early Cretaceous in age (prior to 125 million years ago). The finds consist of a lower leg (left tibia) of a metriacanthosaurid and cranial elements of another metriacanthosaurid (Buffetaut and Suteethorn, 2007, Chanthasit and Suteethorn, 2013). Seven theropods plus a bird from the Early Cretaceous Sao Khua Formation (ca. 130–125 million years ago) include a compsognathid (Buffetaut and Ingavat, 1984); several teeth of a spinosaurid (Siamosaurus suteethorni, Buffetaut and Ingavat, 1986); the postcranial skeleton of an ostrich-mimic dinosaur (Kinnareemimus khonkaenensis, Buffetaut et al., 2009); a maxilla of a carcharodontosaurid (Buffetaut and Suteethorn, 2012); the putative basal tyrannosauroid Siamotyrannus isanensis (Buffetaut et al., 1996), for which new studies suggest that it may not be a tyrannosauroid but an allosauroid (e.g. Rauhut, 2003a, Carrano et al., 2012) or that it could be a basal coelurosaur (Samathi, 2013, Samathi and Chanthasit, 2017); a Mesozoic bird (Buffetaut et al., 2005a); and two newly reported mid-sized, probable megaraptoran coelurosaurs, which look similar to Fukuiraptor from Japan (Samathi et al., 2019). Two theropods from the Early Cretaceous Khok Kruat Formation (ca. 125–113 million years ago) include an undescribed partial postcranial skeleton of a spinosaurid (Buffetaut et al., 2005b, Milner et al., 2007) and an undescribed partial skull and postcranial skeleton of a carcharodontosaurian (Azuma et al., 2011).
Other theropods from Southeast Asia have been recovered from Laos, Malaysia, and Myanmar. These include one spinosaurid from Laos named Ichthyovenator laosensis from the “Grès supérieurs” Formation reported in 2012 (Allain et al., 2012, Allain, 2014) and two spinosaurid teeth from the late Early Cretaceous of the Malay Peninsula (Sone et al., 2015). One coelurosaur and several enantiornithine birds preserved in amber were also recently reported from the mid-Cretaceous of Myanmar (Xing et al., 2016a, Xing et al., 2016b, Xing et al., 2017, Xing et al., 2018, Xing et al., 2019). Theropods from the Xinlong Formation of southern China, which belongs to the same paleobiogeographic province as Southeast Asia, includes the carcharodontosaur Datanglong guangxiensis (Mo et al., 2014b) and several teeth of carcharodontosaurids and spinosaurids (Buffetaut et al., 2008, Mo et al., 2014a, Mo et al., 2016).
Here, we review theropod dinosaurs found on the Southeast Asian mainland, as well as southern China, focusing mainly on Thailand from where most of the theropod faunas have been reported. We discuss their affinities based on recent information and recent finds, as well as comment on their evolution and paleobiogeography. Our review also focuses on the geologic age of, in particular, the finds from northeastern Thailand (Khorat Plateau). There, a circa 3–4 km thick sequence of continental redbeds (Morley et al., 2011), the Khorat Group, that has been divided into several formations, yields the dinosaur fossils, including those of theropods. However, as in other parts of the world, the continental redbed sequences of the Khorat Plateau have been notoriously difficult to date, leading to sometimes strongly differing age assignments of the fossils in the older and more recent literature.
Section snippets
Institutional abbreviations
DIP: Dexu Institute of Palaeontology, Chaozhou, China; GMG: Geological Museum of Guangxi Zhuang Autonomous Region, southern China; HPG: Hupoge Amber Museum, Tengchong City Amber Association, China; IVPP: Institute of Vertebrate Paleontology and Paleoanthropology, Beijing, China; MDS: Dinosaur Museum, Savannakhet, Lao PDR; PRC: Paleontological Research and Education Center, Mahasarakham University, Maha Sarakham, Thailand; SM: Sirindhorn Museum, Department of Mineral Resources, Kalasin,
Theropod dinosaurs in Thailand
The first dinosaur bone from Thailand was found in 1976 by a geologist from the Department of Mineral Resources, Bangkok, in the course of a uranium survey at Phu Wiang District, Khon Kaen Province. The discovery consisted of a fragment of a sauropod femur from the Sao Khua Formation (Buffetaut, 1982). Since then, a collaboration led by the Department of Mineral Resources, Thailand, and the Centre National de la Recherche Scientifique, France, has led to the discovery of many dinosaur remains
Theropod dinosaur in Laos
The spinosaurid from Laos, Ichthyovenator laosensis Allain, Xaisanavong, Richir, and Khentavong, 2012, is from the “Grès supérieurs” Formation. This formation might be the same age as the Khok Kruat (Buffetaut, 1991) and Phu Pan Formations and is probably late Barremian to early Cenomanian in age (Allain et al., 2012, Allain, 2014). A non-marine bivalve assemblage recovered in the Tang Vay area suggests Ichthyovenator is Aptian in age (Allain et al., 2012).
Material: Holotype: partially
Theropod dinosaur in Malaysia
Spinosaurid teeth from the Malay Peninsula (Sone et al., 2015).
Material: Teeth UM10575 and UM10576 (Sone et al., 2015).
Occurrence: The Tembeling Group, which correlates to the Khorat Group. The exact locality is kept secret for protection and conservation (Sone et al., 2015).
Comments: The teeth show sharp vertical ridges and serrated carinae with minute denticles and display a veined micro-ornament over the surface (information from Sone et al., 2015). The teeth show spinosaurid characters
Theropod dinosaurs in Myanmar
Enantiornithine birds (Xing et al., 2016a, Xing et al., 2017, Xing et al., 2018, Xing et al., 2019).
Material: DIP-V-15100, DIP-V-15101, DIP-V-15102, DIP-V-15105, and HPG-15-1.
Occurrence: All material comes from the mid-Cretaceous Angbamo site, Hukawng Valley, Tanai Township, Myitkyina District, Kachin Province of Myanmar, which might be in the early Cenomanian (98.8 ± 0.6 Ma) or Albian–Cenomanian (105–95 Ma, based on ammonite biostratigraphy and palynology) (Shi et al., 2012, Xing et al., 2016a).
The Xinlong Formation
The Early Cretaceous Xinlong Formation of the Napai Basin and Datang Basin, Guangxi, southern China belongs to the same paleobiogeographic province as Southeast Asia. Therefore, adding theropod records from this formation to the present work can help to understand the diversity and paleobiogeography of this region. The Xinlong Formation was found to be equivalent to the Khok Kruat Formation in Thailand and the ‘Grès supérieurs’ Formation in Laos which are Aptian in age (Mo et al., 2016). From
Theropod footprints
Theropod footprints in Southeast Asia are mainly from Thailand with one report from Laos and one recent report from Malaysia (Buffetaut et al., 1985, Liard et al., 2015, Le Loeuff et al., 2009, Kozu et al., 2017, Allain et al., 1997, Akhir et al., 2015), see Table 1.
Theropod Diversity (Table 2)
Theropod faunas from the Late Jurassic to the Early Cretaceous of Southeast Asia include several clades, among them Metriacanthosauridae, Spinosauridae, Carcharodontosauria, Ornithomimosauria, as well as basal Coelurosauria and birds.
In Thailand, The Phu Kradung Formation yields two metriacanthosaurids. However, the Kham Phok and Phu Noi metriacanthosaurids might belong to the same taxon. The reported metriacanthosaurids, together with the mamenchisaurids found at the Phu Noi Locality, comprise
Disclosure of interest
The authors declare that they have no competing interest.
Acknowledgments
The authors would like to thank directors and staff of the Sirindhorn Museum, Paleontological Research and Education Center, Mahasarakham University, and Phu Wiang Fossil Research Center and Dinosaur Museum for hospitality and help during the visits. Thanks to Varavudh Suteethorn and Suravech Suteethorn (both PRC MSU) for general discussion, help, and support. This work was done during the tenure of A. Samathi on a scholarship to study in Germany from the Ministry of Science and Technology,
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Age, depositional history, and paleoclimatic setting of Early Cretaceous dinosaur assemblages from the Sao Khua Formation (Khorat Group), Thailand
2022, Palaeogeography, Palaeoclimatology, PalaeoecologyCitation Excerpt :Furthermore, based on the results of the KS-Test, we interpret that youthful zircon populations reflect episodic volcanism in the adjacent South China-Vietnam South Borneo Arc (SC-V-SB Arc) and the erosion of the developing fold belt (for instance, the Ban Don Group of Southern Vietnam (and similar terranes) (Hennig-Breitfeld et al., 2021; Nguyen et al., 2021; Schmidt et al., 2021). Our refined temporal framework for the Sao Khua Formation most closely aligns with the age estimate provided by Hasegawa et al. (2010) (late Valanginian/Barremian) as opposed to most published age estimates, which have proposed younger (e.g., Hauterivian (Racey et al., 1996); Hauterivian–late Barremian (Meesook, 2000; Nulay and Arjwech, 2021; Tumpeesawan et al., 2010; Samathi et al., 2019a, 2019b, 2021); or considerably older (Jurassic, Sattayarak (1983)) depositional ages. Upon investigation into exposed strata of the Sao Khua Formation, we find more diversity of paleosol types than originally described (Meesook, 2000).
New insights about theropod palaeobiodiversity in the Iberian Peninsula and Europe: Spinosaurid teeth (Theropoda, Megalosauroidea) from the Lower Cretaceous of La Rioja (Spain)
2020, Cretaceous ResearchCitation Excerpt :Other studies do also note the presence of the spinosaurid Spinosaurus in coastal environments (see Candeiro et al., 2018 and references). However, Southeast Asian spinosaurids seem to have inhabited fluvial environments with no marine influence (Buffetaut and Suteethorn, 1999; Buffetaut et al., 2005; Samathi et al., 2019). In Europe, spinosaurids are restricted to the western part of the continent, with remains found in Portugal, United Kingdom and Spain (Candeiro et al., 2017).
Preface for Palaeobiodiversity of SE Asia, issue 1
2019, Annales de Paleontologie