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Earth-Science Reviews 207 (2020) 103215

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Earth-Science Reviews

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Constraining assembly time of some blocks on eastern margin of Pangea T using Permo- non-marine tetrapod records ⁎ Jun Liua,b,c, , Jian Yia,c, Jian-Ye Chena a Key Laboratory of Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, b CAS Center for Excellence in Life and Paleoenvironment, Beijing 100044, China c College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing 100049, China

ARTICLE INFO ABSTRACT

Keywords: The supercontinent Pangea was mainly formed during the , but when it reached maximum land is Pangea unsure because the configuration of the East Asian blocks during the Permo-Triassic is still highly debated. Fossil East Asia tetrapods provide one of the best calibrations to the time of connection between continents, but the data of Assembly Permian Permo-Triassic tetrapods have rarely been applied in the previous reconstructions. Here we review the oldest Triassic records of non-marine tetrapods on the East Asia blocks and use them to discuss the timing of connection Non-marine tetrapods between continents during the Permian and Triassic. The distribution of Seymouriamorpha shows the Kazakhstanian was connected to the Baltica by land from the Cisuralian. The diverse Dashankou Fauna indicates that the Alxa Block amalgamated to Pangea at least in the (older than 266 Ma). The questionable footprints and late Permian assemblage from the North China show the North China Craton may have been connected to the main part of Pangea from the Guadalupian, and became part of Pangea at least by 256 Ma. The late Permian Laotian tetrapods support the hypothesis that the South China Block already collided with the North China and the Indochina blocks by the end of the Permian, and there was a land route for the migration of non-marine tetrapods.

1. Introduction Permian mass extinction (Faure et al., 1995). The configuration of major continental blocks in Pangea has been widely accepted although the Paleogeographic maps, showing ancient mountain ranges, shor- connection time of Kazakhstania and Baltica varies (Metcalfe, 2013; elines, shallow seas and deep ocean basins, are widely used in paleo- Muttoni et al., 2009; Torsvik and Cocks, 2017), but that of the East Asian climatology, plate tectonic reconstructions, paleobiogeography, re- blocks in Pangea is still highly debated (e.g., Domeier and Torsvik, 2014; source exploration and geodynamics (Cao et al., 2017). These maps are Metcalfe, 2011; Metcalfe, 2013; Stampfli et al., 2013; Torsvik and Cocks, reconstructed traditionally by using paleomagnetism, biogeography 2017; Zhao et al., 2018). The East Asia blocks include many blocks and and faunal links, abiotic links, and continent–ocean configurations terranes, but in this paper we mainly discuss those with Permo-Triassic (Ross, 1999). Starting from Stampfli and Borel (2002), the plate model non-marine tetrapod fossils: North China (NCB, also known as the Sino- accounting of plate kinematics and geodynamic forces was applied to Korean Block), South China (SCB, including Yangtze and Cathaysia), the early and Paleozoic reconstructions (Domeier and Tarim, Alxa (Alashan)-Hexi corridor-Qilian (AHQ), and Indochina (ICB, Torsvik, 2014). Because some fundamental data are absent and some also known as Annamia) (Fig. 1). are conflicting, different teams often produce different maps forthe In previous reconstructions, Pangea reached its maximum land by the same age, e.g., Permo-Triassic (Huang et al., 2018; Li and Powell, 2001; , and the East Asian blocks did not amalgamate to form part Metcalfe, 2011; Scotese and McKerrow, 1990; Sengör, 1987; Shi, 2006; of Pangea by that time in some paleogeographical maps (Domeier and Stampfli and Borel, 2002; Torsvik and Cocks, 2017; Zhao et al., 2018). Torsvik, 2014; Golonka, 2007; Metcalfe, 2011; Metcalfe, 2013; Scotese, During this period, the continents and terranes amalgamated as a 2004; Torsvik and Cocks, 2017). Because the process and timing of some supercontinent, Pangaea. The consolidation of Pangea had a deep impact key events are still unclear, the paleogeographical maps vary in the po- on ecosystems, e.g., the formation of coal (Nelsen et al., 2016), the end- sitions of many blocks, especially for the East Asian blocks.

⁎ Corresponding author at: Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China. E-mail address: [email protected] (J. Liu). https://doi.org/10.1016/j.earscirev.2020.103215 Received 19 January 2020; Received in revised form 5 May 2020; Accepted 10 May 2020 Available online 01 June 2020 0012-8252/ © 2020 Elsevier B.V. All rights reserved. J. Liu, et al. Earth-Science Reviews 207 (2020) 103215

Fig. 1. Orthographic projection of the present day Eastern Asia and adjacent areas showing the boundaries of some Paleozoic blocks and terranes and some Permian tetrapod localities. The blocks and terrane boundaries are modified from Domeier (2018) and Xiao et al. (2018)..

Traditionally, the biogeographic data were applied to determine During the early , tetrapods were known only in and test paleogeographic linkages between continental blocks (Cao Euramerica and Australia, but may have been distributed worldwide et al., 2017; McKerrow and Cocks, 1986; Shi, 2006). The timing of through equatorial regions (Thulborn et al., 1996). However, no Car- collage is generally inferred from sedimentary and metamorphic rock boniferous tetrapods are currently known from the NCB, SCB and ICB records. To supplement the knowledge, fossil tetrapods provide one of (Carroll, 2009 and see the data below), which may due to the isolation the best calibrations to the time of connection between continents. In of these blocks from Laurussia and although they lay in this paper, we summarize the current Permian and Triassic tetrapod equatorial region (Metcalfe, 2013; Torsvik and Cocks, 2017). Amniotes records of East Asia, use them to support the assembly of the East Asian appeared no later than the Late Carboniferous (~315 Ma) from Nova blocks with the Pangea and calibrate the times of assembly. The results Scotia, Canada (Carroll, 1964). During that time, the NCB, SCB, and ICB can be used to test and improve the reconstructed maps mainly based were isolated from Pangea, so the terrestrial tetrapods in these regions on the geological and paleomagnetic data. should have migrated from Pangea. As mentioned above, if the reached new continents after the blocks had collided and there was a land bridge, the fossil age can be 2. Method used as a time calibration of the land connection and the collision. However, if the animals had sweepstakes routes such as rafting on trees, In one of his most celebrated papers, G. G. Simpson (1940) dis- fossil calibration for collision is compromised. A good criterion for the cussed how some species had crossed regions that are now land bridge is that it does tend to transport integrated faunules impassable to them. He proposed land bridges and sweepstakes routes (Simpson, 1940). Another criterion is that the large-sized terrestrial as possible dispersal mechanisms. The rafting was used to explain how animals (e.g., length greater than 25 cm) passed the barrier and migrated from Africa to Madagascar. For ancient groups, appeared on a new continent. This kind of is too heavy to have a their distribution can only be understood in the light of plate tectonics. sweepstakes process. However, sometimes large animals can swim for The movement of plates changed the distribution of the tetrapods, ~100 km to reach isolated islands. Examples can be seen from living especially terrestrial tetrapods (Hedges et al., 1996). Most terrestrial and fossil mammals, e.g., Pliometanastes had arrived North America species can migrate from one block to another only when two blocks about 9 million years ago, prior to formation of the Isthmus of Panama collide and become connected. For example, some Gondwanan lineages about 2.7 million years ago, by island-hopping across the Central dispersed into Asia when peninsular collided with the Asian plate American Seaway (Tedford et al., 2004). So the second criterion should (Karanth, 2006). However, the sweepstakes process maybe a common be used in cautious. mechanism for small animals if there were suitable ocean currents and wind (Ali and Huber, 2010; Hawlitschek et al., 2016; Houle, 1998).

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3. Review of the Permian and Triassic non-marine tetrapods Lucaogou Formation of Urumqia, Xinjiang, China (Zhang et al., 1984), records of East Asia therefore, is the oldest Permian tetrapod in this region. The Lucaogou Formation in Xinjiang was regarded as upper Permian or later Gufeng In East Asia, Permian and Triassic tetrapods were reported only stage (Guadalupian), but it should be correlated to the Sakmarian to from China, Mongolia, and Thailand (Buffetaut et al., 2009a; Artinskian (Cisuralian) based on an IDTIMS age of 281.39 ± 0.10 Ma Gubin and Sinitza, 1993; Li et al., 2008). from the top of the Hongyanchi For mation and Gzhelian age (~300 Ma) from the lower part of the Daheyan Formation (Yang et al., 3.1. North and South foothills of the Bogda Mountain 2010), therefore, the age of Urumqia liudaowanensis could be older than 285 Ma (Fig. 2). No fossil tetrapod is known between the Lucaogou The Permo-Triassic deposits are well exposed on two sides of the Formation and the top of the Quanzijie Formation, which produced a Bogda Mountain, within the Junggar Basin and Turpan-Hami Basin. dicynodont Kunpania scopulusa (Sun, 1978). The taxonomical status of The tetrapods appeared in the Lucaogou, Quanzijie, Wutonggou, this species has been highly debated (Kammerer et al., 2011; Lucas, Guodikeng, Jiucaiyuan, Shaofanggou, Karamay, and Huangshanjie 1998, 2005), and an ongoing study may shed light on this question. formations, from old to young (Fig. 2). Urumqia liudaowanensis from the

Fig. 2. Typical North China Permian to Triassic stratigraphic sequences, tetrapod-bearing strata in bold..

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3.2. Tarim Basin 3.5. North China

No tetrapod fossils has been reported from the Permian and Triassic The Permo-Triassic tetrapods in this area appeared in the Xiashihezi rocks of the Tarim Basin. However, fragmentary tetrapod bones were (Lower Shihhotse), Shangshihezi (Upper Shihhotse), Sunjiagou, collected from the Lower Triassic Ehuobulak Group of Kuqa County, Heshanggou, Ermaying, and Tongchuan formations, from old to young Xinjiang. (Fig. 2). The only alleged Carboniferous tetrapod record in China was a slab of footprints from Baode, Shanxi, China (Fig. 1), in which the largest 3.3. Mongolia digit lengths are 70–80 mm, and total width of the three digits is 120 mm (Fig. 4). The original authors were unsure about the local The only Permo-Triassic tetrapod locality in Mongolia is Sain-sar- geology, and estimated the age based on the fossil plants in this area bulak in the Noyan Somon Depression. The tetrapod fossils were in- (Lucas and Ataabadi, 2010). We revisited the locality in 2018 and formally reported from the NoyanSomon and DalanShandakhuduk discovered that the slab was collected from the lower part of the Xia- formations, including a few skeletons of hedini from the shihezhi Formation, which is the lowest level in this small valley. The upper part of NoyanSomon Formation (Gubin and Sinitza, 1993). Xiashihezhi Formation was correlated to the Roadian to (lower Comparing with Lystrosaurus in Xinjiang, its age should be Changh- to middle Gudalupian) based on flora (Wang, 2010), but the Artinskin singian or . to Roadian based on spores and pollen (Liu et al., 2015). So, the age of the slab is conservatively estimated as the base of Roadian (~273 Ma). The slab, if really representing tetrapod footprints, is the oldest tetrapod 3.4. Alxa-Hexi corridor-Beishan record in North China. Other than this slab, the oldest tetrapods in this area belong to the From the late Carboniferous to the Permian, the North Qilian area Honania Assemblage (Jiyuan Fauna) from the upper part of the deposited the following sequence: Taiyuan, Shanxi, Dahuanggou, Shangshihezi Formation (Fig. 1), which is correlated to the Cistecephalus Yaogou, Qingtoushan and Sunan formations (Liu et al., 2012)(Fig. 2). Zone of , approximately 256 Ma (Liu, 2018a; Liu et al., The Taiyuan and Shanxi formations comprise marine facies, and ter- 2014; Rubidge et al., 2016). Originally, this assemblage was only restrial facies began to develop from the Dahuanggou Formation. There known from one locality in Jiyuan, Henan, but it is also discovered from are two Permian tetrapod fossil localities on the north margin of the Shouyan, Shanxi recently (Yi and Liu, 2020). Qilian Mountain: the Dashankou and Lugou localities, which lie in the Hexi corridor (North Qilian belt). 3.6. South China The Dashankou locality is close to Jiayuguan City, Gansu Province, China (Fig. 1), and a diverse tetrapod fauna is from the Qingtoushan There is no Permian tetrapod record in South China. The oldest Formation (Liu et al., 2012). This fauna includes diverse groups tetrapod records are marine from the Nanlinghu and (Table 1) with a large variation of body size (Fig. 3). Most of them are Jialingjiang formations, () in age (Cheng et al., likely carnivorous (including insectivorous, piscivorous), while Gan- 2019; Li et al., 2002; Motani et al., 2015). The terrestrial tetrapods are surhinus qingtoushanensis could be herbivorous. The skull length varies known from the Guanling Formation, the Falang Formation, and the from a few to more than 30 cm, and the largest skull reaches 40 cm in Badong Formation of South China (Li et al., 2017; Li et al., 2007; Liu length (Cheng and Ji, 1996; Li and Cheng, 1999; Liu, 2018b). This is and Wang, 2005; Zhang, 1975). The oldest terrestrial tetrapod in one of the oldest assemblages of the world. The age of the marine Triassic deposits is Pectodens zhenyuensis from the Member II of fauna is regarded as Roadian (Liu et al., 2009) or Wordian (Olroyd and the Guanling Formation, , from Luoping, Sidor, 2017; Rubidge, 2005), older than 266 Ma. (Li et al., 2017), with an age of 244.0 ± 1.3 Ma (Wang et al., The Lugou locality lies within Sunan County, Gansu Province, China 2014). The Member II of Badong Formation produced Lotosaurus and only a late Permian dicynodont, Turfanodon sunanensis, was pro- adentus, Yuanansuchus laticeps, and Y. maopingchangensis (Liu, 2016; Liu duced from the top of the Sunan Formation (Kammerer et al., 2011; Li and Wang, 2005; Zhang, 1975). Traditionally, the Badong Formation and Liu, 2015; Li et al., 2000). was assigned an Anisian to the in age, but recently Member II Another locality in this region, the Beishan locality, lies within was dated as the Ladinian or even in age by the laser ablation Beishan Hills, close to Mazongshan Town, Subei County, Gansu inductively coupled plasma mass spectroscopy [LA-ICP-MS] method Province, China. It produced plants, fish and tetrapods such as (Hagen et al., 2018; Wang et al., 2019). No matter what age it is, it Beishanodon youngi. The fossils were from the Lower Triassic dark shales should be younger than the Anisian (Liu, 2018a). of the Hongyanjing Formation. B.youngi is the second Triassic species in China, and is closely related to Sinognathus gracilis from the 3.7. Southeastern Asia Ermaying Formation (Gao et al., 2010). The oldest tetrapod records from the Southeast Asia include a Table 1 chroniosuchian Laosuchus naga (Arbez et al., 2018) and two dicynodont Fauna list of the Dashankou tetrapod Fauna. species (Counillonia superoculis and Repelinosaurus robustus)(Olivier et al., 2019) from the Purple Claystone Formation of the Luang Prabang Therapsida Sinophonus yumenensis Cheng and Ji, 1996 Basin, Laos (Bercovici et al., 2012; Rossignol et al., 2016). The chron- Biseridens qilianicus Cheng and Li, 1997 iosuchian skull measures 26 cm in length, while the dicynodont Raranimus dashankouensis Liu et al., 2009 are ~16 cm in basal length (Fig. 5). Although a maximum depositional Temnospondyli age of 251.0 ± 1.4 Ma was dated on detrital zircons by LA-ICP-MS Anakamacops petrolicus Li and Cheng, 1999 Chronisuchia method (Rossignol et al., 2016), a late Permian age is preferred (Liu, Ingentidens corridoricus Li and Cheng, 1999 2020) and a minimum age of 252 Ma is adopted here. The Red Clays- Phratochronis qilianensis Li and Cheng, 1999 tone Formation above the Purple Claystone Formation from Laos pro- Yumenerpeton yangi Jiang et al., 2017 duced a large dicynodont, but perhaps is Late Triassic in age (Bercovici Reptilia et al., 2012; Rossignol et al., 2016). Belebey chengi Müller et al., 2008 Gansurhinus qingtoushanensis Reisz et al., 2011 Thailand also produced fossil tetrapods in the Huai Hin Lat and Nam Phong formations, which are the latest Triassic in age (Buffetaut et al.,

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Fig. 3. Tetrapods of the Dashankou Fauna from the Qingtoushan Formation of Yumen, Gansu, China. Therapsida: (A) Sinophonus yumenensis, (B) Biseridens qilianicus, (C) Raranimus dashankouensis; Reptilia: (D) Gansurhinus qingtoushanensis, (E) Belebey chengi; Chronisuchia: (F) Phratochronis qilianensis, (G) Ingentidens corridoricus; Temnospondyli: (H) Anakamacops petrolicus. Scale bars equal to 1 cm except for H (4 cm).

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Fig. 4. Slab of footprints from the Xiashihezi Formation of Baode, Shanxi, China.

2009a). The tetrapods include temnospondyl Cyclotosaurus (Ingavat and Janvier, 1981), Proganochelys ruchae (De Lapparent, 1985), phytosaurs (Buffetaut and Ingavat, 1982), and (Buffetaut and Suteethorn, 1998; Buffetaut et al., 2009b).

4. Discussion

4.1. Urumqia liudaowanensis and the early Permian Kazakhstania

The Urumqi locates at the southern border of the Junggar Basin, in contact with the North Tian Shan, which consists of a to Carboniferous accretionary complex and a volcanic arc. From the late Carboniferous to the early Permian, this area changed from shallow sea to land: the Tashikula Formation is marine facies, the Wulabo and Fig. 5. Late Permian tetrapods from China and Laos. Dicynodontia: (A) Jingjingzigou formations are delta and littoral deposits, and the Taoheodon baizhijun from the of Shouyan, Shanxi, China; Lucaogou Formation is a lacustrine deposit. This is part of the Tianshan (B) Counillonia superoculis, and (C) Repelinosaurus robustus from the Purple and Junggar orogenic collage in the late Paleozoic, all oceans are closed Claystone Formation of the Luang Prabang Basin, Laos. Chronisuchia: (D) at the end of the Carboniferous in this orogenic collage, and this area Laosuchus naga from the Purple Claystone Formation of the Luang Prabang was amalgamated to Kazakhstania (Han and Zhao, 2018). This area was Basin, Laos. Scale bars equal 5 cm. (For interpretation of the references to a lake within the Bogda basin in the early Permian. The Mongolian colour in this figure legend, the reader is referred to the web version ofthis locality also located near the southern margin of Mongolian continent, article.) it needs no discussion here for its later age. U. liudaowanensis is similar to Utegenia shpinari from Southern Torsvik (2014) or Torsvik and Cocks (2017), but different from that of Kazakhstan and was regarded as a junior synonym of the latter Metcalfe (2013). (Ivakhnenko, 1987). U. shpinari may be the basal most member of the Seymouriamorpha and is of the late Carboniferous or the early Permian in age; most other seymouriamorphs are distributed in the lower Per- 4.2. Tarim Craton mian of Europe and Tadzhikistan (Laurin, 1996). The new age of Ur- umqia is concordant with the ages of other species. The distribution of The timing of collision between Tarim and Kazakhstania is con- this group in the early Permian also indicated that the Kazakhstania tested. Some suggested in the late Carboniferous-earliest Permian was connected to the Baltica Craton (Fig. 6), and became part of Pangea (~300 Ma) (Gao et al., 2011; Gao et al., 2006); and some held in the at least from the early Permian, similar to the view of Domeier and late Permian to Triassic (Xiao et al., 2013; Zhang et al., 2007). Or, it

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Fig. 6. Simplified paleogeographic reconstructions of part of the Pangea in (A) early Guadalupian (~266 Ma) and (B) end-Permian (~252 Ma),showingthe connection of lands (modified after figure 11of Xiao et al., 2015). could be a diachronic process: it occurred in the late Carboniferous to tetrapods can only migrate via Tarim Craton. This was a scenario held by Permian in the east and in the end-Permian to Middle Triassic in the most studies (Huang et al., 2018; Metcalfe, 2013; Shi, 2006; Torsvik and west (Xiao et al., 2015). Now, the timing of collision between Tarim and Cocks, 2017; Xiao et al., 2015; Zhao et al., 2018). Kazakhstania is generally accepted as at least by ~285 Ma based on geological evidence (Domeier and Torsvik, 2014; Zhao et al., 2018), and paleomagnetic data (Huang et al., 2018). Following this hypoth- 4.4. Beishanodon youngiand Beishan Orogenic Collage esis, the tetrapods should have existed in this block from the early Permian. The only known Early Triassic tetrapod bone agrees with this The Beishan locality lies within the Mazongshan arc, north to the hypothesis, and the future discovery of Permian tetrapods may be used Hongyuanjing basin (Gao et al., 2010; Tian et al., 2015). The fossil to further test this hypothesis. shows this part was connected to the North China Block by the Early Triassic. This agrees with the reconstructed history of accretionary or- 4.3. Dashankou tetrapod fauna and assembly of the Alxa block to Pangea ogenesis of this region by Tian et al. (2015). However, Xiao et al. (2018) proposed the amalgamation of the Dunhuang Block with the Mongolia The Hexi corridor, Qilian, and Qaidam were united to the Alxa block collage was in the early Permian to the Middle Triassic, so the Dun- by at least the Devonian (Domeier, 2018; Duan et al., 2015; Song et al., huang Block could have amalgamated with the Huaniushan–Dundun- 2013), so this region is referred only as the Alxa block plus Qaidam shan arc in the Guadalupian, and tetrapods could also exist in the BOC block here. (e.g., Hongyuanjin basin) during that time. This can be tested by the The Dashankou tetrapod fauna is characterized by the Sinophoneus fossil tetrapods from the BOC. Assemblage, which resembles the Russian Ocher Subassemblage (on Baltica Craton), and is more primitive than the South African Eodicynodon Assemblage. The age of the Eodicynodon Assemblage Zone 4.5. Assembly of the Alxa Block to the North China Block is ~266 Ma (Lucas, 2018; Rubidge et al., 2016), so a minimum age for the Dashanhou Fauna is estimated as 266 Ma. The high diversity and Traditionally, the position of the Alxa terrane relative to the main the large body sizes of the Dashankou Fauna show that the tetrapods body of the NCB is thought to have been almost stable at least since had freely exchanged on land between the Alxa terrane and Baltica mid-Paleozoic (Huang et al., 2001; Huang et al., 1999; Liu et al., 2016). craton in the mid-Guadalupian (~266 Ma). So the Alxa terrane should However, based on the paleopoles reconstructed from Alxa, Yuan and have amalgamated to be part of the Pangea at least by this time. Yang (2015) proposed that the Alxa terrane may have been separated How did the Alxa Block connect to the Baltica craton? This needs a from the main body of the NCB until the late Permian. route via the Qaidam and Dunhuang blocks or the Beishan Orogenic If the identification of the above-mentioned footprints (Lucas and Collage (BOC), across Kazakhstania (Domeier, 2018; Tian et al., 2015; Ataabadi, 2010) is correct, the tetrapods had reached the NCB by Xiao et al., 2018)(Fig. 1). The Dunhuang Block, commonly presumed to ~273 Ma. Because the large size of the trackmaker, the possibility of be of the Tarim affinity, was also proposed as a Paleozoic orogenic belt sweepstakes is fairly low, so the alleged footprints could support the and part of the Central Asian Orogenic Belt (Zhao et al., 2016). There is no connection of the NCB to Pangea by that time. However, this evidence available data on the collision between the Dunhuang and Alxa blocks, but is weak and lacks the support from other faunas, so it will not be they should have connected by land in the Guadalupian (before 266 Ma) counted here. Diverse late Permian tetrapods provide a constraining (Fig. 6A). If the Dunhuang Block was still completely separated from the age (256 Ma) for the amalgamation of the NCB with the Alxa and Beishan Orogenic Collage by the Paleo-Asian Ocean (Tian et al., 2015), the Pangea.

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4.6. Laotian tetrapods and their implications for the end-Permian Indochina blocks by the end of Permian, and there was land route paleogeographical map for the migration of non-marine tetrapods. (5) Pangea already reached its maximum land by the end Permian not The Laotian tetrapods were known for more than 100 years, but the Late Triassic. only until much later did paleontologists pay attention to its paleo- geographical implications (Arbez et al., 2018; Battail, 2009; Buffetaut, Declaration of Competing Interest 1989; Lucas, 1997; Olivier et al., 2019). The tetrapods were collected from the Luang Prabang Basin, Laos, which lies in the Indo-China Block The authors acclaim no conflict of interest. (or Annamia craton). This place belongs to the Luang Prabang-Loei back-arc basin between the Central Indochina fragment and Simao and Acknowledgments West Indochina fragments (Wang et al., 2018). There were two possible routes for tetrapods from the Pangea to the This research was supported by Strategic Priority Research Program ICB at the end-Permian: (1) the NCB connected with the ICB, via the of Chinese Academy of Sciences (XDB26000000) and National Natural SCB; (2) via Cimmerian continental strip (Olivier et al., 2019). The Science Foundation of China (41572019, 41661134047). Thanks for large size of the tetrapods and two different clades guarantee the pre- the comments from Lin Wei and Shang Qing-Hua for an earlier version sence of a land bridge between the ICB and NCB, and the route from the of the manuscript, and from Spence Lucas and Sebastian Voigt for the SCB is preferred. This conclusion is also supported by the close re- submitted manuscript. Photos in figure 5 were kindly provided by lationship of Laotian tetrapods with those from Russia and China (Arbez Chloe Olivier, Thomas Arbez and Christian Sidor. et al., 2018; Liu, 2020). The timing of collision between the North China and South China References blocks has been extensively discussed in the earlier literature. Some proposed a Mesozoic age e.g., (Li et al., 1989; Li et al., 1993), and some Ali, J.R., Huber, M., 2010. Mammalian biodiversity on Madagascar controlled by ocean paleogeographic maps adopted this view (Scotese, 2004; Torsvik and currents. Nature 463, 653–656. Cocks, 2017). Based on paleomagnetic evidence, Zhao and Coe (1987) Arbez, T., Sidor, C.A., Steyer, J.S., 2018. Laosuchus naga gen. et sp. nov., a new chron- iosuchian from South-East Asia (Laos) with internal structures revealed by micro-CT first proposed NCB and SCB having the first contact at the present scan and discussion of its palaeobiology. J. Syst. Palaeontol. 1–18. eastern end of the Qinling-Dabie Fold Belt during the Late Permian. Battail, B., 2009. Late Permian dicynodont fauna from Laos. Geol. Soc. Spec. 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