sign in sign up
<<
Home , Ginglymodi, Halecostomi, Holostei, Lepidotes, Non Sung District, Semionotiformes

Original

Ginglymodian fi shes (, ) from : An overview

Lionel Cavin1*, Uthumporn Deesri2,3, Varavudh Suteethorn2 Received: 30 July 2013; Accepted: 15 October 2013

Abstract Isolated ganoid fi sh scales are not uncommon in deposits of Thailand. Traditionally referred to ‘semionotiform’ or -like fi shes, they were not assigned to well-defi ned taxa and are of little use for palaeontological reconstructions. During the last fi fteen years, however, the discovery of well-preserved articulated fi sh specimens, with ganoid squamations, allowed us to properly defi ne new taxa, to search for phylogenetic relationships and to address the place of these fi shes in palaeoenvironments. So far two genera and three of ginglymodians have been named on the basis on material from the Phu Kradung Formation, but at least nine different taxa have been recognized ranging from the Late to the Aptian. Phylogenetic analyses of Thaiichthys and Isanichthys indicate that they belong, or are closely related, to the Lepisosteiformes. The palaeogeographical distribution of the four known Isanichthys species is restricted to the northern margin of the Tethys during the Middle to the basal . The palaeobiogeographical signal of Thaiichthys is more ambiguous, its closest relatives having been found in the early of western Gondwana (South America and Africa). Keywords: Cretaceous, , palaeobiogeography, phylogeny Introduction articulated material, in particular if no articulated cranial Mesozoic deposits worldwide, both marine and freshwater material is preserved. Relatively complete specimens in in origin, have yielded isolated ganoid scales commonly anatomical connexion of ginglymodians, however, are referred to ‘semionotiforms’ or Lepidotes-like fi shes. known for a long time in various Lagerstätten worldwide, These scales can generally be distinguished from ganoid and their study have enabled researchers to propose scales of non-neopterygian fi shes (the ‘palaeonisciforms’) phylogenetic relationships within the group, and to consider because the abdominal fl ank scales usually bear a pair relationships of these fi shes with other actinopterygians. of processes on their anterior margin in complement Classically, the Holostei gathered the living to the dorsal process1,2. Their abundance in the and Lepisosteidae, together with some extinct groups. record is due in part to their strong mineralization, being Patterson (1973)7 questioned this concept: he regarded constituted of a bony basal plate cover with an enamel the Holostei as a non-monophyletic group and consid- layer. Except some research that attempt to identify ered the Semionotidae as , i.e. as closer to the systematic affi nities of isolated scales on the basis than to (Lepisosteidae). Alternatively, Olsen of the micro-ornamentation of the enamel layer3,4.5.6, a and McCune (1991)8 regarded the and the taxonomical assignment is usually not possible with no Lepisosteidae as forming a with the Macrosemiidae

1 Department of Geology and Palaeontology, Muséum d’Histoire naturelle, CP 6434, 1211 Genève 6, Switzerland 2 Palaeontological Research and Education Centre, Mahasarakham University, Khamrieng, Kantharawichai, Mahasarakham 44150, Thailand 3 Department of Biology, Faculty of Science, Mahasarakham University, Khamrieng, Kantharawichai, Mahasarakham 44150, Thailand * Corresponding author: [email protected] Vol 33. No 4, July-August 2014 Ginglymodian fi shes (Actinopterygii, Holostei) from Thailand: An overview 349

(the ), which is resolved as the sister- Phra Wihan, Sao Khua, Phu Phan and Khok Kruat forma- group of teleosts. In their scheme, Halecomorphi ( tions, in ascending . Chonglakmani (2011)18 included and relatives) are located as the sister-group of Semiono- in Khorat Group the underlaying Nam Phong Formation. tiformes + Teleostei. During the last decades, however, Ginglymodians have been found in the Phu Kradung, most molecular phylogenetic studies have resuscitated the Sao Khua and the Khok Kruat formations, as well as the Holostei (for instance Venkatesh et al. 20019 , Inoue in the underlying Huai Hin Lat Formation. et al. 200310, Near et al. 201211; Betancur et al. 201312), The Phu Kradung Formation comprises fl uvial channel and this confi guration was subsequently also found in sandstone, siltstone and mudstone with intermittent morphological studies, which included fossil taxa13,14. calcretes. The formation was deposited in a mainly In his review of the Lepisosteiformes Grande (2010)13 lake-dominated fl oodplain cut by meandering and occa- proposed a classifi cation, in which the Holostei include sionally braided river channels. The formation is sandier the Halecomorphi and a clade called Ginglymodi, which in its upper part. The age of most of this formation is gathers gars together with several extinct taxa. Most considered to be Late Jurassic on the basis of recent recent studies have found the monophyly of Ginglymodi vertebrate discoveries19,20,21 but the upper part, however, containing the Lepisosteiformes, ‘macrosemiiformes’ and is now regarded as basal Cretaceous in age based ‘semionotiformes’, but there is still no consensus about on palynological evidence17. Similar ages have been the respective composition and relationships of the lat- established on the basis of the assemblages22. The ter two groups (compared for instance López-Arbarello Sao Khua Formation comprises dominantly fl oodplain 20122 with Cavin et al. 201314 and Deesri et al. 201415). deposits including sandstone, siltstone and mudstone, An overview of Jurassic and Cretaceous bony together with common calcretes, and was deposited fi sh record from Thailand has been published in 20091. in a low-energy fl uvial setting comprising meandering However, new discoveries and new analyses of the channels and extensive fl ood plains. Based on paly- data have been made during the last years, and a new nomorphs, a Berriasian- early Barremian age is sug- synthesis is now necessary. Here, we present an overview gested for this formation17. The Khok Kruat Formation of the discoveries of ginglymodians in Thailand, and we comprises sandstone, conglomerate, siltstone, shale and discuss their impact in the evolutionary history of the intermittent palaeosols deposited in a dominantly fl uvial group within the frame recent phylogenies. environment. This formation is considered as Aptian in age on the basis of palynomorphs23 and vertebrates24,25. Geological setting Thailand consists of two continental blocks or microcon- The Ginglymodian Thai fossil record tinents: the eastern part, on which is located the Khorat Since thirty years, the Thai-French collaboration team Plateau, belongs to the Indochina block, and the western has studied non marine vertebrate from Thailand, part, including the southern Peninsula, belongs to the ranging in age from the Late Triassic to the late Early Cre- ‘Shan-Thai’ or ‘Sibumasu’. The Shan-Thai block has taceous. Bony fi shes have been discovered regularly, and yielded very few ginglyomodian fi sh remains so far, except were mostly represents by isolated ganoid scales referred scales and fragmentary elements found in the Mab Ching to semionotid-like fi shes25. From 2002 to 2007, excava- locality in the Khlong Min Formation, Middle or Late Juras- tions were repeatedly conducted in the site of Phu Nam sic in age1. Most of the ginglymodian remains have been Jun, Lao Yai, Kalasin Province, a locality disco found in the Khorat Plateau in NE Thailand, which contains vered some years before by local people26. So far this site non-marine sediments deposited during the Mesozoic. has yielded only bony fi shes, and this discovery triggered Racey (2009)16 and Racey and Goodall(2009)17 restricteded specifi c researches on this group of vertebrates. Most of the Khorat Group to fi ve formations: the Phu Kradung, the specimens belong to a ginglymodian fi sh, Thaiich- 350 Cavin et al. J Sci Technol MSU thys buddhabutrensis (Cavin, Suteethorn, Khansubha, Formation known by articulated material (Thaiichthys Buffetaut & Tong, 2003)27, which was described in detail and Isanichthys) or if they correspond to new taxa. only recently14. A taphonomical and morphometric study Outside the Phu Kradung Formation, numerous other of the T. buddhabutrensis assemblage was published by localities from the Mesozoic of Thailand have yielded Deesri et al.28 in 2009. In the meantime, a single speci- isolated ginglymodian remains from different stratigraphic men from the Phu Nam Jun site was referred to another horizons. The locality of Huai Pha Pheng, in the Late ginglymodian, Isanichthys palustris Cavin and Suteethorn, Triassic Huai Hin Lat Formation, is one of the rare lo- 2006. In 2008, a new site called Phu Noi, situated in calities, which has yielded articulated material with the the Phu Kradung Formation but located stratigraphically recent discovery of several subcomplete ray-fi nned fi shes. lower than the site of Phu Nam Jun22 has been regularly Some of these specimens are probably ginglymodians, excavated. A very rich vertebrate assemblage, compri but their study is too preliminary to permit well-supported sing , turtles, crocodiles and different dinosaurs, assignations. Other discoveries are mostly isolated scales, has been discovered22. Several ginglymodian specimens rare teeth and ossifi cations. Although this kind of were also recovered. Although they show some variability preservation precludes the recognition of taxa, we can in their skull bone pattern, they have been all referred to assess the diversity on the basis of the association of a species of Isanichthys distinct from I. palustris, I. lert- teeth and scales, and on the morphology of the latters. boosi Deesri, Lauprasert, Suteethorn, Wongko and Cavin The locality of Mab Ching, in the Khlong Min Forma- (2014)15. Isolated discoveries of articulated ginglymodians tion, has yielded an association of large ganoid smooth have been done in other localities from the Phu Kradung scales with a conical tooth, which indicate the occur- Formation. An almost complete specimen was discovered rence of a ginglymodian taxon likely different from those in the Ban Nong Rua site, , of the other localities (Ginglymodi indet. 1 in table 1). Province, and has been described Two localities from the Sao Khua Formation (Phu Phan in a PhD Thesis29. It likely represents a new taxon and Thong and Phu Wiang) have yielded smooth ganoid scales its phylogenetic relationships are currently investigated. associated with button-like teeth. This kind of teeth has Two specimens closely related to Thaiichthys have never been found in the Phu Kradung Formation so far, and been discovered in localities other than Phu Nam Jun: a it likely indicates the occurrence of another taxon of gingly- specimen referred to Thaiichthys cf. buddhabutrensis was modian in this formation (Ginglymodi indet. 2 in table 1). found in the locality of Chong Chat, , In the Khok Kruat Formation, two kinds of scales have Nong Bua Lamphu Province29 and one specimen from been found in the Khok Pha Suam locality: one kind has a the locality of Kham Phok, Nong Sung District, Mukdahan smooth surface and the other has parallel ridges Province, was regarded as a juvenile of T. buddhabut- on the surface. These scales, never found associated with rensis30. It should be noticed here that both specimens button-like teeth, indicates the occurrence of two taxa in show differences with individuals from the type locality, this locality (Ginglymodi indet. 3 and 4 in table 1). The but because this species possesses a wide range of site of Lam Pao Dam, in the Khok Kruat Formation, has morphological variability, and because only one speci- yielded large ganoid scales ornamented with parallel rows men from each locality is available, we prefer to avoid of small pits, as well as a large button-like tooth (Gingly- erecting new taxa pending the discovery of new material. modi indet. 5 in table 1). The locality of Ban Saphan Hin Other localities from the Phu Kradung Formation, such in the Khok Kruat Formation, Mueang Nakhon Ratchasima as Khok Sanam, Dan Luang, Phu Dan Kaeng, Lam District, Nakhon Ratchasima Province, has yielded the Payang (Phu Klang) and Wan Din So contain smooth posterior part of an articulated fi sh, as well as numerous ganoid scales, but we are not able so far to determine isolated skull bones29. We cannot determine so far if these if they can be referred to one of the taxon from this elements correspond to one of the three other taxa pre Vol 33. No 4, July-August 2014 Ginglymodian fi shes (Actinopterygii, Holostei) from Thailand: An overview 351 sent in other localities of the Khok Kruat Formation, or if monophyletic Lepisosteiformes, but the taxa placed in they correspond to a new taxon. Thus, at least three gin- the Semionotiformes in López-Arbarello’s study are not glymodian taxa are present in the Khok Kruat Formation. resolved as a clade, but are situated in various positions The study of the micro-ornamentation of the ena as ‘basal’ ginglymodians. Most of the recent phylogenetic mel surface of these scales is a research in prog- analyses resolve the genera Scheenstia and Lepidotes ress29, which should allow assessing more effec- as basal Lepisosteiformes (Figure 1). Cladistic analyses tively the diversity of ginglymodians in each formation. that include both genera defi ned on the basis of Thai So far ten distinct ginglymodian species can be distin- material, Thaiichthys and Isanichthys, indicate 1) that guished in Thailand ranging from the Late Triassic Huai Thaiichthys is a Lepisosteiformes situated as the sister- Hin Lat Formation to the Aptian Khok Kruat Formation. group of the Western Gondwanian pair Pliodetes and Three species have been named from the Phu Kradung Araripelepidotes14,15, and 2) that Isanichthys, is resolved Formation, and a forth is pending. The diversity in the either in a polytomy with other ginglymodians14,15,36,37 or Khok Kruat Formation is apparently also rather important; as basal Lepisosteiformes2,15. It is interesting to note that although the material is not complete enough for erec both Thai genera belong to the Lepisosteiformes (or are ting new taxa. Strangely enough, the lowest diversity is closely related to Lepisosteiformes for Isanichthys in observed in the Sao Khua Formation, which otherwise some of the studies), a clade containing the living gars has yielded the most diverse assemblages31. This restricted now to North and Central America. ‘Lepidotes’ paucity is possibly related to taphonomical features of luchowensis Wang, 1974, from the Middle Jurassic of the vertebrate localities that have been excavated in this Sichuan, China, and ‘Lepidotes’ latifrons Woodward, formation until now. One exception is the locality of Phu 1893 from the Middle Jurassic of UK are included in the Phok (Sakhon Nakhon Province), which is the type locality Isanichthys15. The palaeogeographical distribu- of the sinamiid Siamamia, a halecomorph fi sh represented tion of the four species of Isanichthys is broad, but it is by isolated bones and some articulated cranial material32. concentrated to the northern margin of the Tethys from the Middle Jurassic to the basal Cretaceous (Figure Evolutionary history 1). The palaeobiogeographical pattern associated with The resurrection of Holostei on the basis of morphologi- Thaiichthys is more confuse. This genus appears to be cal characters, including data from fossil taxa, was fi rst related to a lineage from western Gondwana, which split proposed by Grande(2005, 2010)33,13. This clade includes into the pair genera Pliodetes and Araripelepidotes in the Halecomorphi and the Ginglymodi, the latter compris- the early Late Cretaceous, probably in connection with ing the Macrosemiiformes, the Semionotiformes and the the opening of the South Atlantic. Because these three Lepisosteiformes. In Grande’s study, the Semionotiformes genera are freshwater, or possibly brackish for Araripel- consist of the genus (represented by S. el- epidotes, we suggest that the observed phylogenetic egans (Newberry, 1888)34 only. Cavin(2010)35 proposed a pattern is related to the fragmentation of Pangea and phylogenetic analysis of a set of Mesozoic ginglymodians, that the split should be as old as Triassic in age. Vi- and found that several Cretaceous taxa are stem lepiso- cariant events associated with the opening of the South steids, a pattern also found in López-Arbarello (2012)2, Atlantic are also observed in other Lepisosteiformes, in although different taxa are placed as stem lepisosteids. particular the split of sister species within both obaichthyid In López-Arbarello’s analysis2, the Semionotiformes are genera, Obaichthys and Dentilepisosteus (Figure 1). monophyletic and gathers genera ranging from the Trias- It is worth noting that the general patterns proposed sic to the Cretaceous, and includes the Macrosemiidae here are still weakly supported because their rest on few (which, consequently, lost an ordinal rank as part of the taxa and on a weakly supported phylogeny. But they Semionotiformes). Recent analyses14,36,15 recover the constitute hypotheses to be tested in further studies. 352 Cavin et al. J Sci Technol MSU

Conclusion 2. López-Arbarello A. Phylogenetic interrelationships Ginglymodian fi shes are relatively common in the Me- of ginglymodian fi shes (Actinopterygii, ). sozoic sediments of Thailand. Although only two genera PLoS ONE 2012; 7(7), e39370. doi,10.1371/journal. and three species have been properly identifi ed so far, pone.0039370. new material currently studied and recent fi eld disco 3. Gayet M, Meunier F. Apport à l’étude de veries indicate that much more taxa were present. The l’ornementation de la ganoïne dans la détermination knowledge of the Thai ginglymodian fossil record has de l’appartenance générique et/ou spécifi que des twofold impacts: 1) It allows to better understand the écailles isolées. Comptes Rendus de l’Académie des palaeoecology of the vertebrates assemblages in the Sciences, Paris 1986; 303, sér. II, 13, 1259-1262. Mesozoic of South-East Asia, in particular by showing 4. Gayet M, Meunier F. A propos du genre Paralepi- the existence of ginglymodian fi shes with peculiar posi- dosteus (Ginglymodi, Lepisosteidae) du Crétacé tions within the trophic web: Thaiichthys was probably a Gondwanien. Cybium 2001; 25 (2): 153-159. grazing herbivorous or detritus feeder, while Isanichthys 5. Brito P, Meunier F. The morphology and histology was likely a predator; 2) It contributes to depict the of the scales of Aspidorhynchidae (Actinopterygii, evolutionary history of the ginglymodians as a whole, Halecostomi). Geobios 2000; 33(1): 105-111. which were fi shes with a wide variety of morphology, 6. Meunier F. The , from the Paleozoic to ecology and complex palaeobiogeographical history. the extant time, through histology and palaeohistology of bony tissues. Comptes Rendus Paleovol 2011; Acknowledgments 10: 347-355. The authors thank all of the members of the Palaeontolog- 7. Patterson C. Interrelationships of holosteans. In: ical Research and Education Center (PRC) of Mahasara- Greenwood, P. H., Miles, R. S. and C. Patterson kham University, students of Biology Department, Faculty (eds) Interrelationships of fi shes, Academic Press, of Science of Mahasarakham University, and especially London 1973; 233-305. the members of the staff of Sirindhorn Museum and of 8. Olsen PE, McCune AR. Morphology of the Semiono- the Department of Mineral Resources (DMR) of Thailand tus elegans group from the Early Jurassic part of the who participated to the fi eldworks. The fi eld work was Newark Supergroup of Eastern North Americ with funded by the department of Mineral Resources and the comments on the Semionotidae (Neopterygii). Palaeontological Research and Education Center. UD was Journal of Vertebrate Paleontology 1991; 11(3): 269- supported by the Research in Higher Education Project 292. year 2014/Higher Education Research Promotion (HERP) 9. Venkatesh B, Erdmann MV, Brenner S. Molecular and National Research University Development in the of- synapomorphies resolve evolutionary relationships fi ce of the Higher Education Commission. We thank Julien of extant jawed vertebrates. PNAS 2001; 98(20): Claude (ISE-M, Montpellier) for reviewing this article. 11382–11387. 10. Inoue JG, Miya M, Tsukamoto K, Nishida M. Basal References actinopterygian relationships, a mitogenomic per- 1. Cavin L, Deesri U, Suteethorn V. The Jurassic and spective on the phylogeny of the ‘ancient fi sh’. Cretaceous bony fi sh record (Actinopterygii, Dipnoi) Molecular Phylogenetics and Evolution 2003; 26: from Thailand. In: Buffetaut, E., Cuny, G., Le Loeuff, 110–120. J. and Suteethorn, V. Late Palaeozoic and Mesozoic 11. Near TJ, Eytan RI, Dornburg A, Kuhn, KL, Moore Continental Ecosystems of SE Asia. Geological JA, Davis MP, Wainwright PC, Friedman M, Smith Society of London, Special Publication 2009; 315: WL. Resolution of ray-fi ned fi sh phylogeny and timing 123-137. of diversifi cation. PNAS 2012; 109(34): 13698-13703. Vol 33. No 4, July-August 2014 Ginglymodian fi shes (Actinopterygii, Holostei) from Thailand: An overview 353

12. Betancur-R R, Broughton RE, Wiley EO, Carpenter Formation of northeastern Thailand. Bulletin de la K, López JA, Li C et al. The tree of life and a new Société Géologique de 2007; 178: 497-502. classifi cation of bony fi shes. PLOS Currents Tree of 21. Tong H, Claude J, Naksri W, Suteethorn V, Buf- Life. DOI, 10.1371/currents.tol.53ba26640df0ccaee fetaut E, Khansubha S, Wongko K, Yuangdetkla P. 75bb165c8c26288. 2013. Basilochelys macrobios n. ge. and n. sp., a large 13. Grande L. An empirical synthetic pattern study of cryptodiran turtle from the Phu Kradung Formation gars (Lepisosteiformes) and closely related species, (latest Jurassic-earliest Cretaceous) of the Khorat based mostly on skeleton anatomy. The resurrection Plateau, NE Thailand. In: Buffetaut, E., Cuny, G., Le of Holostei. Copeia 10. American Society of Ichthyolo- Loeuff, J. and Suteethorn, V. (eds) Late Palaeozoic gists and Herpetologists, special Publication 2010; 6 and Mesozoic Ecosystem in SE Asia. Geological (suppl. 2a): 1-871. Society, London, Special Publications 2009; 315: 14. Cavin L, Deesri U, Suteethorn V. Osteology and rela- 151-170. tionships of Thaiichthys nov. gen., a ginglymodi from 22. Cuny G, Liard R, Deesri U, Liard T, Khamha S, Su- the Late Jurassic – of Thailand. teethorn V. Shark faunas from the Late Jurassic-Early Palaeontology 2013; 56(1): 183-208. Cretaceous of northeastern Thailand. Paläontolo- 15. Deesri U, Lauprasert K, Suteethorn V, Wongko K, gische Zeitschrift. DOI 10.1007/s12542-013-0206-0. Cavin L. A new species of the ginglymodian fi sh 2013. Isanichthys from the Late Jurassic Phu Kradung For- 23. Sattayarak N, Srigulawong S, Patarametha M. Sub- mation, northeastern Thailand. Acta Palaeontologica surface stratigraphy of the non-marine Mesozoic Polonica 2014; 59(2): 313-331. Khorat Group, northeastern Thailand. GEOSEA VII 16. Racey A. Mesozoic red bed sequences from SE Abstracts, Bangkok, 1991; p.36. Asia and the signifi cance of the Khorat Group of NE 24. Cappetta H, Buffetaut E, Suteethorn V. A new Thailand. In: Buffetaut, E., Cuny, G., Le Loeuff, J. and hybodont from the Lower Cretaceous of Thailand. Suteethorn, V. (eds) LatePalaeozoic and Mesozoic Neues Jahrbuch für Geologie und Paläontologie, Ecosystems in SE Asia. Geological Society, London, Abhandlungen 1990; 11: 659-666. Special Publications 2009; 315: 41-67. 25. Buffetaut E, Suteethorn V. The biogeographical sig- 17. Racey A, Goodall J. Palynology and stratigraphy of nifi cance of the Mesozoic vertebrates from Thailand. the Mesozoic Khorat Group red bed sequences from In: Hall, R. and Holloway, J.D. (eds), Biogeography Thailand. In: B Buffetaut, E., Cuny, G., Le Loeuff, and Geological Evolution of SE Asia, Backhuys, J. and Suteethorn, V. (eds) Late Palaeozoic and Leiden 1998; 83-90. Mesozoic in SE Asia. Geological Society, London, 26. Cavin L, Suteethorn V, Buffetaut E, Chitsing S, Special Publications 2009; 315: 69-83. Lauprasert K, Le Loeuff J, Lutat P, Philippe M, Richter 18. Chonglakmani C. Chapter 6 Triassic. In: Ridd, M.F., U, Tong H. A new fi sh locality from the continental Late Barber, A.J. and Crow, M.J. (eds) The Geology of Jurassic–Early Cretaceous of Northeastern Thailand. Thailand. Geological Society, London 2011; 137-150. Revue de Paléobiologie 2004; 9: 161-167. 19. Buffetaut E, Suteethorn V, Tong H. The fi rst thyreo 27. Cavin L, Suteethorn V, Khansubha S, Buffetaut E, phoran dinosaur from Southeast Asia, a stegosaur Tong H. A new Semionotid (Actinopterygii, Neop- vertebra from the Late Jurassic Phu Kradung Forma- terygii) from the Late Jurassic–Early Cretaceous of tion of Thailand. Neues Jahrbuch für Geologie und Thailand. Comptes Rendus Palévol 2003; 2: 291-297. Paläontologie Monatshefte 2001; 95-102. 28. Deesri U, Cavin L, Claude J, Suteethorn V, Yuangdet- 20. Buffetaut E, Suteethorn V. A sinraptorid theropod kla P. Morphometric and taphonomic study of a ray- (Dinosauria, Saurischia) from the Phu Kradung fi nned fi sh assemblage (Lepidotes buddhabutrensis, 354 Cavin et al. J Sci Technol MSU

Semionotidae) from the Late Jurassic-earliest Creta- 33. Grande L. Phylogenetic study of gars and closely ceous of NE Thailand. In: Buffetaut, E., Cuny, G., Le related species, based mostly on skeletal morpho Loeuff, J. and Suteethorn, V. (eds) Late Palaeozoic logy. The resurrection of Holostei. In: Poyato-Ariza, and Mesozoic Ecosystems in SE Asia. Geological F.J. (ed.) Fourth International Meeting on Mesozoic Society, London, Special Publications 2009; 315: - Systematics, homology and Nomenclature. 115-124. UAM, Mirafl ores de la Sierra, Madrid, Spain 2005; 29. Deesri U. Osteology and scale microstructure of 119-121. Late Jurassic-Early Cretaceous Ginglymodians from 34. Newberry JS. Fossil fi shes and fossil plants of the Thailand, Evolutionary, Taxonomical and Palaeogeo- Triassic rocks of New Jersey and the Connecticut graphical implicantions. PhD Thesis. Mahasarakham Valley. Monographs of the United States Geological University. Thailand. 2013; 266 pp. Survey 188; 14: xiv+152 pp., pls. 1-26. 30. Deesri U, Suteethorn V, Liard R, Cavin L. First disco 35. Cavin L. Diversity of Mesozoic semionotiform fi shes very of a juvenile Thaiichthys (Actinopterygii, Holostei) and the origin of gars (Lepisosteidae). Naturwissen- from the Late Jurassic - Early Cretaceous of Thailand. schaften 2010; 97: 1035-1040. Journal of Science and Technology, Mahasarakham 36. Gibson SZ. and Evolutionary History of University. In press. Lophionotus (Neopterygii: Semionotiformes) from the 31. Buffetaut E, Cuny G, Le Loeuff J, Suteethorn V. Late Western United States. Copeia 2013; 4: 582–603. Palaeozoic and Mesozoic continental ecosystems of 37. Cavin L, Suteethorn V. A new Semionotiform (Acti SE Asia, an introduction. Geological Society, London, nopterygii, Neopterygii) from Upper Jurassic–Lower Special Publications 2009; 315: 1-5. Cretaceous deposits of North-East Thailand, with 32. Cavin L, Suteethorn V, Buffetaut E, Claude J, comments on the relationships of Semionotiforms. Cuny G, Le Loeuff J, Tong H. The fi rst sinamiid Palaeontology 2006; 49: 339-353. fi sh (Holostei, Halecomorpha) from South-east Asia (Early Cretaceous of Thailand). Journal of Vertebrate Paleontology 2007; 27(4): 827-837. Vol 33. No 4, July-August 2014 Ginglymodian fi shes (Actinopterygii, Holostei) from Thailand: An overview 355 Button-like teeth (Ginglymodi indet. 3) (Ginglymodi indet. 4) Smooth scales (Ginglymodi indet. 1) (Ginglymodi indet. 5) Smooth scales (Ginglymodi indet. 2) with inclusion of new discoveries. 1 g. 6 and table 1) fi Isanichthys palustris Isanichthys lertboosi (juvenile) buddhabutrensis . cf. New taxon Thaiichthys buddhabutrensis T. buddhabutrensis T Lam Pao Dam Large pitted scales Large button-like tooth Khok Pha Suam Ridged scales Kham Phok Chong Chat Phu Noi Ban Nong Rua Khok Sanam, etc. Thai fossil record of ginglymodians based on data from Cavin et al. (2009: KM Mab ChingHHL Huai Pha Pheng Ginglymodi indet. Large smooth scales Conical tooth FmsKK Localities Ban Saphan Hin Posterior part of a body Articulated materialSK Phu Wiang Various skull bones Isolated remains Smooth scales PK Phu Nam Jun Table 1 356 Cavin et al. J Sci Technol MSU

Figure 1 Plot of the of the Lepisosteiformes (from Deesri et al.201415) against palaeogeography. The upper row shows schematic evolution of the palaeogeographical pattern in the Late Jurassic and in the Cretaceous. The lower row shows the phylogenetic tree included in the palaeogeography. Vicariant events are favoured over dispersal events.