Polyglyphanodontia

Polyglyphanodontia, also known as the Borioteiioidea, is an extinct clade of lizards from the Cretaceous that includes around a dozen genera. Polyglyphanodontians were the dominant group of lizards in North America[1] and Asia[2] during the Late Cretaceous. Most polyglyphanodontians are Late Cretaceous in age, though the oldest one, Kuwajimalla kagaensis, is known from the Early Cretaceous (Valanginian to Hauterivian) Kuwajima Formation (Japan).[3] Early Cretaceous South American taxon Tijubina, and possibly also Olindalacerta, might also fall within Polyglyphanodontia or be closely allied to the group, but if so, they would be two of only three Gondwanan examples of an otherwise Laurasian clade[4] (the third one, and the only unambiguous one, being Bicuspidon hogreli from the Kem Kem Beds of Morocco).[5] They produced a remarkable range of forms. Chamopsiids, including Chamops, were characterized by large, blunt, crushing teeth, and were most likely omnivores. Macrocephalosaurus, from the Gobi Desert, was a specialized herbivore; it grew to roughly a meter long and had multicusped, leaf-shaped teeth like those of modern iguanas. Polyglyphanodon, from the Maastrichtian of Utah, was another herbivore, but its teeth formed a series of transverse blades, similar to those of Trilophosaurus. Peneteius had remarkable, multicusped teeth, similar to those of mammals. The polyglyphanodontids first appear in the latter part of the Early Cretaceous in North America, and became extinct during the Cretaceous-Paleogene extinction event. Polyglyphanodontians closely resembled the teiid lizards, and purported teiid lizards from the Late Cretaceous appear to be polyglyphanodontians.[6] The only species known to have survived the Cretaceous was Chamops, which survived until the very early Ypresian (early Eocene).[7][8]

Polyglyphanodontia
Temporal range: Early Cretaceous-Early Eocene, Hauterivian–Ypresian
Skeleton of the polyglyphanodontian Polyglyphanodon sternbergi
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Squamata
Clade: Scincogekkonomorpha
Clade: †Polyglyphanodontia
Alifanov, 2000
Synonyms
  • Borioteiioidea Nydam et al., 2007

Classification
Tianyusaurus fossil

A large-scale phylogenetic analysis of squamates conducted by Conrad (2008) found polyglyphanodontians (called Polyglyphanodontidae by Conrad) to be closely related to teiid lizards. In the strict consensus tree recovered in the analysis, polyglyphanodontids were part of a polytomy (unresolved evolutionary relationship) with teiids, gymnophthalmids, Chamops and lacertids; in the Adams consensus tree polyglyphanodontids were the sister group to teiids.[9] (Some other studies that recognize a close relationship between polyglyphanodontians and teiids use the name Borioteiioidea rather than Polyglyphanodontia, although Borioteiioidea encompasses only North American polyglyphanodontians.)[10] Conrad's analysis also recovered Sineoamphisbaena, a Cretaceous lizard that resembles legless amphisbaenian lizards, as a member of Polyglyphanodontidae.[9] On the other hand, a later large-scale phylogenetic analysis of fossil and living squamates published in 2012 by Gauthier et al. found that Polyglyphanodontia was not particularly closely related to teiids, but rather that it was the sister taxon of the clade containing the extinct marine mosasaurs, their closest relatives and the major lizard group Scleroglossa. Because the first scleroglossans appear in the Late Jurassic, polyglyphanodontians must also have originated in the Late Jurassic if this phylogeny is correct. However, polyglyphanodontians are limited to the Cretaceous, meaning that a long ghost lineage may exist.[6] The primary analysis of Gauthier et al. (2012) did not find polyglyphanodontians and Sineoamphisbaena to be closely related; however, the authors noted that when all snake-like squamates and mosasaurs were removed from the analysis, and burrowing squamates were then added individually to it, Sineoamphisbaena grouped with polyglyphanodontians. Gauthier et al. (2012) considered it possible that Sineoamphisbaena was a burrowing polyglyphanodontian,[6] though, agreeing with.[9] Subsequent analyses based on [9] have also supported Sineoamphisbaenia as a polyglyphanodontian (the combined molecular and morphological analysis of Müller et al.;[11] in the morphology only analyses carried out by Wiens et al. but not their combined molecular and morphological analyses;[12] and in morphology only analyses carried out by [13][14] who were describing new polyglyphanodontians from Asia [Funuisaurus and Tianyusaurus]).

A phylogenetic analysis conducted by Reeder et al. (2015) based on combined molecular and morphological data (based on [6]) recovered Polyglyphanodontia as members of Toxicofera; specifically, it was recovered as the sister group of Iguania.[15] Lee (2009)[16] analyzed a combined molecular and morphological data set (based on earlier studies by Lee[17][18]) and also had found polyglyphanodontians as the sister to iguanians but Lee's finding was not acknowledged by.[15] Combined molecular and morphological data analyses using Conrad's data [11][12] consistently found polyglyphanodontians as the sisters to teiioids.

References
  1. Longrich N. R., Bhullar A.-B. S.; et al. (2012). "Mass extinction of lizards and snakes at the Cretaceous-Paleogene boundary". Proceedings of the National Academy of Sciences. 109 (52): 21396–21401. Bibcode:2012PNAS..10921396L. doi:10.1073/pnas.1211526110. PMC 3535637. PMID 23236177.
  2. Gao K.; Hou L. (1996). "Systematics and taxonomic diversity of squamates from the Upper Cretaceous Djadochta Formation, Bayan Mandahu, Gobi Desert, People's Republic of China". Canadian Journal of Earth Sciences. 33 (4): 578–598. Bibcode:1996CaJES..33..578G. doi:10.1139/e96-043.
  3. Susan E. Evans & Makoto Manabe (2008). "An early herbivorous lizard from the Lower Cretaceous of Japan". Palaeontology. 51 (2): 487–498. doi:10.1111/j.1475-4983.2008.00759.x.
  4. Tiago R. Simões, Michael W. Caldwell and Alexander W. A. Kellner (2015). "A new Early Cretaceous lizard species from Brazil, and the phylogenetic position of the oldest known South American squamates". Journal of Systematic Palaeontology. 13 (7): 601–614. doi:10.1080/14772019.2014.947342.
  5. Romain Vullo; Jean-Claude Rage (2018). "The first Gondwanan borioteiioid lizard and the mid-Cretaceous dispersal event between North America and Africa". The Science of Nature. 105 (11–12): Article 61. Bibcode:2018SciNa.105...61V. doi:10.1007/s00114-018-1588-3. PMID 30291449.
  6. Gauthier, J. A.; Kearney, M.; Maisano, J. A.; Rieppel, O.; Behlke, A. D. B. (2012). "Assembling the Squamate Tree of Life: Perspectives from the Phenotype and the Fossil Record". Bulletin of the Peabody Museum of Natural History. 53: 3–308. doi:10.3374/014.053.0101.
  7. D. E. Russell. 1967. Le Paleocene continental d'Amerique du nord. Memoires du Museum National d'Histoire Naturelle. Serie C., Sciences de la Terre 16(2):37-99
  8. Marsh, O.C. (1892). "Notice of new reptiles from the Laramie formation". American Journal of Science. 43.
  9. Conrad J (2008). "Phylogeny and systematics of Squamata (Reptilia) based on morphology" (PDF). Bulletin of the American Museum of Natural History. 310: 1–182. doi:10.1206/310.1. hdl:2246/5915.
  10. Nydam, R. L.; Caldwell, M. W.; Fanti, F. (2010). "Borioteiioidean lizard skulls from Kleskun Hill (Wapiti Formation; upper Campanian), west-central Alberta, Canada". Journal of Vertebrate Paleontology. 30 (4): 1090–1099. doi:10.1080/02724634.2010.483539.
  11. Müller Johannes; Hipsley Christy; Head Jason; Kardjilov Nikolay; Hilger André; Wuttke Michael; Reisz Robert (2011). "Eocene lizard from Germany reveals amphisbaenian origins". Nature. 473 (7347): 364–367. Bibcode:2011Natur.473..364M. doi:10.1038/nature09919. PMID 21593869.
  12. John Wiens, Caitlin Kuczynski, Ted Townsend, Tod Reeder, Daniel Mulcahy, and Jack Sites, Jr. "Combining phylogenomics and fossils in higher-level squamate reptile phylogeny: molecular data change the placement of fossil taxa. Syst Biol 59 (6), 674-688 (2010).
  13. Mo JY, Xu X, Evans SE (2010). "The evolution of the lepidosaurian lower temporal bar: new perspectives from the Late Cretaceous of South China". Proc R Soc Lond B. 277 (1679): 331–336. doi:10.1098/rspb.2009.0030. PMC 2684617. PMID 19324758.
  14. XU L., WU, Junchang L., JIA, Zhang J., PU, Zhang X. (2014). "A New Lizard (Lepidosauria: Squamata) from the Upper Cretaceous of Henan, China". Acta Geologica Sinica. 88 (4): 1041–1050. doi:10.1111/1755-6724.12271.CS1 maint: multiple names: authors list (link)
  15. Tod W. Reeder; Ted M. Townsend; Daniel G. Mulcahy; Brice P. Noonan; Perry L. Wood Jr.; Jack W. Sites Jr.; John J. Wiens (2015). "Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa". PLoS ONE. 10 (3): e0118199. Bibcode:2015PLoSO..1018199R. doi:10.1371/journal.pone.0118199. PMC 4372529. PMID 25803280.
  16. Michael S. Y. Lee (2009). "Hidden support from unpromising data sets strongly unites snakes with anguimorph 'lizards'". J Evol Biol. 22 (6): 1308–1316. doi:10.1111/j.1420-9101.2009.01751.x. PMID 19490385.
  17. Lee Michael S.Y. (2005). "Squamate phylogeny, taxon sampling and data congruence". Org Divers Evol. 5: 25–45. doi:10.1016/j.ode.2004.05.003.
  18. Lee Michael S.Y. (2005). "Molecular evidence and marine snake origins". Biology Letters. 1 (2): 227–230. doi:10.1098/rsbl.2004.0282. PMC 1626205. PMID 17148173.
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