Fossil Friday: Is the Four-Legged Snake Tetrapodophis a Missing Link or Not?
The origin of snakes is a hotly debated topic in evolutionary biology. There are controversies about their phylogenetic position, their origin from aquatic vs burrowing (fossorial) ancestors, and of course transitional fossils (Caldwell & Lee 1997, Rieppel et al. 2003, Brandley et al. 2008, ApesteguÃa & Zaher 2006, Lee et al. 2007, Caldwell 2007, 2008, Zaher et al. 2009, 2023, Longrich et al. 2012, Zaher & Scanferla 2012, Palci et al. 2013a, 2013b, Palci 2014, Caldwell et al. 2015, Hsiang et al. 2015, Yi & Norell 2015, Da Silva et al. 2018, Xing et al. 2018, Garbergoglio 2019a, 2019b, 2019c). While some scientists are convinced that “snakes and mosasauroids/coniasaurs share a limbed common ancestor that was likely aquatic, not fossorial” (Caldwell 2008), other scientists strictly claim that “lizards could not have transitioned to snakes by any other evolutionary path than through fossoriality” (Da Silva et al. 2018). Either way, there should have existed transitional forms, which exhibit at least some typical features of basal snake anatomy combined with four fully developed fore and hind limbs to document the transition of limbless snakes from still quadrupedal walking ancestors.
Assumed Mesozoic stem snakes like the Madtsoiidae and Simoliophiidae as well as the very primitive Cretaceous genera Coniophis, Dinilysia, and Najash did only possess small external hind limbs (of which vestigial remnants are also retained in some living snakes like boids), but show no evidence of forelimbs (Tchernov et al. 2000, Zaher & Rieppel 2002, Rieppel et al. 2003, ApesteguÃa & Zaher 2006, Garberoglio et al. 2019c).
In their study of the evolution of the snake body plan Garberoglio et al. (2019c) commented:
This basal position of the limbed terrestrial and marine snakes indicates that snakes retain sizeable external hindlimbs and sacral contacts for a substantial time after their origin—from approximately 170 Ma to the youngest confirmed legged snakes, the simoliophiids, at approximately 100 Ma. This indicates that (i) the reduction and loss of the pectoral girdle and forelimbs probably occurred much earlier, given the definitive absence of these structures in simoliophiids and the lack of evidence for their presence in Najash, Dinilysia, and madtsoiids, and was probably a major event in the early radiation of snakes, and one that occurred well before crown (modern) snake origins; (ii) the “forelimb-absent and hindlimb-reduced” morphology was a stable and successful body plan, rather than a transient phase between limbed and limbless conditions; and (iii) the origin of crown snakes was characterized by a major reduction in the hindlimb and pelvis (including loss of sacral contacts).
This All Sounds Well and Good
That is, until you find that other modern studies (e.g., Tchernov et al. 2000, Rieppel et al. 2002, Zaher & Rieppel 2002, Longrich et al. 2012, Zaher & Scanferia 2012, Palci et al. 2013b in part, Hsiang et al. 2015, Garbergoglio et al. 2019a contra Garbergoglio et al. 2019c) do not even agree that the fossil Madtsoniidae and Simoliophidae or even Dinilysia are stem group snakes at all, but recover them nested among living snakes, which would imply that the hind limb reduction occurred more than once in crown group snake. Probably, mainstream evolutionary biologists would respond that such a reduction is simple and therefore not unlikely, as is proven by the fact that several other groups of amphibians and reptilians have produced legless forms. However, this is exactly the point: convergence is all over the place in the animal kingdom and is nothing but incongruent similarity that contradicts a unique nested hierarchy of similarities predicted by Darwinism. Furthermore, there is experimental evo-devo evidence from transgenic mice that the reduction of limbs is not a simple task at all but requires multiple codependent mutations (compare Kvon et al. 2016). Together with the complex changes in skull kinetics of snakes, this raises a significant waiting time problem, because even the geologically available window of time of several million years is orders of magnitude too short to accommodate the genetic changes for this re-engineering of the body plan, based on the mathematical toolkit of mainstream population genetics.
Anyway, because of the fact that no known fossil stem snakes had retained four functional limbs, the discovery of a putative four-legged snake was a real sensation. Eight years ago my good colleague Prof. David Martill, with whom I co-authored several articles and monograph of the fossils of the Lower Cretaceous Crato Formation in Brazil (Martill et al. 2007), described from this locality the long sought missing link of snake evolution in the prestigious journal Science (Martill et al. 2015; also see this YouTube video): the beautifully and completely preserved fossil shows a very elongate snake-like animal with small fore and hind limbs, both with five well-developed digits. They named the 110-million-year-old fossil Tetrapodophis, which means four-legged snake. Their phylogenetic analysis placed it at the very base of the snake lineage. The scientists also identified burrowing adaptations that arguably would support an origin of snakes from fossorial rather than aquatic ancestors. Of course, the discovery was much celebrated in news reports as “a snake version of Archaeopteryx” (e.g., University of Portsmouth 2015, Christakou 2015a, Evans 2015, Yong 2015)
Yong (2015) commented in National Geographic:
Their analysis produced a family tree in which Tetrapodophis came after the earliest known snakes like Eophis, Parviraptor, and Diablophis, but is still very much a snake. But how could that be? Eophis and the others only have two legs, so how could four-legged Tetrapodophis have come after them? The answer is that evolution doesn’t proceed along simple, straight lines.
This is typical Darwinist doublespeak for the simple fact that the fossil does not fit within a Darwinian scenario without ad hoc hypotheses to explain away conflicting data. But in this case there is another little problem: all known fossils of the Jurassic earliest stem snakes (Portugalophis, Eophis, Parviraptor, and Diablophis) do not even have the body region preserved, which would show if they possessed limbs or not (see Caldwell et al. 2015), so that the above statement is also incorrect. Never believe Darwinist pop science journalists, who are notoriously sloppy with facts. But apart from this glitch, the real blows against Tetrapodophis were yet to come.
A First Blow
The first blow came with a very heated discussion about the legal status of the fossil (Christakou 2015b), even though there was no proof that any laws were broken with the export and sale of the fossil. I personally consider this discussion as very much ridiculous, given the fact that countless fossils from the quarries in Brazil were grinded for cement production or used for pavements, without any local scientist caring for this natural heritage. Now, museums around the world, which invested a lot of money and preparation effort to preserve these fossils for science, are now asked to repatriate the collections to Brazilian museums with a very poor track record of proper conservation and curation, all in the name of protecting alleged national treasures of poor countries from evil Western exploitation. Unfortunately, the pervasive woke agenda has taken hold of paleontology as well, which impairs scientific progress and has made fossil collecting to a very frustrating endeavour.
A Second Blow
The second blow came with another silly dispute about the deposition of the specimen. The specimen was deposited in a public museum in Germany as a loan by a private collector, who temporarily limited access to the specimen because it was damaged during the examination with synchrotron micro-CT scanning. Even though access was later restated and the specimen was also meticulously documented with drawings and photographs in the original publication, some scientists made absurd statements like “if the fossil can’t be studied, it doesn’t exist … I don’t want to mention the name Tetrapodophis ever again” (Gauthier quoted in Gramling 2016). Other researchers said the “original paper should not have been published, because the fossil was not officially deposited in a museum or other repository”. This reflects a common politically correct sentiment among scientists against private collections, even though many private fossil collections are better curated and more easily accessible to scientists than public collections in certain dubious countries with a reputation of sloppiness and corruption. Nevertheless, these bigoted scientists rather would leave sensational finds of high scientific value undescribed forever, than having scientific publications based on material in private collections. I find this bizarre, scientifically reprehensible, and politically highly problematic (anti private property in favor of statist bureaucracy). After all, using the same kind of reasoning we could purge all paleontological knowledge from science that is based on museum specimens that were destroyed during World War II, or lost by the post when they were sent on loan to foreign scientists, or simply cannot be relocated in the museum archives anymore, which happens way more often than many people may think. Many scientists seem to have lost their common sense nowadays.
Last but Not Least
The third and truly fatal blow came with several studies by paleontologist Michael Caldwell and his colleagues, who strongly disputed the fossorial adaptation of Tetrapodophis in favor of an aquatic adaptation, and also disputed the snake-affinity in favor of a determination as dolichosaurid lizard (Lee et al. 2016, Caldwell et al. 2016, 2021, Paparella et al. 2018), which is an extinct group of marine reptiles. The press immediately reported that the big splash was a case of mistaken identity (Geggel 2016), and that the assumed missing link between lizards and snakes has been debunked (Hodžić_2021, Lyle 2021, Strauss 2022).
One might think that this misidentification is a minor issue because dolichosaurids are often considered as close relatives of snakes within a clade Pythonomorpha. However, this opens a whole new can of worms, as the status of Pythonomorpha is subject of considerable scientific controversy itself. The Wikipedia article on Pythonomorpha has a good summary of this mess, of which the long story short is as follows: Pythonomorpha was proposed by the famous 19th century paleontologist Edward Drinker Cope (1869) to include mosasaurs and snakes as assumed close relatives, while snakes were believed to have evolved from burrowing lizards. This was rejected by most 20th century experts, who instead suggested that the closest relatives of mosasaurs are monitor lizards (e.g., Russell 1967). Then, Pythonomorpha was resurrected by Michael Caldwell and Michael Lee in the late 1990s (Caldwell & Lee 1997, Lee 1997, 1998, Caldwell 1999, Lee & Caldwell 2000) to include the aquatic extinct reptile groups aigialosaurs, mosasaurs, dolichosaurs, coniasaurs, as well as snakes, which were postulated as derived from aquatic ancestors (also see Pierce & Caldwell 2004, Palci & Caldwell 2007, Caldwell 2008, Paparella et al. 2018). This was again questioned by anatomical evidence (Zaher & Rieppel 1999) and more recent phylogenetic studies, which again placed mosasaurs with monitor lizards and snakes with skinks (Conrad 2008) or with anguinids and iguanas (Simões et al. 2020), or placed mosasaurs more basal than monitor lizards and snakes (Gauthier et al. 2012). On the other hand, more recent analyses (Palci et al. 2013a, Palci 2014, Martill et al. 2015, Reeder et al. 2015) recovered snakes and mosasaurs as sister groups and thus supported a monophyly of Pythonomorpha. The latter would also be congruent with the growing for a monophyletic group called Toxicofera, which would include all squamates with poison glands such as iguanas, monitor lizards, gila monsters, and snakes, under exclusion of skinks, gekkos, and true lizards. However, as always in phylogenetics there is also strong conflicting evidence presented by opponents of the Toxicofera hypothesis (Hargreaves et al. 2015, Mongiardino Koch & Gauthier 2018, Joffroy 2022). If we would draw a least common denominator of all published trees of the past decades, the result would be an unresolved polytomy (as I described in a recent Fossil Friday article about arachnid phylogeny), which rather agrees with the expectations of creationists than those of evolutionary biologists.
A New Turn of Events
Considering this uncertainty it is hardly a surprise that this year a new study brought a new turn of events and again confirmed the status of Tetrapodophis as most basal snake (Zaher et al. 2023). The authors remarked that “Our scorings of Tetrapodophis were based on first-hand observations of the original specimen, and our interpretations of the anatomy and attendant scoring disagree significantly with those of Caldwell et al. (2021), as do our phylogenetic results.” It does not take too much of a prophetic gift to predict that we will not have to wait very long for a rebuttal by Caldwell and his team.
Whoever may ultimately win this dispute, there is no doubt that the true reason for all this uncertainty is the simple fact that the pattern of similarities between different animal groups does not fall into a neatly ordered set of nested hierarchies as often pretended by hardcore Darwinists. In reality, incongruent patterns are all over the place. A snake-like body plan is a good example: Among tetrapod vertebrates a snake-like body only originated in caecilians (Gymnophiona) among lissamphibians and in squamates among amniotes. However, within amniotes this body plan originated not less than 26 times independently (Brandley et al. 2008, Evans 2015). Tetrapodophis could be a missing link to at least two of them respectively, or just might be number 27, who knows. What is also interesting is that scientists found “statistically significant support for at least six examples of the re-evolution of lost digits in the forelimb and hind limb” ((Brandley et al. 2008), which shows the level of absurdity that is readily accepted only to preserve the evolutionary paradigm.
References
ApesteguÃa S & Zaher H 2006. A Cretaceous terrestrial snake with robust hindlimbs and a sacrum. Nature 440(7087), 1037–1040. DOI: https://doi.org/10.1038/nature04413
Brandley MC, Huelsenbeck JP, Wiens JJ 2008. Rates and patterns in the evolution of snake-like body form in squamate reptiles: evidence for repeated re-evolution of lost digits and long-term persistence of intermediate body forms. Evolution 62(8), 2042–2064. DOI: https://doi.org/10.1111/j.1558-5646.2008.00430.x
Caldwell MW 1999. Squamate phylogeny and the relationships of snakes and mosasauroids. Zoological Journal of the Linnean Society 125(1), 115–147. DOI: https://doi.org/10.1111/j.1096-3642.1999.tb00587.x
Caldwell MW 2007. Snake phylogeny, origins, and evolution: the role, impact, and importance of fossils (1869–2006). pp. 253–302 in: Anderson JS & Sues H-D (eds). Major Transitions in Vertebrate Evolution. Indiana University Press: Indiana, 417 pp.
Caldwell MW 2008. Squamate phylogeny and the relationships of snakes and mosasauroids. Zoological Journal of the Linnean Society 125(1), 115–147. DOI: https://doi.org/10.1111/j.1096-3642.1999.tb00587.x
Caldwell MW & Lee MSY 1997. A snake with legs from the marine Cretaceous of the Middle East. Nature 386, 705–709. DOI: https://doi.org/10.1038/386705a0
Caldwell MW, Nydam RL, Palci A & ApesteguÃa S 2015. The oldest known snakes from the Middle Jurassic-Lower Cretaceous provide insights on snake evolution. Nature Communications 6: 5996, 1–11. DOI: https://doi.org/10.1038/ncomms6996
Caldwell MW, Reisz RR, Nydam RL, Palci A & Simões TR 2016. Tetrapodophis amplectus (Crato Formation, Lower Cretaceous, Brazil) is not a snake. Society of Vertebrate Paleontology 76th Annual Meeting Program & Abstracts, 108. https://vertpaleo.org/wp-content/uploads/2021/03/SVP-2016-Program-Book-v10-with-covers.pdf
Caldwell MW, Simões TR, Palci A, Garberoglio FF, Reisz RR, Lee MSY & Nydam RL 2021. Tetrapodophis amplectus is not a snake: re-assessment of the osteology, phylogeny and functional morphology of an Early Cretaceous dolichosaurid lizard. Journal of Systematic Palaeontology 19(13), 893–952. DOI: https://doi.org/10.1080/14772019.2021.1983044
Christakou A 2015a. Four-legged fossil snake is a world first. Nature July 23, 2015. https://doi.org/10.1038/nature.2015.18050
Christakou A 2015b. Four-legged snake fossil sparks legal investigation. Nature August 4, 2015. DOI: https://doi.org/10.1038/nature.2015.18116
Conrad JL 2008. Phylogeny And Systematics Of Squamata (Reptilia) Based On Morphology. Bulletin of the American Museum of Natural History 310, 1–182. DOI: https://doi.org/10.1206/310.1Cope ED 1869. On the reptilian orders Pythonomorpha and Streptosauria. Proceedings of the Boston Society of Natural History 12, 250–266. https://www.biodiversitylibrary.org/partpdf/243473
Da Silva FO, Fabre A-C, Savriama Y, Ollonen J, Mahlow K, Herrel A, Müller J & Di-Poï N 2018. The ecological origins of snakes as revealed by skull evolution. Nature Communications 9: 376, 1–11. DOI: https://doi.org/10.1038/s41467-017-02788-3
Evans S 2015. Four legs too many? Science 349(6246), 374–375. DOI: https://doi.org/10.1126/science.aac5672
Garberoglio FF, Gómez RO, ApesteguÃa S, Caldwell MW, Sánchez ML & Veiga G 2019a. A new specimen with skull and vertebrae of Najash rionegrina (Lepidosauria: Ophidia) from the early Late Cretaceous of Patagonia. Journal of Systematic Palaeontology 17(18), 1313–1330. DOI: https://doi.org/10.1080/14772019.2018.1534288
Garberoglio FF, Gómez RO, Simões TR, Caldwell MW & ApesteguÃa S 2019b. The evolution of the axial skeleton intercentrum system in snakes revealed by new data from the Cretaceous snakes Dinilysia and Najash. Scientific Reports 9(1): 1276, 1–10. DOI: https://doi.org/10.1038/s41598-018-36979-9
Garberoglio FF, ApesteguÃa S, Simões TR, Palci A, Gómez RO, Nydam RL, Larsson HCE, Lee MSY & Caldwell Michael W 2019c. New skulls and skeletons of the Cretaceous legged snake Najash, and the evolution of the modern snake body plan. Science Advances 5(11): eaax5833, 1–8. DOI: https://doi.org/10.1126/sciadv.aax5833
Gauthier JA, Kearney M, Maisano JA, Rieppel O & Behkke ADB 2012: Assembling the Squamate Tree of Life: Perspectives from the Phenotype and the Fossil Record. Bulletin of the Peabody Museum of Natural History 53(1), 3–308. DOI: https://doi.org/10.3374/014.053.0101
Geggel L 2016. Mistaken Identity? Debate Over Ancient 4-Legged Snake Heats Up. LiveScience October 28, 2016. https://www.livescience.com/56685-debate-about-four-legged-snake-fossil.html
Gramling C 2016. ‘Four-legged snake’ may be ancient lizard instead. Science 354(6312), 536–537. DOI: https://doi.org/10.1126/science.354.6312.536 (Update Science November 11, 2016. https://www.science.org/content/article/update-controversial-four-legged-snake-may-be-ancient-lizard-instead)
Hargreaves AD, Tucker AS & Mulley JF 2015. A Critique of the Toxicoferan Hypothesis. pp. 1-15 in: Gopalakrishnakone P & Malhotra A (eds). Evolution of Venomous Animals and Their Toxins. Toxinology. Springer: Dordrecht (NL). DOI: https://doi.org/10.1007/978-94-007-6727-0_4-1Hodžić J 2021. Debunked: controversial fossil linking lizards to first snakes. BigThink November 27, 2021. https://bigthink.com/the-past/tetrapodophis/
Hsiang AY, Field DJ, Webster TH, Behlke ADB, Davis MB, Racicot RA & Gauthier JA 2015. The origin of snakes: revealing the ecology, behavior, and evolutionary history of early snakes using genomics, phenomics, and the fossil record. BMC Evolutionary Biology 15: 87, 1–22. DOI: https://doi.org/10.1186/s12862-015-0358-5
Joffroy K 2022. Squamata phylogenomics and molecular evolution of venom proteins in Toxicofera. Master thesis Christian-Albrechts-University Kiel, 77 pp. https://pure.mpg.de/rest/items/item_3430012_2/component/file_3430013/content
Kvon EZ et al. 2016. Progressive Loss of Function in a Limb Enhancer during Snake Evolution. Cell 167(3), 633–642. DOI: https://doi.org/10.1016/j.cell.2016.09.028
Lee MSY 1997. The phylogeny of varanoid lizards and the affinities of snakes. Philosophical Transactions of the Royal Society of London B 352(1349), 53–91. DOI: https://doi.org/10.1098/rstb.1997.0005
Lee MSY 1998. Convergent evolution and character correlation in burrowing reptiles: towards a resolution of squamate relationships. Biological Journal of the Linnean Society 65(4), 369–453. DOI: https://doi.org/10.1006/bijl.1998.0256
Lee MSY & Caldwell MW 2000. Adriosaurus and the Affinities of Mosasaurs, Dolichosaurs, and Snakes. Journal of Paleontology 74(5), 915–937. https://www.jstor.org/stable/1306991
Lee MSY, Hugall AF, Lawson R & Scanlon JD 2007. Phylogeny of snakes (Serpentes): combining morphological and molecular data in likelihood, Bayesian and parsimony analyses. Systematics and Biodiversity 5(4), 371–389. DOI: https://doi.org/10.1017/S1477200007002290
Lee MSY, Palci A, Jones MEH, Caldwell MW, Holmes JD & Reisz RR 2016. Aquatic adaptations in the four limbs of the snake-like reptile Tetrapodophis from the Lower Cretaceous of Brazil. Cretaceous Research 66, 194–199. DOI: https://doi.org/10.1016/j.cretres.2016.06.004
Longrich NR, Bhullar B-AS & gauthier JA 2012. A transitional snake from the Late Cretaceous period of North America. Nature 488 (7410), 205–208. DOI: https://doi.org/10.1038/nature11227
Lyle A 2021. Paleontologists debunk fossil thought to be missing link between lizards and first snakes. University of Alberta Folio November 18, 2021. https://www.ualberta.ca/folio/2021/11/paleontologists-debunk-fossil.htmlMartill DM, Bechly G & Loveridge RF (eds) 2007. The Crato Fossil Beds of Brazil: Window into an Ancient World. Cambridge University Press: Cambridge (UK), xvi+625 pp. https://www.cambridge.org/at/universitypress/subjects/earth-and-environmental-science/palaeontology-and-life-history/crato-fossil-beds-brazil-window-ancient-world?format=HB&isbn=9780521858670
Martill DM, Tischlinger H & Longrich NR 2015. A four-legged snake from the Early Cretaceous of Gondwana. Science 349(6246): 416–419. DOI: https://doi.org/10.1126/science.aaa9208
Mongiardino Koch N & Gauthier JA 2018. Noise and biases in genomic data may underlie radically different hypotheses for the position of Iguania within Squamata. PLoS ONE 13(8): e0202729, 1–29. DOI: https://doi.org/10.1371/journal. pone.0202729
Palci A 2014. On the Origin and Evolution of the Ophidia. PhD thesis University of Alberta, viii+474 pp. https://era.library.ualberta.ca/items/08055594-6458-4185-a058-0dfe62841f24
Palci A & Caldwell MW 2007. Vestigial forelimbs and axial elongation in a 95 million-year-old non-snake squamate. Journal of Vertebrate Paleontology 27(1), 1–7. DOI: https://doi.org/10.1671/0272-4634(2007)27[1:VFAAEI]2.0.CO;2
Palci A, Caldwell MW & Albino AM 2013a. Emended diagnosis and phylogenetic relationships of the Upper Cretaceous fossil snake Najash rionegrina ApesteguÃa and Zaher, 2006. Journal of Vertebrate Paleontology 33(1), 131–140. DOI: https://doi.org/10.1080/02724634.2012.713415
Palci A, Caldwell MW & Nydam RL 2013b. Reevaluation of the anatomy of the Cenomanian (Upper Cretaceous) hind-limbed marine fossil snakes Pachyrhachis, Eupodophis, and Haasiophis. Journal of Vertebrate Paleontology 33(6), 1328–1342. DOI: https://doi.org/10.1080/02724634.2013.779880
Paparella I, Palci A, Nicosia U & Caldwell MW 2018 A new fossil marine lizard with soft tissues from the Late Cretaceous of southern Italy. Royal Society Open Science 5: 172411, 1–27. DOI: https://doi.org/10.1098/rsos.172411
Pierce SE & Caldwell MW 2004. Redescription and phylogenetic position of the Adriatic (Upper Cretaceous; Cenomanian) dolichosaur Pontosaurus lesinensis (Kornhuber, 1873). Journal of Vertebrate Paleontology 24(2), 373–386. DOI: https://doi.org/10.1671/1960
Reeder TW, Townsend TM, Mulcahy DG, Noonan BP, Wood PL, Sites JW & Wiens JJ 2015. Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa. PLOS One 10(3): e0118199, 1–22. DOI: https://doi.org/10.1371/journal.pone.0118199
Rieppel O, Kluge AG & Zaher H 2002. Testing the phylogenetic relationships of the Pleistocene snake Wonambi naracoortensis Smith. Journal of Vertebrate Paleontology 22(4), 812–829. DOI: https://doi.org/10.1671/0272-4634(2002)022[0812:TTPROT]2.0.CO;2
Rieppel O, Zaher H, Tchernov E & Polcyn MJ 2003. The anatomy and relationships of Haasiophis terrasanctus, a fossil snake with well-developed hind limbs from the mid-Cretaceous of the Middle East. Journal of Paleontology 77(3), 536–558. DOI: https://doi.org/10.1666/0022-3360(2003)077<0536:TAAROH>2.0.CO;2
Russell DA 1967. Systematics and morphology of American mosasaurs (Reptilia, Sauria). Bulletin of the Peabody Museum of Natural History 23, vii+240 pp. https://elischolar.library.yale.edu/peabody_museum_natural_history_bulletin/23/
Simões TR, Vernygora O, Caldwell MW & Pierce SE 2020. Megaevolutionary dynamics and the timing of evolutionary innovation in reptiles. Nature Communications 11: 3322, 1–14. DOI: https://doi.org/10.1038/s41467-020-17190-9
Strauss K 2022. Researchers discover controversial four-legged ‘snake’ is a different ancient animal. Phys.org January 5, 2022. https://phys.org/news/2022-01-controversial-four-legged-snake-ancient-animal.html
Tchernov E, Rieppel O, Zaher H, Polcyn MJ & Jacobs LL 2000. A fossil snake with limbs. Science 281(5460), 2010–2012. DOI: https://doi.org/10.1126/science.287.5460.2010
University of Portsmouth 2015. Scientists Discover Four-Legged Snake Fossil. SciTechDaily August 2, 2015. https://scitechdaily.com/scientists-discover-four-legged-snake-fossil/Zaher H & Rieppel O 1999. Tooth implantation and replacement in squamates, with special reference to mosasaur lizards and snakes. American Museum Novitates 3271, 1–19. https://www.biodiversitylibrary.org/bibliography/91347
Zaher H & Rieppel O 2002. On the phylogenetic relationships of the Cretaceous snakes with legs, with special reference to Pachyrhachis problematicus (Squamata, Serpentes). Journal of Vertebrate Paleontology 22(1), 104–109. DOI: https://doi.org/10.1671/0272-4634(2002)022[0104:OTPROT]2.0.CO;2
Zaher H & Scanferla CA 2012. The skull of the Upper Cretaceous snake Dinilysia patagonica Smith-Woodward, 1901, and its phylogenetic position revisited. Zoological Journal of the Linnean Society 164(1), 194–238. DOI: https://doi.org/10.1111/j.1096-3642.2011.00755.x
Zaher H, ApesteguÃa S & Scanferla CA 2009. The anatomy of the Upper Cretaceous snake Najash rionegrina ApesteguÃa & Zaher, 2006, and the evolution of limblessness in snakes. Zoological Journal of the Linnean Society 156(4), 801–826. DOI: https://doi.org/10.1111/j.1096-3642.2009.00511.x