Testudines (
turtles ,
tortoises , and
terrapins ). However, it was later suggested that the anapsid-like turtle skull is due to
reversion rather than to anapsid descent. The majority of modern paleontologists believe that the Testudines are descended from
diapsid reptiles that lost their temporal fenestrae. More recent morphological
phylogenetic studies with this in mind placed turtles firmly within
diapsids ,
[2] [3] [4] [5] [6] or, more commonly, within
Archelosauria .
[7]
Phylogenetic position of turtles
All
lepidosaurs (
lizards ,
snakes , and
tuataras ).
[14] However, in a later paper from the same authors, published in 2014, based on more extensive data, the archosauromorph hypothesis is supported.
[7]
Reanalysis of prior phylogenies suggests that they classified turtles as anapsids both because they assumed this classification (most of them were studying what sort of anapsid turtles are) and because they did not sample fossil and extant taxa broadly enough for constructing the
Permian-Triassic extinction event
.
Despite the molecular studies, there is evidence that contradicts their classification as diapsids. All known diapsids excrete uric acid as nitrogenous waste (uricotelic), and there is no known case of a diapsid reverting to the excretion of urea (ureotelism), even when they return to semi-aquatic lifestyles. Crocodilians, for example, are still uricotelic, although they are also partly ammonotelic, meaning they excrete some of their waste as ammonia. Ureotelism appears to be the ancestral condition among primitive amniotes, and it is retained by mammals, which likely inherited ureotelism from their synapsid and therapsid ancestors. Ureotelism therefore would suggest that turtles were more likely anapsids than diapsids. The only known uricotelic chelonian is the desert tortoise , which likely evolved it recently as adaptation to desert habitats. Some desert mammals are also uricotelic, so since practically all known mammals are ureotelic, uricotelic adaptation is a likely result of convergence among desert species. Therefore, turtles would have to be the only known case of a uricotelic reptile reverting to ureotelism.[citation needed ]
Anapsida in modern taxonomy
Anapsida is still sporadically recognized as a valid group, but this is not favoured by current workers.[17] [18] Anapsids in the traditional meaning of the word are not a clade, but rather a paraphyletic group composed of all the early reptiles retaining the primitive skull morphology, grouped together by the absence of temporal openings.[17] [18] Gauthier, Kluge and Rowe (1988) attempted to redefine Anapsida so it would be monophyletic, defining it as the clade containing "extant turtles and all other extinct taxa that are more closely related to them than they are to other reptiles".[19]
This definition explicitly includes turtles in Anapsida; because the phylogenetic placement of turtles within Amniota is very uncertain, it is unclear what taxa, other than turtles themselves, would be included in such defined Anapsida, and whether its content would be similar to the Anapsida of tradition. Indeed, Gauthier, Kluge and Rowe (1988) themselves included only turtles and Captorhinidae in their Anapsida, while excluding the majority of anapsids in the traditional sense of the word from it.[19]
Temporal openings in traditional anapsids
Tsuji and Müller (2009) noted that the name Anapsida implies a morphology (lack of temporal openings) that is in fact absent in the skeletons of a number of taxa traditionally included in the group.
See also
References
External links