Kallokibotion: a late-surviving, basal, hard-shell turtle

A recent paper by Pérez-Garcia and Codrea 2018
on the basal, but late-surviving turtle, Kallokibotion bajazidi, (Nopsca 1923a,b) brings us more specimens and more hypotheses of turtle relationships. This taxon has been described as ‘an enigma’, perhaps because all currently known specimens combine Late Triassic traits with a Latest Cretaceous occurrence The visibility of the naris in lateral view goes back to the pre-turtle, Elginia. Transitional taxa, like Meiolania (Fig. 2) and Proganochelys, do not have this trait. Essentially this taxon links the Proganochelys clade to all later hard-shell turtles. (Remember, soft-shell turtles had a separate origin).

Figure 2. Kallokibotion skull in 5 views. Note the twin nares.

Figure 1. Kallokibotion skull in 5 views. Note the twin nares. This skull retains many meiolanid traits.

Gaffney and Meylan 1992 report, “Kallokibotion is a cryptodire because it has the otic trochlea synapomorphy of cryptodires, and it is a member of the Selmacryptodira because it has a posterior pterygoid process under the middle ear. It lacks the posterior temporal emargination synapomorphic of the Daiocryptodira and lies outside that group.”

I’m not going to discuss the suprageneric taxa listed above
because no prior turtle phylogenies recognized the diphyletic nature of soft-shell and hard-shell turtles.

Figure 3. Subset of the large reptile tree (LRT, 1199 taxa) with the addition of three basal turtles

Figure 2. Subset of the large reptile tree (LRT, 1199 taxa) with the addition of three basal turtles

Kallokibotion links
Niolamia and Miolania (Fig. 3 ) with higher turtles, skipping over Proganochelys (Late Triassic, Fig. 4), the traditional baalmost turtle.


Figure 3. Kallokibotion compared to Meiolania.

Figure 3. Kallokibotion compared to Meiolania.

Kallokibotion was recognized as a member of Meiolaniformes
in several recently published papers (see Rabi et al., 2013b; Sterli & de la Fuente, 2013; Sterli et al., 2015a, b). Chubutemys is considered a meiolaniform without horns, but workers who proposed this clade did not realize meiolaniids were basalmost hard-shell turtles.

Figure 2. The skull of Proganochelys, a basal turtle without skull invagation and without horns.

Figure 4. The skull of Proganochelys, a basal turtle close to Kallokibation, Note the squamosal and supratemporal and compare those to Kallokibation in  Fig. 1.

Figure 2. Kallokibation carapace and plastron from Gaffney and Melton 1992.

Figure 5. BMNH R4918 specimen of Kallokibation carapace and plastron from Gaffney and Melton 1992. This is the first of the tested taxa that has a short, unarmored tail. The skull was relatively small and unarmored.

Pérez-Garcia and Codrea 2018
discovered new specimens (Fig. 1) that “not only reveal detailed cranial and postcranial elements poorly known until now, refuting previous hypotheses about the anatomy of this taxon, but also allow us to identify numerous hitherto unknown characters.” The new information, “shows Kallokibotion as the sister taxon of the crown Testudines.”

A lot of discussion about crown-group turtles goes out the window
when taxa are included that split hard from soft-shell taxa before turtles had shells. Thus all known turtles, and several non-turtle pareiasaurs, fall under the current definition of crown turtles in the LRT.

(Fig. 3) Kallokibotion does indeed nest close to the meiolanids and Kayentachelys, but other taxa intervene. The loss of horns and a long armored tail on higher hard-shell turtles, like Kallokibotion, can be attributed to neotony.

Pérez-Garcia and Codrea had no idea what turtles are.
They nest the pre-lepidosauriform, Owenetta, the pareiasaur, Anthodon, the plesiosaur, Simosaurus, and the rhynchocephalian, Sphenodon as progressively more distant outgroup taxa. Throw those out and, except for the meiolanids, their pruned turtle cladogram is similar to the same subset of the LRT.

Dyke GJ et al. (20 co-authors) 2014. Thalassodromeus sebesensis—a new name for an old turtle. Comment on ‘Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur’, Grellet-Tinner and Codrea. Gondwana Research. doi:10.1016/j.gr.2014.08.004.
Gaffney ES and Meylan PA 1992. The Transylvanian turtle, Kallokibotion, a primitive cryptodire of Cretaceous Age. American Museum Novitates (3040).
Nopcsa F 1923a. On the geological importance of the primitive reptilian fauna of the Uppermost Cretaceous ofHungary; with a description of a new tortoise (Kallokibotion). Quart. Jour. Geol. Soc. 79(1): 100—116.
Pérez-García A and Vlad Codrea 2018. New insights on the anatomy and systematics of Kallokibotion Nopcsa, 1923, the enigmatic uppermost Cretaceous basal turtle (stem Testudines) from Transylvania. Zoological Journal of the Linnean Society. 182(2):419–443. doi:10.1093/zoolinnean/zlx037.
Zhou CF et al. 2015. A sinemydid turtle from the Jehol Biota provides insights into the basal divergence of crown turtles, Scientific Reports (2015). DOI: 10.1038/srep16299

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