Temporal range: Campanian-Maastrichtian
~84–66 Ma
B. variolosa skeleton, Royal Tyrrell Museum
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Testudines
Suborder: Cryptodira
Family: Nanhsiungchelyidae
Genus: Basilemys
Hay 1902
Type species
Basilemys variolosa
(Cope, 1876)
  • B. gaffneyi Sullivan et al. 2012
  • B. morrinensis Mallon & Brinkman 2018
  • B. praeclara Hay 1911
  • B. sinuosa Riggs 1906
  • B. variolosa (Cope 1876) (type)

Basilemys (IPA: [bæsɪlɛmiːs]) is a large, terrestrial trionychoid turtle from the Upper Cretaceous.[1] In Greek, the word "Basil" means royal or kingly and the word "Emys" means turtle. Therefore, Basilemys means King Turtle. Basilemys has been found in rocks dating to the Campanian and Maastrichtian subdivisions of the Late Cretaceous and is considered to be the largest terrestrial turtle of its time.[2] Basilemys has solely been found in North America.[3] The family Nanhsiungchelyidae, which is the family Basilemys belongs to, made its first appearance in the Lower Cretaceous, in what we now call Asia. Because of Basilemys, we know that this family appeared in North America in the Upper Cretaceous.[4] It is possible that Basilemys and other nanhsiungchelyids are immigrants from Asia. They might have arrived in North America by passing through what we now call the Bering Strait somewhere during the Cretaceous.[4] In an analysis made by Sukhanov et al. on a new Nansiunghelyid turtle from the Upper Cretaceous of Mongolia, it was demonstrated that Asian nanhsiungchelyids gave rise to the North American nanhsiungchelyids.[5] Basilemys shares some traits with another member of nanhsiungchelyidae, Zangerlia, which is similar to Basilemys in, for example, skull proportions.[6] However, Basilemys has a more complex triturating surface that includes well-defined pockets on the dentary, and it also has tooth-like projections on the triturating surface of the maxilla.[6]

From the species in nanhsiungchelyidae, Basilemys is considered to be most similar to tortoises.[3] Many paleontologists have described the behaviors of Basilemys to likely be comparable to that of tortoises, due to living in terrestrial habitats and consuming tough plants.[3] Moreover, the complex triturating surface of Basilemys indicates that they are similar to tortoises in being terrestrial herbivores.[6] Basilemys is easily distinguishable from other fossil turtles due to how thick its shell is, the intricate sculpture of rows of triangular tubercles separated by pits, and its reduced inframarginal scales.[2] The fossil record is abundant with material from the shell and with appendicular, but cranial and cervical material is quite rare for Basilemys.[1]

Geological Information and Discovery

Turtles were prominent members of the Upper Cretaceous and thus, their specimens found throughout North America are useful in defining biogeographic patterns.[7] In 1924, a partial skeleton and crushed skull of Basilemys was collected by C.M. Sternberg from the Horseshoe Canyon Formation.[1] However, this specimen was poorly preserved and information of the skull could not be analyzed.[1] In another part of Canada, a Basilemys specimen from the Frenchman Formation of Saskatchewan retained most of the neck and cranial fragments.[1] This was groundbreaking because of how scarce cranial and cervical material is for Basilemys. Most of the well-preserved specimens of Basilemys are of the carapace, the hard upper shell of a turtle. Apart from the type species Basilemys variolosa, five other species have been described to date. These include B. gaffneyi, B. morrinensis, B. nobilis, B. praeclara and B. sinuosa. [1] In 2018, a nearly complete shell of B. morrinensis was found from the Horseshoe Canyon Formation of Alberta.[3] Basilemys specimens have also been found from the Oldman Formation and Dinosaur Park Formation.[6]

Other shell fragments of Basilemys have been found in the El Gallo Formation, located in Baja California.[7] Like the Horseshoe Canyon Formation and the Frenchman Formation, the El Gallo Formation is a rich fossiliferous geological unit.[7] The three shell fragments that were found in the El Gallo Formation represent the first record of Basilemys from the Upper Cretaceous of Mexico.[7] Due to the ornamentation of the shell fragments, researchers were able to identify them as Basilemys.[7] These ornamentations include shallow pits and small pyramidal elevations.[7] Basilemys remains have also been recovered from the Aguja Formation, which stretches from Texas to Mexico.[8] Here, shell fragments of Basilemys were found that are similar to the shell fragments found in the El Gallo Formation, also possessing the previously mentioned pits.[8]

Description and Paleobiology


The proportions of Basilemys' skull are very similar to those of Adocus, Baptemys and Zangerlia.[1] These genera have cheek and temporal regions that are deeply emarginated.[1] The cheek emargination is short and deep and reaches above the level of the ventral edge of the orbit.[1] The ventro-posterior corner of Basilemys' skull lacks a posterior projection into the temporal emargination which is unlike the genus Adocus.[1] Additionally, the temporal emargination of Basilemys projects forward and reaches the anterior edge of the cheek emargination.[1] The deep cheek and temporal emarginations found in Basilemys are not seen in the genus Nanhsiungchelys, which is part of the same family Basilemys belongs to; Nanhsiungchelyidae.[1] Near the center of individual bones, the skull roof of Basilemys is developed and there are striations that extend outwards from these central regions.[1] In contrast, the skull roof of Nanhsiungchelys is covered by sculpture that matches the carapace.[1] Meanwhile, in Adocus, the skull roof bones are smooth.[1]

Resembling both Adocus and Zangerlia, Basilemys has a small external narial opening.[1] Basilemys has a deep premaxillary border that is just below the external narial opening.[1] In addition, the premaxillae of Basilemys are paired instead of fused to form a single element like in the Trionychia family.[1] Similar to other trionychoids, Basilemys' orbits have large openings.[1] At the antero-ventral edge of the orbit, a groove on the external surface of the maxilla borders it.[1] The orbit is also extensively floored by the palate which is a condition that is seen in Adocus, Baptemys, and Dermatemys.[1]

The triturating surface of the maxilla is visible on both sides of the skull of Basilemys and has a well developed maxillary tooth.[1] The tooth is elongated and has a blade-like structure. It also borders a deep, circular cavity medially.[1] A deep labial ridge is also present.[1] The lower jaw of Basilemys is short and deep.[1] A sharp symphyseal hook is present on the lower jaw.[1] The coronoid process of Basilemys is low and is located near the posterior end of the jaw.[1]

Carapace and plastron

Underside of B. gaffneyi shell
Underside of B. gaffneyi shell

Turtle shells are a key feature in the identification and differentiation of turtles. The shells are made up of two portions: the carapace and the plastron. The carapace is the hard dorsal shell of a turtle while the plastron is the ventral surface of the shell. Turtle shells are important because they, to varying degrees, protect them from harm. The surface texture of a Basilemys carapace consists of many small, shallow pits that are arranged in a chain-link pattern.[3] These shallow pits are bordered by low, tetrahedal protrusions.[3] In Nanhsiungchelys, the pits are arranged more tightly and are in transverse rows over the costals, which heavily contrasts the arrangement seen in Basilemys.[3] The pattern of the carapace in Basilemys can be described as being nearly isotropic.[3]

The researchers who found the carapace of B. morrinensis from the Horseshoe Canyon Formation were able to reconstruct it through drawings and distinguish the various features of the carapace. The midline of the carapace consists of vertebral scales.[3] On B. morrinensis there are five of these scales.[3] On the lateral sides of the vertebral scales, there are costals and pleural scales that make up another portion of the carapace.[3] The borders of the carapace are made up of the peripherals and marginal scales.[3] The pygal bone sits at the posterior end of the shell, and the suprapygal is right above it.[3]

Basilemys' plastron is octagonal and elongated.[3] The various parts that make up the plastron can be seen in the reconstion made by the researchers who recovered the B. morrinensis specimen from the Horseshoe Canyon Formation. Like the carapace, the plastron is made up of many bony elements, and the plastron can be divided into five distinct parts. At the anterior end of the plastron, the first part is the epiplastron.[3] Following the epiplastron is the entoplastron.[3] The next two divisions after the entoplastron are the hyoplastron and the hypoplastron.[3] From the reconstructed images of the plastron of B. morrinensis, the hyoplastron and hypoplastron appear to be the largest sections of the plastron. At the posterior end of the plastron, the last division is the xiphiplastron.[3] The scales that make up the plastron include the humeral-, axillary-, pectoral-, abdominal-, femoral- and anal scales.[3]


In a histological analysis of the shell bones of two groups of stem-trionychians, Adocidae and Nanhsiungchelyidae, it was found that the shell bones of the Nanhsiungchelyids have a diploe structure and cortical bone layers that frame the interior cancellous bone.[9] The osteoderms of Basilemys were found to have pronounced and elaborate sculpturing patterns.[9] The sample from the North American Basilemys showcased a highly organized “spindle-shaped” ornamentation pattern.[9] In addition, the sculpturing pattern of Basilemys is made of irregular grooves and pits of external bone surface which is what is typically described as the “pock-mark” surface.[9]


Based on the specimens found from the family Nanhsiungchelyidae, researchers have been able to create cladograms based on their findings. The placement of Basilemys puts it closest to the genera Nanhsiungchelys and Zangerlia. Both within its family, Nanhsiungchelyidae. With close examination of shell histology, along with skull and neck proportions, it was determined that, of the two, Basilemys is more closely related to Zangerlia.[10] Based on similar characteristics, Adocus was recovered as relatively closely related to Basilemys. However, most cladograms place Adocus outside of Nanhsiungchelyidae.[4]








  1. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z Brinkman, Donald (1998). "The skull and neck of the Cretaceous turtle Basilemys (Trionychoidea, Nanhsiungchelyidae), and the interrelationships of the genus". Paludicola. 1 (4): 150–157.
  2. ^ a b Brinkman, Donald; Nicholls, Elizabeth L. (1993). "New Specimen of Basilemys praeclara Hay and Its Bearing on the Relationships of the Nanhsiungchelyidae (Reptilia: Testudines)". Journal of Paleontology. 67 (6): 1027–1031. doi:10.1017/S002233600002535X.
  3. ^ a b c d e f g h i j k l m n o p q r s Mallon, Jordan; Brinkman, Donald (2018). "Basilemys morrinensis, a new species of nanhsiungchelyid turtle from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada". Journal of Paleontology. 38 (2).
  4. ^ a b c Hirayama, R.; Sakurai, K.; Chitoku, G.; Kawakami, G.; Kito, N. (2001). "Anomalochelys angulata, an unusual land turtle of Family Nanhsiungchelyidae (Superfamily Trionychoidea; Order Testudines) from the Upper Cretaceous of Hokkaido, North Japan". Russian Journal of Herpetology. 8 (2): 127–138.
  5. ^ Sukhanov, V.B.; Danilov, I.G.; Syromyatnikova, E.V. (2008). "The description and phylogenetic position of a new nanhsiungchelyid turtle from the Late Cretaceous of Mongolia". Acta Palaeontol. 53 (4): 601–614. doi:10.4202/app.2008.0405. S2CID 86585072.
  6. ^ a b c d Brinkman, Donald (2003). "A review of nonmarine turtles from the Late Cretaceous of Alberta". Canadian Journal of Earth Sciences. 40 (4): 557–571. Bibcode:2003CaJES..40..557B. doi:10.1139/e02-080.
  7. ^ a b c d e f Lopez-Conde, O.A.; Sterli, J.; Chavarria-Arellano, M.L.; Brinkman, D.B.; Montellano-Ballesteros, M. (2018). "Turtles from the Late Cretaceous (Campanian) of El Gallo Formation, Baja California, Mexico". Journal of South American Earth Sciences. 88: 693–699. Bibcode:2018JSAES..88..693L. doi:10.1016/j.jsames.2018.10.005. S2CID 134734488.
  8. ^ a b Sankey, Julia (2006). "Turtles of the upper Aguja Formation (late Campanian), Big Bend National Park, Texas". New Mexico Museum of Natural History and Science Bulletin. 35: 235–243.
  9. ^ a b c d Scheyer, T.M.; Syromyatnikova, E.V.; Danilov, I.G. (2017). "Turtle shell bone and osteoderm histology of Mesozoic and Cenozoic stem-trionychian Adociadae and Nanhsiungchelyidae (Cryptodira: Adocusia) from Central Asia, Mongolia, and North America". Fossil Record. 20 (1): 69–85. doi:10.5194/fr-20-69-2017.
  10. ^ Brinkman, D.; Peng, J-H. (1996). "A new species of Zangerlia (Testudines: Nanhsiungchelyidae) from the Upper Cretaceous redbeds at Bayan Mandahu, Inner Mongolia, and the relationships of the genus". Canadian Journal of Earth Sciences. 33 (4): 526–540. Bibcode:1996CaJES..33..526B. doi:10.1139/e96-041.