Turtle shell bone and osteoderm histology of Mesozoic and Cenozoic stem-trionychian Adocidae and Nanhsiungchelyidae (Cryptodira: Adocusia) from Central Asia, Mongolia, and North America

The record of fossil turtles from the Cretaceous and Cenozoic of Asia and North America is very rich, including several lineages of cryptodiran turtles. Here we survey the shell bone histology of two important closely related groups of stem trionychians, the Adocidae and Nanhsiungchelyidae, which have representatives in both Asia and North America. All studied taxa show shell bones in which the diploe is framed by well-developed cortical compact bone layers. Taxa of both groups also express external regular surface sculpturing of their shell bones, and in the case of the nanhsiungchelyid genus <i>Basilemys</i> also on the osteoderms, which is also reflected in the internal histological bone structures. Besides similarities of the regular ornamentation patterns, both groups share a number of microanatomical and histological characters such as the zonation of external cortex with rather homogeneous fine-fibred interwoven structural fibres (ISF) in the more internal zone and a dominance of vertically oriented fibres in the ISF and the presence of growth marks in the more external zone. On the other hand, growth marks, i.e. lines of arrested growth, which are visible as wavy lines in thin sections, extend subparallel to the external bone surface in adocids, but they are not parallel/subparallel in nanhsiungchelyids. Thickness and structure of bone trabeculae in the cancellous interior regions depends on the shell bone thickness of the individual samples. The internal cortices of all taxa except the North American samples of <i>Adocus</i> usually consist of parallel-fibred bone that locally grades into lamellar bone. Secondary bone remodelling is more frequent in nanhsiungchelyids compared to adocids, and Sharpey's fibres that extend perpendicular to the bone margins extending across subparallel growth marks are more commonly found in adocids. In addition, bone histology served to identify trionychid specimens in the adocid and nanhsiungchelyid samples, especially as bone surface sculpturing patterns were weathered or eroded in those cases. The histological data present thus supplement the numerous previously reported differences in external shell morphology between adocids and nanhsiungchelyids on the one hand and trionychids and carettochelyids on the other