Unexpected morphological variability in the eggshells of the South American caimans Caiman latirostris and Caiman yacare.

Eggshell morphology is a valuable indicator of the local conditions within the nests of modern crocodilians and birds. In contrast to these latter, the anatomical structure of the eggshells of most crocodilian species is practically unknown. Here, we provide the first characterization of crocodilian eggshells, using x-ray micro-CT scans. We studied eggshells of Caiman latirostris and Caiman yacare from various developmental stages that coincide with the beginning of embryonic ossification. The new 3D renderings revealed complex ornamentation, unique among crocodilians, and amphora-shaped pore canals, some of which converge in single pore openings. We also documented a high density of pore canals with a gas diffusion capacity 45 times higher than the average predicted for modern avian eggshells. The external ornamentation and the thickness of the compact layer of the eggshells (i.e. excluding ornamentation) showed ontogenetic and interspecific differences that could be related to nesting materials and nesting areas selected by each species. The shell features described here evidence a greater structural complexity than previously recognized in phylogenetically close, sympatric crocodilian species. Further comprehensive morphological analyses on other modern and fossil crocodilian eggshells using micro-CT technology will shed new light on the evolution of reproductive strategies in this intriguing archosaur clade.

Development of the chondrocranium of two caiman species, Caiman latirostris and Caiman yacare.

Little is known about the embryonic development and variation of the chondrocranium in Crocodylia and there are no works on any Caiman species. Due to the importance of cranial features in the systematics of this clade, investigating the development of the skull in embryonic stages is essential. In this study, we present for the first time the development of the cartilaginous skull of two extant Caiman species. Anatomical descriptions of the embryonic chondrocranium of Caiman latirostris and Caiman yacare were made, paying special attention to their inter- and intraspecific variation. For this purpose, pre-hatching ontogenetic cranial series of these two caiman species were prepared with a double staining and diaphanization technique. The main differences with other crocodylian species were observed in the palatoquadrate, and interspecific variation within the genus was recorded in the hyobranchial apparatus and larynx. Some characters may be distinctive of Caiman (posterior and ventral surface of the otic process of the palatoquadrate articulated with the dorsal process of the columella auris, and otic process articulated with the lateral wall of the auditory capsule), Alligatoridae (presence of an epiphanial foramen) or C. latirostris and C. yacare (Corpus hyoidei with different number and position of foramina and different shapes of its anterior contour and anterior and posterior notch, different degrees of broadening of the distal end of the Cornu branchiale I, and presence/absence of a notch in the posteroventral surface of the cricoid). Homologies of the elements belonging to the hyobranchial apparatus could not be confirmed. As in other tetrapods the trachea consists of incomplete cartilaginous rings. Morphological changes and dissimilarities found in this study are useful as a context to start studying phylogenetic constraints. Moreover, in a heterochronic context, variations may be involved.

Skull ontogeny of extant caimans: a three-dimensional geometric morphometric approach.

Ontogenetic variation of cranial characters used in crocodylian phylogenetic systematics has never been studied. Furthermore, the relationship between diet and skull morphological transformation during ontogeny has not been properly explored yet. We quantify the inter- and intraspecific skull morphological variation in extant caiman species focusing on those areas relevant to systematics and, also investigate the relation between diet and morphological changes during ontogeny. We applied a three-dimensional approach of geometric morphometrics on post-hatching ontogenetic cranial series of Caiman latirostris and C. yacare. In order to incorporate incomplete material, we additionally tested four different methods of missing landmark estimation and apply the thin-plate spline interpolation. We detected morphological changes between species and during ontogeny (snout and pterygoid flanges increase their proportions and, orbits, temporal fenestrae, skull roof and foramen magnum decrease their relative size) that constitutes part of a general morphological change in the cranial ontogeny of crocodylians. Moreover, the negative allometry of the fenestrae and neurocranium and the positive allometry of the splanchnocranium in both caiman species are the plesiomorphic condition, at least, for tetrapods. Shape changes during growth were found to be related to ontogenetic changes in the diet. Dissimilarities between species seem to be related to different mechanical requirements and different use of the habitat. We found inter- and intraspecific variation in some morphological characters with systematic implications (the contact of nasals with naris, the contact of prefrontals in the midline, and the bones that border the suborbital fenestra and the proportion in which one of them participates) that are not currently considered in phylogenetic analyses.

A new Palaeocene crocodylian from southern Argentina sheds light on the early history of caimanines.

Caimanines are crocodylians currently restricted to South and Central America and the oldest members are from lower Palaeocene localities of the Salamanca Formation (Chubut Province, Argentina). We report here a new caimanine from this same unit represented by a skull roof and partial braincase. Its phylogenetic relationships were explored in a cladistic analysis using standard characters and a morphogeometric two-dimensional configuration of the skull roof. The phylogenetic results were used for an event-based supermodel quantitative palaeobiogeographic analysis. The new species is recovered as the most basal member of the South American caimanines, and the Cretaceous North American lineage 'Brachychampsa and related forms' as the most basal Caimaninae. The biogeographic results estimated north-central North America as the ancestral area of Caimaninae, showing that the Cretaceous and Palaeocene species of the group were more widespread than thought and became regionally extinct in North America around the Cretaceous-Palaeocene boundary. A dispersal event from north-central North America during the middle Late Cretaceous explains the arrival of the group to South America. The Palaeogene assemblage of Patagonian crocodylians is composed of three lineages of caimanines as a consequence of independent dispersal events that occurred between North and South America and within South America around the Cretaceous-Palaeogene boundary.