Cranial and endocranial anatomy of a three-dimensionally preserved teleosauroid thalattosuchian skull.

Thalattosuchians represent one of the several independent transitions into the marine realm among crocodylomorphs. The extent of their aquatic adaptations ranges from the semiaquatic teleosauroids, superficially resembling extant gharials, to the almost cetacean-like pelagic metriorhynchids. Understanding the suite of osteological, physiological, and sensory changes that accompanied this major transition has received increased attention, but is somewhat hindered by a dearth of complete three-dimensionally preserved crania. Here, we describe the cranial and endocranial anatomy of a well-preserved three-dimensional specimen of Macrospondylus bollensis from the Toarcian of Yorkshire, UK. The trigeminal fossa contains two similar-sized openings separated by a thin lamina of prootic, a configuration that appears unique to a subset of teleosauroids. Macrospondylus bollensis resembles other thalattosuchians in having pyramidal semicircular canals with elongate cochlear ducts, enlarged carotid canals leading to an enlarged pituitary fossa, enlarged orbital arteries, enlarged endocranial venous sinuses, reduced pharyngotympanic sinuses, and a relatively straight brain with a hemispherical cerebral expansion. We describe for the first time the olfactory region and paranasal sinuses of a teleosauroid. A relatively large olfactory region suggests greater capacity for airborne olfaction in teleosauroids than in the more aquatically adapted metriorhynchoids. Additionally, slight swellings in the olfactory region suggest the presence of small salt glands of lower secretory capacity than those of metriorhynchoids. The presence of osteological correlates for salt glands in a teleosauroid corroborates previous hypotheses that these glands originated in the common ancestor of Thalattosuchia, facilitating their rapid radiation into the marine realm.

Inner ear sensory system changes as extinct crocodylomorphs transitioned from land to water.

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.

Evolutionary structure and timing of major habitat shifts in Crocodylomorpha.

Extant crocodylomorphs are semiaquatic ambush predators largely restricted to freshwater or estuarine environments, but the group is ancestrally terrestrial and inhabited a variety of ecosystems in the past. Despite its rich ecological history, little effort has focused on elucidating the historical pattern of ecological transitions in the group. Traditional views suggested a single shift from terrestrial to aquatic in the Early Jurassic. However, new fossil discoveries and phylogenetic analyses tend to imply a multiple-shift model. Here we estimate ancestral habitats across a comprehensive phylogeny and show at least three independent shifts from terrestrial to aquatic and numerous other habitat transitions. Neosuchians first invade freshwater habitats in the Jurassic, with up to four subsequent shifts into the marine realm. Thalattosuchians first appear in marine habitats in the Early Jurassic. Freshwater semiaquatic mahajangasuchids are derived from otherwise terrestrial notosuchians. Within nearly all marine groups, some species return to freshwater environments. Only twice have crocodylomorphs reverted from aquatic to terrestrial habitats, both within the crown group. All living non-alligatorid crocodylians have a keratinised tongue with salt-excreting glands, but the lack of osteological correlates for these adaptations complicates pinpointing their evolutionary origin or loss. Based on the pattern of transitions to the marine realm, our analysis suggests at least four independent origins of saltwater tolerance in Crocodylomorpha.

Giant taxon-character matrices: the future of morphological systematics.

Simões et al. () argued that large matrices are linked to the construction of "problematic" characters, and that those characters negatively affect tree topology. In their re-evaluation of two squamate datasets, however, Simões et al. () simply eliminated what they termed "problematic" characters, rather than recode them. This practice ignores potential sources of phylogenetic information and, if it were to be more widely followed, would inhibit the advancement of the field of systematics. Here, we defend the necessity and inevitability of large morphological (phenomic) datasets and discuss best practices for morphological data collection in contemporary phylogenetics.

The utility of captive animals in actualistic research: A geometric morphometric exploration of the tooth row of Alligator mississippiensis suggesting ecophenotypic influences and functional constraints.

Captive broad snouted crocodylians are generally thought to have wider, shorter rostra than their wild counterparts. Interpreted to reflect morphological change in response to the conditions of captivity, this qualitative pattern could affect the utility of these animals in a variety of fields of research. However, due to relative ease of access and availability of life history data, captive animals are often utilized in actualistic research. Thus, this issue should be addressed in more detail. Here we explore snout shape variation between captive and wild members of Alligator mississippiensis using two-dimensional (2D) morphometric techniques. Several landmark schemesare used to assess the utility of different aspects of morphology in distinguishing the groups. While statistical analyses consistently differentiated between the groups, the area of morphospace occupied by wild members of A. mississippiensis generally overlapped with the larger area encompassing the captive specimens. This indicates that the captive condition is not as uniform as previously thought and instead encompasses a large spectrum of morphologies, ranging from the stereotypical broad, shortened snouts to outlines that are indistinguishable from the wild morphotype. These results align well with the interpretation that this change reflects an extreme example of ecophenotypy, since ranched, farmed, or zoo organisms are held in an array of enclosures, ranging from indoor, climate controlled pens to outdoor, more natural areas. This variation in environments should be reflected in different reactions to the animals' surroundings, resulting in a broad spectrum of morphotypes. While wild specimens are still preferred, especially for fine scale analyses, these results indicate that not all captive members of A. mississippiensis exhibit the extreme morphological alterations often cited in the literature. Weighing the conditions in which the animals are held and exploring the possibility of morphological differences against the benefits of using captive specimens should be part of any actualistic study. J. Morphol. 277:866-878, 2016. © 2016 Wiley Periodicals, Inc.

What's in an Outgroup? The Impact of Outgroup Choice on the Phylogenetic Position of Thalattosuchia (Crocodylomorpha) and the Origin of Crocodyliformes.

Outgroup sampling is a central issue in phylogenetic analysis. However, good justification is rarely given for outgroup selection in published analyses. Recent advances in our understanding of archosaur phylogeny suggest that many previous studies of crocodylomorph and crocodyliform relationships have rooted trees on outgroup taxa that are only very distantly related to the ingroup (e.g., Gracilisuchus stipanicicorum), or might actually belong within the ingroup. Thalattosuchia, a group of Mesozoic marine crocodylomorphs, has a controversial phylogenetic position--they are recovered as either the sister group to Crocodyliformes, in a basal position within Crocodyliformes, or nested high in the crocodyliform tree. Thalattosuchians lack several crocodyliform apomorphies, but share several character states with derived long-snouted forms with a similar ecological habit, suggesting their derived position may be the result of convergent evolution. Several of these "shared" characters may result from ambiguously worded character state definitions--structures that are superficially similar but anatomically different in detail are identically coded. A new analysis of crocodylomorphs with increased outgroup sampling recovers Thalattosuchia as the sister group to Crocodyliformes, distantly related to long-snouted crocodyliforms. I also demonstrate that expanding the outgroup sampling of previously published matrices results in the recovery of thalattosuchians as sister to Crocodyliformes. The exclusion of thalattosuchians from Crocodyliformes has numerous implications for large-scale evolutionary trends within the group, including extensive convergence in the evolution of the secondary palate characteristic of the group. These results demonstrate the importance of careful outgroup sampling and character construction, and their profound effect on the position of labile clades.

Efficacy or convenience? Model-based approaches to phylogeny estimation using morphological data.

Model-based approaches (e.g. maximum likelihood, Bayesian inference) are widely used with molecular data, where they might be more appropriate than maximum parsimony for estimating phylogenies under various models of molecular evolution. Recently, there has been an increase in the application of model-based approaches with morphological (mainly fossil) data; however, there is some doubt as to the effectiveness of the model of morphological evolution. The input parameters (prior probabilities) for the model are unclear, particularly when concerned with unobserved character states. Despite this, some systematists are suggesting superiority of these model-based methods over maximum parsimony based on, for example, increased resolution or, in the current study, the preferred phylogenetic placement of an iconic taxon. Here, we revisit a recently published analysis implying such superiority and document the discrepancies between parsimony-based and model-based approaches to phylogeny estimation. We find that although some taxa are shifted back to their "traditional" phylogenetic placement, other clades are disturbed. The model-based phylogenies are better resolved; however, due to the lack of an appropriate model of morphological evolution, the increase in resolving power is probably not meaningful. Similarly, some of the preferred phylogenetic positions of taxa, particularly of labile taxa such as Archaeopteryx, are based solely on analyses employing maximum parsimony as the optimality criterion. Poor resolution and labile taxa indicate a need for further examination of the morphology and not a change in method.