Crown Mammalia Research Articles

Origins of mammalian vertebral function revealed through digital bending experiments.

Unravelling the functional steps that underlie major transitions in the fossil record is a significant challenge for biologists owing to the difficulties of interpreting functional capabilities of extinct organisms. New computational modelling approaches provide exciting avenues for testing function in the fossil record. Here, we conduct digital bending experiments to reconstruct vertebral function in non-mammalian synapsids, the extinct forerunners of mammals, to provide insights into the functional underpinnings of the synapsid-mammal transition. We estimate range of motion and stiffness of intervertebral joints in eight non-mammalian synapsid species alongside a comparative sample of extant tetrapods, including salamanders, reptiles and mammals. We show that several key aspects of mammalian vertebral function evolved outside crown Mammalia. Compared to early diverging non-mammalian synapsids, cynodonts stabilized the posterior trunk against lateroflexion, while evolving axial rotation in the anterior trunk. This was later accompanied by posterior sagittal bending in crown mammals, and perhaps even therians specifically. Our data also support the prior hypothesis that functional diversification of the mammalian trunk occurred via co-option of existing morphological regions in response to changing selective demands. Thus, multiple functional and evolutionary steps underlie the origin of remarkable complexity in the mammalian backbone.

Temporal framework for the Yanliao Biota and timing of the origin of crown mammals

Establishing the temporal sequence of the Middle-Late Jurassic Yanliao Biota is essential as it anchors the timing of many key evolutionary innovations in vertebrates. Lack of sufficiently reliable high-precision ages of fossil-bearing horizons hinders our ability to reconstruct the tempo and mode of vertebrate evolution. Here, we frame a temporal sequence of Yanliao Biota with precise age constraints for iconic vertebrates, proposing that the major vertebrate-bearing strata span from 164 Ma to 157 Ma in age. The increasing ecological diversity of mammaliaforms is well illustrated by the Middle-Late Jurassic Yanliao Biota and Early Cretaceous Jehol Biota. Incorporation of the updated temporal frameworks in Bayesian tip-dated mammaliaform phylogeny reveals that Triassic haramiyidans are separate from Jurassic taxa and unrelated to crown Mammalia. Tip-dated phylogeny supports a long-fuse model for mammal evolution, featured by a Late Triassic root and Middle-Late Jurassic interordinal diversification of crown Mammalia, showing consistency with molecular-based timetrees in divergence timing.

Non-mammaliaform cynodonts from western Gondwana and the significance of Argentinean forms in enhancing understanding of the group

Terrestrial Triassic assemblages are an important component of the heritage of Argentina, particularly because of their diverse record of fossil vertebrates. Triassic basins from western Argentina have a good representation of therapsids, a lineage that includes a series of stem extinct groups plus crown Mammalia. Because of their diversity and abundance, non-mammaliaform cynodonts are the most conspicuous therapsids in the Argentinean Triassic. Cynodonts have been intensively researched in Argentina since the 1960s and continue to provide key advances in the understanding of the evolution and biology of the lineage. Here we offer a summary of Triassic Argentinean non-mammaliaform cynodonts, presenting a historical context of research on this group, and highlighting seminal contributions of Triassic Argentinean cynodonts to the global knowledge of the group. To provide a Gondwanan context we review the cynodont record in the two other places where the lineage is best represented: Brazil and southern Africa (including Madagascar), highlighting commonalities in the Argentinean record with these places. Considering the important role of non-mammaliaform cynodonts in the interpretation of the origin of mammals, we provide a historical account on the contribution of Argentinean, and Gondwanan, cynodonts to the understanding of this topic. This is supplemented with a summary of the progression of phylogenetic ideas and hypotheses for all non-mammaliaform cynodonts and for the most prolific subgroups represented in western Gondwana.

Tip dating supports novel resolutions of controversial relationships among early mammals.

The estimation of the timing of major divergences in early mammal evolution is challenging owing to conflicting interpretations of key fossil taxa. One contentious group is Haramiyida, the earliest members of which are from the Late Triassic. Many phylogenetic analyses have placed haramiyidans in a clade with multituberculates within crown Mammalia, thus extending the minimum divergence date for the crown group deep into the Triassic. A second taxon of interest is the eutherian Juramaia from the Middle–Late Jurassic Yanliao Biota, which is morphologically very similar to eutherians from the Early Cretaceous Jehol Biota and implies a very early origin for therian mammals. Here, we apply Bayesian tip-dated phylogenetic methods to investigate these issues. Tip dating firmly rejects a monophyletic Allotheria (multituberculates and haramiyidans), which are split into three separate clades, a result not found in any previous analysis. Most notably, the Late Triassic Haramiyavia and Thomasia are separate from the Middle Jurassic euharamiyidans. We also test whether the Middle–Late Jurassic age of Juramaia is ‘expected’ given its known morphology by assigning an age prior without hard bounds. Strikingly, this analysis supports an Early Cretaceous age for Juramaia, but similar analyses on 12 other mammaliaforms from the Yanliao Biota return the correct, Jurassic age. Our results show that analyses incorporating stratigraphic data can produce results very different from other methods. Early mammal evolution may have involved multiple instances of convergent morphological evolution (e.g. in the dentition), and tip dating may be a method uniquely suitable to recognizing this owing to the incorporation of stratigraphic data. Our results also confirm that Juramaia is anomalous in exhibiting a much more derived morphology than expected given its age, which in turn implies very high rates of evolution at the base of therian mammals.

The many faces of synapsid cranial allometry

AbstractPrevious studies of cranial shape have established a consistent interspecific allometric pattern relating the relative lengths of the face and braincase regions of the skull within multiple families of mammals. In this interspecific allometry, the facial region of the skull is proportionally longer than the braincase in larger species. The regularity and broad taxonomic occurrence of this allometric pattern suggests that it may have an origin near the base of crown Mammalia, or even deeper in the synapsid or amniote forerunners of mammals. To investigate the possible origins of this allometric pattern, we used geometric morphometric techniques to analyze cranial shape in 194 species of nonmammalian synapsids, which constitute a set of successive outgroups to Mammalia. We recovered a much greater diversity of allometric patterns within nonmammalian synapsids than has been observed in mammals, including several instances similar to the mammalian pattern. However, we found no evidence of the mammalian pattern within Therocephalia and nonmammalian Cynodontia, the synapsids most closely related to mammals. This suggests that the mammalian allometric pattern arose somewhere within Mammaliaformes, rather than within nonmammalian synapsids. Further investigation using an ontogenetic series of the anomodontDiictodon felicepsshows that the pattern of interspecific allometry within anomodonts parallels the ontogenetic trajectory ofDiictodon.This indicates that in at least some synapsids, allometric patterns associated with ontogeny may provide a “path of least resistance” for interspecific variation, a mechanism that we suggest produces the interspecific allometric pattern observed in mammals.

First record of a basal mammaliamorph from the early Late Triassic Ischigualasto Formation of Argentina.

We describe a new probainognathian cynodont, Pseudotherium argentinus, from the early Late Triassic Ischigualasto Formation of Argentina. Pseudotherium adds to a growing assemblage of small Triassic cynodonts that offers new insight into events leading up to the origin of crown Mammalia and the successively more inclusive Mammaliaformes and Mammaliamorpha. Using high-resolution X-ray computed tomography, we illustrate and describe the holotype and only known specimen, which consists of a well-preserved isolated skull. It preserves apomorphic features of the orbit and braincase. Prefrontal and vestigial postorbital bones are present, despite the absence of an ossified postorbital bar. As in Brasilitherium riograndensis, thin turbinal-like bones are present in the nasopharyngeal passage, and we discuss impediments to establishing their identity and function. Compared to more basal cynodonts, the cochlea is elongated but uncoiled and in this and other features it resembles basal mammaliamorphs. Our analysis found weak support for Pseudotherium as the sister taxon of Tritylodontidae. However, a broader assessment of its relationships in light of additional character data from the literature and unpublished computed tomography data suggest that it may be more realistic to view the relationships of Pseudotherium as an unresolved polytomy with tritylodontids, and the taxa referred to as tritheledontids and brasilodontids (groups of variable membership and questionable monophyly). Thus, Pseudotherium may lie just inside or just outside of Mammaliamorpha, and there is also weak character support for its sister taxon relationship with Brasilitherium. Our results amplify previous conclusions that phylogenetic relationships in this adaptive radiation of small cynodonts will remain somewhat uncertain until more complete specimens are recovered, and until high-resolution CT scans of existing specimens become available to the larger community. Toward that goal, we make the CT dataset for the holotype of Pseudotherium argentinus publically available under a Creative Commons license at www.DigiMorph.org.

Invasion of Ancestral Mammals into Dim-light Environments Inferred from Adaptive Evolution of the Phototransduction Genes

Nocturnality is a key evolutionary innovation of mammals that enables mammals to occupy relatively empty nocturnal niches. Invasion of ancestral mammals into nocturnality has long been inferred from the phylogenetic relationships of crown Mammalia, which is primarily nocturnal, and crown Reptilia, which is primarily diurnal, although molecular evidence for this is lacking. Here we used phylogenetic analyses of the vision genes involved in the phototransduction pathway to predict the diel activity patterns of ancestral mammals and reptiles. Our results demonstrated that the common ancestor of the extant Mammalia was dominated by positive selection for dim-light vision, supporting the predominate nocturnality of the ancestral mammals. Further analyses showed that the nocturnality of the ancestral mammals was probably derived from the predominate diurnality of the ancestral amniotes, which featured strong positive selection for bright-light vision. Like the ancestral amniotes, the common ancestor of the extant reptiles and various taxa in Squamata, one of the main competitors of the temporal niches of the ancestral mammals, were found to be predominate diurnality as well. Despite this relatively apparent temporal niche partitioning between ancestral mammals and the relevant reptiles, our results suggested partial overlap of their temporal niches during crepuscular periods.

A Cretaceous eutriconodont and integument evolution in early mammals.

The Mesozoic era (252-66 million years ago), known as the domain of dinosaurs, witnessed a remarkable ecomorphological diversity of early mammals. The key mammalian characteristics originated during this period and were prerequisite for their evolutionary success after extinction of the non-avian dinosaurs 66 million years ago. Many ecomorphotypes familiar to modern mammal fauna evolved independently early in mammalian evolutionary history. Here we report a 125-million-year-old eutriconodontan mammal from Spain with extraordinary preservation of skin and pelage that extends the record of key mammalian integumentary features into the Mesozoic era. The new mammalian specimen exhibits such typical mammalian features as pelage, mane, pinna, and a variety of skin structures: keratinous dermal scutes, protospines composed of hair-like tubules, and compound follicles with primary and secondary hairs. The skin structures of this new Mesozoic mammal encompass the same combination of integumentary features as those evolved independently in other crown Mammalia, with similarly broad structural variations as in extant mammals. Soft tissues in the thorax and abdomen (alveolar lungs and liver) suggest the presence of a muscular diaphragm. The eutriconodont has molariform tooth replacement, ossified Meckel's cartilage of the middle ear, and specialized xenarthrous articulations of posterior dorsal vertebrae, convergent with extant xenarthran mammals, which strengthened the vertebral column for locomotion.

A new arboreal haramiyid shows the diversity of crown mammals in the Jurassic period

A major unsolved problem in mammalian evolution is the origin of Allotheria, including Multituberculata and Haramiyida. Multituberculates are the most diverse and best known Mesozoic era mammals and ecologically resemble rodents, but haramiyids are known mainly from isolated teeth, hampering our search for their phylogenetic relationships. Here we report a new haramiyid from the Jurassic period of China, which is, to our knowledge the largest reported so far. It has a novel dentition, a mandible resembling advanced multituberculates and postcranial features adapted for arboreal life. Our phylogenetic analysis places Haramiyida within crown Mammalia, suggesting the origin of crown Mammalia in the Late Triassic period and diversification in the Jurassic, which contrasts other estimated divergence times of crown Mammalia. The new haramiyid reveals additional mammalian features of the group, helps to identify other haramiyids represented by isolated teeth, and shows again that, regardless of various phylogenetic scenarios, a complex pattern of evolution involving many convergences and/or reversals existed in Mesozoic mammals.

Mammalian hairs in Early Cretaceous amber

Two mammalian hairs have been found in association with an empty puparium in a approximately 100-million-year-old amber (Early Cretaceous) from France. Although hair is known to be an ancestral, ubiquitous feature in the crown Mammalia, the structure of Mesozoic hair has never been described. In contrast to fur and hair of some Jurassic and Cretaceous mammals preserved as carbonized filaments, the exceptional preservation of the fossils described here allows for the study of the cuticular structure. Results show the oldest direct evidence of hair with a modern scale pattern. This discovery implies that the morphology of hair cuticula may have remained unchanged throughout most of mammalian evolution. The association of these hairs with a possible fly puparium provides paleoecological information and indicates peculiar taphonomic conditions.

Natural History of Human’s Most Distant Living Mammalian Relatives

In comparison to the more than 1,200 genera of placentals and marsupials, the three extant monotreme genera make up only a tiny part of modern mammalian diversity. However, the significance of monotremes to mammalian evolutionary history and biology is far more than can be measured by their few genera. The evolutionary stature of monotremes as the first group to diverge among extant mammal lineages has been corroborated by an overwhelming body of scientific evidence from fossils, comparative morphology, and molecular information on the genomic scale. With their highly specialized physiological and functional adaptations, distinctive developmental pattern and karyotypes, these egg-laying mammals push the envelope of human imagination regarding what is possible for mammalian biological diversity. For evolutionists, monotremes represent the extant phylogenetic bracket for the common ancestor of all modern mammals (crown Mammalia). For physiologists and functional morphologists, monotremes give meaning to mammalian structural and functional diversities. For ecologists and conservation biologists, the monotremes are an integral element of an ecosystem on Gondwanaland from the deep times of the Earth. Since the echidna and platypus became known to western science in 1792 and 1800, respectively, monotremes have always fascinated the scientific community and the public (Moyal 2001). But monotremes are more than a curiosity—they are an indispensable group for all scientists in pursuit of any area of mammalian comparative biology to consider. In Echidna—An Extraordinary Egg-Laying Mammal, Michael Augee, Brett Gooden, and Anne Musser have written an excellent primer on the biology and natural history of the shortbeaked echidna—Tachyglossus aculeatus or the spiny anteater. This book of 136 pages is a concise, informative, and immensely readable review of the structure, function, physiology, behavior, and life history of the echidna. It is well aimed at a general scientific readership and contains very interesting information on all aspects of the natural history and biology J Mammal Evol (2007) 14:283–285 DOI 10.1007/s10914-007-9059-1