Jurassic shuotheriids show earliest dental diversification of mammaliaforms.

Shuotheriids are Jurassic mammaliaforms that possess pseudotribosphenic teeth in which a pseudotalonid is anterior to the trigonid in the lower molar, contrasting with the tribosphenic pattern of therian mammals (placentals, marsupials and kin) in which the talonid is posterior to the trigonid1-4. The origin of the pseudotribosphenic teeth remains unclear, obscuring our perception of shuotheriid affinities and the early evolution of mammaliaforms1,5-9. Here we report a new Jurassic shuotheriid represented by two skeletal specimens. Their complete pseudotribosphenic dentitions allow reidentification of dental structures using serial homology and the tooth occlusal relationship. Contrary to the conventional view1,2,6,10,11, our findings show that dental structures of shuotheriids can be homologized to those of docodontans and partly support homologous statements for some dental structures between docodontans and other mammaliaforms6,12. The phylogenetic analysis based on new evidence removes shuotheriids from the tribosphenic ausktribosphenids (including monotremes) and clusters them with docodontans to form a new clade, Docodontiformes, that is characterized by pseudotribosphenic features. In the phylogeny, docodontiforms and 'holotherians' (Kuehneotherium, monotremes and therians)13 evolve independently from a Morganucodon-like ancestor with triconodont molars by labio-lingual widening their posterior teeth for more efficient food processing. The pseudotribosphenic pattern passed a cusp semitriangulation stage9, whereas the tribosphenic pattern and its precursor went through a stage of cusp triangulation. The two different processes resulted in complex tooth structures and occlusal patterns that elucidate the earliest diversification of mammaliaforms.

Attenuated evolution of mammals through the Cenozoic.

The Cenozoic diversification of placental mammals is the archetypal adaptive radiation. Yet, discrepancies between molecular divergence estimates and the fossil record fuel ongoing debate around the timing, tempo, and drivers of this radiation. Analysis of a three-dimensional skull dataset for living and extinct placental mammals demonstrates that evolutionary rates peak early and attenuate quickly. This long-term decline in tempo is punctuated by bursts of innovation that decreased in amplitude over the past 66 million years. Social, precocial, aquatic, and herbivorous species evolve fastest, especially whales, elephants, sirenians, and extinct ungulates. Slow rates in rodents and bats indicate dissociation of taxonomic and morphological diversification. Frustratingly, highly similar ancestral shape estimates for placental mammal superorders suggest that their earliest representatives may continue to elude unequivocal identification.

Brawn before brains in placental mammals after the end-Cretaceous extinction.

Mammals are the most encephalized vertebrates, with the largest brains relative to body size. Placental mammals have particularly enlarged brains, with expanded neocortices for sensory integration, the origins of which are unclear. We used computed tomography scans of newly discovered Paleocene fossils to show that contrary to the convention that mammal brains have steadily enlarged over time, early placentals initially decreased their relative brain sizes because body mass increased at a faster rate. Later in the Eocene, multiple crown lineages independently acquired highly encephalized brains through marked growth in sensory regions. We argue that the placental radiation initially emphasized increases in body size as extinction survivors filled vacant niches. Brains eventually became larger as ecosystems saturated and competition intensified.

Exploring the ontogenetic development of the inner ear in Aardvarks.

The aardvark is the last living Tubulidentata, an order of afrotherian mammals. Afrotheria is supported strongly by molecular analyses, yet sparingly by morphological characters. Moreover, the biology of the aardvark remains incompletely known. The inner ear, and its ontogeny in particular, has not been studied in details yet, though it bears key ecomorphological characters and phylogenetical signal. The aim of this study is to decipher and discuss the ontogenetic development of the different areas of the inner ear of Orycteropus afer. We focused in particular on their relative size and morphological rates of development. Specimens were scanned with 3D imaging techniques. 3D and 2D geometric morphometrics coupled with qualitative descriptions of the petrosal ossification allowed us to evidence several stages through development. Based on our sample, the cochlea is the first structure of the inner ear to reach adult size, but it is the last one to acquire its adult morphology close to parturition. In contrast, after a delayed growth spurt, the semicircular canals reach their mature morphology before the cochlea, concomitantly with the increase of petrosal ossification. The ontogeny of the aardvark inner ear shows similarities with that of other species, but the apex of the cochlea presents some autapomorphies. This work constitutes a first step in the study of the ontogeny of this sensorial organ in Afrotheria.

Folding Stability of Pax9 Intronic G-Quadruplex Correlates with Relative Molar Size in Eutherians.

Eutherian dentition has been the focus of a great deal of studies in the areas of evolution, development, and genomics. The development of molar teeth is regulated by an antero-to-posterior cascade mechanism of activators and inhibitors molecules, where the relative sizes of the second (M2) and third (M3) molars are dependent of the inhibitory influence of the first molar (M1). Higher activator/inhibitor ratios will result in higher M2/M1 or M3/M1. Pax9 has been shown to play a key role in tooth development. We have previously shown that a G-quadruplex in the first intron of Pax9 can modulate the splicing efficiency. Using a sliding window approach with we analyzed the association of the folding energy (Mfe) of the Pax9 first intron with the relative molar sizes in 42 mammalian species, representing 9 orders. The Mfe of two regions located in the first intron of Pax9 were shown to be significantly associated with the M2/M1 and M3/M1 areas and mesiodistal lengths. The first region is located at the intron beginning and can fold into a stable G4 structure, whereas the second is downstream the G4 and 265 bp from intron start. Across species, the first intron of Pax9 varied in G-quadruplex structural stability. The correlations were further increased when the Mfe of the two sequences were added. Our results indicate that this region has a role in the evolution of the mammalian dental pattern by influencing the relative size of the molars.

Macroscopic and microscopic morphology of the trachea and lungs of giant anteater (Myrmecophaga tridactyla) / Morfologia macroscópica e microscópica da traqueia e pulmões do tamanduá-bandeira (Myrmecophaga tridactyla)

Giant anteater (Myrmecophaga tridactyla) is a wild mammal distributed in Central and South America; nowadays, it is classified as an endangered species. Research about the macroscopic and histomorphological aspects of its respiratory tract is scarce, and, sometimes, it limits the treatment provided to sick animals and impairs species preservation. Thus, the present study aims to describe the macroscopic and microscopic morphology of its lower respiratory tract, including trachea and lungs. To do so, 12 adult giant anteaters from "Centro de Triagem de Animais Silvestres de Goiânia" (CETAS-GO), Goiás State, Brazil, were used in the research after natural death or euthanasia. Three of these animals were used for macroscopic assessments; they were fixed in 10% buffered formalin and dissected. Trachea and lung tissue samples were collected from nine animals right after death and fixed in 10% buffered formalin for histomorphological analysis; they were processed, embedded in paraffin, and inked with hematoxylin-eosin (HE), periodic acid-Schiff (PAS), and Masson's trichrome. The macroscopic analysis showed that the trachea in this species is proportionally short and presents from 19 to 27 tracheal cartilages. The right lung presents four lobes and the left one, two. The microscopic analysis evidenced respiratory epithelium of the ciliated cylindrical pseudostratified type, without evident goblet cells in the mucosa layer of the trachea and bronchi. The pulmonary visceral pleura is thick, similar to other large domestic mammals - complete septa extend from the pulmonary visceral pleura. In conclusion, the macroscopy and histomorphology of giant anteater's lower respiratory tract, represented by trachea and lungs, are similar to that of other domestic and wild mammals. Pulmonary histomorphology is mainly similar to that of pigs and ruminants: it has thick visceral pleura that emits complete septa of conjunctive tissue, which enable lobular parenchymal architecture.(AU)

O tamanduá-bandeira (Myrmecophaga tridactyla) é um mamífero silvestre com distribuição na América Central e do Sul e, atualmente, encontra-se classificado como ameaçado de extinção. Pesquisas acerca dos aspectos macroscópicos e histomorfológicos do seu sistema respiratório são escassas, o que, por vezes, limita o tratamento e o manejo de eventuais animais doentes, bem como a preservação da espécie. Desse modo, o objetivo deste estudo foi descrever a morfologia macroscópica e microscópica do aparelho respiratório inferior do tamanduá-bandeira, incluindo traqueia e pulmões. Para tanto, foram utilizados 12 tamanduás-bandeiras adultos, provenientes do Centro de Triagem de Animais Silvestres de Goiânia (CETAS-GO), Goiás, Brasil, após morte natural ou eutanásia. Destes, três foram utilizados para o estudo macroscópico, sendo fixados em formalina tamponada a 10% e dissecados. Para a análise histomorfológica, amostras teciduais da traqueia e do pulmão foram colhidas de nove animais logo após o óbito, fixadas em formalina tamponada a 10%, processadas, incluídas em parafina e coradas com hematoxilina e eosina (HE), ácido periódico-Schiff (PAS) e tricrômico de Masson. À análise macroscópica notou-se que a traqueia é proporcionalmente curta, apresentando 19 a 27 cartilagens traqueais. O pulmão direito apresenta quatro lobos e o esquerdo dois. À análise microscópica foi constatado epitélio respiratório do tipo pseudoestratificado cilíndrico ciliado, sem células caliciformes evidentes na camada mucosa da traqueia e dos brônquios. A pleura visceral pulmonar é espessa, assim como nos grandes mamíferos domésticos, e, a partir desta, estendem-se septos completos. Conclui-se que a macroscopia e a histomorfologia do sistema respiratório inferior do tamanduá-bandeira, representado pela traqueia e pulmões, são semelhantes àquelas de outros mamíferos domésticos e silvestres. A histomorfologia pulmonar é especialmente semelhante à de suínos e ruminantes, com a pleura visceral espessa e emitindo septos completos de tecido conjuntivo, que conferem arquitetura parenquimal lobular.(AU)

Numerical Analysis of the Cerebral Cortex in Diprotodontids (Marsupialia; Australidelphia) and Comparison with Eutherian Brains.

Diprotodontids are a diverse group of Australian metatherians, which occupy a range of ecological niches from nectar and pollen-feeders to grazers and folivores. The group encompasses small-brained nectar-feeding species (Tarsipes) and large-brained grazing and browsing species (macropods). This group of Australian metatherians therefore represents an opportunity to examine how the cerebral cortex has expanded in an adaptive radiation quite independent of that occurring among eutherians. We have used the Nelson Brain Collection and online resources to perform a quantitative analysis of the isocortex, hippocampal formation and olfactory structures in diprotodontids. We found that the scaling relationship between iso- and periallocortical grey matter and brain size, and between subcortical white matter and iso- and periallocortex grey matter, are both almost identical among diprotodontids and eutherians. By contrast, the relationship between gyrification and brain size is strikingly different between diprotodontids and eutherians, with gyrification being much lower for a given brain size among the diprotodontids, although gyrification is much more varied among macropods than other diprotodontids. The scaling of iso- and periallocortical volume with dorsal striatal and dorsal thalamic volume is almost identical among the diprotodontids and eutherians, but the claustrum is smaller, and amygdala larger, for a given brain size among diprotodontids than eutherians. The hippocampal formation and central olfactory areas (anterior olfactory region and piriform cortex) both scale more steeply with brain size among diprotodontids compared to eutherians. Our findings suggest that, although white matter scaling is identical among all therians, there are significant differences between diprotodontids and eutherians in the way that cortical folding and expansion of allocortical structures occurs with brain enlargement.

Phylogenetic tree of Litopterna and Perissodactyla indicates a complex early history of hoofed mammals.

The Litopterna is an extinct clade of endemic South American ungulates that range from Paleocene up to late Pleistocene times. Because of their unique anatomy, litopterns are of uncertain phylogenetic affinities. However, some nineteenth century authors, considered litopterns as related to perissodactyl ungulates, a hypothesis recently sustained by molecular data. The aim of the present contribution is to include litopterns and other South American related taxa in a comprehensive phylogenetic analysis together with several extant and extinct basal perissodactyl ungulates. The analysis resulted in the nesting of litopterns and kin as successive stem-clades of crown Perissodactyla. Further, litopterns are not phylogenetically grouped with any North American basal ungulate, in agreement with some previous proposals. Presence of pan-perissodactyls in South America and India indicates that southern continents probably played an important role in the early evolution of hoofed mammals.

RADIOGRAPHIC ANATOMY OF THE FORELIMB IN THE NORTHERN TAMANDUA (TAMANDUA MEXICANA).

The thoracic limb anatomy of anteaters in the family Myrmecophagidae is specialized for accessing termite and ant nests and for defense purposes. In the case of the northern tamandua (Tamandua mexicana), the forelimbs are also adapted for arboreal and terrestrial locomotion. Unfortunately, this species faces many conservation threats, such as habitat loss and traffic accidents, and injured individuals are frequently taken to wildlife rehabilitation centers. However, lack of knowledge of the radiographic osteoanatomy of this species may prevent appropriate management of injuries and thereby reduce the chances of successful release and survival. In order to fill this knowledge gap, this article describes for the first time the radiographic anatomy of the thoracic limb of the northern tamandua using four standard views and one additional view. The additional orthogonal view helps visualize structures, such as the hamatus process and the sesamoid bone, that are otherwise difficult to visualize due to the natural forearm position of anteaters. Additionally, some fractures and physeal growth plates were identified in one juvenile individual. Further radiographic investigations should be conducted on anteaters to provide more tools for diagnosis, treatment, and rehabilitation of these animals.

Quantitative analysis of cerebellar morphology in monotreme, metatherian and eutherian mammals.

To superficial inspection, the mammalian cerebellum appears to be a stereotypical structure that varies little in morphology across mammals. In the present study, the volumes of components of the corpus cerebelli, foliation of the cerebellar cortex and the volumes of the pontine and deep cerebellar nuclei have been measured and compared in three species of monotreme, 90 species of marsupial and 57 species of eutherian mammal. In all three mammalian groups, the volume of the corpus cerebelli scales isometrically with brain volume, and pontine nuclear volume also scales isometrically with cerebellar volume. The ratio of hemisphere to vermal cerebellar cortex is comparable in all mammals at small cerebellar volume, but elaboration of cerebellar hemispheres is largely confined to large cerebella of eutherian mammals. At small cerebellar volumes, diprotodontid metatherians have proportionally large cerebellar hemispheres compared to non-diprotodontid metatherians, and metatherian cerebella in general have a high volume of central white matter for a given cerebellar cortex volume compared to eutherians. The degree of foliation of the cerebellum scales similarly in therian mammals, but is relatively low in the monotremes for the volume of their corpus cerebelli. Among metatherians, cerebellar foliation is stronger among diprotodontid as compared to non-diprotodontids. Although the cerebellum has a similar structure in all mammals, there are subtle differences in structure between different mammal groups with possible functional implications.

Brain anatomy of two-toed sloth (Choloepus didactylus, Linnaeus, 1758): A comparative gross anatomical study of extant xenarthrans.

The neural system plays an important role in understanding some features of animals. Anatomical complexity correlates with the increase of functional capacity. Xenarthrans include anteaters (Vermilingua), armadillos (Cingulata) and sloths (Folivora). This group is the base of eutherian mammals, and understanding the anatomy of its neural system could provide data for functional and evolutionary interpretations. The gross anatomy of the xenarthran brain is recorded. Four extant families of Pilosa and two families of Cingulata were sampled. Usual dissection procedures were used, and the brains were analysed macroscopically. The brain of two-toed sloth, three-toed sloth, six-banded armadillo, giant anteater and collared anteater are gyrencephalic. Pygmy anteater, nine-banded armadillo, great long-nosed armadillo, southern naked-tailed armadillo and giant armadillo are lissencephalic. In most species, the rhinal fissure presents two segments, rostral and caudal (except in Vermilingua and three-toed sloth). The diencephalon and brainstem present similar anatomy. The cerebellum is wide and presents four lobes (rostral, central, caudal and floccular). Its average volume is 12.16% (Folivora), 14.26% (Vermilingua) and 18.61% (Cingulata). Among these groups, there is a statistical difference between Folivora/Cingulata concerning the cerebellum average. The general pattern of the brain of the xenarthrans is similar to that of other mammals.

Virtual endocranial and inner ear endocasts of the Paleocene 'condylarth' Chriacus: new insight into the neurosensory system and evolution of early placental mammals.

The end-Cretaceous mass extinction allowed placental mammals to diversify ecologically and taxonomically as they filled ecological niches once occupied by non-avian dinosaurs and more basal mammals. Little is known, however, about how the neurosensory systems of mammals changed after the extinction, and what role these systems played in mammalian diversification. We here use high-resolution computed tomography (CT) scanning to describe the endocranial and inner ear endocasts of two species, Chriacus pelvidens and Chriacus baldwini, which belong to a cluster of 'archaic' placental mammals called 'arctocyonid condylarths' that thrived during the ca. 10 million years after the extinction (the Paleocene Epoch), but whose relationships to extant placentals are poorly understood. The endocasts provide new insight into the paleobiology of the long-mysterious 'arctocyonids', and suggest that Chriacus was an animal with an encephalization quotient (EQ) range of 0.12-0.41, which probably relied more on its sense of smell than vision, because the olfactory bulbs are proportionally large but the neocortex and petrosal lobules are less developed. Agility scores, estimated from the dimensions of the semicircular canals of the inner ear, indicate that Chriacus was slow to moderately agile, and its hearing capabilities, estimated from cochlear dimensions, suggest similarities with the extant aardvark. Chriacus shares many brain features with other Paleocene mammals, such as a small lissencephalic brain, large olfactory bulbs and small petrosal lobules, which are likely plesiomorphic for Placentalia. The inner ear of Chriacus also shares derived characteristics of the elliptical and spherical recesses with extinct species that belong to Euungulata, the extant placental group that includes artiodactyls and perissodactyls. This lends key evidence to the hypothesized close relationship between Chriacus and the extant ungulate groups, and demonstrates that neurosensory features can provide important insight into both the paleobiology and relationships of early placental mammals.

External and digestive system morphology of the Tamandua tetradactyla.

Collared anteater is present throughout the Brazilian territory, but its morphology is little known. This study aimed to trace its external biometric profile and its digestive system. Six cadavers (4 ± 0.16 kg) from Paraíba, Brazil, were used. The thoracic limbs were more robust when compared to pelvic. The length of the tail is below that described in specimens from south-eastern Brazil, probably due to the greater use of the tail suspension function in the leafier forests in this region. The tongue had no papillae and had keratinized squamous epithelium. The oesophagus was tubular with keratinized squamous epithelium. The stomach was single-chambered with simple prismatic epithelium and presence of tubular glands in the lamina propria. The pyloric region was thicker and had larger folds of mucosa; the greater curvature was approximately three times greater than the smaller. The small intestine was larger than that found in other Xernathra species such as sloth, an herbivore animal; in the large intestine, the caecum was small. The colon was subdivided ascending, transverse and descending, with simple prismatic epithelium. This study is the first to bring the description of the external and digestive system morphometrics of collared anteater adult and pup. The anatomical characteristics of the digestive system show that the digestion of the anteaters, although not using the fermentation, is not as rapid as in other Xenarthras, probably due to the material of the exoskeleton of the insects by them consumed. The data generate subsidies for the clinic, surgery, preservation and nutrition of this species.

Anatomical and radiographic study of the scapula in juveniles and adults of Tamandua mexicana (Xenarthra: Myrmecophagidae).

Tamandua mexicana is a mammal of the superorder Xenarthra geographically located between Mexico and Peru that remains insufficiently studied. These species are characterized by their movement, resting, and eating of termites and ants in arboreal and terrestrial environments, and therefore, their thoracic limbs have been adapted morphologically to move in these environments. Tamandua mexicana is a species constantly threatened by different causes, and their thoracic limbs can be affected in the scapular and shoulder regions as a result; therefore, specific anatomical knowledge of the scapula allows for better clinical procedures, surgical approaches, radiological diagnosis and muscular reconstructions in extinct species of anteaters. The main objective of this study was to report the anatomical and radiographic features of the scapula in juvenile and adult specimens of T. mexicana. Gross dissections and radiographic study were performed in six cadavers. The scapula had large differences compared with other mammals, such as two spines, a foramen for the suprascapular nerve, a long acromion, a cranial transverse scapular ligament and caudolateral fossa for the origin of the muscles teres major, tensor fasciae antebrachii and subscapular. In juvenile specimens, a scapular cartilage, an acromial cartilage, craniomedial and caudolateral coracoscapular ligaments forming the foramen for the suprascapular nerve, and progressive ossification of those ligaments and the cranial transverse scapular ligament were observed. These findings were corroborated by radiography; therefore, these characteristics change with age. The bony reliefs of the scapula of T. mexicana corroborate the specialization of the shoulder in this species.

The brain of the tree pangolin (Manis tricuspis). VI. The brainstem and cerebellum.

The brainstem (midbrain, pons, and medulla oblongata) and cerebellum (diencephalic prosomere 1 through to rhombomere 11) play central roles in the processing of sensorimotor information, autonomic activity, levels of awareness and the control of functions external to the conscious cognitive world of mammals. As such, comparative analyses of these structures, especially the understanding of specializations or reductions of structures with functions that have been elucidated in commonly studied mammalian species, can provide crucial information for our understanding of the behavior of less commonly studied species, like pangolins. In the broadest sense, the nuclear complexes and subdivisions of nuclear complexes, the topographical arrangement, the neuronal chemistry, and fiber pathways of the tree pangolin conform to that typically observed across more commonly studied mammalian species. Despite this, variations in regions associated with the locus coeruleus complex, auditory system, and motor, neuromodulatory and autonomic systems involved in feeding, were observed in the current study. While we have previously detailed the unusual locus coeruleus complex of the tree pangolin, the superior olivary nuclear complex of the auditory system, while not exhibiting additional nuclei or having an altered organization, this nuclear complex, particularly the lateral superior olivary nucleus and nucleus of the trapezoid body, shows architectonic refinement. The cephalic decussation of the pyramidal tract, an enlarged hypoglossal nucleus, an additional subdivision of the serotonergic raphe obscurus nucleus, and the expansion of the superior salivatory nucleus, all indicate neuronal specializations related to the myrmecophagous diet of the pangolins.

Quantitative analysis of forebrain pallial morphology in monotremes and comparison with that in therians.

We have made quantitative volumetric analyses of cerebral cortical (pallial) structures in the brains of three species of monotreme (Ornithorhynchus anatinus, Tachyglossus aculeatus, Zaglossus bruijni) and compared the findings with similar measurements in a range of therian mammals (6 marsupials and 50 placentals). We have found that although the iso- and periallocortical grey matter volume of the monotremes is about what would be expected for their brain size, the proportion of iso- and periallocortical white matter in monotremes is substantially lower than that in the forebrains of therians. This suggests that the forebrains of the three monotremes have fewer association, commissural and/or projection connections than those of similarly sized forebrains of therian mammals. We also found that the iso- and periallocortex of the platypus is relatively smooth-surfaced compared to similarly sized brains of therian mammals, with a distinct caudal shift in the positioning of cortical white matter in the forebrain, consistent with expansion of the posterior thalamic radiation. Central laminated olfactory structures (anterior olfactory nucleus and piriform cortex) are large in the tachyglossid monotremes (Tachyglossus aculeatus and Zaglossus bruijni) and large in xenarthran placental mammals, suggesting convergence of the forebrain structure of monotreme formivores with that of similarly specialized therians like the xenarthrans Myrmecophaga tridactyla and Dasypus novemcinctus.

Rapid morphological evolution in placental mammals post-dates the origin of the crown group.

Resolving the timing and pattern of early placental mammal evolution has been confounded by conflict among divergence date estimates from interpretation of the fossil record and from molecular-clock dating studies. Despite both fossil occurrences and molecular sequences favouring a Cretaceous origin for Placentalia, no unambiguous Cretaceous placental mammal has been discovered. Investigating the differing patterns of evolution in morphological and molecular data reveals a possible explanation for this conflict. Here, we quantified the relationship between morphological and molecular rates of evolution. We show that, independent of divergence dates, morphological rates of evolution were slow relative to molecular evolution during the initial divergence of Placentalia, but substantially increased during the origination of the extant orders. The rapid radiation of placentals into a highly morphologically disparate Cenozoic fauna is thus not associated with the origin of Placentalia, but post-dates superordinal origins. These findings predict that early members of major placental groups may not be easily distinguishable from one another or from stem eutherians on the basis of skeleto-dental morphology. This result supports a Late Cretaceous origin of crown placentals with an ordinal-level adaptive radiation in the early Paleocene, with the high relative rate permitting rapid anatomical change without requiring unreasonably fast molecular evolutionary rates. The lack of definitive Cretaceous placental mammals may be a result of morphological similarity among stem and early crown eutherians, providing an avenue for reconciling the fossil record with molecular divergence estimates for Placentalia.

Evolutionary Tinkering of the Mandibular Canal Linked to Convergent Regression of Teeth in Placental Mammals.

Loss or reduction of teeth has occurred independently in all major clades of mammals [1]. This process is associated with specialized diets, such as myrmecophagy and filter feeding [2, 3], and led to an extensive rearrangement of the mandibular anatomy. The mandibular canal enables lower jaw innervation through the passage of the inferior alveolar nerve (IAN) [4, 5]. In order to innervate teeth, the IAN projects ascending branches directly through tooth roots [5, 6], bone trabeculae [6], or bone canaliculi (i.e., dorsal canaliculi) [7]. Here, we used micro-computed tomography (µ-CT) scans of mandibles, from eight myrmecophagous species with reduced dentition and 21 non-myrmecophages, to investigate the evolutionary fate of dental innervation structures following convergent tooth regression in mammals. Our observations provide strong evidence for a link between the presence of tooth loci and the development of dorsal canaliculi. Interestingly, toothless anteaters present dorsal canaliculi and preserve intact tooth innervation, while equally toothless pangolins do not. We show that the internal mandibular morphology of anteaters has a closer resemblance to that of baleen whales [7] than to pangolins. This is despite masticatory apparatus resemblances that have made anteaters and pangolins a textbook example of convergent evolution. Our results suggest that early tooth loci innervation [8] is required for maintaining the dorsal innervation of the mandible and underlines the dorsal canaliculi sensorial role in the context of mediolateral mandibular movements. This study presents a unique example of convergent redeployment of the tooth developmental pathway to a strictly sensorial function following tooth regression in anteaters and baleen whales.

Serial Homology and Correlated Characters in Morphological Phylogenetics: Modeling the Evolution of Dental Crests in Placentals.

Accurate modeling of the complexity of morphological evolution is crucial for morphological phylogenetics and for performing tests on a wide variety of evolutionary scenarios. In this context, morphological integration and the problem of correlated categorical characters represent a major challenge. In particular, the magnitude and implications of correlations among serially homologous structures such as teeth have been much debated but were never tested statistically within a broad phylogenetic context. Here, we present a large-scale empirical study analyzing the serial variation of cingular crests on successive molars (M1, M2, and M3) of 274 placental species in a phylogenetic context. Both likelihood analyses and analysis of phylogenetic co-distributions demonstrated highly correlated evolution in the entire sample and thus the non-independence of these serial features at a macroevolutionary scale. Likelihood analyses show that their serial variation should be better scored within a single composite character model with constrained paths for transitions enabling simultaneous changes on all three molars, which suggests a strong developmental or genetic integration. These results are congruent with current genetic and developmental knowledge related to dental morphological variation and call into question the frequent use of separate characters scored on serially homologous structures of the dentition in phylogenetic analyses. Overall, they provide long-overdue and clear empirical evidence that in-depth studies of patterns of integration constitute an essential step toward more realistic character construction and modeling. This approach is critical for more accurate morphological phylogenetics and, more generally, for testing macroevolutionary scenarios on groups of correlated characters.

The brain of the tree pangolin (Manis tricuspis). IV. The hippocampal formation.

Employing a range of standard and immunohistochemical stains we provide a description of the hippocampal formation in the brain of the tree pangolin. For the most part, the architecture, chemical neuroanatomy, and topological relationships of the component parts of the hippocampal formation of the tree pangolin were consistent with that observed in other mammalian species. Within the hippocampus proper fields CA1, 3, and 4 could be identified with certainty, while CA2 was tentatively identified as a small transitional zone between the CA1 and CA3 fields. Within the dentate gyrus evidence for adult hippocampal neurogenesis at a rate comparable to other mammals was observed. The subicular complex and entorhinal cortex also exhibited divisions typically observed in other mammalian species. In contrast to many other mammals, an architecturally and neurochemically distinct CA4 field was observed, supporting Lorente de Nó's proposed CA4 field, at least in some mammalian species. In addition, up to seven laminae were evident in the dentate gyrus. Calretinin immunostaining revealed the three sublamina of the molecular layer, while immunostaining for vesicular glutamate transporter 2 and neurofilament H indicate that the granule cell layer was composed of two sublamina. The similarities and differences observed in the tree pangolin indicate that the hippocampal formation is an anatomically and neurochemically conserved neural unit in mammalian evolution, but minor changes may relate to specific life history features and habits of species.