Thesis

Région auditive des Artiodactyles : signal phylogénétique et écologique

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Abstract

La mise en évidence par la biologie moléculaire et par les données paléontologiques de l'appartenance des cétacés au groupe des artiodactyles constitue une des avancées majeures de ces 30 dernières années en mammalogie. Il n'y a cependant pas à l'heure actuelle de consensus quant aux relations phylogénétiques basales des artiodactyles fondées sur des caractères morphologiques et l'histoire évolutive du groupe est de fait, ponctuée de nombreux points d'interrogation. Cette thèse explore une source de caractères phylogénétiques prometteuse : la région auditive (os pétreux, bulle auditive, osselets de l'oreille moyenne, oreille interne) à partir notamment des nouvelles perspectives offertes par l'imagerie µCT Scan. Les principaux objectifs de cette thèse sont (1) de déterminer le signal phylogénétique porté par la région auditive chez les artiodactyles afin d’apporter une nouvelle source de caractères aux analyses et (2) d’explorer le signal écologique porté par les différents éléments de cette région sensorielle dédiée à l’audition (oreille externe, moyenne et canal cochléaire du labyrinthe osseux) et à l’équilibrioception (vestibule et canaux semi-circulaires du labyrinthe osseux). La première partie de cette thèse (I) nous emmène au Togo, où de nombreux restes inédits de la région auditive de « baleines à pattes » (Protocetidae Stromer, 1908) ont été récoltés. D’un point de vu anatomique, ces restes fossiles ont permis de documenter et de décrire pour la première fois le stapes, l’incus et le labyrinthe osseux d’un protocète ; des éléments indispensables pour comprendre leur audition. L’analyse morpho-fonctionnelle indique qu’une audition optimale était probablement possible dans l’air et dans l’eau pour ces cétacés semi-aquatiques. De plus, la morphologie de leur cochlée indique que leur capacité auditive était proche de celle de leurs cousins terrestres et que les spécialisations relatives aux capacités auditives remarquables des cétacés modernes (i.e. sensibilité aux infra- ou ultrasons) se sont opérées après la séparation historique entre les mysticètes et les odontocètes.La deuxième partie de ce travail (II) se concentre sur les origines de l’amphibiose au sein des Cetancodonta, à travers l’étude de plusieurs familles fossiles, connues pour leurs liens étroits au milieu aquatique. L’étude de la région auditive des hippopotamoïdes (Anthracotheriidae + Hippopotamidae), révèle que l’adaptation à un mode de vie semi-aquatique est apparue plusieurs fois, de façon convergente, dans son histoire évolutive et semble d’ailleurs indiquer une origine terrestre pour ce groupe. Quant au raoellidé Indohyus, son complexe pétro-tympanique présente une combinaison de caractères suggérant un certain degré d’adaptation au milieu aquatique, mais l’étude fonctionnelle de sa cochlée indique que ce taxon ne pouvait très probablement pas entendre de façon efficace sous l’eau. Pour finir, le dernier point de cette thèse explore également le potentiel phylogénétique de la région auditive à travers une analyse construite sur des caractères morphologiques du pétreux et du labyrinthe osseux à l’échelle des artiodactyles. Pour la première fois, les résultats de notre analyse concordent avec ceux des analyses moléculaires. Parmi les points les plus notables, le clade des Cetancodonta est bien soutenu par la morphologie du pétreux et la position d’Indohyus suggère fortement que les raoellidés sont des cétacés.Ainsi, la région auditive s’avère être un élément essentiel d’un point de vu phylogénétique et morphofonctionnel. En effet, comme nous avons pu le voir tout au long de cette thèse, lorsque la nature complexe et variée de la région auditive est appréhendée dans son ensemble, elle permet d’inférer l’écologie d’un taxon donné et d’en apprendre davantage sur ses relations de parenté. Par conséquent, la région auditive est encore loin d’avoir dit ses derniers mots... et nous n’avons pas encore fini d’en entendre parler.

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Cetaceans originated in south Asia more than 50 million years ago (mya), from a small quadrupedal artiodactyl ancestor [1-3]. Amphibious whales gradually dispersed westward along North Africa and arrived in North America before 41.2 mya [4]. However, fossil evidence on when, through which pathway, and under which locomotion abilities these early whales reached the New World is fragmentary and contentious [5-7]. Peregocetus pacificus gen. et sp. nov. is a new protocetid cetacean discovered in middle Eocene (42.6 mya) marine deposits of coastal Peru, which constitutes the first indisputable quadrupedal whale record from the Pacific Ocean and the Southern Hemisphere. Preserving the mandibles and most of the postcranial skeleton, this unique four-limbed whale bore caudal vertebrae with bifurcated and anteroposteriorly expanded transverse processes, like those of beavers and otters, suggesting a significant contribution of the tail during swimming. The fore- and hind-limb proportions roughly similar to geologically older quadrupedal whales from India and Pakistan, the pelvis being firmly attached to the sacrum, an insertion fossa for the round ligament on the femur, and the retention of small hooves with a flat anteroventral tip at fingers and toes indicate that Peregocetus was still capable of standing and even walking on land. This new record from the southeastern Pacific demonstrates that early quadrupedal whales crossed the South Atlantic and nearly attained a circum-equatorial distribution with a combination of terrestrial and aquatic locomotion abilities less than 10 million years after their origin and probably before a northward dispersal toward higher North American latitudes. VIDEO ABSTRACT.
Article
Anthracotheres dispersed from Asia toward Africa at least three times: at the Eocene/Oligocene transition, during the early Miocene and later during the Miocene. Those dispersals are important datum events for African Tertiary biochronology. New fossil remains of early Libycosaurus, the genus implicated in the late Miocene dispersal, are described from a new Tunisian locality of the Kasserine area. The new fossils enhance the hypodigm of Libycosaurus algeriensis and increase the resolution of the phylogenetic position of this species using cladistics analysis. The inclusion of the genus Libycosaurus within the well-described Merycopotamus lineage allows us to constrain its dispersal time. Dispersal of this anthracothere from the Indian sub-continent to Africa was probably facilitated by sea level decrease during the early Tortonian, just preceding the Hipparion dispersal event. This new age estimation refines the resolution of the succession of late Miocene deposits in Maghreb and frames the date of the onset of the Sahara.
Article
We describe Tupelocetus palmeri, a new genus and species of archaeocete whale, based on a single specimen from the Cross Member of the middle Eocene Tupelo Bay Formation (Bartonian). The holotype consists of a partial cranium with complete petrosals, posterior processes of tympanics, the posterior ends of the nasals, a part of the right orbit, and what are interpreted as the right and left P2s. The incomplete skull was collected from the Martin Marietta Cross Quarry (Berkeley County, South Carolina), which has also furnished the holotype of Carolinacetus gingerichi. The new species differs from all other protocetids in having a deep cavity on the occiput, as well as having the following combination of features: large nasal processes of frontals, single-cusped P2, and premaxilla terminates at level of P2 or P3. A phylogenetic analysis places T. palmeri in a large polytomy along the cetacean stem, more closely related to crown Cetacea than Maiacetus, Artiocetus, and Rodhocetus, but more basal than Georgiacetus, Babiacetus, and Eocetus. Another member of this polytomy is Natchitochia jonesi, and although specimens of Natchitochia and T. palmeri do not share any elements, what is preserved suggests that they are similar in size and among the largest of all protocetids. Although we are unable to differentiate these two species, we outline future data that could resolve this question.
Article
Studying ontogeny in both extant and extinct species can unravel the mechanisms underlying mammal diversification and specialization. Among mammalian clades, Cetartiodactyla encompass species with a wide range of adaptations, and ontogenetic evidence could clarify longstanding debates on the origins of modern specialized families. Here, we study the evolution of dental eruption patterns in early diverging cetartiodactyls to assess the ecological and biological significance of this character and shed new light on phylogenetic issues. After investigation of the ontogenetic dental series of 63 extinct genera, our parsimony reconstructions of eruption state evolution suggest that the eruption of molars before permanent premolars represents a plesiomorphic condition within Cetartiodactyla. This result substantially differs from a previous study based on modern species only. As a result, the presence of this pattern in most ruminants might represent an ancestral condition contributing to their specialized herbivory, rather than an original adaptation. In contrast, the late eruption of molars in hippopotamoids is more likely related to biological aspects, such as increases in body mass and slower pace of life. Our study mainly shows that eruption sequences reliably characterize higher level cetartiodactyl taxa and could represent a new source of phylogenetic characters, especially to disentangle the origin of hippopotamoids and cetaceans.
Article
The auditory ossicles - malleus, incus and stapes - are the smallest bones in mammalian bodies and enable stable sound transmission to the inner ear. Sperm whales are one of the deepest diving aquatic mammals that produce and perceive sounds with extreme loudness greater than 180 dB and frequencies higher than 30 kHz. Therefore, it is of major interest to decipher the microstructural basis for these unparalleled hearing abilities. Using a suite of high-resolution imaging techniques, we reveal that auditory ossicles of sperm whales are highly functional, featuring an ultra-high matrix mineralization that is higher than their teeth. On a micro-morphological and cellular level, this was associated with osteonal structures and osteocyte lacunar occlusions through calcified nanospherites (i.e. micropetrosis), while the bones were characterized by a higher hardness compared to a vertebral bone of the same animals as well as to human auditory ossicles. We propose that the ultra-high mineralization facilitates the unique hearing ability of sperm whales. High matrix mineralization represents an evolutionary conserved or convergent adaptation to middle ear sound transmission. © 2018 The Author(s) Published by the Royal Society. All rights reserved.
Article
Cetartiodactyla comprises one of the most diverse mammal radiations. Currently, 23 families, 131 genera and more than 330 species are recognized. Several studies have been trying to resolve its phylogenetic relationships. The most comprehensive dated phylogenetic hypothesis available includes only 55% of the extant species, precluding a clear understanding of ecological and evolutionary patterns in Cetartiodactyla. Here, we gathered all mitochondrial genetic data available in GenBank to build a robust Cetartiodactyla calibrated phylogenetic tree using 21 fossil calibration points. We found mitogenomic data for 225 species and included other 93 species from which there was at least one mitochondrial gene available. Using a Bayesian approach, we generated a dated tree comprising 90% of the extant Cetartiodactyla species (n = 318). The major lineages showed robust support and families divergence times are congruent with the available fossil evidence and with previously published phylogenetic hypotheses. By making available a dated phylogeny with extensively sampled clades, we expect to foster future studies on the origin, tempo and mode of Cetartiodactyla diversification.
Article
Much evidence for non-human culture comes from vocally learned displays, such as the vocal dialects and song displays of birds and cetaceans. While many oscine birds use song complexity to assess male fitness, the role of complexity in humpback whale (Megaptera novaeangliae) song is uncertain owing to population-wide conformity to one song pattern. Although songs change gradually each year, the eastern Australian population also completely replaces their song every few years in cultural ‘revolutions’. Revolutions involve learning large amounts of novel material introduced from the Western Australian population. We examined two measures of song structure, complexity and entropy, in the eastern Australian population over 13 consecutive years. These measures aimed to identify the role of complexity and information content in the vocal learning processes of humpback whales. Complexity was quantified at two hierarchical levels: the entire sequence of individual sound ‘units’ and the stereotyped arrangements of units which comprise a ‘theme’. Complexity increased as songs evolved over time but decreased when revolutions occurred. No correlation between complexity and entropy estimates suggests that changes to complexity may represent embellishment to the song which could allow males to stand out amidst population-wide conformity. The consistent reduction in complexity during song revolutions suggests a potential limit to the social learning capacity of novel material in humpback whales.
Article
The external auditory meatus (EAM) in many species of mysticete whales is filled with a waxy ear plug. Though this lamellated structure is often used to age a whale, its formation and development remain undescribed. It is thought that growth layer groups (GLGs) are laid down annually, thereby increasing the size of this structure. Since some mysticete whales are migratory and many undergo molting, we hypothesized that the cyclical production of these GLGs may be related to these processes. The epithelia of both EAM and glove finger (a part of the tympanic membrane protruding into the EAM) of one juvenile and multiple adult bowhead whales from both fall (October: non‐molting) and spring (May: molting) seasons were dissected and examined anatomically and histologically. These tissue samples were compared with the adult oral epithelia at the same time periods. These epithelia shared a similar basic broad structure, though there were differences in thickness and presence of intraepithelial structures. All epithelia in the October specimens were rich in both glycogen and lipid. The parakeratinized epithelium of the oral cavity in the juvenile and some May specimens shed via the production of several superficial epithelial fissures. Other adult May specimens exhibited deep epithelial fissures, reminiscent of pressure ulcers, which would cause the detachment of the entire epithelium from the dermis. We propose that sloughed epithelial lining is the source of the GLGs in the ear plug. Correlating a potential molting sequence with these observations explained the presence of epidermal glycogen, deep epidermal fissures and dermal glycolipid, and to some extent calls into question the origin and structure of the ear plug itself. Further morphological characterization of ear plugs in bowheads is needed to better understand cell origin and ear plug formation.
Article
Terrestrial artiodactyls (even‐toed ungulates) inhabit some of the world's most extreme environments, including arid deserts and high elevations. As medium‐to‐large‐bodied mammals, artiodactyls have a suite of specialized physiologies to facilitate occupation of regions unavailable to other large mammals. One such physiology is selective brain cooling, wherein reduction of brain temperature below core body temperature has been demonstrated to reduce evaporative water loss. This physiology is enabled by an arterial heat‐exchanger called the carotid rete. The ubiquity of the carotid rete throughout the clade, as well as its evolutionary history, is currently uninvestigated. Here, I use osteological correlates to survey clade‐wide presence and morphology of the carotid rete, prior to conducting a preliminary evolutionary analysis. Nearly all living artiodactyls possess a carotid rete and are capable of selective brain cooling; however, major arteries supplying the rete are derived from different embryonic aortic arches on a suborder‐specific basis. Ancestral character estimation infers this pattern of variation to be the result of independent evolutionary processes, suggesting carotid rete homoplasy arising via parallelism. This is a surprising finding given the role this structure plays in driving a physiology that has been implicated in mitigating artiodactylan responses to extreme environmental conditions. Future studies should incorporate extinct species represented in the fossil record to better parse between parallel and convergent mechanisms, as well as to better understand the relationship between the carotid rete, selective brain cooling, and survivorship of climate perturbation. Anat Rec, 2018.
Chapter
Quantification and analysis of shape is an important component of many paleoecological studies. Geometric morphometrics is a powerful shape analysis tool that allows its user to compare entire regions of morphology, visualize shape differences between groups, and create visualizations based on real data. This method is rapidly becoming the standard for data collection and analysis in many fields such as anthropology, biology, ecology, forensics, paleontology, and zoology. Here, the basic procedures of geometric morphometrics are reviewed and a case study on the ecomorphology of the cervid calcaneus is provided to illustrate how geometric morphometrics can be used in paleoecological studies.
Chapter
The goal of ecomorphology is to identify morphological variation that is related to ecology (e.g., dietary preference or locomotor habits), with the aim of inferring ecological traits from morphological traits. This chapter reviews the basic principles of ecomorphology and provides many examples of ecomorphic studies in a range of taxa, with an emphasis on bovids (antelope and relatives). The focus of this chapter is on “applied” ecomorphology, which refers to studies that use ecomorphology as a tool to reconstruct environments. The chapter summarizes some of the strengths and weaknesses of the applied ecomorphic approach, and discusses future directions for studies using this methodology.
Article
Narwhals (Monodon monoceros) and belugas (Delphinapterus leucas) are the only extant members of the Monodontidae, and are charismatic Arctic‐endemic cetaceans that are at risk from global change. Investigating the anatomy and sensory apparatuses of these animals is essential to understanding their ecology and evolution, and informs efforts for their conservation. Here, we use X‐ray CT scans to compare aspects of the endocranial and inner ear labyrinth anatomy of extant monodontids and use the overall morphology to draw larger inferences about the relationship between morphology and ecology. We show that differences in the shape of the brain, vasculature, and neural canals of both species may relate to differences in diving and other behaviors. The cochleae are similar in morphology in the two species, signifying similar hearing ranges and a close evolutionary relationship. Lastly, we compare two different methods for calculating 90var – a calculation independent of body size that is increasingly being used as a proxy for habitat preference. We show that a ‘direct’ angular measurement method shows significant differences between Arctic and other habitat preferences, but angle measurements based on planes through the semicircular canals do not, emphasizing the need for more detailed study and standardization of this measurement. This work represents the first comparative internal anatomical study of the endocranium and inner ear labyrinths of this small clade of toothed whales.
Article
Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ∼MP7 and ∼MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.
Article
Almost all mammals communicate using sound, but few species produce complex songs. Two baleen whales sing complex songs that change annually, though only the humpback whale (Megaptera novaeangliae) has received much research attention. This study focuses on the other baleen whale singer, the bowhead whale (Balaena mysticetus). Members of the Spitsbergen bowhead whale population produced 184 different song types over a 3-year period, based on duty-cycled recordings from a site in Fram Strait in the northeast Atlantic. Distinct song types were recorded over short periods, lasting at most some months. This song diversity could be the result of population expansion, or immigration of animals from other populations that are no longer isolated from each other by heavy sea ice. However, this explanation does not account for the within season and annual shifting of song types. Other possible explanations for the extraordinary diversity in songs could be that it results either from weak selection pressure for interspecific identification or for maintenance of song characteristics or, alternatively, from strong pressure for novelty in a small population.
Article
Common hippos (Hippopotamus amphibius) live in murky waters and produce a variety of acoustic signals including underwater click trains considered to be social in function. We tested the hypothesis that click trains may function for underwater detection. We used observational and experimental methods involving 16 captive hippos to document the occurrence of click trains in different contexts and describe the acoustic parameters of the clicks. Male and female hippos produced click trains correlated with searching underwater for food items placed in their pools. Males produced click trains when alone supporting the hypothesis that these signals function for detection and are not only social in function. The frequency bandwidth of individual clicks varied and most were below 10 000 Hz. Click train production by hippos during underwater searches suggests a rudimentary form of echo-ranging that may function when other sensory systems are limited in their aquatic environment. © 2018
Article
MC5R is one of five melanocortin receptor genes found in placental mammals. MC5R plays an important role in energy homeostasis and is also expressed in the terminal differentiation of sebaceous glands. Among placental mammals there are multiple lineages that either lack or have degenerative sebaceous glands including Cetacea (whales, dolphins, and porpoises), Hippopotamidae (hippopotamuses), Sirenia (manatees and dugongs), Proboscidea (elephants), Rhinocerotidae (rhinos), and Heterocephalus glaber (naked mole rat). Given the loss or diminution of sebaceous glands in these taxa, we procured MC5R sequences from publicly available genomes and transcriptomes, supplemented by a newly generated sequence for Choeropsis liberiensis (pygmy hippopotamus), to determine if this gene remains intact or is inactivated in association with loss/reduction of sebaceous glands. Our data set includes complete MC5R sequences for 114 placental mammal species including two individuals of Mammuthus primigenius (woolly mammoth) from Oimyakon and Wrangel Island. Complete loss or inactivation of the MC5R gene occurs in multiple placental lineages that have lost sebaceous glands (Cetacea, West Indian manatee, African elephant, white rhinoceros) or are characterized by unusual skin (pangolins, aardvarks). Both M. primigenius individuals share inactivating mutations with the African elephant even though sebaceous glands have been reported in the former. MC5R remains intact in hippopotamuses and the naked mole rat, although slightly elevated dN/dS ratios in these lineages allow for the possibility that the accumulation of inactivating mutations in MC5R may lag behind the relaxation of purifying selection. For Cetacea and Hippopotamidae, the absence of shared inactivating mutations in two different skin genes (MC5R, PSORS1C2) is consistent with the hypothesis that semi-aquatic lifestyles were acquired independently in these clades following divergence from a common ancestor.
Article
Sauropterygia, a successful clade of marine reptiles abundant in aquatic ecosystems of the Mesozoic, inhabited nearshore to pelagic habitats over >180 million years of evolutionary history [1]. Aquatic vertebrates experience strong buoyancy forces that allow movement in a three-dimensional environment, resulting in structural convergences such as flippers and fish-like bauplans [2, 3], as well as convergences in the sensory systems. We used computed tomographic scans of 19 sauropterygian species to determine how the transition to pelagic lifestyles influenced the evolution of the endosseous labyrinth, which houses the vestibular sensory organ of balance and orientation [4]. Semicircular canal geometries underwent distinct changes during the transition from nearshore Triassic sauropterygians to the later, pelagic plesiosaurs. Triassic sauropterygians have dorsoventrally compact, anteroposteriorly elongate labyrinths, resembling those of crocodylians. In contrast, plesiosaurs have compact, bulbous labyrinths, sharing some features with those of sea turtles. Differences in relative labyrinth size among sauropterygians correspond to locomotory differences: bottom-walking [5, 6] placodonts have proportionally larger labyrinths than actively swimming taxa (i.e., all other sauropterygians). Furthermore, independent evolutionary origins of short-necked, large-headed "pliosauromorph" body proportions among plesiosaurs coincide with reductions of labyrinth size, paralleling the evolutionary history of cetaceans [7]. Sauropterygian labyrinth evolution is therefore correlated closely with both locomotory style and body proportions, and these changes are consistent with isolated observations made previously in other marine tetrapods. Our study presents the first virtual reconstructions of plesiosaur endosseous labyrinths and the first large-scale, quantitative study detailing the effects of increasingly aquatic lifestyles on labyrinth morphology among marine reptiles.
Chapter
The primate peripheral auditory organ closely resembles that of other terrestrial mammals. Acoustic communication has an important role in primate communities, and hearing characteristics are well-known for several species. In this chapter, morphological variation of the primate outer, middle, and inner ears is reviewed and is related to auditory data known from experimental work. Differences can be discerned among various primate groups, the greatest differences being between small, mainly nocturnal strepsirrhines and larger, mainly diurnal haplorhines. The evolutionary history of primate hearing is discussed in relation to different hypotheses of primate origins with the view that, as in the earliest mammals, the earliest primates were nocturnal and had good high-frequency hearing. Increased sensitivity to lower frequencies evolved later, although relatively early in the history of primates. This was made possible by an elongation of the cochlea and the disappearance of the secondary spiral lamina. The body size and ecology of primates is related to their ear size, and the role of hearing together with other sensory modalities, mainly vision and olfaction, is discussed.
Article
The Hippopotamidae have been a major component of the African wetland fauna for the last 7 million years, following the ‘Hippopotamine Event,’ i.e., the sudden emergence in the fossil record of the subfamily Hippopotaminae, including both extant species. The general dearth of African fossiliferous deposits dated between 9.5 Ma and 7.5 Ma concealed until now the evolution that led to the Hippopotamine Event and the subsequent success of these large semiaquatic herbivores. Part of this evolution is unveiled by the hippopotamid dental remains found at Chorora, a late Miocene site of the southern Afar Depression in Ethiopia spanning most of the fossil-depleted time interval. Although fragmentary, these remains represent a new, mid-sized hippopotamid species dated to ca. 8 Ma, as well as a somewhat younger, larger form. A cladistic analysis of a large array of cetartiodactyls indicates that the Chorora taxa were basal to the latest Miocene hippopotamines. The new species displays a mosaic of dental characters that support the attribution of the new species to a new genus within Hippopotaminae. The new fossils also clarify the course of early hippopotamine dental evolution. The Chorora hippopotamids suggest that transition to a marked abundance of hippopotamines with their unique dental pattern in African ecosystems occurred within a relatively short time interval, most probably between 8 Ma and 7.5 Ma. http://zoobank.org/urn:lsid:zoobank.org:pub:47B9381F-E3B5-40C9-B9AB-51CC3D0D3A8A SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Boisserie, J.-R., G. Suwa, B. Asfaw, F. Lihoreau, R. L. Bernor, S. Katoh, and Y. Beyene. 2017. Basal hippopotamines from the upper Miocene of Chorora, Ethiopia. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1297718.
Article
Theories of ecological diversification make predictions about the timing and ordering of character state changes through history. These theories are testable by "reconstructing" ancestor states using phylogenetic trees and measurements of contemporary species. Here we use maximum likelihood to estimate and evaluate the accuracy of ancestor reconstructions. We present likelihoods of discrete ancestor states and derive probability distributions for continuous ancestral traits. The methods are applied to several examples: diets of ancestral Darwin's finches; origin of inquilinism in gall wasps; microhabitat partitioning and body size evolution in scrubwrens; digestive enzyme evolution in artiodactyl mammals; origin of a sexually selected male trait, the sword, in platies and swordtails; and evolution of specialization in Anolis lizards. When changes between discrete character states are rare, the maximum-likelihood results are similar to parsimony estimates. In this case the accuracy of estimates is often high, with the exception of some nodes deep in the tree. If change is frequent then reconstructions are highly uncertain, especially of distant ancestors. Ancestor states for continuous traits are typically highly uncertain. We conclude that measures of uncertainty are useful and should always be provided, despite simplistic assumptions about the probabilistic models that underlie them. If uncertainty is too high, reconstruction should be abandoned in favor of approaches that fit different models of trait evolution to species data and phylogenetic trees, taking into account the range of ancestor states permitted by the data.
Book
Organized by functional neurologic system, the 3rd edition of this authoritative reference provides the most up-to-date information on neuroanatomy, neurophysiology, neuropathology, and clinical neurology as it applies to small animals, horses, and food animals. Accurate diagnosis is emphasized throughout with practical guidelines for performing neurologic examinations, interpreting examination results, and formulating effective treatment plans. In-depth disease descriptions, color images, and video clips reinforce important concepts and assist with diagnosis and treatment. Expert authors bring more than 50 years of experience in veterinary neuroanatomy and clinical neurology to this book - Dr. Alexander DeLahunta and Dr. Eric Glass offer their unique insights from both academic and practitioner perspectives. Disease content is presented in a logical case study format with three distinct parts: Description of the disorder Neuroanatomic diagnosis (including how it was determined, the differential diagnosis, and any available ancillary data) Course of the disease (providing final clinical or necropsy diagnosis and a brief discussion of the syndrome) More than 600 full-color photographs and line drawings, plus approximately 150 high-quality radiographs, visually reinforce key concepts and assist in reaching accurate diagnoses. The book comes with free access to 370 video clips on Cornell University's website that directly correlate to the case studies throughout the book and clearly demonstrate nearly every recognized neurologic disorder. High-quality MR images of the brain are presented alongside correlating stained transverse sections for in-depth study and comparison. Vivid photos of gross and microscopic lesions clearly illustrate the pathology of many of the disorders presented in the book.