Hypsodonty in ungulates: an adaptation for grass consumption or for foraging in open habitat?

Hypsodont (i.e. high-crowned) teeth have been interpreted as an indicator of feeding preferences and habitat selection in ungulates. For this reason, the degree of hypsodonty has been used for estimating the diet of ancient taxa and in palaeoenvironmental reconstructions. The goal of this study is to elucidate the relative importance of grass consumption and open habitat foraging in the development of hypsodont teeth, using novel computer techniques of knowledge discovery applied to a dataset of 134 species of artiodactyls and perissodactyls distributed among thirteen families. The results obtained suggest that high-crowned teeth represent an adaptation for feeding in an open habitat, although the minimum threshold of hypsodonty seems to increase with the relative length of the anterior part of the jaw. On the contrary, there is no direct relationship between the degree of hypsodonty and the percentage of grass consumed, except for the correspondence between grazing and dwelling in open habitats. A relatively wide muzzle evidences an adaptation for grass foraging in open and mixed habitats, but there are some non-grazing species from a closed habitat that also show wide muzzles. Thus, the hypsodonty index, combined with the length of the anterior part of the jaw and the width of the muzzle, allows accurate inferences on the ecological preferences of extinct ungulates.     

Relationships between oral morphology and feeding style in the Ungulata: a phylogenetically controlled evaluation

In ungulates it is argued that specialization in the consumption of a particular type of food (feeding style) is reflected in morphological adaptations of the organs involved in the selection, processing and digestion of food. We analysed the differences in size and morphology of some oral traits that have been functionally related to food-selection ability (muzzle width, incisor-arcade shape, incisor shape), prehension of food (incisor protrusion), food comminution (molar occlusal surface area, hypsodonty (high-crowned molars)) and intake rate (incisor breadth) between ungulate species with different feeding styles (browser, mixed feeder, grazer). Grazers were characterized by large-body-size species. After controlling only for body mass, we found that grazers had wider muzzles and incisors, more-protruding incisors and more-bulky and higher-crowned molars than did mixed feeders and browsers. When the analyses took into account both body mass and phylogeny, only body mass and two out of the three hypsodonty indexes used remained significantly different between feeding styles. Browsers were smaller, on average, than mixed feeders and grazers, whilst grazers and mixed feeders did not differ in size. Also, browsers had shorter and less-bulky molars than did mixed feeders and grazers; the latter two feeding styles did not differ from each other in any of the hypsodonty indexes. We conclude that the adaptation to different dietary types in most of the oral traits studied is subsumed by the effects of body mass and the sharing of common ancestors. We hypothesize that differences in the ability to exploit different food resources primarily result from differences in body mass between species, and also discuss why hypsodonty characterizes feeding styles.     

On the relationship between hypsodonty and feeding ecology in ungulate mammals,and its utility in palaeoecology

High‐crowned (hypsodont) teeth are widely found among both extant and extinct mammalian herbivores. Extant grazing ungulates (hoofed mammals) have hypsodont teeth (a derived condition), and so extinct hypsodont forms have usually been presumed to have been grazers. Thus, hypsodonty among ungulates has, over the past 150 years, formed the basis of widespread palaeoecological interpretations, and has figured prominently in the evolutionary study of the spread of grasslands in the mid Cenozoic. However, perceived inconsistencies between levels of hypsodonty and dental wear patterns in both extant and extinct ungulates have caused some workers to reject hypsodonty as a useful predictive tool in palaeobiology, a view that we consider both misguided and premature. Despite the acknowledged association between grazing and hypsodonty, the quantitative relationship of hypsodonty to the known ecology of living ungulate species, critical in making interpretations of the fossil record, was little studied until the past two decades. Also, much of the literature on ungulate ecology relevant to understanding hypsodonty has yet to be fully incorporated into the perspectives of palaeontologists. Here we review the history and current state of our knowledge of the relationship between hypsodonty and ungulate ecology, and reassert the value of hypsodonty for our understanding of ungulate feeding behaviour. We also show how soil consumption, rather than the consumption of grass plants per se, may be the missing piece of the puzzle in understanding the observed correlation between diets, habitats, and hypsodonty in ungulates. Additionally, we show how hypsodonty may impact life‐history strategies, and resolve some controversies regarding the relevance of hypsodonty to the prediction of the diets of extinct species. This in turn strengthens the utility of hypsodonty in the determination of past environmental conditions, and we provide a revised view of a traditional example of evolutionary trends in palaeobiology, that of the evolution of hypsodonty in horses and its correlation with the Miocene spread of grasslands in North America.     

Testing the Role of Cursorial Specializations as Adaptive Key Innovations in Paleocene-Eocene Ungulates of North America

Episodes of rapid faunal turnover in the fossil record are often used to examine processes driving macroevolutionary changes, such as competitive exclusion. The sudden appearance in the earliest Eocene of North America of artiodactyls and perissodactyls, and subsequent decline of endemic “condylarths” constitutes such an episode. It has been suggested that the specializations for high speed locomotion (cursoriality) that are present in artiodactyls and perissodactyls were key innovations of these orders accounting for their success in the Eocene and onwards. A quantitative geometric morphometric analysis of distal femoral articular morphology was used to examine changes in locomotor specializations in North American ungulates across the Paleocene-Eocene boundary. “Condylarths” were found to have displayed a broad range of locomotor adaptions, including cursoriality. The early Eocene had the broadest disparity in terms of taxonomic and locomotor contributions to morphological diversity. Changes in locomotor variety were associated with the disappearance of arboreal taxa, primarily “condylarths.” The initial impact of artiodactyls and perissodactyls in North America on existing locomotor diversity was limited and does not support a competitive exclusion hypothesis.     

Structural Morphology of Molars in Large Mammalian Herbivores: Enamel Content Varies between Tooth Positions

The distribution of dental tissues in mammalian herbivores can be very different from taxon to taxon. While grazers tend to have more elaborated and complexly folded enamel ridges, browsers have less complex enamel ridges which can even be so far reduced that they are completely lost. The gradient in relative enamel content and complexity of structures has so far not been addressed within a single species. However, several studies have noted tooth position specific wear rates in small mammals (rabbits, guinea pigs) which may be related to individual tooth morphology. We investigate whether differentiated enamel content by tooth position is also to be found in large herbivores. We use CT-scanning techniques to quantify relative enamel content in upper and lower molar teeth of 21 large herbivorous mammal species. By using a broad approach and including both perissodactyls and artiodactyls, we address phylogenetic intraspecific differences in relative enamel content. We find that enamel is highly unevenly distributed among molars (upper M1, M2, M3 and lower m1, m2, m3) in most taxa and that relative enamel content is independent of phylogeny. Overall, relative enamel content increases along the molar tooth row and is significantly higher in lower molars compared to upper molars. We relate this differential enamel content to prolonged mineralisation in the posterior tooth positions and suggest a compensatory function of m3 and M3 for functional losses of anterior teeth.     

The carotid rete and artiodactyl success

Since the Eocene, the diversity of artiodactyls has increased while that of perissodactyls has decreased. Reasons given for this contrasting pattern are that the evolution of a ruminant digestive tract and improved locomotion in artiodactyls were adaptively advantageous in the highly seasonal post-Eocene climate. We suggest that evolution of a carotid rete, a structure highly developed in artiodactyls but absent in perissodactyls, was at least as important. The rete confers an ability to regulate brain temperature independently of body temperature. The net effect is that in hot ambient conditions artiodactyls are able to conserve energy and water, and in cold ambient conditions they are able to conserve body temperature. In perissodactyls, brain and body temperature change in parallel and thermoregulation requires abundant food and water to warm/cool the body. Consequently, perissodactyls occupy habitats of low seasonality and rich in food and water, such as tropical forests. Conversely, the increased thermoregulatory flexibility of artiodactyls has facilitated invasion of new adaptive zones ranging from the Arctic Circle to deserts and tropical savannahs.     

Muscles of the masticatory apparatus in two genera of hyraces (Procavia and Heterohyrax)

Subungulate hyraces are similar to the condition assumed to have characterized primitive ungulates and subungulates by virtue of their small body size, relatively unspecialized cranial and postcranial anatomy, and primitive type of lophodont dentition. The muscles of mastication of Procavia habessinica and Heterohyrax brucei are here compared with those of other mammals, both with ungulates, as an example of more specialized mammals, and with opossums, as an example of more generalized mammals, to determine aspects of hyrax myology that represent the retention of a condition primitive for herbivorous mammals. The masticatory muscles of hyraces retain the primitive ungulate/subungulate condition in the large, complexly subdivided temporalis, and in the enlarged, pinnated, bilayered medial pterygoid. The medial pterygoid originates from the pterygoid hamulus, a condition that may also be primitive for this assemblage. The large complex superficial masseter is derived compared with the condition in ruminant artiodactyls, but may represent the condition primitive for perissodactyls. The architectural modifications of this muscle in hyraces may represent adaptations to allow a wide gape threat display. Hyraces possess a posterior belly of the digastric alone, paralleling the condition in some perissodactyls. They possess a large and complexly subdivided styloglossus, which may be a shared derived character of subungulates. Hyraces are unique among ungulates and subungulates in the extreme reduction of the anterior hyoid cornua, and may be unique among mammals in the development of paired lingual processes from the ceratohyal ossifications.     

Inferring habitat and feeding behaviour of early Miocene notoungulates from Patagonia

Cassini, G.H., Mendoza, M., Vizcaíno, S.F. & Bargo, M.S. 2011: Inferring habitat and feeding behaviour of early Miocene notoungulates from Patagonia. Lethaia, Vol. 44, pp. 153–165. Notoungulates, native fossil mammals of South America, have been usually studied from a taxonomic point of view, whereas their palaeobiology has been largely neglected. For example, morpho‐functional or eco‐morphological approaches have not been rigorously applied to the masticatory apparatus to propose hypothesis on dietary habits. In this study, we generate inferences about habitat and feeding preferences in five Santacrucian genera of notoungulates of the orders Typotheria and Toxodontia using novel computer techniques of knowledge discovery. The Santacrucian (Santa Cruz Formation, late‐early Miocene) fauna is particularly appropriate for this kind of studies due to its taxonomic richness, diversity, amount of specimens recorded and the quality of preservation. Over 100 extant species of ungulates, distributed among 13 families of artiodactyls and perissodactyls, were used as reference samples to reveal the relationships between craniodental morphology and ecological patterns. The results suggest that all Santacrucian notoungulates present morphologies characteristic of open habitats’ extant ungulates. Although the Toxodontia exhibits the same morphological pattern of living mixed‐feeders and grazers, the Typotheria shows exaggerated traits of specialized grazer ungulates. □Craniodental morphology, ecomorphology, fossil ungulates, knowledge discovery, South America.     

Mandibles and molars of the wood mouse, Apodemus sylvaticus (L.): integrated latitudinal pattern and mosaic insular evolution

Aim The distinct nature of island populations has traditionally been attributed either to adaptation to particular insular conditions or to random genetic effects. In order to assess the relative importance of these two disparate processes, insular effects were addressed in the European wood mouse, Apodemus sylvaticus (Linnaeus, 1758). Location Wood mice from 33 localities on both mainland and various Atlantic and western Mediterranean islands were considered. This sampling covers only part of the latitudinal range of A. sylvaticus but included the two main genetic clades identified by previous studies. Islands encompass a range of geographical conditions (e.g. small islands fringing the continent through large and isolated ones). Methods The insular syndrome primarily invokes variations in body size, but ecological factors such as release from competition, niche widening and food availability should also influence other characters related to diet. In the present study, the morphology of the wood mice was quantified based on two characters involved in feeding: the size and shape of the mandibles and first upper molars. The size of the mandible is also a proxy for the body size of the animal. Patterns of morphological differentiation of both features were estimated using two‐dimensional outline analysis based on Fourier methods. Results Significant differences between mainland and island populations were observed in most cases for both the mandibles and molars. However, molars and mandibles displayed divergent patterns. Mandible shape diverged mostly on islands of intermediate remoteness and competition levels, whereas molars exhibited the greatest shape differentiation on small islands, such as Port‐Cros and Porquerolles. A mosaic pattern was also displayed for size. Body and mandible size increased on Ibiza, but molar size remained similar to mainland populations. Mosaic patterns were, however, not apparent in the mainland populations. Congruent latitudinal variations were evident for the size and shape of both mandibles and molars. Main conclusions Mosaic evolution appears to characterize insular divergence. The molar seems to be more prone to change with reduced population size on small islands, whereas the mandible could be more sensitive to peculiar environmental conditions on large and remote islands.     

Virtual endocranial cast of earliest Eocene Diacodexis (Artiodactyla, Mammalia) and morphological diversity of early artiodactyl brains

The study of brain evolution, particularly that of the neocortex, is of primary interest because it directly relates to how behavioural variations arose both between and within mammalian groups. Artiodactyla is one of the most diverse mammalian clades. However, the first 10 Myr of their brain evolution has remained undocumented so far. Here, we used high-resolution X-ray computed tomography to investigate the endocranial cast of Diacodexis ilicis of earliest Eocene age. Its virtual reconstruction provides unprecedented access to both metric parameters and fine anatomy of the most complete endocast of the earliest artiodactyl. This picture is assessed in a broad comparative context by reconstructing endocasts of 14 other Early and Middle Eocene representatives of basal artiodactyls, allowing the tracking of the neocortical structure of artiodactyls back to its simplest pattern. We show that the earliest artiodactyls share a simple neocortical pattern, so far never observed in other ungulates, with an almond-shaped gyrus instead of parallel sulci as previously hypothesized. Our results demonstrate that artiodactyls experienced a tardy pulse of encephalization during the Late Neogene, well after the onset of cortical complexity increase. Comparisons with Eocene perissodactyls show that the latter reached a high level of cortical complexity earlier than the artiodactyls.     

Variation in mandible shape and body size of house mice Mus musculus in five separate New Zealand forest habitats

This study investigates variation in house mouse Mus musculus body size and mandible shape across New Zealand, using geometric morphometrics and biomechanical advantage analyses. The Mus phylogroups currently known in New Zealand include Mus musculus domesticus, M. m. musculus and M. m. castaneus. We examined samples of house mice inhabiting five different podocarp and beech forest environments across the North and South Islands (Pureora Forest, Zealandia Wildlife Sanctuary, Craigieburn Forest Park, Eglinton Valley and Hollyford Valley). Significant variation in mandible shape and body size was found between all five forest populations. South Island mice had larger bodies and greater mechanical advantage in the temporalis muscle compared with their North Island counterparts. Zealandia Sanctuary mouse mandibles were broader and shorter than South Island mouse mandibles, and had greater masseter muscle advantage. Centroid size and body weight, but not head-body length, varied significantly with two distinct genetic haplotypes. Finally, annual rainfall was the most significant covariate with mandible shape. Higher rainfall locations were generally associated with soft-food related mandible shapes, while lower rainfall correlated with hard-food mandible shapes. This preliminary investigation provides the framework for further research into mandible shape and body size variation in New Zealand house mice.     

西伯利亚南部旧石器时代晚期哺乳动物群动态学

旧石器时代晚期至全新世发生了广泛而迅速的动物群更替,为研究动物群对快速环境变化的反应提供了有益的信息。自中新世以来中亚地区一直是欧亚大陆向更开阔和干旱环境转变的焦点。利用已发表的俄罗斯贝加尔湖和阿尔泰山区域旧石器时代晚期的文献资料,通过对大型食草动物臼齿齿冠高度、食性和体型大小的分析,绘制了环境变化图。结果表明,在旧石器时代最晚期(晚冰期),平均齿冠高度、平均体型大小均有所减小,食物取向从食草向食树叶转变,这意味着气候变得温暖,降雨量增加,森林也有可能扩展。常见种(出现于超过25%的地点)更强烈地表现出这一趋势。齿冠高度随海拔高度而增大,提示降雨量减少。从齿冠高度得出的降雨量数值与根据其他指标所估算的数值之间的偏差可能随海拔高度而增大。影响西伯利亚南部猛犸象动物群消亡的主要变化可能开始于末次冰期冰盛期(LGM)之后。猛犸象和奇蹄动物出现频率的降低与大型哺乳动物的绝灭相一致,它们和偶蹄类的差异则反映了气候的变化。人类的过度开发可能给区域环境带来了更大的压力,使得哺乳动物的种群数量下降,导致它们更迅速地绝灭。     

Phylogenetic and environmental components of morphological variation: skull, mandible, and molar shape in marmots (Marmota, Rodentia)

The phenotype is a product of its phylogenetic history and its recent adaptation to local environments, but the relative importance of the two factors is controversial. We assessed the effects of diet, habitat, elevation, temperature, precipitation, body size, and mtDNA genetic divergence on shape variation in skulls, mandibles, and molars, structures that differ in their genetic and functional control. We asked whether these structures have adapted to environment to the same extent and whether they retain the same amount of phylogenetic signal. We studied these traits in intra- and interspecific populations of Eurasian marmots whose last common ancestor lived 2-5 million years ago. Path Analysis revealed that body size explained 10% of variation in skulls, 7% in mandibles, and 15% in molars. Local vegetation explained 7% of variation in skulls, 11% in mandibles, and 12% in molars. Dietary category explained 25% of variation in skulls, 11% in mandibles, and 9% in molars. Cyt b mtDNA divergence (phylogeny) explained 15% of variation in skulls, 7% in mandibles, and 5% in molars. Despite the percentages of phylogenetic variance, maximum-likelihood trees based on molar and skull shape recovered most phylogenetic groupings correctly, but mandible shape did not. The good performance of molars and skulls was probably due to different factors. Skulls are genetically and functionally more complicated than teeth, and they had more mathematically independent components of variation (5-6-in skulls compared to 3-in molars). The high proportion of diet-related variance was not enough to mask the phylogenetic signal. Molars had fewer independent components, but they also have less ecophenotypic variation and evolve more slowly, giving each component a proportionally stronger phylogenetic signal. Molars require larger samples for each operational taxonomic unit than the other structures because the proportion of within-taxon to between-taxon variation was higher. Good phylogenetic signal in quantitative skeletal morphology is likely to be found only when the taxa have a common ancestry no older than hundreds of thousands or millions of years (1% to 10% mtDNA divergence)--under these conditions skulls and molars provide stronger signal than mandibles.     

Developmental constraints revealed by co-variation within and among molar rows in two murine rodents

SUMMARY Morphological integration corresponds to interdependency between characters that can arise from several causes. Proximal causes of integration include that different phenotypic features may share common genetic sets and/or interact during their development. Ultimate causes may be the prolonged effect of selection favoring integration of functionally interacting characters, achieved by the molding of these proximal causes. Strong and direct interactions among successive teeth of a molar row are predicted by genetic and developmental evidences. Functional constraints related to occlusion, however, should have selected more strongly for a morphological integration of occluding teeth and a corresponding evolution of the underlying developmental and genetic pathways. To investigate how these predictions match the patterns of phenotypic integration, we studied the co‐variation among the six molars of the murine molar row, focusing on two populations of house mice (Mus musculus domesticus) and wood mice (Apodemus sylvaticus). The size and shape of the three upper and lower molars were quantified and compared. Our results evidenced similar patterns in both species, size being more integrated than shape among all the teeth, and both size and shape co‐varying strongly between adjacent teeth, but also between occluding teeth. Strong co‐variation within each molar row is in agreement with developmental models showing a cascade influence of the first molar on the subsequent molars. In contrast, the strong co‐variation between molars of the occluding tooth rows confirms that functional constraints molded patterns of integration and probably the underlying developmental pathways despite the low level of direct developmental interactions occurring among molar rows. These patterns of co‐variation are furthermore conserved between the house mouse and the wood mouse that diverged >10 Ma, suggesting that they may constitute long‐running constraints to the diversification of the murine rodent dentition.     

More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen

Recent phylogenetic analyses of DNA sequences suggest that cetaceans (whales) and hippopotamid artiodactyls (hippos) are extant sister taxa. Consequently, the shared aquatic specializations of these taxa may be synapomorphies. This molecular view is contradicted by paleontological data that overwhelmingly support a monophyletic Artiodactyla (even-toed ungulates) and a close relationship between Cetacea and extinct mesonychian ungulates. According to the fossil evidence, molecular, behavioral, and anatomical resemblances between hippos and whales are interpreted as convergences or primitive retentions. In this report, competing interpretations of whale origins are tested through phylogenetic analyses of the blood-clotting protein gene gamma- fibrinogen from cetaceans, artiodactyls, perissodactyls (odd-toed ungulates), and carnivores (cats, dogs, and kin). In combination with published DNA sequences, the gamma-fibrinogen data unambiguously support a hippo/whale clade and are inconsistent with the paleontological perspective. If the phylogeny favored by fossil evidence is accepted, the convergence at the DNA level between Cetacea and Hippopotamidae is remarkable in its distribution across three genetic loci: gamma-fibrinogen, the linked milk casein genes, and mitochondrial cytochrome b.      

Correlations between feeding type and mandibular morphology in the sika deer

Morphological comparisons of the sika deer Cervus nippon mandible and molars were conducted between two (northern and southern) Japanese subspecific lineages and among local populations of different ('grazer' or 'intermediate feeder') feeding types. The northern lineage showed greater M₁ breadth, M₃ hypsodonty and mandibular corpus height than the southern lineage. Such differences were not observed between the 'grazer' and 'intermediate feeder' populations of the northern lineage. However, a northern population, which inhabits a particularly harsh environment (Kinkazan Island), had the largest values of relative molar size and hypsodonty, although this was not statistically significant. These results imply that, in the Japanese sika deer, the selective pressures acting on the current 'grazer' populations are not strong enough to bring out noticeable adaptive change in molar size and hypsodonty, but adaptive change in these traits may occur in an environment that promotes excessive molar wear, more than that seen in the current sika deer habitats of Japan. Combined with what is known of the Pleistocene history of the sika deer, we infer that the ancestral population of the northern Japanese lineage likely acquired their relatively larger and more hypsodont molars in an extremely harsh environment during the last or previous glacial periods.     

Patterns of size variation and correlation in the dentition of the red colobus monkey (Colobus badius)

There have been numerous studies on variability and correlation in dental crown size, but the significance of the resulting patterns remains unclear. Regions of low variation and high correlation have been hypothesized to represent the poles of Butler's morphological fields, to be related to absolute tooth size, or to be related to morphological complexity of the teeth and functional efficiency. Variation and correlation of tooth lengths and breadths were investigated in 138 red colobus monkeys to further assess the relations among size associations, crown morphology, and absolute tooth size. In the maxilla and mandible, the postcanine teeth are the most highly correlated and least variable, followed by the incisors, then the canines. There are also lower correlations between premolars and molars than within either group. While there appears to be a relation between degree of morphological differentiation and levels of correlation and variation, there are no notable differences in the correlation of opponents along the dental arcade, which is the most important functional consideration. This suggests that different levels of correlation and variation within upper or lower teeth are “artifacts” of tooth dimensions that contribute to different geometric designs in different tooth groups as the germs develop. This morphological effect is coupled with the influence of integration fields, indicated by higher variability and lower correlations of the third molar, the largest or most molarized tooth. It is concluded that there are wide functional tolerances in occlusion with respect to the gross dimensions of dental crowns and their interrelationships.     

Post‐weaning ontogeny of the mandible in fossorial water voles: ecological and evolutionary implications

Ventura, J. and Casado‐Cruz, M. 2011. Post‐weaning ontogeny of the mandible in fossorial water voles: ecological and evolutionary implications. —Acta Zoologica (Stockholm) 92 : 12–20. Geometric morphometrics was applied to the mandible of fossorial water voles (Arvicola terrestris monticola) to determine size and shape variations in this structure during post‐weaning ontogeny. The sample consisted of collection specimens obtained in the Aran Valley (Spain), which were grouped into six age classes. Mandible size and shape did not differ significantly between sexes, but between age classes. Mandible size accounted significantly for the shape variation. After the size‐related differences were removed, the mandible shape did not show significant sexual dimorphism but differences by age remained significant. The main shape changes occur between the third and tenth weeks of life and are related to the shift from suckling to a herbivorous diet. Although mandible shape was less remodelled after that age, an appreciable variation also occurs during adulthood. Age‐related changes lead to enhancing the digging potential of the mandible, which in adults becomes a robust structure with an increased surface and stronger crests for muscle insertion. As part of the mandible shape variation was not related to the size‐dependent adjustment and diet does not vary significantly between juvenile and adult voles, shape changes that occur during adulthood can be related to the mechanical stress derived from digging activities.     

Tooth Enamel Microstructure of Living and Extinct Hyracoids Reveals Unique Enamel Types Among Mammals

Among medium- to large-sized terrestrial ‘ungulates,’ there is often a relationship between increasing body size, correlated changes in diet, and increased complexity of the enamel microstructures [notably the development of Hunter-Schreger bands (HSB)]. An exhaustive survey of the enamel microstructures of living and extinct Hyracoidea demonstrates, however, that the Schmelzmuster within this order of mammals is generally one-layered and formed by radial enamel despite a large range of body sizes and dietary adaptations; HSB are remarkably absent. Radial enamel is characteristic of early diverging hyracoids, as well as more derived members of the extinct families Geniohyidae and Pliohyracidae, and the extant Procaviidae. Only some large ‘Saghatheriidae,’ and all members of the family Titanohyracidae, developed a more complex enamel microstructure (i.e., with prisms decussating), a unique condition among Mammalia that we name ‘bundled enamel’ (BE). This structure is reminiscent to some degree of both the ‘Pyrotherium enamel’ and the ‘3D enamel’ of proboscideans. Hyracoids with BE represented a major component of the diversity of mid- to large-sized herbivores during the Paleogene in Africa. Like HSB, which are developed by most other ‘ungulates,’ the BE is regarded as a device for resisting propagation of cracks during mastication. Hyracoids never developed however the ‘modified radial enamel’ that is characteristic of most large and hypsodont perissodactyls and artiodactyls that entered Africa during the Miocene.     

Mandible shape in marsupial and placental carnivorous mammals: a morphological comparative study using geometric morphometrics

We analysed mandible shape of the orders Dasyuromorpha, Didelphimorphia, and Carnivora using two‐dimensional geometric morphometrics, in order to explore the relationship between shape, size, and phylogeny. We studied 541 specimens, covering most of the genera of the terrestrial Carnivora (115 species) and a wide sample of marsupials (36 species). The observed shape variation had an ecological component. As an example, omnivorous carnivores have thick mandibles and large talonids in the carnassials, while hypercarnivores possess short mandibles and reduced talonids. There is also a discrimination between different taxonomic groups (i.e. marsupials and Carnivora), indicating some kind of constraint. Size explains a large percentage of total variance (large species had shorter and stronger mandibles, with anteriorly displaced carnassials), was significant when phylogeny was taken into account with a comparative method, but not when size and shape were optimized on the phylogeny. Carnivora presents a larger disparity and variation in body size, which could be related to the difference in teeth replacement. The optimization of mandible shape on the phylogenetic tree indicates that functional aspects, such as diet, are a key factor in the evolution of the carnivore mandible, but also that there is a phylogenetic pattern that cannot be explained by differences in diet alone. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 836–855.