Mixed evidence for early bursts of morphological evolution in extant clades

Macroevolutionary theory predicts high rates of evolution should occur early in a clade's history as species exploit ecological opportunity. Evidence from the fossil record has shown a high prevalence of early bursts in morphological evolution, but recent work has provided little evidence for early high rates in the evolution of extant clades. Here, I test the prevalence of early bursts in extant data using phylogenetic comparative methods. Existing models are extended to allow a shift from a background Brownian motion (BM) process to an early burst process within subclades of phylogenies, rather than an early burst being applied to an entire phylogenetic tree. This nested early burst model is compared to other modes of evolution that can occur within subclades, such as evolution with a constraint (Ornstein‐Uhlenbeck model) and nested BM rate shift models. These relaxed models are validated using simulations and then are applied to body size evolution of three major clades of amniotes (mammals, squamates and aves) at different levels of taxonomic organization (order, family). Applying these unconstrained models greatly increases the support for early bursts within nested subclades, and so early bursts are the most common model of evolution when only one shift is analysed. However, the relative fit of early burst models is worse than models that allow for multiple shifts of the BM or OU process. No single‐shift or homogenous model is superior to models of multiple shifts in BM or OU evolution, but the patterns shown by these multirate models are generally congruent with patterns expected from early bursts.     

Evolution of morphology and locomotor performance in anurans: relationships with microhabitat diversification

The relationships between morphology, performance, behavior and ecology provide evidence for multiple and complex phenotypic adaptations. The anuran body plan, for example, is evolutionarily conserved and shows clear specializations to jumping performance back at least to the early Jurassic. However, there are instances of more recent adaptation to habit diversity in the post‐cranial skeleton, including relative limb length. The present study tested adaptive models of morphological evolution in anurans associated with the diversity of microhabitat use (semi‐aquatic arboreal, fossorial, torrent, and terrestrial) in species of anuran amphibians from Brazil and Australia. We use phylogenetic comparative methods to determine which evolutionary models, including Brownian motion (BM) and Ornstein‐Uhlenbeck (OU) are consistent with morphological variation observed across anuran species. Furthermore, this study investigated the relationship of maximum distance jumped as a function of components of morphological variables and microhabitat use. We found there are multiple optima of limb lengths associated to different microhabitats with a trend of increasing hindlimbs in torrent, arboreal, semi‐aquatic whereas fossorial and terrestrial species evolve toward optima with shorter hindlimbs. Moreover, arboreal, semi‐aquatic and torrent anurans have higher jumping performance and longer hindlimbs, when compared to terrestrial and fossorial species. We corroborate the hypothesis that evolutionary modifications of overall limb morphology have been important in the diversification of locomotor performance along the anuran phylogeny. Such evolutionary changes converged in different phylogenetic groups adapted to similar microhabitat use in two different zoogeographical regions.     

Phylogenetic topology mapped onto dietary ecospace reveals multiple pathways in the evolution of the herbivorous niche in African Bovidae

Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy. More recently, mesowear and stable isotope analyses have been used to great effect to reconstruct dietary preferences. We use these dietary proxies to construct a morphology‐free dietary ecospace and examine the topology of the phylogenetic relationships of African bovids mapped onto this ecospace. The reconstructed dietary ecospace provides evidence for four distinct dietary classes: species with C3‐ or C4‐dominated diets that produce low or high occlusal relief, likely related to diets high or low in abrasives, respectively. We detected no evidence for a discrete mixed feeder category; the species often categorized as such represent the end members of groups of species with either C3‐ or C4‐ dominated diets. Our analysis reveals high variability within the C4 grazing ecospace, and phylogenetic evidence indicates at least two pathways to grazing, likely related to the abrasive qualities of ingested food, which may be determined by the moisture content or the height of consumed grasses. These different pathways probably contribute to the high diversity of African grazers, both today and in the fossil record. C3 browsers (non‐frugivores) also display a high degree of variation, but there are no species associated with highly abrasive diets and there is evidence for only a single evolutionary pathway. We find evidence for only one evolutionary route towards frugivory, which includes species with diets that produce both high and low occlusal reliefs. The cause of abrasive wear in frugivores may be related to grit and/or the hard parts of fruits, but this requires further examination.     

Arthropod remains in the oral cavities of fossil reptiles support inference of early insectivory

Inference of feeding preferences in fossil terrestrial vertebrates (tetrapods) has been drawn predominantly from craniodental morphology, and less so from fossil specimens preserving conclusive evidence of diet in the form of oral and/or gut contents. Recently, the pivotal role of insectivory in tetrapod evolution was emphasized by the identification of putative insectivores as the closest relatives of the oldest known herbivorous amniotes. We provide the first compelling evidence for insectivory among early tetrapods on the basis of two 280-million-year-old (late Palaeozoic) fossil specimens of a new species of acleistorhinid parareptile with preserved arthropod cuticle on their toothed palates. Their dental morphology, consisting of homodont marginal dentition with cutting edges and slightly recurved tips, is consistent with an insectivorous diet. The intimate association of arthropod cuticle with the oral region of two small reptiles, from a rich fossil locality that has otherwise not produced invertebrate remains, strongly supports the inference of insectivory in the reptiles. These fossils lend additional support to the hypothesis that the origins and earliest stages of higher vertebrate evolution are associated with relatively small terrestrial insectivores.     

Relative embryo length as an adaptation to habitat and life cycle in Apiaceae

? The factors driving the evolution of the relative embryo length in Apiaceae were examined. We tested the hypothesis that seeds with large relative embryo length, because of more rapid germination, are beneficial in dry and open habitats and for short-lived species. We also analyzed to what extent delayed germination as a result of embryo growth can be considered a dormancy mechanism. ? Hypotheses were tested by correlating the relative embryo length with other plant traits, habitat and climatic variables. The adaptive nature of the relative embryo length was determined by comparing the performance of a pure drift, Brownian motion (BM) model of trait evolution with that of a selection-inertia, Ornstein-Uhlenbeck (OU) model. ? A positive correlation of the relative embryo length with germination speed and negative correlations with the amount of habitat shade, longevity and precipitation were found. An OU model, in which the evolution of longer embryos corresponded to a transition to habitats of high light, or to a short life cycle, outperformed significantly a BM model. ? The results indicated that the relative embryo length may have evolved as an adaptation to habitat and life cycle, whereas dormancy was mainly related to temperature at the sampling sites.     

Geometric morphometrics of mandibles for dietary differentiation of Bovidae (Mammalia: Artiodactyla)

The mammalian family Bovidae has been widely studied in ecomorphological research, with important applications to paleoecological and paleohabitat reconstructions. Most studies of bovid craniomandibular features in relation to diet have used linear measurements. In this study, we conduct landmark-based geometric-morphometric analyses to evaluate whether different dietary groups can be distinguished by mandibular morphology. Our analysis includes data for 100 species of extant bovids, covering all bovid tribes and 2 dietary classifications. For the first classification with 3 feeding categories, we found that browsers (including frugivores), mixed feeders, and grazers are moderately well separated using mandibular shape. A finer dietary classification (frugivore, browser, browser–grazer intermediate, generalist, variable grazer, and obligate grazer) proved to be more useful for differentiating dietary extremes (frugivores and obligate grazers) but performed equally or less well for other groups. Notably, frugivorous bovids, which belong in tribe Cephalophini, have a distinct mandibular shape that is readily distinguished from all other dietary groups, yielding a 100% correct classification rate from jackknife cross-validation. The main differences in mandibular shape found among dietary groups are related to the functional needs of species during forage prehension and mastication. Compared with browsers, both frugivores and grazers have mandibles that are adapted for higher biomechanical demand of chewing. Additionally, frugivore mandibles are adapted for selective cropping. Our results call for more work on the feeding ecology and functional morphology of frugivores and offer an approach for reconstructing the diet of extinct bovids.     

Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogency, and behavioural ecology

Sexual dimorphism in craniodental features is investigated in a sample of 45 carnivore species in relation to allometry, phylogeny, and behavioural ecology. Dimorphism is more pronounced in both upper and lower canine size and strength than in carnassial size, skull dimensions and biomechanical features, but all dimorphism indices covary. As with most morphological characters, differences in canine sexual dimorphism are significantly related to phylogeny, estimated from either taxonomic rankings or a limited matrix of molecular distances; in particular, mustelids, felids and procyonids are more dimorphic than other carnivore families. Thus, because of problems related to species dependence in comparative data, remaining analyses are based on phylogenetically transformed values using a spatial autoregressive method.
In contrast to other mammals, sexual dimorphism in carnivore canines is not correlated with differences in body weight, skull length or basicranial axis length. Nor is it correlated with categorical variables of activity pattern, habitat, or diet. In our Carnivore sample, canine dimorphism is related only to breeding system: uni-male, group-living (harem) species have significantly greater canine dimorphism than multi-male, multi-female groups and monogamous pair-bonding species. By contrast, dimorphism in carnassial size is related to dietary differences, specifically greater dimorphism in meat-eating species, and not breeding patterns. Dimorphism in estimates of jaw muscle size suggest functional demands from both diet and breeding type. It is concluded that, befitting patterns of heterodont dentition, sexual selection influences variation in canine dimorphism while feeding ecology is related to carnassial dimorphism.     

The isotopic ecology of African mole rats informs hypotheses on the evolution of human diet

The diets of Australopithecus africanus and Paranthropus robustus are hypothesized to have included C4 plants, such as tropical grasses and sedges, or the tissues of animals which themselves consumed C4 plants. Yet inferences based on the craniodental morphology of A. africanus and P. robustus indicate a seasonal diet governed by hard, brittle foods. Such mechanical characteristics are incompatible with a diet of grasses or uncooked meat, which are too tough for efficient mastication by flat, low-cusped molars. This discrepancy, termed the C4 conundrum, has led to the speculation that C4 plant underground storage organs (USOs) were a source of nutrition for hominin species. We test this hypothesis by examining the isotopic ecology of African mole rats, which consume USOs extensively. We measured delta18O and delta13C of enamel and bone apatite from fossil and modern species distributed across a range of habitats. We show that delta18O values vary little and that delta13C values vary along the C3 to C4/CAM-vegetative axis. Relatively high delta13C values exist in modern Cryptomys hottentotus natalensis and Cryptomys spp. recovered from hominin-bearing deposits. These values overlap those reported for A. africanus and P. robustus and we conclude that the USO hypothesis for hominin diets retains certain plausibility.     

Allometry, sexual dimorphism, and phylogeny: A cladistic analysis of extant African papionins using craniodental data

This study conducts a phylogenetic analysis of extant African papionin craniodental morphology, including both quantitative and qualitative characters. We use two different methods to control for allometry: the previously described narrow allometric coding method, and the general allometric coding method, introduced herein. The results of this study strongly suggest that African papionin phylogeny based on molecular systematics, and that based on morphology, are congruent and support a Cercocebus/Mandrillus clade as well as a Papio/Lophocebus/Theropithecus clade. In contrast to previous claims regarding papionin and, more broadly, primate craniodental data, this study finds that such data are a source of valuable phylogenetic information and removes the basis for considering hard tissue anatomy “unreliable” in phylogeny reconstruction. Among highly sexually dimorphic primates such as papionins, male morphologies appear to be particularly good sources of phylogenetic information. In addition, we argue that the male and female morphotypes should be analyzed separately and then added together in a concatenated matrix in future studies of sexually dimorphic taxa. Character transformation analyses identify a series of synapomorphies uniting the various papionin clades that, given a sufficient sample size, should potentially be useful in future morphological analyses, especially those involving fossil taxa.     

Fine-scale phenotypic change across a species transition zone in the genus neotoma: disentangling independent evolution from phylogenetic history

Zones of secondary contact between closely related species provide a rare opportunity to examine evidence of evolutionary processes that reinforce species boundaries and/or promote diversification. Here, we report on genetic and morphological variation in two sister species of woodrats, Neotoma fuscipes and N. macrotis, across a 30-km transition zone in the Sierra Nevada of California. We assessed whether these lineages readily hybridize, and whether their morphology suggests ecological interactions favoring phenotypic diversification. We combined measurements of body size and 11 craniodental traits from nine populations with genetic data to examine patterns of variation within and between species. We used phylogenetic autocorrelation methods to estimate the degree to which phenotypic variation in our dataset arose from independent evolution within populations versus phylogenetic history. Although no current sympatry or hybridization was evident, craniodental morphology diverged in both lineages near their distributional limits, whereas body size converged. The shift in craniodental morphology arose independently within populations whereas body size retained a strong phylogenetic signal, yet both patterns are consistent with expectations of phenotypic change based on different models of resource competition. Our findings demonstrate the importance of examining a suite of morphological traits across contact zones to provide a more complete picture of potential ecological interactions: competition may drive both diversification and convergence in different phenotypic traits.     

A Comparative Test of Adaptive Explanations for Hypsodonty in Ungulates and Rodents

Hypsodonty has long been recognized as an adaptation for grazing: grazing is suggested to increase tooth wear due to endogenous (e.g., fiber, silica) and/or exogenous (e.g., dust, grit) properties of ingested food. However, it is unknown whether tooth crown height is correlated with the mastication of high fiber or silica in grasses, the ingestion of external abrasives, or both. Furthermore, comparative studies of hypsodonty have not explicitly taken into account phylogenetic biases due to shared ancestry in tooth morphology and/or feeding behavior. This study highlights the relationship between molar crown height and feeding habits in African ungulates and South American rodents when phylogenetic effects are controlled. Among ungulates, high hypsodonty indices are significantly associated with specific plant and foraging height preferences, while habitat and climate show no correlation with tooth crown height. For rodents, grass-eating species are significantly more hypsodont than frugivorous or folivorous species, and arboreal rodents are less hypsodont than terrestrial species. These results as well as those of a posteriori analyses controlling for aspects of the behavioral ecology (e.g., grass-eating, substrate preference) of the sample species confirm the role of both diet and grit in shaping the evolution of cheek tooth crown height in herbivorous mammals.     

Functional morphology of the bovid astragalus in relation to habitat: Controlling phylogenetic signal in ecomorphology

Bovid astragali are one of the most commonly preserved bones in the fossil record. Accordingly, astragali are an important target for studies seeking to predict the habitat preferences of fossil bovids based on bony anatomy. However, previous work has not tested functional hypotheses linking astragalar morphology with habitat while controlling for body size and phylogenetic signal. This article presents a functional framework relating the morphology of the bovid astragalus to habitat‐specific locomotor ecology and tests four hypotheses emanating from this framework. Highly cursorial bovids living in structurally open habitats are hypothesized to differ from their less cursorial closed‐habitat dwelling relatives in having (1) relatively short astragali to maintain rotational speed throughout the camming motion of the rotating astragalus, (2) a greater range of angular excursion at the hock, (3) relatively larger joint surface areas, and (4) a more pronounced “spline‐and‐groove” morphology promoting lateral joint stability. A diverse sample of 181 astragali from 50 extant species was scanned using a Next Engine laser scanner. Species were assigned to one of four habitat categories based on the published ecological literature. A series of 11 linear measurements and three joint surface areas were measured on each astragalus. A geometric mean body size proxy was used to size‐correct the measurement data. Phylogenetic generalized least squares (PGLS) was used to test for differences between habitat categories while controlling for body size differences and phylogenetic signal. Statistically significant PGLS results support Hypotheses 1 and 2 (which are not mutually exclusive) as well as Hypothesis 3. No support was found for Hypothesis 4. These findings confirm that the morphology of the bovid astragalus is related to habitat‐specific locomotor ecology, and that this relationship is statistically significant after controlling for body size and phylogeny. Thus, this study validates the use of this bone as an ecomorphological indicator. J. Morphol. 275:1201–1216, 2014. © 2014 Wiley Periodicals, Inc.     

Evolving between land and water: key questions on the emergence and history of the Hippopotamidae (Hippopotamoidea,Cetancodonta, Cetartiodactyla)

The fossil record of the Hippopotamidae can shed light on three major issues in mammalian evolution. First, as the Hippopotamidae are the extant sister group of Cetacea, gaining a better understanding of the origin of the Hippopotamidae and of their Paleogene ancestors will be instrumental in clarifying phylogenetic relationships within Cetartiodactyla. Unfortunately, the data relevant to hippopotamid origins have generally been ignored in phylogenetic analyses of cetartiodactyls. In order to obtain better resolution, future analyses should consider hypotheses of hippopotamid Paleogene relationships. Notably, an emergence of the Hippopotamidae from within anthracotheriids has received growing support, leading to reconciliation between genetic and morphological evidence for the clade Cetancodonta (Hippopotamidae + Cetacea). Secondly, full account needs to be taken of the Hippopotamidae when studying the impact of environmental change on faunal evolution. This group of semi‐aquatic large herbivores has a clear and distinct ecological role and a diverse and abundant fossil record, particularly in the African Neogene. We examine three major phases of hippopotamid evolution, namely the sudden appearance of hippopotamines in the late Miocene (the “Hippopotamine Event”), the subsequent rampant endemism in African basins, and the Pleistocene expansion of Hippopotamus. Each may have been influenced by multiple factors, including: late Miocene grass expansion, African hydrographical network disruption, and a unique set of adaptations that allowed Hippopotamus to respond efficiently to early Pleistocene environmental change. Thirdly, the fossil record of the Hippopotamidae documents the independent emergence of adaptive character complexes in relation to semiaquatic habits and in response to insular isolation. The semiaquatic specializations of fossil hippopotamids are particularly useful in interpreting the functional morphology and ecology of other, extinct groups of large semiaquatic herbivores. Hippopotamids can also serve as models to elucidate the evolutionary dynamics of island mammals.     

The subfamilies and tribes of the family Bovidae

In this paper 112 skeletal characters in 27 living species of bovids are used in cladistic and phenetic analyses of the relationships among the tribes in the family. Consideration and modification of the cladistic analysis leads to the conclusion that bovids cluster around four foci in ascending evolutionary sequence: Boselaphini and allies; Antilopini and some Neotragini; the Caprinae; and a group of African antelopes containing Aepyceros , Alcelaphini, Reduncini and Hippotragini. This conclusion is quite closely compatible with the phenetic distance analysis of the same data, provided die latter is read as if primitive or early bovids share more similarities than divergently advanced ones and hence associate more closely. Given the primitiveness of Boselaphini and allies, the crucial finding is that Caprinae link with African antelopes and that Antilopini are more remote. Cladistic and phenetic analyses of 32 characters in 12 extinct bovid species produce similar groupings, but also throw doubt on the classification of Aepyceros , Reduncini and Hippotragini alongside Alcelaphini within a clade of African antelopes. As a result of these two sets of studies, of living and of extinct bovids, minimal alterations are proposed to the arrangement of bovid tribes. In addition, Saiga is placed in the Antilopini, and, with less assurance, Pelea in the Neotragini, Aepyceros in the Alcelaphinae, and Pantholops in the Caprinae. The contribution of the fossil record to understanding bovid evolution is considered.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Mammalian body size changes and Plio-Pleistocene environmental shifts: implications for understanding hominin evolution in eastern and southern Africa

This study examines geographic and temporal variation in three mammalian taxa co-occurring in eastern and southern Africa. The selected taxa-the spotted hyena (Crocuta crocuta), the plains zebra (Equus burchellii), and the impala (Aepyceros melampus)--are geographically widespread in modern times and are abundant in eastern and southern African Plio-Pleistocene fossil sites. Craniodental measurements of modern conspecifics from known geographic locations are compared using multivariate statistical methods to discern patterns of modern geographic variation within taxa. Modern and fossil samples are statistically compared to assess the nature and extent of inferred shifts in body size, both between modern samples and through time in each region. These results indicate that modern spotted hyenas and plains zebras exhibit mainly size variation between regions, with southern African samples possessing statistically larger craniodental metrics than eastern African samples. Comparison of fossil and modern samples reveals that the fossil assemblages do not show the same pattern of geographic variation. Significant temporal changes are more numerous between fossil and modern eastern African samples, and these changes are not mirrored by similar changes in the southern African samples. The changes experienced by taxa in eastern Africa appear to have been more extreme and wide-ranging than those in southern Africa, a presumed refugium. This result accords well with genetic studies of several large mammal species and paleoenvironmental studies suggesting that eastern African localized environments were more affected by tectonism and volcanism than were those in southern Africa. This study suggests that different evolutionary scenarios may have existed within Africa during the Plio-Pleistocene, but that both regions played unique and complementary roles in the evolution of African hominins and the broader faunal community.     

Dietary classification of extant kangaroos and their relatives (Marsupialia: Macropodoidea)

Kangaroos and kin (superfamily Macropodoidea) are the principal endemic herbivores of Australia and the most diverse radiation of marsupial herbivores ever to have evolved. As is typical of other herbivore groups (e.g. bovids), dietary niches span fruit, fungi, dicot leaves and monocot grasses in both specialists and generalists, but to date dietary classification has been largely ad hoc and poorly tied to actual dietary ecological data. Here we provide a simple dietary classification of the Macropodoidea based on an extensive literature survey. Intake of four major dietary items – grasses, dicot leaves, fruits and seeds, and fungi – was assessed using proportional intake for 19 species and categorical (ranked intake) data for 37 species. Statistical comparisons with cluster and principal components analyses aligned species into four dietary groups. Members of the first group have diets that primarily consist of fungi and fruits. Relative proportions of grasses to dicot leaves separate the remaining species into browser (more than 70% dicots), grazer (more than 70% grasses) and mixed feeder groups. Comparison of our diet‐based classification with a prevailing scheme based on dental morphology suggests that most species with what has traditionally been viewed as a ‘browser‐grade dentition’ are actually mixed feeders. This suggests that either morphology and diet are not tightly linked or that morphological differences between the dentitions of browsers and mixed feeders are subtle and have been overlooked. A positive correlation was found between body mass and average proportional intake of grass in the diet of macropodoids. This parallels the situation found in bovids, as well as the percentage cut‐off between dietary groups. These trends suggest that some underlying ecophysiological constraints may influence food choice in mammalian herbivores providing useful pointers to the diets of extinct taxa.     

Ecomorphological study of large canids from the lower Pleistocene of southeastern Spain

An ecomorphological analysis of the skeletal remains of large canids, Canis (Xenocyon) falconeri and Canis etruscus (Mammalia, Carnivora, Canidae), preserved in an assemblage of large mammals from the lower Pleistocene site at Venta Micena (Guadix–Baza Basin, Orce, Granada, southeastern Spain) is reported. Mean body mass of adult individuals was estimated to be around 10 kg for C. etruscus and approximately 28 kg for C. falconeri using multiple regression. A comparative study of tooth measurements in modem canids, using principal components and discriminant function analysis, infers quite different ecomorphological adaptations and feeding behavior for both fossil species. The craniodental morphology of C. falconeri is similar to that shown by extant hypercarnivorous canids whose diet include more than 70% of vertebrate meat, whereas C. etruscus shows a cranial morphology similar to those of modem omnivorous species, thus indicating a dietary niche in which vertebrate meat represented less than 70% of its diet, with other feeding resources making up the balance. These results suggest that there was a marked ecological segregation between both sympatric species of large canids. The find of a complete skull of C. falconeri showing bilateral asymmetry and marked dental anomalies could suggest high levels of genetic homozygosis in the population which inhabited this region during early Pleistocene times, possibly as a consequence of isolation and the low number of individuals. This may have subsequently led to the extinction of C. falconeri in the Western fringe of Europe. The survival of this pathological individual to adulthood indicates that this species may have developed cooperative behavior similar to that of modem African wild dogs.     

Phylogenetic relationships among the Lorisoidea as indicated by craniodental morphology and mitochondrial sequence data

The phylogeny of the Afro-Asian Lorisoidea is controversial. While postcranial data attest strongly to the monophyly of the Lorisidae, most molecular analyses portray them as paraphyletic and group the Galagidae alternately with the Asian or African lorisids. One of the problems that has bedevilled phylogenetic analysis of the group in the past is the limited number of taxa sampled for both ingroup families. We present the results of a series of phylogenetic analyses based on 635 base pairs (bp) from two mitochondrial genes (12S and 16S rRNA) with and without 36 craniodental characters, for 11 galagid and five lorisid taxa. The outgroup was the gray mouse lemur (Microcebus murinus). Analyses of the molecular data included maximum parsimony (MP), neighbor joining (NJ), maximum likelihood (ML), and Bayesian methods. The model-based analyses and the combined "molecules+morphology" analyses supported monophyly of the Lorisidae and Galagidae. The lorisids form two geographically defined clades. We find no support for the taxonomy of Galagidae as proposed recently by Groves [Primate Taxonomy, Washington, DC: Smithsonian Institution Press. 350 p, 2001]. The taxonomy of Nash et al. [International Journal of Primatology 10:57-80, 1989] is supported by the combined "molecules+morphology" analysis; however, the model-based analyses suggest that Galagoides may be an assemblage of species united by plesiomorphic craniodental characters.     

Phylogenetic patterns of trait and trait plasticity evolution: Insights from amphibian embryos

Environmental variation favors the evolution of phenotypic plasticity. For many species, we understand the costs and benefits of different phenotypes, but we lack a broad understanding of how plastic traits evolve across large clades. Using identical experiments conducted across North America, we examined prey responses to predator cues. We quantified five life‐history traits and the magnitude of their plasticity for 23 amphibian species/populations (spanning three families and five genera) when exposed to no cues, crushed‐egg cues, and predatory crayfish cues. Embryonic responses varied considerably among species and phylogenetic signal was common among the traits, whereas phylogenetic signal was rare for trait plasticities. Among trait‐evolution models, the Ornstein–Uhlenbeck (OU) model provided the best fit or was essentially tied with Brownian motion. Using the best fitting model, evolutionary rates for plasticities were higher than traits for three life‐history traits and lower for two. These data suggest that the evolution of life‐history traits in amphibian embryos is more constrained by a species’ position in the phylogeny than is the evolution of life history plasticities. The fact that an OU model of trait evolution was often a good fit to patterns of trait variation may indicate adaptive optima for traits and their plasticities.