Correlates between calcaneal morphology and locomotion in extant and extinct carnivorous mammals

Locomotor mode is an important component of an animal's ecology, relating to both habitat and substrate choice (e.g., arboreal versus terrestrial) and in the case of carnivores, to mode of predation (e.g., ambush versus pursuit). Here, we examine how the morphology of the calcaneum, the ‘heel bone’ in the tarsus, correlates with locomotion in extant carnivores. Other studies have confirmed the correlation of calcaneal morphology with locomotion behaviour and habitat. The robust nature of the calcaneum means that it is frequently preserved in the fossil record. Here, we employ linear measurements and 2D‐geometric morphometrics on a sample of calcanea from eighty‐seven extant carnivorans and demonstrate a signal of correlation between calcaneal morphology and locomotor mode that overrides phylogeny. We used this correlation to determine the locomotor mode, and hence aspects of the palaeobiology of, 47 extinct carnivorous mammal taxa, including both Carnivora and Creodonta. We found ursids (bears), clustered together, separate from the other carnivorans. Our results support greater locomotor diversity for nimravids (the extinct ‘false sabertooths’, usually considered to be more arboreal), than previously expected. However, there are limitations to interpretation of extinct taxa because their robust morphology is not fully captured in the range of modern carnivoran morphology.     

Costs of carnivory: tooth fracture in Pleistocene and Recent carnivorans

Large, carnivorous mammals often break their teeth, probably as a result of tooth to bone contact that occurs when carcasses are consumed more fully, a behaviour likely to occur under conditions of food stress. Recent studies of Pleistocene predators revealed high numbers of teeth broken in life, suggesting that carcass utilization and, consequently, food competition was more intense in the past than at present. However, the putative association between diet and tooth fracture frequency was based on a small sample of large, highly carnivorous species. In the present study, a greater diversity of extant carnivorans is sampled, including insectivorous, omnivorous, and carnivorous forms, ranging in size from weasels to tigers. Species that habitually consume hard foods (bones, shells) had the highest fracture frequencies, followed by carnivores, and then insectivorous and/or omnivorous species. Predator and prey sizes were not associated with tooth fracture frequency, but more aggressive species did break their teeth more often. Comparison of the modern sample with five Pleistocene species confirms the previous finding of higher tooth breakage in the past, although some extant species have fracture frequencies that approach those of extinct species. Thus, the Pleistocene predator guild appears to have been characterized by relatively high levels of competition that are rarely observed today.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 68–81.     

Postcranial element shape and function: assessing locomotor mode in extant and extinct mustelid carnivorans

Assessment of locomotor modes in fossil taxa must often be made on the basis of heavily fragmented postcranial material. Previous authors have used quantitative methods to determine locomotor function from whole postcranial elements. The goals of this project were to assess the ability of element shape to discern between locomotor modes through landmark analysis, and to apply the results to assessment of fossils. Results suggest that element shape is a good predictor of function, but that different elements have different predictive capacities for each locomotor mode. Additionally, a relationship between size and shape exists that appears to drive morphological differentiation in the group. Finally, data from the extant sample were applied to fossil material of the extinct Plio-Pleistocene taxon Trigonictis . The results suggest that the locomotor mode of Trigonictis was generalized and probably an intermediate between the half-bound locomotion found in weasels and ferrets and the scansorial locomotion of martens and fishers.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 895–914.     

The evolution of encephalization in caniform carnivorans

A weighted-average model, which reliably estimates endocranial volume from three external measurements of the neurocranium of extant taxa in the mammalian order Carnivora, was tested for its applicability to fossil taxa by comparing model-estimated endocranial volumes to known endocast volumes. The model accurately reproduces endocast volumes for a wide array of fossil taxa across the crown radiation of the Carnivora, three stem carnivoramorphan taxa, and Pleistocene fossils of two extant species. Applying this model to fossil taxa without known endocast volumes expanded the sample of fossil taxa with estimated brain volumes in the carnivoran suborder Caniformia from 11 to 60 taxa. This then allowed a comprehensive assessment of the evolution of relative brain size across this clade. An allometry of brain volume to body mass was calculated on phylogenetically independent contrasts for the set of extant taxa, and from this, log-transformed encephalization quotients (logEQs) were calculated for all taxa, extant, and fossil. A series of Mann-Whitney tests demonstrated that the distributions of logEQs for taxa early in caniform evolutionary history possessed significantly lower median logEQs than extant taxa. Median logEQ showed a pronounced shift around the Miocene-Pliocene transition. Support tests, based on likelihood ratios, demonstrated that the variances of these distributions also were significantly lower than among modern taxa, but logEQ variance increased gradually through the history of the clade, not abruptly. Reconstructions of ancestral logEQs using weighted squared-change parsimony demonstrate that increased encephalization is observed across all major caniform clades (with the possible exception of skunks) and that these increases were achieved in parallel, although an "ancestor-descendant differencing" method could not rule out drift as a hypothesis. Peculiarities in the estimated logEQs for the extinct caniform family Amphicyonidae were also investigated; these unusual patterns are likely due to a unique allometry in scaling brain to body size in this single clade.     

Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.     

Getting a grip on the evolution of grasping in musteloid carnivorans: a three‐dimensional analysis of forelimb shape

The ability to grasp and manipulate is often considered a hallmark of hominins and associated with the evolution of their bipedal locomotion and tool use. Yet, many other mammals use their forelimbs to grasp and manipulate objects. Previous investigations have suggested that grasping may be derived from digging behaviour, arboreal locomotion or hunting behaviour. Here, we test the arboreal origin of grasping and investigate whether an arboreal lifestyle could confer a greater grasping ability in musteloid carnivorans. Moreover, we investigate the morphological adaptations related to grasping and the differences between arboreal species with different grasping abilities. We predict that if grasping is derived from an arboreal lifestyle, then the anatomical specializations of the forelimb for arboreality must be similar to those involved in grasping. We further predict that arboreal species with a well‐developed manipulation ability will have articulations that facilitate radio‐ulnar rotation. We use ancestral character state reconstructions of lifestyle and grasping ability to understand the evolution of both traits. Finally, we use a surface sliding semi‐landmark approach capable of quantifying the articulations in their full complexity. Our results largely confirm our predictions, demonstrating that musteloids with greater grasping skills differ markedly from others in the shape of their forelimb bones. These analyses further suggest that the evolution of an arboreal lifestyle likely preceded the development of enhanced grasping ability.     

Postcranial morphology and the locomotor habits of living and extinct carnivorans

Members of the order Carnivora display a broad range of locomotor habits, including cursorial, scansorial, arboreal, semiaquatic, aquatic, and semifossorial species from multiple families. Ecomorphological analyses from osteological measurements have been used successfully in prior studies of carnivorans and rodents to accurately infer the locomotor habits of extinct species. This study uses 20 postcranial measurements that have been shown to be effective indicators of locomotor habits in rodents and incorporates an extensive sample of over 300 individuals from more than 100 living carnivoran species. We performed statistical analyses, including analysis of variance (ANOVA) and stepwise discriminant function analysis, using a set of 16 functional indices (ratios). Our ANOVA results reveal consistent differences in postcranial skeletal morphology among locomotor groups. Cursorial species display distal elongation of the limbs, gracile limb elements, and relatively narrow humeral and femoral epicondyles. Aquatic and semiaquatic species display relatively robust, shortened femora and elongate metatarsals. Semifossorial species display relatively short, robust limbs with enlarged muscular attachment sites and elongate claws. Both semiaquatic and semifossorial species have relatively elongate olecranon process of the ulna and enlarged humeral and femoral epicondyles. Terrestrial, scansorial, and arboreal species are characterized by having primarily intermediate features, but arboreal species do show relatively elongate manual digits. Morphological indices effectively discriminate locomotor groups, with cursorial and arboreal species more accurately classified than terrestrial, scansorial, or semiaquatic species. Both within and between families, species with similar locomotor habits converge toward similar postcranial morphology despite their independent evolutionary histories. The discriminant analysis worked particularly well to correctly classify members of the Canidae, but not as well for members of the Mustelidae or Ursidae. Results are used to infer the locomotor habits of extinct carnivorans, including members of several extinct families, and also 12 species from the Pleistocene of Rancho La Brea. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

Fruit consumption by carnivores in Mediterranean Europe

• 1 We reviewed patterns of fruit consumption amongst 10 species of mesocarnivores: red fox Vulpes vulpes, weasel Mustela nivalis, stoat Mustela erminea, polecat Mustela putorius, stone marten Martes foina, pine marten Martes martes, Eurasian badger Meles meles, common genet Genetta genetta, Egyptian mongoose Herpestes ichneumon and wildcat Felis silvestris in Mediterranean Europe.
• 2 The 65 reviewed studies recorded 79 different fruits eaten by carnivores, 58 of which were identified to species. Most records (63%) were of fleshy fruits with high pulp content. The frequency of occurrence of fruit items varied widely amongst species and regions. Four of the carnivore species (red fox, stone marten, badger and common genet) included more than 30 fruit species in their diet.
• 3 A longitudinal pattern was detected in the consumption of fruit in the Mediterranean region, with the frequency of occurrence of fruit consumption increasing towards the east.

     

Wing shape variation in the medium ground finch (Geospiza fortis): an ecomorphological approach

Wing design in birds is subject to a suite of interacting selective pressures. As different performance traits are favoured in different ecological settings, a tight link is generally expected between variation in wing morphology and variation in ecological parameters. In the present study, we document aspects of variation in wing morphology in the medium ground finch ( Geospiza fortis ) on Isla Santa Cruz in the Galápagos. We compare variation in body size, simple morphometric traits (body mass, last primary length, wing length, wing chord, and wing area) and functional traits (wing loading, aspect ratio and wing pointedness) across years, among populations, and between sexes. Functional traits are found to covary across years with differences in climatic conditions, and to covary among populations with differences in habitat structure. In dry years and arid locations, wing aspect ratios are highest and wings are more pointed, consistent with a need for a low cost of transport. In wet years and cluttered habitats, wing loading is lowest and wings are more rounded, suggesting enhanced capabilities for manoeuvrability. Sexes differ in wing loading, with males having lower wing loadings than females. Superior manoeverability might be favoured in males for efficient territory maintenance. Lastly, in contrast to functional traits, we found little consistent inter-annual or inter-site variation in simple morphometric traits.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 129–138.     

Differences in mandibular disparity between extant and extinct species of metatherian and placental carnivore clades

There are a number of studies relating to skull morphology differences within the carnivoran clades of both placentals and metatherians. It is difficult to compare these studies because of differences in taxonomic sampling, for example some include fossil taxa while others include non‐carnivoran placentals. As a consequence, we studied mandible morphology in a broad range of both extant and extinct carnivorous species, including Carnivora, Marsupialia and Sparassodonta to test for differences between these clades. We used geometric morphometrics and two disparity indexes, the variance and Procrustes distances mean. When including fossil species, we found no significant differences for both disparities in some analyses, except after the exclusion of the sabretooth morphotype. This can be explained by the extreme morphology of this morphotype, which increases the variance and reduces the disparity effect of the other species in the analyses. Using Procrustes distances, we found significant differences in disparity distances between Carnivora and Metatheria for most of the analyses. We also found significant differences using the variance index in some analyses. The mandibular disparity in Carnivora is greater than in carnivorous metatherian mammals for most of the cases and this can be related with differences in evolutionary history and constraints of both groups. The pattern found in the mandible is similar to that found in the face of the skull but was not observed in the braincase, due to differences in skull function and mandible function.     

The structure of the mandibular corpus and its relationship to feeding behaviours in extant carnivorans

Assuming some optimization of bone structure to applied mechanical loadings in vivo , different killing and feeding behaviours in carnivores should be reflected in observed differences in cross-sectional shape of their mandibular corpora. Section moduli are used to gauge the magnitudes of bending moments in the mandibular corpus and, when dentary length is controlled, the magnitudes of forces applied to the corpus. Comparisons are made of section moduli at the P₃P₄ and P₄M₁ interdental gaps among canids, felids and hyaenids; in canids only, the M₁M₂ interdental gap was also studied. Local variations in loadings are identified by comparing the section moduli at adjacent loci along the corpus within each family.
The findings of this study show that the precarnassial corpora of canids and hyaenids have greater strength in bending than the corpora of felids of similar body weight. This is taken to reflect relatively greater bending moments under loading in the corpora of canids and hyaenids due, in part, to their elongate dentaries (relative to body weight). Relative to dentary length , however, the precarnassial corpora of felids and hyaenids have much greater strength in bending than the corpora of canids. These scaling relationships appear to reflect the high customary forces (i.e. not moments) applied to the precarnassial corpora of felids and hyaenids with sustained canine killing bites and with bone ingestion using the premolars, respectively. An increase in bending strength of the corpus caudal to the camassial blade in canids is interpreted to be an adaptation for bone-crushing with the postcarnassial molars.     

Deep ecomorphological and genetic divergence in Steller's Jays (Cyanocitta stelleri,Aves: Corvidae)

The relationship between ecology and morphology is a cornerstone of evolutionary biology, and quantifying variation across environments can shed light on processes that give rise to biodiversity. Three morphotypes of the Steller''s Jay (Cyanocitta stelleri) occupy different ecoregions in western North America, which vary in climate and landcover. These morphotypes (Coastal, Interior, Rocky Mountain) differ in size, plumage coloration, and head pattern. We sampled 1080 Steller''s Jays from 68 populations (plus 11 outgroups) to address three main questions using data on morphology, plumage, genetics (mtDNA, microsatellites), and ecological niches: (1) How do phenotypic and genetic traits vary within and among populations, morphotypes, and ecoregions? (2) How do population‐level differences in Steller''s Jays compare with other sister species pairs of North American birds? (3) What can we infer about the population history of Steller''s Jays in relation to past climates, paleoecology, and niche evolution? We found substantial morphological, genetic, and ecological differentiation among morphotypes. The greatest genetic divergence separated Coastal and Interior morphotypes from the Rocky Mountain morphotype, which was associated with warmer, drier, and more open habitats. Microsatellites revealed additional structure between Coastal and Interior groups. The deep mtDNA split between Coastal/Interior and Rocky Mountain lineages of Steller''s Jay (ND2 ~ 7.8%) is older than most North American avian sister species and dates to approximately 4.3 mya. Interior and Rocky Mountain morphotypes contact across a narrow zone with steep clines in traits and reduced gene flow. The distribution of the three morphotypes coincides with divergent varieties of ponderosa pine and Douglas fir. Species distribution models support multiple glacial refugia for Steller''s Jays. Our integrative dataset combined with extensive geographic sampling provides compelling evidence for recognizing at least two species of Steller''s Jay.     

Changing environmental gradients over forty years alter ecomorphological variation in Guadalupe Bass Micropterus treculii throughout a river basin

Understanding the degree of intraspecific variation within and among populations is a key aspect of predicting the capacity of a species to respond to anthropogenic disturbances. However, intraspecific variation is usually assessed at either limited temporal, but broad spatial scales or vice versa, which can make assessing changes in response to long‐term disturbances challenging. We evaluated the relationship between the longitudinal gradient of changing flow regimes and land use/land cover patterns since 1980 and morphological variation of Guadalupe Bass Micropterus treculii throughout the Colorado River Basin of central Texas. The Colorado River Basin in Texas has experienced major alterations to the hydrologic regime due to changing land‐ and water‐use patterns. Historical collections of Guadalupe Bass prior to rapid human‐induced change present the unique opportunity to study the response of populations to varying environmental conditions through space and time. Morphological differentiation of Guadalupe Bass associated with temporal changes in flow regimes and land use/land cover patterns suggests that they are exhibiting intraspecific trait variability, with contemporary individuals showing increased body depth, in response to environmental alteration through time (specifically related to an increase in herbaceous land cover, maximum flows, and the number of low pulses and high pulses). Additionally, individuals from tributaries with increased hydrologic alteration associated with urbanization or agricultural withdrawals tended to have a greater distance between the anal and caudal fin. These results reveal trait variation that may help to buffer populations under conditions of increased urbanization and sprawl, human population growth, and climate risk, all of which impose novel selective pressures, especially on endemic species like Guadalupe Bass. Our results contribute an understanding of the adaptability and capacity of an endemic population to respond to expected future changes based on demographic or climatic projection.     

Postcanine dental form in the mustelidae and viverridae (Carnivora: Mammalia)

This study investigates whether the gross morphology of mustelid and viverrid postcanine dentitions corresponds with differences in diet. For each species, the predominant foods ingested are used to form predictions of dental form and measurements of the carnassial and molar teeth determine the extent of shearing and crushing surfaces on the postcanine teeth. Principal components analysis distinguishes species according to morphological differences in the dentition and these differences are compared with predictions of dental form based on diet. Dietarily specialized species are more likely to be correspondingly specialized in the dentition and species with varied food sources are more likely to possess dental characteristics that are generalized in function. Consumers of foods with high fracture resistance, such as vertebrate tissue and hard-surfaced invertebrates, possess specialized shearing or crushing postcanine teeth. On the other hand, species that consume foods of lesser fracture resistance, such as fruit and soft invertebrates, differ greatly in dental form and are more generalized in dental function. A few species possess postcanine dentitions that do not correspond with diet; the absence of dental-dietary correlation in these species suggests that other factors, such as phylogeny, are important in determining dental form.     

Ecological consequences of ontogenetic changes in head shape and bite performance in the Jamaican lizard Anolis lineatopus

It has been documented extensively that body size affects the physiology and musculoskeletal function of organisms. However, less well understood is how body size affects the ecology of organisms through its effects on physiology and performance. We explored the effects of body size on morphology and performance in different ontogenetic classes and sexes of a common Anolis lizard ( A. lineatopus ). Next, we tested whether these morphological and performance differences may affect functional aspects of the diet such as prey size and prey hardness. Our data showed that males, females and juveniles differ significantly in head size, head shape and bite force. Multiple regression models indicated that head shape and bite force are significantly correlated to prey size and hardness. Yet juveniles had relatively large heads and bit disproportionately hard for their size, allowing them to eat prey as large as those of females. However, for a given prey size, males and females ate more robust prey than did juveniles. Additionally, males ate relatively harder prey than did juveniles. These data suggest that: (1) body size affects the dietary ecology of animals through its effect on head size and bite force; (2) changes in head morphology independent of changes in overall size also have important effects on performance and diet.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 443–454.