Canine tooth strength and killing behaviour in large carnivores

Bending strength of upper canine teeth is examined among living canids, felids, hyaenids and several extinct species including sabretooth cats, borophagine dogs and the dire wolf, Canis dirus. The tooth is modelled as a cantilever with an elliptical cross-section. Using beam theory, the bending strength of the upper canine is calculated given a constant force applied to the canine tip. Results indicate that felids and hyaenids have relatively stronger canines than canids, particularly in bending about the anteroposterior (AP) rather than the mediolateral axis. It is suggested that canine shape reflects the forces produced during killing and feeding. As shown by an analysis of jaw muscle moment arms, felids and hyaenids have relatively stronger bites than canids. Moreover, the canines of hyaenids and felids are perhaps more likely to contact bone during feeding and killing and consequently may be subjected to larger and more frequent bending moments about the AP axis. The canines of sabretooth cats are shown to be more similar in shape and strength characteristics to those of living canids than felids, whereas those of the borophagine dogs and the dire wolf are closer to modern hyaenas.     

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.     

Locomotor correlates of the scapholunar of living and extinct carnivorans

The relationship of carpal morphology to ecology and habitat is under studied in carnivorans and more generally in mammals. Here, we use 3D-scanning techniques to assess the usefulness of a carpal bone, the scapholunar, in carnivorans to reflect ecology and habitat, and to reconstruct the ecology of five extinct carnivorans from two fossil sites: Rancho La Brea and Natural Trap Cave. We 3D-scanned scapholunars and measured articular surface areas and angles between articular facets using GeoMagic and Rhino 3D-software. We analyzed the difference in these metrics using multivariate analysis of variance and discriminant function analysis. Results show that the scapholunar reflects ecological signal, with clear groupings of cursorial carnivorans and grappling/climbing carnivorans; however, phylogenetic signal was also present in the results with hyaenids, canids, and large felids in distinct morphospaces. Extinct species Miracinonyx trumani (American cheetah) and Smilodon fatalis (sabertooth cat) showed surprising results with M. trumani grouping with pantherines instead of Acinonyx or Puma, suggesting it runs but still retains the ability to grapple prey. S. fatalis groups with pantherines, but also shows some unique adaptations, suggesting it had a different range of wrist motion than living cats. Overall, the scapholunar is a good indicator of ecology and functional morphology and can be another tool to use in modern and fossil carnivorans to reconstruct extinct ecologies and locomotor behaviors.     

Mid-Pliocene Carnivora from the Woranso-Mille Area,Afar Region,Ethiopia

The Woranso-Mille paleontological study area, located in the central Afar region of Ethiopia, is one of the most important Pliocene sites in eastern Africa. Since the Woranso-Mille Research Project began its investigation in 2005, more than 7,500 mammalian fossils, including hominins, have been collected from 80 vertebrate localities. This paper provides a preliminary assessment of the Woranso-Mille carnivoran record, a record that is of great interest given the high level of species richness of African carnivorans during the middle Pliocene. Craniodental and postcranial material of canids, lutrine mustelids, viverrids, herpestids, machairodontine and feline felids, and hyaenids has been recovered. Thus, this diverse fauna includes not only the largest carnivorans from this time period (e.g., Enhydriodon and Homotherium), but also some of the smallest, including mongooses, civets, genets, and felids, some of which represent new species. However, the diversity of small taxa does not yet approach that found in the roughly contemporaneous Upper Laetolil Beds of Tanzania. In contrast, lutrine mustelids are better represented at Woranso-Mille than at Kanapoi (Kenya) or Laetoli (Tanzania), which is to be expected given the diversity of habitats represented at these sites. While more material from these sites and others are necessary to truly understand the increased diversity within the early to middle Pliocene eastern African carnivoran guild, it is clear that the material from Woranso-Mille has the potential to fill many of the gaps in our knowledge of carnivorans during this time period.     

Auditory ossicles from southwest Asian Mousterian sites

The present study describes and analyzes new Neandertal and early modern human auditory ossicles from the sites of Qafzeh and Amud in southwest Asia. Some methodological issues in the measurement of these bones are considered, and a set of standardized measurement protocols is proposed. Evidence of erosive pathological processes, most likely attributed to otitis media, is present on the ossicles of Qafzeh 12 and Amud 7 but none can be detected in the other Qafzeh specimens. Qafzeh 12 and 15 extend the known range of variation in the fossil H. sapiens sample in some metric variables, but morphologically, the new specimens do not differ in any meaningful way from living humans. In most metric dimensions, the Amud 7 incus falls within our modern human range of variation, but the more closed angle between the short and long processes stands out. Morphologically, all the Neandertal incudi described to date show a very straight long process. Several tentative hypotheses can be suggested regarding the evolution of the ear ossicles in the genus Homo. First, the degree of metric and morphological variation seems greater among the fossil H. sapiens sample than in Neandertals. Second, there is a real difference in the size of the malleus between Neandertals and fossil H. sapiens, with Neandertals showing larger values in most dimensions. Third, the wider malleus head implies a larger articular facet in the Neandertals, and this also appears to be reflected in the larger (taller) incus articular facet. Fourth, there is limited evidence for a potential temporal trend toward reduction of the long process within the Neandertal lineage. Fifth, a combination of features in the malleus, incus, and stapes may indicate a slightly different relative positioning of either the tip of the incus long process or stapes footplate within the tympanic cavity in the Neandertal lineage.     

Comparisons of canid and felid social systems from an evolutionary perspective

Canids and felids are compared regarding evolutionary history, distribution, habitat preferences, morphology and behaviour. Factors permitting and promoting sociality in both families are discussed, and communication mechanisms within each sensory modality compared. A solitary existence is compatible with felid specializations, the mother and young being the basic social unit. Canid specializations have permitted the development of a pair bond and male provisioning of young. As an adaptation to hunting large herbivorous prey, an increase in size occurred in the felids, but canids developed sophisticated pack-hunting techniques. Group structure in canids is based on long-term affiliations between a pair and matured offspring, while in the lion, the only truly social felid, it is based on a mother and maturing daughters.     

Evolution of community composition in several carnivore palaeoguilds from the European Pleistocene: the role of interspecific competition

This study focuses on ecological processes such as competition or predation from an evolutionary perspective. First, we attempt to test the idea that species with similar feeding requirements tend to coexist by separating morphologically or behaviourally. Then, the Barton–David test was applied to several carnivore communities (felids and canids) separated in time. Although the preservation bias of the fossil record renders our conclusions tentative, the general equal size–ratio pattern in most of the guilds examined indicates that inter-specific competition for prey species seems to be a good candidate to explain the evolution of guild composition and morphological traits throughout the Pleistocene for the two groups considered, felids and canids.     

Comparative Scaling of Humeral Cross-Sections of Felids and Canids Using Radiographic Images

The cortical thickness of long bones can be an effective indicator of locomotor modes and other stresses encountered by bone. Felids and canids are two carnivoran families that have similar levels of phylogenetic diversity and overlap in body size, but differ in their locomotor habits. Many canids and felids are cursorial, but felids also climb more frequently than canids. Felids also display a secondary use for their forelimbs not observed in any canids: they use their forelimbs to grasp and subdue prey. Large felids use their forelimbs much more extensively to subdue prey than do large canids and, therefore, should have proportionately greater forces applied to their forelimbs. This study uses a non-invasive radiographic approach to examine the differences in cortical thickness in the humerus between the Felidae and Canidae, as well as between size groups within these two families. Results show few significant differences between the two families, with a slight trend toward more positive allometry in the felids. Overall, radiographic measurements were found to be better predictors of body mass than either prey killing behavior or locomotor mode in these two carnivoran families. One canid that demonstrated exceptionally high cortical area was the bush dog, Speothos venaticus. The rarely observed bush dog has been postulated to swim and dig regularly, and it may be that the thickened cortical bone reflects these behaviors.     

A phylogenetic hypothesis for species of the genus Taenia (Eucestoda : Taeniidae)

Cladistic analysis of a numerical data matrix describing 27 characters for species of Taenia resulted in 4 most parsimonious phylogenetic trees (174 steps; consistency index = 0.28; homoplasy index = 0.72; retention index = 0.48). Monophyly for Taenia is diagnosed by the metacestode that is either a cysticercus or a form derived from a bladder-like larva; no other unequivocal synapomorphies are evident. Tree structure provides no support for recognition of a diversity of tribes or genera within the Taeniinae: Fimbriotaeniini and Taeniini have no phylogenetic basis. Hydatigera, Fimbriotaenia, Fossor, Monordotaenia, Multiceps, Taeniarhynchus, Tetratirotaenia must be subsumed within Taenia as synonyms. Taenia saginata and Taenia asiatica are sister species and distantly related to Taenia solium. Cospeciation with respect to carnivorous definitive hosts and Taenia appears to be limited. Although felids are putative ancestral hosts, contemporary associations appear to have resulted from extensive host-switching among felids, canids, hyaenids, and others. In contrast, relationships with herbivorous intermediate hosts are indicative of more pervasive coevolution; rodents as intermediate hosts are postulated as ancestral for the Taeniidae, Taenia + Echinococcus. Patterns appear consistent with rapid shifts between phylogenetically unrelated carnivores but among those that historically exploited a common prey resource within communities in specific biogeographic regions.     

Evolution of skull shape in carnivores 1. Representative modern carnivores

Fifteen variables, selected primarily to reflect functionally significant aspects of cranial morphology, were measured on one skull each of 62 species of modern carnivores, including viverrids, canids, mustelids and felids. To allow comparisons between species of different sizes without the potentially confounding effects of allometric shape changes, the measurements were transformed to dimensionless variables, based on the residuals from allometric equations. Fourteen out of 15 of the transformed variables distinguish one or more of the four family groups and the rotated first two axes of a principal components analysis distinguish all four families from each other. The following functional hypotheses are proposed: mustelids and felids have the most powerful bites and canids the weakest among the four family groups studied; mustelids and, to a lesser degree, felids have more powerful neck musculature than do canids and viverrids; and visual abilities are best developed among felids and least developed among mustelids. The first two functional hypotheses suggest possible differences in killing behaviour, which are supported by a preliminary survey of the literature on such behaviour. Allometric analysis of the 15 cranial measures shows that the neurocranial components scale with negative allometry, while most of the other measures scale approximately isometrically.     

Development of a model based method for investigating facet articulation

Reported investigations of facet articulation in the human spine have often been conducted through the insertion of pressure sensitive film into the joint space, which requires incision of the facet capsule and may alter the characteristics of interaction between the facet surfaces. Load transmission through the facet has also been measured using strain gauges bonded to the articular processes. While this method allows for preservation of the facet capsule, it requires extensive instrumentation of the spine, as well as strain-gauge calibration, and is highly sensitive to placement and location of the strain gauges. The inherently invasive nature of these techniques makes it difficult to translate them into medical practice. A method has been developed to investigate facet articulation through the application of test kinematics to a specimen-specific rigid-body model of each vertebra within a lumbar spine segment. Rigid-body models of each vertebral body were developed from CT scans of each specimen. The distances between nearest-neighboring points on each facet surface were calculated for specific time frames of each specimen's flexion/extension test. A metric describing the proportion of each facet surface within a distance (2 mm) from the neighboring surface, the contact area ratio (CAR), was calculated at each of these time frames. A statistically significant difference (p<0.037) was found in the CAR between the time frames corresponding to full flexion and full extension in every level of the lumbar spine (L1-L5) using the data obtained from the seven specimens evaluated in this study. The finding that the contact area of the facet is greater in extension than flexion corresponds to other findings in the literature, as well as the generally accepted role of the facets in extension. Thus, a biomechanical method with a sufficiently sensitive metric is presented as a means to evaluate differences in facet articulation between intact and treated or between healthy and pathologic spines.     

In the Pursuit of the Predatory Behavior of Borophagines (Mammalia,Carnivora, Canidae): Inferences from Forelimb Morphology

Here, we perform an ecomorphological study on the major bones (humerus, radius, and ulna) of the carnivoran forelimb using three-dimensional geometric morphometrics. More specifically, we test the association between forelimb morphology and predatory behavior. Our results suggest that the main morphological adaptions of carnivorans to different predatory behaviors relate to: (i) the capacity to perform long and efficient runs as in pounce/pursuit and pursuit predators; (ii) the ability to maneuver as in occasional predators; and (iii) the capacity to exert and resist large loads as in ambushing predators. We used borophagine canids as a case study, given the controversy on the predatory behavior of this extinct subfamily. Our results indicate that borophagines displayed a limited set of adaptions towards efficient running, including reduced joint mobility in both the elbow and the wrist, aspects in which they resemble the living canids. Furthermore, they had forelimbs as powerful as those of the extant ambushing carnivorans (i.e., most felids). This combination of traits suggests that the predatory behavior of borophagines was unique among carnivorans, as it was not fully equivalent to any of the living species.     

Carnivora from the Plio-Pleistocene hominin site of Drimolen, Gauteng, South Africa

Drimolen is one of the newest and most productive hominin sites in South Africa, and is dated on faunal grounds between 2.0 Ma to 1.5 Ma. This paper provides the first overview of the Carnivora from Drimolen, updating the previously published preliminary faunal list, and describing all currently prepared craniodental and postcranial material. The Drimolen specimens are described in comparison with other modern and fossil South African carnivore material. The carnivores cover a range of taxa including hyaenids, felids, canids and herpestids. Most notable amongst these are the sabretooth Dinofelis aff. piveteaui craniodental and postcranial remains, which are described in detail, and a Chasmaporthetes nitidula cranium. The genus Chasmaporthetes is found at three other sites in the area - Sterkfontein, Swartkrans and Coopers D. There are two models for the geographic origin of Dinofelis piveteaui, in that it may have arisen in either eastern or southern Africa. These possibilities are discussed in the light of the new South African Dinofelis material, as the Drimolen material appears to represent a more primitive form with affinities with D. piveteaui. Fossil leopard material from Kromdraai B and Drimolen is also discussed, as the metapodia assigned to P. pardus from these two sites are very small, but lie within the variation of modern leopards. Such size differences in fossil postcrania may have implications for the niches that these animals may have occupied in the past.     

Antibodies to Toxoplasma gondii and Neospora caninum in captive neotropical and exotic wild canids and felids

This study was designed to detect antibodies to Toxoplasma gondii and Neospora caninum in wild captive carnivores maintained in Brazilian zoos. Blood samples were collected from 142 Brazilian wild felids and 19 exotic felids in zoos, and 3 European wolves (Canis lupus) and 94 Brazilian wild canids maintained in captivity in Brazilian zoos of S?o Paulo, Mato Grosso states and Federal District. One hundred and two (63.4%) and 70 (50.3%) of the 161 wild felids tested were seropositive for T. gondii and N. caninum by indirect immunofluorescent assay test (IFAT), respectively. Among sampled wild canids, 49 (50.5%) and 40 (41.2%) animals were seropositive for T. gondii and N. caninum antigens by IFAT, respectively. To our knowledge, this is the first serological detection of antibodies to N. caninum in Brazilian wild captive felids and bush dogs (Speothos venaticus (Lund)).     

Building large trees by combining phylogenetic information: a complete phylogeny of the extant Carnivora (Mammalia)

One way to build larger, more comprehensive phylogenies is to combine the vast amount of phylogenetic information already available. We review the two main strategies for accomplishing this (combining raw data versus combining trees), but employ a relatively new variant of the latter: supertree construction. The utility of one supertree technique, matrix representation using parsimony analysis (MRP), is demonstrated by deriving a complete phylogeny for all 271 extant species of the Carnivora from 177 literature sources. Beyond providing a 'consensus' estimate of carnivore phylogeny, the tree also indicates taxa for which the relationships remain controversial (e.g. the red panda; within canids, felids, and hyaenids) or have not been studied in any great detail (e.g. herpestids, viverrids, and intrageneric relationships in the procyonids). Times of divergence throughout the tree were also estimated from 74 literature sources based on both fossil and molecular data. We use the phylogeny to show that some lineages within the Mustelinae and Canidae contain significantly more species than expected for their age, illustrating the tree's utility for studies of macroevolution. It will also provide a useful foundation for comparative and conservational studies involving the carnivores.     

Chromosomal localization of satellite DNA sequences among 22 species of felids and canids (Carnivora)

In situ hybridization was carried out using cloned satellite DNAs from the domestic cat and domestic dog as probes to metaphase chromosomes from 12 species of felids and 10 species of canids. Autoradiographic silver grains along metaphase chromosomes were counted and analyzed with regard to the mean number of grains per cell in each species, their chromosomal location, and their presence or absence on specific autosomes or sex chromosomes, where known. Among the felids and canids there was a 7.6- and 8.9-fold statistically significant difference, respectively, in the mean number of grains per cell between the species having the minimum and maximum values. Among the felids, most grains occurred on the telomeres of D- and E-group chromosomes, although departures from this general pattern also occurred. For example, the Asian golden cat and the Bornean bay cat showed substantial labeling at the centromeric region of chromosome A1, and a number of species showed some labeling at the short-arm telomeres of B-group chromosomes. Among the canids, about 90% of all grains were located at autosomal centromeres, and grains were absent from the sex chromosomes. Grains are usually distributed at chromosomal locations that stain C-band positive; however, certain C-band-positive regions without grains probably do not contain the particular satellites studied here.     

Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores

Among the hunting strategies employed by members of the order Carnivora (Mammalia), two, stalk and ambush and sustained pursuit, are particularly prevalent among larger species of the order. It has been difficult to identify morphological traits that support this distinction and ecological observations have shown that most carnivorans adopt a continuum of strategies, depending on available habitat and prey. In this paper, the shape of the distal humerus articulation is analysed, with the aim of exploring the use of the forelimb in prey procurement, and as a guide to such behaviour among fossil carnivorans. The results suggest that manual manipulation and locomotion are conflicting functions. Elbow‐joint morphology supports a division between grapplers (i.e. ambushers) and nongrapplers (i.e. pursuers). Joints of the former are characterized by being relatively wide and the latter, by being relatively narrow and box‐like with pronounced stabilizing features. At intermediate and large body sizes, carnivorans show a pattern suggesting mutually exclusive feeding strategies that involve either grappling with prey or sustained pursuit. The former allows for large body sizes, such as pantherine felids and ursids; the latter includes species of only moderate size, such as hyenids and canids. Elbow‐joint morphology is closely linked to phylogeny, but the morphology of the cheetah converges with that of nongrapplers, showing that strong selective forces may override the phylogenetic component. Two taxa of giant mustelids from the Miocene were analysed to test whether this sort of analysis is applicable to carnivorans of the past. The African Late Miocene species Ekorus ekakeran has a joint morphology comparable to that of modern‐day nongrapplers. Two joint morphologies were found in the North American Late Oligocene‐Early Miocene Megalictis ferox. The first morphology is comparable to that of modern pantherine cats and the second forms an intermediate between grapplers and nongrapplers that is not present in the modern carnivoran fauna. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 91–104.     

Comparing Linkage Designs Based on Land Facets to Linkage Designs Based on Focal Species

Least-cost modeling for focal species is the most widely used method for designing conservation corridors and linkages. However, these designs depend on today''s land covers, which will be altered by climate change. We recently proposed an alternative approach based on land facets (recurring landscape units of relatively uniform topography and soils). The rationale is that corridors with high continuity of individual land facets will facilitate movement of species associated with each facet today and in the future. Conservation practitioners might like to know whether a linkage design based on land facets is likely to provide continuity of modeled breeding habitat for species needing connectivity today, and whether a linkage for focal species provides continuity and interspersion of land facets. To address these questions, we compared linkages designed for focal species and land facets in three landscapes in Arizona, USA. We used two variables to measure linkage utility, namely distances between patches of modeled breeding habitat for 5–16 focal species in each linkage, and resistance profiles for focal species and land facets between patches connected by the linkage. Compared to focal species designs, linkage designs based on land facets provided as much or more modeled habitat connectivity for 25 of 28 species-landscape combinations, failing only for the three species with the most narrowly distributed habitat. Compared to land facets designs, focal species linkages provided lower connectivity for about half the land facets in two landscapes. In areas where a focal species approach to linkage design is not possible, our results suggest that conservation practitioners may be able to implement a land facets approach with some confidence that the linkage design would serve most potential focal species. In areas where focal species designs are possible, we recommend using the land facet approach to complement, rather than replace, focal species approaches.     

Cats under cover: Habitat models indicate a high dependency on woodlands by Atlantic Forest felids

Four Neotropical small and medium felids—the ocelot, jaguarundi, margay, and southern tiger cat—have overlapping geographic distributions in the endangered Atlantic Forest. Local studies show that these felids avoid areas with high human impact, but the three smaller ones use human‐modified areas more frequently than do ocelots. To understand how landscape changes affect the habitat distribution of these four felids in the Atlantic Forest of Argentina, we used maximum entropy models to analyze the effect of environmental and anthropogenic factors. We estimated niche breadth and overlap among these felids. The conversion of the native forest to land uses without trees was the most important variable that determined the habitat distribution of the four species. For all four species, the optimal habitat covered about 1/3 of the study area and corresponds mainly to the native forest areas. Nearly 50% of these areas had some level of protection. The niche width was higher for the small felids than for ocelots. Niche overlap was high for all species pairs, but higher among the small felids and lower for each of these with the ocelot. The four felids were negatively affected by native forest loss, with ocelots being more sensitive than the smaller felids. The conversion of unprotected forest areas to other types of land uses would imply a greater habitat loss for these felids. The protection of current remnants of Atlantic Forest in Argentina is important for the long‐term conservation of the four felids. Abstract in Spanish is available with online material.     

Modular Wear Facet Nomenclature for mammalian post-canine dentitions

Abstract

Dental wear facets on the occlusal surface of premolars and molars are traces of their main function, the mastication and therefore reflect masticatory movements and also paramasticatory (i.e. non-dietary use of teeth) behavior. Here we present the Modular Wear Facet Nomenclature applicable to most mammalian dentitions. Topographic positions of wear facets in relation to the major cusps and crests of the teeth are used to designate the areas of the occlusal surface the facets occupy (e.g. their mesial, distal, lingual, or buccal position). Previous published systems for labeling wear facets have been inconsistent with each other. Therefore, we provide a synoptic review of the most widely-used terminologies, and introduce the alternative Modular Wear Facet Nomenclature. This nomenclature aims to overcome the difficulties caused by the existing inconsistent wear facet terminologies. Our new approach is applicable to dentitions where the occlusal morphology does not change significantly for most of the lifetime of the animal. In those dentitions, the primary occlusal surfaces are not significantly modified as wear facets become more extensive with wearing. This appears to be a common pattern in pre-tribosphenic, tribosphenic molars, and the teeth derived from tribosphenic precursors (e.g. bunodont molar morphologies). In teeth where the secondary occlusal surface is functionally intensely modified (i.e. high-crowned and evergrowing teeth with large areas of dentine exposed) any facet labeling system appears to be challenging, since the identification of individual facets is blurred and their spatial position may be indeterminable.