Convergence and divergence in prey of sympatric canids and felids: opportunism or phylogenetic constraint?

Since the canids and felids diverged in the mid‐Eocene or earlier, each family has developed a suite of morphological and behavioural adaptations for obtaining and consuming prey. We here distinguish between prey taxa captured and eaten as a result of these phylogenetic adaptations, and those because they are fortuitously encountered, and argue that such supplementary prey, often opportunistically caught, create a buffer between sympatric, and potentially competitive, canids and felids and thus enhance coexistence. We base our analysis on dietary data derived from the stomach contents of four sympatric canid and felid species in the Free State Province, South Africa (canids: Cape fox Vulpes chama and black‐backed jackal Canis mesomelas; felids: African wild cat Felis silvestris lybica and caracal Caracal caracal), and from results of studies on these species elsewhere in southern Africa. The two canid species preyed heavily on invertebrates, and thus opportunistically, while the felids (especially the caracal) concentrated on mammals, prey they are phylogenetically adapted to capture. Only three species of mammalian prey are shared by the four species. The ratio of opportunistically‐to‐phylogenetically mediated prey taxa used (the O/P ratio) differ between the species, with the black‐backed jackal having the most opportunistically caught taxa in its diet, and the caracal the least. As predicted, a comparison of this data with those from dietary studies of the same species carried out elsewhere indicates that the number of opportunistically obtained prey taxa varies more than those resulting from phylogenetic adaptations. The largest canid had the widest food spectrum (35 prey taxa) while the smallest felid had the most restricted one (11 prey taxa). We argue that using the O/P distinction allows a better understanding of changes in food niche breadth of particular species, especially in xeric areas, and gives a better indication of possible exploitative competition for food by sympatric carnivores than when regarding all prey taxa as actively pursued. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 527–538.     

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.     

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.     

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.     

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.     

Forelimb indicators of prey‐size preference in the Felidae

The forelimbs, along with the crania, are an essential part of the prey‐killing apparatus in cats. Linear morphometrics of the forelimbs were used to determine the morphological differences between felids that specialize on large prey, small prey, or mixed prey. We also compared the scaling of felid forelimbs to those of canids to test whether prey capture strategies affect forelimb scaling. Results suggest that large prey specialists have relatively robust forelimbs when compared with smaller prey specialists. This includes relatively more robust humeri and radii, relatively larger distal ends of the humerus, and relatively larger articular areas of the humerus and radius. Large prey specialists also had relatively longer olecranon processes of the ulna and wider proximal paws. These characters are all important for subduing large prey while the cat positions itself for the killing bite. Small prey specialists have relatively longer distal limb elements for swift prey capture, and mixed prey specialists had intermediate values with relatively more robust metacarpals. Arboreal felids also had more robust limbs. They had relatively longer proximal phalanges for better grip while climbing, and a relatively short brachial index (radius to humerus ratio). Additionally, we found that felids and canids differ in forelimb scaling, which emphasizes the dual use of forelimbs for locomotion and prey capture in felids. This morphometric technique worked well to separate prey‐size preference in felids, but did not work as well to separate locomotor groups, as scansorial and terrestrial felids were not clearly distinguished. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Evolution of heterochromatin-associated satellite DNA loci in felids and canids (Carnivora)

Cloned satellite DNAs that hybridize primarily to C-band-positive regions of felid and canid chromosomes were used to probe the organization of satellite families in the genomes of 16 species of felids and 15 species of canids. Southern-blot and quantitative dot-blot experiments demonstrated that satellite families within the great cats (panthera lineage) vary considerably in regard to amount and/or sequence mismatch and vary some-what in regard to restriction patterns. Satellite families within the canids appeared to be more uniform in regard to both amount/sequence and restriction patterns, although some canid species did differ significantly from the consensus in both respects. Even though intrafamilial satellite restriction patterns were generally similar, every species could be shown to have a unique, characteristic pattern.     

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.     

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)).     

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.     

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.     

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.     

The geography of diet variation in Neotropical Carnivora

1. Mammalian carnivores (order Carnivora) perform important regulatory functions in terrestrial food webs. Building a comprehensive knowledge of the dietary patterns of carnivorans and the factors determining such patterns is essential for improving our understanding of the role of carnivorans in ecosystem functioning.
2. In the Neotropics, there are 64 extant species of terrestrial Carnivora, but information on their trophic ecology is diffuse. We compiled and analysed the available quantitative dietary data for Neotropical carnivorans, aiming to detect patterns of intraspecific and interspecific dietary variation at a large geographical scale.
3. The resulting database encompasses information on trophic interactions of 37 native carnivoran species from six families across 14 countries. There are clear geographical biases towards southern Brazil, Chile, and Argentina, and a noticeable knowledge gap within the Amazon. Also, most studies are focused on canids and felids, especially Puma concolor, Panthera onca, Cerdocyon thous, Leopardus pardalis, and Chrysocyon brachyurus, whereas for 27 native species, we found no quantitative dietary information.
4. Neotropical carnivorans consume species from at least 651 genera of vertebrates, invertebrates, and plants. We found clear species-specific dietary patterns and marked differences between Neotropical felids and canids. Although predators generally exhibit high levels of consistency in their diets regarding prey body mass, we detected significant intraspecific variation for all species analysed across study sites.
5. Body mass imposes strong constraints on prey use, but biogeographical differences in prey availability and human influence may drive the geographical variation we found. Overall, observed patterns show not only similarities with resource-use patterns found for carnivorans in other continents, such as nestedness driven by body mass, but also differences, such as high levels of frugivory and consumption of invertebrates by canids. Assessing resource-use patterns is the first step towards a better understanding of processes underlying the organisation of trophic interactions, and is imperative for addressing impacts of defaunation on ecosystems and for informing conservation efforts.

     

DNA extraction from hair shafts of wild Brazilian felids and canids

Wild felids and canids are usually the main predators in the food chains where they dwell and are almost invisible to behavior and ecology researchers. Due to their grooming behavior, they tend to swallow shed hair, which shows up in the feces. DNA found in hair shafts can be used in molecular studies that can unravel, for instance, genetic variability, reproductive mode and family structure, and in some species, it is even possible to estimate migration and dispersion rates in given populations. First, however, DNA must be extracted from hair. We extracted successfully and dependably hair shaft DNA from eight wild Brazilian felids, ocelot, margay, oncilla, Geoffroy's cat, pampas cat, jaguarundi, puma, and jaguar, as well as the domestic cat and from three wild Brazilian canids, maned wolf, crab-eating fox, and hoary fox, as well as the domestic dog. Hair samples came mostly from feces collected at the S?o Paulo Zoo and were also gathered from non-sedated pet or from recently dead wild animals and were also collected from museum specimens. Fractions of hair samples were stained before DNA extraction, while most samples were not. Our extraction protocol is based on a feather DNA extraction technique, based in the phenol:chloroform:isoamyl alcohol general method, with proteinase K as digestive enzyme.     

Incus facet morphology in carnivorous mammals from different ecosystems: Taxonomy vs. habitat

This study is prompted by the discovery of an incus of Hyaenodon, the first known auditory ossicle of this genus and thus of any hyaenodont mammal so far. A large set of incudes of recent Carnivora, including felids, hyaenids, viverrids, herpestids, nandiniid and canids of different ecosystems, was set up for morphological comparison. This study examines especially the incudo-mallear facet. Typically, the incudo-mallear facet is composed of: (1) three articular surfaces in felids, (2) a U-shaped surface in hyaenids and (3) four surfaces in canids. Both taxonomy (on family level) and habitat (open, closed or mixed habitat preference) might have an impact on the morphology of the incus facets, the former having a major impact in our sample. The Hyaenodon incus is small, delicate and possesses an incudo-mallear facet of a general saddle-shape with two articulation facets, a large superior articulation area and a circular, inferior articulation area. Herein, its general morphology and facet shape is most similar to the felid incus morphology.     

Ecogeographical Variation in Skull Shape of South-American Canids: Abiotic or Biotic Processes?

Species morphological changes can be mutually influenced by environmental or biotic factors, such as competition. South American canids represent a quite recent radiation of taxa that evolved forms very disparate in phenotype, ecology and behaviour. Today, in the central part of South America there is one dominant large species (the maned wolf, Chrysocyon brachyurus) that directly influence sympatric smaller taxa via interspecific killing. Further south, three species of similar sized foxes (Lycalopex spp.) share the same habitats. Such unique combination of taxa and geographic distribution makes South American dogs an ideal group to test for the simultaneous impact of climate and competition on phenotypic variation. Using geometric morphometrics, we quantified skull size and shape of 431 specimens belonging to the eight extant South American canid species: Atelocynus microtis, Cerdocyon thous, Ch. brachyurus, Lycalopex culpaeus, L. griseus, L. gymnocercus, L. vetulus and Speothos venaticus. South American canids are significantly different in both skull size and shape. The hypercarnivorous bush dog is mostly distinct in shape from all the other taxa while a degree of overlap in shape—but not size—occurs between species of the genus Lycalopex. Both climate and competition impacts interspecific morphological variation. We identified climatic adaptations as the main driving force of diversification for the South American canids. Competition has a lower degree of impact on their skull morphology although it might have played a role in the past, when canid community was richer in morphotypes.     

Traditional Versus Multivariate Methods for Identifying Jaguar,Puma, and Large Canid Tracks

ABSTRACT The jaguar (Panthera onca) and puma (Puma concolor) are the largest felids of the American Continent and live in sympatry along most of their distribution. Their tracks are frequently used for research and management purposes, but tracks are difficult to distinguish from each other and can be confused with those of big canids. We used tracks from pumas, jaguars, large dogs, and maned wolves (Chrysocyon brachyurus) to evaluate traditional qualitative and quantitative identification methods and to elaborate multivariate methods to differentiate big canids versus big felids and puma versus jaguar tracks (n = 167 tracks from 18 zoos). We tested accuracy of qualitative classification through an identification exercise with field-experienced volunteers. Qualitative methods were useful but there was high variability in accuracy of track identification. Most of the traditional quantitative methods showed an elevated percentage of misclassified tracks (≥20%). We used stepwise discriminant function analysis to develop 3 discriminant models: 1 for big canid versus big felid track identification and 2 alternative models for jaguar versus puma track differentiation using 1) best discriminant variables, and 2) size-independent variables. These models had high classification performance, with <10% of error in the validation procedures. We used simpler discriminant models in the elaboration of identification keys to facilitate track classification process. We developed an accurate method for track identification, capable of distinguishing between big felids (puma and jaguar) and large canids (dog and maned wolf) tracks and between jaguar and puma tracks. Application of our method will allow a more reliable use of tracks in puma and jaguar research and it will help managers using tracks as indicators of these felids' presence for conservation or management purposes.     

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.     

Cephalisation bei Caniden

Cephalisation in Canidae Studied were the intraspecific and interspecific relationships between brain weight and body weight in Canidae. Inclination values of the allometric lines: intraspecific – 0,25, interspecific 0,571. Most of the studied species have nearly the same degree of cephalisation, deviations from the interspecific allometric line are small; the cephalisation of Nyctereutes and Otocyon is less developed than in the other canid species. There is no difference in cephalisation between most canids and felids. Mustelids show a high variability in cephalisation. Criticized were the data of Bauchot (1985) and Gittleman (1986) on brain weight and body weight.     

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.