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.     

Body size and species-richness in carnivores and primates.

We use complete species-level phylogenies of extant Carnivora and Primates to perform the first thorough phylogenetic tests, in mammals, of the hypothesis that small body size is associated with species-richness. Our overall results, based on comparisons between sister clades, indicate a weak tendency for lineages with smaller bodies to contain more species. The tendency is much stronger within caniform carnivores (canids, procyonids, pinnipeds, ursids and mustelids), perhaps relating to the dietary flexibility and hence lower extinction rates in small, meat-eating species. We find significant heterogeneity in the size-diversity relationship within and among carnivore families. There is no significant association between body mass and species-richness in primates or feliform carnivores. Although body size is implicated as a correlate of species-richness in mammals, much of the variation in diversity cannot be attributed to size differences.     

Basicranial axis length v. skull length in analysis of carnivore skull shape

Skull length is the measurement most commonly used as a standard against which other aspects of cranial morphology are compared to derive an index of relative size or proportions. However, skull length is composed of two different functional components, facial skull and cerebral skull, which vary independently and have different scaling relationships with body size. An analysis of carnivore skull shape with measurements standardized against basicranium length produced very different results than an analysis using skull length as the standard. For example, expressions of relative size of cranial measurements were reduced by 13% in mustelids and increased by 20% in canids, reflecting removal of jaw length (short in mustelids and long in canids) from the comparative standard (basicranial axis length). Cranial measurements scale with higher allometric exponents against basicranial axis length than against skull length.     

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.     

Chromosomal rearrangements and karyotype evolution in carnivores revealed by chromosome painting

Chromosomal evolution in carnivores has been revisited extensively using cross-species chromosome painting. Painting probes derived from flow-sorted chromosomes of the domestic dog, which has one of the most rearranged karyotypes in mammals and the highest dipoid number (2n=78) in carnivores, are a powerful tool in detecting both evolutionary intra- and inter-chromosomal rearrangements. However, only a few comparative maps have been established between dog and other non-Canidae species. Here, we extended cross-species painting with dog probes to seven more species representing six carnivore families: Eurasian lynx (Lynx lynx), the stone marten (Martes foina), the small Indian civet (Viverricula indica), the Asian palm civet (Paradoxurus hermaphrodites), Javan mongoose (Hepestes javanicas), the raccoon (Procyon lotor) and the giant panda (Ailuropoda melanoleuca). The numbers and positions of intra-chromosomal rearrangements were found to differ among these carnivore species. A comparative map between human and stone marten, and a map among the Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis), stone marten and human were also established to facilitate outgroup comparison and to integrate comparative maps between stone marten and other carnivores with such maps between human and other species. These comparative maps give further insight into genome evolution and karyotype phylogenetic relationships among carnivores, and will facilitate the transfer of gene mapping data from human, domestic dog and cat to other species.     

Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora)

The Carnivora occupy a wide range of feeding niches in concordance with the enormous diversity in their skull and dental form. It is well established that differences in crown morphology are linked to variations in the material properties of the foods ingested and masticated. However, how tooth root form is related to dietary specialization is less well known. In the present study, we investigate the relationship between tooth root morphology and dietary specialization in terrestrial carnivores (canids, felids, hyaenids, and ursids). We specifically address the question of how variation in tooth root surface area is related to bite force potentials as one of the crucial masticatory performance parameters in feeding ecology. We applied computed tomography imaging to reconstruct and quantify dental root surface area in 17 extant carnivore species. Moreover, we computed maximal bite force at several tooth positions based on a dry skull model and assessed the relationship of root surface area to skull size, maximal bite force, food properties, and prey size. We found that postcanine tooth root surface areas corrected for skull size serve as a proxy for bite force potentials and, by extension, dietary specialization in carnivores. Irrespective of taxonomic affinity, species that feed on hard food objects have larger tooth roots than those that eat soft or tough foods. Moreover, carnivores that prey on large animals have larger tooth root surface areas. Our results show that tooth root morphology is a useful indicator of bite force production and allows inferences to be made about dietary ecology in both extant and extinct mammals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 456–471.     

Body size of insular carnivores: evidence from the fossil record

Aim Our goals here are to: (1) assess the generality of one aspect of the island rule – the progressive trend towards decrease in size in larger species – for fossil carnivores on islands; (2) offer causal explanations for this pattern and deviations from it – as far as fossil carnivores are concerned; and (3) estimate the speed of this trend. Location Oceanic and oceanic‐like islands world‐wide. Methods Body size estimates of fossil insular carnivores and of their phylogenetically closest mainland relative were obtained from our own data and the published literature. Our dataset consisted of 18 species from nine islands world‐wide. These data were used to test whether the body size of fossil insular carnivores varies as a function of body size of the mainland species in combination with characteristics of the island ecosystem. Results Dwarfism was observed in two canid species. Moderate decrease in body mass was observed in one hyena species. Gigantism was observed in one otter species. Moderate body mass increase was observed in two otter species, one galictine mustelid and perhaps one canid. Negligible or no change in body mass at all was observed in five otter species, three galictine mustelids and one genet. Size changes in teeth do not lag behind in comparison to skeletal elements in the dwarfed canids. The evolutionary speed of dwarfism in a canid lineage is low. Main conclusions Size change in fossil terrestrial insular carnivores was constrained by certain ecological conditions, especially the availability of prey of appropriate body size. When such alternative prey was not available, the carnivores retained their mainland size. The impact of competitive carnivores seems negligible. The case of (semi‐)aquatic carnivores is much less clear. The species that maintained their ancestral body mass may have changed their diet, as is evidenced by their dentition. Among the otters, one case of significant size increase was observed, perhaps best explained as being due to it entering the niche of an obligate aquatic otter. Dwarfism was not observed in otters. The island rule seems to apply to fossil carnivores, but with exceptions. The dependency of the island rule on resource availability is emphasized by the present study.     

The monophyletic origin of delayed implantation in carnivores and its implications

Abstract. In several carnivores a newly fertilized egg enters diapause instead of being directly implanted into the uterus, a phenomenon called delayed implantation. Several hypotheses have been forwarded to explain the utility of this prolonged gestation period, but all of these depend on several independent origins of the character. Here, we conduct a phylogenetic reconstruction of the evolution of delayed implantation in the Carnivora that reveals one basal origin, with additional transitions all having occurred within the Mustelidae. Hence, previous hypotheses relating to its evolution become untestable. Further analyses revealed that the presence or absence of delayed implantation is unrelated to the timing of mating season and birth season. Instead, mustelids with direct implantation are smaller than those with delayed implantation. We therefore suggest that delayed implantation has been selected against in small species due to the relatively higher fecundity costs of a prolonged gestation period.     

From Desert to Rainforest: Phenotypic Variation in Functionally Important Traits of Bushy-Tailed Woodrats (Neotoma cinerea) Across Two Climatic Extremes

Changes in body size inversely related to ambient temperatures have been described in woodrats (Neotoma) over time scales ranging from decades to millennia. However, climate-mediated variation in other traits has not been evaluated, and the effects of precipitation have been overlooked. We assessed variation in skull morphology among bushy-tailed woodrats (Neotoma cinerea) over two sampling transects spanning coastal rainforest and interior desert environments to determine whether skull morphology varied with climate. We also tested whether previously described size-temperature relationships could be generalized to our study populations. In both transects, linear measurements of functionally significant traits differed between coastal and interior populations. Geometric morphometric analyses of shape confirmed some of those differences and revealed additional patterns of skull variation. Variation in some linear measurements, including body size, was predicted by climate. However, body and skull size, as well as measurements of skull components, displayed varying responses. Although longitudinal patterns of body size variation supported Bergmann’s rule, skull size variation was only weakly associated with climate. The strongest phenotypic responses to climate were those of auditory, dental, and palatal skull traits. Altogether, our findings suggest that geographic variation in temperature and precipitation mediated selective heterogeneity and plasticity in skull traits associated with food processing and sensory organs in N. cinerea. This was consistent with our expectation of resource-dependent phenotypic variation among populations in environments with highly contrasting climatic regimes.     

Sparassocynus (Marsupialia, Didelphidae), a peculiar mammal from the late Cenozoic of Argentina

The extinct genus Sparassocynus is known from the Montehermosan, Chapadmalalan, and Uquian stages of Argentina, spanning the Pliocene-Pleistocene transition, and a relative occurs in the older Huayquerian. After a taxonomict review, the morphology of skull, mandible, and dentition is described. The short, triangular face, wide zygomata, broad cranium with large completely bone-enclosed hypo- and epitympanic sinuses, and carnassialized molars are unique among known American marsupials. All these characters are functionally convergent toward living Australian phascogaline dasyurids, small marsupial carnivores, although phylogenetically Sparassocynus clearly represents a moderately specialized offshoot of the American family Didelphidae. It represents a small steppe carnivore now ecologically replaced, at least in part, by placental carnivores that spread from North America in the late Pliocene and Pleistocene.     

Surplus killing by carnivores

In several field observations, foxes, Spotted hyaenas and other carnivores killed many more prey individuals than they could eat. Functional and causal aspects of this phenomenon are discussed and the conclusion is reached that these surplus kills are the consequence of behavioural compromises in both predator and prey to meet opposing environmental requirements.  

Summary:

(1) Observations are reported in which carnivores killed considerably more prey animals than they could possibly eat, and causal and functional aspects of this behaviour are discussed. The species concerned were especially foxes and Spotted hyaenas, and references are quoted about surplus killing by other Canidae, Felidae and Ursidae.
(2) It is argued that satiation in carnivores does not inhibit further catching and killing , but it probably does inhibit searching and hunting. Thus carnivores are able to procure an "easy prey" but normally satiation limits numbers killed.
(3) Many, if not all, carnivores possess behaviour patterns which allow utilization of a kill at a later time, or allow other members of the same social unit or offspring to use the food.
(4) Several prey species showed a lack of anti-predator reaction under particular climatological circumstances; it is argued that this lack of response usually has survival value. Sometimes anti-predator behaviour is accidentally made ineffective.
(5) Surplus kills are made possible by (2) and (4) above, and only very rare circumstances give a predator access to so many prey that (3) is ineffective. It is suggested that surplus kills are the consequence of behavioural compromises in both predator and prey to meet opposing environmental requirements.     

Canine size, shape, and bending strength in primates and carnivores

Anthropoid primates are well known for their highly sexually dimorphic canine teeth, with males possessing canines that are up to 400% taller than those of females. Primate canine dimorphism has been extensively documented, with a consensus that large male primate canines serve as weapons for intrasexual competition, and some evidence that large female canines in some species may likewise function as weapons. However, apart from speculation that very tall male canines may be relatively weak and that seed predators have strong canines, the functional significance of primate canine shape has not been explored. Because carnivore canine shape and size are associated with killing style, this group provides a useful comparative baseline for primates. We evaluate primate maxillary canine tooth size, shape and relative bending strength against body size, skull size, and behavioral and demographic measures of male competition and sexual selection, and compare them to those of carnivores. We demonstrate that, relative to skull length and body mass, primate male canines are on average as large as or larger than those of similar sized carnivores. The range of primate female canine sizes embraces that of carnivores. Male and female primate canines are generally as strong as or stronger than those of carnivores. Although we find that seed-eating primates have relatively strong canines, we find no clear relationship between male primate canine strength and demographic or behavioral estimates of male competition or sexual selection, in spite of a strong relationship between these measures and canine crown height. This suggests either that most primate canines are selected to be very strong regardless of variation in behavior, or that primate canine shape is inherently strong enough to accommodate changes in crown height without compromising canine function.     

Carnivore olfactory bulb size: allometry, phylogeny and ecology

Olfactory bulb size was measured in 146 species of Carnivora in order to examine whether recently observed functional patterns for overall brain size were similar for component parts of the brain. Comparative measures were analysed in relation to various allometric characters (body, brain and skull size), phylogeny, behaviour and ecology. Olfactory bulbs are significantly and positively correlated with all allometric variables, but indices of skull size correlate slightly more closely than other variables. This probably relates to functional aspects of skull size, facial proportions, and anterior elements of the brain. Phylogenetic associations were examined by two comparative methods: the method of independent contrasts and phylogenetic autoregression. Both revealed similar phylogenetic correlation at generic and familial levels. Using calculated values from either method, relative olfactory bulb size only correlates with zonation among seven behavioural and ecological variables; aquatic otters have smaller bulb sizes than carnivores of other zonal types. This agrees with discussion about the diminution of olfactory communication in aquatic environments. Also, olfactory bulb size correlates with home range size, which is consistent with a recent model on the use of olfaction for foraging in designated home ranges. Generally, comparative differences in olfactory bulb size in carnivores do not associate with functional variables found in other comparative studies. Nevertheless, future analyses of specific brain components in mammals may be more useful than overall brain size for testing evolutionary hypotheses of mammalian brain size.     

Geographic Variation in the Skull Morphometry of Four Populations of Batrachuperus karlschmidti(Urodela: Hynobiidae)

Geographic variation of morphology is an important topic of evolutionary biology, and research on geographic variation can provide insights on the formation, evolution, and adaptation of species and subspecies. The vertebrate skull is a developmentally and functionally complex morphological structure with multiple functions, that is susceptible to vary according to selection pressure. In this study, geographic variations in skull morphology of Batrachuperus karlschmidti from four different geographic populations(Shade, Gexi,Shangluokema, and Xinduqiao) were examined via geometric morphometrics. No significant differences were found among these populations with regard to skull size; however, significant variation was found in skull shape. The most notable shape changes are the relative sizes and positions of the frontal, maxilla,pterygoid, and vomer. Skull shape changes were not related to allometry. However, due to limitation of sample populations and size, the results of this study need to be further verified by more sample populations and individuals in the future. The results of this study contribute to our knowledge about these aspects of morphological variability in this species as well as in hynobiid salamanders.     

狒狒圈舍废弃栖杠的嗅觉丰容对圈养食肉动物行为的影响

单调的饲养环境是导致圈养食肉动物表现刻板行为的主要原因之一。为减少圈养动物刻板行为的发生,探索便于操作、经济且有效的丰容方式是动物饲养管理者面临的一个挑战。本研究以狒狒圈舍废弃栖杠作为嗅觉丰容材料,对北京动物园小型(9种)、中型(4种)和大型(7种)食肉动物进行嗅觉丰容实验。在非投喂时段,采用扫描动物取样和瞬时记录法,分别对28只动物在丰容前后的5种行为进行观察和比较分析。结果显示,小型和中型食肉动物在丰容后对旧设施的使用频次显著下降(小型：P=0.026;中型：P=0.038),而大型食肉动物无显著变化(P> 0.05)。仅小型食肉动物的刻板性走动、休息和观望行为频次显著下降(走动：P=0.023;休息：P=0.023;观望：P=0.017),其他类群的各类行为均无显著变化(P> 0.05)。同时,丰容后小型食肉动物的行为多样性指数显著降低(P=0.008),主要表现为动物花费较多的时间探究新的丰容设施;中型和大型食肉动物的行为多样性指数变化不显著。结果表明,使用带有狒狒身体气味的栖杠进行嗅觉丰容,仅对小型食肉动物有效,对中型和大型食肉动物无效。研究认为,以其他动物圈舍的...     

Endozoochorous seed dispersal by sympatric mustelids, Martes melampus and Mustela itatsi, in western Tokyo, central Japan

We investigated seed dispersal by two sympatric mustelid species, the Japanese marten (Martes melampus) and Japanese weasel (Mustela itatsi), along an intercity forest path in western Tokyo, central Japan, from Jul 2007 to Jul 2008. We aimed to investigate the effect of food/habitat preference of these mustelids (martens are semi-arboreal frugivores while weasels are terrestrial carnivores) on their seed dispersal characteristics, which determine their efficacy as seed dispersers. In total, we analyzed 478 fecal samples collected from the two mustelids (N_marten = 381, N_weasel = 97). The proportions of feces containing seeds for martens and weasels were 81.4% and 55.7%, respectively. The number of plant species whose seeds were found within the feces were 28 and 17, respectively. Almost all seeds within feces of both mustelids were intact. The number of plant species whose seeds were found within a single fecal sample ranged from one to four, but no significant difference was detected between the two mustelids. However, marten feces contained a significantly greater number of seeds of most plant species as well as total number of seeds than did weasel feces. The numbers of plant species and seeds represented in marten feces varied seasonally, but those represented in weasel feces did not. Our findings suggest the possibility that both mustelids act in some ways as seed dispersers, although martens seem to disperse a greater diversity and total amount of seeds.     

Differences in diet and trophic interactions of Patagonian carnivores between areas with mostly native or exotic prey

The prey base for Patagonian carnivores has been altered greatly over the past 150 years due to widespread overgrazing by livestock, invasions by exotic wildlife, and hunting. On ranches in northern Patagonia carnivores consume mostly exotic species, and native herbivores are ecologically extinct in their role as prey. In this study we compare diets of the culpeo (Lycalopex culpaeus), chilla (L. griseus), puma (Puma concolor), Geoffroy's cat (Leopardus geoffroyi), colocolo (L. colocolo), and hog-nosed skunks (Conepatus chinga and C. humboldtii) in a reserve with a mostly native prey base to their diets on the ranches, and evaluate how differences in prey bases affect trophic interactions among carnivores. Carnivores in the reserve consumed mostly native prey. Dietary overlap among carnivores was not significant on the reserve, but was highly significant on the ranches. This homogenization of diets where densities of native species are reduced could lead to stronger negative interactions among carnivores, altering the composition of the carnivore assemblage to the detriment of the more specialized species. Study of carnivore diets may be a relatively quick way to evaluate the conservation status and ecological functionality of prey assemblages in Patagonia and other areas where these have been altered.     

An evaluation of the systematic value of skull morphology in the Trimeresurus radiation (Serpentes: Viperidae: Crotalinae) of Asian pitvipers

Trimeresurus (in its widest sense) is a very diverse and widespread radiation of Asian pitvipers, which has been subject to numerous taxonomic revisions, some of which have been based on characteristics of the skull. In order to evaluate the taxonomic utility of such characters, we conducted a comparison of the skulls of 57 specimens representing nine genera, and two currently unassigned species that are very closely related to each other. A canonical variate analysis reveals three distinct phenetic groups: the Protobothrops group, a group containing Ovophis monticola , ' Ovophis ' okinavensis , and ' Trimeresurus ' gracilis , and finally a group comprising the remaining species and characterized by considerable overlap between most genera with the exception of the monotypic Peltopelor and Himalayophis . Agreement between phenetic similarity based on skull characteristics and phylogenetic relationships based on molecular evidence varies between different groups: the morphological similarity of the skull of Protobothrops sieversorum to the other Protobothrops species is congruent with their recent synonymization while the phenetic similarity among the species within the second group does not reflect current molecular phylogenetic relationships and indicates that convergent or parallel evolution may be responsible for at least some of the phenetic similarity detected among skulls of the Asian pit vipers examined. A test of phylogenetic independence, however, indicates that there is still a significant phylogenetic signal that can be recovered from several skull characteristics. Thus, we conclude that skull morphology can contribute to an overall understanding of pitviper taxonomy, but that it would be unwise to rely on skull characteristics alone.     

Spatial and temporal adjustments allowing the coexistence among carnivores in Liguria (N-W Italy)

We investigated the seasonal spatial and temporal co-occurrence of three carnivore species in Liguria region (NW Italy)—the red fox (Vulpes vulpes), the European badger (Meles meles) and the wolf (Canis lupus)—using the information provided by camera-trapping monitoring. Data were collected from January 2013 to January 2015 by positioning camera traps in 200 sample stations. During 3479 trap days, we collected 1048 independent videos of target carnivore species, which revealed a general spatial coexistence among carnivores with some differences in seasonal occurrence of species. The red fox and the European badger showed temporal segregation, as their activity patterns suggested a differential use of night-time in all seasons. Activity patterns of the red fox and the wolf revealed moderate-high overlap and similar density distributions in all seasons except during winter. Coexistence between these species may be allowed by temporal segregation during winter and spatial segregation during spring. Finally, results regarding the European badger and the wolf suggest a moderate temporal segregation with a marked avoidance effect for the European badger induced by the presence of tracks left by wolves. Programmes aimed at carnivore conservation, and management should treat the entire guild, as it has been demonstrated that populations of different carnivores interact with each other in complex ways and that fine-scale mechanisms regulating carnivore assemblage influence different aspects of natural communities.     

Baculum length and copulatory behaviour in carnivores and pinnipeds (Grand Order Ferae)

Relationships between baculum length, body weight and copulatory behaviour were examined in 66 species of carnivores and pinnipeds (Grand Order Ferae). Elongated bacula occur in most carnivore species of the families Ursidae, Canidae, Procyonidae and Mustelidae as well as in all pinnipeds studied. By contrast, members of the family Felidae have short bacula in relation to their body weights. Elongate bacula are found in carnivores and pinnipeds with a prolonged single intromission (PI) copulatory pattern. This finding agrees with results of a previous study of baculum length and PI copulatory patterns in primates. The enlarged baculum may serve to strengthen the penis and protect the urethra during prolonged intromissions. The distal pole of the baculum may also assist sperm transport since in some species it projects beyond the tip of the penis and probably contacts the female's os cervix during copulation. It is possible that stimulation of the female's genitalia by the baculum might also be important in mammals which are induced ovulators (e.g. Mustelidae). However, it is notable that elongation of the baculum has also occurred in some groups where females ovulate spontaneously (e.g. Canidae, Primates).