Resolution of recent radiations within three evolutionary lineages of felidae using mitochondrial restriction fragment length polymorphism variation

Patterns of mitochondrial restriction fragment length polymorphism (RFLP) variation were used to resolve more recent relationships among the species of the Felidae ocelot lineage, domestic cat lineage, and pantherine lineage. Twenty-five of 28 restriction enzymes revealed site variation in at least 1 of 21 cat species. The ocelot lineage was resolved into three separate sistertaxa groups: Geoffroy's cat (Oncifelis geoffroyi) and kodkod (O. guigna), ocelot (Leopardus pardalis) and margay (L. wiedii), and pampas cat (Lynchailurus colocolo) and most of the tigrina samples (Leopardus tigrina). Within the domestic cat lineage, domestic cat (Felis catus), European wild cat (F. silvestris), and African wild cat (F. libyca) formed a monophyletic trichotomy, which was joined with sand cat (F. margarita) to a common ancestor. Jungle cat (F. chaus) and black-footed cat (F. nigripes) mtDNAs diverged earlier than those of the other domestic cat lineage species and are less closely related. Within the pantherine lineage, phylogenetic analysis identified two distinct groups, uniting lion (P. leo) with leopard (P. pardus) and tiger (P. tigris) with snow leopard (P. uncia).To whom correspondence should be addressed.     

On the tryptic peptides from hemoglobin chains of six carnivores

Summary The amino acid compositions of the tryptic peptides of the following carnivore hemoglobin chains have been determined: gray fox (Urocyon cineroargenteus); raccoon (Procyon lotor); polar bear (Thalarctos maritimus); coati mundi (Nasua nasua) chain; coati mundi (Nasua narica) two chains; cat (Felis catus) chain; and lion (Panthera leo) chain. These provide a basis for future sequencing of these hemoglobins and construction of an evolutionary tree. The specific results are summarized in the following article (Stenzel and Brimhall, 1977).     

Phylogenetics of Panthera,including Panthera atrox,based on craniodental characters

The phylogenetic position of Panthera atrox within Felidae is still controversial despite many morphological and molecular studies addressing its relationships. This is in part due to the lack of consensus on a tree for Panthera. These inconsistencies suggest the need for further analysis and perhaps even different methodology to better understand pantherine evolution. Morphologic characters from the skull and dentary were analysed within Panthera to elucidate pantherine phylogeny. Extant taxa included Panthera leo (African lion), Panthera tigris (tiger), Panthera onca (jaguar), Panthera pardus (leopard), Uncia uncia (snow leopard) and Neofelis nebulosa (clouded leopard). Four outgroups were used: Crocuta crocuta (spotted hyena), Metailurus spp., Proailurus lemanensis and Pseudaelurus validus. Our study found a clade containing Panthera leo, Panthera tigris and Panthera atrox, suggesting that Panthera atrox is more closely related to the African lion and the tiger than the jaguar, in contrast to what has been recently proposed. Moreover, gross morphological similarities between Panthera atrox and Panthera onca are more likely the result of convergent hunting styles and/or prey selection, rather than phylogenetic affinity.     

Seroprevalence and genomic divergence of circulating strains of feline immunodeficiency virus among Felidae and Hyaenidae species

Feline immunodeficiency virus (FIV) infects numerous wild and domestic feline species and is closely related to human immunodeficiency virus (HIV) and simian immunodeficiency virus (SIV). Species-specific strains of FIV have been described for domestic cat (Felis catus), puma (Puma concolor), lion (Panthera leo), leopard (Panthera pardus), and Pallas' cat (Otocolobus manul). Here, we employ a three-antigen Western blot screening (domestic cat, puma, and lion FIV antigens) and PCR analysis to survey worldwide prevalence, distribution, and genomic differentiation of FIV based on 3,055 specimens from 35 Felidae and 3 Hyaenidae species. Although FIV infects a wide variety of host species, it is confirmed to be endemic in free-ranging populations of nine Felidae and one Hyaenidae species. These include the large African carnivores (lion, leopard, cheetah, and spotted hyena), where FIV is widely distributed in multiple populations; most of the South American felids (puma, jaguar, ocelot, margay, Geoffroy's cat, and tigrina), which maintain a lower FIV-positive level throughout their range; and two Asian species, the Pallas' cat, which has a species-specific strain of FIV, and the leopard cat, which has a domestic cat FIV strain in one population. Phylogenetic analysis of FIV proviral sequence demonstrates that most species for which FIV is endemic harbor monophyletic, genetically distinct species-specific FIV strains, suggesting that FIV transfer between cat species has occurred in the past but is quite infrequent today.     

Implications of Diet for the Extinction of Saber-Toothed Cats and American Lions

The saber-toothed cat, Smilodon fatalis, and American lion, Panthera atrox, were among the largest terrestrial carnivores that lived during the Pleistocene, going extinct along with other megafauna ∼12,000 years ago. Previous work suggests that times were difficult at La Brea (California) during the late Pleistocene, as nearly all carnivores have greater incidences of tooth breakage (used to infer greater carcass utilization) compared to today. As Dental Microwear Texture Analysis (DMTA) can differentiate between levels of bone consumption in extant carnivores, we use DMTA to clarify the dietary niches of extinct carnivorans from La Brea. Specifically, we test the hypothesis that times were tough at La Brea with carnivorous taxa utilizing more of the carcasses. Our results show no evidence of bone crushing by P. atrox, with DMTA attributes most similar to the extant cheetah, Acinonyx jubatus, which actively avoids bone. In contrast, S. fatalis has DMTA attributes most similar to the African lion Panthera leo, implying that S. fatalis did not avoid bone to the extent previously suggested by SEM microwear data. DMTA characters most indicative of bone consumption (i.e., complexity and textural fill volume) suggest that carcass utilization by the extinct carnivorans was not necessarily more complete during the Pleistocene at La Brea; thus, times may not have been “tougher” than the present. Additionally, minor to no significant differences in DMTA attributes from older (∼30–35 Ka) to younger (∼11.5 Ka) deposits offer little evidence that declining prey resources were a primary cause of extinction for these large cats.     

Diet quality in a wild grazer declines under the threat of an ambush predator

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator–prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.     

Diet of the feral cat,Felis catus,in central Australian grassland habitats: do cat attributes influence what they eat?

The house cat Felis catus was introduced to Australia as a pet and means of rodent control over 200 years ago, but now has established feral populations and has become a serious threat to native wildlife. Using stomach content analysis of 73 feral cats from semi-arid grassland habitats in Queensland, Australia, we aimed to identify dominant prey groups in the cats' diet and to explore associations between the diversity of prey eaten and attributes of the cats including body size, condition, sex, age and coat colour. We also sought to determine any relationships between cat size and the size of the dominant prey in the diet, the long-haired rat Rattus villosissimus. Mammals and reptiles were the dominant prey, with R. villosissimus occurring in 60 % of samples and comprising more than half of all prey by volume. Birds and terrestrial invertebrates were the next most important contributors to the diet, but fish, frogs and freshwater crustaceans also were surprisingly well represented. The dietary diversity of cats was largely unrelated to any of the cat attributes that we measured, although a positive relationship emerged between cat head width and the range of prey types eaten. Our study was conducted during a population irruption of R. villosissimus and confirms that cats are able to exploit an abundant focal prey resource when the opportunity occurs. Further research now is needed to explore associations between diet and cat attributes during periods when rats are scarce.     

Radiographs Reveal Exceptional Forelimb Strength in the Sabertooth Cat,Smilodon fatalis

Background

The sabertooth cat, Smilodon fatalis, was an enigmatic predator without a true living analog. Their elongate canine teeth were more vulnerable to fracture than those of modern felids, making it imperative for them to immobilize prey with their forelimbs when making a kill. As a result, their need for heavily muscled forelimbs likely exceeded that of modern felids and thus should be reflected in their skeletons. Previous studies on forelimb bones of S. fatalis found them to be relatively robust but did not quantify their ability to withstand loading.

Methodology/Principal Findings

Using radiographs of the sabertooth cat, Smilodon fatalis, 28 extant felid species, and the larger, extinct American lion Panthera atrox, we measured cross-sectional properties of the humerus and femur to provide the first estimates of limb bone strength in bending and torsion. We found that the humeri of Smilodon were reinforced by cortical thickening to a greater degree than those observed in any living felid, or the much larger P. atrox. The femur of Smilodon also was thickened but not beyond the normal variation found in any other felid measured.

Conclusions/Significance

Based on the cross-sectional properties of its humerus, we interpret that Smilodon was a powerful predator that differed from extant felids in its greater ability to subdue prey using the forelimbs. This enhanced forelimb strength was part of an adaptive complex driven by the need to minimize the struggles of prey in order to protect the elongate canines from fracture and position the bite for a quick kill.     

Spatio-Temporal Variation in Predation by Urban Domestic Cats (Felis catus) and the Acceptability of Possible Management Actions in the UK

Urban domestic cat (Felis catus) populations can attain exceedingly high densities and are not limited by natural prey availability. This has generated concerns that they may negatively affect prey populations, leading to calls for management. We enlisted cat-owners to record prey returned home to estimate patterns of predation by free-roaming pets in different localities within the town of Reading, UK and questionnaire surveys were used to quantify attitudes to different possible management strategies. Prey return rates were highly variable: only 20% of cats returned ≥4 dead prey annually. Consequently, approximately 65% of owners received no prey in a given season, but this declined to 22% after eight seasons. The estimated mean predation rate was 18.3 prey cat⁻¹ year⁻¹ but this varied markedly both spatially and temporally: per capita predation rates declined with increasing cat density. Comparisons with estimates of the density of six common bird prey species indicated that cats killed numbers equivalent to adult density on c. 39% of occasions. Population modeling studies suggest that such predation rates could significantly reduce the size of local bird populations for common urban species. Conversely, most urban residents did not consider cat predation to be a significant problem. Collar-mounted anti-predation devices were the only management action acceptable to the majority of urban residents (65%), but were less acceptable to cat-owners because of perceived risks to their pets; only 24% of cats were fitted with such devices. Overall, cat predation did appear to be of sufficient magnitude to affect some prey populations, although further investigation of some key aspects of cat predation is warranted. Management of the predation behavior of urban cat populations in the UK is likely to be challenging and achieving this would require considerable engagement with cat owners.     

Comparative analysis of the diet of feral and house cats and wildcat in Europe

Differences in availability of food resources are often manifested in the differentiation of feeding habits of closely related mammal species. Therefore, we assumed that the diet composition and trophic niche of house (i.e., highly dependent on human households), feral (i.e. independent on human households) domestic cats (Felis silvestris catus) and wildcats (Felis s. silvestris) differs. Based on the literature data from Europe (53 study sites), we compared the diet of these three felids analyzed with use of indirect methods (stomach and scat analyses). In the case of the house cat, we additionally compared consumption data obtained directly from prey brought home. Data were expressed as the relative frequency of occurrence to compare dietary patterns. The main prey of the three cat types were small mammals in different ratios. According to the stomach and scat samples, the diet composition of the cat types showed differences in the consumption of rodents, insectivores, wild ungulates, and household food, supporting the “dietary differences originate from varying resources” hypothesis. More opportunistic house cats had a broader trophic niche than feeding specialist wildcats, while feral cats had an intermediate position. The trophic niche breadth of all three cat types increased along a latitudinal gradient from northern to southern areas of Europe. The predation of the house cat which was examined from prey brought home differed from the data obtained by indirect diet analysis; however, it yielded similar results to the diet of the feral cat and the wildcat. Due to their high numbers and similarity of its diet to the wildcat, house cats are a threat to wild animals; therefore, their predation pressure needs to be further investigated.     

Evidence for African origins of founders of the asiatic lion species survival plan

The Asiatic lion (Panthera leo persica) exists in the wild as a single relict population of approximately 250 individuals in the protected Gir Forest Sanctuary in western India. In 1981, a species survival plan (SSP) for the Asiatic lion was established by the American Association of Zoological Parks and Aquariums to manage the 200 + descendants of Asiatic lions in captivity in western zoological facilities. This captive population was derived from seven founders. In order to compare the genetic structure of the Gir Forest population with that of the captive SSP population, a genetic survey of 46 electrophoretic allozyme systems resolved from extracts of lion blood was undertaken by using 29 SSP Asiatic lions and 28 wild-caught or captive-bred lions maintained at the Sakkarbaug Zoo in India but originally derived from the Gir Forest. The Gir lion population was found to be genetically monomorphic at each of 46 allozyme loci. This was in contrast to several African lion (Panthera leo leo) populations, which show moderate levels of allozyme variation at the same loci. The SSP lion population was polymorphic at three allozyme loci (IDHI, TF, and PTI) for alleles that were previously found only in African lion populations. Pedigree analysis of the genetic transmission of these three biochemical loci demonstrated that two of the five primary founder animals of the SSP Asiatic lion population (a breeding pair originally imported from the Trivandrum Zoo in southern India) were descendants of the African subspecies. Three other founder animals were pure Asian. A retrospective SSP pedigree analysis of two morphologic characters (prominent abdominal fold and pairing of infraorbital foramen) that are partially diagnostic for persica vs leo was consistent with this conclusion as well. The implications for the management of small captive populations of threatened species and of the Asiatic lion SSP population are discussed.     

Environmental factors influencing spotted hyena and lion population biomass across Africa

The spotted hyena (Crocuta crocuta Erxleben) and the lion (Panthera leo Linnaeus) are two of the most abundant and charismatic large mammalian carnivores in Africa and yet both are experiencing declining populations and significant pressures from environmental change. However, with few exceptions, most studies have focused on influences upon spotted hyena and lion populations within individual sites, rather than synthesizing data from multiple locations. This has impeded the identification of over‐arching trends behind the changing biomass of these large predators. Using partial least squares regression models, influences upon population biomass were therefore investigated, focusing upon prey biomass, temperature, precipitation, and vegetation cover. Additionally, as both species are in competition with one other for food, the influence of competition and evidence of environmental partitioning were assessed. Our results indicate that spotted hyena biomass is more strongly influenced by environmental conditions than lion, with larger hyena populations in areas with warmer winters, cooler summers, less drought, and more semi‐open vegetation cover. Competition was found to have a negligible influence upon spotted hyena and lion populations, and environmental partitioning is suggested, with spotted hyena population biomass greater in areas with more semi‐open vegetation cover. Moreover, spotted hyena is most heavily influenced by the availability of medium‐sized prey biomass, whereas lion is influenced more by large size prey biomass. Given the influences identified upon spotted hyena populations in particular, the results of this study could be used to highlight populations potentially at greatest risk of decline, such as in areas with warming summers and increasingly arid conditions.     

Fréquences géniques chez le chat domestique,Felis catus,dans quatre communes rurales françaises

Gene frequencies in the domestic cat, Felis catus, in four French rural areas — Frequencies of eight genes (seven responsible for coat colour and one for hair length) are analyzed in French populations of domestic cats (Felis catus); for this purpose, exhaustive samplings have been done in four rural areas. Sex-ratio values are in favour of females. Our data are added to those obtained by other authors in order to perform a comparative analysis of rural and urban areas. This analysis reveals a disadvantage of the Orange allele in urban areas, whereas no tendency is shown for the other genes studied. These results are discussed in relation to the spatial heterogeneity of populations.     

Testing Predictions of the Prey of Lion Derived From Modeled Prey Preferences

Abstract: Apex predators are often threatened with extinction, and reintroduction is one method conservation managers are using to secure their persistence. Yet the ability to predict what these predators will eat upon reintroduction is lacking. Here we test predictions of the diet of the lion (Panthera leo), derived from dietary electivity index and optimality theory, using independent data collected from reintroduced and resident populations. We solved the Jacobs’ index preference equation for each prey species of the lion using values calculated by Hayward and Kerley (2005) and prey abundance data from 4 reintroduction sites and one resident lion population over several years. We then compared these estimates with actual kill data gathered from each site and time period, using the log-likelihood ratio and linear regression. The model precisely predicted the observed number of kills in 9 of the 13 tests. There was a highly significant linear relationship between the number of lion kills predicted to occur at a site and the number observed for all but one site (x̄r² = 0.612; β = 1.03). Predicting predator diet will allow conservation managers to stop responding and start planning in advance for reintroductions and environmental variation. Furthermore, ensuring that sufficient food resources are available is likely to increase the success of reintroduction projects. In addition, managers responsible for threatened prey species will be able to predict the vulnerability of these species to predation in the event of predator reintroductions or changes in abundance. These methods are applicable to virtually all large predators that have been sufficiently studied.     

Dietary partitioning of three large carnivores in Majete Wildlife Reserve,Malawi

Between 2011 and 2012, the carnivore guild in Majete Wildlife Reserve (MWR), Malawi, was restored following the reintroduction of lion (Panthera leo) and leopard (Panthera pardus). The aim of this study was to describe and compare the diet of lion, leopard and resident spotted hyaena (Crocuta crocuta) using scat analysis. Lions and spotted hyaenas displayed the greatest dietary overlap (O_ab = 0.88) and selected mainly medium- to large-bodied prey species. Lions had a mean preferred prey weight of 120.33 ± 42.14 kg (SE), with warthog (Phacochoerus africanus) and waterbuck (Kobus ellipsiprymnus) making up 60.64% of relative biomass consumed. Spotted hyaenas had a mean preferred prey weight of 102.40 ± 41.69 kg and had a more generalised diet (B_a = 0.46) compared to lions (B_a = 0.36). In contrast, leopards occupied a dietary niche substantially lower than that of lions and spotted hyaenas, selecting relatively smaller prey with a mean preferred prey weight of 27.50 ± 6.74 kg. Our results suggest that coexistence between the resident hyaena and reintroduced lion and leopard in MWR is facilitated by dietary partitioning. We advise long-term monitoring of reintroduced carnivores in small, enclosed reserves to assess their impacts on predator and prey populations.     

Evolutionary analysis of a large mtDNA translocation (numt) into the nuclear genome of the Panthera genus species

Translocation of cymtDNA into the nuclear genome, also referred to as numt, has been reported in many species, including several closely related to the domestic cat (Felis catus). We describe the recent transposition of 12,536 bp of the 17 kb mitochondrial genome into the nucleus of the common ancestor of the five Panthera genus species: tiger, P. tigris; snow leopard, P. uncia; jaguar, P. onca; leopard, P. pardus; and lion, P. leo. This nuclear integration, representing 74% of the mitochondrial genome, is one of the largest to be reported in eukaryotes. The Panthera genus numt differs from the numt previously described in the Felis genus in: (1) chromosomal location (F2-telomeric region vs. D2-centromeric region), (2) gene make up (from the ND5 to the ATP8 vs. from the CR to the COII), (3) size (12.5 vs. 7.9 kb), and (4) structure (single monomer vs. tandemly repeated in Felis). These distinctions indicate that the origin of this large numt fragment in the nuclear genome of the Panthera species is an independent insertion from that of the domestic cat lineage, which has been further supported by phylogenetic analyses. The tiger cymtDNA shared around 90% sequence identity with the homologous numt sequence, suggesting an origin for the Panthera numt at around 3.5 million years ago, prior to the radiation of the five extant Panthera species.     

Relationship between moonlight and nightly activity patterns of the ocelot (Leopardus pardalis) and some of its prey species in Formosa,Northern Argentina

The moon can profoundly influence the activity patterns of animals. If predators are more successful under bright moonlight, prey species are likely to respond by shifting their own activity patterns (predator-avoidance hypothesis). However, the assumption that prey will necessarily avoid full-moon nights does not take into account that moonlight also allows prey to more easily detect predators, and to forage more efficiently. Thus, nightly activity patterns could depend on night vision capabilities (visual-acuity hypothesis). To consider the possible influences of moonlight and to distinguish between these hypotheses, we used camera-trapping records of a predator, the ocelot (Leopardus pardalis), and several of its night-active prey to compare activity patterns under different moonlight conditions. The ocelots' activity patterns were not strongly related to moonlight, but showed a slight tendency for higher activity during brighter nights. Tapeti rabbits (Sylvilagus brasiliensis) and brocket deer (Mazama americana) showed a clear preference for brighter nights. White-eared opossums (Didelphis albiventris) also showed a trend to be less active in new moon light. In contrast, smaller grey four-eyed opossums (Philander opossum) and the poor eye-sight nine-banded armadillo (Dasypus novemcinctus) showed similar activity patterns across all moon phases. Since activity patterns of most prey species were not shifted away from the activity of the ocelot, the differences between species are probably linked to their night vision capabilities, and emphasise the need for more information on the visual system of these taxa. Their activity patterns seem to be less strongly linked to avoidance of predation than previously thought, suggesting that foraging and predator detection benefits may play a more important role than usually acknowledged.     

To Kill,Stay or Flee: The Effects of Lions and Landscape Factors on Habitat and Kill Site Selection of Cheetahs in South Africa

Understanding how animals utilize available space is important for their conservation, as it provides insight into the ecological needs of the species, including those related to habitat, prey and inter and intraspecific interactions. We used 28 months of radio telemetry data and information from 200 kill locations to assess habitat selection at the 3^rd order (selection of habitats within home ranges) and 4^th order (selection of kill sites within the habitats used) of a reintroduced population of cheetahs Acinonyx jubatus in Phinda Private Game Reserve, South Africa. Along with landscape characteristics, we investigated if lion Panthera leo presence affected habitat selection of cheetahs. Our results indicated that cheetah habitat selection was driven by a trade-off between resource acquisition and lion avoidance, and the balance of this trade-off varied with scale: more open habitats with high prey densities were positively selected within home ranges, whereas more closed habitats with low prey densities were positively selected for kill sites. We also showed that habitat selection, feeding ecology, and avoidance of lions differed depending on the sex and reproductive status of cheetahs. The results highlight the importance of scale when investigating a species’ habitat selection. We conclude that the adaptability of cheetahs, together with the habitat heterogeneity found within Phinda, explained their success in this small fenced reserve. The results provide information for the conservation and management of this threatened species, especially with regards to reintroduction efforts in South Africa.     

Prey availability and intraguild competition regulate the spatiotemporal dynamics of a modified large carnivore guild

Effective conservation management requires an understanding of the spatiotemporal dynamics driving large carnivore density and resource partitioning. In African ecosystems, reduced prey populations and the loss of competing guild members, most notably lion (Panthera leo), are expected to increase the levels of competition between remaining carnivores. Consequently, intraguild relationships can be altered, potentially increasing the risk of further population decline. Kasungu National Park (KNP), Malawi, is an example of a conservation area that has experienced large‐scale reductions in both carnivore and prey populations, leaving a resident large carnivore guild consisting of only leopard (Panthera pardus) and spotted hyena (Crocuta crocuta). Here, we quantify the spatiotemporal dynamics of these two species and their degree of association, using a combination of co‐detection modeling, time‐to‐event analyses, and temporal activity patterns from camera trap data. The detection of leopard and spotted hyena was significantly associated with the detection of preferred prey and competing carnivores, increasing the likelihood of species interaction. Temporal analyses revealed sex‐specific differences in temporal activity, with female leopard activity patterns significantly different to those of spotted hyena and male conspecifics. Heightened risk of interaction with interspecific competitors and male conspecifics may have resulted in female leopards adopting temporal avoidance strategies to facilitate coexistence. Female leopard behavioral adaptations increased overall activity levels and diurnal activity rates, with potential consequences for overall fitness and exposure to sources of mortality. As both species are currently found at low densities in KNP, increased risk of competitive interactions, which infer a reduction in fitness, could have significant implications for large carnivore demographics. The protection of remaining prey populations is necessary to mitigate interspecific competition and avoid further alterations to the large carnivore guild.