Evolutionary changes in craniomandibular shape in the great cats (Neofelis Griffith and Panthera Oken)

Evolutionary shape changes in skull and mandibular anatomy was analysed in 223 specimens of pantherine felids (Neofelis nebulosa, Panthera leo, Panthera onca, Panthera pardus, Panthera tigris, Panthera uncia) compared to a small‐felid outgroup, consisting of 86 specimens of nine different species, using digital surface morphometry on 25 (skull) and 17 (mandible) landmarks. Shape evolution in the pantherine species is complex and nonlinear, and involves both large‐scale and small‐scale shape changes. Shape changes frequently differ among the ingroup species, but the four large Panthera species (leo, onca, pardus, tigris) bear some resemblance to each other. The leopard and jaguar bear the closest resemblance to each other, and several shape changes are common to the lion and tiger, but have probably evolved convergently as a result of large size. The lion has undergone the largest and most numerous shape changes from a small‐felid outgroup. Certain shape changes in the skull and, in some respects, the mandible of the clouded leopard bear resemblance to those in the four large Panthera species. The snow leopard is often regarded as the most primitive of the extant Panthera, and skull and mandibular shape changes often diverge markedly from those observed in the other five ingroup taxa; its overall skull shape is rather similar to the small‐felid outgroup. This indicates that the shape changes in the clouded leopard are convergent with those of the four large Panthera species. Landmark integration showed no significant correlation with molecular phylogeny, chiefly owing to the snow leopard being placed among the four large Panthera species. A traditional phylogenetic topology with the snow leopard as the basal‐most species of Panthera yielded a weak but nonsignificant phylogenetic signal. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 766–778.     

Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters

Molecular phylogenetic studies of the extant Pantherinae have resulted in a variety of different hypotheses of relationships. This study presents the results of a cladistic study encompassing 45 osteological and dental characters in the skull and mandible, as well as 13 soft‐tissue and behavioural characters. Analyzing extant pantherines with osteological data only resulted in two equally parsimonious trees, which differed only with respects to the jaguar, a taxon which shows morphological affinity to the tiger as well as the lion + leopard. Addition of soft‐tissue characters resolved this ambiguity, and led to markedly improved bootstrap values. The inclusion of fossil taxa did not have an impact on topology, but was important for a correct understanding of character evolution, due to the fossils having a combination of characters unlike those of any extant taxon. The clouded leopard is the most basal pantherine, followed by the snow leopard. The large pantherines are a well supported group, to which the snow leopard does not belong, contrary to some molecular studies. Panthera palaeosinensis is no tiger, but may be close to the stem group from which the tiger evolved. P. atrox and P. spelaea are not on the lion lineage, as traditionally assumed, but are successive outgroups to the lion + leopard, although the position of P. spelaea is tentative, but is supported by other lines of evidence such as brain anatomy. © The Willi Hennig Society 2008.     

Highly conserved D-loop-like nuclear mitochondrial sequences (Numts) in tiger (Panthera tigris)

Using oligonucleotide primers designed to match hypervariable segments I (HVS-1) ofPanthera tigris mitochondrial DNA (mtDNA), we amplified two different PCR products (500 bp and 287 bp) in the tiger (Panthera tigris), but got only one PCR product (287 bp) in the leopard (Panthera pardus). Sequence analyses indicated that the sequence of 287 bp was a D-loop-like nuclear mitochondrial sequence (Numts), indicating a nuclear transfer that occurred approximately 4.8–17 million years ago in the tiger and 4.6–16 million years ago in the leopard. Although the mtDNA D-loop sequence has a rapid rate of evolution, the 287-bp Numts are highly conserved; they are nearly identical in tiger subspecies and only 1.742% different between tiger and leopard. Thus, such sequences represent molecular ‘fossils’ that can shed light on evolution of the mitochondrial genome and may be the most appropriate outgroup for phylogenetic analysis. This is also proved by comparing the phylogenetic trees reconstructed using the D-loop sequence of snow leopard and the 287-bp Numts as outgroup.     

The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications

By the late 19th and early 20th centuries, authors described several specimens belonging to a very large felid, the size of a lion, from some Late Pleistocene localities at southern Chile and Argentina. These remains were considered as belonging to large and now extinct subspecies of jaguar. In the present contribution based on qualitative and quantitative postcranial and cranial characters, we conclude that that the “Patagonian Panthera” should be considered as representing remains of the American Cave Lion Panthera atrox. Evidence at hand indicates that Panthera atrox was a very large lion-like felid, with reddish skin, that inhabited dry open habitats. Furthermore, in contrast to living jaguars, transported and accumulated its preys in rocky dens. The presence of at least two pantherine lineages (i.e. P. onca and P. atrox) in the Pleistocene of South America indicates that the lineage has a long and complex history in the New World, and that the history of the clade in this landmass is still far from being well understood.     

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.     

What is Panthera palaeosinensis?

The pantherine cat Panthera palaeosinensis from putative Plio-Pleistocene deposits in North China is one of the oldest known species of Panthera , but its relationship to other Pantherinae is still debated. I compare the holotype skull of P. palaeosinensis with 508 skulls of extant and extinct pantherine cats and show that, when controlling for size, the skull morphology of P. palaeosinensis is most similar to that of the lion Panthera leo or leopard Panthera pardus . Results support the hypothesis that P. palaeosinensis represents a form closely related either to the early lion or leopard clade or to the ancestor of the genus Panthera and suggest an Asian origin for Panthera .     

Modelling differential extinctions to understand big cat distribution on Indonesian islands

Aims To model differential extinction rates for island populations of tigers Panthera tigris and leopards P. pardus. Location Indonesia. Methods We built VORTEX population models of tiger and leopard populations on an island the size of Bali (3632 km²), using data from the literature. Results The tiger populations were less extinction prone than the leopard populations. This was unexpected as tigers had the smaller population sizes and, as such, might be assumed to be more extinction prone. We identified several aspects of tiger breeding biology that explain the result. Main conclusions Sea level reconstructions suggest that both tiger and leopard would have been present in Java, Sumatra and Bali at the end of the last glacial. Our model provides a plausible mechanism based on population ecology to explain why these leopard populations were more extinction prone than the tiger populations. In addition it illustrates the potential utility of population ecology models in understanding historical patterns in biogeography.     

Lost populations and preserving genetic diversity in the lion Panthera leo: Implications for its ex situ conservation

Two of the eight recognized lion subspecies, North African Barbary lion (Panthera leo leo) and South African Cape lion (Panthera leo melanochaita), have become extinct in the wild in the last 150 years. Based on sequences of mitochondrial DNA (mtDNA) control region (HVR1) extracted from museum specimens of four Barbary and one Cape lion, the former was probably a distinct population characterized by an invariable, unique mtDNA haplotype, whilst the latter was likely a part of the extant southern African lion population. Extinction of the Barbary line, which may still be found in “generic” zoo lions, would further erode lion genetic diversity. Therefore, appropriate management of such animals is important for maintaining the overall genetic diversity of the species. The mtDNA haplotype unique to the Barbary lion, in combination with the small size of the HVR1 analyzed (c. 130 bp), makes it possible and cost-effective to identify unlabelled Barbary specimens kept in museums and “generic” captive lions that may carry the Barbary line. An initial study of five samples from the lion collection of the King of Morocco, tested using this method, shows that they are not maternally Barbary.     

Only the largest terrestrial carnivores increase their dietary breadth with increasing prey richness

1. Animals should adapt their foraging habits, changing their dietary breadth in response to variation in the richness and availability of food resources. Understanding how species modify their dietary breadth according to variation in resource richness would support predictions of their responses to environmental changes that alter prey communities.
2. We evaluated relationships between the dietary breadth of large terrestrial carnivores and the local richness of large prey (defined as the number of species). We tested alternative predictions suggested by ecological and evolutionary theories: with increasing prey richness, species would (1) show a more diverse diet, thus broadening their dietary breadth, or (2) narrow their dietary breadth, indicating specialisation on a smaller number of prey.
3. We collated data from 505 studies of the diets of 12 species of large terrestrial mammalian carnivores to model relationships between two indices of dietary breadth and local prey richness.
4. For the majority of species, we found no evidence for narrowing dietary breadth (i.e. increased specialisation) with increasing prey richness. Although the snow leopard and the dhole appeared to use a lower number of large prey species with increasing prey richness, larger sample sizes are needed to support this result.
5. With increasing prey richness, the five largest carnivores (puma Puma concolor, spotted hyaena Crocuta crocuta, jaguar Panthera onca, lion Panthera leo, and tiger Panthera tigris), plus the Eurasian lynx Lynx lynx and the grey wolf Canis lupus (which are usually top predators in the areas from which data were obtained), showed greater dietary breadth and/or used a greater number of large prey species (i.e. increased generalism).
6. We suggest that dominant large carnivores encounter little competition in expanding their dietary breadth with increasing prey richness; conversely, the dietary niche of subordinate large carnivores is limited by competition with larger, dominant predators. We suggest that, over evolutionary time, resource partitioning is more important in shaping the dietary niche of smaller, inferior competitors than the niche of dominant ones.

     

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.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

华南虎、东北虎、孟加拉虎的D-100p和ND5序列及其在系统进化分析中的应用

本文采用PCR和质粒克隆测序方法,获得了华南虎线粒体D-loop区的480bp序列和东北虎、孟加拉虎线粒体D-loop区的503bp序列;同时还获得了这三个虎亚种和金钱豹线粒体ND5基因5’端309bp的部分序列。根据D-loop序列分析,华南虎与孟加拉虎、东北虎的平均距离(p-distance)分别为0．11088和O．11087,而东北虎与孟加拉虎间的平均距离为0．00994;根据ND5序列分析,华南虎与孟加拉虎、东北虎的平均距离分别为0．11434和0．11758,而东北虎与孟加拉虎间的平均距离为0．00324。三个虎亚种的mtDNA D-loop和ND5序列比较表明,华南虎是这三个虎亚种中最为古老的亚种。     

Brain size of the lion (Panthera leo) and the tiger (P. tigris): implications for intrageneric phylogeny, intraspecific differences and the effects of captivity

Intraspecific encephalization of the lion and the tiger is investigated for the first time using a very large sample. Using cranial volume as a measure of brain size, the tiger has a larger brain relative to greatest length of skull than the lion, the leopard and the jaguar. The Asian lion has a relatively much smaller brain compared with those of sub-Saharan lions, between which there are few differences. The Balinese and Javan tigers had relatively larger brains compared with those of Malayan and Sumatran tigers, even although these four putative subspecies occupy adjacent ranges in south-eastern Asia. Differences in brain size do not appear to correlate with any known differences in behaviour and ecology and, therefore, may reflect only chance differences in intrageneric and intraspecific phylogeny. However, captive-bred big cats generally have a reduced brain size compared with that of wild animals, so that an animal's life history and living conditions may affect brain size and, hence, functional or environmental explanations should be considered when linking brain size differences to intraspecific phylogenies.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 85–93.     

A quantitative study of copulatory behaviour of large Felidae

A total of 109 copulations was observed in six male-female pairs from four species of large Felidae. The mean intromission durations were 3.0 sec for Asian leopards (Panthera pardus), 3.3 sec for African leopards (Panthera pardus), 12.9 sec for snow leopards (Uncia uncia), 2.3 sec for spotted jaguars (Panthera onca), 3.3 sec for black jaguars (Panthera onca), and 12.4 sec for Siberian tigers (Panthera tigris). Behavioural patterns were qualitatively similar across species; all displayed a copulatory pattern with no lock, no intra-vaginal thrusting, ejaculation on a single insertion, and multiple ejaculations. Whereas domestic cats are reported to assume a neck grip and to tread prior to insertion, these larger Felidae generally did so after intromission had been achieved. After copulation, females of some pairs swiped at the male and displayed a rolling after-reaction.     

Non-Invasive Pregnancy Diagnosis in Big Cats using the PGFM (13,14-dihydro-15-keto-PGF2α) Assay

Non-invasive monitoring of hormones using feces has become a vital tool for reproductive management and reliable pregnancy diagnosis in big cats. Previous studies described the PGF_2α metabolite (PGFM) as an indicator of pregnancy in various feline species. The present study aimed to standardize pregnancy detection in big cats like the tiger (Panthera tigris), jaguar (Panthera onca) and lion (Panthera leo) using fecal samples. High-performance liquid chromatography (HPLC) and liquid chromatography-mass spectrometry (LCMS) were performed to identify PGFM in feces. An EIA developed against 9α,11α-dihydroxy-15-oxo-prost-5-en-1-oic acid-BSA was used to assay PGFM in fecal samples of the Bengal tiger, Asiatic lion and jaguar. The PGFM levels increased after 9 weeks of pregnancy and remained elevated until parturition. All animals showed elevated levels of PGFM in the last trimester of pregnancy, thus making PGFM a reliable tool for pregnancy diagnosis during this period that can be useful in captive breeding programs in these species.     

Development and characterization of novel microsatellite markers in the Asiatic lion (Panthera leo persica)

All cat species, except domestic cats, are either threatened or endangered. The Asiatic lion (Panthera leo persica), a critically endangered species, is driven to a single habitat zone in Gir forests in India for its survival. The present status of this big cat compels us to acquire deeper knowledge of its genetic variability and phylogenetic relationships. Here we report the isolation of highly polymorphic microsatellite loci from a partial genomic library of the Asiatic lion. These loci were characterized and have shown high levels of variation ranging from six to 11 alleles per locus in the population studied.     

Age Estimation of African Lions Panthera leo by Ratio of Tooth Areas

Improved age estimation of African lions Panthera leo is needed to address a number of pressing conservation issues. Here we present a formula for estimating lion age to within six months of known age based on measuring the extent of pulp closure from X-rays, or Ratio Of tooth AReas (ROAR). Derived from measurements taken from lions aged 3–13 years for which exact ages were known, the formula explains 92% of the total variance. The method of calculating the pulp/tooth area ratio, which has been used extensively in forensic science, is novel in the study of lion aging. As a quantifiable measure, ROAR offers improved lion age estimates for population modeling and investigations of age-related mortality, and may assist national and international wildlife authorities in judging compliance with regulatory measures involving age.     

Sympatric Asian felid phylogeography reveals a major Indochinese–Sundaic divergence

The dynamic geological and climatological history of Southeast Asia has spawned a complex array of ecosystems and 12 of the 37 known cat species, making it the most felid‐rich region in the world. To examine the evolutionary histories of these poorly studied fauna, we compared phylogeography of six species (leopard cat Prionailurus bengalensis, fishing cat P. viverrinus, Asiatic golden cat Pardofelis temminckii, marbled cat P. marmorata, tiger Panthera tigris and leopard P. pardus) by sequencing over 5 kb of DNA each from 445 specimens at multiple loci of mtDNA, Y and X chromosomes. All species except the leopard displayed significant phylogenetic partitions between Indochina and Sundaland, with the central Thai–Malay Peninsula serving as the biogeographic boundary. Concordant mtDNA and nuclear DNA genealogies revealed deep Indochinese–Sundaic divergences around 2 MYA in both P. bengalensis and P. marmorata comparable to previously described interspecific distances within Felidae. The divergence coincided with serial sea level rises during the late Pliocene and early Pleistocene, and was probably reinforced by repeated isolation events associated with environmental changes throughout the Pleistocene. Indochinese–Sundaic differentiations within P. tigris and P. temminckii were more recent at 72–108 and 250–1570 kya, respectively. Overall, these results illuminate unexpected, deep vicariance events in Southeast Asian felids and provide compelling evidence of species‐level distinction between the Indochinese and Sundaic populations in the leopard cat and marbled cat. Broader sampling and further molecular and morphometric analyses of these species will be instrumental in defining conservation units and effectively preserving Southeast Asian biodiversity.