Mitogenomic analyses of caniform relationships

Extant members of the order Carnivora split into two basal groups, Caniformia (dog-like carnivorans) and Feliformia (cat-like carnivorans). In this study we address phylogenetic relationships within Caniformia applying various methodological approaches to analyses of complete mitochondrial genomes. Pinnipeds are currently well represented with respect to mitogenomic data and here we add seven mt genomes to the non-pinniped caniform collection. The analyses identified a basal caniform divergence between Cynoidea and Arctoidea. Arctoidea split into three primary groups, Ursidae (including the giant panda), Pinnipedia, and a branch, Musteloidea, which encompassed Ailuridae (red panda), Mephitidae (skunks), Procyonidae (raccoons) and Mustelidae (mustelids). The analyses favored a basal arctoid split between Ursidae and a branch containing Pinnipedia and Musteloidea. Within the Musteloidea there was a preference for a basal divergence between Ailuridae and remaining families. Among the latter, the analyses identified a sister group relationship between Mephitidae and a branch that contained Procyonidae and Mustelidae. The mitogenomic distance between the wolf and the dog was shown to be at the same level as that of basal human divergences. The wolf and the dog are commonly considered as separate species in the popular literature. The mitogenomic result is inconsistent with that understanding at the same time as it provides insight into the time of the domestication of the dog relative to basal human mitogenomic divergences.     

Phylogenetic relationships within caniform carnivores based on analyses of the mitochondrial 12S rRNA gene

The complete 12S rRNA gene of 32 carnivore species, including four feliforms and 28 caniforms, was sequenced. The sequences were aligned on the basis of their secondary structures and used in phylogenetic analyses that addressed several evolutionary relationships within the Caniformia. The analyses showed an unresolved polytomy of the basic caniform clades; pinnipeds, mustelids, procyonids, skunks,Ailurus (lesser panda), ursids, and canids. The polytomy indicates a major diversification of caniforms during a relatively short period of time. The lesser panda was distinct from other caniforms, suggesting its inclusion in a monotypic family, Ailuridae. The giant panda and the bears were joined on the same branch. The skunks are traditionally included in the family Mustelidae. The present analysis, however, showed a less close molecular relationship between the skunks and the remaining Mustelidae (sensu stricto) than between Mustelidae (sensu stricto) and Procyonidae, making Mustelidae (sensu lato) paraphyletic. The results suggest that the skunks should be included in a separate family, Mephitidae. Within the Pinnipedia, the grouping of walrus, sea lions, and fur seals was strongly supported. Analyses of a combined set of 12S rRNA and cytochromeb data were generally consistent with the findings based on each gene.     

Molecular phylogeny of the Arctoidea (Carnivora): effect of missing data on supertree and supermatrix analyses of multiple gene data sets

Phylogenetic relationships of 79 caniform carnivores were addressed based on four nuclear sequence-tagged sites (STS) and one nuclear exon, IRBP, using both supertree and supermatrix analyses. We recovered the three major arctoid lineages, Ursidae, Pinnipedia, and Musteloidea, as monophyletic, with Ursidae (bears) strongly supported as the basal arctoid lineage. Within Pinnipedia, Phocidae (true seals) were sister to the Otaroidea [Otariidae (fur seals and sea lions) and Odobenidae (walrus)]. Phocid subfamily and tribal designations were supported, but the otariid subfamily split between fur seals and sea lions was not. All family designations within Musteloidea were strongly supported: Mephitidae (skunks), Ailuridae (monotypic red panda), Mustelidae (weasels, badgers, otters), and Procyonidae (raccoons). A novel hypothesis for the position of the red panda was recovered, placing it as branching after Mephitidae and before Mustelidae+Procyonidae. Within Mustelidae, subfamily taxonomic changes are considered. This study represents the most comprehensive sampling to date of the Caniformia in a molecular study and contains the most complete molecular phylogeny for the Procyonidae. Our data set was also used in an empirical examination of the effect of missing data on both supertree and supermatrix analyses. Sequence for all genes in all taxa could not be obtained, so two variants of the data set with differing amounts of missing data were examined. The amount of missing data did not have a strong effect; instead, phylogenetic resolution was more dependent on the presence of sufficient informative characters. Supertree and supermatrix methods performed equivalently with incomplete data and were highly congruent; conflicts arose only in weakly supported areas, indicating that more informative characters are required to confidently resolve close species relationships.     

A phylogeny of the Caniformia (order Carnivora) based on 12 complete protein-coding mitochondrial genes

Evolutionary relationships of the order Carnivora have been extensively studied. However, phylogenetic studies based on different types of data, species samples, and methods of analysis provide contradictory results. Consequently, phylogenetic relationships of Carnivora remain contentious. Here, the sequence of 12 mitochondrial genes (10,842 nucleotides) from a total of 38 carnivore species was used to investigate the phylogeny of the caniform (dog-like) carnivores. An analysis using maximum parsimony, maximum likelihood, and Bayesian approaches provided a unique and well-supported solution to most contentious relationships within Caniformia. The clade Arctoidea was shown to consist of three major monophyletic groups: Pinnipedia, Ursidae, and Musteloidea. Within Pinnipedia, the families Otariidae and Odobenidae formed a clade, sister to Phocidae. Within Musteloidea, there was a sister relationship between true mustelids (i.e., excluding the skunks) and procyonids, and between ailurids and mephitids (skunks). Despite a high level of confidence obtained at most nodes, uncertainty remained about the relative position of the three major arctoid clades.     

Molecular phylogenetic study on the origin and evolution of Mustelidae

The family Mustelidae, which consists of Mustelinae, Lutrinae, Melinae, and Taxidiinae, is the largest family among Carnivora and is a highly diverse group. Recent molecular phylogenetic studies have clarified the phylogenetic relations among Mustelidae, but there remain several unresolved problems, particularly concerning the deep branchings. Whereas many studies support the monophyly of Mustelidae+Procyonidae among Musteloidea, the relations between Mustelidae+Procyonidae, Ailuridae, and Miphitidae are still unclear. To address these problems, we inferred a tree on the basis of the sequences of mitochondrial genomes and of multiple nuclear genes using the maximum likelihood method. Our results strongly support the hypothesis that the Taxidiinae branched at first, followed by the branching of the Melinae. After that, Mustelinae diversified, and Lutrinae evolved within Mustelinae. With respect to the deep branchings in Musteloidea, the Ailuridae/Mephitidae monophyly tree and the Mephitidae-basal tree are indistinguishable in log-likelihood score, and this problem remains unresolved.     

Ancestral state reconstruction of body size in the Caniformia (Carnivora, Mammalia): the effects of incorporating data from the fossil record

A recent molecular phylogeny of the mammalian order Carnivora implied large body size as the ancestral condition for the caniform subclade Arctoidea using the distribution of species mean body sizes among living taxa. "Extant taxa-only" approaches such as these discount character state observations for fossil members of living clades and completely ignore data from extinct lineages. To more rigorously reconstruct body sizes of ancestral forms within the Caniformia, body size and first appearance data were collected for 149 extant and 367 extinct taxa. Body sizes were reconstructed for four ancestral nodes using weighted squared-change parsimony on log-transformed body mass data. Reconstructions based on extant taxa alone favored large body sizes (on the order of 10 to 50 kg) for the last common ancestors of both the Caniformia and Arctoidea. In contrast, reconstructions incorporating fossil data support small body sizes (< 5 kg) for the ancestors of those clades. When the temporal information associated with fossil data was discarded, body size reconstructions became ambiguous, demonstrating that incorporating both character state and temporal information from fossil taxa unambiguously supports a small ancestral body size, thereby falsifying hypotheses derived from extant taxa alone. Body size reconstructions for Caniformia, Arctoidea, and Musteloidea were not sensitive to potential errors introduced by uncertainty in the position of extinct lineages relative to the molecular topology, or to missing body size data for extinct members of an entire major clade (the aquatic Pinnipedia). Incorporating character state observations and temporal information from the fossil record into hypothesis testing has a significant impact on the ability to reconstruct ancestral characters and constrains the range of potential hypotheses of character evolution. Fossil data here provide the evidence to reliably document trends of both increasing and decreasing body size in several caniform clades. More generally, including fossils in such analyses incorporates evidence of directional trends, thereby yielding more reliable ancestral character state reconstructions.     

Tandemly repeated satellite DNA ofDolichopoda schiavazzii: A test for models on the evolution of highly repetitive DNA

The evolutionary relationships among the Carnivora were studied in a phylogenetic analysis based on the complete mitochondrial cytochromeb gene. The study, which addressed primarily the relationships among the Caniformia, included 4 feliform and 26 caniform species, with 9 pinnipeds. The analysis identified five caniform clades: Canidae, Ailuridae (with the monotypic lesser panda), Musteloidea (Mustelidae+Procyonidae), Ursidae (including the giant panda), and Pinnipedia. The closest relatives of the Pinnipedia among terrestrial caniforms were not identified conclusively. Our analysis shows that the skunks are only distantly related to remaining mustelids (Mustelidae sensu stricto) and that the family Mustelidae, including the skunks, is paraphyletic. The relationship among the five caniform clades was unresolved, suggesting an evolutionary separation within a relatively short period of time. Based on distance values, we propose that this primary diversification took place 45 million years ago.     

Incorporating indels as phylogenetic characters: Impact for interfamilial relationships within Arctoidea (Mammalia: Carnivora)

Insertion and deletion events (indels) provide a suite of markers with enormous potential for molecular phylogenetics. Using many more indel characters than those in previous studies, we here for the first time address the impact of indel inclusion on the phylogenetic inferences of Arctoidea (Mammalia: Carnivora). Based on 6843 indel characters from 22 nuclear intron loci of 16 species of Arctoidea, our analyses demonstrate that when the indels were not taken into consideration, the monophyly of Ursidae and Pinnipedia tree and the monophyly of Pinnipedia and Musteloidea tree were both recovered, whereas inclusion of indels by using three different indel coding schemes give identical phylogenetic tree topologies supporting the monophyly of Ursidae and Pinnipedia. Our work brings new perspectives on the previously controversial placements among Arctoidea families, and provides another example demonstrating the importance of identifying and incorporating indels in the phylogenetic analyses of introns. In addition, comparison of indel incorporation methods revealed that the three indel coding methods are all advantageous over treating indels as missing data, given that incorporating indels produces consistent results across methods. This is the first report of the impact of different indel coding schemes on phylogenetic reconstruction at the family level in Carnivora, which indicates that indels should be taken into account in the future phylogenetic analyses.     

Phylogeny of the Carnivora (Mammalia): Congruence vs Incompatibility among Multiple Data Sets

The purpose of this study was to determine the higher-level phylogenetic relationships among Carnivora, using a conditional data combination (CDC) approach to analyzing multiple data sets. New nucleotide sequences (851 base pairs from intron I of the transthyretin gene) among 22 representatives of the 11 families of Carnivora were generated and analyzed in concert with, and comparison to, other mitochondrial and morphological character data. Conditional data combination analyses of the four independent data sets (transthyretin intron I, cytochromeb,partial 12S rRNA, and morphology) indicate that the phylogenetic results derived from each generally agree, with two exceptions. The first exception, signal heterogeneity in comparisons involving transthyretin and morphology, provides an example where phylogenetic conclusions drawn from total evidence analyses may differ from conclusions drawn from CDC analyses. The second exception demonstrates that while a CDC method may reject the null hypothesis of homogeneity for a particular partition, including that partition in combined analyses, may nevertheless provide an overall increase in phylogenetic signal, in terms of nodal support for most associations, without altering the topology derived from the combined homogeneous data partitions. Phylogenetic reconstruction among the feliform families supports a sister-group relationship between the hyaenas (Hyaenidae) and mongooses (Herpestidae) and places the African palm civet (Nandinia) as basal to all other living Feliformia. Among the caniform families, CDC analyses strongly support the previously enigmatic red panda (Ailurus) as a monotypic lineage that is sister to Musteloideasensu stricto(mustelids plus procyonids), in addition to pinniped monophyly and a sister-group relationship between the walrus and sea lions.     

Phylogenetic reconstruction of carnivore social organizations

It is generally assumed that carnivore social organizations evolved directionally from a solitary ancestor into progressively more advanced forms of group living. Although alternative explanations exist, this evolutionary hypothesis has never been tested. Here, I used literature data and maximum likelihood reconstruction on a complete carnivore phylogeny to test this hypothesis against two others: one assuming directional evolution from a non-solitary ancestor, and one assuming parallel evolutions from a socially flexible ancestor, that is, an ancestor with abilities to live in a variety of social organizations. The phylogenetic reconstructions did not support any of the three hypotheses of social evolution at the root of Carnivora. At the family level, however, there was support for a non-solitary and socially flexible ancestor to Canidae, a socially flexible or solitary ancestor to Mustelidae, a solitary or socially flexible ancestor to Mephitidae, a solitary or group living ancestor to Phocidae, a group living ancestor to Otariidae and a solitary ancestor to Ursidae, Felidae, Herpestidae and Viverridae. There was equivocal support for the ancestral state of Procyonidae and Hyaenidae. It is unclear whether the common occurrence of a solitary ancestry at the family level was caused by a solitary ancestor at the root of Carnivora or by multiple transitions into a solitary state. The failure to support a solitary ancestor to Carnivora calls for caution when using this hypothesis in an evolutionary framework, and I suggest continued investigations of the pathways of the evolution of carnivore social organizations.     

Morphological integration in the carnivoran skull

The correlated evolution of traits may be a principal factor in morphological evolution, but it is typically studied in genetic or developmental systems. Most studies examining phenotypic trait correlations, through analysis of morphological integration, consider only few taxa, with limited ability to test hypotheses of the influence of trait integration on morphological variation and diversity. The few comparative studies in less inclusive groups have yielded varying relationships of integration to the key factors of phylogeny and diet. In this paper, I present analyses of cranial morphological integration in 30 species from the mammalian order Carnivora, spanning eight extant families and a wide range of ecological and morphological diversity. Fifty-five cranial landmarks were captured through three-dimensional digitization of 15-22 specimens for each species. Using a node-based phylogenetic distance matrix, a significant correlation was found between similarity in patterns of integration and phylogenetic relatedness within Felidae (cats) and Canidae (dogs), but not within more inclusive clades, when size-related variation was removed. When size was included, significant correlations were found across all Caniformia, Musteloidea, Mustelidae, and Felidae. There was a significant correlation between phylogeny and morphological integration only within the higher-level clade Feliformia (cats, civets, mongooses, and hyaenas) when a branch-length-based phylogenetic distance matrix was analyzed, with and without size. In contrast, diet was significantly correlated with similarity in morphological integration in arctoid carnivorans (bears, raccoons, and weasels), but had no significant relationship with integration in feliforms or canids. These results support the proposition that evolutionary history is correlated with cranial integration across large clades, although in some smaller clades diet also exerts significant influence on the correlated evolution of traits.     

Phylogeny of the caniform carnivora: evidence from multiple genes

The monophyletic group Caniformia in the order Carnivora currently comprises seven families whose relationships remain contentious. The phylogenetic positions of the two panda species within the Caniformia have also been evolutionary puzzles over the past decades, especially for Ailurus fulgens (the red panda). Here, new nuclear sequences from two introns of the β-fibrinogen gene (β-fibrinogen introns 4 and 7) and a complete mitochondrial (mt) gene (ND2) from 17 caniform representatives were explored for their utilities in resolving higher-level relationships in the Caniformia. In addition, two previously available nuclear (IRBP exon 1 and TTR intron 1) data sets were also combined and analyzed simultaneously with the newly obtained sequence data in this study. Combined analyses of four nuclear and one mt genes (4417 bp) recover a branching order in which almost all nodes were strongly supported. The present analyses provide evidence in favor of Ailurus fulgens as the closest taxon to the procyonid-mustelid (i.e., Musteloidea sensu stricto) clade, followed by pinnipeds (i.e., Otariidae and Phocidae), Ursidae (including Ailuropoda melanoleuca), and Canidae, the most basal lineage in the Caniformia. The potential utilities of different genes in the context of caniform phylogeny were also evaluated, with special attention to the previously unexplored β-fibrinogen intron 4 and 7 genes.     

The ancestral carnivore karyotype (2n = 38) lives today in ringtails

Chromosome painting was used to investigate the conservation of high-resolution longitudinal 4',6-diamidino-2-phenylindole (DAPI)/G bands in Carnivore chromosomes. Cat (Felis catus) and raccoon dog (Nyctereutes procyonoides) painting probes were hybridized to the ringtail (Bassaricus astutus), dwarf mongoose (Helogale parvula), and Malagasy civet (Fossa fossa) to identify homologous chromosome elements. The patterns of chromosome segment homology among Carnivore species allowed us to reconstruct and propose the disposition of a high-resolution banded ancestral carnivore karyotype (ACK). Three bi-armed chromosomes consistently found among Caniformia species are represented as 6 homologous acrocentric chromosomes among Feliformia species of Carnivora. However, reexamination of the most basal of Feliformia species, the African palm civet Nandinia, revealed the presence of the 3 heretofore Caniformia bi-armed chromosomes. Because these 3 bi-armed chromosomes are found in both Caniformia and Feliformia lineages, they are presumed ancestral for all Carnivora, suggesting that the ACK chromosome number would be 38, rather than the previously supposed 42. Banded chromosomes of the ACK are used to evaluate the consistency between recently determined molecular phylogenetic relationships and postulated cytogenetic dynamics in the same Carnivore species.     

Whence the red panda?

The evolutionary history of the red panda (Ailurus fulgens) plays a pivotal role in the higher-level phylogeny of the "bear-like" arctoid carnivoran mammals. Characters from morphology and molecules have provided inconsistent evidence for placement of the red panda. Whereas it certainly is an arctoid, there has been major controversy about whether it should be placed with the bears (ursids), ursids plus pinnipeds (seals, sea lions, walrus), raccoons (procyonids), musteloids (raccoons plus weasels, skunks, otters, and badgers [mustelids]), or as a monotypic lineage of uncertain phylogenetic affinities. Nucleotide sequence data from three mitochondrial genes and one nuclear intron were analyzed, with more complete taxonomic sampling of relevant taxa (arctoids) than previously available in analyses of primary molecular data, to clarify the phylogenetic relationships of the red panda to other arctoid carnivorans. This study provides detailed phylogenetic analyses (both parsimony and maximum-likelihood) of primary character data for arctoid carnivorans, including bootstrap and decay indices for all arctoid nodes, and three statistical tests of alternative phylogenetic hypotheses for the placement of the red panda. Combined phylogenetic analyses reject the hypotheses that the red panda is most closely related to the bears (ursids) or to the raccoons (procyonids). Rather, evidence from nucleotide sequences strongly support placement of the red panda within a broad Musteloidea (sensu lato) clade, including three major lineages (the red panda, the skunks [mephitids], and a clearly monophyletic clade of procyonids plus mustelids [sensu stricto, excluding skunks]). Within the Musteloidea, interrelationships of the three major lineages are unclear and probably are best considered an unresolved trichotomy. These data provide compelling evidence for the relationships of the red panda and demonstrate that small taxonomic sample sizes can result in misleading or possibly erroneous (based on prior modeling, as well as conflict between the results of our analyses of less and more complete data sets) conclusions about phylogenetic relationships and taxonomy.     

Evidence from nuclear DNA sequences sheds light on the phylogenetic relationships of Pinnipedia: single origin with affinity to Musteloidea

Considerable long-standing controversy and confusion surround the phylogenetic affinities of pinnipeds, the largely marine group of "fin-footed" members of the placental mammalian order Carnivora. Until most recently, the two major competing hypotheses were that the pinnipeds have a single (monophyletic) origin from a bear-like ancestor, or that they have a dual (diphyletic) origin, with sea lions (Otariidae) derived from a bear-like ancestor, and seals (Phocidae) derived from an otter-, mustelid-, or musteloid-like ancestor. We examined phylogenetic relationships among 29 species of arctoid carnivorans using a concatenated sequence of 3228 bp from three nuclear loci (apolipoprotein B, APOB; interphotoreceptor retinoid-binding protein, IRBP; recombination-activating gene 1, RAG1). The species represented Pinnipedia (Otariidae: Callorhinus, Eumetopias; Phocidae: Phoca), bears (Ursidae: Ursus, Melursus), and Musteloidea (Mustelidae: Mustela, Enhydra, Melogale, Martes, Gulo, Meles; Procyonidae: Procyon; Ailuridae: Ailurus; Mephitidae: Mephitis). Maximum parsimony, maximum likelihood, and Bayesian inference phylogenetic analyses of separate and combined datasets produced trees with largely congruent topologies. The analyses of the combined dataset resulted in well-resolved and well-supported phylogeny reconstructions. Evidence from nuclear DNA evolution presented here contradicts the two major hypotheses of pinniped relationships and strongly suggests a single origin of the pinnipeds from an arctoid ancestor shared with Musteloidea to the exclusion of Ursidae.     

Molecular systematics of the Hyaenidae: relationships of a relictual lineage resolved by a molecular supermatrix

The four extant species of hyenas (Hyaenidae; Carnivora) form a morphologically and ecologically heterogeneous group of feliform carnivorans that are remnants of a formerly diverse group of mammalian predators. They include the aardwolf (Proteles cristatus), a termite-feeding specialist, and three species with a craniodental morphology adapted to cracking the bones of prey and/or carcasses, the spotted hyena (Crocuta crocuta), brown hyena (Parahyaena brunnea), and striped hyena (Hyaena hyaena). Hyenas have been the subject of a number of systematic studies during the last two centuries, due in large part to the extensive fossil record of the group, with nearly 70 described fossil species. Morphological studies incorporating both fossil and living taxa have yielded different conclusions regarding the evolutionary relationships among living hyenas. We used a molecular supermatrix comprised of seven nuclear gene segments and the complete mitochondrial cytochrome b gene to evaluate phylogenetic relationships among the four extant hyaenid species. We also obtained sequence data from representative species of all the main families of the Feliformia (Felidae, Herpestidae, and Viverridae) to estimate the sister group of the Hyaenidae. Maximum parsimony and maximum likelihood analyses of the supermatrix recovered identical topologies. Furthermore, Bayesian phylogenetic analyses of the supermatrix, with among-site rate variation among data partitions parameterized in three different ways, also yielded the same topology. For each phylogeny reconstruction method, all but two nodes received 100% bootstrap or 1.00 posterior probability nodal support. Within the monophyletic Hyaenidae, Parahyaena and Hyaena were joined together, with Crocuta as the sister to this clade, and Proteles forming the most basal lineage. A clade containing two species of mongoose (core Herpestidae) plus Cryptoprocta ferox (currently classified in Viverridae) was resolved as the sister group of Hyaenidae. The pattern of relationships among the three bone-cracking hyaenids (Crocuta, Hyaena, and Parahyaena) is incongruent with recent cladistic assessments based on morphology and suggests the need to reevaluate some of the morphological characters that have been traditionally used to evaluate relationships among hyenas. Divergence time estimates based on a Bayesian relaxed molecular clock indicates that hyaenids diverged from their feliform sister group 29.2 MYA, in the Middle Oligocene. Molecular clock estimates also suggest that the origin of the aardwolf is much more recent (10.6 MYA) than that implied by a cladistic analysis of morphology ( approximately 20 MYA) and suggests that the aardwolf is possibly derived from a bone and meat eating lineage of hyaenids that were present in the Late Miocene. [Hyaenidae; phylogeny; cytochrome b; nuclear gene segments; Proteles; Crocuta; Hyaena; Parahyaena.].     

Evolutionary and biogeographic history of weasel-like carnivorans (Musteloidea)

We analyzed a concatenated (8492 bp) nuclear-mitochondrial DNA data set from 44 musteloids (including the first genetic data for Lyncodon patagonicus) with parsimony, maximum likelihood, and Bayesian methods of phylogenetic and biogeographic inference and two Bayesian methods of chronological inference. Here we show that Musteloidea emerged approximately 32.4-30.9 million years ago (MYA) in Asia, shortly after the greenhouse-icehouse global climate shift at the Eocene-Oligocene transition. During their Oligocene radiation, which proceeded wholly or mostly in Asia, musteloids diversified into four primary divisions: the Mephitidae lineage separated first, succeeded by Ailuridae and the divergence of the Procyonidae and Mustelidae lineages. Mustelidae arose approximately 16.1 MYA within the Mid-Miocene Climatic Optimum, and extensively diversified in the Miocene, mostly in Asia. The early offshoots of this radiation largely evolved into badger and marten ecological niches (Taxidiinae, Melinae, Mellivorinae, Guloninae, and Helictidinae), whereas the later divergences have adapted to other niches including those of weasels, polecats, minks, and otters (Mustelinae, Ictonychinae, and Lutrinae). Notably, and contrary to traditional beliefs, the morphological adaptations of badgers, martens, weasels, polecats, and minks each evolved independently more than once within Mustelidae. Ictonychinae (which is most closely related to Lutrinae) arose approximately 9.5-8.9 MYA, most likely in Asia, where it diverged into the Old World Ictonychini (Vormela, Poecilictis, Ictonyx, and Poecilogale) and New World Lyncodontini (Lyncodon and Galictis) lineages. Ictonychini presumably entered Africa during the Messinian Salinity Crisis (at the Miocene-Pliocene transition), which interposed the origins of this clade (approximately 6.5-6.0 MYA) and its African Poecilictis-Ictonyx-Poecilogale subclade (approximately 4.8-4.5 MYA). Lyncodontini originated approximately 2.9-2.6 MYA at the Pliocene-Pleistocene transition in South America, slightly after the emergence of the Panamanian land bridge that provided for the Great American Biotic Interchange. As the genera Martes and Ictonyx (as currently circumscribed) are paraphyletic with respect to the genera Gulo and Poecilogale, respectively, we propose that Pekaniaand Poecilictis be treated as valid genera and that "Martes"pennanti and "Ictonyx"libyca, respectively, be assigned to these genera.     

Phylogenetic systematics and tempo of evolution of the Viverrinae (Mammalia, Carnivora, Viverridae) within feliformians: implications for faunal exchanges between Asia and Africa

We reconstructed the phylogeny of the subfamily Viverrinae (Mammalia, Carnivora, Viverridae) using a approximately 3kb data set in order to reassess timing and patterns of faunal exchanges between Asia and Africa. Maximum parsimony, maximum likelihood, and Bayesian analyses of separated and combined matrices (cytochrome b, transthyretin intron I and IRBP exon 1 [IRBP1]) recovered all the well-supported relationships within feliformian lineages. In addition, IRBP1 supported paraphyly of genus Herpestes and contributed to the resolution of equivocal hypotheses within Viverridae, including (1) the monophyly of Viverrinae, and (2) Viverricula sister-group of the other terrestrial civets (Civettictis and Viverra). The combined analysis yielded a robust phylogeny, recovering monophyly of Prionodontidae and yielding high posterior probabilities for nodes (1) (Prionodontidae, Felidae) and (2) ((Felidae, Prionodontidae), ((Hyaenidae, (Herpestidae, Eupleridae)), Viverridae)). Using a fossil cross-validation method, we estimated the emergence of Viverridae at 34.29Myr, with a separation between the three traditional subfamilies Hemigalinae, Paradoxurinae, and Viverrinae during the Late Oligocene-Early Miocene. The terrestrial civets and the splits between (1) Civettictis and Viverra and (2) Poiana and Genetta were estimated to appear during the Middle Miocene. Parsimony- and maximum likelihood-based methods yielded unambiguous ancestral area reconstructions, including the Asian origin of the family Viverridae, the subfamily Viverrinae, the terrestrial civets and the clade (Civettictis, Viverra). On the grounds of genetic distances, morphological divergence, and divergence time estimates, we propose the erection of the subfamily Genettinae (including Genetta and Poiana). Our analyses suggested two independent migration events from Asia to Africa, during the Middle Miocene (Civettictis) and between the Late Oligocene and Middle Miocene (Genettinae). These results are in agreement with the hypothesis of Miocene routes from Asia to Africa-via the Arabian microplate-that would have involved several independent events of migrations. Couched in the context of the viverrid fossil record, our study calls for a revision of the paleontological data in order to fully appreciate the complexity of Afro-Asian faunal exchanges.     

A new basal caniform (Mammalia: Carnivora) from the middle Eocene of North America and remarks on the phylogeny of early carnivorans

Background

Despite a long history of research, the phylogenetic origin and initial diversification of the mammalian crown-group Carnivora remain elusive. Well-preserved fossil materials of basal carnivorans are essential for resolving these issues, and for constraining the timing of the carnivoran origin, which constitutes an important time-calibration point in mammalian phylogenetics.

Methodology/Principal Findings

A new carnivoramorphan from the middle Eocene of southern California, Lycophocyon hutchisoni, is described. The new taxon exhibits stages of dental and basicranial evolution that are intermediate between earlier carnivoramorphans and the earliest representatives of canoid carnivorans. The evolutionary affinity of the new taxon was determined by a cladistic analysis of previously-published and newly-acquired morphological data for 30 Paleogene carnivoramorphans. The most-parsimonious trees identified L. hutchisoni as a basal caniform carnivoran, and placed (1) Tapocyon robustus, Quercygale angustidens, “Miacis” sylvestris, “M.” uintensis, and “M.” gracilis inside or outside the Carnivora, (2) nimravids within the Feliformia, and (3) the amphicyonid Daphoenus outside the crown-group Canoidea. Parsimony reconstructions of ancestral character states suggest that loss of the upper third molars and development of well-ossified entotympanics that are firmly fused to the basicranium (neither condition is observed in L. hutchisoni) are not associated with the origin of the Carnivora as traditionally thought, but instead occurred independently in the Caniformia and the Feliformia. A discriminant analysis of the estimated body weight and dental ecomorphology predicted a mesocarnivorous diet for L. hutchisoni, and the postcranial morphology suggests a scansorial habit.

Conclusions/Significance

Lycophocyon hutchisoni illuminates the morphological evolution of early caniforms leading to the origin of crown-group canoids. Considerable uncertainty remains with respect to the phylogenetic origin of the Carnivora. The minimum date of caniform-feliform divergence is provisionally suggested to be either 47 million years ago or 38 million years ago, depending on the position of “Miacis” sylvestris within or outside the Carnivora, respectively.     

A Total Evidence Phylogeny of the Arctoidea (Carnivora: Mammalia): Relationships Among Basal Taxa

A total evidence phylogenetic analysis was performed for 14 extant and 18 fossil caniform genera using a data matrix of 5.6 kbp of concatenated sequence data from six independent loci and 80 morphological characters from the cranium and dentition. Maximum parsimony analysis recovered a single most parsimonious cladogram (MPC). The topology of the extant taxa in the MPC agreed with previous molecular phylogenies. Phylogenetic positions for fossil taxa indicate that several taxa previously described as early members of extant families (e.g., Bathygale and Plesictis) are likely stem taxa at the base of the Arctoidea. Taxa in the “Paleomustelidae” were found to be paraphyletic, but a monophyletic Oligobuninae was recovered within this set of taxa. This clade was closely related to the extant genera Gulo and Martes, therefore, nested within the extant radiation of the family Mustelidae. This analysis provides a resolution to several discrepancies between phylogenies considering either fossil taxa or extant taxa separately, and provides a framework for incorporating fossil and extant taxa into comprehensive combined evidence analyses.