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Molecular systematics of the Canidae.

Despite numerous systematic studies, the relationships among many species within the dog family, Canidae, remain unresolved. Two problems of broad evolutionary significance are the origins of the taxonomically rich canidae fauna of South America and the development in three species of the trenchant heel, a unique meat-cutting blade on the lower first molar. The first problem is of interest because the fossil record provides little evidence for the origins of divergent South American species such as the maned wolf and the bush dog. The second issue is problematic because the trenchant heel, although complex in form, may have evolved independently to assist in the processing of meat. We attempted to resolve these two issues and five other specific taxonomic controversies by phylogenetic analysis of 2,001 base pairs of mitochondrial DNA (mtDNA) sequence data from 23 canidae species. The mtDNA tree topology, coupled with data from the fossil record, and estimates of rates of DNA sequence divergence suggest at least three and possibly four North American invasions of South America. This result implies that an important chapter in the evolution of modern canids remains to be discovered in the fossil record and that the South American canidae endemism is as much the result of extinction outside of South America as it is due to speciation within South America. The origin of the trenchant heel is not well resolved by our data, although the maximum parsimony tree is weakly consistent with a single origin followed by multiple losses of the character in several extant species. A combined analysis of the mtDNA data and published morphological data provides unexpected support for a monophyletic South American canidae clade. However, the homogeneity partition tests indicate significant heterogeneity between the two data sets.

Animals↗

The pattern of phylogenomic evolution of the Canidae.

Canidae species fall into two categories with respect to their chromosome composition: those with high numbered largely acrocentric karyotypes and others with a low numbered principally metacentric karyotype. Those species with low numbered metacentric karyotypes are derived from multiple independent fusions of chromosome segments found as acrocentric chromosomes in the high numbered species. Extensive chromosome homology is apparent among acrocentric chromosome arms within Canidae species; however, little chromosome arm homology exists between Canidae species and those from other Carnivore families. Here we use Zoo-FISH (fluorescent in situ hybridization, also called chromosomal painting) probes from flow-sorted chromosomes of the Japanese raccoon dog (Nyctereutes procyonoides) to examine two phylogenetically divergent canids, the arctic fox (Alopex lagopus) and the crab-eating fox (Cerdocyon thous). The results affirm intra-canid chromosome homologies, also implicated by G-banding. In addition, painting probes from domestic cat (Felis catus), representative of the ancestral carnivore karyotype (ACK), and giant panda (Ailuropoda melanoleuca) were used to define primitive homologous segments apparent between canids and other carnivore families. Canid chromosomes seem unique among carnivores in that many canid chromosome arms are mosaics of two to four homology segments of the ACK chromosome arms. The mosaic pattern apparently preceded the divergence of modern canid species since conserved homology segments among different canid species are common, even though those segments are rearranged relative to the ancestral carnivore genome arrangement. The results indicate an ancestral episode of extensive centric fission leading to an ancestral canid genome organization that was subsequently reorganized by multiple chromosome fusion events in some but not all Canidae lineages.

Animals↗

Immunogenetic evidence for the phylogenetic sister group relationship of dogs and bears (Mammalia, Carnivora: Canidae and Crsidae). a comparative determinant analysis of carnivoran albumin, c3 complement and immunoglobulin micro-chain.

Thirty-seven antigenic determinants were identified in the albumins, the immunoglobulin micro- and IgG(Fc) chains, and the C3 proteins of 51 carnivoran (sub)species from 31 genera, and in 12 noncarnivoran mammals. In addition to 19 determinants plesiomorphic for Carnivora as an order, 18 synapomorphic epitopes of carnivoran families revealed nine phylogenetic reaction groups: (1) canids, (2) ursids, (3) the racoon, (4) the Weddell seal, (5) the lesser panda, (6) the harbour seal, (7) mustelids, (8) viverrids and hyaenas, and (9) felids. These data identify Canoidea (Canidae, Ursidae, Phocidae, Procyonidae, Ailuridae, Mustelidae) and Feloidea (Viverridae, Hyaenidae, Felidae) as two fundamentally differentiated lineages of Carnivora, and confirm the inclusion of seals among the former. The Ursidae are the sister group of the Canidae. The antigenic determinants in the studied proteins do not subdivide the Canidae, Ursidae and Felidae into immunologically differentiated lineages.

Albumins↗

[Infectious pathology of Canidae and Felidae in zoological parks].

The Canidae (36 species) and Felidae (34-37 species) are two families of carnivores represented by numerous exotic species in zoos or wildlife reserves. To some extent, the diseases of these species are similar to those of dogs and cats, and are therefore relatively well known. However, there are differences in sensitivity to infectious agents, treatments and vaccines. Canidae and Felidae may also act as carriers or even vectors of zoonoses, such as leptospirosis, rabies, salmonellosis, toxoplasmosis and tuberculosis. Due to their behaviour patterns and morphological adaptations, these species are capable of transmitting various opportunistic infections by biting or scratching. These characteristics mean that Canidae and Felidae are difficult to keep in captivity, and require special health precautions, particularly protection from contact with stray carnivores.

Animals↗

Isolation and molecular evolution of the selenocysteine tRNA (Cf TRSP) and RNase P RNA (Cf RPPH1) genes in the dog family, Canidae.

In an effort to identify rapidly evolving nuclear sequences useful for phylogenetic analyses of closely related species, we isolated two genes transcribed by RNA polymerase III (pol III), the selenocysteine tRNA gene (TRSP) and an RNase P RNA (RPPH1) gene from the domestic dog (Canis familiaris). We focus on genes transcribed by pol III because their coding regions are small (generally 100-300 base pairs [bp]) and their essential promoter elements are located within a couple of hundred bps upstream of the coding region. Therefore, we predicted that regions flanking the coding region and outside of the promoter elements would be free of constraint and would evolve rapidly. We amplified TRSP from 23 canids and RPPH1 from 12 canids and analyzed the molecular evolution of these genes and their utility as phylogenetic markers for resolving relationships among species in Canidae. We compared the rate of evolution of the gene-flanking regions to other noncoding regions of nuclear DNA (introns) and to the mitochondrial encoded COII gene. Alignment of TRSP from 23 canids revealed that regions directly adjacent to the coding region display high sequence variability. We discuss this pattern in terms of functional mechanisms of transcription. Although the flanking regions evolve no faster than introns, both genes were found to be useful phylogenetic markers, in part, because of the synapomorphic indels found in the flanking regions. Gene trees generated from the TRSP and RPPH1 loci were generally in agreement with the published mtDNA phylogeny and are the first phylogeny of Canidae based on nuclear sequences.

Animals↗

[Formation sources and topographic characteristics of the pelvic plexus in furry animals of Canidae and Mustelidae families].

Using preparation methods, the topography and the formation sources of pelvic plexus (PP) were studied in animals of Canidae (silver-black fox, blue fox) and Mustelidae (sable, American mink) families. Maximal differences in PP topography and PP-forming nerves were found between species (Canidae-Mustelidae), whereas the intraspecies differences were less pronounced. Intraspecies sex-associated differences in PP topography are associated with only quantitative parameters, while those in PP formation sources and in peculiarities of PP-forming nerves' branching pattern are qualitative in nature. There is a distinct correlation between PP structural type (diffuse, compact, mixed) and peculiarities of its ganglionic apparatus: in diffuse type the juxta- and intraogan position of ganglia is prevalent, while in compact type the extraorgan position is more common.

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Comparison of the nuclear organiser region activity in four taxa of the family Canidae.

Four species of the family Canidae were cytogenetically studied. The activity of NORs was detected with the use of silver staining. The number of NORs was characteristic for a given karyotype. For the dog found on autosomes 7, 17, 20 and on sex chromosome (Y), for the racoon dog on autosomes 1, 4, 13 and on sex chromosome (Y), for the silver fox only on autosomes 8, 9, 13 and for the blue fox on autosomes 13, 15, 17, 18, 20 and 22. The results demonstrate that NOR activity is similar in all the analysed species of the Canidae. Simultaneously, NOR activity for a medium-sized chromosome pair is distinctly higher than for two other autosome pairs (the longest and the smallest pair). Considerable variability was observed within individuals.

Journal Article↗

Comparative anatomy of the cardiac foramen ovale in cats (Felidae), dogs (Canidae), bears (Ursidae) and hyaenas (Hyaenidae).

The structure of the foramen ovale from 16 species representing 4 carnivore families, the Felidae, Canidae, Ursidae and Hyaenidae, was studied using the scanning electron microscope. The Felidae were represented by 9 domestic cat fetuses (Felis catus), 2 snow leopard neonates (Uncia uncia), an ocelot neonate (Leopardus pardalis), 2 lion neonates (Panthera leo), a panther neonate (Panthera pardus) and 3 tigers (Neofelis tigris), comprising 2 fetuses and a neonate. The Canidae were represented by a golden jackal neonate (Canis aureus), a newborn wolf (Canis lupus), 8 domestic dog fetuses (Canis familiaris), 3 red fox neonates (Vulpes vulpes) and a dhole neonate (Cuon alpinus). The Ursidae were represented by a brown bear neonate (Ursus arctos), a day-old grizzly bear cub (Ursus arctos horribilis), a polar bear neonate (Ursus maritimus), and 2 additional bear fetuses (species unknown). The Hyaenidae were represented by a striped hyaena neonate (Hyaena hyaena). In each species, the foramen ovale, when viewed from the terminal part of the caudal vena cava, had the appearance of a short tunnel. A thin fold of tissue, the developed remains of the embryonic septum primum, extended from the distal end of the caudal vena cava for a variable distance into the lumen of the left atrium and contributed towards the 'tunnel' appearance in all specimens. It constituted a large proportion of the tube, and its distal end was straight-edged. There was fibrous material underlying the endothelium of the flap, the apparent morphology of which suggested that it comprised cardiac muscle.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

[Modular evolution of Bsp-repeats: formation of subrepeats and monomers preceded the divergence of four species of Canidae].

Determination of two nucleotide sequences of satellite-like DNA (Bsp-repeats) segments from the raccoon-like dog genome and analysis of these and three other homologous sequences from distantly related genomes of Canidae (silver fox, domestic dog, and gray fox) show that all these DNA fragments consist of five types of subrepeats (A, B, C, D, and E) and two types of monomers-ACB and ACED. Raccoon-like dog repeated segment of 1620 bp is composed of both type of monomers: ACBACED. Different monomers of the same type could be distinguished by subtypes of subrepeats. Building of a cladogram of 18 subrepeats and analysis of the structures of monomers and higher-order repeated units allowed us to reveal probable evolution events which have determined the hierarchical order in Bsp-repeat organization. Approximate periods of time for these evolution events were calculated from the number of substitutions per branch. Five main types of subrepeats and two types of monomers have been generated and propagated through several steps before divergence of the main phylogenetic trees of Canidae. Combinatorial variability appears to play a fundamental role in forming the hierarchical layers in Bsp-repeats structure.

Animals↗

A molecular phylogeny of the Canidae based on six nuclear loci.

We have reconstructed the phylogenetic relationships of 23 species in the dog family, Canidae, using DNA sequence data from six nuclear loci. Individual gene trees were generated with maximum parsimony (MP) and maximum likelihood (ML) analysis. In general, these individual gene trees were not well resolved, but several identical groupings were supported by more than one locus. Phylogenetic analysis with a data set combining the six nuclear loci using MP, ML, and Bayesian approaches produced a more resolved tree that agreed with previously published mitochondrial trees in finding three well-defined clades, including the red fox-like canids, the South American foxes, and the wolf-like canids. In addition, the nuclear data set provides novel indel support for several previously inferred clades. Differences between trees derived from the nuclear data and those from the mitochondrial data include the grouping of the bush dog and maned wolf into a clade with the South American foxes, the grouping of the side-striped jackal (Canis adustus) and black-backed jackal (Canis mesomelas) and the grouping of the bat-eared fox (Otocyon megalotis) with the raccoon dog (Nycteruetes procyonoides). We also analyzed the combined nuclear+mitochondrial tree. Many nodes that were strongly supported in the nuclear tree or the mitochondrial tree remained strongly supported in the nuclear+mitochondrial tree. Relationships within the clades containing the red fox-like canids and South American canids are well resolved, whereas the relationships among the wolf-like canids remain largely undetermined. The lack of resolution within the wolf-like canids may be due to their recent divergence and insufficient time for the accumulation of phylogenetically informative signal.

Animals↗

Variation of short tandem repeats within and between species belonging to the Canidae family.

Frequency distribution and allele size in 20 canine microsatellite loci were analyzed in 33 flat-coated retrievers, 32 dachshunds, 10 red foxes, and 10 Arctic foxes. Overall, the major difference between the two dog breeds was the relative allele frequencies rather than the size ranges of alleles at the individual locus. The average heterozygosity within the two dog breeds was not significantly different. Since the average heterozygosity at several polymorphic loci is a relative measure of heterogeneity within the population, analysis of heterozygosity within microsatellite loci is suggested as a measure for the diversity of populations. Eighty percent (16 of 20) of the canine microsatellite primer pairs amplified corresponding loci in the two fox species. This reflects a very high sequence conservation within the Canidae family relative to findings in, for instance, the Muridae family. This indicates that it will be possible to utilize the well-characterized fox karyotype instead of the dog karyotype as a step towards physical mapping of the dog genome. Analysis of exclusion power and probabilities of genetic identity between unrelated animals by use of the seven most informative loci demonstrated that it will be possible to assemble a panel of microsatellite loci that is effective for parentage analysis in all breeds.

Alleles↗

Lectin histochemistry of snout skin and foot pads in the wolf and the domesticated dog (Mammalia: Canidae).

The distribution and selectivity of complex carbohydrates in the snout skin and the foot pads of the wolf and the domesticated dog were studied by means of light-microscopic histochemical methods, particularly lectin histochemistry. In the snout, moderate to strong staining reactions were confined to intercellular substances among the epidermal cells, containing neutral glycoconjugates (glycoproteins, glycolipids) with various saccharide residues (alpha-D-mannose, beta-N-acetyl-D-glucosamine, alpha-N-acetyl-D-galactosamine, beta-D-galactose). In the foot pads, distinct reactions were found in neutral glycoconjugates of intercellular substances of the stratum corneum, with alpha-N-acetyl-D-galactosaminyl and beta-N-acetyl-D-glucosaminyl residues, but only in the wolf, whereas in the dog such substances were concentrated in the spinosal and basal epidermal layers, additionally marked by D-galactose. The eccrine glands exhibited high amounts of mainly neutral glycoconjugates in the secretory portion, especially in the wolf. A clear spectrum of saccharide residues was demonstrable in the dark cells (alpha-D-mannose, beta-N-acetyl-D-glucosamine, alpha-N-acetyl-D-galactosamine, alpha-D-galactose), and in the clear cells which lacked mannose and contained beta-D-galactose. In contrast to the wolf, strong reactions for glycoconjugates were visible among intercellular substances of intradermal excretory duct cells in the dog. Only in the wolf did the periphery of the intracorneal excretory ducts and the outer surface of the foot pads show clearly positive reactions for glycoconjugates. The results obtained are discussed with regard to the specific functions of the body regions investigated and the behavioural biology of the Canidae. It is obvious that differences in amounts, quality, and localization of the glycoconjugates produced exist between the wolf and its domesticated descendant.

Animals↗

Natural spillover of a distinctly Canidae-associated biotype of rabies virus into an expanded wildlife host range in southern Africa.

Rabies enzootics in southern Africa are associated with two genetically distinct groups of viruses, thought to be adapted to two different sets of host species. The virus groups are referred to as the canid biotype (infecting carnivores of the family Canidae) and the viverrid biotype (infecting carnivores of the subfamily Viverrinae). Cross- or spillover infections of one biotype into the host range of the other are thought to occur from time to time. However, very little is known about this phenomenon and its role in the epidemiology of rabies in southern Africa. We have investigated spillover by monoclonal antibody and nucleic acid sequence analysis of a wide range of virus isolates. Although the inverse had been documented, this report constitutes the first evidence of spillover of canid biotype viruses into viverrid hosts. Our genetic analysis was focused specifically on the G-L intergenic region of the virus genome, thought to be a remnant or pseudogene and it was indicated that, with respect to this region of the genome, spillover does not influence the phylogeny of virus isolates.

Animals↗

Inheritance of litter size at birth in farmed arctic foxes (Alopex lagopus, Canidae, Carnivora).

Natural populations of the arctic fox (Alopex lagopus, Canidae, Carnivora) differ drastically in their reproductive strategy. Coastal foxes, which depend on stable food resources, produce litters of moderate size. Inland foxes feed on small rodents, whose populations are characterized by cycling fluctuation. In the years with low food supply, inland fox populations have a very low rate of reproduction. In the years with high food supply, they undergo a population explosion. To gain insight into the genetic basis of the reproductive strategy of this species, we performed complex segregation analysis of the litter size in the extended pedigree of the farmed arctic foxes involving 20,665 interrelated animals. Complex segregation analysis was performed using a mixed model assuming that the trait was under control of a major gene and a large number of additive genetic and random factors. To check the significance of any major gene effect, we used Elston-Stewart transmission probability test. Our analysis demonstrated that the inheritance of this trait can be described within the frameworks of a major gene model with recessive control of low litter size. This model was also supported by the pattern of its familial segregation and by comparison of the distributions observed in the population and that expected under our model. We suggest that a system of balanced polymorphism for litter size in the farmed population might have been established in natural populations of arctic foxes as a result of adaptation to the drastic fluctuations in prey availability.

Animals↗

Comparative chromosomal localization of the canine-derived BAC clones containing LEP and IGF1 genes in four species of the family Canidae.

In the present report we show the chromosomal localization of two BAC clones, carrying the leptin (LEP) and insuline-like growth factor 1 (IGF1) genes, respectively, in four species belonging to the family Canidae: the dog, red fox, arctic fox and the Chinese raccoon dog. The assignments are in agreement with earlier data obtained from comparative chromosome painting for the dog, red fox and arctic fox.

Animals↗

Chromosomal evolution of the Canidae. I. Species with high diploid numbers.

The Giemsa banding patterns of seven canid species, including the grey wolf (Canis lupus), the maned wolf (Chrysocyon brachyurus), the bush dog (Speothos venaticus), the crab-eating fox (Cerdocyon thous), the grey fox (Urocyon cinereoargenteus), the bat-eared fox (Otocyon megalotis), and the fennec (Fennecus zerda), are presented and compared. Relative to other members of Canidae, these species have high diploid complements (2n greater than 64) consisting of largely acrocentric chromosomes. They show a considerable degree of chromosome homoeology, but relative to the grey wolf, each species is either missing chromosomes or has unique chromosomal additions and rearrangements. Differences in chromosome morphology among the seven species were used to reconstruct their phylogenetic history. The results suggest that the South American canids are closely related to each other and are derived from a wolf-like progenitor. The fennec and the bat-eared fox seem to be recent derivatives of a lineage that branched early from the wolf-like canids and which also includes the grey fox.

Animals↗