Effects of Coyote Population Reduction on Swift Fox Demographics in Southeastern Colorado

ABSTRACT The distribution and abundance of swift foxes (Vulpes velox) has declined from historic levels. Causes for the decline include habitat loss and fragmentation, incidental poisoning, changing land use practices, trapping, and predation by other carnivores. Coyotes (Canis latrans) overlap the geographical distribution of swift foxes, compete for similar resources, and are a significant source of mortality amongst many swift fox populations. Current swift fox conservation and management plans to bolster declining or recovering fox populations may include coyote population reduction to decrease predation. However, the role of coyote predation in swift fox population dynamics is not well-understood. To better understand the interactions of swift foxes and coyotes, we compared swift fox population demographics (survival rates, dispersal rates, reproduction, density) between areas with and without coyote population reduction. On the Piñon Canyon Maneuver Site, Colorado, USA, we monitored 141 swift foxes for 65,226 radio-days from 15 December 1998 to 14 December 2000 with 18,035 total telemetry locations collected. Juvenile swift fox survival rate was increased and survival was temporarily prolonged in the coyote removal area. Adult fox survival patterns were also altered by coyote removal, but only following late-summer coyote removals and, again, only temporarily. Coyote predation remained the main cause of juvenile and adult fox mortality in both areas. The increase in juvenile fox survival in the coyote removal area resulted in a compensatory increase in the juvenile dispersal rate and an earlier pulse in dispersal movements. Adult fox dispersal rate was more consistent throughout the year in the coyote removal area. Coyote removal did not influence the reproductive parameters of the swift foxes. Even though juvenile survival increased, swift fox density remained similar between the areas due to the compensatory dispersal rate among juvenile foxes. We concluded that the swift fox population in the area was saturated. Although coyote predation appeared additive in the juvenile cohort, it was compensatory with dispersal.     

Phylogenetic relationships of the fox-like canids: mitochondrial DNA restriction fragment, site and cytochrome b sequence analyses

The fox-like canids include taxa from the genera Alopex, Otocyon, Fennecus, Urocyon and Vulpes . Previous morphological analysis indicated that species from the latter three genera are very similar and should be included in the same genus whereas Alopex and Otocyon are sufficiently different to be included in separate genera. Using phylogenetic methods, we analyse mitochondrial DNA (mtDNA) restriction fragment and restriction site data, and 402 bp of cytochrome b sequence variation in fox-like canids. Our results suggest that Alopex lagopus , the arctic fox, is actually a very close relative of the swift fox, a species in the genus Vulpes . Similarly, the fennec, Fennecus zerda is related to the co-existing desert species, the Blanford's fox, Vulpes cana . The grey fox, Urocyon cinereoargenteus , and the bat-eared fox, Otocyon megalotis , are not closely related to each other or to any of the sampled fox taxa. Our results indicate that desert adaptations have evolved independently at least twice in the Canidae, and that Pleistocene glaciations and character divergence may be important causes of morphological change in canids.     

Genetic diversity in a reintroduced swift fox population

Swift fox (Vulpes velox) were historically distributed in southwestern South Dakota including the region surrounding Badlands National Park (BNP). The species declined during the mid-1800s, largely due to habitat loss and poisoning targeted at wolves (Canis lupus) and coyotes (Canis latrans). Only a small population of swift foxes near Ardmore, which is located in Fall River County, South Dakota, persisted. In 2003, a reintroduction program was initiated at BNP with swift foxes translocated from Colorado and Wyoming. Foxes released in the years 2003, 2004 and 2005 were translocated from Colorado (BNP-Colorado) whereas in 2006, released foxes were translocated from Wyoming (BNP-Wyoming). Our objective was to evaluate genetic diversity and structure of the restored swift fox population in the area surrounding BNP compared to source fox populations in an area of Colorado and Wyoming, as well as the local swift fox population neighboring BNP near Ardmore in Fall River County, South Dakota. A total of 400 swift foxes (28 released in 2003, 28 released in 2004, 26 released in 2005, 26 released in 2006, 252 wild-born foxes, 40 individual foxes from the Ardmore area of South Dakota) was genotyped using twelve microsatellite loci. We report mean gene diversity values of 0.778 (SD = 0.156) for the BNP-Colorado population, 0.753 (SD = 0.165) for the BNP-Wyoming population, 0.751 (SD = 0.171) for the BNP population, and 0.730 (SD = 0.166) for the Fall River population. We also obtained F_st values ranging from 0.014 to 0.029 for pair-wise comparisons of fox populations (BNP, Fall River, BNP-Wyoming, BNP-Colorado). We conclude that the reintroduced fox population around BNP has high genetic diversity comparable to its source populations in Colorado and Wyoming. Although genetic diversity indicates that the reintroduction was successful, additional time is necessary to fully evaluate long-term genetic maintenance and interconnectivity among these populations.     

Mitochondrial DNA phylogeny and morphological diversity in the genus Mastus (Beck, 1837): a study in a recent (Holocene) island group (Koufonisi, south-east Crete)

Three endemic Cretan land snail species of the genus Mastus (Beck, 1837) from the island group of Koufonisi (south-east Crete) and the eastern part of Crete, were studied by multivariate analysis of shell morphology and analysis of mtDNA sequences. The phylogeny of the populations studied and the processes effecting the genetic and morphological diversity of the species were investigated. Extremely high mtDNA sequence divergence was observed both within and between populations. The Cretan populations could not be distinguished morphologically, while the populations of the islets were more distinct. We argue that the active geological past of the area (including sea level changes) and the long-term presence of humans has produced a mixing up of Mastus populations leading to the accumulation of high divergence of mtDNA sequences on a small spatial scale. The limited morphological diversity and the distinct shell 'identity' of the islets' populations can be attributed to the selective pressures of the island group.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 383–399.     

Adaptive divergence despite strong genetic drift: genomic analysis of the evolutionary mechanisms causing genetic differentiation in the island fox (Urocyon littoralis)

The evolutionary mechanisms generating the tremendous biodiversity of islands have long fascinated evolutionary biologists. Genetic drift and divergent selection are predicted to be strong on islands and both could drive population divergence and speciation. Alternatively, strong genetic drift may preclude adaptation. We conducted a genomic analysis to test the roles of genetic drift and divergent selection in causing genetic differentiation among populations of the island fox (Urocyon littoralis). This species consists of six subspecies, each of which occupies a different California Channel Island. Analysis of 5293 SNP loci generated using Restriction‐site Associated DNA (RAD) sequencing found support for genetic drift as the dominant evolutionary mechanism driving population divergence among island fox populations. In particular, populations had exceptionally low genetic variation, small N_e (range = 2.1–89.7; median = 19.4), and significant genetic signatures of bottlenecks. Moreover, islands with the lowest genetic variation (and, by inference, the strongest historical genetic drift) were most genetically differentiated from mainland grey foxes, and vice versa, indicating genetic drift drives genome‐wide divergence. Nonetheless, outlier tests identified 3.6–6.6% of loci as high F_ST outliers, suggesting that despite strong genetic drift, divergent selection contributes to population divergence. Patterns of similarity among populations based on high F_ST outliers mirrored patterns based on morphology, providing additional evidence that outliers reflect adaptive divergence. Extremely low genetic variation and small N_e in some island fox populations, particularly on San Nicolas Island, suggest that they may be vulnerable to fixation of deleterious alleles, decreased fitness and reduced adaptive potential.     

Effects of Habitat on Competition Between Kit Foxes and Coyotes

Abstract: San Joaquin kit foxes (Vulpes macrotis mutica) are an endangered species with a narrow geographic range whose natural populations are limited by predation by coyotes (Canis latrans). In the warm, arid grassland and shrubland habitats where kit foxes occur, coyotes are more cover dependent than kit foxes, creating the possibility of habitat segregation. Effects of habitat variation on coyote and kit fox competition are unknown. We assessed exploitation and interference competition between coyotes and kit foxes in grassland and shrubland habitats to determine if such competition varies among habitats. With respect to exploitation competition, we evaluated habitat and spatial partitioning, diet, prey abundance, and survival for kit foxes and coyotes at the Lokern Natural Area in central California, USA, from January 2003 through June 2004. Kit foxes partitioned habitat, space, and diet with coyotes. Coyotes primarily used shrubland habitats whereas kit foxes selectively used burned grasslands. Kit foxes and coyotes had high dietary overlap with regards to items used, but proportional use of items differed between the 2 species. Kit foxes selected for Heermann's kangaroo rats (Dipodomys heermanni), which were closely tied to shrub habitats. With respect to interference competition, predation was the primary source of mortality for kit foxes, and survival of individual kit foxes was inversely related to proportion of shrub habitat within their home ranges. Our results suggest that a heterogeneous landscape may benefit kit foxes by providing habitat patches where predation risk may be lower.     

A MORPHOLOGIC AND GENETIC STUDY OF THE ISLAND FOX,UROCYON LITTORALIS

The Island Fox, Urocyon littoralis, is a dwarf form found on six of the Channel Islands located 30–98 km off the coast of southern California. The island populations differ in two variables that affect genetic variation: effective population size and duration of isolation. We estimate that the effective population size of foxes on the islands varies from approximately 150 to 1,000 individuals. Archeological and geological evidence suggests that foxes likely arrived on the three northern islands minimally 10,400–16,000 years ago and dispersed to the three southern islands 2,200–4,300 years ago. We use morphometrics, allozyme electrophoresis, mitochondrial DNA (mtDNA) restriction-site analysis, and analysis of hypervariable minisatellite DNA to measure variability within and distances among island fox populations. The amount of within-population variation is lowest for the smallest island populations and highest for the mainland population. However, the larger populations are sometimes less variable, with respect to some genetic measures, than expected. No distinct trends of variability with founding time are observed. Genetic distances among the island populations, as estimated by the four techniques, are not well correlated. The apparent lack of correspondence among techniques may reflect the effects of mutation rate and colonization history on the values of each genetic measure.     

SPECIATION AND POPULATION GENETIC STRUCTURE IN TROPICAL PACIFIC SEA URCHINS

Unlike populations of many terrestrial species, marine populations often are not separated by obvious, permanent barriers to gene flow. When species have high dispersal potential and few barriers to gene flow, allopatric divergence is slow. Nevertheless, many marine species are of recent origin, even in taxa with high dispersal potential. To understand the relationship between genetic structure and recent species formation in high dispersal taxa, we examined population genetic structure among four species of sea urchins in the tropical Indo-West Pacific that have speciated within the past one to three million years. Despite high potential for gene flow, mtDNA sequence variation among 200 individuals of four species in the urchin genus Echinometra shows a signal of strong geographic effects. These effects include (1) substantial population heterogeneity; (2) lower genetic variation in peripheral populations; and (3) isolation by distance. These geographic patterns are especially strong across scales of 5000-10,000 km, and are weaker over scales of 2500-5000 km. As a result, strong geographic patterns would not have been readily visible except over the wide expanse of the tropical Pacific. Surface currents in the Pacific do not explain patterns of gene flow any better than do patterns of simple spatial proximity. Finally, populations of each species tend to group into large mtDNA regions with similar mtDNA haplotypes, but these regional boundaries are not concordant in different species. These results show that all four species have accumulated mtDNA differences over similar spatial and temporal scales but that the precise geographic pattern of genetic differentiation varies for each species. These geographic patterns appear much less deterministic than in other well-known coastal marine systems and may be driven by chance and historical accident.     

Genotyping-By-Sequencing (GBS) Detects Genetic Structure and Confirms Behavioral QTL in Tame and Aggressive Foxes (Vulpes vulpes)

The silver fox (Vulpes vulpes) offers a novel model for studying the genetics of social behavior and animal domestication. Selection of foxes, separately, for tame and for aggressive behavior has yielded two strains with markedly different, genetically determined, behavioral phenotypes. Tame strain foxes are eager to establish human contact while foxes from the aggressive strain are aggressive and difficult to handle. These strains have been maintained as separate outbred lines for over 40 generations but their genetic structure has not been previously investigated. We applied a genotyping-by-sequencing (GBS) approach to provide insights into the genetic composition of these fox populations. Sequence analysis of EcoT22I genomic libraries of tame and aggressive foxes identified 48,294 high quality SNPs. Population structure analysis revealed genetic divergence between the two strains and more diversity in the aggressive strain than in the tame one. Significant differences in allele frequency between the strains were identified for 68 SNPs. Three of these SNPs were located on fox chromosome 14 within an interval of a previously identified behavioral QTL, further supporting the importance of this region for behavior. The GBS SNP data confirmed that significant genetic diversity has been preserved in both fox populations despite many years of selective breeding. Analysis of SNP allele frequencies in the two populations identified several regions of genetic divergence between the tame and aggressive foxes, some of which may represent targets of selection for behavior. The GBS protocol used in this study significantly expanded genomic resources for the fox, and can be adapted for SNP discovery and genotyping in other canid species.     

Genetic variability among endangered Chinese giant salamanders, Andrias davidianus

The endangered Chinese giant salamander (Andrias davidianus) is endemic to mainland China. Genetic divergence among six populations of the species was investigated by means of isozyme electrophoresis and mitochondrial DNA (mtDNA) sequences. Forty allozyme loci were resolved for all populations; the amount of genetic divergence among populations was comparable to that in other amphibians. mtDNA sequences showed a similar level of divergence. The population from Huangshan is distinct from other populations, indicating the existence of localized divergence. Both allozyme and mtDNA data failed to associate the populations into a pattern corresponding to the three Chinese river systems, which may be the consequence of human relocation. Conservation policies should emphasize the protection of localized populations and cessation of human-facilitated introductions. Future studies should focus on investigating the divergence among localized populations from isolated mountain regions, particularly using more fine-grained techniques such as microsatellite DNA.     

Genetic and phenotypic variation among geographically isolated populations of the globally threatened Dupont's lark Chersophilus duponti

Identifying genetically and phenotypically distinct populations of threatened species is critical if we are to delineate appropriate plans for their conservation. We conducted an integrated analysis of population genetic structure, historical demographic events, current gene flow (all based on mtDNA sequences) and morphological variation of three geographically separated groups of populations of Dupont's lark Chersophilus duponti, located in the Iberian Peninsula (three populations), Morocco (two populations), and Tunisia (one population). Unusually, this lark species is the only one among the genus Chersophilus. Our results revealed the early historical divergence of an eastern Dupont's lark lineage (in Tunisia) and a western lineage (in Morocco and Spain), consistent with subspecies taxonomy and distribution. The western lineage subsequently split into two lineages, following the isolation of Iberian and African populations. Such pattern of historical differentiation caused great population genetic structure, with differences among geographic areas explaining more than 80% of total genetic variation. Mismatch distributions and coalescent estimates of divergence time showed that lineage divergence was associated with sudden population expansion events, which apparently took place during the last glaciation, when steppe habitats were widespread across the Mediterranean region. Extant populations from different geographic areas hardly shared any haplotype (only one out of 16 ND2 haplotypes was shared by Tunisian and Moroccan Dupont's larks), and consequently gene flow between geographic areas was found to be virtually absent. Apart from showing great genetic differentiation, Dupont's larks from different geographic areas were morphologically distinct, showing substantial variation in body size and feeding-related traits (length of feet and bill). We conclude that Dupont's lark populations isolated in the Iberian Peninsula, Morocco, and Tunisia are distinct evolutionary entities and should be considered as such in conservation plans. Such circumstance sets a daunting conservation challenge that exemplifies the need of incorporating knowledge of historical processes to our general understanding of the demography of threatened species.     

Population structure of two rabies hosts relative to the known distribution of rabies virus variants in Alaska

For pathogens that infect multiple species, the distinction between reservoir hosts and spillover hosts is often difficult. In Alaska, three variants of the arctic rabies virus exist with distinct spatial distributions. We tested the hypothesis that rabies virus variant distribution corresponds to the population structure of the primary rabies hosts in Alaska, arctic foxes (Vulpes lagopus) and red foxes (Vulpes vulpes) to possibly distinguish reservoir and spillover hosts. We used mitochondrial DNA (mtDNA) sequence and nine microsatellites to assess population structure in those two species. mtDNA structure did not correspond to rabies virus variant structure in either species. Microsatellite analyses gave varying results. Bayesian clustering found two groups of arctic foxes in the coastal tundra region, but for red foxes it identified tundra and boreal types. Spatial Bayesian clustering and spatial principal components analysis identified 3 and 4 groups of arctic foxes, respectively, closely matching the distribution of rabies virus variants in the state. Red foxes, conversely, showed eight clusters comprising two regions (boreal and tundra) with much admixture. These results run contrary to previous beliefs that arctic fox show no fine‐scale spatial population structure. While we cannot rule out that the red fox is part of the maintenance host community for rabies in Alaska, the distribution of virus variants appears to be driven primarily by the arctic fox. Therefore, we show that host population genetics can be utilized to distinguish between maintenance and spillover hosts when used in conjunction with other approaches.     

Landscape genetics of the nonnative red fox of California

Invasive mammalian carnivores contribute disproportionately to declines in global biodiversity. In California, nonnative red foxes (Vulpes vulpes) have significantly impacted endangered ground‐nesting birds and native canids. These foxes derive primarily from captive‐reared animals associated with the fur‐farming industry. Over the past five decades, the cumulative area occupied by nonnative red fox increased to cover much of central and southern California. We used a landscape‐genetic approach involving mitochondrial DNA (mtDNA) sequences and 13 microsatellites of 402 nonnative red foxes removed in predator control programs to investigate source populations, contemporary connectivity, and metapopulation dynamics. Both markers indicated high population structuring consistent with origins from multiple introductions and low subsequent gene flow. Landscape‐genetic modeling indicated that population connectivity was especially low among coastal sampling sites surrounded by mountainous wildlands but somewhat higher through topographically flat, urban and agricultural landscapes. The genetic composition of populations tended to be stable for multiple generations, indicating a degree of demographic resilience to predator removal programs. However, in two sites where intensive predator control reduced fox abundance, we observed increases in immigration, suggesting potential for recolonization to counter eradication attempts. These findings, along with continued genetic monitoring, can help guide localized management of foxes by identifying points of introductions and routes of spread and evaluating the relative importance of reproduction and immigration in maintaining populations. More generally, the study illustrates the utility of a landscape‐genetic approach for understanding invasion dynamics and metapopulation structure of one of the world's most destructive invasive mammals, the red fox.     

Population genetic structure of Bryde’s whales (<Emphasis Type="Italic">Balaenoptera brydei</Emphasis>) at the inter-oceanic and trans-equatorial levels

Bryde’s whales (Balaenoptera brydei) differ from other typical baleen whale species because they are restricted to tropical and warm temperate waters in major oceans, and frequent trans-equatorial movement has been suggested for the species. We tested this hypothesis by analyzing genetic variation at 17 microsatellite loci (N = 508) and 299 bp of mitochondrial DNA (mtDNA) control region sequences (N = 472) in individuals obtained from the western North Pacific, South Pacific, and eastern Indian Ocean. Combined use of microsatellite and mtDNA markers allowed us to distinguish between contemporary gene flow and ancestral polymorphism and to describe sex-specific philopatry. A high level of genetic diversity was found within the samples. Both nuclear and mtDNA markers displayed similar population structure, indicating a lack of sex-specific philopatry. Spatial structuring was detected using both frequency-based population parameters and individual-based Bayesian approaches. Whales in the samples from different oceanic regions came from genetically distinct populations with evidence of limited gene flow. We observed low mtDNA sequence divergence among populations and a lack of concordance between geographic and phylogenetic position of mtDNA haplotypes, suggesting recent separation of populations rather than frequent trans-equatorial and inter-oceanic movement. We conclude that current gene flow between Bryde’s whale populations is low and that effective management actions should treat them as separate entities to ensure continued existence of the species.     

Vicariance and dispersal across an intermittent barrier: population genetic structure of marine animals across the Torres Strait land bridge

Biogeographic barriers, some transitory in duration, are likely to have been important contributing factors to modern marine biodiversity in the Indo-Pacific region. One such barrier was the Torres Strait land bridge between continental Australia and New Guinea that persisted through much of the late Pleistocene and separated Indian and Pacific Ocean taxa. Here, we examine the patterns of mitochondrial DNA diversity for marine animals with present-day distributions spanning the Torres Strait. Specifically, we investigate whether there are concordant signatures across species, consistent with either vicariance or recent colonization from either ocean basin. We survey four species of reef fishes (Apogon doederleini, Pomacentrus coelestis, Dascyllus trimaculatus, and Acanthurus triostegus) for mtDNA cytochrome oxidase 1 and control region variation and contrast these results to previous mtDNA studies in diverse marine animals with similar distributions. We find substantial genetic partitioning (estimated from F-statistics and coalescent approaches) between Indian and Pacific Ocean populations for many species, consistent with regional persistence through the late Pleistocene in both ocean basins. The species-specific estimates of genetic divergence, however, vary greatly and for reef fishes we estimate substantially different divergence times among species. It is likely that Indian and Pacific Ocean populations have been isolated for multiple glacial cycles for some species, whereas for other species genetic connections have been more recent. Regional estimates of genetic diversity and directionality of gene flow also vary among species. Thus, there is no apparent consistency among historical patterns across the Torres Strait for these co-distributed marine animals.     

Land Use as a Driver of Patterns of Rodenticide Exposure in Modeled Kit Fox Populations

Although rodenticides are increasingly regulated, they nonetheless cause poisonings in many non-target wildlife species. Second-generation anticoagulant rodenticide use is common in agricultural and residential landscapes. Here, we use an individual-based population model to assess potential population-wide effects of rodenticide exposures on the endangered San Joaquin kit fox (Vulpes macrotis mutica). We estimate likelihood of rodenticide exposure across the species range for each land cover type based on a database of reported pesticide use and literature. Using a spatially-explicit population model, we find that 36% of modeled kit foxes are likely exposed, resulting in a 7-18% decline in the range-wide modeled kit fox population that can be linked to rodenticide use. Exposures of kit foxes in low-density developed areas accounted for 70% of the population-wide exposures to rodenticides. We conclude that exposures of non-target kit foxes could be greatly mitigated by reducing the use of second-generation anticoagulant rodenticides in low-density developed areas near vulnerable populations.     

Gene flow among San Joaquin kit fox populations in a severely changed ecosystem

The San Joaquin kit fox (Vulpes macrotis mutica) was once ubiquitous throughout Californias San Joaquin Valley and its surrounds. However, most of its habitat has been lost to irrigated agriculture, urban development, and oil fields. The remaining foxes are concentrated in six areas, although there are several small pockets of foxes throughout the Valley. To help conserve kit foxes, we sought an ecological understanding of the level of genetic variation remaining in these locations and the extent of gene flow among them. We collected tissue from 317 kit foxes from 8 sites and estimated genetic variability in and gene flow among sites using data from 8 polymorphic, microsatellite markers. We found no differences in both observed and expected heterozygosity between locations using Bonferonni corrected paired t-tests. We found differences in mean number of alleles per locus, even after we used Monte Carlo simulations to adjust for sample size differences. Population subdivision was low among sites (F_st=0.043), yet a matrix of pairwise F_st values was correlated with a matrix of pairwise geographic distances. An assignment test classified only 45% of the individuals to the site where they were captured. Overall, these data suggest that kit fox dispersal between locations may still maintain genetic variation throughout most of the areas we sampled.     

Resource partitioning among cape foxes,bat-eared foxes,and black-backed jackals in South Africa

Cape foxes (Vulpes chama) and bat-eared foxes (Otocyon megalotis) are sympatric with black-backed jackals (Canis mesomelas) over much of southern Africa, although competition with and/or predation by jackals may suppress local populations of both fox species. From 2005 to 2008, we captured, radio-collared, and monitored 11 cape foxes, 22 bat-eared foxes, and 15 black-backed jackals on a game ranch in South Africa to investigate their spatial, habitat, temporal, and dietary resource overlap. Mean annual home-range sizes were 27.7 km² for cape foxes, 5.0 km² for bat-eared foxes, and 17.8 km² for jackal family groups. Home ranges overlapped completely between species, although core areas overlapped less (<45%), with cape foxes and jackals overlapping the least (12%). When active, cape foxes, but not bat-eared foxes, used core areas of jackal groups less than expected. Additionally, both fox species used jackal core areas less than expected for their den sites, suggesting areas outside jackal core areas were used as refuges by foxes. Strong levels of habitat partitioning were not apparent at the study site or home-range levels, although habitat selection for den sites differed between jackals and cape foxes. Jackals were the most diurnal across seasons, whereas cape foxes were the most nocturnal. Diets overlapped little (R₀ = 0.20–0.34) among the canid species, with bat-eared foxes overlapping the least with the others. Jackals killed at least 5 collared bat-eared foxes and 1 collared cape fox, indicating potential interference competition, probably for exclusive use of territorial space rather than over shared resources. We conclude that bat-eared foxes coexisted with jackals primarily by their dietary specialization and group living. Cape foxes coexisted with jackals by exhibiting high levels of spatial, habitat, temporal, and dietary partitioning. Surprisingly, the fox species exhibited positive associations with each other. Our results show the mechanisms that may allow jackals to suppress fox populations, yet also show how foxes, in turn, use different mechanisms to coexist with a dominant canid. © 2012 The Wildlife Society.     

Mitochondrial Diversity and Distribution of African Green Monkeys (Chlorocebus Gray, 1870)

African green monkeys (Chlorocebus) represent a widely distributed and morphologically diverse primate genus in sub‐Saharan Africa. Little attention has been paid to their genetic diversity and phylogeny. Based on morphological data, six species are currently recognized, but their taxonomy remains disputed. Here, we aim to characterize the mitochondrial (mt) DNA diversity, biogeography and phylogeny of African green monkeys. We analyzed the complete mitochondrial cytochrome b gene of 126 samples using feces from wild individuals and material from zoo and museum specimens with clear geographical provenance, including several type specimens. We found evidence for nine major mtDNA clades that reflect geographic distributions rather than taxa, implying that the mtDNA diversity of African green monkeys does not conform to existing taxonomic classifications. Phylogenetic relationships among clades could not be resolved suggesting a rapid early divergence of lineages. Several discordances between mtDNA and phenotype indicate that hybridization may have occurred in contact zones among species, including the threatened Bale monkey (Chlorocebus djamdjamensis). Our results provide both valuable data on African green monkeys’ genetic diversity and evolution and a basis for further molecular studies on this genus. Am. J. Primatol. 75:350‐360, 2013. © 2013 Wiley Periodicals, Inc.     

Molecular patterns of differentiation in canyon treefrogs (Hyla arenicolor): evidence for introgressive hybridization with the Arizona treefrog (H. wrightorum) and correlations with advertisement call differences

Detection of genetic and behavioural diversity within morphologically similar species has led to the discovery of cryptic species complexes. We tested the hypothesis that US populations of the canyon treefrog (Hyla arenicolor) may consist of cryptic species by examining mate‐attraction signals among three divergent clades defined by mtDNA. Using a multi‐locus approach, we re‐analysed phylogenetic relationships among the three clades and a closely related, but morphologically and behaviourally dissimilar species, the Arizona treefrog (H. wrightorum). We found evidence for introgression of H. wrightorum’s mitochondrial genome into H. arenicolor. Additionally, the two‐clade topology based on nuclear data is more congruent with patterns of call variation than the three‐clade topology from the mitochondrial dataset. The magnitude of the call divergence is probably insufficient to promote isolation of the nuclear DNA‐defined clades should they become sympatric, but further divergence in call properties significant in species identification could promote speciation in the future.