Limited mitochondrial DNA variation within South Africa's black rhino (Diceros bicornis minor) population and implications for management

The taxonomy of African black rhinoceros (Diceros bicornis) remains unresolved. Maintaining levels of genetic diversity and species rescue by reintroduction and restocking requires its resolution. We compared the sequences of the mitochondrial DNA (mtDNA) control region for a total of 101 D. bicornis from three subspecies: D. b. minor, D. b. michaeli and D. b. bicornis. A single unique haplotype was found within the 65 D. b. minor samples from KwaZulu‐Natal (KZN) Province, South Africa, 55 of which came from Hluhluwe‐iMfolozi Game Park (HiP) and Mkuzi Game Reserve (MGR) source populations. However, six different haplotypes were represented in eleven D. b. minor samples from Zimbabwe. Similarly, published autosomal microsatellite data indicate low levels of diversity within the KZN D. b. minor populations. The low levels of mtDNA diversity within the KZN metapopulation point to the possible need for genetic supplementation. However, there is a need to determine whether the low levels of genetic variation within KZN D. b. minor are a result of the recent bottleneck or whether KZN historically always had low diversity.     

The Divergence of the Dolly Varden Char Salvelinus malma in the Asian Northern Pacific Populations Inferred from the PCR–RFLP Analysis of Mitochondrial DNA

Genetic differentiation of the Dolly Varden char Salvelinus malmaWalbaum was studied in five populations from the western part of the Northern Pacific. Using restriction analysis (RFLP), we examined polymorphism of three mitochondrial DNA (mtDNA) fragments amplified in polymerase chain reaction (PCR). MtDNA haplotypes were shown to fall into two phylogenetic groups, which probably reflect the existence of two previously described subspecies of Asian Dolly Varden,S. malma malma and S. malma krascheninnikovi. The divergence of mtDNA nucleotide sequences in the Dolly Varden subspecies (about 4%) corresponds to the differences between the valid char species from the genus Salvelinus.     

Is the Guadeloupean racoon (Procyon minor) really an endemic species? New insights from molecular and chromosomal analyses

The Guadeloupean racoon has been chosen as the emblematic species of the Parc National de Guadeloupe because of its supposed endemic specificity. However, its taxonomic position is not clearly defined. At what period did this animal colonize Guadeloupe and how it reached the island remain unsolved questions. To clarify these points, insular and continental individuals (Procyon lotor), coming from five localities and belonging to three subspecies, were compared using molecular (sequencing of a 294-bp of the left domain of the mtDNA control region) and chromosomal (C, G and R karyotypes) analyses. Genetic distance between the Guadeloupean racoon and lotor subspecies from Virginia and Maryland was very short (mean sequence divergence between haplotypes equals to 1.46%). A four times higher distance was found between P. l. lotor and P. l. pallidus, a continental subspecies inhabiting Arizona than between P. l. lotor and Procyon minor. A neighbour-joining phenogram is proposed; the Guadeloupean haplotype is on a branch close to the one clustering P. l. lotor from the East coast whereas the haplotype from Arizona diverged earlier. These results strongly suggest that the Guadeloupean racoon is included in the P. lotor species. Moreover, no differences were found between P. lotor and P. minor karyotypes, again suggesting a high genetic homogeneity between the two taxa. Furthermore, comparison of the control region variability within Procyon with intra- and interspecific variability observed in other Carnivores, clearly strengthens the hypothesis that P. minor is conspecific with P. lotor. These results imply that the taxonomic status of the Guadeloupean racoon should be revised. In a biogeographic perspective, it is suggested that the racoon has been recently introduced in Guadeloupe by man with individuals coming from the east coast of the USA. The consequences of these results for conservation of the racoon in Guadeloupe, and more generally, the definition and the use of the subspecies as a unit of conservation are discussed.     

Phylogeographic differentiation in the mitochondrial control region in the koala, Phascolarctos cinereus (Goldfuss 1817)

The koala, Phascolarctos cinereus, is a geographically widespread species endemic to Australia, with three currently recognized subspecies: P.c. adustus, P.c. cinereus, and P.c. victor. Intraspecific variation in the mitochondrial DNA (mtDNA) control region was examined in over 200 animals from 16 representative populations throughout the species’ range. Eighteen different haplotypes were defined in the ≈ 860 bp mtDNA control region, as determined by heteroduplex analysis/temperature gradient gel electrophoresis (HDA/TGGE). Any single population typically possessed only one or two haplotypes yielding an average within-population haplotypic diversity of 0.180 ± 0.003, and nucleotide diversity of 0.16%. Overall, mtDNA control region sequence diversity between populations averaged 0.67%, and ranged from 0% to 1.56%. Nucleotide divergence between populations averaged 0.51%, and ranged from 0% to 1.53%. Neighbour-joining methods revealed limited phylogenetic distinction between geographically distant populations of koalas, and tentative support for a single evolutionarily significant unit (ESU). This is consistent with previous suggestions that the morphological differences formalized by subspecific taxonomy may be interpreted as clinal variation. Significant differentiation in mtDNA-haplotype frequencies between localities suggested that little gene flow currently exists among populations. When combined with microsatellite analysis, which has revealed substantial differentiation among koala populations, we conclude that the appropriate short-term management unit (MU) for koalas is the local population.     

Phylogeography of the Black-tailed Godwit Limosa limosa: substructuring revealed by mtDNA control region sequences

Black-tailed (Limosa limosa) and Hudsonian Godwits (L. haemastica) are sometimes described as a superspecies. The Black-tailed Godwit is further split into three subspecies on the basis of morphological differences (L. l. limosa, L. l. islandica and L. l. melanuroides). We studied variation in partial mtDNA control region sequences among Black-tailed and Hudsonian Godwits which showed 5% divergence. Black-tailed and Hudsonian Godwits were thus clearly differentiated and the separate species status for the two taxa is validated. All three subspecies described for the Black-tailed Godwit had unique haplotypes but the genetic distances were small (0.3–0.6%). Despite small genetic differences we could not detect any substantial gene flow between any of the subspecies as haplotypes were private to each subspecies. Thus, genetic variation within Black-tailed Godwits showed a clear geographic structure. We found a high proportion of rare private haplotypes in three fringe populations of the nominate subspecies of the Black-tailed Godwit (L. l. limosa) where godwits breed in low numbers, but no genetic variation at all in a sample from the Netherlands where godwits are abundant. This suggests that Dutch Godwits may have been affected by a founder effect.     

MITOCHONDRIAL DNA HOMOGENEITY IN THE PHENOTYPICALLY DIVERSE REDPOLL FINCH COMPLEX (AVES: CARDUELINAE: CARDUELIS FLAMMEA-HORNEMANNI)

Breeding redpoll finches (Aves: Carduelinae) show extensive plumage and size variability and, in many cases, a plumage polymorphism that is not related to age or sex. This has been ascribed to extreme phenotypic variation within a single taxon or to moderate variability within distinct taxa coupled with hybridization. The predominant view favors the recognition of two largely sympatric species: Carduelis flammea, comprised of four well-marked subspecies—flammea, cabaret, islandica, and rostrata; and C. hornemanni, comprised of two subspecies—hornemanni and exilipes. We studied representative samples of these putative subspecies (except islandica) for variation in mitochondrial DNA (mtDNA). Using 20 informative restriction enzymes that recognized 124 sites (642 base pairs [bp] of sequence or ≈ 3.7% of the molecule), we identified 17 RFLP haplotypes in the 31 individuals surveyed. The haplotypes formed a simple phylogenetic network with most clones diverging by a single site difference from a common haplotype found in almost half of the individuals. Within populations and taxa, levels of mtDNA diversity were similar to those observed in other avian species. The pattern of mtDNA divergence among populations was statistically unrelated to their geographic or traditional taxonomic relationships, and the estimated distance between the two traditionally recognized species was very small relative to those typically observed among avian sister species.     

Genetic differentiation and intraspecific structure of Eastern Tropical Pacific spotted dolphins,Stenella attenuata,revealed by DNA analyses

Mitochondrial DNA (mtDNA) control region sequences and microsatellite loci length polymorphisms were used to investigate genetic differentiation in spotted dolphins (Stenella attenuata) in the Eastern Tropical Pacific and to examine the intraspecific structure of the coastal subspecies (Stenella attenuata graffmani). One-hundred and thirty-five animals from several coastal areas and 90 offshore animals were sequenced for 455 bp of the mitochondrial control region, resulting in 112 mtDNA haplotypes. Phylogenetic analyses and the existence of shared haplotypes between the two subspecies suggest recent and/or current gene flow. Analyses using χ², F _ST (based on haplotype frequencies) and Φ_ST values (based on frequencies and genetic distances between haplotypes) yielded statistically significant separation (randomized permutation values P<0.05) among four different coastal populations and between all but one of these and the offshore subspecies (overall F _ST=0.0691). Ninety-one coastal animals from these four geographic populations and 50 offshore animals were genotyped for seven nuclear microsatellite loci. Analysis using F _ST values (based on allelic frequencies) yielded statistically significant separation between most coastal populations and offshore animals, although no coastal populations were distinguished. These results argue for the existence of some genetic isolation between offshore and inshore populations and among some inshore populations, suggesting that these should be treated as separate units for management purposes.     

Evolution of the Cyprinella lutrensis species group. III. Geographic variation in the mitochondrial DNA of Cyprinella lutrensis— the influence of Pleistocene glaciation on population dispersal and divergence

We employed restriction site variation in mitochondrial (mt)DNA to determine if significant phylogeographic structure occurs in the North American cyprinid fish Cyprinella lutrensis. Digestion patterns from 16 restriction endonucleases identified fifty mtDNA haplotypes among 127 individuals of Cyprinella lutrensis assayed from localities in the Gulf Coastal Plain, the Great Plains, and the Central Lowlands. Nucleotide sequence divergence among haplotypes was highly variable (mean ± SE: 2.87%± 0.08; range: 0.14–9.24%). Maximum-parsimony analysis and the neighbour joining method of tree construction revealed three major groupings (clades) of haplotypes that differed in geographic distribution. Divergence estimates between the basal clade, comprised of haplotypes primarily from the Brazos River in east Texas, and the remaining two clades, place C. lutrensis in the western Gulf Coastal Plain prior to Pleistocene glaciation. Nucleotide sequence divergence between the second clade, comprised of haplotypes from the Trinity and Calcasieu rivers in east Texas and southwestern Louisiana, respectively, and the third clade (comprised primarily of haplotypes from localities north of Texas and affected directly by Pleistocene glaciation), suggest that C. lutrensis colonized gladated regions to the north during the mid- to late Pleistocene. This hypothesis is supported by levels of intrapopulational nucleotide diversity in geographic localities outside of Texas and by geological evidence. Despite marked geographic variation in morphometries, meristics, and nuptial coloration, mtDNA variation in glaciated regions was not geographically structured, and subspecies of C. lutrensis were not identifiable by phylogenetic analysis of mtDNA.     

Intraspecific genetic differentiation of the Siberian Rubythroat (Luscinia calliope): Data of sequencing the mtDNA cytochrome b gene

For the first time, genetic diversity and intraspecific structure of Luscinia calliope was studied according to the data of the mtDNA cytochrome b gene sequencing. The strong differentiation of haplotypes of the Siberian Rubythroat into western and eastern groups, which include subspecies according to their geographical attachment, was revealed. A high-haplotypic (Hd = 0.986) and nucleotide (π= 0.00875) variety was shown for the species as a whole. We revealed considerable genetic distances (D = 0.016) between the western and eastern haplotypes that were four times higher than the intraspecific distances in terms of cytochrome b for passerines (D = 0.004). For three birds from Transbaikal, significant genetic divergence was detected, which could indirectly indicate the existence of the hybrid zone of several subspecies in this part of the area.     

Mitochondrial DNA variation in the hybridizing fire-bellied toads, Bombina bombina and B. variegata

Using five restriction enzymes, geographical variation of mitochondrial DNA (mtDNA) in Bombina bombina and B. variegata was studied in samples from 20 locations. Each restriction enzyme produced a species-specific fragment pattern. B. bombina haplotypes A and B were closely related to each other. In contrast, haplotypes A and B of B. variegata formed two distinct lineages. A very distinctive haplotype (C) was found in the Carpathian Mountains, whereas two other haplotypes, D and E (differing by a single AvaI site), were present in western Europe and the Balkans, respectively. Populations polymorphic for haplotypes D and E occurred in the central Balkans where the haplotypes could replace each other clinally. mtDNA sequence divergence between B. bombina and B. variegata was estimated as 6.0-8.1% and 4.7-5.2% between type C and types D/E of B. variegata. The latter divergence is contrary to allozyme and morphological data that place the western and Carpathian B. v. variegata together (Nei's D = 0.07) and separate them from the Balkan subspecies B. v. scabra (Nei's D = 0.18). Broad interspecific correlation among morphology, allozymes and mtDNA types in European fire-bellied toads argues that, despite continuous hybridization (interrupted perhaps during Pleistocene glacial maxima), little or no mtDNA introgression between the species has occurred outside the narrow hybrid zones that separate these parapatric species.     

Mitochondrial DNA polymorphisms in the two subspecies of Drosophila sulfurigaster: relationship between geographic structure of population and nucleotide diversity.

Recent empirical and theoretical studies on mitochondrial DNA (mtDNA) variation in higher animals have suggested that the extent of mtDNA polymorphism is largely affected by spatial population subdivision. To examine this we studied mtDNA polymorphism in two subspecies of Drosophila sulfurigaster: D. s. albostrigata and D. s. bilimbata. Drosophila sulfurigaster albostrigata is mainly distributed on the mainland of Southeast Asia. In contrast, D. s. bilimbata forms discontinuous populations on many islands scattered in the Pacific Ocean. Because of the difference in their distribution patterns, the two subspecies are thought to be different in the extent of spatial population subdivision. mtDNA was isolated from greater than 50 isofemale strains for each subspecies and were analyzed by eight restriction endonucleases. Nucleotide diversity within a population was higher in D. s. albostrigata than in D. s. bilimbata. However, haplotype diversity was 1.6 times greater in D. s. bilimbata (0.85) than in D. s. albostrigata (0.53). The large difference in overall heterogeneity was attributed to the difference in interpopulational nucleotide diversity. For the two subspecies the proportion of interpopulational gene diversity in a subdivided population was calculated to be 0.54 in D. s. bilimbata and 0.40 in D. s. albostrigata. These observations indicate that spatial population subdivision is a major factor in determining mtDNA polymorphism in these subspecies. The extent of mtDNA divergence between the subspecies was very high. The average nucleotide divergence between them was 7.6%, which is almost the interspecific level reported for other Drosophila species. The cause of the high degree of mtDNA divergence is discussed.     

Morphological evolution and genetic differentiation in Daphnia species complexes

Despite many ecological and evolutionary studies, the history of several species complexes within the freshwater crustacean genus Daphnia (Branchiopoda, Anomopoda) is poorly understood. In particular, the Daphnia longispina group, comprising several large-lake species, is characterized by pronounced phenotypic plasticity, many hybridizing species and backcrossing. We studied clonal assemblages from lakes and ponds comprising daphnids from several species complexes. In order to reveal patterns of reticulate evolution and introgression among species, we analysed three data sets and compared nuclear, mtDNA and morphological divergence using animals from 158 newly established clonal cultures. By examining 15 nuclear and 11 mitochondrial (12S/16S rDNA) genetic characters (allozymes/restriction enzymes), and 48 morphological traits, we found high clonal diversity and discontinuities in genotypic and morphological space which allowed us to group clones by cytonuclear differentiation into seven units (outgroup D. pulex). In contrast to six groups emerging from nuclear divergence (related to three traditional species, D. cucullata, D. galeata, D. hyalina and three pairwise intermediate hybrids), a seventh group of clones was clearly resolved by morphological divergence: distinct mtDNA haplotypes within one nuclear defined cluster, ‘D. hyalina’, resembled traditional D. hyalina and D. rosea phenotypes, respectively. In other nuclear defined clusters, association between mtDNA haplotype and morphology was low, despite hybridization being bidirectional (reciprocal crosses). Morphological divergence was greatest between young sister species which are separated on the lake/pond level, suggesting a significant role for divergent selection during speciation along with habitat shifts. Phylogenetic analyses were restricted to four cytonuclear groups of clones related to species. mtDNA and nuclear phylogenies were consistent in low genetic divergence and monophyly of D. hyalina and D. rosea. Incongruent patterns of phylogenies and different levels of genetic differentiation between traditional species suggest reticulate evolutionary processes.     

Phylogeography of the Alpine salamander, Salamandra atra (Salamandridae) and the influence of the Pleistocene climatic oscillations on population divergence

Fifty individuals of the endemic Alpine salamander, Salamandra atra, representing 13 populations throughout the range of the two currently recognized subspecies, atra and aurorae, were examined for sequence variation in a large portion (1050 bp) of the mitochondrial cytochrome b gene. We revealed a large number of mitochondrial DNA (mtDNA) haplotypes (10). Interpopulation sequence divergence was very low, ranging from 0 to 3.1%. The relationships among haplotypes were poorly resolved. The divergence time estimate between several mtDNA haplotypes suggested a pre-Pleistocene differentiation approximately 3 million years ago. Moreover, the impact of the Pleistocene glaciations on the phylogeographical patterns appears to have been secondary, although a somewhat reduced genetic variability was found in populations living in areas that were directly affected by the glaciation.     

Phylogeography and conservation genetics of the hellbender salamander (Cryptobranchus alleganiensis)

We investigated hellbender phylogeography through phylogenetic analyses of individuals sampled from 16 locations throughout their range in the eastern United States. Analyses were conducted on concatenated cytochrome-oxidase I (COI), cytochrome-b (Cytb) and NADH dehydrogenase subunit 4 (ND4) mtDNA sequence, totaling 2160 nucleotides. Hellbender haplotypes differed by 0.1 to 5.8% maximum likelihood (ML) corrected sequence divergence. Phylogenetic analyses reveal that hellbenders are separated into 8 reciprocally monophyletic populations or clades differentiated by a minimum of 0.7 to 5.4% sequence divergence, each of which constitutes a separate Management Unit (MU). High among population divergence and reciprocal monophyly suggest that female-mediated gene flow is severely restricted or non-existent among each MU. Hellbenders are currently divided into two subspecies, Cryptobranchus alleganiensis alleganiensis and C. a. bishopi based on morphological characters. The phylogenetic analyses presented here strongly indicate that these subspecies are paraphyletic. Management priorities for the hellbender should be reconsidered in light of these new molecular data. Results from Bayesian rooting indicate the root of the hellbender mtDNA tree lies on the branch leading to hellbender haplotypes from the Current, Eleven Point and New Rivers. The rooted tree suggests that a common ancestor in the southern Ozarks and/or southern Appalachians gave rise to northern hellbender populations, consistent with a Pleistocene refuge hypothesis.     

A genetic comparison of two alleged subspecies of Philippine cynomolgus macaques

Two subspecies of cynomolgus macaques (Macaca fascicularis) are alleged to co‐exist in the Philippines, M. f. philippensis in the north and M. f. fascicularis in the south. However, genetic differences between the cynomolgus macaques in the two regions have never been studied to document the propriety of their subspecies status. We genotyped samples of cynomolgus macaques from Batangas in southwestern Luzon and Zamboanga in southwestern Mindanao for 15 short tandem repeat (STR) loci and sequenced an 835 bp fragment of the mtDNA of these animals. The STR genotypes were compared with those of cynomolgus macaques from southern Sumatra, Singapore, Mauritius and Cambodia, and the mtDNA sequences of both Philippine populations were compared with those of cynomolgus macaques from southern Sumatra, Indonesia and Sarawak, Malaysia. We conducted STRUCTURE and PCA analyses based on the STRs and constructed a median joining network based on the mtDNA sequences. The Philippine population from Batangas exhibited much less genetic diversity and greater genetic divergence from all other populations, including the Philippine population from Zamboanga. Sequences from both Batangas and Zamboanga were most closely related to two different mtDNA haplotypes from Sarawak from which they are apparently derived. Those from Zamboanga were more recently derived than those from Batangas, consistent with their later arrival in the Philippines. However, clustering analyses do not support a sufficient genetic distinction of cynomolgus macaques from Batangas from other regional populations assigned to subspecies M. f. fascicularis to warrant the subspecies distinction M. f. philippensis. Am J Phys Anthropol 155:136–148, 2014. © 2014 Wiley Periodicals, Inc.     

The role of niche divergence and phenotypic adaptation in promoting lineage diversification in the Sage Sparrow (Artemisiospiza belli,Aves: Emberizidae)

Niche divergence or conservatism and phenotypic adaptation are important in lineage diversification. We used mitochondrial DNA (mtDNA), morphology and ecological niche models to examine these processes in three subspecies of Sage Sparrow (Artemisiospiza belli belli, A. b. canescens and A. b. nevadensis) that breed in bioclimatically diverse ecoregions in western North America. Overall, mtDNA and morphology are congruent with subspecies, ecoregion and bioclimatic niche. Niche divergence, rather than niche conservatism, accompanied by phenotypic adaptation, is associated with lineage diversification between subspecies. This diversification has occurred with and without physical barriers or accompanying genetic divergence. Populations of A. b. canescens are divided by a montane barrier into two bioclimatic regions (San Joaquin Valley, Mojave Desert), where they are indistinguishable phenotypically, but show distinctive genetic patterns. Although there is no physical barrier between A. b. canescens in the San Joaquin Valley and A. b. belli in the Coast Ranges, these populations occupy different bioclimatic niches and are phenotypically, but not genetically, diagnosable. Niche overlap is greatest between A. b. canescens from the Mojave Desert and A. b. nevadensis from the Great Basin, yet these subspecies maintain distinctive phenotypes and mtDNA, even in local secondary contact and sympatry. Palaeoclimatic niche models for the Last Glacial Maximum (c. 21 000 bp ) and the Last Interglacial (c. 120 000 bp ) suggest that ecoregionally distinct populations of Artemisiospiza belli experienced different Pleistocene range fluctuations and glacial refugia, with temporal niche conservatism. Populations probably reached their current distributions as favourable climates and habitats expanded after the last glaciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Mitochondrial DNA variation, population structure, and evolution of mountain sheep in the south-western United States and Mexico

In the south-western United States and Mexico, mountain sheep were found to have low levels of mtDNA sequence divergence among haplotypes and low overall nucleotide diversity. Potential causes for this low diversity are discussed. Significant differences in mtDNA haplotype distributions over short distances and high values of N_ST on a local scale appear to be a result of the tendency of female mountain sheep to disperse less frequently and over shorter distances than males. A lack of concordance between mtDNA haplotype distributions and assumed subspecies boundaries suggest that some taxonomic labels need to be revised.     

Differentiation of Dolly Varden Char <Emphasis Type="Italic">Salvelinus malma</Emphasis> from Asia and North America Inferred from PCR-RFLP Analysis of Mitochondrial DNA

Genetic differentiation of Dolly Varden char Salvelinus malma Walbaum from the Asian and North American Pacific coasts was studied. We examined restriction fragment length polymorphism of three mitochondrial DNA (mtDNA) fragments amplified in polymerase chain reaction, which encoded four NADH dehydrogenase subunits, the cytochrome b gene, and a D-loop segment. The mtDNA haplotypes were shown to form three phylogenetic groups, whose geographic distribution corresponded to three Dolly Varden subspecies: S. malma malma, S. malma krascheninnikovi, and S. malma lordi. The nucleotide sequence divergence between S. malma malma and S. malma krascheninnikovi was 3.8%; between S. malma malma and S. malma lordi, 3.1%; and between S. malma krascheninnikovi and S. malma lordi, 2.5%. The northern Dolly Varden S. malma malma from Asia was shown to be genetically identical to that from North America.__________Translated from Genetika, Vol. 41, No. 5, 2005, pp. 626–634.Original Russian Text Copyright © 2005 by Oleinik, Skurikhina, Brykov, Crane, Wenburg.     

Integrated genetic and morphological data support eco‐evolutionary divergence of Angolan and South African populations of Diplodus hottentotus

The genus Diplodus presents multiple cases of taxonomic conjecture. Among these the D. cervinus complex was previously described as comprising three subspecies that are now regarded as separate species: Diplodus cervinus, Diplodus hottentotus and Diplodus omanensis. Diplodus hottentotus exhibits a clear break in its distribution around the Benguela Current system, prompting speculation that Angolan and South African populations flanking this area may be isolated and warrant formal taxonomic distinction. This study reports the first integrated genetic [mitochondrial (mt)DNA and nuclear microsatellite] and morphological (morphometric, meristic and colouration) study to assess patterns of divergence between populations in the two regions. High levels of cytonuclear divergence between the populations support a prolonged period of genetic isolation, with the sharing of only one mtDNA haplotype (12 haplotypes were fully sorted between regions) attributed to retention of ancestral polymorphism. Fish from the two regions were significantly differentiated at a number of morphometric (69·5%) and meristic (46%) characters. In addition, Angolan and South African fish exhibited reciprocally diagnostic colouration patterns that were more similar to Mediterranean and Indian Ocean congeners, respectively. Based on the congruent genetic and phenotypic diversity we suggest that the use of hottentotus, whether for full species or subspecies status, should be restricted to South African D. cervinus to reflect their status as a distinct species‐like unit, while the relationship between Angolan and Atlantic–Mediterranean D. cervinus will require further demo‐genetic analysis. This study highlights the utility of integrated genetic and morphological approaches to assess taxonomic diversity within the biogeographically dynamic Benguela Current region.     

ASSOCIATIONS BETWEEN MITOCHONDRIAL AND NUCLEAR GENOTYPES IN CUTTHROAT TROUT HYBRID SWARMS

We examined mtDNA and nuclear allozyme genotypes in hybrid populations formed from interbreeding of westslope cutthroat trout (Oncorhynchus clarki lewisi) and Yellowstone cutthroat trout (O. c. bouvieri). These subspecies show substantial genetic divergence (Nei's D = 0.30; mtDNA P = 0.02). Diagnostic alleles at multiple nuclear loci and two distinct mtDNA haplotypes segregate in the hybrids. Nuclear and mtDNA genotypes are largely randomly associated, although there is slight disequilibrium in both nuclear and cytonuclear measures in some samples. Consistent positive gametic disequilibria for three pairs of nuclear loci confirm one previously reported linkage, and indicate two more. Allele frequencies provide no evidence for selection on individual chromosome segments. However, westslope mtDNA haplotype frequencies exceed westslope nuclear allele frequencies in all samples. This may be explained by differences in the frequency of occurrence of reciprocal F₁ matings, by viability, fertility, or sex ratio differences in the progeny of reciprocal matings, or by weak selection on mtDNA haplotypes.