Taxonomic status and evolutionary history of the Saxicola torquata complex

We explored variation in mitochondrial ND2 gene sequences from 171 stonechats (Saxicola torquata complex) collected from 27 Eurasian and 3 African localities. We found that two widespread neighboring Eurasian subspecies of Siberian stonechat, Saxicola maura maura and S. m. stejnegeri, although very similar in appearance, are not each other’s closest relatives. Eastern Asian S. m. stejnegeri appears to have split from other Palearctic, African, and island stonechats well before differentiation occurred among the latter taxa. Our data indicate that European (S. t. rubicola), central Palearctic (S. m. maura) and eastern Palearctic (S. m. stejnegeri) are independently evolving, and could each warrant separate species status, as assumed earlier for S. rubicola, S. maura and S. torquata. However, we found three localities at which individuals from different major groups occurred. Thus, although these are likely phylogenetic species it is unclear whether they are biological species. There was little phylogeographic structure within the three major Palearctic clades, although samples from Spain might be showing incipient divergence. We maintain that the topology of a mtDNA gene tree is a valid means to discern taxonomic limits.     

Moult and basal metabolic costs in males of two subspecies of stonechats: the European Saxicola torquata rubicula and the East African S. t. axillaris

The circannual patterns in resting metabolic rate (RMR) of males of two subspecies of stonechats, the European Saxicola torquata rubicula and the East African S. t. axillaris, are compared. As the birds from the two subspecies were raised and kept under comparable laboratory conditions, differences in metabolic rate between the two subspecies had to be genetically determined. RMR peaked during moult in both subspecies. During the rest of the year RMR was fairly constant in both subspecies and assumed to reflect basal metabolic rate (BMR). African stonechats had a 22% lower mass specific BMR than European stonechats, which is thought to reflect a genetical physiological adaptation to the differences in environmental circumstances they experience in the field. A low BMR makes an animal more susceptible to cold. Hence, the relatively high plumage mass in the African compared to the European stonechat may be functionally linked to its relatively low BMR. Moult costs, calculated from the plumage masses and the differences in RMR inside and outside the moult period, tended to be higher in the European compared to the African stonechats. These data and an interspecific comparison of moult costs over various species of birds support the earlier notion by Lindström et al. (1993) that moult costs are more closely linked with BMR than with body mass or rate of moult. The relation between moult costs and BMR and the fact that the efficiency of moult is extremely low (3.8 and 6.4% for European and African stonechats, respectively) suggest that the maintenance of specific tissues necessary for moult is a large cost factor. Alternatively, impaired insulation during moult may necessitate an increased metabolic capacity which may be associated with an increased RMR.     

A molecular phylogeny of the Sylvia cantillans complex: cryptic species within the Mediterranean basin

The subalpine warbler Sylvia cantillans is formally considered a polytypic species, with four subspecies, European S. c. cantillans, albistriata, moltonii (recently resumed name: subalpina) and North African S. c. inornata. They are very similar in external morphology but clearly differ in their vocalizations. We evaluated their uncertain taxonomic status reconstructing the phylogenetic and phylogeographic relationships among populations sampled across major biogeographical areas in the European species’ range, using nucleotide sequences of the mitochondrial cytochrome b gene (mtDNA cyt b). A variety of phylogenetic analyses concordantly led to identify four major groups, only partially corresponding to the three European nominal subspecies. Phylogenetic trees showed a monophyletic group including all moltonii individuals, well diverged from all other taxa. Populations taxonomically assigned to cantillans were polyphyletic being split into two distinct clades (western and southern cantillans), with monophyletic albistriata closely related to southern cantillans. Individuals of moltonii and southern cantillans sampled in sites of sympatry in central Italy were assigned to their respective groups, with perfect concordance between phenotypic and genetic identifications. All findings indicate that moltonii should be ranked as a distinct species. Former subspecies cantillans is polyphyletic, but additional data are needed to define the taxonomic status of its two clades. Albistriata is phylogenetically related to southern cantillans and should be provisionally kept as a subspecies of S. cantillans. The cantillans complex thus provides an interesting case-study illustrating geographical structuring across small geographical ranges, and it exemplifies speciation through differentiation in allopatry leading to reproductive isolation after a secondary contact.     

Die Jugendmause europ?ischer und afrikanischer Schwarzkehlchen (Saxicola torquata rubicula undaxillaris) sowie von F1-Hybriden

Summary In a constant 12 h, 50 min equatorial photoperiod postjuvenile moult started earlier and lasted shorter in European stonechats than in their African conspecifics. F₁-hybrids showed an intermediary time course of moult indicating that the differences found between these two subspecies are genetically determined.     

A survey of equid mitochondrial DNA: Implications for the evolution, genetic diversity and conservation of Equus

The evolution, taxonomy and conservation of the genus Equuswere investigated by examining the mitochondrial DNA sequences of thecontrol region and 12S rRNA gene. The phylogenetic analysis of thesesequences provides further evidence that the deepest node in thephylogeny of the extant species is a divergence between twolineages; one leading to the ancestor of modern horses (E.ferus, domestic and przewalskii) and the other to thezebra and ass ancestor, with the later speciation events of the zebrasand asses occurring either as one or more rapid radiations, or withextensive secondary contact after speciation. Examination of the geneticdiversity within species suggested that two of the E. hemionussubspecies (E. h. onager and E. h. kulan) onlyrecently diverged, and perhaps, are insufficiently different to beclassified as separate subspecies. The genetic divergence betweendomestic and wild forms of E. ferus (horse) and E.africanus (African ass) was no greater than expected within anequid species. In E. burchelli (plains zebra) there was anindication of mtDNA divergence between populations increasing withdistance. The implications of these results for equid conservation arediscussed and recommendations are made for conservation action.     

Partial reproductive isolation of subspecies of the California vole,Microtus californicus

Summary A partial reproductive barrier found between subspecies of the California vole indicates that there may be obstacles to free interbreeding within this species in nature. Both the reduced fertility between subspecies and the sterility of hybrid males found in laboratory crosses would be obstacles to gene exchange between contiguous subspecies populations. Combative behavior engaged in by approximately half of the mixed subspecies pairs in the laboratory indicate that there might also be behavioral constraints to their breeding. Substantial differentiation has already occurred between M.c. californicus and stephensi in some of the faster evolving structural gene loci coding for enzymes not involved in glucose metabolism. Due to significant differences in body size, skull characteristics, and organ weights, the two subspecies are clearly distinguishable on the basis of these morphophysiological characters at this stage of reproductive isolation. Thus, there is good reason to believe that these populations of the California vole are diverging to the species level. I am continuing the investigation of the intraspecific divergence with a comparative study of the geographically intervening subspecies M.c. sanctidiegi, to determine its interfertility with californicus and stephensi. The speciation process has been studied extensively in drosophilids (Ayala, 1975) and there have been many studies of geographical variation and speciation in the genus Peromyscus (Dice, 1940, 1968), but it is not likely that the same model of speciation will apply equally to all organisms (White, 1973; Dobzhansky, 1972). The California vole promises to provide another important case study of the speciation process and of the nature of genetic changes involved in it.     

Programmed and flexible: long‐term Zugunruhe data highlight the many axes of variation in avian migratory behaviour

Studies of Zugunruhe – the ‘migratory restlessness’ behaviour of captive birds – have been integral to our understanding of animal migration, revealing an inherited propensity to migrate and an endogenous timing and navigation system. However, differences between Zugunruhe in captivity and migration in the wild call for more data, in particular on variation within and among taxa with diverse migration strategies. Here, we characterise Zugunruhe in a long‐term dataset of activity profiles from stonechats (genus Saxicola) with diverse migratory phenotypes (976 migration periods from 414 birds), using a flexible and consistent quantitative approach based on changepoint analysis. For east African, Austrian, Irish, and Siberian stonechats and hybrids, we report key inter‐population differences in the occurrence, timing, and intensity of Zugunruhe. In line with expectations, we found the highest Zugunruhe intensity in the longest‐distance migrants, more variable patterns in short‐distance migrants, and intermediate characteristics of hybrids relative to their parental groups. Inter‐population differences imply high evolutionary lability of Zugunruhe timing within a robustly structured annual cycle. However, counter to theory, Irish partial migrants showed no segregation between migrant and resident individuals, and previously reported nocturnal restlessness was confirmed for resident African stonechats. Further features of nocturnal restlessness that did not align with migratory behaviour of stonechats were juvenile nocturnal restlessness even prior to postjuvenile moult, and protandry in spring, although stonechats winter in heterosexual pairs. Importantly, Zugunruhe of all populations declined with age, and the intensity of an individual bird's Zugunruhe was correlated with activity levels during other parts of the annual cycle. Our results confirm endogenous, population‐specific migration programmes but also reveal apparent discrepancies between Zugunruhe and migration in the wild. We thus highlight both the continued potential of Zugunruhe study and the need for circumspect interpretation when using migratory restlessness to make inferences about migration in the wild.     

Correlated patterns of genetic diversity and differentiation across an avian family

Comparative studies of closely related taxa can provide insights into the evolutionary forces that shape genome evolution and the prevalence of convergent molecular evolution. We investigated patterns of genetic diversity and differentiation in stonechats (genus Saxicola), a widely distributed avian species complex with phenotypic variation in plumage, morphology and migratory behaviour, to ask whether similar genomic regions have become differentiated in independent, but closely related, taxa. We used whole‐genome pooled sequencing of 262 individuals from five taxa and found that levels of genetic diversity and divergence are strongly correlated among different stonechat taxa. We then asked whether these patterns remain correlated at deeper evolutionary scales and found that homologous genomic regions have become differentiated in stonechats and the closely related Ficedula flycatchers. Such correlation across a range of evolutionary divergence and among phylogenetically independent comparisons suggests that similar processes may be driving the differentiation of these independently evolving lineages, which in turn may be the result of intrinsic properties of particular genomic regions (e.g. areas of low recombination). Consequently, studies employing genome scans to search for areas important for reproductive isolation or adaptation should account for corresponding regions of differentiation, as these regions may not necessarily represent speciation islands or evidence of local adaptation.     

Intraspecific comparison of population structure, genetic diversity, and dispersal among three subspecies of Townsend’s big-eared bats, Corynorhinus townsendii townsendii, C. t. pallescens, and the endangered C. t. virginianus

Townsend’s big-eared bat, Corynorhinus townsendii, is distributed broadly across western North America and in two isolated, endangered populations in central and eastern United States. There are five subspecies of C. townsendii; C. t. pallescens, C. t. australis, C. t. townsendii, C. t. ingens, and C. t. virginianus with varying degrees of concern over the conservation status of each. The aim of this study was to use mitochondrial and microsatellite DNA data to examine genetic diversity, population differentiation, and dispersal of three C. townsendii subspecies. C. t. virginianus is found in isolated populations in the eastern United States and was listed as endangered under the Endangered Species Act in 1979. Concern also exists about declining populations of two western subspecies, C. t. pallescens and C. t. townsendii. Using a comparative approach, estimates of the genetic diversity within populations of the endangered subspecies, C. t. virginianus, were found to be significantly lower than within populations of the two western subspecies. Further, both classes of molecular markers revealed significant differentiation among regional populations of C. t. virginianus with most genetic diversity distributed among populations. Genetic diversity was not significantly different between C. t. townsendii and C. t. pallescens. Some populations of C. t. townsendii are not genetically differentiated from populations of C. t. pallescens in areas of sympatry. For the western subspecies gene flow appears to occur primarily through male dispersal. Finally, geographic regions representing significantly differentiated and genetically unique populations of C. townsendii virginianus are recognized as distinct evolutionary significant units.     

Gonadal status upon spring arrival in long-distance and short-distance migrating stonechats (Saxicola torquata)

Long-distance migration is often associated with relatively short breeding seasons and a start of reproductive activities shortly after arrival. The full activation of the reproductive system from the regressed state takes, however, several weeks and must, therefore, be initiated in the winter quarters or during spring migration. Hence, long-distance migrants face a potential conflict between the energetic and temporal requirements of migration and the preparation for reproduction. We studied long-distance migratory Siberian stonechats in northern Kazakhstan and short- distance migratory European stonechats in Slovakia. We hypothesized that migratory distance and gonadal status at the time of arrival are related. We found that males of both populations arrived with gonads that were not fully developed. However, the populations neither differed in gonadal state at the time of arrival, nor in the rate of testicular development to the fully active state at the time of egg laying. The rate of the last stages of gonadal development may be determined by physiological constraints rather than by a trade-off between migration and reproduction. Within populations, passage migrants and local breeders could not be distinguished on the basis of their testicular development. However, passage migrants showed higher variation in gonadal size than local breeders, which could relate to the differences in migratory distance and hence reproductive timing.     

The mitochondrial cytochrome c oxidase I gene reveals phylogeographic structure in the African Goshawk Accipiter tachiro (Accipitridae)

We used a 298?bp fragment of the mitochondrial cytochrome c oxidase subunit I gene (COI) to examine sequence variation in (mostly) museum specimens of the African Goshawk Accipiter tachiro. Our results showed two clades with high bootstrap support in a phylogenetic analysis and two groups in a nonmetric multidimensional scaling (NMDS) analysis. Each of the two phylogenetic clades corresponded to one of the NMDS groups. One clade comprised haplotypes of the subspecies A. t. lopezi, A. t. macroscelides, A. t. toussenelli and A. t. canescens and corresponded to the morphospecies A. toussenelii. This taxon has a more north-western distribution. The second clade comprised haplotypes of the subspecies A. t. sparsimfasciatus, A. t. pembaensis and A. t. tachiro and corresponded to the morphospecies A. tachiro, which has a more south-eastern distribution. Furthermore, one branch corresponded to the morphospecies A. t. unduliventer, which is confined to the Ethiopian highlands. The genetic divergence observed among the three A. tachiro morphospecies appeared concordant with the ecological and morphological divergence and suggests the existence of three putative species. Within A. tachiro and A. toussenelii there is substantial morphological, but very little genetic, differentiation among subspecies.     

Zugunruhe of migratory and non-migratory birds in a circannual context

Recent findings in non-migratory birds have reopened questions about the interpretation and seasonal organization of Zugunruhe . I address the relationship between Zugunruhe and migration by comparing underlying circannual patterns in captive populations of migratory and non-migratory stonechats. Zugunruhe was highly variable and lacked clear periodicity, indicating its sensitivity to external cues. Patterns of Zugunruhe were similar in African residents, European short-distance migrants, and Siberian long-distance migrants, revealing no major difference in circannual organization. Moult was regulated independently of Zugunruhe and timed more rigidly, particularly in stonechats from equatorial Africa. The persistent and variable circannual patterns of Zugunruhe suggest that non-migratory and migratory stonechats have similar underlying programs but have modified the expression of actual migration. The findings, together with published observations from other species, emphasize the importance of considering programs for migration in a wide range of species, without losing sight of its environmental context.     

Geographical Variation of Allele Frequencies at the LDH-A* Locus in the Arctic Grayling Thymallus arcticus Pall. and in the European Grayling Thymallus thymallus L.

The allele frequencies of LDH-A* locus were studied in the populations of Siberian grayling from the Kozhym River (Pechora basin) and in the population of European grayling from Pechora, Mezen', and Vym' rivers (Northern Dvina basin). In samples of both species (n = 134), three LDH-A phenotypes have been identified in total, which proved to be under the control of two alleles: LDH-A*100 and LDH-A*50. The alternative alleles of LDH-A* locus were identified in the populations of Siberian grayling from Kozhym River and in the population of European grayling from the same river and other Pechora tributaries, namely, LDH-A*100 and LDH-A*50 in the Siberian and the European grayling, respectively. However, in the European grayling populations from the Mezen' and Vym' rivers, both alleles occur at the frequencies of the rare LDH-A*100 allele of 0.143 and 0.222, respectively. According to the published data, the frequency of LDH-A*100 allele increases in the European grayling populations of northwestern (Finland) and southern Europe (France) rivers, reaching 0.872 and 1.000 in Rhone and Loire, respectively, i.e., the values characteristic of the Siberian grayling populations.     

Phylogeography of the blue tit ( Parus teneriffae -group) on the Canary Islands based on mitochondrial DNA sequence data and morphometrics

An analysis of the sequences of the mitochondrial cytochrome b gene (1005 bp) of the Parus teneriffae-group from the Canary Islands and North Africa revealed new insights into the phylogeography of this taxon. The origin of the radiation on the Canarian Archipelago was apparently one of the central islands—Tenerife or Gran Canaria. The populations on El Hierro (P. t. ombriosus) and La Palma (P. t. palmensis) represent distinct monophyletic lineages. Blue tits from Gran Canaria are genetically distinct from those of La Gomera and Tenerife (P. t. teneriffae), which supports the results of other studies and suggests the existence of an—until now—undescribed taxon there. In contrast, the populations on the eastern islands of Fuerteventura and Lanzarote (P. t. degener) could not be distinguished from North African blue tits (P. t. ultramarinus), and these populations should be subsumed under the subspecies ultramarinus. Taxonomic recommendations based on these results include the distinction of the northern European P. caeruleus from P. teneriffae, including blue tits from North Africa and the Canary Islands, the treatment of degener and ultramarinus as synonymous (P. teneriffae ultramarinus) and a new blue tit taxon on the island of Gran Canaria (P. t. hedwigii nov. ssp.), which is formally described. The genetic results are in parts supported by bioacoustic and morphological data.     

From Asia to Europe across Siberia: phylogeography of the Siberian spined loach (Teleostei,Cobitidae)

We assessed the genetic variability of the Siberian spined loach Cobitis melanoleuca across its unusually broad distribution that encompasses areas greatly affected by Pleistocene glaciations. Due to extensive morphological variation among their populations, the taxonomic status of C. melanoleuca is complicated. It is unclear whether C. melanoleuca represents a single taxonomic unit or contains several species or subspecies. Our analyses showed low genetic variability in all populations without any phylogenetic structure. The absence of molecular distinctiveness indicates the conspecificity of all C. melanoleuca populations. Only a few common haplotypes shared by East Asian, Siberian and European populations were found at high frequency in the nuclear genes analysed. At the mitochondrial level, Siberian populations shared haplotypes with populations located at both extremes of the species’ range suggesting central populations as a source of current mitochondrial variability. Unimodal mismatch distributions and significant values from neutrality tests support a recent expansion of C. melanoleuca. Our time estimates suggest a postglacial colonisation of European waters around 1.0 MYA, indicating that C. melanoleuca may represent the last cobitid immigrant in Europe that used the northern route across Siberia to expand its range.     

Ecological traits influence the current distribution and range of an island endemic bird

Aim  The aim of this paper is to investigate the causes of the current restricted distribution of a narrow-range endemic bird species, the Canary Islands stonechat, Saxicola dacotiae .
Location  Eastern islands of the Canary Islands archipelago.
Methods  We compared climatic patterns (temperature and rainfall), habitat and microhabitat structure, food availability during a full annual cycle, and the abundance of native avian competitors and predators inside and outside the species' range. Three study areas, located in similar habitats on nearby islands, were studied: northern Fuerteventura, close to the northern border of the species' range; southern Lanzarote, 22 km from the nearest site occupied by stonechats; and the Lobos islet, 10 km from the nearest occupied site and 2 km from the coast of Fuerteventura.
Results  The cover of suitable habitats (slopes with high cover of large shrubs, stony fields and ravines) and microhabitats (shrubs and boulders) and the abundance of arthropods during the breeding period of Canary Islands stonechats were lower outside than inside the species' range. Temperature, rainfall and the abundance of competitors and predators inside and outside the species' range did not differ significantly.
Main conclusions  Ecological requirements explaining the distribution of the Canary Islands stonechat within its range seem to be the main factor hindering its settlement on nearby islands. Geological and palaeoclimatic processes, as well as past and current human impact, could also have constrained the distribution of this narrow-range endemic bird species.     

Following the cold: geographical differentiation between interglacial refugia and speciation in the arcto‐alpine species complex Bombus monticola (Hymenoptera: Apidae)

Cold‐adapted species are expected to have reached their largest distribution range during a part of the Ice Ages whereas postglacial warming has led to their range contracting toward high‐latitude and high‐altitude areas. This has resulted in an extant allopatric distribution of populations and possibly to trait differentiations (selected or not) or even speciation. Assessing inter‐refugium differentiation or speciation remains challenging for such organisms because of sampling difficulties (several allopatric populations) and disagreements on species concept. In the present study, we assessed postglacial inter‐refugia differentiation and potential speciation among populations of one of the most common arcto‐alpine bumblebee species in European mountains, Bombus monticola Smith, 1849. Based on mitochondrial DNA/nuclear DNA markers and eco‐chemical traits, we performed integrative taxonomic analysis to evaluate alternative species delimitation hypotheses and to assess geographical differentiation between interglacial refugia and speciation in arcto‐alpine species. Our results show that trait differentiations occurred between most Southern European mountains (i.e. Alps, Balkan, Pyrenees, and Apennines) and Arctic regions. We suggest that the monticola complex actually includes three species: B. konradini   stat.n. status distributed in Italy (Central Apennine mountains), B. monticola with five subspecies, including B. monticola mathildis   ssp.n. distributed in the North Apennine mountains ; and B. lapponicus . Our results support the hypothesis that post‐Ice Age periods can lead to speciation in cold‐adapted species through distribution range contraction. We underline the importance of an integrative taxonomic approach for rigorous species delimitation, and for evolutionary study and conservation of taxonomically challenging taxa.     

Phylogeography of the Siberian newt Salamandrella keyserlingii by mitochondrial DNA sequence analysis

Differentiation of geographical populations of the Siberian newt Salamandrella keyserlingii throughout the species range was analyzed using a fragment of the cytochrome b gene. The population of the Primorye region (Russian Far East) is separated to the greatest extent; the Japanese and South Kuril populations are the next most separate. These populations are possibly subspecies. Geographical differentiation of populations in the Siberian part of the species range is lower, lacks a clinal variation, and is irregular. The molecular variation of S. keyserlingii supports the hypothesis that several primary vicarious refugia of pre-Pleistocene differentiation of a common ancestor of Salamandrella occurred in the southeastern part of its current distribution range and that northern and western regions were gradually colonized via repeated steps of expansion and retreat in the Siberian part of the modern species range.