Genetic structure of the <Emphasis Type="Italic">Anthyllis vulneraria</Emphasis> L. s. l. species complex in Estonia based on AFLPs

Anthyllis vulneraria L. (Fabaceae) s. lato includes many cryptic taxa, ranging from 25 to 60 subspecies according to different authors. The delimitation of intraspecific taxa of A. vulneraria s. lato has always been complicated and inconsistent. Different data sets (multivariate analyses of morphological variation, allozymes, chloroplast SSRs and ITS) have not resolved the existing problem with distinguishing some subspecies. We used the amplified fragment length polymorphism (AFLP) analysis to describe the differentiation in this species complex and to characterize variation on a geographic scale. Some correlation was found between genetic variability and geographic distribution (western-eastern directional variation), but AFLP data analysis did not reveal clear intraspecific structure of the seven analysed taxa. The analysed specimens did not comprise groups correlated with the subspecies.     

Size variation facilitates population divergence but does not explain it all: an example study from a widespread African monkey

Subspecific variation is widespread in vertebrates. Within Africa, several mammals have extensive geographic distributions with attendant morphological, ecological, and behavioural variations, which are often used to demarcate subspecies. In the present study, we use a primate species, the vervet monkey, Cercopithecus aethiops, as a case study for intraspecific divergence in widespread mammals, assessed through hard tissue morphology. We examine intraspecific differences in size, shape, and non‐allometric shape from a taxonomic perspective, and discuss the macroevolutionary implications of findings from microevolutionary analyses of geographic variation. A geometric morphometric approach was used, employing 86 three‐dimensional landmarks of almost 300 provenanced crania. Many of the taxonomic differences in skull morphology between vervet populations appear to be related to geographic proximity, with subspecies at opposite extremes of a west‐to‐east axis showing greatest divergence, and populations from central and south Africa being somewhat intermediate. The classification rate from discriminant analyses was lower than that observed in other African primate radiations, including guenons as a whole and red colobus. Nonetheless, taxonomic differences in shape were significant and not simply related to either geography or size. Thus, although shifts in size may be an important first step in adaptation and diversification, with size responding more quickly than shape to environmental change, the six vervet taxa currently recognized (either as species or subspecies) are not simply allometrically scaled versions of one another and are probably best viewed as subspecies. Holding allometry constant when examining inter‐population differences in shape may thus help to reveal the early stages of evolutionary divergence. The vervet case study presented here hence has relevance for future studies examining intraspecific differentiation in other large mammals, particularly through the methods used to identify small but biologically meaningful divergence, with attendant implications for conservation planning. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 823–843.     

How species longevity, intraspecific morphological variation, and geographic range size are related: a comparison using late Cambrian trilobites

Phenotypic variation is fundamental to evolutionary change. Variation not only evinces the connectivity of populations but it is also associated with the adaptability and evolvability of taxa. Despite the potential importance of morphological variation in structuring evolutionary patterns, little is known about how relative differences in intraspecific morphological variation and its geographic structure are linked to differences in species longevity. This study offers a novel combination of analyses that reveal the quantitative relationships among intraspecific variation, geographic range size and duration in the fossil record using late Cambrian trilobites. Results show that geographic range size and duration are positively correlated. Surprisingly, longer lived species tend to have less intraspecific variation. Phylogenetic effects were also explored and found not to determine the association between these variables. However, the distribution of geographic range sizes shows strong phylogenetic signal. In light of previous work, one possible explanation for these results is that species with shorter durations have comparatively higher rates of morphological evolution, reflected in higher phenotypic variation overall.     

Systematics and evolutionary relationships of the mountain lizard Liolaemus monticola (Liolaemini): how morphological and molecular evidence contributes to reveal hidden species diversity

The delimitation of species is a major issue in systematic biology and has been a re-emerging discipline in the last decade. A number of studies have shown that the use of multiple data sets is critical for the identification of cryptic species, particularly in groups with complex evolutionary histories. Liolaemus monticola is a montane lizard species distributed in central Chile (32°–42°S), with four described subspecies in a latitudinal gradient from north to south: L. m. monticola , L. m. chillanensis , L. monticola ssp. and L. m. villaricensis . In order to test the systematic status and phylogenetic relationships of the taxa included in the L. monticola group, we analysed morphological (morphometric and meristic) and molecular (allozyme and mitochondrial DNA) data sets. The results of the morphological analyses showed that meristic variables correctly assigned individuals with higher accuracy than did morphometric characters. The results of the analyses of allozyme data revealed eight diagnostic loci that are evidence for significant differences among the four L. monticola subspecies. Phylogenetic analyses with mitochondrial DNA data, including additional species, showed that the L. monticola group is polyphyletic. We postulate that the four current subspecies represent independent evolutionary lineages and must be raised to the specific level as L. monticola , L. chillanensis and L. villaricensis . The taxonomic status of the unnamed L. monticola ssp. remains unresolved, although we provide a preliminary proposal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 635–650.     

Phylogeny and cryptic diversity in geckos (Phyllopezus; Phyllodactylidae; Gekkota) from South America's open biomes

The gecko genus Phyllopezus occurs across South America's open biomes: Cerrado, Seasonally Dry Tropical Forests (SDTF, including Caatinga), and Chaco. We generated a multi-gene dataset and estimated phylogenetic relationships among described Phyllopezus taxa and related species. We included exemplars from both described Phyllopezus pollicaris subspecies, P. p. pollicaris and P. p.przewalskii. Phylogenies from the concatenated data as well as species trees constructed from individual gene trees were largely congruent. All phylogeny reconstruction methods showed Bogertia lutzae as the sister species of Phyllopezus maranjonensis, rendering Phyllopezus paraphyletic. We synonymized the monotypic genus Bogertia with Phyllopezus to maintain a taxonomy that is isomorphic with phylogenetic history. We recovered multiple, deeply divergent, cryptic lineages within P. pollicaris. These cryptic lineages possessed mtDNA distances equivalent to distances among other gekkotan sister taxa. Described P. pollicaris subspecies are not reciprocally monophyletic and current subspecific taxonomy does not accurately reflect evolutionary relationships among cryptic lineages. We highlight the conservation significance of these results in light of the ongoing habitat loss in South America's open biomes.     

Stable climate corridors promote gene flow in the Cape sand snake species complex (Psammophiidae)

Climate shifts during the Quaternary Period have driven changes in regional range dynamics for many species, influencing population structure of species and in some cases promoting speciation. Within southern Africa, the psammophine snakes Psammophis trinasalis and P. namibensis were historically considered subspecies of P. leightoni but were elevated to species rank based on ecological differences. Preliminary phylogenetic analyses suggested intraspecific, not interspecific genetic variation between these taxa, but this finding was based on very limited data and could not be confirmed. To assess the level of genetic differentiation within the P. leightoni species complex, we explored the evolutionary history of these snakes by combining phylogenetic analyses, species distribution modelling and an examination of morphology. We generated a comprehensive, multi-gene phylogeny for Psammophis that included wider geographic sampling of the three species in the complex. Using this phylogeny, Bayesian and distance-based species delimitation analyses showed intraspecific, not interspecific divergences between taxa in the complex, suggesting that they collectively represent a single taxon. Furthermore, non-metric multidimensional scaling analysis of scalation characters showed no differences between the species. Moreover, palaeo-modelling at three time periods since the last interglacial period suggest that there have been varying levels of connectivity between these taxa, which has likely facilitated gene flow between them. Given the evidence, we propose that the P. leightoni complex represents a single species and therefore formally synonymise the three species.     

Phylogeography of Aglais urticae (Lepidoptera) based on DNA sequences of the mitochondrial COI gene and control region

Mitochondrial DNA (mtDNA) sequences of the COI gene and the control region were used to examine the genetic population structure of Aglais urticae L. (Lepidoptera) over its entire geographic range, i.e., the Palaearctic. The phylogenetic relationships within and between A. urticae subspecies were determined and patterns of mtDNA divergence and ecological differentiation were compared. High gene flow together with a recent and sudden population expansion characterise the genetic population structure of this species. No geographically induced differentiation was observed, nor were subspecies identified as separate evolutionary units. The discrepancy between the genetic and ecological variation is most likely due to the slower rate of mtDNA evolution compared to ecological differentiation. The control region proved to be a less useful molecular marker for the population genetics and the phylogenetic reconstruction of closely related taxa in A. urticae than it has for other species. The extreme bias in adenine and thymine content (A+T=90.91%) probably renders this region highly susceptible to homoplasy, resulting in a less informative molecular marker.     

Allopatric origin of cryptic butterfly species that were discovered feeding on distinct host plants in sympatry

Surveys of tropical insects are increasingly uncovering cryptic species – morphologically similar yet reproductively isolated taxa once thought to comprise a single interbreeding entity. The vast majority of such species are described from a single location. This leaves us with little information on geographic range and intraspecific variation and limits our ability to infer the forces responsible for generating such diversity. For example, in herbivorous and parasitic insects, multiple specialists are often discovered within what were thought to be single more generalized species. Host shifts are likely to have contributed to speciation in these cases. But when and where did those shifts occur, and were they facilitated by geographic isolation? We attempted to answer these questions for two cryptic species within the butterfly Cymothoe egesta that were recently discovered on different host plants in central Cameroon. We first used mtDNA markers to separate individuals collected on the two hosts within Cameroon and then extended our analysis to incorporate individuals collected across the entire pan‐Afrotropical range of the original taxon. To our surprise, we found that the species are almost entirely allopatric, dividing the original range and overlapping only in the narrow zone of West‐Central Africa where they were first discovered in sympatry. This finding, combined with analyses of genetic variation within each butterfly species, strongly suggests that speciation occurred in allopatry, probably during the Pleistocene. We discuss the implications of our results for understanding speciation among other cryptic species recently discovered in the tropics and argue that more work is needed on geographic patterns and host usage in such taxa.     

Getting western: biogeographical analysis of morphological variation,mitochondrial haplotypes and nuclear markers reveals cryptic species and hybrid zones in the Junonia butterflies of the American southwest and Mexico

The American southwest and northern Mexico has a great degree of endemic diversity compared with the rest of North America. The Pleistocene glaciations and the dispersal of species from glacial refuges in this region have been important engines for the production of biodiversity in the region. The New World Junonia are a recent radiation of butterflies that are thought to have spent time in these refuges during periods of glacial advancement. We have reconstructed the plausible movements and the contemporary geographic distributions of the five taxa (J. coenia, J. grisea, J. litoralis, J. nigrosuffusa and J. zonalis) that occur in the American southwest and northern Mexico using phenotypic and genotypic information primarily from specimens preserved in museum collections, supplemented with additional contemporary collections. Evidence of cryptic species and hybridization events were observed using mitochondrial haplotypes, genotypic variation at the nuclear wingless locus, multilocus DNA fingerprinting, patterns of larval host plant use, variation in life‐history traits and morphological characteristics. Based on these lines of evidence, and in spite of low levels of hybridization between them, we argue that all five Junonia taxa are independent evolutionary lineages. Junonia grisea and J. coenia are morphologically very similar, but differences in morphology, life‐history traits, nuclear genotypes and mitochondrial haplotypes suggest that they are a cryptic species pair, thus elevating J. grisea comb.n. to a full species when it had previously been considered to be a subspecies of J. coenia.     

Cranial ontogeny of Papio baboons (Papio hamadryas)

Cranial form in subspecies of Papio baboons (Papio hamadryas) varies in relation to size, geography, and sex. However, knowledge about this variation is based mainly on adults, precluding direct assessments of the evolutionary factors that are ultimately responsible for adult shape variation. Consequently, this study tests hypotheses about the development of size and shape differences among subspecies of Papio baboons, anticipating limited evolutionary divergences in the ontogenetic pathways leading to adult endpoints. Geometric morphometric and bivariate allometric analyses are used to explore developmental size and shape variation. Allometric scaling in adult Papio baboons occurs because both sexes and all subspecies follow similar developmental pathways to a variety of adult forms. However, complex allometry contributes to form differences, producing potentially important shape differences that emerge during development. Modest shape differences that are statistically independent of size distinguish chacma baboons (P. h. ursinus) from other forms. A small-headed subspecies, the Kinda baboon (P. h. kindae), also presents a distinctive ontogeny, and may provide insights into the evolution of size change in this species. Variation among subspecies that is statistically independent of size involves the rostrum, zygomatic breadths, and cranial flexion. These features may be related to diet, but the precise biomechanical correlates of baboon form variation remain unclear.     

Cryptic species as a window into the paradigm shift of the species concept

The species concept is the cornerstone of biodiversity science, and any paradigm shift in the delimitation of species affects many research fields. Many biologists now are embracing a new “species” paradigm as separately evolving populations using different delimitation criteria. Individual criteria can emerge during different periods of speciation; some may never evolve. As such, a paradigm shift in the species concept relates to this inherent heterogeneity in the speciation process and species category—which is fundamentally overlooked in biodiversity research. Cryptic species fall within this paradigm shift: they are continuously being reported from diverse animal phyla but are poorly considered in current tests of ecological and evolutionary theory. The aim of this review is to integrate cryptic species in biodiversity science. In the first section, we address that the absence of morphological diversification is an evolutionary phenomenon, a “process” counterpart to the long‐studied mechanisms of morphological diversification. In the next section regarding taxonomy, we show that molecular delimitation of cryptic species is heavily biased towards distance‐based methods. We also stress the importance of formally naming of cryptic species for better integration into research fields that use species as units of analysis. Finally, we show that incorporating cryptic species leads to novel insights regarding biodiversity patterns and processes, including large‐scale biodiversity assessments, geographic variation in species distribution and species coexistence. It is time for incorporating multicriteria species approaches aiming to understand speciation across space and taxa, thus allowing integration into biodiversity conservation while accommodating for species uncertainty.     

Range size heritability in Carnivora is driven by geographic constraints

Range size heritability refers to an intriguing pattern where closely related species occupy geographic ranges of similar extent. Its existence may indicate selection on traits emergent only at the species level, with interesting consequences for evolutionary processes. We explore whether range size heritability may be attributable to the fact that range size is largely driven by the size of geographic domains (i.e., continents, biomes, areas given by species' climatic tolerance) that tend to be similar in phylogenetically related species. Using a well-resolved phylogeny of Carnivora, we show that range sizes are indeed constrained by geographic domains and that the phylogenetic signal in range sizes diminishes if the domain sizes are accounted for. Moreover, more detailed delimitation of species' geographic domain leads to a weaker signal in range size heritability, indicating the importance of definition of the null model against which the pattern is tested. Our findings do not reject the hypothesis of range size heritability but rather unravel its underlying mechanisms. Additional analyses imply that evolutionary conservatism in niche breadth delimits the species' geographic domain, which in turn shapes the species' range size. Range size heritability patterns thus emerge as a consequence of this interplay between evolutionary and geographic constraints.     

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.     

Sequence variation in the coding region of the melanocortin-1 receptor gene (MC1R) is not associated with plumage variation in the blue-crowned manakin (Lepidothrix coronata)

Avian plumage traits are the targets of both natural and sexual selection. Consequently, genetic changes resulting in plumage variation among closely related taxa might represent important evolutionary events. The molecular basis of such differences, however, is unknown in most cases. Sequence variation in the melanocortin-1 receptor gene (MC1R) is associated with melanistic phenotypes in many vertebrate taxa, including several avian species. The blue-crowned manakin (Lepidothrix coronata), a widespread, sexually dichromatic passerine, exhibits striking geographic variation in male plumage colour across its range in southern Central America and western Amazonia. Northern males are black with brilliant blue crowns whereas southern males are green with lighter blue crowns. We sequenced 810 bp of the MC1R coding region in 23 individuals spanning the range of male plumage variation. The only variable sites we detected among L. coronata sequences were four synonymous substitutions, none of which were strictly associated with either plumage type. Similarly, comparative analyses showed that L. coronata sequences were monomorphic at the three amino acid sites hypothesized to be functionally important in other birds. These results demonstrate that genes other than MC1R underlie melanic plumage polymorphism in blue-crowned manakins.     

Geographic variation and the evolution of song in Mesoamerican rufous‐naped wrens Campylorhynchus rufinucha

Speciation may be influenced by geographic variation in animal signals, particularly when those signals are important in reproductive decisions. Here, we describe patterns of geographic variation in the song of rufous‐naped wrens Campylorhynchus rufinucha. This species complex is a morphologically variable taxon confined to tropical dry forest areas from Mexico to northwestern Costa Rica. Morphological and genetic analyses suggest that there are at least three partially isolated groups within the complex, including a secondary‐contact zone in coastal western Chiapas between the subspecies C. r. humilus and C. r. nigricaudatus. Based on recordings throughout their geographic range, we investigate the effects of historical isolation on song structure and analyze whether genetic differences or climatic conditions explain observed patterns of variation. Our findings, based on a culturally‐transmitted and sexually‐selected trait, support the hypothesis that three evolutionary units exist within this taxon. Our results suggest that song differences between genetic groups were influenced by historical isolation. We report a strong relationship between vocal dissimilarity and genetic distance, suggesting that differences in vocal characteristics are probably affected by the same factors that drive genetic divergence. We argue that the evolution of song in this taxon is influenced by vicariant events, followed by accumulation of changes in song structure due to several possible factors: cultural drift in song structure; genetic drift in features related to song production; or natural selection acting on features that influence songs, such as body and beak size.     

Testing evolutionary hypotheses for DNA barcoding failure in willows

The goal of DNA barcoding is to enable the rapid identification of taxa from short diagnostic DNA sequence profiles. But how feasible is this objective when many evolutionary processes, such as hybridization and selective sweeps, cause alleles to be shared among related taxa? In this issue of Molecular Ecology, Percy et al. (2014) test the full suite of seven candidate plant barcoding loci in a broad geographic sample of willow species. They show exceptional plastid haplotype sharing between species across continents, with most taxa not possessing a unique barcode sequence. Using population genetic and molecular dating analyses, they implicate hybridization and selective sweeps, but not incomplete lineage sorting, as the historical processes causing widespread haplotype sharing among willow taxa. This study represents an exceptional case of how poorly barcoding can perform, and highlights methodological issues using universal organellar regions for species identification.     

Specialist and generalist symbionts show counterintuitive levels of genetic diversity and discordant demographic histories along the Florida Reef Tract

Specialist and generalist life histories are expected to result in contrasting levels of genetic diversity at the population level, and symbioses are expected to lead to patterns that reflect a shared biogeographic history and co-diversification. We test these assumptions using mtDNA sequencing and a comparative phylogeographic approach for six co-occurring crustacean species that are symbiotic with sea anemones on western Atlantic coral reefs, yet vary in their host specificities: four are host specialists and two are host generalists. We first conducted species discovery analyses to delimit cryptic lineages, followed by classic population genetic diversity analyses for each delimited taxon, and then reconstructed the demographic history for each taxon using traditional summary statistics, Bayesian skyline plots, and approximate Bayesian computation to test for signatures of recent and concerted population expansion. The genetic diversity values recovered here contravene the expectations of the specialist–generalist variation hypothesis and classic population genetics theory; all specialist lineages had greater genetic diversity than generalists. Demography suggests recent population expansions in all taxa, although Bayesian skyline plots and approximate Bayesian computation suggest the timing and magnitude of these events were idiosyncratic. These results do not meet the a priori expectation of concordance among symbiotic taxa and suggest that intrinsic aspects of species biology may contribute more to phylogeographic history than extrinsic forces that shape whole communities. The recovery of two cryptic specialist lineages adds an additional layer of biodiversity to this symbiosis and contributes to an emerging pattern of cryptic speciation in the specialist taxa. Our results underscore the differences in the evolutionary processes acting on marine systems from the terrestrial processes that often drive theory. Finally, we continue to highlight the Florida Reef Tract as an important biodiversity hotspot.

     

Infraspecific classification reflects genetic differentiation in the widespread Petunia axillaris complex: A comparison among morphological,ecological, and genetic patterns of geographic variation

Plants are subjected to natural selection during expansion of their geographic range or when facing changes in environmental conditions, which in turn may affect phenotypic diversity. Studies on geographic phenotypic variation provide insight into the evolutionary processes and have long contributed to better understanding of diversification within and among species. Petunia axillaris is widely distributed in temperate South America, occurring throughout the Pampas region. The current taxonomy of this species recognizes three allopatric subspecies, which occupy nearly adjacent territories, according to floral morphology: P. axillaris subsp. axillaris, P. axillaris subsp. parodii, and P. axillaris subsp. subandina. In this study, we sampled the three P. axillaris subspecies and used both molecular markers (from the plastid and nuclear genomes) and morphological measurements to investigate how genetic diversity and morphological variation correlate to ecological variables and the geographic context. We used different forms of Mantel tests (partial and correlograms) to investigate the geographic distribution patterns in distinct types of similarity/dissimilarity among the populations and their relationships. We also modeled the morphological variation in P. axillaris as a function of the genetic marker frequencies in the populations. We found that the morphological differences leading to the recognition of different subspecies of P. axillaris reflect historical processes of isolation and that adaptation to different ecological conditions faced by each lineage is perhaps not merely a consequence of phenotypic plasticity. These findings suggest that differences between subspecies could represent an incipient speciation stage.     

Testing reticulate evolution of four Vitis species from East Asia using restriction‐site associated DNA sequencing

Reticulate evolution is an important driving force of angiosperm evolution. It has been proposed as an important evolutionary process in Vitis L. subgenus Vitis. Events of natural hybridization and introgression of several taxa native to North America have been hypothesized and discussed. However, there is no convincing evidence of reticulate evolution reported for closely related Vitis taxa from East Asia. We aim to explore natural hybridization and introgression among four closely related Vitis taxa from East Asia (V. amurensis Ruprecht, V. romanetii Romanet du Caillaud, V. shenxiensis C. L. Li, and V. piasezkii Maximowicz) with the restriction‐site associated DNA sequencing technique. A total of 46 accessions, covering the potential morphological and geographic variation of each species, are sequenced. Our results show a complex evolutionary pattern of the four Vitis species. The phylogenetic inference suggests that V. amurensis is monophyletic, however, V. romanetii, V. shenxiensis, and V. piasezkii do not appear to be monophyletic. Significant signals of introgression in some accessions have been detected by population structure analyses. D‐statistics analysis and population structure analyses support the presence of introgression between V. shenxiensis/V. piasezkii and V. romanetii in sympatric populations, but a strong signal of admixture has not been recognized between distantly located populations. Our results provide strong evidence of reticulate evolution among V. romanetii, V. shenxiensis, and V. piasezkii.     

Molecular systematics,hybridization, and phylogeography of the Bufo americanus complex in Eastern North America

We reconstruct phylogenetic relationships among a well-studied group of toads and find relationships that differ greatly from the current taxonomic understanding. We use mitochondrial sequences encoding ND1, tRNA(Leu(UUR)), and part of 16S to infer relationships among members of the Bufo americanus complex. Focusing on the four taxa that historically have been most problematic due to morphological similarity and hybridization in sympatry, we sample 150 individuals from multiple populations across each species' geographic range. Our evidence conflicts with previous taxonomic hypotheses that were based on ability to hybridize, geographic distribution, and call variation. First, sequences from B. fowleri do not comprise the sister clade to sequences of B. woodhousii; therefore the previous classifications of B. fowleri as sister species to, or eastern subspecies of, B. woodhousii are both called into question. Second, sequences from B. americanus are more closely related to those of B. woodhousii than to those of B. terrestris, indicating that similar advertisement call characteristics evolved independently. Third, sequences of B. fowleri are paraphyletic, with sequences of B. terrestris embedded within. Lastly, sequences from B. fowleri cluster into three distinct mitochondrial clades, with some divergences corresponding to greater than 2mya. These clades are somewhat geographically structured, suggesting divergence in allopatry during the Pleistocene. These mitochondrial divergences are not accompanied by known phenotypic differences, however, suggesting either evolutionary stasis in morphology and behavior, cryptic phenotypic evolution, or that hybridization in secondary contact has homogenized phenotypic differences that may have arisen in allopatry.