General Lineage Concept Research Articles

Diversification of the African legless skinks in the subfamily Acontinae (Family Scincidae)

Cladogenic diversification is often explained by referring to climatic oscillations and geomorphic shifts that cause allopatric speciation. In this regard, southern Africa retains a high level of landscape heterogeneity in vegetation, geology, and rainfall patterns. The legless skink subfamily Acontinae occurs broadly across the southern African subcontinent and therefore provides an ideal model group for investigating biogeographic patterns associated with the region. A robust phylogenetic study of the Acontinae with comprehensive coverage and adequate sampling of each taxon has been lacking up until now, resulting in unresolved questions regarding the subfamily’s biogeography and evolution. In this study, we used multi-locus genetic markers (three mitochondrial and two nuclear) with comprehensive taxon coverage (all currently recognized Acontinae species) and adequate sampling (multiple specimens for most taxa) of each taxon to infer a phylogeny for the subfamily. The phylogeny retrieved four well-supported clades in Acontias and supported the monophyly of Typhlosaurus. Following the General Lineage Concept (GLC), many long-standing phylogenetic enigmas within Acontias occidentalis and the A. kgalagadi, A. lineatus and A. meleagris species complexes, and within Typhlosaurus were resolved. Our species delimitation analyses suggest the existence of hidden taxa in the A. occidentalis, A. cregoi and A. meleagris species groups, but also suggest that some currently recognized species in the A. lineatus and A. meleagris species groups, and within Typhlosaurus, should be synonymised. We also possibly encountered “ghost introgression” in A. occidentalis. Our inferred species tree revealed a signal of gene flow, which implies possible cross-over in some groups. Fossil evidence calibration dating results showed that the divergence between Typhlosaurus and Acontias was likely influenced by cooling and increasing aridity along the southwest coast in the mid-Oligocene caused by the opening of the Drake Passage. Further cladogenesis observed in Typhlosaurus and Acontias was likely influenced by Miocene cooling, expansion of open habitat, uplifting of the eastern Great Escarpment (GE), and variation in rainfall patterns, together with the effect of the warm Agulhas Current since the early Miocene, the development of the cold Benguela Current since the late Miocene, and their co-effects. The biogeographic pattern of the Acontinae bears close resemblance to that of other herpetofauna (e.g., rain frogs and African vipers) in southern Africa.

Integrative approach resolves the taxonomy of Eulaema cingulata (Hymenoptera, Apidae), an important pollinator in the Neotropics

Species delimitation is a rich scientific field that often uses different sources of data to identify independently evolving lineages that might be recognized as species. Here, we use an integrative approach based on morphometrics, COI-barcoding, and phylogenomics using ultraconserved elements (UCEs) to investigate whether the orchid bee species Eulaema cingulata (Fabricius, 1804) and E. pseudocingulata Oliveira, 2006 represent a single variable taxon or two different species. We analyzed 126 specimens across the geographical range of these nominal species to test species hypotheses using the general lineage concept. We found substantial overlap in wing and head morphometrics, and both taxa form one phylogenetic lineage based on COI mitochondrial and UCE data. Our results support the recognition of both forms as members of the same evolutionary unit and E. pseudocingulata is herein recognized as a junior synonym of E. cingulata.

A new species of the genus Oligoryzomys (Rodentia: Cricetidae) from Peru

Abstract The genus Oligoryzomys is the most speciose genus in the tribe Oryzomyini; however, under the general lineage concept of species, at least nine lineages are pending review and naming. In this study, I carry out qualitative and quantitative morphological analyses of specimens of Oligoryzomys andinus, O. aff. andinus, and O. arenalis, to delimit these three lineages. I show that O. aff. andinus is morphologically distinct both from O. andinus and O. arenalis, corroborating that the lineage merits recognition as a species, as suggested by previously published molecular analyses (Hurtado and D’Elía 2019). I therefore describe O. aff. andinus as a new species endemic to the Polylepis forests on the western slope of the Andes and coastal Lomas in central Peru. Finally, I propose two hypotheses about the biogeographical history of O. andinus, O. arenalis, and this new species.

Ecological and spatial patterns associated with diversification of South American Physaria (Brassicaceae) through the general concept of species

Analyzing the roles of ecology and geography on speciation and lineage diversification can shed light on the processes that generate and maintain biodiversity. Additionally, lineages rapidly diversifying across unstable habitats provide substantial challenges for resolving evolutionary histories and delimiting species. Physaria is represented in South American by six species distributed from southern Bolivia to northern central Argentina and growing in highlands of the southern-central Andes, but also along the hills and lowlands of central-eastern Argentina. This biogeographical variability, not common among other South American crucifers (Brassicaceae), prompted us to conduct different climatic niche and geographical range comparisons to study the potential roles of ecology and geography through the diversification of the group. However, the remarkable similarity between these species, coupled with the continuous variability of the diagnostic morphological characters, blurs the species boundaries. Therefore, in order to identify independent evolving lineages, we first employed species delimitation methods together with the general lineage concept of species, and used molecular sequences from nuclear ribosomal and chloroplast loci. Secondly, and in the light of the lineages obtained, we explored the roles of geography and ecology on the diversification of South American Physaria and tested for presence of phylogenetic niche-conservatism or niche-divergence patterns, as well as potential ecological speciation. Lineages identified by these delimitation methods were highly congruent with described species; nevertheless, some morphospecies were included under the same independent evolutionary lineage. Our results suggest that the climatic niche divergence along the heterogeneous landscape apparently was a major factor promoting diversification of the South American Physaria. Divergence was registered mainly on the temperature dimension, which promoted shifts between cold-temperate habitats associated with the highlands of the central-southern Andes and warm lowlands from central-eastern Argentina, i.e., the Monte and Dry Chaco ecoregions. In addition, some degree of niche divergence along the precipitation gradient was also secondarily recovered. Allopatry and dispersal capabilities also seem to be associated with the diversification of the group, presumably through the Late Pliocene-Pleistocene, and promoted by glacial cycles and climatic oscillations during the Quaternary. Results of these analyses are also discussed in a general context, which will contribute to the understanding of the evolutionary and ecological patterns of South American Brassicaceae.

Addressing the Linnean shortfall in a cryptic species complex

AbstractBiodiversity is being lost at an alarming rate, but the rate of this loss is likely to be underestimated as a result of a deficit in taxonomic knowledge (i.e. the Linnean shortfall). This knowledge gap is more extensive for morphologically indistinct taxa. The advancement of molecular techniques and delimitation methods has facilitated the identification of such cryptic species, but a majority of these taxa remain undescribed. To investigate the effects of taxonomic uncertainty on understanding of biodiversity, we applied the general lineage concept of species to an amphipod species complex, the Gammaruslacustris lineage that occupies springs of the northern Chihuahuan Desert, which is emerging in contemporary times. We investigated species boundaries using a validation-based approach and examined genetic structure of the lineage using a suite of microsatellite markers to identify independently evolving metapopulations. Our results show that each spring contains a genetically distinct population that is geographically isolated from other springs, suggesting evolutionary independence and status as separate species. Additionally, we observed subtle interspecific morphological variation among the putative species. We used multiple lines of evidence to formally describe four new species (Gammarus langi sp. nov., G. percalacustris sp. nov., G. colei sp. nov. and G. malpaisensis sp. nov.) endemic to the northern Chihuahuan Desert. Cryptic speciation is likely to be high in other aquatic taxa within these ecosystems, and across arid landscapes throughout North America and elsewhere, suggesting that the magnitude of the Linnean shortfall is currently underestimated in desert springs worldwide.

Phylogenomics and species delimitation for effective conservation of manta and devil rays.

Practical biodiversity conservation relies on delineation of biologically meaningful units. Manta and devil rays (Mobulidae) are threatened worldwide, yet morphological similarities and a succession of recent taxonomic changes impede the development of an effective conservation strategy. Here, we generate genome-wide single nucleotide polymorphism (SNP) data from a geographically and taxonomically representative set of manta and devil ray samples to reconstruct phylogenetic relationships and evaluate species boundaries under the general lineage concept. We show that nominal species units supported by alternative data sources constitute independently evolving lineages, and find robust evidence for a putative new species of manta ray in the Gulf of Mexico. Additionally, we uncover substantial incomplete lineage sorting indicating that rapid speciation together with standing variation in ancestral populations has driven phylogenetic uncertainty within Mobulidae. Finally, we detect cryptic diversity in geographically distinct populations, demonstrating that management below the species level may be warranted in certain species. Overall, our study provides a framework for molecular genetic species delimitation that is relevant to wide-ranging taxa of conservation concern, and highlights the potential for genomic data to support effective management, conservation and law enforcement strategies.

A molecular phylogeny of the genus Laubuka (Teleostei: Cyprinidae) in Sri Lanka reveals multiple origins and a cryptic species

The freshwater-fish genus Laubuka contains ∼13 species distributed through the lowlands of tropical South and South-east Asia. Four of these species (Laubuka lankensis, L. varuna, L. ruhuna, and L. insularis) are reported as endemic to Sri Lanka, a remarkable datum given the island’s small size. We sampled populations of Laubuka at 56 locations in 14 of the island’s principal river basins and analysed their morphological, meristic, phylogenetic, and phylogeographic relationships using the mitochondrial genes cytochrome b and cytochrome c oxidase subunit 1 and the nuclear recombination activating protein 1. We investigated the geographic structure, and delineated species using molecular species delimitation methods and morphological analysis within the general lineage concept of species. Molecular and morphological analyses failed to identify L. ruhuna and L. insularis as distinct species; we show them to be synonyms of L. varuna and L. lankensis, respectively. Taxonomic inflation in previous literature is attributed as a result of limited sampling and negligence of size allometry. Additionally, a third, cryptic species discovered in this study is described as L. hema sp. nov. which is evidently confined to the headwaters of the Gal basin in eastern Sri Lanka. Phylogenetic and haplotype network analyses suggest phylogeographic structure within both the southwest-endemic L. varuna and the dry-zone endemic L. lankensis, but the former shows strong phylogeographic structure between adjacent basins. The Sri Lankan species of Laubuka do not form a monophyletic group: they stem from two dispersal or vicariance events, one involving the lineage that led to L. varuna in south-western Sri Lanka, and the other, of [L. lankensis + L. hema] in the dry zone. http://zoobank.org/urn:lsid:zoobank.org:pub:D2A29329-E0E7-447E-B316-81049212CCCBhttp://zoobank.org/urn:lsid:zoobank.org:act:920E5DC0-CEDC-4243-98C7-12CCF829CBCD

Speciation Associated with Shifts in Migratory Behavior in an Avian Radiation.

Migratory animals move up to thousands of kilometers every year [1]. Losses of migration (i.e., migratory drop-offs) occur when individuals of a migratory species stop migrating and establish founder sedentary populations, a phenomenon documented in birds [2-5] and butterflies [6]. In theory, losses-and also gains-of migration might promote speciation if sedentary and migratory populations become reproductively isolated [7-9]. Because migratory and sedentary strategies involve alternative physiological, behavioral, and morphological traits [10-13], divergence along multiple axes of organismal function is expected to accompany switches in migratory behavior, potentially accelerating speciation. We present evidence of speciation driven by a migratory drop-off in the fork-tailed flycatcher (Tyrannus savana) resulting in reproductive isolation likely driven by changes in breeding schedules (allochronic speciation [13-15]) and geographic isolation of breeding grounds (allopatric speciation [16]). Phylogenetic analyses across New World flycatchers (Tyrannidae) showed that an association between speciation and drop-offs is also observable at a macroevolutionary scale. Loss of migration was significantly more frequent than its gain, and speciation rates of migratory and partially migratory lineages (i.e., species having both migratory and sedentary populations) exceeded those of sedentary lineages. Models of trait evolution indicated that partial migration is an intermediate step between migratory and sedentary states in this family. Given that partial migration is widespread across migratory animals (e.g., of all migratory birds, ca. 51% are partially migratory [5]), speciation via switches in migratory behavior might be an important yet overlooked mechanism of animal diversification.

Reconciling molecules and morphology in North AmericanMartes

Molecular investigations recently resurrected the Pacific marten (Martes caurina) as a species distinct from the American pine marten (M. americana). Previously, 2 hybrid zones for these species were identified, 1 in the northern Rocky Mountains and another on Kuiu Island off the coast of Southeast Alaska. We test the molecular perspectives on species status and hybridization using geometric morphometrics to characterize size and shape differences between North American marten species from 6 genetically defined populations: M. americana from interior Alaska, insular M. caurina from Admiralty Island in Southeast Alaska, M. caurina from Colorado, the 2 previously identified hybrid populations, and a third suspected hybrid population on neighboring Kupreanof Island. We found significant differences in shape supporting the designation of 2 morphological species, with M. caurina exhibiting a broader cranial shape and shorter rostrum. We also found significant size differences between species, and also between insular and mainland populations of M. caurina. Morphological intergradation of M. americana and M. caurina in both insular and mainland hybrid zones identified potentially admixed individuals in 2 of the 3 examined contact zones. Under the general lineage concept, our results support specieslevel classification of both M. americana and M. caurina and suggest that hybridization is geographically limited in scope or may not be effectively detected with morphometrics.

Multivariate species boundaries and conservation of harlequin poison frogs.

In this study, we present an iterative method for delimiting species under the general lineage concept (GLC) based on the multivariate clustering of morphological, ecological and genetic data. Our rationale is that distinct multivariate groups correspond to evolutionarily independent metapopulation lineages because they reflect the common signal of different secondary defining properties (environmental and genetic distinctiveness, phenotypic diagnosability, etc.) that imply the existence of barriers preventing or limiting gene exchange. We applied this method to study a group of endangered poison frogs, the Oophaga histrionica complex. In our study case, we used next-generation targeted amplicon sequencing to obtain a robust genetic data set that we combined with patterns of morphological and ecological features. Our analyses revealed the existence of at least five different species in the histrionica complex (three, new to science), some of them, occurring in small isolated populations outside any protected areas. The lineage delimitation proposed here has important conservation implications as it revealed that some of the Oophaga species should be considered among the most vulnerable of the Neotropical frogs. More broadly, our study exemplifies how multiple-amplicon and multivariate statistical techniques can be integrated to successfully identify species and their boundaries.

Revision of the Eurycea quadridigitata (Holbrook 1842) Complex of Dwarf Salamanders (Caudata: Plethodontidae: Hemidactyliinae) with a Description of Two New Species

Abstract: The Eurycea quadridigitata complex is currently composed of the nominate species and E. chamberlaini, with no other species recognized. However, recent molecular studies have revealed at least five genetic lineages within this species complex, with one lineage more closely related to the neotenic Eurycea species of central Texas and E. chamberlaini nested within E. quadridigitata sensu lato. We use large-scale geographic sampling in combination with a multilocus species delineation method and morphology to test whether these genetic lineages represent distinct species under the general lineage concept of species. We describe two new species of salamander from this complex, resurrect and elevate a former subspecies to full species status, add to the diagnosis of E. chamberlaini, and redefine E. quadridigitata in the context of this revision. All five species are diagnosable from one another through a number of meristic, morphometric, molecular, and ecological criteria.

Unpacking the species conundrum: philosophy, practice and a way forward.

The history of ecology and evolutionary biology is rife with attempts to define and delimit species. However, there has been confusion between concepts and criteria, which has led to discussion, debate, and conflict, eventually leading to lack of consistency in delimitation. Here, we provide a broad review of species concepts, a clarification of category versus concept, an account of the general lineage concept (GLC), and finally a way forward for species discovery and delimitation. Historically, species were considered as varieties bound together by reproduction. After over 200 years of uncertainty, Mayr attempted to bring coherence to the definition of species through the biological species concept (BSC). This has, however, received much criticism, and the last half century has spawned at least 20 other concepts. A central philosophical problem is that concepts treat species as 'individuals' while the criteria for categorization treats them as 'classes'. While not getting away from this problem entirely, the GLC attempts to provide a framework where lineage divergence is influenced by a number of different factors (and correlated to different traits) which relate to the different species concepts. We also introduce an 'inclusive' probabilistic approach for understanding and delimiting species. Finally, we provide aWallacean (geography related) approach to the Linnaean problem of identifying and delimiting species, particularly for cases of allopatric divergence, and map this to the GLC. Going one step further, we take a morphometric terrain approach to visualizing and understanding differences between lineages. In summary, we argue that while generalized frameworks may work well for concepts of what species are, plurality and 'inclusive' probabilistic approaches may work best for delimitation.

A novel genus and cryptic species harboured within the monotypic freshwater crayfish genus Tenuibranchiurus Riek, 1951 (Decapoda: Parastacidae).

Identifying species groups is an important yet difficult task, with there being no single accepted definition as to what constitutes a species, nor a set of criteria by which they should be delineated. Employing the General Lineage Concept somewhat circumvents these issues, as this concept allows multiple concordant lines of evidence to be used as support for species delimitation, where a species is defined as any independently evolving lineage. Genetically diverse groups have previously been identified within the monotypic parastacid genus Tenuibranchiurus Riek, 1951, but no further investigation of this diversity has previously been undertaken. Analysis of two mitochondrial DNA gene regions has previously identified two highly divergent groups within this taxon, representing populations from Queensland (Qld) and New South Wales (NSW), respectively. Additional testing within this study of both mitochondrial and nuclear DNA through species discovery analyses identified genetically diverse groups within these regions, which were further supported by lineage validation methods. The degree of genetic differentiation between Qld and NSW populations supports the recognition of two genera; with Qld retaining the original genus name Tenuibranchiurus, and NSW designated as Gen. nov. until a formal description is completed. Concordance between the species discovery and lineage validation methods supports the presence of six species within Tenuibranchiurus and two within Gen. nov. The recognition of additional species removes the monotypy of the genus, and the methods used can improve species identification within groups of organisms with taxonomic problems and cryptic diversity.

Variation of plumage patterns, geographic distribution and taxonomy of the Unicolored Blackbird (Aves: Icteridae).

The Unicolored Blackbird Agelasticus cyanopus (Vieillot, 1819) is a marsh bird with four allopatric subspecies restricted to lowlands in South America east of the Andes. I conducted a taxonomic revision of the species based on analysis of external morphological characters of 288 study skins, including all types available. My revision shows that: 1) Leistes unicolor Swainson, 1838, is a senior synonym of A. c. xenicus (Parkes, 1966) and, therefore, the correct name of the taxon should be A. c. unicolor (Swainson, 1838); 2) the range of A. c. unicolor (Swainson, 1838) is much wider than previously thought, extending from the mouth of the Rio Amazonas to the state of São Paulo, in southeastern Brazil, where it intergrades with A. c. atroolivaceus (zu Wied-Neuwied, 1831); 3) A. c. atroolivaceus extends its range well beyond the coast of Rio de Janeiro, reaching the coast of São Paulo, the central part of Minas Gerais, Bahia and Espírito Santo; and 4) specimens attributed to A. c. beniensis are highly variable, so this name must be considered a subjective junior synonym of the nominotypical taxon. Under the Biological Species Concept, two broadly parapatric species should be recognized, A. cyanopus and A. atroolivaceus (including unicolor as a subspecies). Under the Phylogenetic Species Concept or the General Lineage Concept of Species, the best taxonomic treatment is to recognize three species: A. cyanopus, A. atroolivaceus, and A. unicolor.

Phylogenetic Pattern, Evolutionary Processes and Species Delimitation in the Genus Echinococcus.

An accurate and stable alpha taxonomy requires a clear conception of what constitutes a species and agreed criteria for delimiting different species. An evolutionary or general lineage concept defines a species as a single lineage of organisms with a common evolutionary trajectory, distinguishable from other such lineages. Delimiting evolutionary species is a two-step process. In the first step, phylogenetic reconstruction identifies putative species as groups of organisms that are monophyletic (share a common ancestor) and exclusive (more closely related to each other than to organisms outside the group). The second step is to assess whether members of the group possess genetic exchangeability (where cohesion is maintained by gene flow among populations) or ecological exchangeability (where cohesion is maintained because populations occupy the same ecological niche). Recent taxonomic reviews have recognized nine species within the genus Echinococcus. Phylogenetic reconstructions of the relationships between these putative species using mtDNA and nuclear gene sequences show that for the most part these nine species are monophyletic, although there are important incongruences that need to be resolved. Applying the criteria of genetic and ecological exchangeability suggests that seven of the currently recognized species represent evolutionarily distinct lineages. The species status of Echinococcus canadensis and Echinococcus ortleppi could not be confirmed. Coalescent-based analyses represent a promising approach to species delimitation in these closely related taxa. It seems likely, from a comparison of sister species groups, that speciation in the genus has been driven by geographic isolation, but biogeographic scenarios are largely speculative and require further testing.

A cryptic new species of Indigo Snake (genus Drymarchon) from the Florida Platform of the United States.

Indigo Snakes (genus Drymarchon) occur from northern Argentina northward into to the United States, where they inhabit southern Texas and disjunct populations in Mississippi, Florida and Georgia. Based on allopatry and morphological differences Collins (1991) hypothesized that the two United States taxa-the Western Indigo Snake, D. melanurus erebennus (Cope, 1860), and the Eastern Indigo Snake, D. couperi (Holbrook, 1842)-deserved full species recognition. Building upon this hypothesis with molecular and morphological analyses we illustrate that D. couperi is split into two distinct lineages. Based on the General Lineage Concept of Species, we describe the lineage that occurs along the Gulf coast of Florida and Mississippi as a new species, Drymarchon kolpobasileus. The new species is distinguished from D. couperi by a suite of morphological features, including a shorter and shallower head, deeper and shorter 7^th infralabial scales, and shorter temporal scales. Overall, the presence of a deep 7^th infralabial scale provides the best univariate identifier of D. kolpobasileus sp. nov. This study illustrates the usefulness of using both morphological and genetic data in refining accurate descriptions of geographical distributions.

A palaeontological case study for species delimitation in diverging fossil lineages

A general lineage concept is widely adopted in studies of species delimitation for extant taxa with DNA sequence data. In the general lineage concept, species are separately evolving metapopulations that acquire sets of properties during the process of speciation. Murine rodents from the Miocene Siwalik Group of Pakistan have an excellent fossil record showing temporal trends of morphological change, and are therefore ideal to observe the process of lineage divergence as fossil evidence. Here, we review the evolution of Siwalik murine rodents in the sense of the general lineage concept and draw lines of evidence for lineage separation. For currently available datasets, we applied the initial split criterion as well as three operational criteria (phenetic, diagnosable and ecological properties). In early Siwalik murine rodents, the acquisition of distinct phenetic properties emerges subsequent to the event of the evolutionary split. The phenetic evidence occurs closer in time to ‘true’ divergence, while distinct diagnosable and ecological properties fully arise later. Within a sequence of time-controlled samples, these observations explicitly show that phenetic properties (here, shape data and a combination of linear measurements) more precisely delimit the divergence of fossil species than does the fixation of diagnostic properties.

Primate Taxonomy: Inflation or Real?

This article reviews changes in primate taxonomy, especially those pertaining to the meaning of the term species, since its inception two and a half centuries ago. Despite continuing discoveries and the involvement of competent practitioners, the adoption of the polytypic species concept, especially underpinned by the biological species concept, ensured that primate taxonomy was in a sorry state by the middle of the twentieth century. In the latter half of the twentieth century, a gradual rethinking of the nature of species took place, and many different species concepts were proposed. The phylogenetic species concept has been widely adopted over the past ∼20 years, sustained by a gradual realization that species are evolutionary lineages. This review provides examples of how the old way of thinking about species hampered our understanding of primate biodiversity and of how the phylogenetic species concept (or the diagnosability criterion under the general lineage concept) has clarified matters, opening them up for discussion. The adoption of this evolutionary view of species has implications for conservation, particularly because it increases recognition of biodiversity.

Molecular Assessment and Taxonomic Status of the Rapid Racerunner (Eremias velox complex) with Particular Attention to the Populations in Northwestern China

The rapid racerunner, Eremias velox, is a widely distributed lizard from the northern Caucasus across entire Central Asia eastward to China. It is increasingly common to accept E. velox as a species complex in its entire range. To date, published morphological and molecular systematic hypotheses of this complex are only partially congruent, and its taxonomic status and evolutionary history are still far from clear. The mitochondrial cytochrome b gene and 12S rRNA sequences were used to evaluate the taxonomy of this complex, with particular attention to the phylogenetic placement of populations in northwestern China. Examination of the phylogenetic analyses recovers seven distinct, biogeographically discrete, and well-supported clades, revealing genetically identifiable populations corresponding to some previously morphology-defined subspecies. Chinese E. v. roborowskii appears to have split from other Central Asian rapid racerunner lizards well before differentiation occurred among the latter taxa. Specifically, we corroborate that there are two subspecies occurring in China, i.e., E. v. velox and E. v. roborowskii. We recommend a novel subspecific status for the phenotypically and genetically distinct populations in southern Aral Sea region of Uzbekistan previously assigned to E. v. velox. Finally, each of the three independently evolving lineages from Iranian Plateau should be recognized as three species new to science under the general lineage concept.

Genetic variation corroborates subspecific delimitation in the Namib fog-basking beetle, Onymacris unguicularis (Haag) (Tenebrionidae, Coleoptera)

The fog-basking beetle, Onymacris unguicularis (Haag, 1875), is currently listed as a polytypic form comprising two subspecies. A flightless substrate specialist, the beetleis endemic to vegetationless dunes in the Namib, where southern populations constitute the nominate subspecies, O. u. unguicularis, and populations some 300 km to the north compose O. u. schulzeae Penrith, 1984. Their taxonomic descriptions are based on minor differences in pronotal and prosternal shape, and the phylogenetic validity of these subspecies has yet to be ascertained. Here we reassess the polytypic status of O. unguicularis by (1) examining diagnostic phenotypic characters in conjunction with a geometric morphometric analysis, and (2) conducting phylogenetic analysis of mitochondrial DNA sequences. Our results confirm pronotal and prosternal differences, which are complemented by geometric morphometric resolution of the subspecies. Phylogenetic analysis recovered two reciprocally monophyletic lineages that exhibit perfect phylogeographic congruence with phenotypic variation. Our genetic data identify southern and northern populations as distinct lineages, corroborate morphometric data regarding subspecific delimitation, and therefore support the recognition of O. u. unguicularis and O. u. schulzeae as valid taxa under the general lineage concept.