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Species-level taxonomy is traditionally based on herbarium collections that typically include few, or even single, representatives per site. This can lead to underestimation of diversity when there are sympatric populations of superficially similar plants belonging to different lineages. Satyrium longicauda (Orchidaceae) represents a taxonomic challenge for the delimitation of species boundaries due to the high degree of morphological variation detected within and among populations. Currently, just two varieties are accepted based mainly on length differences of the lateral sepal and nectar spur. However, there is extensive morphological variation within South African populations and evidence for several pollination ecotypes, indicating that this taxon represents an actively diverging species complex. Here, we evaluate intraspecific morphological variation through uni- and multivariate morphometrics and analyse internal transcribed spacer sequences for individuals sampled from 36 sites, including 14 sites where divergent morphotypes occur sympatrically. Morphometric analyses of 1802 individuals revealed the presence of eight morphotypes based on vegetative and floral characters. Up to six morphologically and genetically distinct morphotypes can coexist in sympatry. Morphological and genetic distances among populations were significantly correlated. Phylogenetic analyses of 120 accessions indicated that neither of the two varieties nor S. longicauda as a species is monophyletic, and provided evidence for the monophyly of some of the morphotypes including the newly described S. cernuiflorum. The presence of distinct morphological and genetic sympatric variants, which in several cases scale up to distinct evolutionary lineages, is consistent with the existence of different taxa according to morphological and biological species concepts. Our results therefore confirm that taxonomy based mainly on herbarium collections can grossly under-estimate actual diversity of disparate lineages, although further work is required to finalize taxonomic decisions. These findings have implications for efforts to estimate species diversity in groups that are in the process of diversifying and for conservation practice. [ABSTRACT FROM AUTHOR]

Incomplete descriptions of nominal taxa are one of the most significant obstacles in modern taxonomy, including the taxonomy of Tardigrada. Another major problem in tardigrade systematics is the lack of tests for the reliability of genetic markers in species delineation. Here, we employ an integrative taxonomy approach to redescribe the nominal taxon for the P. areolatus complex, Paramacrobiotus areolatus. Moreover, we obtained multilocus DNA sequences for another 16 populations representing 9–12 Paramacrobiotus species collected from Europe, North America, Africa and Australia, enabling us to reconstruct the most extensive phylogeny of the genus to date. The identification of a pair of potentially cryptic dioecious P. areolatus complex species with divergent genetic distances in ITS2 (1.4%) and COI (13.8%) provided an opportunity to test the biological species concept for the first time in the history of tardigrade taxonomy. Intra- and interpopulation crosses did not differ in reproductive success in terms of F1 offspring. However, because of the low F1 family sizes, we were unfortunately unable to test F1 hybrid fertility. Although our results are only partially conclusive, they offer a baseline not only for further taxonomic and phylogenetic research on the areolatus complex, but also for studies on species delineation in tardigrades in general. [ABSTRACT FROM AUTHOR]

Delineation of species is fundamental to organizing and understanding biological diversity. The most widely applied criterion for distinguishing species is the Biological Species Concept (BSC), which defines species as groups of interbreeding individuals that remain reproductively isolated fromother suchgroups. The BSC has broad appeal; however, many organisms, most notably asexual lineages, cannot be classified according to the BSC. Despite their exclusively asexual mode of reproduction, Bacteria and Archaea can transfer and exchange genes though homologous recombination. Here we show that barriers to homologous gene exchange define biological species in prokaryotes with the same efficacy as in sexual eukaryotes. By analyzing the impact of recombination on the polymorphisms in thousands of genome sequences, we find that over half of named bacterial species undergo continuous recombination among sequenced constituents, indicative of true biological species. However, nearly a quarter of named bacterial species show sharp discontinuities and comprise multiple biological species. These interruptions of gene flow are not a simple function of genome identity, indicating that bacterial speciation does not uniformly proceed by the gradual divergence of genome sequences. The same genomic approach based on recombinant polymorphisms retrieves known species boundaries in sexually reproducing eukaryotes. Thus, a single biological species definition based on gene flow, once thought to be limited only to sexually reproducing organisms, is applicable to all cellular lifeforms. [ABSTRACT FROM AUTHOR]

Modern multilocus molecular techniques are a powerful tool in the detection and analysis of cryptic taxa. However, its shortcoming is that with allopatric populations it reveals phylogenetic lineages, not biological species. The increasing power of coalescent multilocus analysis leads to the situation in which nearly every geographically isolated or semi-isolated population can be identified as a lineage and therefore raised to species rank. It leads to artificial taxonomic inflation and as a consequence creates an unnecessary burden on the conservation of biodiversity. To solve this problem, we suggest combining modern lineage delimitation techniques with the biological species concept. We discuss several explicit principles on how genetic markers can be used to detect cryptic entities that have properties of biological species (i.e. of actually or potentially reproductively isolated taxa). Using these principles we rearranged the taxonomy of the butterfly species close to Polyommatus ( Agrodiaetus) ripartii. The subgenus Agrodiaetus is a model system in evolutionary research, but its taxonomy is poorly elaborated because, as a rule, most of its species are morphologically poorly differentiated. The taxon P. ( A.) valiabadi has been supposed to be one of the few exceptions from this rule due to its accurately distinguishable wing pattern. We discovered that in fact traditionally recognized P. valiabadi is a triplet of cryptic species, strongly differentiated by their karyotypes and mitochondrial haplotypes. [ABSTRACT FROM AUTHOR]

Determining which forms of reproductive isolation have the biggest impact on the process of divergence is a major goal of speciation research. These barriers are often divided into those that affect the potential for hybridization (premating isolation), and those that occur after mating (postmating isolation), and much debate has surrounded the relative importance of these categories. Within the species Mimulus aurantiacus, red- and yellow-flowered ecotypes occur in the southwest corner of California, and a hybrid zone occurs where their ranges overlap. We show that premating barriers are exclusively responsible for isolation in this system, with both ecogeographic and pollinator isolation contributing significantly to total isolation. Postmating forms of reproductive isolation have little or no impact on gene flow, indicating that hybrids likely contribute to introgression at neutral loci. Analysis of molecular variation across thousands of restriction-site associated DNA sequencing (RAD-seq) markers reveals that the genomes of these taxa are largely undifferentiated. However, structure analysis shows that these taxa are distinguishable genetically, likely due to the impact of loci underlying differentiated adaptive phenotypes. These data exhibit the power of divergent natural selection to maintain highly differentiated phenotypes in the face of gene flow during the early stages of speciation. [ABSTRACT FROM AUTHOR]

In Australia's arid and semi-arid zone, most aquatic habitats are nonpermanent. Although approximately 70% of its land surface belongs to these zones, very little is known of the iconic 'large branchiopods' that inhabit these important and widespread habitats. In the present study, we investigated 737 Australian specimens of the spinicaudatan taxa Caenestheria and Caenestheriella with a combination of one mitochondrial (cytochrome oxidase subunit I; COI) and three nuclear (elongation factor 1α, internal transcribed spacer 2, and 28S) markers to assess the diversity of species, their phylogenetic relationships, and phylogeographical history. The initial species delimitation was based on COI employing a combination of phylogenetic analyses and two automated approaches to species delimitation (general mixed Yule coalescent model and Automated Barcode Gap Discovery). The outcome was tested by the nuclear markers and considered under differing species concepts. The number of delineated species ranged from 14-27, in no case being in full agreement with any of the two automated approaches. The lower numbers resulted if inferred reproductive isolation, as required for the biological or Hennigian species concept, was employed. Although nuclear markers did not indicate ongoing reproduction, the lack of sympatric co-occurrences inhibited inferences of definitive reproductive isolation in several instances. If monophyly or an 'independent evolutionary fate' was employed, as required for the phylogenetic or evolutionary species concepts, the species' distribution was of no importance and up to 27 species could be delimitated. Because the Australian representatives of both studied genera could not be clearly separated from each other but constitute a single monophyletic clade separated from all available non- Australian representatives of these genera, we describe a new spinicaudatan genus Ozestheria gen. nov. to accommodate these species. Populations revealed relatively small levels of genetic differentiation over large areas of central and eastern Australia. By far the most pronounced levels of genetic differentiation were observed towards the north-eastern regions, a pattern possibly explainable by ecological conditions and the movement of nomadic water birds that disperse resting eggs. © 2015 The Linnean Society of London [ABSTRACT FROM AUTHOR]

Subspecies lie at the interface between systematics and population genetics, and represent a unit of biological organization in zoology that is widely used in the disciplines of taxonomy and conservation biology. In this review, we explore the utility of subspecies in relation to their application in systematics and biodiversity conservation, and briefly summarize species concepts and criteria for their diagnosis, particularly from an invertebrate perspective. The subspecies concept was originally conceived as a formal means of documenting geographical variation within species based on morphological characters; however, the utility of subspecies is hampered by inconsistencies by which they are defined conceptually, a lack of objective criteria or properties that serve to delimit their boundaries, and their frequent failure to reflect distinct evolutionary units according to population genetic structure. Moreover, the concept has been applied to populations largely comprising different components of genetic diversity reflecting contrasting evolutionary processes. We recommend that, under the general lineage (unified) species concept, the definition of subspecies be restricted to extant animal groups that comprise evolving populations representing partially isolated lineages of a species that are allopatric, phenotypically distinct, and have at least one fixed diagnosable character state, and that these character differences are (or are assumed to be) correlated with evolutionary independence according to population genetic structure. Phenotypic character types include colour pattern, morphology, and behaviour or ecology. Under these criteria, allopatric subspecies are a type of evolutionarily significant unit within species in that they show both neutral divergence through the effects of genetic drift and adaptive divergence under natural selection, and provide an historical context for identifying biodiversity units for conservation. Conservation of the adaptedness and adaptability of gene pools, however, may require additional approaches. Recent studies of Australian butterflies exemplify these points. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ••, ••-••. [ABSTRACT FROM AUTHOR]

We reconsider the taxonomy of a group of closely related Ostrinia spp., illustrating how useful Mayr’s biological species concept remains for studying speciation patterns and processes. We review and re-analyse recent data on Ostrinia scapulalis, Ostrinia nubilalis, Ostrinia narynensis and Ostrinia orientalis, along with those obtained over > 45 years in the former Soviet Union. The ten species of the ‘trilobed uncus’ group in the Ostrinia genus are classified into subgroups according to male mid-tibia morphology. However, none of the characters that further discriminate between them (female sex pheromones, male genitalia and calling time) varies together with male mid-tibia morphology, and neither do molecular markers. Moreover, male mid-tibia morphology appears to depend on only two diallelic loci and seems to be unrelated to reproductive isolation between Ostrinia taxa. By contrast, reproductive isolation is strongly related to host-plant type. In accordance with Mayr’s species concept, we thus propose a revision of the trilobed uncus Ostrinia spp. based primarily on host-plant type. We propose that O. narynensis Mutuura & Munroe, 1970 ( syn. nov.) and O. orientalis Mutuura & Munroe, 1970 ( syn. nov.) be synonymized with O. scapulalis (Walker, 1859). We further demonstrate that O. nubilalis auctt. pro parte feeding on mugwort, hop, and several other dicotyledons (previously called the ‘ O. nubilalis mugwort-race’ in France) also belongs to O. scapulalis. Consequently, we propose that only O. nubilalis specimens feeding on maize (the former French ‘ O. nubilalis maize-race’) belong to O. nubilalis (Hübner, 1796). The implications of this revision are discussed. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 49–72. [ABSTRACT FROM AUTHOR]

The original description of the slugs Arion ( Carinarion) fasciatus, Arion ( Carinarion) silvaticus and Arion ( Carinarion) circumscriptus was based on subtle differences in body pigmentation and genital anatomy. However, body pigmentation in these slugs may be influenced by their diet, whereas the genital differences between the species could not be confirmed by subsequent multivariate morphometric analyses. Hence, the status of the three nominal morphospecies remains controversial, with electrophoretic studies based on albumen gland proteins and allozymes also providing conflicting results. These studies suggested that Carinarion species are difficult to reconcile with the biological species concept because there is evidence of interspecific hybridization in places where these predominantly self-fertilizing slugs apparently outcross. Therefore, in the present study, the three Carinarion species are evaluated under a phylogenetic species concept, using nucleotide sequences of the nuclear ribosomal internal transcribed spacer 1 (ITS-1) and the mitochondrial 16S rDNA. ITS-1 showed no species specific variation. However, 16S rDNA yielded five haplotype groups. Three of these grouped haplotypes by species, whereas the two others joined haplotypes of different species and included all haplotypes that were shared by species (22% of all haplotypes). Hence, the three nominal Carinarion species appear to be inconsistent with a phylogenetic species concept. The present data also confirmed that North American Carinarion populations are genetically impoverished and may be not sufficiently representative with respect to the taxonomy of Carinarion. In conclusion, we currently regard Carinarion as a single species-level taxon, whose taxonomically deceiving, correlated phenotypic and genetic intraspecific variation is caused by sustained self-fertilization. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89, 589–604. [ABSTRACT FROM AUTHOR]

In the early 1930s, the synthesis of Darwinian natural selection, mutation, and Mendelian genetics gave rise to the paradigm of ‘modern Darwinism’, also known as ‘neo-Darwinism’. This has contributed greatly to our understanding. But increasing knowledge of other mechanisms, including endosymbiosis, genetic and genomic duplication, polyploidy, hybridization, epigenetics, horizontal gene transfer in prokaryotes, and the modern synthesis of embryonic development and evolution, has widened our horizons to a diversity of possibilities for change. All of these can be gathered under the umbrella concept of ‘genomic creativity’, which, in partnership with natural selection, affords a more comprehensive modern explanation of evolution. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 655–672. [ABSTRACT FROM AUTHOR]