Species and speciation.

This chapter discusses two species models, which are diametrically opposed. The first, often called the 'biological species concept', defines species in terms of 'reproductive isolation', convinced that species arise when subsets of a population are split off and remain geographically isolated over evolutionary time. If and when such new species are reunited with their founder population, interbreeding does not occur, or if it does, infertile progeny result. Hence, from the biological species concept, natural selection is a primary agent of change and directly selects for new species. In this sense, species are the direct products of natural selection and they are therefore 'adaptive devices'. When applying this species concept, it has been impossible to separate some sibling species of fruit flies in the genus Bactrocera where distinct morphological species can be similar in molecular analyses of certain DNA sequences, while similar species morphologically are distinct in the same molecular characters. A radically different model, the 'recognition concept of species', relies heavily on a knowledge of species ecology and behaviour, particularly in their natural habitat. The principal points in this concept are given. In contrast to the now-outdated biological species concept that leads one to depend on laboratory-based research to define species, the recognition concept requires workers to undertake extensive field research in the habitat of the taxon under investigation. In translating this approach to research in the insect family Tephritidae, particularly the Dacinae, some 35 years of field surveys have been undertaken throughout the Indian subcontinent, South-east Asia and the South Pacific region. These surveys included trapping using male lure traps and host fruit collections of commercial/edible fruits. The results of this work have included the provision of specimens of almost all known species for morphological descriptions (c.800 species), material for male pheromone chemistry, and data on host fruit relationships and biogeographical studies.

Predictors of Taxonomic Inflation and its Role in Primate Conservation

ABSTRACTSpecies are the main unit used to measure biodiversity, but different preferred diagnostic criteria can lead to very different delineations. For instance, named primate species have more than doubled since 1982. Such increases have been termed “taxonomic inflation” and have been attributed to the widespread adoption of the ‘phylogenetic species concept’ (PSC) in preference to the previously popular ‘biological species concept’ (BSC). Criticisms of the PSC have suggested taxonomic inflation may be biased toward particular taxa and have unfavourable consequences for conservation. Here, we explore predictors of taxonomic inflation across primate taxa since the initial application of the PSC nearly 40 years ago. We do not find evidence that diversification rate, the rate of lineage formation over evolutionary time, is linked to inflation, contrary to expectations if the PSC identifies incipient species. We also do not find evidence of research effort in fields where work has been suggested to motivate splitting being associated with increases in species numbers among genera. To test the suggestion that splitting groups is likely to increase their perceived risk of extinction, we test whether genera that have undergone more splitting have also observed a greater increase in their proportion of threatened species since the introduction of the PSC. We find no cohesive signal of inflation leading to higher threat probabilities across primate genera. Overall, this analysis sends a positive message that threat statuses of primate species are not being overwhelmingly affected by splitting in line with what has recently been reported for birds. Regardless, we echo warnings that it is unwise for conservation to be reliant on taxonomic stability. Species (however defined) are not independent from one another, thus, monitoring and managing them as such may not meet the overarching goal of conserving biodiversity.

The evolving species concepts used for yeasts: from phenotypes and genomes to speciation networks

Here we review how evolving species concepts have been applied to understand yeast diversity. Initially, a phenotypic species concept was utilized taking into consideration morphological aspects of colonies and cells, and growth profiles. Later the biological species concept was added, which applied data from mating experiments. Biophysical measurements of DNA similarity between isolates were an early measure that became more broadly applied with the advent of sequencing technology, leading to a sequence-based species concept using comparisons of parts of the ribosomal DNA. At present phylogenetic species concepts that employ sequence data of rDNA and other genes are universally applied in fungal taxonomy, including yeasts, because various studies revealed a relatively good correlation between the biological species concept and sequence divergence. The application of genome information is becoming increasingly common, and we strongly recommend the use of complete, rather than draft genomes to improve our understanding of species and their genome and genetic dynamics. Complete genomes allow in-depth comparisons on the evolvability of genomes and, consequently, of the species to which they belong. Hybridization seems a relatively common phenomenon and has been observed in all major fungal lineages that contain yeasts. Note that hybrids may greatly differ in their post-hybridization development. Future in-depth studies, initially using some model species or complexes may shift the traditional species concept as isolated clusters of genetically compatible isolates to a cohesive speciation network in which such clusters are interconnected by genetic processes, such as hybridization.

Some Examples of the Use of Molecular Markers for Needs of Basic Biology and Modern Society

Application of molecular genetic markers appeared to be very fruitful in achieving many goals, including (i) proving the theoretic basements of general biology and (ii) assessment of worldwide biodiversity. Both are provided in the present meta-analysis and a review as the main signal. One of the basic current challenges in modern biology in the face of new demands in the 21st century is the validation of its paradigms such as the synthetic theory of evolution (STE) and biological species concept (BSC). Another of most valuable goals is the biodiversity assessment for a variety of social needs including free web-based information resources about any living being, renovation of museum collections, nature conservation that recognized as a global project, iBOL, as well as resolving global trading problems such as false labeling of species specimens used as food, drug components, entertainment, etc. The main issues of the review are focused on animals and combine four items. (1) A combination of nDNA and mtDNA markers best suits the identification of hybrids and estimation of genetic introgression. (2) The available facts on nDNA and mtDNA diversity seemingly make introgression among many taxa obvious, although it is evident, that introgression may be quite restricted or asymmetric, thus, leaving at least the “source” taxon (taxa) intact. (3) If we consider sexually reproducing species in marine and terrestrial realms introgressed, as it is still evident in many cases, then we should recognize that the BSC, in view of the complete lack of gene flow among species, is inadequate because many zoological species are not biological ones yet. However, vast modern molecular data have proven that sooner or later they definitely become biological species. (4) An investigation into the fish taxa divergence using the BOLD database shows that most gene trees are basically monophyletic and interspecies reticulations are quite rare.

An overview of speciation and species limits in birds

Accurately determining avian species limits has been a challenge and a work in progress for most of a century. It is a fascinating but difficult problem. Under the biological species concept, only lineages that remain essentially independent when they are in sympatry are clearly species. Otherwise, there is no clear line yet found that marks when a pair of diverging lineages (e.g., in allopatry) become different enough to warrant full biological species status. Also, with more data, species limits often require reevaluation. The process of divergence and speciation is itself very complex and is the focus of intense research. Translating what we understand of that process into taxonomic names can be challenging. A series of issues are important. Single-locus criteria are unlikely to be convincing. Genetic independence is not a species limits requirement, but the degree of independence (gene flow) needs to be considered when there is opportunity for gene flow and independence is not complete. Time-based species (limits determined by time of separation) are unsatisfactory, though integrating time more effectively into our datasets is warranted. We need to disentangle data signal due to neutral processes vs. selection and prioritize the latter as the main driver of speciation. Assortative mating is also not likely to be an adequate criterion for determining species limits. Hybridization and gene flow are more important than ever, and there is a condition not being treated evenly in taxonomy: evolutionary trysts of 2 or more lineages stuck together through gene flow just short of speciation over long periods. Comparative methods that use what occurs between good species in contact to infer species limits among allopatric forms remain the gold standard, but they can be inaccurate and controversial. Species-level taxonomy in birds is likely to remain unsettled for some time. While the study of avian speciation has never been more exciting and dynamic, there is no silver bullet for species delimitation, nor is it likely that there will ever be one. Careful work using integrative taxonomy in a comparative framework is the most promising way forward.

Coiling directions in the planktonic foraminifer Pulleniatina: A complex eco-evolutionary dynamic spanning millions of years

Planktonic foraminifera are heterotrophic sexually reproducing marine protists with an exceptionally complete fossil record that provides unique insights into long-term patterns and processes of evolution. Populations often exhibit strong biases towards either right (dextral) or left (sinistral) shells. Deep-sea sediment cores spanning millions of years reveal that some species show large and often rapid fluctuations in their dominant coiling direction through time. This is useful for biostratigraphic correlation but further work is required to understand the population dynamical processes that drive these fluctuations. Here we address the case of coiling fluctuations in the planktonic foraminifer genus Pulleniatina based on new high-resolution counts from two recently recovered sediment cores from either side of the Indonesian through-flow in the tropical west Pacific and Indian Oceans (International Ocean Discovery Program Sites U1486 and U1483). We use single-specimen stable isotope analyses to show that dextral and sinistral shells from the same sediment samples can show significant differences in both carbon and oxygen isotopes, implying a degree of ecological separation between populations. In one case we detect a significant difference in size between dextral and sinistral specimens. We suggest that major fluctuations in coiling ratio are caused by cryptic populations replacing one another in competitive sweeps, a mode of evolution that is more often associated with asexual organisms than with the classical ‘biological species concept’.

Discriminating arboviral species

Mosquito-borne arboviruses, including a diverse array of alphaviruses and flaviviruses, lead to hundreds of millions of human infections each year. Current methods for species-level classification of arboviruses adhere to guidelines prescribed by the International Committee on Taxonomy of Viruses (ICTV), and generally apply a polyphasic approach that might include information about viral vectors, hosts, geographical distribution, antigenicity, levels of DNA similarity, disease association and/or ecological characteristics. However, there is substantial variation in the criteria used to define viral species, which can lead to the establishment of artificial boundaries between species and inconsistencies when inferring their relatedness, variation and evolutionary history. In this study, we apply a single, uniform principle – that underlying the Biological Species Concept (BSC) – to define biological species of arboviruses based on recombination between genomes. Given that few recombination events have been documented in arboviruses, we investigate the incidence of recombination within and among major arboviral groups using an approach based on the ratio of homoplastic sites (recombinant alleles) to non-homoplastic sites (vertically transmitted alleles). This approach supports many ICTV-designations but also recognizes several cases in which a named species comprises multiple biological species. These findings demonstrate that this metric may be applied to all lifeforms, including viruses, and lead to more consistent and accurate delineation of viral species.

Species limits and taxonomy in birds

Despite the acknowledged importance of defining avian species limits to scientific research, conservation, and management, in practice, they often remain contentious. This is true even among practitioners of a single species concept and is inevitable owing to the continuous nature of the speciation process, our incomplete and changing understanding of individual cases, and differing interpretations of available data. This issue of Ornithology brings together several papers on species limits, some more theoretical and general, and others case studies of specific taxa. These are viewed primarily through the lens of the biological species concept (BSC), by far the most widely adopted species concept in influential ornithological works. The more conceptual contributions focus on the importance of the integrative approach in species delimitation; the importance of considering selection with the increasing use of genomic data; examinations of the effectiveness of the Tobias et al. character-scoring species limits criteria; a review of thorny issues in species delimitation using examples from Australo-Papuan birds; and a review of the process of speciation that addresses how population divergence poses challenges. Case studies include population genomics of the American Kestrel (Falco sparverius); an integrative taxonomic analysis of Graceful Prinia (Prinia gracilis) that suggests two species are involved; and a reevaluation of species limits in Caribbean Sharp-shinned Hawk (Accipiter striatus) taxa.

Species

This chapter studies the systematists' perspective on species concepts and the role of species in systematics. No matter how sophisticated the tools and methods enhancing the conceptualization of reality may become in the future, systematists will still be constrained by their perceptions. In their more modest, empirical view, systematists embrace their perceived reality and prefer species concepts that incorporate tools for identifying and delimiting species as empirical hypotheses, thereby providing them with efficacious working terminal elements for phylogenetic analysis and classification of more inclusive taxa. It is fortunate that cladists employed the notion of a “phylogenetic” species concept based on diagnosability before more metaphysically inclined authors appropriated the term for concepts founded on monophyly or common ancestors. As noted, Willi Hennig's species concept was a version of the “biological” species concept, and it fell to his followers to develop a species concept that is well suited to cladistic principles. Among the earliest of the post-Hennigian empiricists was American Museum ichthyologist Donn Rosen. Rosen's concept, sometimes called the apomorphic concept because of its requirement that every recognized species must have its own derived character state, accomplished two key advances for systematics: it proposed a cladistic criterion for recognizing species, and it defined species as the minimal units of analysis, as far as taxonomy is concerned, thus setting a lower bound for systematic inquiry.