Subspecies Taxonomy Research Articles

Registros más occidentales de Gonatodes albogularis fuscus (Squamata: Sphaerodactylidae) en Honduras y comentarios sobre su taxonomía y conservación

Gonatodes albogularis fuscus (Hallowell, 1855) is known mainly from the lowlands from Chiapas in Mexico to western Venezuela and Colombia and in several Caribbean islands. In Honduras it has been reported from the center, south, east and north, however there are no reports from the west of the country. We document this species for the first time in the departments of Comayagua and Intibucá in the center-west and comment on its distribution in Honduras. In addition, we suggest the need to reassess the taxonomy of subspecies of G. albogularis and comment about conservation problems derived from competition and predation by feral and invasive species and habitat modification.

Global conservation genomics of blue whales calls into question subspecies taxonomy and refines knowledge of population structure

AbstractBlue whales (Balaenoptera musculus) are the largest living animal and, like other baleen whales, became endangered due to whaling. Here, we used population genomics to infer the number, distribution and other characteristics of subspecies and populations. We used the largest DNA dataset in blue whales, both in terms of genomic markers (16,661 SNPs and mtDNA) and geographic coverage (n = 276 for SNPs; n = 531 for mtDNA). We found greatest divergence among the eastern Pacific, Indo‐western Pacific and Antarctic blue whales. There were indications that natural selection in different environments promoted divergence among these groupings. Within these regions, there was divergence between the eastern North and eastern South Pacific, and among the eastern Indian Ocean, the western South Pacific and the northern Indian Ocean. There was no divergence within the Antarctic. These findings are consistent with the current classification of Antarctic and Indo‐western Pacific blue whales in the Southern Hemisphere as different subspecies but call into question the subspecies taxonomy of eastern Pacific blue whales. The study shows that opposite breeding seasons on either side of the equator do not necessarily inhibit connectivity across the equator, and reinforces that population structure needs to be well understood to conserve the diversity within species.

Subspecific Synonym of Monochamus alternatus (Coleoptera: Cerambycidae): Population Genetics and Morphological Reassessment.

Monochamus alternatus Hope, 1842, is a major forest pest that hosts the pathogenic pinewood nematode (PWN), Bursaphelenchus xylophilus (Steiner and Buhrer, 1934) Nickle 1970. Taxonomically, M. alternatus is currently divided into two subspecies, based on morphology and geography: Monochamus alternatus alternatus Hope, 1842 in China, Taiwan, Tibet, Vietnam, and Laos and Monochamus alternatus endai Makihara, 2004 in South Korea and Japan. Despite their economic importance, the subspecies taxonomy of M. alternatus has never been tested after the first description. In this study, we aimed to reassess the subspecies taxonomy of M. alternatus using molecular and morphological data. For morphological analysis, we examined three major morphological characters (pronotal longitudinal band, granulation on humeri, and elytral proximomedial spine) from 191 individuals from China, Korea, and Taiwan. Population genetic structures were examined using 85 de novo sequences and 82 public COI sequences from China, Korea, Japan, Malaysia, Taiwan, and a few intercepted specimens from the United States. All the genetic data were aligned as three different multiple sequence alignments. Individuals from each subspecies were morphologically and genetically scattered, not clustered according to subspecies in any of the analyses. Therefore, a new synonymy is proposed: Monochamus alternatus Hope, 1842 = Monochamus alternatus endai, syn. n. This study suggests a more robust classification of M. alternatus for the first time and ultimately will pose a substantial impact on implementing quarantine or forestry policies.

Subspecies Taxonomy and Inter-Population Divergences of the Critically Endangered Yellow-Breasted Bunting: Evidence from Song Variations.

Simple SummaryIdentifying the taxonomic status of subspecies or population with independent evolutionary tendencies is important for the targeted conservation of endangered species. Two subspecies of the critically endangered, Yellow-breasted Bunting Emberiza aureola have long been classified: E. a. aureola and E. a. ornata. However, populations distributed in Hokkaido, Japan, are sometimes considered another subspecies of E. a. insulana. By comparing vocal divergences, we found that the song of the Yellow-breasted Bunting Emberiza aureola has subspecies-specific properties and that insulana can be classified acoustically as a subspecies. Morphological and genetic differences should be tested further to confirm its subspecies status. Our results not only confirm the subspecies but also provide key evidence for targeted taxon conservation efforts for this critically endangered bird species, given that several Japanese populations have disappeared.The critically endangered Yellow-breasted Bunting has undergone population collapse globally because of illegal hunting and habitat deterioration. It was listed as critically endangered (CR) by the International Union for Conservation of Nature (IUCN) in 2017 and designated a Class I (highest level) national conservation bird species in China in 2021. Birdsong in the breeding season is the main communicative signal under sexual selection, and song variations have long been considered critical evidence of divergence among subspecies or populations. We compared the songs of 89 males from 18 populations to test subspecies taxonomy. We found that songs of the Yellow-breasted Bunting Emberiza aureola are subspecies specific and that three subspecies can be clearly discriminated by song divergences. Moreover, an analysis of multiple vocal traits supports the claim that insulana is distinct from aureola and ornata. Finally, at the geographic population level, populations can be clearly classified in accordance with the three subspecies, although the aureola population in Xinjiang, China is differentiated from other populations of the same subspecies. The results of this study demonstrate that all populations and subspecies are unique and should be protected to maintain intraspecies song diversity. In addition, several specific populations, such as insulana populations in Japan and the Xinjiang, China population of aureola, need to be paid special attention to prevent the extinction of unique or local taxa.

Geographical and population diversity of African honeybees, Apismellifera L. (Hymenoptera: Apidae): A review

This review was prepared to summarize the information on geographic and genetic diversity of Apismellifera L. of African honeybees and the progresses made through time and applications of various advanced tools in their taxonomy at subspecies level. African honeybees have developed distinct morphology, behavior and physiology as they evolved in tropical favorable climate. The variations among the African honeybee populations have helped to produce geographic subspecies based on the morphological characteristics. However, the introduction of molecular taxonomic tools has come up with discrepancies in the number of subspecies and lineages, and subspecies composition of evolutionary lineages in Africa and other parts of the world. It has also brought up controversies in the evolutionary history of A. mellifera reporting different location as the center of origin of the species at different times: Asia, Africa and Europe. It is not only the tools that are used, but also the genetic variations within subspecies, within colonies in an apiary site and within individuals of the same colony temporally are the factors for continually appearing discrepancies. Natural hybridization between populations of adjacent subspecies along the geographic transition zones and introgression of genes through introduction of colonies that is mainly driven by human demand for desirable traits of honeybees have been among the challenges for discrimination of honeybee subspecies. Therefore, in future genetic diversity and taxonomic investigations, considering these natural variations at subspecies level and developing standardized and integrated procedures from sampling to analysis methods would help to produce generally recognized data base that is critically important to resolve the problem of inconsistencies in taxonomy and nomenclature of honeybee subspecies of Arica and elsewhere.

Integrating voice and phenotype in a revision of the brush cuckoo Cacomantis variolosus (Aves: Cuculidae) complex

Brush cuckoos of the Cacomantis variolosus complex, which range from Southeast Asia to Australia and the Solomon Islands, have undergone much taxonomic upheaval. Here we examine 389 vocal recordings, 832 skins, and records of brood parasitism and habitat partitioning to shed light on their species and subspecies taxonomy. Bioacoustic analysis revealed seven distinct vocal groups. Among morphological markers, shape and proportions of the tail were found to be supporting indicators, in addition to plumage tone and pattern. Integration of the resulting data set distinguished six species-level taxa within the complex: sepulcralis in the Philippines and Sundaland east to central Nusa Tenggara, virescens in Sulawesi and the Sula Archipelago, aeruginosus in the Moluccas, variolosus in east Nusa Tenggara, the Moluccas, north and east Australia, New Guinea and Bismarck Archipelago, blandus in the Admiralty Islands, and addendus in the Solomon Islands. Our review of infra-specific differentiation among species leads us to distinguish 13 subspecies. All taxa identified are listed in a summary classification of the complex. Taxon diversity is greatest in the Moluccas, where two habitat-partitioned species and five allopatric subspecies occur, of which one species and three subspecies are endemic to this region.

Genetic differentiation of autochthonous sable populations in Western and Eastern Siberia

Morphological variation and active migration of sables (Martes zibellina Linnaeus, 1758) cause confusion in this species’ intraspecific taxonomy. Four to 17 subspecies have been described thus far. In this study, we clarified sable population structure using 11 microsatellite loci in 665 specimens from 33 sable populations. According to subspecies taxonomy, we expect to find four genetic groups. Our results confirmed the presence of two genetic groups in the territories of Western Siberia (subspecies Martes zibellina zibellina Linnaeus, 1758) and the Western Altai Mountains (subspecies Martes zibellina averini Bashanov, 1943). Another genetic group is formed by the populations of the Central Siberian Plateau and presumably represents the subspecies Martes zibellina yeniseensis Ognev, 1925. Previous descriptions of the area occupied by this subspecies include the mountain regions of Southern Siberia. We found a few genetic groups in the Baikal region, whereas only one subspecies Martes zibellina princeps Birula, 1918 had been described previously.

Range-wide population genetic analysis of Seaside Sparrows (Ammospiza maritima) supports at least five distinct population segments that do not align with current subspecies descriptions

AbstractAs an obligate salt marsh species, Seaside Sparrows (Ammospiza maritima) are vulnerable to numerous threats including climate change, coastal erosion, sea-level rise, and both natural and anthropogenic disasters. Of the 9 recognized subspecies, 2 are extinct and 1 is endangered. Previous genetic analyses of mitochondrial DNA (mtDNA) and microsatellite loci showed that current taxonomy does not accurately reflect underlying genetic diversity, with possible consequences for the distribution of conservation resources. To further inform Seaside Sparrow management, we comprehensively describe genetic structure among 24 range-wide sampling locations that include all extant subspecies. We inferred population structure from several thousand single-nucleotide polymorphisms collected from 272 individuals via restriction-site-associated DNA sequencing. Principal components, pairwise FST values, and clustering approaches suggest that Seaside Sparrows on the Atlantic and Gulf Coasts are distinct and consist of at least 5 genetic clusters: 1 in southern Texas, 1 ranging from Aransas County, Texas, to Mississippi; 1 in western Florida; and 2 or 3 genetic groups intermixed along a gradient on the Atlantic Coast. These genetic clusters are not consistent with current subspecies taxonomy and could be used as distinct population segments (DPSs) to inform the most efficient allocation of resources to Seaside Sparrow conservation. Our results regarding the endangered subspecies, A. m. mirabilis, from southern Florida are inconclusive due to low sample size, but indicate that it is distinct and may represent a sixth DPS. Based on our genetic results, we recommend additional song and morphometric analyses in western Florida and a closer study of the boundary between the breeding distributions of A. m. maritima and A. m. macgillivraii to ensure the proper identification of DPSs.

Phylogeography of the iconic Australian red-tailed black-cockatoo (Calyptorhynchus banksii) and implications for its conservation.

Advances in sequencing technologies have revolutionized wildlife conservation genetics. Analysis of genomic data sets can provide high-resolution estimates of genetic structure, genetic diversity, gene flow, and evolutionary history. These data can be used to characterize conservation units and to effectively manage the genetic health of species in a broad evolutionary context. Here we utilize thousands of genome-wide single-nucleotide polymorphisms (SNPs) and mitochondrial DNA to provide the first genetic assessment of the Australian red-tailed black-cockatoo (Calyptorhynchus banksii), a widespread bird species comprising populations of varying conservation concern. We identified five evolutionarily significant units, which are estimated to have diverged during the Pleistocene. These units are only partially congruent with the existing morphology-based subspecies taxonomy. Genetic clusters inferred from mitochondrial DNA differed from those based on SNPs and were less resolved. Our study has a range of conservation and taxonomic implications for this species. In particular, we provide advice on the potential genetic rescue of the Endangered and restricted-range subspecies C. b. graptogyne, and propose that the western C. b. samueli population is diagnosable as a separate subspecies. The results of our study highlight the utility of considering the phylogeographic relationships inferred from genome-wide SNPs when characterizing conservation units and management priorities, which is particularly relevant as genomic data sets become increasingly accessible.

Superspecific Taxonomy of the Far Eastern Unionids (Bivalvia, Unionidae): Review and Analysis

On the basis of previously published materials, which include data on the morphology of the shells of adult mollusks and larvae (glochidia), as well as the results of molecular phylogenetic studies, a critical analysis of the subspecies taxonomy of the Far Eastern Unionidae Rafinesque, 1820, namely, the subfamilies Unioninae Rafinesque, 1820 and Anodontinae Rafinesque, 1820, has been performed. It is noted that the previously described subfamily Nodulariinae Starobogatov et Zatrawkin, 1987 can be regarded as the tribe Nodulariini. The systematic position of all genera of Unionidae inhabiting the Russian Far East is discussed. An updated list of the genera, tribes, and subfamilies of Unionidae is given for this region.

Phylogeography of silver‐studded blue, Plebejus subsolanus (Lepidoptera: Lycaenidae), in the central mountainous regions of Japan

AbstractThe silver‐studded blue, Plebejus subsolanus, is widely distributed in the Russian Altai mountains, northeastern China, the Korean Peninsula, and the Japanese archipelago. In Japan, the species is distributed across wide elevation ranges from the lowlands of Hokkaido to the subalpine zone of Honshu. Current subspecies classification in Japan is as follows: ssp. iburiensis, occurring in lowland grasslands in Hokkaido; ssp. yaginus in lower mountain grasslands in Honshu; and ssp. yarigadakeanus in higher mountain grasslands in Honshu. The habitat of this species has been markedly reduced due to recent habitat destruction and land‐use changes. Here, we undertook phylogeographic analyses of two subspecies, ssp. yaginus and yarigadakeanus in the central mountainous regions of Japan, based on two mitochondrial gene sequences, in order to collect information for establishing effective conservation strategies. From 57 samples from the four mountain ranges, we obtained a haplotype network comprised of 12 haplotypes. Because of the haplotype network topology, the geographic distribution of haplotypes and the correspondence of haplotype divergence to subspecies taxonomy, we provisionally divided the haplotypes into three haplogroups: YR1 and YR2, which comprised ssp. yarigadakeanus, and YG, which comprised ssp. yaginus. Mitochondrial DNA genetic differentiation generally agreed with morphological subspecies classification. The haplotype network suggested that ssp. yarigadakeanus populations had multiple origins, and the subspecies character of “bright blue of the male's wings” was assumed to have evolved independently in each subalpine meadow. We found that P. subsolanus was genetically differentiated depending upon the elevation at each mountain region, suggesting that each haplogroup should be a conservation unit.

The Eriochilus dilatatus (Orchidaceae) complex in Western Australia: subspecies taxonomy is not supported by consistent differences in morphology or distribution

A detailed study of taxonomic features of the Eriochilus dilatatus (Orchidaceae) complex (white bunny orchids) in Western Australia found that there were no consistent differences among most subspecies when herbarium specimens or living plants were compared. These subspecies were originally segregated primarily by differences in leaf size and shape and the number of flowers produced, but a critical examination of herbarium specimens found that these features were highly inconsistent within taxa. These features were also found to be highly variable over time and space within populations of living plants. The distribution patterns, habitat preferences and flowering times of these taxa were found to overlap, even for subspecies brevifolius and orientalis, which occupy the northern and eastern limits of the distribution of this species. Eriochilus dilatatus subsp. magnus and subsp. multiflorus were shown to be synonyms of subsp. dilatatus, whereas subsp. undulatus and subsp. orientalis are synonymised under subsp. brevifolius. As a result of this study, the two recognised subspecies are subsp. dilatatus and subsp. brevifolius, which can be readily separated by plant height, flower numbers and leaf morphology, except for a few intermediate plants where ranges overlap. New keys and descriptions to these taxa are provided. The reasons for previous taxonomic confusion in this group and in many other Western Australian orchids are discussed and research approaches to resolve these issues are suggested.

Allometry and morphometry of blue whales photographed in the Gulf of California: insights into subspecies taxonomy in the Eastern North Pacific

Allometry and morphometry of blue whales photographed in the Gulf of California: insights into subspecies taxonomy in the Eastern North Pacific

Phylogeography of southern brown and golden bandicoots: implications for the taxonomy and distribution of endangered subspecies and species

Southern brown (Isoodon obesulus) and golden (Isoodon auratus) bandicoots are iconic Australian marsupials that have experienced dramatic declines since European settlement. Conservation management programs seek to protect the remaining populations; however, these programs are impeded by major taxonomic uncertainties. We investigated the history of population connectivity to inform subspecies and species boundaries through a broad-scale phylogeographic and population genetic analysis of Isoodon taxa. Our analyses reveal a major east–west phylogeographic split within I. obesulus/I. auratus, supported by both mtDNA and nuclear gene analyses, which is not coincident with the current species or subspecies taxonomy. In the eastern lineage, all Tasmanian samples formed a distinct monophyletic haplotype group to the exclusion of all mainland samples, indicative of long-term isolation of this population from mainland Australia and providing support for retention of the subspecific status of the Tasmanian population (I. o. affinis). Analyses further suggest that I. o. obesulus is limited to south-eastern mainland Australia, representing a significant reduction in known range. However, the analyses provide no clear consensus on the taxonomic status of bandicoot populations within the western lineage, with further analyses required, ideally incorporating data from historical museum specimens to fill distributional gaps.

Discordance between genomic divergence and phenotypic variation in a rapidly evolving avian genus (Motacilla)

Generally, genotypes and phenotypes are expected to be spatially congruent; however, in widespread species complexes with few barriers to dispersal, multiple contact zones, and limited reproductive isolation, discordance between phenotypes and phylogeographic groups is more probable. Wagtails (Motacilla) are a genus of birds with striking plumage pattern variation across the Old World. Up to 13 subspecies are recognized within a single species, yet previous studies using mitochondrial DNA have supported polyphyletic phylogeographic groups that are inconsistent with subspecies plumage characteristics. In this study, we investigate the link between phenotypes and genotype by taking a phylogenetic approach. We use genome-wide SNPs, nuclear introns, and mitochondrial DNA to estimate population structure, isolation by distance, and species relationships. Together, our genetic sampling includes complete species-level sampling and comprehensive coverage of the three most phenotypically diverse Palearctic species. Our study provides strong evidence for species-level patterns of differentiation, however population-level differentiation is less pronounced. SNPs provide a robust estimate of species-level relationships, which are mostly corroborated by a combined analysis of mtDNA and nuclear introns (the first time-calibrated species tree for the genus). However, the mtDNA tree is strongly incongruent and is considered to misrepresent the species phylogeny. The extant wagtail lineages originated during the Pliocene and the Eurasian lineage underwent rapid diversification during the Pleistocene. Three of four widespread Eurasian species exhibit an east-west divide that contradicts both subspecies taxonomy and phenotypic variation. Indeed, SNPs fail to distinguish between phenotypically distinct subspecies within the M. alba and M. flava complexes, and instead support geographical regions, each of which is home to two or more different looking subspecies. This is a major step towards our understanding of wagtail phylogeny compared to previous analyses of fewer species and considerably less sequence data.

Phylogeography of the bobwhite (Colinus) quails

ABSTRACTThe bobwhites (Colinus) consist of 3 grassland‐associated, allopatric species of New World quails (family Odontophoridae): the northern bobwhite (C. virginianus), distributed from the eastern United States to Guatemala; the black‐throated bobwhite (C. nigrogularis), which occurs in scattered localities in the Yucatán Peninsula, Nicaragua, and Honduras; and the crested bobwhite (C. cristatus), whose range stretches from Guatemala to northern Brazil. We used mitochondrial DNA (mtDNA) sequences from both the control region and the ND2 gene to study the phylogenetic relationships and phylogeography of the 3 bobwhite species. We developed ecological niche models to evaluate conclusions inferred from genetic data. Colinus was composed of 2 deeply divergent lineages, 1 represented by the crested bobwhite and the other by northern and black‐throated bobwhites, both of which were genetically distinct from each other. Although the northern bobwhite had high haplotype diversity, this species exhibited evidence of past demographic and geographic expansion, no phylogeographic structure, and no congruence between genetic variation and subspecies taxonomy. Ecological niche modeling was congruent with a recent range expansion for the northern bobwhite from Late Pleistocene refugia in México and the southern United States. The black‐throated bobwhites from the Yucatán Peninsula were distinct from those in Nicaragua, and as a whole this species displayed little evidence of recent expansion. Ecological niche modeling suggested that the current, fragmented distribution of the black‐throated bobwhite has existed for the past 130,000 years. Although the crested bobwhite displayed little evidence of population expansion, the mtDNA data revealed 3–4 geographically and genetically distinct lineages. Results of niche modeling suggest that the crested bobwhite had a much wider distribution in Central and South America during the Last Glacial Maximum. Given the sensitivity of all 3 bobwhite species to climatic cycles, managers should consider impacts of climate change in addition to the loss of habitat when crafting conservation plans. © 2015 The Wildlife Society.

Variability of cytochrome b gene and adjacent part of tRNA-Thr gene of mitochondrial DNA in the northern mole vole Ellobius talpinus (Mammalia, Rodentia)

The Northern mole vole E. talpinus, despite its wide distribution, is characterized by a stable karyotype (2n = NF = 54) and slight morphological polymorphism. We made a preliminary analysis of a mitochondrial DNA fragment to clarify the level of genetic variation and differentiation of E. talpinus The complete cytochrome b gene (cyt b, 1143 bp) and a short part of its flanking gene tRNA-Thr (27 bp) were sequenced. We studied 16 specimens of E. talpinus from eight localities, including Crimea, the Volga region, the Trans-Volga region, the Southern Urals, Western Siberia, and Eastern Turkmenistan. Mitotypes of E. talpinus were distributed on a ML dendrogram as four distinct clusters: the first (I) contains specimens from the Crimea, the second (II) combines individuals from the Volgograd region, the left bank of the Don River; the third (III) includes those from the Trans-Volga region, Southern Urals, the left bank of the Irtysh River, and Eastern Turkmenistan; the fourth (IV) are those from the right bank of the Irtysh River. These clusters were relatively distant from each other: the mean genetic distances (D) between them are 0.021–0.051. The Eastern mole vole E. tancrei differed from E. talpinus population groups 1.5–2 times more (D = 0.077–0.084) than the latters did among themselves. Such variations indirectly proved the unity of E. talpinus, despite its high intraspecific differentiation for the studied fragment of mitochondrial DNA. This differentiation apparently occurred because of the long isolation of E. talpinus population groups by geographic barriers, in particular, the large rivers that completely separate the species range meridionally (the Volga River, the Irtysh River). Sociality and underground lifestyle could accelerate the fixation of mutations in disjunct populations. The composition and distribution of intraspecific groups of E. talpinus, which were identified in analysis of the mitochondrial DNA fragment, do not coincide with the subspecies taxonomy. The subspecies E. t. talpinus is actually a complex taxon, including two or three genetically discrete forms (III, IV, and probably II). Moreover, one of the forms (III) occupies the territory where three subspecies, E. t. talpinus, E. t. rufescens, and E. t. transcaspiae, were described.

Taxonomic checklist of the day geckos of the genera Phelsuma GRAY, 1825 and Rhoptropella HEWITT, 1937 (Squamata: Gekkonidae)

The day geckos of the genus Phelsuma have their center of diversity in Madagascar and the adjacent archipelagos in the western Indian Ocean, especially the Mascarenes, Comoros and Seychelles. Their brilliant colours, diurnal activity, and interesting behaviours have resulted in a long lasting high demand for the pet trade. Accordingly, the international trade of all Phelsuma species is regulated by the Convention on International Trade in Endangered Species of Wild Fauna and Floras (CITES) where all species are included in CITES appendix II. In order to facilitate operating and control of these regulations by national and international nature conservation authorities we provide an updated taxonomic checklist for all species and subspecies of the genus Phelsuma and for Rhoptropella ocellata (a taxon resurrected from the synonymy of Phelsuma). Since a few species (especially P. dubia, P. grandis and P. laticauda) have become invasive across different countries and political territories, updated distributional data is also provided. Although a substantial number of the currently accepted taxa is well defined, the taxonomy of several species and subspecies is in need of revision and taxonomic changes are expected in future. We designate Phelsuma ornatum Gray, 1825 as type species of the genus Colobopus Fitzinger, 1843 in order to allow an unambiguous interpretation of this genus name.

Taxonomic checklist of chameleons (Squamata: Chamaeleonidae)

Due to their often brilliant colours, diurnal activity, and fascinating behaviours there is a long lasting high demand for chameleons in the pet trade. Accordingly, the international trade of most chameleon genera is regulated by the Convention on International Trade in Endangered Species of Wild Fauna and Floras (CITES). In order to facilitate operating and control of these regulations by national and international nature conservation authorities an updated taxonomic checklist of the family Chamaeleonidae (202 species plus 23 subspecies in 12 genera) is provided. A comparison with the last taxonomic checklist published in 1997 (132 species plus 39 subspecies in six genera) demonstrates the enormous progress in chameleon taxonomy and systematics in recent years. Although a substantial number of the currently accepted taxa are well defined, the taxonomy of several species and subspecies is in need of revision and many new species both from Africa and Madagascar still await their scientific description.

Subspecies and the philosophy of science

Phylogenetic methods increasingly are brought to bear on questions of subspecies taxonomy, but several recent examples highlight the need for a clear and consistent philosophical approach to how genetic data are used to assess subspecies limits. Such standards are crucial conceptually, whether or not taxonomic decisions affect conservation decisions, as they might in a recent study focused on the California Gnatcatcher (Polioptila californica), a taxon currently protected under the U.S. Endangered Species Act. It is also crucial that any adopted framework allows each of a full range of alternatives to be either supported or rejected. In this spirit, in addition to recommending best practices, I propose an amendment to the phylogenetic species concept to include a subspecies category.