Stipa adamii sp. nov. (Poaceae) from the western Tian‐Shan,and some remarks on the taxa of the section Smirnovia occurring in Kazakhstan

Stipa adamii M. Nobis sp. nov. (Poaceae), endemic to Kazakhstan, is described and illustrated. The species belongs to the section Smirnovia Tzvel. and is distributed in the central Karatau Mountains (western Tian‐Shan). It is morphologically close to Stipa karataviensis Roshev. but is easily distinguishable by longer glumes, lemmas, awns and the callus. The awn is scabrous in the lower part and the callus is bearded in S. adamii whereas both organs are glabrous in S. karataviensis. Characters distinguishing S. karataviensis and S. adamii are presented in a table, together with photographs of the callus. The analysis of morphological characters of the taxa provides new information on their variability. A key to the species of Stipa sect. Smirnovia occurring in Kazakhstan, typification of the name S. manrakica and synonymization of the name S. saurica as a syn. nov. are included.     

A revision of the Nasa ranunculifolia group (Nasa ser. Grandiflorae pro parte,Loasaceae)

The monophyletic Nasa ranunculifolia group (Nasa ser. Grandiflorae pro parte) is revised on the basis of extensive field studies and a revision of copious herbarium material. All species of this group are from the high Andes of Peru and Ecuador (mostly 3000–4300 m). The overall degree of morphological divergence is much lower than in other groups of Nasa and many of the various local ‘races’ are best recognized as ecogeographically isolated subspecies. Thus, six species and 13 subspecies are recognized on the basis of differences in leaf and floral morphology, growth habit, life history and distribution. Nasa ranunculifolia, N. macrantha, N. cymbopetala and N. macrorrhiza are united as subspecies under N. ranunculifolia. An additional five subspecies of N. ranunculifolia are newly described (ssp. pamparomasii, ssp. guzmangoensis, ssp. bolivarensis, ssp. patazensis and ssp. huanucoensis). Nasa rugosa is subdivided into four subspecies, three of them new (ssp. llaqtacochaensis, ssp. gracilipes and ssp. pygmaea). Nasa tulipadiaboli sp. nov. and N. basilica sp. nov. are described as new species. Eight taxa (seven subspecies and one species) are endemic to the Amotape–Huancabamba Zone, and seven of them (six subspecies and one species) are new to science. Only a single species ranges into the Amotape–Huancabamba Zone from the south, underscoring the high levels of endemism (and taxonomic novelty) in this area. Illustrations, distribution maps, a key and diagnoses are given for all taxa recognized. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 47–93.     

Molecular phylogenetics and taxonomic revision of Habenaria section Pentadactylae (Orchidaceae,Orchidinae)

As a step towards a revision of the sectional classification of Neotropical species of Habenaria, we focus here on section Pentadactylae. In its current delimitation, this is the largest of the 14 New World sections and embraces a group of 34 morphologically heterogeneous species. We expanded the sampling of Neotropical species currently placed in this section and performed Bayesian, maximum likelihood and parsimony analyses using nucleotide sequences from one nuclear (internal transcribed spacer, ITS) and three plastid (matK, trnK intron, rps16–trnK) DNA regions. In addition, morphological features of these species were reassessed. Based on our analyses, we propose that Habenaria section Pentadactylae should be recircumscribed to include only seven species: H. pentadactyla (the type species of the section), H. dutrae, H. ekmaniana, H. exaltata, H. henscheniana, H. megapotamensis and H. montevidensis. Thirty‐two species previously assigned to the section grouped within unrelated clades and are therefore excluded from the section. There are no unambiguous morphological synapomorphies for the section, but the group can be confidently recircumscribed and identified on the basis of a combination of diagnostic morphological vegetative and floral characters. Morphological floral features in Habenaria montevidensis are distinct from those of other species in the section, probably as a result of a shift to diurnal pollinators. Following a taxonomic revision of the group, H. crassipes is placed under the synonymy of H. exaltata and neotypes are designated for H. crassipes, H. montevidensis and H. recta (= H. ekmaniana). All species in the section live in marshes or wet grasslands from northern Argentina to central Brazil; most species are concentrated in southern Brazil. Most species are probably rare, and five may be threatened according to the World Conservation Union (IUCN) criteria. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 47–73.     

Genetic patterns in the Lathyrus pannonicus complex (Fabaceae) reflect ecological differentiation rather than biogeography and traditional subspecific division

The endangered European relict species complex Lathyrus pannonicus shows distinct morphological variation, reflected by the number of subspecies recognized, and complicated patterns of genetic variation. The traditionally recognized subspecies appear to possess different ecological preferences and disjunct distributions, particularly in the western range of the species. In this study, L. pannonicus was investigated by the correlation of distance matrices based on phytosociological, ecological, molecular and morphological data. Ecological characteristics of selected stands of L. pannonicus throughout Europe were assessed using ‘Ellenberg values’ of all the constituent taxa in phytosociological relevés. Genetic distances were calculated using recently developed methods to analyse high degrees of intra‐individual nuclear‐encoded internal transcribed spacer variability. We found that the remarkable genetic (and morphological) diversity in L. pannonicus could not be explained solely by the fragmentation of the distributional range. Instead, patterns of morphological and genetic differentiation were a reflection of the moisture regime in the sampled stands. Two major lineages could be identified: (1) a lineage adapted to dry conditions (Ellenberg indicator F‐value ≤ 3.5) and (2) a lineage preferring moist conditions (F‐value ≥ 4.5). Although both lineages occurred in close proximity in the Pannonian area, they appeared to be reproductively isolated in general. Further data are needed to determine whether these genetically and ecologically defined lineages, or ecospecies, within the L. pannonicus species complex can be formalized as (Linnaean) species or subspecies. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 402–421.     

A new hexaploid species of Centaurea section Acrolophus (Asteraceae) from Evvia Island,Greece

Centaurea carystea Trigas & Constantin., a new yellow‐flowered species of Centaurea from Mt. Ochi on Evvia Island (Greece), is described and illustrated. It is a member of the polymorphic C. section Acrolophus and allied to taxa of the C. attica aggregate, C. pelia and C. mantoudii. The new species appears to be a local and threatened endemic, with the total number of individuals known being less than 500. A karyological examination revealed that it is hexaploid, with 2n = 6x = 54, an unusual number in Centaurea, which may indicate a hybrid origin. To further clarify the taxonomic position of C. carystea , we used random amplification of polymorphic DNA markers of 25 plants belonging to nine species, subspecies and varieties morphologically related to the new species, together with two reference taxa. Clustering using the unweighted pair group method with arithmetic mean indicated a discrete position for C. carystea , close to but distinct from the yellow‐flowered C. mantoudii. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 762–774.     

Nutlet micromorphology of Carex section Rhomboidales sensu Kükenthal (Cyperaceae) and its systematic implications

The sectional delimitation of Carex section Rhomboidales sensu Kükenthal (1909) is problematic and, here, we provide contributions to a systematic understanding of nutlet micromorphology in section Rhomboidales. Nutlet micromorphology of 87 samples representing 71 taxa was investigated using scanning electron microscopy and nine nutlet types were recognized. Although some closely related taxa show similarities, nutlet morphology in section Rhomboidales is stable at the species level and is reliable for species identification. Based on the nutlet characters, section Rhomboidales is delimited and emended as having nutlet rhombic–ovoid, trigonous; beaks conspicuous, erect, rarely curved or coiled; periclinal walls of epidermal cells straight, with zero to two silica bodies. The species with nutlets of the C. harlandii‐type and C. thibetica‐type are included, and species of the C. chinensis‐type and C. macrandrolepsis‐type are ascribed to section Mitratae and section Infossae, respectively. Carex longirostrata and C. pseudolongirostrata (= C. nodaeana), formerly ascribed to section Careyanae or section Depauperatae, are well supported as members of section Rhomboidales. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 123–143.     

Clinal variation or validation of a subspecies? A case study of the Graptemys nigrinoda complex (Testudines: Emydidae)

Widely distributed species often display intraspecific morphological variation due to the abiotic and biotic gradients experienced across their ranges. Historically, in many vertebrate taxa, such as birds and reptiles, these morphological differences within a species were used to delimit subspecies. Graptemys nigrinoda is an aquatic turtle species endemic to the Mobile Bay Basin. Colour pattern and morphological variability were used to describe a subspecies (G. n. delticola) from the lower reaches of the system, although it and the nominate subspecies also reportedly intergrade over a large portion of the range. Other researchers have suggested that these morphological differences merely reflect clinal variation. Our molecular data (mtDNA) did not support the existence of the subspecies, as the haplotypes were differentiated by only a few base pairs and one haplotype was shared between the putative subspecies. While there were significant morphological and pattern differences among putative specimens of G. n. nigrinoda, G. n. delticola and G. n. nigrinoda × delticola, these differences probably represent clinal variation as they were also related to environmental variables [i.e. cumulative drainage area and drainage (categorical)]. Specimens occupying slow‐current, high‐turbidity river reaches (e.g. the Tensaw River) exhibited greater relative carapace heights and more dark pigmentation, while specimens occupying fast‐current, clearer rivers (e.g. the upper Alabama, Cahaba and Tallapoosa rivers) exhibited lower carapace heights and more yellow pigmentation. Given the absence of clear molecular and morphological differences that are related to drainage characteristics, we suggest that there is not sufficient evidence for the recognition of G. n. delticola as a distinct subspecies. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 810–822.     

Complex evolutionary relationships in Origanum section Majorana (Lamiaceae)

Origanum (Lamiaceae) comprises a number of essential oil‐rich species that have been used by humans for centuries. Today, the four species of section Majorana (O. onites, O. dubium, O. majorana and O. syriacum) are amongst the most widely used. Despite the importance of this section, phylogenetic relationships and species boundaries amongst its four taxa are unclear. In the present investigation, we used DNA sequence data from two nuclear regions [internal transcribed spacer (ITS) and 1‐deoxy‐d ‐xylulose 5‐phosphate synthase (DXS)] as well as five microsatellite loci to test the taxonomic status of the four species of section Majorana. The combined DNA data revealed O. onites and O. syriacum as the older species in the section. Origanum majorana descends directly from O. syriacum. Origanum dubium was found to be of hybridogenous origin showing attributes of O. onites, O. syriacum and a third, unknown, Origanum species. Both sequence and microsatellite analyses provided evidence for recent hybridization between O. onites and O. dubium in Turkey. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 667–686.     

Molecular phylogenetics of the juno irises,Iris subgenus Scorpiris (Iridaceae), based on six plastid markers

Relationships among the roughly 55 species of Iris subgenus Scorpiris have been studied. A matrix of six plastid DNA regions (matK, rpl14‐rps8 spacer, infA‐rpl36 spacer, trnE‐trnT spacer, trnL intron and trnL‐F spacer) was produced from 57 accessions (52 taxa) and analysed with both parsimony and Bayesian methods. Five major clades are identified, of which four have strong geographical correlations, whereas the fifth corresponds to Iris section Physocaulon. In our results, several species are placed with species not previously considered to be related, although, in some cases, there are morphological characters that suggest that these newly indicated relationships are reasonable. For some of the other oddly grouped species, we can only assume that remarkable parallelisms in morphology have occurred or hybridization is involved. Presently, with plastid DNA as our only comprehensive data resource, we are not able to evaluate more thoroughly these more puzzling associations of species. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 281–300.     

Phylogenetic interrelationships,taxonomy, and reductive evolution in the Neotropical electric fish genus Hypopygus (Teleostei,Ostariophysi, Gymnotiformes)

A phylogenetic reconstruction of the Neotropical electric fish genus Hypopygus based on 47 parsimony‐informative morphological characters is presented. A series of synapomorphies support the hypothesis of monophyly of Hypopygus, and partially resolve species‐level relationships within the genus. Hypopygus species are recognized here as miniaturized fishes based on two criteria; first, a derived condition of diminutive body size, and; second, the presence of a suite of reductive morphological characters, including partial or total losses, simplifications, and reductions of the anal‐fin rays, scales, cranial bones, and laterosensory canal system. Reductive characters associated with miniaturization comprise 45% of the total number of characters in the phylogenetic reconstruction of the genus. Miniaturization and reductive morphological evolution in Hypopygus are discussed here in the phylogenetic context. A taxonomic revision of Hypopygus is presented, in which five new species are described, two species previously assigned to the genus are redescribed, and a single known species of Stegostenopos is redescribed and included in Hypopygus as a junior synonym. Distribution maps and a key for all eight valid species of Hypopygus are provided, based on the examination of 5014 catalogued museum specimens. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1096–1156.     

Fine‐scale patterns of genetic divergence within and between morphologically variable subspecies of the sea urchin Heliocidaris erythrogramma (Echinometridae)

The spatial scale over which genetic divergences occur between populations and the extent that they are paralleled by morphological differences can vary greatly among marine species. In the present study, we use a hierarchical spatial design to investigate genetic structure in Heliocidaris erythrogramma occurring on near shore limestone reefs in Western Australia. These reefs are inhabited by two distinct subspecies: the thick‐spined Heliocidaris erythrogramma armigera and the thin‐spined Heliocidaris erythrogramma erythrogramma, each of which also have distinct colour patterns. In addition to pronounced morphological variation, H. erythrogramma exhibits a relatively short (3–4 days) planktonic phase before settlement and metamorphosis, which limits their capacity for dispersal. We used microsatellite markers to determine whether patterns of genetic structure were influenced more by morphological or life history limitations to dispersal. Both individual and population‐level analyses found significant genetic differentiation between subspecies, which was independent of geographical distance. Genetic diversity was considerably lower within H. e. erythrogramma than within H. e. armigera and genetic divergence was four‐fold greater between subspecies than among populations within subspecies. This pattern was consistent even at fine spatial scales (< 5 km). We did detect some evidence of gene flow between the subspecies; however, it appears to be highly restricted. Within subspecies, genetic structure was more clearly driven by dispersal capacity, although weak patterns of isolation‐by‐distance suggest that there may be other factors limiting gene exchange between populations. Our results show that spatial patterns of genetic structure in Western Australian H. erythrogramma is influenced by a range of factors but is primarily correlated with the distribution of morphologically distinct subspecies. This suggests the presence of reproductive barriers to gene exchange between them and demonstrates that morphological variation can be a good predictor of genetic divergence. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 578–592.     

Systematics of Dendrobiinae (Orchidaceae), with special reference to Australian taxa

The systematics of subtribe Dendrobiinae, Dendrobium, and Australian dendrobiums in particular, are reviewed. Molecular approaches have delimited the subtribe, established Asian and Australasian clades and clarified relationships at some levels within them. At most lower taxonomic levels within Dendrobium s.l., resolution has been partial. Morphological and molecular phylogenetics indicate that Dendrocoryne, Rhizobium and some other sections are not monophyletic, with morphological analyses providing the greatest resolution into groups corresponding with biogeography. For most sections, sampling is too incomplete to determine relationships and monophyly. Distribution, pollination syndromes, breeding systems, fertility and specialized anatomical features contribute little to classification. Natural hybrids are more common than previously stated and may have played a role in the high level of homoplasy and evolution of Australian epiphytes. There is little support for splitting Dendrobium into many genera and species based on terminal clades. The evidence is interpreted as favouring recombination of Cadetia, Flickingeria and Diplocaulobium within the conserved broad genus Dendrobium. Nomenclatural stability is retained in continuing to use the long‐held subgeneric sectional classification, including many of the sections of Schlechter, and infraspecific ranks of subspecies and varieties. In Dendrocoryne, Thelychiton, Tropilis and Tetrabaculum are rejected, as is Dockrillia in Rhizobium. Proposals to elevate infraspecific taxa to species level for variable species complexes are not supported by current morphological or molecular data based on slowly evolving DNA regions. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 105–126.     

Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses

Adults of the Euthalia phemius complex, which is composed of three South‐East Asian nymphalid species, Euthalia phemius, Euthalia ipona, and Euthalia euphemia, were genetically analysed by examining mitochondrial and nuclear genes. The E. phemius complex was also examined morphologically, with particular emphasis on wing markings and male genitalia. No significant differences amongst the three species in the complex were detected with respect to either genetic distance or genital morphology. We therefore conclude that the three currently recognized Euthalia species belong to a single species. Accordingly, E. ipona is synonymized with E. phemius. Euthalia euphemia is treated as a subspecies of E. phemius. Type specimens of all taxa and a synonymic list for the E. phemius complex are also given. In addition, we briefly discuss the evolution and biogeography of the species complex. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 304–327.     

Resurrection in Carapa (Meliaceae): a reassessment of morphological variation and species boundaries using multivariate methods in a phylogenetic context

The taxonomy of the amphi‐Atlantic tree genus Carapa (Meliaceae) has long been controversial. Of the three species currently recognized in the genus, two are known to present substantial morphological variation that has been used in the past to distinguish several taxa, most of which are currently placed in synonymy. Here, a combination of field observations, univariate analyses of leaf, floral and seed characters and principal coordinate analyses of floral characters in the context of a molecular phylogenetic analysis was used to investigate the patterns of variation and delimit morphological species anew in the genus. These results support the recognition of 27 species in Carapa, of which 16 are previously described and 11 are new. In general, phylogenetically related species occurred in the same geographical area, but were morphologically distinct. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 186–221.     

How common are dot‐like distributions? Taxonomical oversplitting in western European Agrodiaetus (Lepidoptera: Lycaenidae) revealed by chromosomal and molecular markers

Approximately 50 taxa of butterflies in Western Europe have been described as new species or elevated to the level of species during the last 40 years. Many, especially those belonging to the genus Agrodiaetus, have unusually localized, ‘dot‐like’ distributional ranges. In the present study, we use a combination of chromosomal and molecular markers to re‐evaluate the species status of Agrodiaetus distributed west of the 17th meridian. The results obtained do not support the current designations of Agrodiaetus galloi, Agrodiaetus exuberans, and Agrodiaetus agenjoi as endemic species with highly restricted distribution ranges, but indicate that these taxa are more likely to be local populations of a widely distributed species, Agrodiaetus ripartii. Agrodiaetus violetae is shown to be a polytypic species consisting of at least two subspecies, including Agrodiaetus violetae subbaeticus comb. nov. and Agrodiaetus violetae violetae. Agrodiaetus violetae is genetically (but not chromosomally) distinct from Agrodiaetus fabressei and has a wider distribution in southern Spain than previously believed. Agrodiaetus humedasae from northern Italy is supported as a highly localized species that is distinct from its nearest relatives. We propose a revision of the species lists for Agrodiaetus taking these new data into account. The results reported in the present study are relevant to animal conservation efforts in Europe because of their implications for IUCN Red List priorities. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 130–154.     

Phylogenetic analysis of the leafhopper genus Apogonalia (Insecta: Hemiptera: Cicadellidae) and comments on the biogeography of the Caribbean islands

A phylogenetic analysis of the leafhopper genus Apogonalia was conducted based on a matrix of 40 terminal taxa and 147 morphological characters. The analysis yielded 1391 equally most‐parsimonious trees, which do not support the monophyly of Apogonalia in the strict consensus. A successive weighting procedure yielded 62 trees in which the genus appeared as a monophyletic group. The strict consensus of these 62 trees is almost entirely dichotomous, showing only two polytomies. The test of phylogenetic integrity was applied for distinct variations of three species: A. germana, A. sanguinipes, and A. histrio. Only for the first species was the conjecture that its variations belong to the same entity corroborated. The best‐supported clade within Apogonalia, which has several synapomorphies and high branch support indices, comprises nine Antillean endemic species. This distributional pattern probably was originated by vicariance in the Late Cretaceous, when the Proto‐Antillean archipelago was pushed north‐eastward by the Caribbean Plate to become the modern Greater Antilles. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 548–570.     

Phylogenetic analysis and differentiation of Veronica subgenus Stenocarpon in the Balkan Peninsula

The Balkan Peninsula is considered the most important refugium for species during the Pleistocene glaciations and today harbours c. 2000 endemic species, but we know surprisingly little about the evolution of taxa in this region. Veronica saturejoides, V. thessalica and V. erinoides are a group of closely related alpine taxa endemic to the Balkan Peninsula. Here, we analyse four DNA regions [the nuclear chalcone synthase intron (CHSi) and ribosomal internal transcribed spacer (ITS) region and the plastid rpoB‐trnC spacer and trnL‐trnL‐trnF region] and amplified fragment length polymorphism (AFLP) fingerprints to provide a phylogenetic hypothesis for the relationships among these taxa. Additionally, we analyse leaf morphological characters used to distinguish the three subspecies of V. saturejoides. The analyses support the distinction of the three subspecies based on previously intuitively suggested characters. Nuclear chalcone synthase intron data indicate that the southern taxa are genetically much more diverse than the more northern V. saturejoides subsp. saturejoides. Phylogenetic relationships inferred from this region and AFLP fingerprints support the monophyly of V. saturejoides. In contrast, plastid DNA regions suggest a closer relationship of V. saturejoides subsp. saturejoides to V. thessalica. The most likely scenario involves introgression into V. saturejoides subsp. saturejoides from V. thessalica. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 616–636.