Molecular phylogenetics and delimitation of species in Cortinarius section Calochroi (Basidiomycota, Agaricales) in Europe

Cortinarius is the most species rich genus of mushroom forming fungi with an estimated 2000 spp. worldwide. However, species delimitation within the genus is often controversial. This is particularly true in the section Calochroi (incl. section Fulvi), where the number of accepted taxa in Europe ranges between c.60 and c.170 according to different taxonomic schools. Here, we evaluated species delimitation within this taxonomically difficult group of species and estimated their phylogenetic relationships. Species were delimited by phylogenetic inference and by comparison of ITS sequence data in combination with morphological characters. A total of 421 ITS sequences were analyzed, including data from 53 type specimens. The phylogenetic relationships of the identified species were estimated by analyzing ITS data in combination with sequence data from the two largest subunits of RNA polymerase II (RPB1 and RPB2). Seventy-nine species were identified, which are believed to constitute the bulk of the diversity of this group in Europe. The delimitation of species based on ITS sequences is more consistent with a conservative morphological species concept for most groups. ITS sequence data from 30 of the 53 types were identical to other taxa, and most of these can be readily treated as synonyms. This emphasizes the importance of critical analysis of collections before describing new taxa. The phylogenetic separation of species was, in general, unambiguous and there is considerable potential for using ITS sequence data as a barcode for the group. A high level of homoplasy and phenotypic plasticity was observed for morphological and ecological characters. Whereas most species and several minor lineages can be recognized by morphological and ecological character states, these same states are poor indicators at higher levels.     

Hidden Diversity in Sardines: Genetic and Morphological Evidence for Cryptic Species in the Goldstripe Sardinella,Sardinella gibbosa (Bleeker, 1849)

Cryptic species continue to be uncovered in many fish taxa, posing challenges for fisheries conservation and management. In Sardinella gibbosa, previous investigations revealed subtle intra-species variations, resulting in numerous synonyms and a controversial taxonomy for this sardine. Here, we tested for cryptic diversity within S. gibbosa using genetic data from two mitochondrial and one nuclear gene regions of 248 individuals of S. gibbosa, collected from eight locations across the Philippine archipelago. Deep genetic divergence and subsequent clustering was consistent across both mitochondrial and nuclear markers. Clade distribution is geographically limited: Clade 1 is widely distributed in the central Philippines, while Clade 2 is limited to the northernmost sampling site. In addition, morphometric analyses revealed a unique head shape that characterized each genetic clade. Hence, both genetic and morphological evidence strongly suggests a hidden diversity within this common and commercially-important sardine.     

A new combination in Hybanthus (Violaceae) from South America

In a survey of the genusHybanthus in Brazil, it was found thatIonidium nanum A. St.-Hil. should be considered distinct fromHybanthus albus (A. St.-Hil.) Baill., based on characteristics of indument, habit and nectariferous appendages, and also habitat and geographical distribution. A new combination ofI. nanum withinHybanthus is therefore proposed.     

Developmental plasticity, morphological variation and evolvability: a multilevel analysis of morphometric integration in the shape of compound leaves

The structure of compound leaves provides flexibility for morphological change by variation in the shapes, sizes and arrangement of leaflets. Here, we conduct a multilevel analysis of shape variation in compound leaves to explore the developmental plasticity and evolutionary potential that are the basis of diversification in leaf shape. We use the methods of geometric morphometrics to study the shapes of individual leaflets and whole leaves in 20 taxa of Potentilla (sensu lato). A newly developed test based on the bootstrap approach suggests that uncertainty in the molecular phylogeny precludes firm conclusions whether there is a phylogenetic signal in the data on leaf shape. For variation among taxa, variation within taxa, as well as fluctuating asymmetry, there is evidence of strong morphological integration. The patterns of variation are similar across all three levels, suggesting that integration within taxa may act as a constraint on evolutionary change.     

海南产鳞始蕨属(鳞始蕨科)的分类学修订

澄清海南有鳞始蕨属Lindsaea Dryand.植物11种和1变种,其中碎叶鳞始蕨L. chingii C. Chr.为海南分布新记录。讨论了团叶鳞始蕨L. orbiculata (Lam.) Mett. ex Kuhn和钱氏鳞始蕨L. chienii Ching的形态变异。4个学名, 即L. annamensis K. U. Kramer、海南鳞始蕨L. hainanensis Ching、封开鳞始蕨L. fengkaiensis B. S. Wang &; S. H. Shi和海岛鳞始蕨L. orbiculata var. commixta (Tagawa) K. U. Kramer处理为异名。列出了分种检索表, 在每一分类群下列有文献引证、标本引证、生境及分布资料。     

Genetic variation of chloroplast DNA in Zingiberaceae taxa from Myanmar assessed by PCR–restriction fragment length polymorphism analysis

We examined genetic variation in 22 accessions belonging to 11 species in four genera of the Zingiberaceae, mainly from Myanmar, by PCR–restriction fragment length polymorphism analysis to investigate their relationships within this family. Two of 10 chloroplast gene regions ( trnS-trnfM and trnK2 – trnQr ) showed differential PCR amplification across the taxa. Restriction enzyme digestion of the PCR products revealed interspecific variability. The restriction patterns were used to classify the regions as either highly conserved or variable across the taxa. None of the regions was highly conserved across the four genera, and the level of conservation varied. The gene region trnS-trnfM appeared to display interspecific variability among most of the species. However, the relative efficiency of different restriction enzymes depended on the gene regions and genera investigated. Cluster analysis revealed interspecific discrimination among the taxa. The two Curcuma species ( Curcuma zedoaria and Curcuma xanthorrhiza ) appeared to be identical, thus supporting their recent classification as synonyms. The results provide the basis for selecting specific combinations of restriction enzymes and gene regions of chloroplast DNA (cpDNA) to identify interspecific variation in the Zingiberaceae and to identify both highly conserved and variable regions. Overall, cpDNA depicted comparatively diverse genetic profile of the studied germplasm. The genetic information revealed here can be applied to the conservation and future breeding of Zingiber and Curcuma species.     

Making the cryptic visible – resolving the species complex of Aphodius fimetarius (Linnaeus) and Aphodius pedellus (de Geer) (Coleoptera: Scarabaeidae) by three complementary methods

Species in cryptic complexes are, per definition, difficult to identify using morphological characters. One such complex was recently detected in the dung beetle Aphodius fimetarius (Linnaeus) sensu lato, an abundant dung beetle with a wide distribution. While the two component taxa, Aphodius fimetarius sensu stricto and Aphodius pedellus (De Geer) exhibit distinctly different karyotypes, the validity of subtle morphological characters proposed to distinguish between them has been debated. Given the variability and minor interspecific differences in external characters, the large‐scale distribution of respective taxa has remained unknown, as have potential differences in ecology and habits. In this study, we ask how A. fimetarius and A. pedellus can best be distinguished, whether the use of different types of characters (karyotypes, DNA sequences and morphological traits) results in consistent species identification, where these species occur and whether they exhibit ecological differences. In total, we inspected a material of 4401 individuals from across the globe, of which 183 were examined for both mtDNA sequences and morphology, 154 for both morphology and karyotype, and 9 (including the recently proposed neotype of Aphodius fimetarius) for all three types of characters. As a marker gene, we sequenced a 590 bp region of the cytochrome c oxidase I gene for 183 individuals. Overall, DNA sequences offered a clear‐cut distinction between taxa: sequences of A. fimetarius and A. pedellus differed by an average pairwise distance of 8.2%, whereas variation within species was only 0.9% for A. fimetarius and 0.5% for A. pedellus. Morphological and chromosomal characters offered species identifications consistent with that of molecular characters: karyotypes identified as A. pedellus consistently fell within one of the molecular clades, whereas karyotypes identified as A. fimetarius fell within the other clade. Likewise, the majority of individuals identified by morphological characters were assigned to the same species by sequence‐based characters. Both taxa thus defined were found to be Holarctic in distribution, with major sympatry within Central and Southern Europe and mixed patterns of sympatry within the US. Northern areas of Europe, Asia and North America are dominated by A. pedellus alone. Within A. pedellus, patterns of sequence diversity were indicative of a recent population expansion. In the western US, the phenology of a population of A. fimetarius was observed to significantly differ from that of a sympatric population of A. pedellus, thereby revealing an ecological difference between the two cryptic taxa. Overall, we conclude that all types of characters offered a consistent classification of the two species. Thus, the laborious karyotyping techniques used to originally establish the presence of two cryptic taxa can now be substituted by characters more easily applied to large ecological samples. Using this approach of integrative taxonomy, we were able to establish the global distribution and species‐specific ecology of these ecologically important cryptic taxa. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:4033473E-8BF7-40F4-852D-916E4F858593 .     

Phylogeny and composition of the Hieroxestinae (Lepidoptera: Tineidae)

The tineid subfamily Hieroxestinae, comprising 275 valid species, is reviewed. The morphology of the group is examined and a fully resolved phylogeny presented for the six constituent genera. Eighteen generic synonyms are noted of which three are new; fourteen specific synonyms are noted, one of them new. Seventy-one new specific combinations are established. A comprehensive checklist of species is provided as an appendix. Three species are left unplaced in Oinophila sensu lato , and two species (one with two subspecies) are transferred to Oecophoridae (Stathmopodinae). The biology of the Hieroxestinae is briefly reviewed. Hieroxestinae larvae are predominantly detritophagous but several species feed facultatively on green plant material, and Opogona sacchari is a pest of seedlings and nursery stock in the U.S.A. and Mediterranean region. The group is predominantly tropical and is distributed throughout the world but is depauperate in the Americas. There has been considerable radiation of Opogona and Amphixystis on the islands of the Indian Ocean that lie within the Malagasy Subregion (Seychelles, Mauritius, Réunion, etc.) where there are fifty-three endemic species. A lesser radiation of Opogona (twenty species) has occurred on St Helena.     

Evolutionary diversification of banded tube-dwelling anemones (Cnidaria; Ceriantharia; Isarachnanthus) in the Atlantic Ocean

The use of molecular data for species delimitation in Anthozoa is still a very delicate issue. This is probably due to the low genetic variation found among the molecular markers (primarily mitochondrial) commonly used for Anthozoa. Ceriantharia is an anthozoan group that has not been tested for genetic divergence at the species level. Recently, all three Atlantic species described for the genus Isarachnanthus of Atlantic Ocean, were deemed synonyms based on morphological simmilarities of only one species: Isarachnanthus maderensis. Here, we aimed to verify whether genetic relationships (using COI, 16S, ITS1 and ITS2 molecular markers) confirmed morphological affinities among members of Isarachnanthus from different regions across the Atlantic Ocean. Results from four DNA markers were completely congruent and revealed that two different species exist in the Atlantic Ocean. The low identification success and substantial overlap between intra and interspecific COI distances render the Anthozoa unsuitable for DNA barcoding, which is not true for Ceriantharia. In addition, genetic divergence within and between Ceriantharia species is more similar to that found in Medusozoa (Hydrozoa and Scyphozoa) than Anthozoa and Porifera that have divergence rates similar to typical metazoans. The two genetic species could also be separated based on micromorphological characteristics of their cnidomes. Using a specimen of Isarachnanthus bandanensis from Pacific Ocean as an outgroup, it was possible to estimate the minimum date of divergence between the clades. The cladogenesis event that formed the species of the Atlantic Ocean is estimated to have occured around 8.5 million years ago (Miocene) and several possible speciation scenarios are discussed.     

Parallel Patterns of Morphological and Behavioral Variation among Host-Associated Populations of Two Gall Wasp Species

A powerful approach to address the general factors contributing to ecological speciation is to compare distantly related taxa that inhabit the same selective environments. In this design, similarities among taxa can elucidate general mechanisms of the process whereas differences may uncover specific factors important to the process for individual taxa. Herein, we present evidence of parallel patterns of morphological and behavioral variation among host-associated populations of two species of cynipid gall wasps, Belonocnema treatae and Disholcaspis quercusvirens, that each exhibit a life cycle intimately tied to the same two host plant environments, Quercus geminata and Q. virginiana. Across both gall-former species we find consistent differences in body size and gall morphology associated with host plant use, as well as strong differences in host plant preference, a measure of habitat isolation among populations. These consistent differences among taxa highlight the important role of host plant use in promoting reproductive isolation and morphological variation among herbivorous insect populations–a prerequisite for ecological speciation.     

A phylogenetic study of the generic relationships within the subtribe Leucophoropterina Schuh (Miridae: Phylinae: Leucophoropterini)

The subtribe Leucophoropterina (Miridae: Phylinae: Leucophoropterini) is a relatively small lineage of mirids comprising 23 genera and 104 species. Most of the species are endemic to the Indo‐Pacific and Australia, and many are considered ant mimics. A phylogenetic analysis including both the Australian and the Indo‐Pacific taxa was conducted for the first time with a dataset of 137 morphological characters coded for all but two known species of Leucophoropterina. The hypothesis by Schuh ( 1984 ) of at least two major lineages of ant‐mimicking taxa based on his analysis of the Indo‐Pacific taxa continues to be supported with our inclusion of Australian taxa. The Australian taxa and Indo‐Pacific taxa of Leucophoropterina do not form independent, geographically based lineages, but instead comprise several lineages containing taxa from both regions. This study provides a basis for future detailed studies of biogeography and ant mimicry in the group.     

Andromonoecy in Solanum lycocarpum A. St. -Hil. (Solanaceae): Floral attributes,visitors and variation in sexual expression over time

Sexual expression in andromonoecious species—those in which a single individual can bear both staminate and hermaphroditic flowers—may vary among reproductive events in the same plant, among individuals and across populations. This variation influences, in turn, the individual contribution of hermaphroditic plants via male and female fitness functions (i.e., Lloydʼs phenotypic gender). However, temporal variation in sexual expression in andromonoecious species and its relationship with seasonal changes in climatic conditions remain poorly understood. Here we analyze floral attributes, visitors and variation in sexual expression in three populations of Solanum lycocarpum A. St. -Hil. Seasonality in the production of floral types, the mating system and floral visitors were also investigated. Hermaphroditic flowers produced more pollen grains, but the pollen of staminate flowers had higher viability. Only hermaphroditic flowers produced fruits, and ovules in staminate flowers were sterile. Solanum lycocarpum is mainly pollinated by large bees with the ability to vibrate flowers. Phenotypic gender varied throughout the year, and the seasonal production of staminate flowers is associated with the local climate. We suggest that the higher and seasonally variable relative abundance of staminate flowers compared to the low and uniform production of hermaphroditic flowers may be explained by (a) the very high energetic costs incurred in producing large fruits, which in turn make hermaphroditic flower production very costly, and (b) the potentially lower energy expenditure of the smaller staminate flowers with reduced pistils and non-viable ovules that allow them to rapidly respond to climate variability.     

NATURAL SELECTION ON HYDROID COLONY MORPHOLOGY BY INTRASPECIFIC COMPETITION

Previous work on colonial hydroids in the genus Hydractinia has demonstrated that colony morphology is highly variable and determines intraspecific competitive ability. Competitive encounters are known to be common in nature, suggesting that intraspecific competition may be a major selective force acting on morphological variation. A replicated common garden experiment demonstrated a genetic basis to morphological variation and two data sets provided correlative support for the hypothesis of selection by intraspecific competition. First, morphologies inferior in competitive ability were less abundant in two adult, postcompetition, samples than in juvenile, precompetition, samples from the same populations. Second, among eight populations, the relative frequency of different morphologies was correlated with the frequency of intraspecific competition observed in each population. The direction of selection by competition on the morphological variation present in this species conflicts with recent predictions based on surveys across diverse taxa, suggesting limitations to the inference of competition as a past selective agent on the basis of present day correlations among species.     

Taxonomic revision of Rhododendron mariae (Ericaceae) and related taxa

Based on a literature survey and a field trip with specimen examination, statistical analysis including principal components analysis (PCA) on morphological characters have been carried out on Rhododendron mariae and related taxa. Cluster analysis and PCA showed three independent groups, which corresponded to three species except R. loniceriflorum . As a consequence, nine taxa are reduced to synonyms of R. mariae , and two new combinations, viz R. mariae subsp. microphyton , and R. mariae subsp. flumineum are made. Rhododendron microphyton var. trichanthum is reduced to a synonym of R. mariae subsp. microphyton , and R. malipoense to a synonym of R. mariae subsp. flumineum . Three names are synonymized and a new combination, viz R. chryocalyx subsp. meridionale , is proposed within R. chryocalyx . The third entity, R. naamkwanense , has got two new synonyms. Rhododendron loniceriflorum , with whitish pubescence on the inner corolla surface, is recognized as the fourth distinct species. Each entity is illustrated with line drawings for identification.     

Patterns of ecomorphological variation in the bats of western Madagascar: Comparisons among and between communities along a latitudinal gradient

The ecomorphology of 10 insectivorous bat species at three study zones in western Madagascar was examined using 567 specimens and based on 6 external, 11 cranial, 12 dental, and 11 wing measurements. The three study sites are located along a cline representing 11.6° of latitude. The southern most site has notable differences in vegetational and climatic regimes than the two more northern sites. Principal component analyses were conducted for each of the four datasets to examine the morphological space occupied by each species at the three sites. Most taxa showed clear intra-site separation and little inter-site variation. The exceptions included extensive morphological overlap in two taxa of Triaenops (cranial, dental, and wing), that have allopatric distributions, and between the sympatric Miniopterus manavi and Myotis goudoti (external, cranial, and dental). In the latter case, there was distinct separation in wing shape between these two taxa, which would allow them to exploit local habitats and prey in different manners. The only species that showed considerable inter-site variation was Hipposideros commersoni, which is sexually dimorphic, with individuals from the south being smaller than those in the north.     

Morphological Variation of the Scorpionfly Panorpa obtusa Cheng (Mecoptera: Panorpidae) with a New Synonym

Background

The overabundance of synonyms is an unavoidable by-product of taxonomic practice in insects. How to reduce or even eliminate synonymy has long been a great challenge for insect taxonomists. The scorpionflies Panorpa obtusa Cheng, 1949 and Panorpa leei Cheng, 1949 (Insecta: Mecoptera: Panorpidae) were originally described from Taibaishan in the Qinling Mountains with identical collection data and both are based on a single gender, the former on a male and the latter on two females. However, whether P. leei is conspecific with P. obtusa or a good species remains an unsolved problem.

Results

On the basis of intensive morphological comparison of 93 males and 53 females of scorpionflies collected from the type locality using light and scanning electron microscopy, we found P. obtusa has considerable morphological variation (especially the wing markings and genitalia in both male and female), and Panorpa leei is totally comprised of one of the morphs of P. obtusa.

Conclusions

In combination with identical type localities and overlapping morphological variation, P. leei Cheng is proposed as a junior subjective synonym of P. obtusa Cheng. To avoid synonyms, taxonomists should pay more attention to individual variation and base decisions on a series of specimens to describe new species.