短孢角毛藻相似种类的形态学和系统学研究

短孢角毛藻的形态特征多变,容易导致物种的错误鉴定,并且其与若干相似种类之间存在物种混淆的情况。基于建立的单克隆培养藻株,结合生活史中形态特征的连续观察,以及基于目标基因序列分析的分子系统学数据,对短孢角毛藻(Chaetoceros brevis Schütt)及其相似种类的形态学和系统学进行了研究。丰富了短孢角毛藻休眠孢子的形态学描述,报道了一个新的特征:休眠孢子壳面上刺的末端具有帽状结构,该特征未报道于其他角毛藻种类,可作为短孢角毛藻的标志性特征。报道了我国角毛藻属的1个新记录种——拟短孢角毛藻(C.pseudobrevis Pavillard),该种具有独特的形态学特征,以及独立的系统学位置,应是一个独立的物种,而非短孢角毛藻的同种异名。垂缘组角毛藻的种类在系统发育树中显示出较好的单系起源,链端角毛及其走势是否可以作为该类群的共享特征,目前尚难做出定论。     

海链藻属重要形态学特征的二次研究——以艾伦海链藻为例

海链藻属（Thalassiosira Cleve）是硅藻门中物种多样性最为丰富的大属之一,广泛分布于全球各类水体。目前沿用的海链藻属分类体系是基于形态特征建立的,尤其是电镜下的超微特征具有重要的分类价值。然而,也有研究认为某些形态特征存在不稳定或变化的情况,应对其开展二次研究和重新评估。故此,本文选择全球近岸广布种艾伦海链藻（T.allen Takano）作为典型代表,基于建立的单克隆培养株系,结合传统形态学方法和现代分子生物学技术,对其重要形态特征进行了二次研究。结果显示：（1）艾伦海链藻壳面孔纹的排列方式存在一定的变化,呈切线形或偏心状;壳缘唇形突的位置多有变化,占据一个壳缘支持突的位置、或位于两个壳缘支持突正中间、或靠近其中一个。艾伦海链藻这两个特征并不具有较高的稳定性,其分类学意义或应降低;（2）壳缘不具有肋纹,此特征相对稳定,可作为艾伦海链藻的识别特征之一;（3）艾伦海链藻具有较保守的遗传多样性,尚未发现（拟）隐形种。结合其它海链藻的形态学报道,我们认为孔纹排列方式易受环境的影响,稳定性较差,在分类时应慎重应用;壳缘唇形突的位置在少数海链藻属物种中易发生变化,分类时应视具体物种而针对性应用;支持突等突起特征较为稳定,在海链藻属物种的分类体系中仍可作为重要依据;艾伦海链藻中壳缘肋纹具有较好的稳定性,可作为分类依据,但在其它海链藻属物种中是否也具有很好的分类价值,还需要更多的研究加以验证;分子信息在形态特征上的二次研究和评价中具有重要应用价值,但目前相关分子数据的缺乏和不足限制了对其进一步的研究。本研究对重新评估艾伦海链藻形态特征的分类学价值、完善现存的分类学体系提供了参考。     

Polyphyletic Alyssum cuneifolium (Brassicaceae) revisited: Morphological and genome size differentiation of recently recognized allopatric taxa

Alyssum cuneifolium has been recognized as a perennial alpine species growing in five isolated European mountain ranges: the Pyrenees, Western Alps, Apennines, Pirin Mts and Mt Smolikas. Recent molecular systematic studies revealed that the disjunct populations from distant mountains are not closely related and belong to five independent species: A. cacuminum (Spain, Pyrenees), A. cuneifolium (Italy, Apennines), A. flexicaule (France, Western Alps), A. pirinicum (Bulgaria, Pirin Mts), and A. spruneri (Greece, Mt Smolikas). The present study brings the thorough morphometric analysis of the segregated taxa. We found minor morphological differences between them. Whereas A. pirinicum can be clearly distinguished, the other taxa are recognizable only at the level of population means of investigated characters. The morphological similarity of these distantly related species is obviously the result of adaptation to similar high‐alpine scree habitats. It is not clear, however, whether this adaptation is environmentally controlled or whether it is also genetically fixed and whether it reflects parallel evolution towards similar morphotypes. The observed morphological patterns and their assumed correlation with environmental factors are discussed using examples from other Alyssum taxa. Three different ploidy levels have been reported for the species under study. In the present article, we examine variation in relative nuclear genome size. The Alpine and Pyrenean species have larger relative monoploid genome sizes than the Apennine and Balkan ones, probably reflecting the evolutionary history of the group. A nomenclatural account of the study species is presented, and lectotypes of A. cuneifolium and of two other names are selected.     

Micro‐morphological characteristics of seed and ovary trichomes of Iranian Glaucium Mill. species (Papaveraceae) and their taxonomic importance

Seeds and trichomes of 15 taxa of the genus Glaucium distributed in Iran were examined by light microscopy (LM) and scanning electron microscopy (SEM). The seeds are slightly shiny to rather shiny, however, strongly shiny seeds were also observed in some species (G. corniculatum and G. grandiflorum). The basic shape of the seeds is semicircular to reniform, but reniform and elongated reniform seeds were found in G. oxylobum and G. elegans, respectively. The sculpturing of the testa surface are verrucate–rugulate (most frequent type), verrucate–granulate, verrucate–perforate, verrucate–lineolate, rugulate–granulate, rugulate and ocellate. Elongated polygonal epidermal cells dominate in most taxa investigated, but there are polygonal and irregular polygonal cells in a few taxa. The investigated species have two types of trichomes on the ovary: tuberculate and non‐tuberculate. The non‐tuberculate trichomes are of two subtypes: pilose and squamiform. Based on the number of cells, the squamiform trichomes were further divided into two groups (uniseriate and multiseriate) differing in shape, size and cellular structure. The results show that the micro‐morphological characteristics of seed and ovary trichomes provide useful and significant information for separation of species and taxa within species and a diagnostic key to the taxa is provided.     

Patterns of allozyme variation in Nordic Pilosella

Patterns of allozyme variation have been investigated in 101 clones representing most of the morphological variation (and taxa) found in the amphi-apomictic genus Pilosella Hill. in the Nordic countries. Levels of variation were very high, both within narrowly defined taxa and within the whole data-set, but the correlation between a morphology-based taxonomy and patterns of allozyme variation was very weak. Almost all enzyme bands were found in more than one taxon and the differences between broadly defined species in terms of the frequency of bands were generally small. Cluster analyses did not perfectly recover any taxon but there were some tendencies for samples of some broadly defined taxa (e.g. P. officinarum s. l., and P. cymosa ssp. cymosa) to cluster together. It is concluded that there is considerable gene-flow among virtually all Nordic morpho-types of Pilosella and this is considered an argument for applying a very broad species concept in this genus.Financial support for the present study has been recieved from Stiftelsen Harald E. Johanssons minnesfond at the Royal Swedish Academy of Sciences.     

Size and shape dimorphism in great ape mandibles and implications for fossil species recognition

Sexual dimorphism is an important source of morphological variation, and species differences in dimorphism may be reflected in magnitude, pattern, or both. While the extant great apes are commonly used as a reference sample for distinguishing between sexual dimorphism and intertaxic variation in the fossil record, few studies have evaluated mandibular dimorphism in these taxa. In this study, percentage, degree, and pattern of mandibular dimorphism are evaluated in Pongo, Gorilla, and Pan. Mandibular dimorphism patterns are explored to determine the extent to which such patterns accurately track great ape phylogeny. Pattern stability is assessed to determine whether there are stable patterns of mandibular size and shape dimorphism that may be usefully applied to hominoid or hominid fossil species recognition studies. Finally, the established patterns of dimorphism are used to address recent debates surrounding great ape taxonomy. Results demonstrate that mandibular dimorphism is universally expressed in size, but only Pongo and Gorilla exhibit shape dimorphism. Pattern similarity tends to be greater between subspecies of the same species than between higher-order taxa, suggesting that within the great apes, there is a relationship between dimorphism pattern and phylogeny. However, this relationship is not exact, given that dimorphism patterns are weakly correlated between some closely related taxa, while great ape subspecies may be highly correlated with taxa belonging to other species or genera. Furthermore, dimorphism patterns are not significantly correlated between great ape genera, even between Gorilla and Pan. Dimorphism patterns are more stable in Gorilla and Pongo as compared to Pan, but there is little pattern stability between species or genera. Importantly, few variables differ significantly between taxa that simultaneously show consistently relatively low levels of dimorphism and low levels of variation within taxa. Combined, these findings indicate that mandibular dimorphism patterns can and do vary considerably, even among closely related species, and suggest that it would be difficult to employ great ape mandibular dimorphism patterns for purposes of distinguishing between intra- and interspecies variation in fossil samples. Finally, the degree of pattern similarity in mandibular dimorphism is lower than previously observed by others for craniofacial dimorphism. Thus, the possibility cannot be ruled out that patterns of craniofacial dimorphism in great apes may be associated with a stronger phylogenetic signal than are patterns of mandibular dimorphism.     

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.     

Assessing the Patterns of Evolution in Anuran Vocal Sexual Signals

Many animal species have evolved signalling traits to mediate various intra-specific interactions. Signals are particularly important for inter-sexual selection, where females use male signalling traits to select mates. Female preferences are therefore a major selective force in the evolution of these male signals, and these preferences can facilitate rapid changes in these traits in an evolutionary timeframe. This introduction of high levels of variation in inter-sexual signals may overshadow any phylogenetic patterns present. Such shadowing effects, however, should be dependant on the characteristics of traits (e.g. morphological, physiological and behavioural). Using male advertisement calls from 72 species of anuran amphibians, we tested the levels of phylogenetic signal present for a variety of call features in relation to trait types, and for calls as whole units using phylogenetic principal components analysis. We found that most call features displayed some level of phylogenetic autocorrelation (or signal), with traits that are dependent on morphology having much stronger phylogenetic signals than those based on behaviour. In addition, when calls were analysed as whole units, closely related species were found to be similar to each other, indicating that phylogenetic patterns had not been cancelled out by selection via female preferences. We suggest that signal functions, such as indicating male quality (e.g. mediated by body size) to potential mates, may place constraints on the amount of variation that can be introduced by female preferences. More research, particularly studies on other taxa, will be required to elucidate whether the patterns found in anurans are general across the animal kingdom.     

Diet-morphology correlations in the radiation of South American geophagine cichlids (Perciformes: Cichlidae: Cichlinae)

Genera within the South American cichlid tribe Geophagini display specialized feeding and reproductive strategies, with some taxa specialized for both substrate-sifting and mouth brooding. Several lineages within the clade also possess an epibranchial lobe (EBL), a unique pharyngeal structure that has been proposed to have a function in feeding and/or mouth brooding. A recently published genus-level phylogeny of Neotropical cichlids was used as the evolutionary framework for investigating the evolution of morphological features presumably correlated with diet and mouth brooding in the tribe Geophagini. We tested for possible associations between the geophagine epibranchial lobe and benthic feeding and mouth brooding. We also addressed whether the EBL may be associated with unique patterns of diversification in certain geophagine clades. Tests of binary character correlations revealed the EBL was significantly associated with mouth brooding. We also tested for a relationship between diet and morphology. We analyzed stomach contents and morphometric variation among 21 species, with data for two additional species obtained from the literature. Principal Components Analysis revealed axes of morphological variation significantly correlated with piscivory and benthivory, and both morphology and diet were significantly associated with phylogeny. These results suggest that the EBL could be an adaptation for either feeding or mouth brooding. The EBL, however, was not associated with species richness or accelerated rates of phyletic diversification.     

DISPARITY: MORPHOLOGICAL PATTERN AND DEVELOPMENTAL CONTEXT

Abstract:  The distribution of organic forms is clumpy at any scale from populations to the highest taxonomic categories, and whether considered within clades or within ecosystems. The fossil record provides little support for expectations that the morphological gaps between species or groups of species have increased through time as it might if the gaps were created by extinction of a more homogeneous distribution of morphologies. As the quantitative assessments of morphology have replaced counts of higher taxa as a metric of morphological disparity, numerous studies have demonstrated the rapid construction of morphospace early in evolutionary radiations, and have emphasized the difference between taxonomic measures of morphological diversity and quantitative assessments of disparity. Other studies have evaluated changing patterns of disparity across mass extinctions, ecomorphological patterns and the patterns of convergence within ecological communities, while the development of theoretical morphology has greatly aided efforts to understand why some forms do not occur. A parallel, and until recently, largely separate research effort in evolutionary developmental biology has established that the developmental toolkit underlying the remarkable breadth of metazoan form is largely identical among Bilateria, and many components are shared among all metazoa. Underlying this concern with disparity is a question about temporal variation in the production of morphological innovations, a debate over the relative significance of the generation of new morphologies vs. differential probabilities of their successful introduction, and the relative importance of constraint, convergence and contingency in the evolution of form.     

Mandibular gland secretions in alloxystine wasps (Hymenoptera,Cynipoidea, Charipidae): do ecological or phylogenetical constraints influence occurrence or composition?

Mandibular gland secretions were investigated in 28 taxa of alloxystine wasps (17 described and 11 potential, hitherto undescribed species) using GC-MS. In two selected species (Alloxysta pleuralis, A. victrix), the chemical patterns of the mandibular gland secretions remained constant and were not qualitatively influenced by different host systems; however, there was a considerable variation in quantity. A cluster analysis revealed the chemical composition to be independent of two different modes of resource utilisation which are characterized by the presence or absence of ants in the respective host system. These results support the idea of the prevalence of historical (phylogenetic) rather than actual ecological constraints. However, the chemotaxonomic applicability of such data on the subfamily level was limited, although a species status could be confirmed in some questionable taxa. By integrating morphological and biological characters, potential relationships of selected species and a hypothetical phylogenetic frame tree are proposed.     

Testa and seed appendage morphology in Aeschynanthus (Gesneriaceae): phytogeographical patterns and taxonomic implications

The current sectional classification of the genus Aeschynanthus Jack, essentially based on seed morphology, presents some problems of species placement. A comparative SEM survey of seed and seed appendages was undertaken in order to assess the value of this classification. Seeds of 99 taxa (that is about two‐thirds of the estimated total) were examined and found to fall into two types, A and B. Type A has spiral testa cell orientation, papillae formed from a single cell and short smooth appendages. Type B is recognized by the straight orientation of the testa cells, combined with the presence of papillae formed from the raised ends of two adjacent cells on the long hair‐like appendages and usually on the testa. Only six of the investigated species did not fall into either category. Three have straight testa cell orientation combined with single‐cell papillae and short smooth appendages; the papillae and appendage characters place them in type A. Three have spiral testa cell orientation and short smooth appendages but the testa cells have slightly raised ends; these are also placed in Type A. The three subtypes in Type A are equivalent to the sections Haplotrichium s.s., Microtrichium and Aeschynanthus, but the divisions are less clear than those within Type B. However, other morphological characters support sectional separation. Type B subdivides into three: two subtypes equivalent to sections Polytrichium and Diplotrichium, and a third encompassing section Xanthanthos together with part of the current sect. Haplotrichium, and here referred to as sect. X. There is sufficient morphological correlation with seed type to make the sectional position of many species clear without recourse to seed, particularly in sects Polytrichium, Diplotrichium, Haplotrichium S.S. and Aeschynanthus. There is strong correlation between seed type and geographical distribution. Sects. Microtrichium and Aeschynanthus, with Type A seed, are essentially Malesian. Groups with Type B seed are largely confined to mainland south and south‐east Asia, except for sect. Polytrichium which is more widespread, possibly due to the greater effectiveness of a coma of hairs in wind dispersal. It is suggested that Type A seed, probably sect. Microtrichium, is the least determined and Type B sect. Polytrichium the most derived seed type. Based on these findings a revised key to the sections is provided.     

Taxonomic value of seed characters in the Erica tetralix L. group (Ericaceae)

A study of the seed morphology of Erica tetralix and the related species E. mackaiana and E. andevalensis was performed to evaluate the taxonomic importance of seed characters. Size and shape of seeds and surface cells were studied, and area, perimeter, main axis lengths, elongation and sinuosity were measured. Levels of intraspecific variation were considered by means of nested analysis of variance. The study of the intraspecific variation of the characters shows significant differences for lower levels, higher than variations between species for most characters. Levels of variation must be based upon the numerical analysis of seed characters for a correct approach to determining systematic value. E. mackaiana and E. andevalensis group together versus E. tetralix for sinuosity of testa cells. This is a good taxonomic character because it shows significant differences between species and no significant differences between populations. E. tetralix has strongly undulated testa cells while E. mackaiana and E. andevalensis have slightly undulated or curved testa cells. These two species can be discriminated by size of the seed, with seeds of 0.37-0.38 mm for E. andevalensis and 0.4 - 0.45 mm for E. mackaiana. Similarities are found in seed morphology for E. mackaiana and E. andevalensis, but we nevertheless consider the species status for the three taxa as the most natural taxonomic assignment. A key to the species is provided.     

Evolutionary history of Trachylepis skinks in the Seychelles islands: introgressive hybridization,morphological evolution and geographic structure

The Seychelles is a remarkably interesting archipelago for evolutionary studies, but only recently have molecular markers been used to explore its biogeographic patterns. Here we used morphological and molecular data to examine diversity and phylogenetic relationships of two endemic skink sister‐species from this archipelago: Trachylepis sechellensis and Trachylepis wrightii. Mitochondrial DNA genealogy rendered a monophyletic T. wrightii nested within a paraphyletic T. sechellensis, whereas nuclear DNA sequences from five unlinked markers reflected the accepted taxonomy. Hybridization and massive mtDNA introgression leading to the complete replacement of the native mtDNA lineage of T. sechellensis in some of the islands were invoked to explain this result, and morphological variation also seemed to reflect this pattern of reticulation. A Mio‐Pliocene divergence between both species is suggested. Multilocus molecular data were used to uncover biogeographic patterns within the archipelago, which reflected shared patterns with other co‐distributed lizard taxa; specifically a north–south marked structure, a close relationship between populations from Fregate and the southern islands, and a detectable isolation within the southern group, between Mahé, and Silhouette and North Islands. Gene flow from these latter islands towards the northern group was also suggested. These results add to the growing body of evidence of the influence of geographic distance and sea‐level oscillations in shaping the genetic structure of Seychellois taxa and of the existence of common biogeographic patterns across the archipelago.     

Genetic and morphological diversity of Trisetacus species (Eriophyoidea: Phytoptidae) associated with coniferous trees in Poland: phylogeny,barcoding, host and habitat specialization

Eriophyoid species belonging to the genus Trisetacus are economically important as pests of conifers. A narrow host specialization to conifers and some unique morphological characteristics have made these mites interesting subjects for scientific inquiry. In this study, we assessed morphological and genetic variation of seven Trisetacus species originating from six coniferous hosts in Poland by morphometric analysis and molecular sequencing of the mitochondrial cytochrome oxidase subunit I gene and the nuclear D2 region of 28S rDNA. The results confirmed the monophyly of the genus Trisetacus as well as the monophyly of five of the seven species studied. Both DNA sequences were effective in discriminating between six of the seven species tested. Host-dependent genetic and morphological variation in T. silvestris and T. relocatus, and habitat-dependent genetic and morphological variation in T. juniperinus were detected, suggesting the existence of races or even distinct species within these Trisetacus taxa. This is the first molecular phylogenetic analysis of the Trisetacus species. The findings presented here will stimulate further investigations on the evolutionary relationships of Trisetacus as well as the entire Phytoptidae family.     

Skeletal indicators of ecological specialization in pika (Mammalia,Ochotonidae)

Pika species generally fall into two ecotypes, meadow‐dwelling (burrowing) or talus‐dwelling, a classification that distinguishes a suite of different ecological, behavioral, and life history traits. Despite these differences, little morphological variation has previously been documented to distinguish among ecotypes. The aim of this study was to test whether postcranial features related to burrowing are present in meadow‐dwelling species and whether talus‐dwelling species exhibit postcranial modifications related to frequent leaping between rocks. To test this, the scapula, humerus, ulna, radius, innominate, femur, tibia, and calcaneus of 15 species were studied and measured. Twenty‐three measurements were taken on 199 skeletons, and 19 indices were constructed from these measurements. Indices were compared between the two ecotypes using Student's t‐test. Comparisons among ecotypes, species, and subgenera were made using one‐way ANOVA with the Tukey honest significant difference post hoc test. Multivariate results were generated using principal components analyses. Thirteen forelimb and hind limb indices proved significant in distinguishing the meadow‐dwelling, talus‐dwelling, and intermediate forms. A number of these indices are associated with burrowing or leaping in other mammals, providing some support for the hypothesis that postcranial modifications in pika are related to locomotor differences. This evidence of morphological responses to ecological specialization will be useful for reconstructing the paleobiology of extinct taxa, assessing the behavioral variability of extant species, and improving our understanding of the evolutionary history of pikas. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Systematic significance of seed morphology in veronica (plantaginaceae): a phylogenetic perspective

BACKGROUND AND AIMS: A new infrageneric rearrangement for Veronica has been proposed based on the most recent evidence from DNA sequence data, morphological evidence, and biogeographical considerations. Looking for morphological synapomorphies for each monophyletic subgenus has been problematic, due to difficulties arising from widespread homoplasy (mainly parallel evolution). In an attempt to overcome these difficulties, previously underexplored morphological characters are starting to be studied in more depth. METHODS: A molecular phylogenetic hypothesis was used based on sequences of ITS (nuclear ribosomal DNA) and plastid trnL-F regions, as a framework to test the use of seed coat ultrastructure (studied under scanning electron microscope) in the systematics of the genus. A sample of 132 taxa representing ten of the 13 subgenera in Veronica, excluding the species of the southern hemisphere Hebe complex and the exclusively North American subgenus Synthyris, was studied. KEY RESULTS AND CONCLUSIONS: The results demonstrate that, in many cases, the ultrastucture of the testa can be employed to assess relationships of taxa within the genus, and the character provides additional support for molecular trees. Further characters relevant for the classification of Veronica, i.e. base chromosome number, iridoid chemical data, life cycle, inflorescence position, have been taken into consideration in a discussion where an attempt is made to highlight the best traits to characterize each subgenus investigated.     

Delineating closely related species with DNA barcodes for routine biological monitoring

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Decoupling of molecular and morphological evolution in deep lineages of a meiobenthic harpacticoid copepod

Molecular and biochemical genetic analyses have revealed that many marine invertebrate taxa, including some well-studied and presumably cosmopolitan species, are actually complexes of sibling species. When morphological differences are slight and estimated divergence times are old, data suggest either unusually high rates of sequence evolution or long-term morphological stasis. Here, five gene regions (mitochondrial cytochrome oxidase subunit I and large-subunit ribosomal 16S rDNA and nuclear ITS1, 5.8S rDNA, and ITS2) were analyzed in four geographic samples of the meiobenthic harpacticoid copepod Cletocamptus deitersi. Molecular sequences revealed four extremely differentiated molecular lineages with unalignable nuclear intergenic spacers and mitochondrial uncorrected divergences reaching 25% (cytochrome oxidase) and 36% (16S rDNA). These levels of divergence are greater than those reported previously for congeneric species in diverse invertebrate taxa, including crustaceans. The nominally intraspecific divergence matches or exceeds the corresponding divergence from a known congener (Cletocamptus helobius). A molecular clock applied to the cytochrome oxidase subunit I data suggests that these lineages split in the Miocene, consistent with the fossil record of a North American Cletocamptus from the same period. Morphological differences among the major lineages are subtle but congruent with the patterns of genetic differentiation. Our conclusion, based on concordant patterns of variation in two mitochondrial and three nuclear gene regions, as well as morphological observations, is that C. deitersi in North America is composed of at least four separate species by the genealogical concordance, phylogenetic, and morphological-species criteria. Alternative explanations for the deep phylogenetic nodes and apparent morphological stasis, including high rates of sequence evolution, balancing selection, and genetic signatures of historical events, are considered unlikely.