On potentials and relevance of the use of copulatory structures in anostracan taxonomy

This paper reviews and extends the knowledge of anostracan penile morphology and its taxonomic significance. Since Linder‘s pioneering classification(based partly on male genital characters), the morphology of both basal and distal penile parts has been applied repeatedly to reorganize or establish higher anostracan taxa, or to extend their diagnoses. In general, the configuration of both basal and distal penile structures follows a constant pattern in well-defined genera. Size and shape of processes and spinulation on the basal (in Linderiella and Chirocephalidae) and the distal parts (in Parartemia), however, may show inter-specific differences. Variability in penile structures in the current genus Brachinella indicates the need for a taxonomic revision of this genus. Similar intra-generic penis morphology may contribute to hybridizability of congeners and validate Dubois‘genus concept for use in anostracans. This revised version was published online in August 2006 with corrections to the Cover Date.     

The variation of the centipede Scolopendra amazonica in Africa

African centipedes labelled Scolopendra amazonica (Buecherl) and S. morsitans L. in the collection of the British Museum (Natural History) have been examined. All are regarded, at present, as S. amazonica but in many respects S. amazonica in Africa shows greater variation than do S. amazonica and S. morsitans in India. North African populations are very distinct.     

A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae)

MIDDLETON, D. J., A systematic survey of leaf and stem anatomical characters in the genus Gaultheria and related genera (Ericaceae). Leaf and stem anatomical characters of 123 species within the Gaultheria group of genera of the tribe Andromedeae are surveyed. A number of characters show considerable variation within a species which limits their taxonomic use. Other characters such as marginal sclerenchyma and, to a lesser extent, the hypodermis, the presence of adaxial stomata, the presence of free fibres and the pith type were of more taxonomic interest. These characters are discussed in relation to the generic and infrageneric classification of the group.     

Revision of Maranta subgen. Maranta (Marantaceae)

Maranta subgen. Maranta includes species related to M. arundinacea L. It is characterized by aerial shoots with a strong monopodial tendency, absence of root tubers, simple inflorescences or few-branched, often diffuse synflorescences, florescences with few, herbaceous spathes, and comparatively large, distinctly pedicellate flowers. There are occasional exceptions to all these characters. Most species are partial selfers, only two or three being allogamous. Sixteen species are recognized, eight of which are new: M. linearis, M. sobolifera, M. lindmanii, M. zingiberina, M. incrassata, M. rupicola, M. amazonica , and M. tuberculata. The new species are described, and all species are defined and discussed, data being given about distribution and habitat. Growth habit and rhizome structure are essential taxonomic characters, but leaf shape and indument distribution are more useful characters for routine identification. Of the species referred to subgen. Automaranta by Schumann in Das Pflanzenreich, three are excluded: M. cordata, M. pohliana , and M. foliosa.     

Quantitative handling of characters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.)

This paper is based on the numerous characters used in a biometric study of the intraspecific variation of Natrix natrix (L.). These characters are drawn from the internal morphology, scalation, colour pattern, body proportions, dentition and dermal sense organs. The criteria involved in selecting these characters for taxonomic work are considered.
The alternative methods for quantifying the above features are discussed and so are the theoretical problems involved in coding the continuous characters.
The value of the characters in studying the geographic variation, ontogenetic variation and sexual dimorphism in N. natrix is indicated.
The use of a wider range of characters than just the scalation and colour pattern is urged when diagnosing the lower taxonomic levels of ophidians. The neglected internal morphology may well be able to supply characters to this end.     

How many species of Diacyclops? New taxonomic characters and species richness in a freshwater cyclopid genus (Copepoda,Cyclopoida)

The genus Diacyclops Kiefer, 1927 emend. Morton (1985) includes more than 100 species, widely distributed in all kinds of freshwater environments, and is the richest genus in the family Cyclopidae. Traditionally, Diacyclops species were defined according to differences in a few morphological characters; some characters (number of antennulary segments, segmentation pattern of swimming legs) are useful only in the separation of species groups, others (length of caudal rami and caudal setae) are highly variable even within the same population. During the study of the D. languidoides-group, minute morphological characters were used to differentiate between species (spinulation pattern and setation of antennary basis, setation and aesthetasc shape of male antennule, setation of mandibulary palp and maxilliped, shape of leg 4 basis) which allow to identify several valid species up to now concealed under the name `Diacyclops languidoides' (Lilljeborg, 1901). The coexistence of up to six congeneric species in the same sampling area: (a) supports the validity of the proposed taxonomic characters, (b) demonstrates that species richness may be highly underestimated in freshwater cyclopoid assemblages in absence of good taxonomic practice; and (c) requires an ecological explanation of species coexistence. The role of morphologically based taxonomy in order to solve general problems of distributional ecology and theoretical biology is explored.     

Normal range and variation in the ontogeny of the Late Paleozoic ammonoid family Marathonitidae

The ontogeny and phylogeny of members of the Late Paleozoic family Marathonitidae are ana-lyzed. The ontogeny of some taxa shows variation in major morphological characters, including the shell shape and sutural outline. The range of variation of the characters examined is defined, and deviations from the normal range are considered to be instrumental in the derivation of new groups of various taxonomic ranks.     

泡沙参复合体（桔梗科）的物种生物学研究：Ⅰ表型的可塑性

表型可塑性是生物变异中由环境引起的一种变异,是植物适应的一种重要方式。对沙参属这样一个形态上复杂多变、分类上很难处理的类群,研究其表型可塑性不仅为探讨性状变异、判断其系统学意义及选择可靠的分类性状提供了有益的资料,而且有助于揭示沙参属植物变异、适应和进化的机制。本文从泡沙参复合体中选择了6个居群,利用播种和移栽试验,通过对不同个体和居群在一致条件下的表现及野外和移栽后的对比,对根、茎、叶、花和果等形态性状的表型可塑性进行了初步的观测分析。结果表明,一些叶片、花部和果实性状具有较大的发育可塑性,尤其是叶形、花萼裂片不仅发育变化大,而且随发育过程定向变化。环境可塑性较大的性状主要是根、茎、花序分枝等性状,而叶片、花部、果实和种子性状的环境饰变能力都较小。最后,对泡沙参复合体形态性状的变异从发育可塑性和环境可塑性的角度进行了讨论。     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae). Ⅰ. Phenotypic Plasticity

Phenotypic plasticity is the environmental modification of genotypic expression and an important means by which individual plants respond to environmental heterogeneity. The study of phenotypic plasticity in the genus Adenophora, which is very complicated taxo nomically because of great morphological variation, proves to be helpful in both investigating the phenotypic variation so as to evaluate potential taxonomic value of their characters and providing important sources of information on the variation, adaptation and evolution of the genus. Twenty-three populations representing all the six species in Adenophora potaninii complex were transplanted into the garden. Of them six populations were selected for study ing their performance in the field and in the garden, in addition to cultivation experiment under different treatments. The results show that there exists considerable developmental plasticity in some leaf, floral and capsule characters. In particular, the leaf shape and length of calyx lobe display significant developmental variation with the maximum being three times as great as the minimum, which is noteworthy because they were previously considered as diagnostic. The characters of root, caudex, stem and inflorescence are found to be very plastic, especially the root diameter, the number of stems, stem height and inflorescence length with great environmental plasticity. In addition, the populations from different habi tats show distinct amounts of plasticity. On the contrary, the characters of leaf, floral, cap sule and seed are less influenced by environments. It seems that the considerable variation in the characters of leaf is attributed mainly to genetic differences. Finally, the phenotypic plasticity of morphological characters of A. potaninii complex and its taxonomic significanceis discussed.     

GENETIC AND ENVIRONMENTAL DETERMINATION OF LEAF EPIDERMAL ANATOMY IN PUCCINELLIA (POACEAE)

Phenotypic plasticity of anatomical leaf epidermal characters was assessed in the halophytic grass genus Puccinellia, to identify those of potential taxonomic utility. Characters with the greatest taxonomic potential are those that exhibit genetic variation and low plasticity. For the analysis, field-collected clones were divided and grown under a series of moisture and salinity regimes. Genetic variation and plasticity in response to environmental variation were assessed for 39 anatomical characters by analysis of among-clone and among-treatment variation, respectively. Both genetic variation and plasticity are widespread among the characters, and both are detected in greater frequency among continuously variable characters than among discretestate characters. Among continuously variable traits, average cell dimensions exhibit more plasticity than do maximum cell dimensions or ratios that reflect shapes and relative sizes. Among discrete-state characters, plasticity in the occurrence of papillae is found in nonstomatal intercostal cellular ranks, but not in costal or stomatal intercostal ranks. In Puccinellia, anatomical characters are, in general, neither more nor less plastic than those of macromorphology.     

Morphological and molecular variation in tiger beetles of the Cicindela hybrida complex: is an ‘integrative taxonomy’ possible?

Current taxon assignments at the species level are frequently discordant with DNA-based analyses. Recent studies on tiger beetles in the Cicindela hybrida complex identified discordance between mtDNA patterns and the entities currently defined by the taxonomic literature. To test the accuracy of morphologically delimited groups, five named taxa (species) from 24 representative sampling sites across Europe were scored for 41 external morphological characters. Three of the named taxa were 'diagnosable', that is, defined by between one and three characters unique to each group. Newly sequenced ITS1 and existing mitochondrial cox1 markers established 20 and 22 different haplotypes, respectively, but only cox1 produced (four) diagnosable units. Phylogenetic analysis and statistical parsimony networks showed poor congruence of character variation with the taxonomic entities (and each other). Variation in morphological characters was therefore tested directly for association with DNA-based nesting groups at various hierarchical levels using permutational contingency analysis. Significant statistical associations of 11 (of 13 variable) morphological characters were observed with nesting groups from ITS1 and mitochondrial DNA markers, predominantly at the 4-step level. The analysis demonstrates the need for formal tests of congruence with morphological variation at the level of individual characters, a step that is omitted from recent studies of 'integrative taxonomy'. In addition, statistical correlation of particular morphological characters with DNA-based nesting groups can identify the lowest hierarchical level at which various character sets show congruence, as a means to define evolutionarily separated entities supported by diverse data sources.     

Morphological characterization of new populations of the copepod Eurytemora velox (Lilljeborg, 1853) (Calanoida,Temoridae) found in Austria and Hungary

The calanoid copepod Eurytemora velox (Lilljeborg, 1853) was found in the backwater system of the Danube River upstream of Vienna (km 1948) in 1994. Since that time, the species has invaded the side arms of the river between kms 1897 and 1988, as confirmed by findings from 10 different localities. In Hungary, the species has been known since 1992. In the present article, we report 15 new sites in the westernmost part of this country.A brief diagnosis of the morphology of male and female of the Austrian specimens is presented, including detailed illustrations of the structures used in this paper for taxonomic analysis. Females show morphological variability mainly on the structure of the fifth leg pair. Right and left legs frequently have a different spinulation pattern. Males show less morphological variability, particularly in the structure of the basis and the distal section of exopod 2 of the fifth left leg. Morphological variation within E. veloxwas determined by comparing specimens from Austria, Hungary and Rumania. The geographic distribution of the species in the Palearctic region, Austria and Hungary is presented.     

Relationship between song characters and morphology in New World pigeons

We studied the pattern of variation in song characters among 16 New World pigeon species belonging to different taxonomic groups defined by morphological characters. Structural, temporal and frequency characters of the song were analysed. Principal components analyses showed that species belonging to the same taxonomic group were also grouped together by their song characters. In addition, individuals were correctly assigned into taxonomic groups by discriminant function analyses in more than 87.8% of cases. These analyses also showed that more than 87.5% of the individuals could be correctly classified by species when all song characters were included. Correct classification of individuals by species and taxonomic groups dropped when character types were analysed separately, thus showing that structural, as well as temporal and frequency characters are fundamental to define species- and group-specific identities of New World pigeon's songs. Correspondence between patterns of vocal and morphological variation found in this study can be a consequence of evolutionary changes in morphology affecting song production, as for example body size changes that constrain the syrinx to produce certain acoustic frequencies.     

Morphology, anatomy and histochemistry of Salicornioideae (Chenopodiaceae) fruits and seeds

BACKGROUND AND AIMS: The subfamily Salicornioideae (Chenopodiaceae) are a taxonomically difficult group largely due to the lack of diagnostic characters available to delineate tribal- and generic-level boundaries; a consequence of their reduced floral and vegetative features. This study examined the variation in fruits and seeds across both tribes of the Salicornioideae to assess if characters support traditional taxonomic sections. METHODS: Light microscopy, environmental scanning electron microscopy and anatomical ultra-thin sectioning were employed to examine variation in fruits and seeds. Sixty-eight representatives across 14 of the 15 genera currently recognized within the tribes Halopeplideae and Salicornieae were examined to determine whether characters support current taxonomic groups. KEY RESULTS: Characters such as seed coat structure, embryo shape, seed orientation, the forms of seed storage proteins and carbohydrates show variation within the Salicornioideae and may be phylogenetically useful. The campylotropous ovule typical of the Chenopodiaceae generally results in a curved embryo; however, many Halosarcia and Sclerostegia species have straight embryos and in Salicornia and Sarcocornia the large peripheral embryo appears bent rather than curved. Seed coat ornamentation of Microcnemum and Arthrocnemum is distinct from other Salicornioideae as the elongated epidermal cells of the exotesta have convex walls. Histochemical stains of anatomical sections of cotyledon cells showed protein bodies were variable in shape, and starch grains were present in some species, namely Salicornia bigelovii, S. europaea and Allenrolfea occidentalis. CONCLUSIONS: While fruits and seeds were found to be variable within the subfamily, no synapomorphic characters support the tribe Halopeplideae as these genera have crustaceous seed coats, curved embryos and abundant perisperm; features characteristic of many of the tribe Salicornieae. The endemic Australian genera are closely related and few seed and fruit characters are diagnostic at the generic level. Nineteen characters identified as being potentially informative will be included in future phylogenetic analyses of the subfamily.     

REVISED CLASSIFICATION OF THE GENUS PACHYMENIA BASED ON MORPHOLOGICAL AND MOLECULAR CHARACTERS

Taxonomic discrimination in the genus Pachymenia (Rhodophyta) in New Zealand is based primarily on phenotypic characters of the thallus. The taxonomic problems raised by this classification method are due to highly variable thallus characters such as blade thickness, blade width, degree of thallus branching, and variation in anatomical characters. Delineation of species is further complicated by a lack of adequate knowledge about the responses of phenotype to environmental variation. There are currently three species recognized in this genus that are endemic to New Zealand: a prostrate species P. crassa , and two erect species, P. laciniata and P. lusoria. In this study, two approaches are used to investigate the current delineation of these species. Morphological and anatomical characters of field collected material and herbarium specimens from throughout the species' distributional ranges were quantified. Multivariate analyses were used to identify discrete phenotypic groups. Species relationships were further analyzed by quantifying the variation found within the internal transcribed spacer region (ITS) of the nuclear ribosomal DNA. The results obtained from both approaches will be discussed with regards to possible re-classification of species relationships within this genus. We suggest that the two erect species should be merged, and the currently recognized P. lusoria should be separated into at least two taxonomic groups.     

Patterns of morphological and geographic variation in trophic bill morphs of the African finch Pyrenestes

Intra- and inter-population variability was studied in three species of tropical African estrildid finches comprising the genus Pyrenestes. Eleven characters were measured on P. ostrinus captured on a study area in Cameroon. Most of these same characters were also measured on museum specimens of this species and P. saguineus and P. minor. Data were analysed using univariate and multivariate statistical methods in order to characterize variation within and between populations and species.
Variation in all three species of Pyrenestes is greatest in bill size, resulting from an exceptional non-sex-linked polymorphism. Bill size differences between morphs are as high as between congeneric species, with extremely large coefficients of variation, while other body characters show comparatively little variation. Sexual dimorphisms and differences in size due to age occur, but contribute little to overall size variation. Distributions of bill characters in each age and sex class are bimodal or greatly skewed, and in some geographical regions tend to be trimodal. Distributions of other body characters tend not to be significantly different from normal. Bill morphs differ in both shape and size and may be separated using principal component analysis. Static allometries of bill morphs differ significantly: relative to body size, bill size increases more rapidly in the large morph. Bill size and shape also vary geographically. The three species differ in mean size but show much overlap. Bill size is negatively correlated with total annual rainfall. In regions characterized by ecotonal transition zones between forest and savanna, tentative evidence suggests that a third, yet larger bill mode occurs. This third mode apparently results from the presence of a distinct larger species of hard-seeded sedge found only in these regions. The taxonomic implications of the polymorphism are discussed.     

Investigation of variation in phenotype and DNA content between single-conidium isolates of single Penicillium strains

Variation in phenotypic properties was examined in three strains of closely related fasciculate species of Penicillium using 114 morphological, physiological, and biochemical characters. Thirty-six of these characters showed variation within single-conidium isolates of the same strain. Conidial sizes and nuclear DNA contents were compared using flow microfluorimetry; these results suggested that significant differences in conidial DNA content are associated with phenotypic variation. The taxonomic significance of the results is discussed.     

The limited value of traditional morphometric features in stromatoporoid taxonomy

The morphological variation of stromatoporoids, which are solitary organisms, is partitioned into its presumably genetic and environmental components. Potentially heritable, environmentally mediated and residual components of morphological variability were estimated in a test set containing Devonian stromatoporoids of the genus Gerronostromaria from southern Poland using analysis of variance. The taxonomic importance of traditional morphometric features is limited, because they are dominated by the intra‐skeletal component of variance. Conventional metrics were therefore replaced by stereological and textural quantities. Both stereological and textural features are dominated by the inter‐skeletal and inter‐locality components of variation and thus may be valuable in taxonomic and environmental studies of stromatoporoids. Statistical analyses of these characters (principal component analysis and cluster analysis) were performed. Of 13 characters considered most useful in taxonomic studies, only five have been used previously in conventional species definitions.