Variation in body size and sexual dimorphism across geographical and environmental space in the frogs Limnodynastes tasmaniensis and L. peronii

This study aimed to identify potential factors responsible for geographically structured morphological variation within the widespread Australian frogs Limnodynastes tasmaniensis Günther and L. peronii Duméril & Bibron. There was support for James's rule, and both latitude and present climate explained large amounts of the variation in body size and shape (particularly in L. peronii ). There was also some support for the influence of several biogeographical barriers. Finally, both species were sexually dimorphic for body size and the degree of sexual size dimorphism (SSD) varied geographically. Climate was an important explanation for SSD variation in L. peronii , while latitude was most important for L. tasmaniensis . Geographical variations in sexual selection via male–male physical competition and climate-related resources are suggested as potential explanations for SSD variation in L. peronii .  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 39–56.     

Divergent and reticulate processes in evolution of Ethiopian Lophuromys flavopunctatus species complex: evidence from mitochondrial and nuclear DNA differentiation patterns

Molecular study of mitochondrial and nuclear genes and cytogenetic analysis were performed to examine possible patterns of speciation in the diverse Lophuromys flavopunctatus species complex of Ethiopia. Phylogenetic analysis of mtDNA data resulted in an unresolved bush of ten deeply diverged haplotype groups corresponding to potential species either well supported by various types of character or 'cryptic'. The cytogenetic analysis showed representatives of five of these mtDNA lineages to share an identical karyotype (2 n  = 70, NFa = 84), that has not been found previously in Ethiopia. One of them, L.  cf.  sikapusi , being a member of the L. flavopunctatus species complex, demonstrates remarkable morphological similarity to representatives of another species complex, L. sikapusi s.l ., which might be considered as a result of convergent evolution in analogous environments. Analysis of RAPD data suggests that at least two mtDNA types might have been subject to interspecific transfer due to hybridization. In the case of two sympatric haplotypes of L. brunneus we may assume that the contemporary pattern of variation between them can be explained by relatively recent hybridization with another distinct species, L. flavopunctatus . The formation of two groups belonging to distinct mitochondrial lineages within northern populations could be associated with more complex processes including ancient hybridization.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 301–316.     

Heritability and genetic correlation of abdominal versus caudal vertebral number in the medaka (Actinopterygii: Adrianichthyidae): genetic constraints on evolution of axial patterning?

Variation in the number of abdominal vs. caudal vertebrae is an important source of morphological diversification of fish. It is not clear, however, whether abdominal and caudal regions evolve independently. Regressions of offspring on parents demonstrated substantial additive genetic variation within populations, i.e. heritability, in both abdominal and caudal vertebral numbers of the medaka ( Oryzias latipes ). However, the heritability of caudal vertebrae tended to be smaller than that of abdominal vertebrae in some estimations, suggesting that abdominal and caudal regions are controlled by separate developmental modules. Furthermore, genetic correlation between abdominal and caudal vertebral numbers, estimated using full-sib family means, was negative but weak, supporting independent evolution. In addition, substantial genetic differentiation among populations was demonstrated in abdominal vertebral numbers, but not in caudal numbers. These results support our view that Jordan's rule, a geographical tendency for fish from higher latitudes to have more vertebrae, in this fish reflects local adaptations of abdominal vertebral numbers. In contrast, the low heritability of caudal vertebrae may reflect the intrinsic invariability of genes associated with a change in caudal vertebral numbers. This genetic constraint may have restricted morphological diversification of not only the medaka, but also the Order Beloniformes as a whole.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 867–874.     

A molecular study of phylogenetic relationships and evolution of antipredator strategies in Australian Diplodactylus geckos, subgenus Strophurus

We present phylogenetic analyses of the lizard genus Diplodactylus subgenus Strophurus using 1646 aligned positions of mitochondrial DNA sequences containing 893 parsimony-informative characters for samples of 12 species of Strophurus and 19 additional Australian gecko species. Sequences from three protein-coding genes (ND1, ND2 and COI) and eight intervening transfer RNA genes were examined using parsimony, maximum-likelihood and Bayesian analyses. Species of Strophurus appeared to form a monophyletic group with the possible exception of S. taenicauda . Strophurus has evolved two distinct defence/display characteristics: caudal glands, which expel an unpalatable substance, and striking mouth colours. Caudal glands appeared to have arisen once in a common ancestor of Strophurus , with dermal augmentation of caudal glands characterizing a subclade within the subgenus. Evolution of yellow and dark-blue mouth colours in Strophurus occurred in the context of diurnal activity and may be interpreted as an augmentation of defensive behavioural displays. Molecular divergence suggests that arboreality evolved in a common ancestor of Oedura and Strophurus approximately 29 Mya and that the caudal glands of Strophurus arose approximately 25 Mya.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 123–138.     

Maximizing information in systematic revisions: a combined molecular and morphological analysis of a cryptic green pitviper complex (Trimeresurus stejnegeri)

Although the integration of DNA information in taxonomy has been invaluable, logistical problems relating to sampling can seriously limit its applicability. Here we describe the analysis of a morphologically cryptic species complex, in which we maximize the information present by using both a DNA phylogeny and a multivariate morphometric approach. The green pitviper Trimeresurus stejnegeri s.l. is widespread in Asia, with a number of described subspecies (some of which are considered full species by some workers) and two new species that have recently been described from Thailand. The phylogeny indicates three clades, which can also be discerned in the principal component analyses of morphological variation. Combining molecular and morphological information permits evaluation of the taxonomic position of populations not represented in the phylogeny − in particular, the subspecies T. s. chenbihui and T. s. yunnanensis . We discuss nomenclatural issues raised by this analysis, although these cannot be fully resolved until the holotypes of these subspecies can be examined. Finally, we apply a molecular clock calibrated in New World pitvipers, and discuss some of the palaeoclimatic changes that might have impacted upon diversification in this group.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 219–235.     

Molecular and morphological data reveal cryptic taxonomic diversity in the terrestrial slug complex Arion subfuscus/fuscus (Mollusca, Pulmonata, Arionidae) in continental north-west Europe

The importance and abundance of cryptic species among invertebrate taxa is well documented. Nowadays, taxonomic, phylogenetic and conservation biological studies frequently use molecular markers to delineate cryptic taxa. Such studies, however, often face the problem of the differential resolution of the molecular markers and techniques involved. This issue is explored in the present study of cryptic taxa within the terrestrial slug complex Arion subfuscus/fuscus in continental north-west Europe. To this end, morphological, allozyme and mitochondrial 16S rDNA sequence data have been jointly evaluated. Using allozyme data and gonad type, two distinct groups were consistently delineated, even under sympatric conditions. The 16S rDNA data strongly supported both those groups and even suggested the presence of three distinct taxa within one of them. However, in view of: (1) the allopatric distribution of three OTUs, (2) the lack of allozyme or morphological differentiation, and (3) the extremely high degree of intraspecific mtDNA variation reported in pulmonate gastropods, they are, for the time being, not regarded as valid species under the biological species concept. By means of 16S rDNA and allozyme data, the position of type and topotype material of A. subfuscus s.s. and A. fuscus relative to the newly defined OTUs was determined, thus clarifying the nomenclature of this species complex. Additionally, gonad type proved to be a useful character for distinguishing the two species in north-west Europe.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 23–38.     

A new species of Arabis (Brassicaceae) from inner Anatolia

A new species of Arabis L., A. erikii Mutlu sp. nov. , is described and illustrated. The species is restricted to inner Anatolia, south-west of Sivas. It grows on steppes. Diagnostic morphological characters are given for discrimination from the most similar taxa.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 251–256.     

Uniformity of karyotypes in Ligularia (Asteraceae: Senecioneae), a highly diversified genus of the eastern Qinghai–Tibet Plateau highlands and adjacent areas

Ligularia , a highly diversified genus in the eastern Qinghai–Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x  = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2 n  = 58 and 87. Other species are only diploid, with 2 n  = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia , are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium , Ligulariopsis , Parasenecio , and Sinacalia . Aneuploid reduction of chromosome number from 2 n  = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 329–342.     

Taxonomic revision of the genus Succisella (Dipsacaceae) in the Iberian Peninsula

A revision of the three endemic Iberian species of Succisella G. Beck ( S. carvalhoana , S. microcephala and S. andreae-molinae ), based on herbarium studies, SEM photographs and field observations, is presented utilizing morphological, palynological, karyological, biogeographical and ecological characters. The distribution of the species in the Iberian Peninsula is shown in a grid map. Full synonymy is given for all taxa.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 351–364.     

Karyomorphology of Incarvillea (Bignoniaceae) and its implications in distribution and taxonomy

The karyomorphology of 11 species of the genus Incarvillea Juss. is reported. The chromosome numbers of all species studied are 2 n  = 22. The interphase nuclei and prophase chromosomes were found to be of the simple chromocentre type and the interstitial type, respectively. The asymmetry of the karyotype of I. arguta (two populations) in subgenus Amphicome is type 2A. The karyotypes of Incarvillea s inensis var. sinensis , I.  s inensis var. przewalskii, and I. olgae in subgenus Incarvillea are of asymmetry type 3A. The remaining nine species and one variety in subgenus Pteroscleris are also of asymmetry 3A. Data on three species and one variety studied are first reports. This study indicates that karyotype variation at the diploid level appears to be the predominant feature of chromosome evolution in the genus Incarvillea . According to this study of karyomorphology, morphological characteristics and geographical distribution, it seems that the three subgenera should be regarded as three independent genera. The geography of the genus is discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 113–121.     

A new Spanish species of Tragopogon (Asteraceae: Lactuceae)

Tragopogon cazorlanum (Asteraceae: Lactuceae) is proposed as a new Spanish species with distribution restricted to the Baetic mountains (south-eastern Spain). Its morphological, palynological, chromosomal, biogeographical and ecological features are discussed, as well as its main relationships and differences with reference to other Spanish species of the genus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 505–511.     

Investigating South American biogeographic history using patterns of skull shape variation on Cerdocyon thous (Mammalia: Canidae)

Patterns of geographic variation of the canid Cerdocyon thous have historically been obscured by its remarkable intraspecific morphological variability. The observed distribution is highly associated with phytophysiognomy, a feature considered highly dynamic along geological time. In the present study, we tested whether vegetation distribution during the Holocene Glacial Maximum of South America (HGM) explains the patterns of morphological variation within Cerdocyon thous . The species was divided in groups according to paleohabitats that could support their presence during the HGM, and then tested for differences in skull morphometrics. The results obtained demonstrate that the climatic changes during the HGM influenced the population structure of this species, resulting in the establishment of geographical groups with different degrees of morphological cohesion. Higher morphological cohesion found in the Northern group might be explained by the marked discontinuity between its geographical range and the rest of the species'distribution. The Eastern and Southern morphological divergence is less striking and, although this could be related to past vegetation distribution, the disappearance of those barriers leads to a population structure that could be slowly breaking down.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 77–84.     

Comparative osteology of the Danio (Cyprinidae: Ostariophysi) axial skeleton with comments on Danio relationships based on molecules and morphology

Danio is a diverse group of small, colourful and easily bred freshwater fishes native to Southern Asia. Biological interest in danios has increased in recent years because the zebrafish, Danio rerio , has become an important model organism, particularly for studies of vertebrate developmental biology and genetics. Though several phylogenetic studies of Danio have been done on a subset of Danio species, the resulting phylogenies conflict in detail. To examine the utility of osteology for systematics of this group at the species level, we studied the axial skeleton for 11 species of Danio. We analyse our morphological data alone and combined with DNA sequence data for five gene sequences generated in earlier phylogenetic studies. The axial skeleton of Danio exhibits 14 characters that prove useful in phylogenetic analysis. Both molecular and morphological data support the monophyly of the danios included in our analysis and both data sets support the monophyly of two subclades: a deep-bodied group and a slender-bodied group. Separate analysis of molecular and morphological data sets show that the molecular data resolves relationships within the slender subclade whereas the topology of the deep-bodied subclade is determined by morphological data.  © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 529–546.     

Evolution of St Helena arborescent Astereae (Asteraceae): relationships of the genera Commidendrum and Melanodendron

The cladistic relationships of endemic Commidendrum (four species) and Melanodendron (one species) from St Helena were inferred from sequences of ITS1 and ITS2 of nuclear ribosomal DNA. Despite showing a range of morphological and ecological variation, the four species of Commidendrum , C. spurium , C. robustum , C. rotundifolium , and C. rugosum , form a closely related monophyletic group with percentage sequence divergence between 0.2 and 0.9%. Melanodendron integrifolium is sister to Commidendrum indicating that the two genera may have evolved from a common ancestor that arrived in St Helena via a single dispersal event. The closest relatives of Commidendrum and Melanodendron appear to be South African, in the predominantly shrubby genus Felicia , although further sampling of South African Astereae is required. We discuss the evolution and adaptive radiation of these rare and threatened species with particular reference to the possible role of heterochrony.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 69–83.     

Patterns of morphological evolution in Marmota (Rodentia, Sciuridae): geometric morphometrics of the cranium in the context of marmot phylogeny, ecology and conservation

Marmots are of great interest for both sociobiologists studying the evolution of mammal societies and conservationists trying to protect them from extinction. In contrast, their phylogeny and morphological evolution are poorly understood and studied. Recently, a phylogenetic analysis using cytochrome b provided the first reconstruction of marmot evolutionary history and suggested that a high level of sociality evolved at least twice independently in the two proposed marmot subgenera. A morphological analysis of the marmot mandible supported this subgeneric classification and showed interesting, and unexpected, patterns in the evolution of marmot skeletal characters. In the present study we investigated a more complex, and potentially informative structure, the ventral cranium. Geometric morphometric techniques were applied in the first analysis of cranial morphology including all marmot species. Three main phenetic groups were found, which reflect phylogeny (subgenus Petromarmota , and Palaearctic subgenus Marmota ) or geographical distribution (Palaearctic vs. Nearctic subgenus Marmota ). Convergence in skeletal characters due to size similarities, a common finding in the sciurid skeleton according to traditional morphological analyses, did not occur in the marmot ventral cranium. Despite a genetic distance between Marmota vancouverensis and Marmota caligata similar to that among different populations of the latter species, the Vancouver Island marmot had the most atypical ventral cranium in the subgenus Petromarmota . This finding confirmed results obtained with the mandible, and emphasized the uniqueness of M. vancouverensis and the usefulness of complementing molecular analyses with morphological studies for a thorough characterization of population divergence, and a careful planning of conservation strategies.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 385–407     

Intraspecific variation in Anolis sagrei mirrors the adaptive radiation of Greater Antillean anoles

Anolis lizards of the Greater Antilles represent one of the classic examples of vertebrate adaptive radiation. The same morphological types ('ecomorphs') have evolved repeatedly in response to similar ecological pressures on different islands. We tested whether patterns of within species diversification were congruent with between species patterns and the processes leading to the adaptive radiation of Greater Antillean anoles by measuring variation in performance-related morphological characters in the brown anole, Anolis sagrei . We measured morphological and genetic variation in two different habitat types on each of five islands in the Bahamas. We estimated population structure and rates of gene flow within and among islands using eight microsatellite markers. Intraspecific variation in performance-related morphological characters was similar to the pattern of interspecific variation that characterizes the adaptive radiation of this group in the Greater Antilles. For example, limb length was correlated with perch diameter within A. sagrei as has also been shown among species of anole. Morphological divergence in traits has occurred despite relatively high levels of gene flow both within and among islands. These results are discussed in the context of the divergence-with-gene-flow model of speciation. The results provide important intraspecific evidence that the diversification of anoles has been shaped by natural selection and show how ecologically-based selection pressures explain diversification at both the population and species levels.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 189–199.     

Multivariate analyses of morphological characters of Leptolaena Thouars s.l. subgenera Mediusella and Xerochlamys (Sarcolaenaceae)

Leptolaena Thouars is one of the first described genera within Sarcolaenaceae, but represents the last great species-level taxonomic challenge in this endemic Malagasy plant family. As currently circumscribed, this genus comprises three subgenera, two of which, Mediusella Cavaco and Xerochlamys (Baker) Cavaco, exhibit patterns of morphological variation that have been understood differently and have led to an unstable taxonomy, such that a consensus on the delimitation of species has yet to be reached. Principal component analysis of morphological characters was performed on 211 herbarium specimens in order to evaluate the morphological variation within the two subgenera and to clarify historically problematical species boundaries. The results showed that ten entities could be distinguished on the basis of vegetative, floral, and fruit characters, and that they provide a robust basis for proposing a new species-level taxonomic framework.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 559–574.     

Galápagos' Opuntia (prickly pear) cacti: extensive morphological diversity, low genetic variability

Due to the pronounced morphological variation and geographical distribution of Galápagos' Opuntia cacti, numerous hypotheses have been advanced regarding their radiation, diversification, and classification. The currently accepted classification is based on morphology and recognizes six species and fourteen varieties, but the plasticity of many of the characteristics renders any morphological taxonomy problematic. Our analysis of previously published morphological data agrees only partially with the current classification. We present the first molecular phylogeny of these plants. Multiple DNA sequences indicate little genetic distinction among the currently identified species, despite restricted gene flow and limited long distance dispersal within the archipelago. No clear relationship exists between morphological and genetic differences. These results suggest that both molecular and morphological data should be used in conservation planning.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 451–461.     

Phylogeography of the freshwater crayfish Cherax destructor Clark (Parastacidae) in inland Australia: historical fragmentation and recent range expansion

The yabby, Cherax destructor Clark, is the most widespread species in the most widespread genus of Australian freshwater crayfish. It has a distribution that spans several distinct drainage basins and biogeographical regions within semiarid and arid inland Australia. Here we report a study designed to investigate patterns of genetic variation within the species and hypotheses put forward to account for its extensive distribution using DNA sequences from the mitochondrial 16S rRNA gene region. Results of phylogenetic analyses contradicted previous allozyme data and revealed relatively deep phylogenetic structure in the form of three geographically correlated clades. The degree of genetic divergences between clades (8–15 bp) contrasted with the relatively limited haplotype diversity within clades (1–3 bp). Network-based analyses confirmed these results and revealed genetic structure on both larger and more restricted geographical scales. Nevertheless some haplotypes and 1-step clades had large distributions, some of which crossed boundaries between river basins and aquatic biogeographical regions. Thus both older and more recent historical processes, including fragmentation on a larger geographical scale and more recent range expansion on a local scale, appear to be responsible for the observed pattern of genetic variation within C. destructor . These results support elements of alternative hypotheses previously put forward to account for the evolutionary history of C. destructor and the origin of its large distribution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 539–550.     

Do taxonomic divisions reflect genetic differentiation? A comparison of morphological and genetic data in Coenonympha tullia (Müller), Satyrinae

Historically, morphological variation has been used to classify many species (and subspecies) of Lepidoptera. However, some of this variation may be unsuitable for inferring the recent evolutionary history of populations. Genetic data provide an alternative. We examine the morphological and genetic variation within and between British subspecies of Coenonympha tullia (Müller 1764) to test the hypothesis that neutral genetic variation corresponds to morphological variation. We find that most morphological and genetic variation occurs within populations and that those populations designated as subspecies based on morphological characters are not necessarily most closely related for mitochondrial DNA or nuclear DNA (amplified fragment length polymorphisms and allozymes). Thus, the notion that wing spot variation reflects population isolation and therefore genetic differentiation does not hold. The present study highlights the need for genetic data where taxonomy may be based on environmentally plastic or locally adapted characters because such characters will not reflect the true population genetic history.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 314–327.