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.     

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.     

A new species of Ilex (Aquifoliaceae) from Espinhaço Range, Bahia, Brazil

A new species, Ilex mucugensis Groppo sp. nov. (Aquifoliaceae), is described and illustrated. The species is known only from 'campos rupestres' (vegetation of rock outcrops at high altitude) in the vicinity of Mucugê, Bahia State, Brazil, in the mountains of the Espinhaço Range. Comments on affinities, geographical distribution and phenology of the new species are also provided.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 153–156.     

Dispersal history of a spider (Stegodyphus lineatus) across contiguous deserts: vicariance and range expansion

Israel marks a crossroads between three continents encompassing several phytogeographical and zoogeographical zones. In this complex area, the flow of species from different biogeographical regions creates opportunities to study how geographical division and colonization routes affect current distribution and structure of resident populations of organisms associated with desert and arid environments, habitats that may have persisted throughout Pleistocene glacial periods. The present paper analyses the population history of the spider Stegodyphus lineatus in the contiguous Negev and Judean deserts in Israel using allozyme and mtDNA variation. The distinct patterns of variation indicate that Judean and Negev populations are vicariant lineages. The residence time was longer in Judea, where populations were more polymorphic for mtDNA, showed isolation by distance and were less structured than in the Negev. The Negev population, possibly linked to other Mediterranean populations of S. lineatus , consisted of two subdivisions derived from a recent eastward expansion across the central Negev watershed. Despite differences in age and level of structure, all lineages show similar dispersal processes dominated by restricted gene flow. The distribution patterns of allozyme and mtDNA markers are unrelated to geographical patterns of precipitation and vegetation. Rather, they follow large-scale topographic features, namely the water divide between Mediterranean and Afro-Syrian rift drainages and between eastern and western Negev drainages.   © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 739–754.     

Cryptic speciation in liverworts – a case study in the Aneura pinguis complex

Bryophytes are amongst the most ancestral terrestrial plants and often have large distribution ranges across continents. Recent biochemical and molecular studies have suggested that many worldwide morphological species of bryophyte may represent genetically divergent and reproductively isolated cryptic species. We tested the cryptic species hypothesis in the thalloid liverwort Aneura pinguis complex. We applied analyses of chloroplast DNA (cpDNA) sequence variation and polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) methods to discriminate between specimens of A. pinguis derived from various habitats in five distant geographical regions in Poland. Of the 19 specimens sequence characterized for the cpDNA tRNA^Leu region, seven haplotypes were identified divided into three nonmonophyletic clusters. The application of developed PCR-RFLP markers confirmed the existence of three tRNA^Leu types of A. pinguis (A–C) within the specimens derived from 21 populations. Sympatric populations of different tRNA^Leu types were found in lowland and mountain regions. No clear correlation between stand type and the presence of two tRNA^Leu types (A, B) was observed, as both were growing on soil, humus, and rocks. The tRNA^Leu type C was found only on humus and its distribution was restricted to low-lying northern populations. The above results indicate that the A. pinguis complex is highly differentiated at the molecular level and may represent three cryptic species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 273–282.     

Morphological and geographic variation of the cycad Dioon edule Lindl. (Zamiaceae): ecological and evolutionary implications

The relationship in geographical distribution and morphological variation of leaflet width and length (diagnostic trait), between and within populations of Dioon edule Lindl., has been investigated throughout its known range in eastern Mexico (from the states of Nuevo León to Veracruz, north to south, respectively). A total of 1832 leaflets were measured for width and length from 154 plants distributed amongst five populations using four leaflet replicas from each of three leaves per plant. For leaflet width and length the variation among populations indicated significant stat-istical differences ( F _4,147 = 125.83; P  < 0.0001; R ² = 92.17% and F _4,147 = 9.04; P  < 0.001; R ² = 26.8%), respectively. With respect to leaflet width, the multiple range test showed three groups with a north to south distributional relationship along the range of the species. The correlation coefficient among paired populations, respect to geographical distance and the absolute value of the mean difference of leaflet width in each population, was positive, and different from zero ( r  = 0.82; P  = 0.013). A great variation of important ecological and evolutionary parameters was shown.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 465–470.     

Genetic cohesion of Eresus walckenaeri (Araneae, Eresidae) in the eastern Mediterranean

The eresid spider genus Eresus is morphologically and ecologically conservative. At least three species occur in Europe. However, deep genetic divergence among geographical samples within two species, E. cinnaberinus and E. sandaliatus , may suggest more cryptic species. In the present study we investigate the genetic cohesion of the third species, Eresus walckenaeri , throughout its eastern Mediterranean distribution range, relative to the E. cinnaberinus–E. sandaliatus species complex. Eresus walckenaeri specimens were monophyletic. Genetic discreteness of E. walckenaeri in a region of sympatry with its sister species in Greece provides evidence for species integrity of E. walckenaeri within the European Eresus species complex. Eresus walckenaeri exhibited high concordance between geographical location and mtDNA genealogy. Two major phylogeographical clades were found in the Greek–Turkish and Syrian–Israel parts of the investigated area, respectively (∼6.5% sequence divergence). Concordance between geography and genetic divergence was further observed between Aegean island samples and their corresponding Greek and Turkish mainland samples, suggesting regional subdivision with gradual but potentially high dispersal propensity. Monophyly and limited regional distribution indicate Mediterranean endemic origin.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 1–9.     

Combining molecular, morphological and ecological data to infer species boundaries in a cryptic tropical pitviper

Few operational methods exist for delimiting species boundaries, and these usually require sampling strategies that are unrealistic for widespread organisms that occur at low densities. Here we apply molecular, morphological and ecological species delimitation criteria to a wide-ranging, fragmented group of Asian green pitvipers, the Popeia popeiorum complex. A mitochondrial DNA phylogeny for the group indicates two well-differentiated clades, corresponding mainly to northern and southern parts of its range. Strong phylogeographical structure within each clade suggests isolation in forest refugia during the Pliocene and a southward colonization of the Sunda islands during the Pleistocene. Multivariate analysis of morphological characters reveals a generally conserved pattern of geographical variation, incongruent with the recovered phylogenetic history. We compare groups delineated by mtDNA variation to morphological and ecological divisions in the complex, and discuss the implications of these for the taxonomy of the group. Discordance between species boundaries inferred from different criteria suggests that combining independent sources of data provides the most reliable estimation of species boundaries in organisms that are difficult to sample in large numbers.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 343–364.     

Multivariate analysis of morphological variation in Cineraria deltoidea (Asteraceae, Senecioneae)

Cluster analysis and principal coordinates analysis were used to investigate phenetic variation in Cineraria deltoidea , a species that ranges from near sea level in KwaZulu-Natal, South Africa, to 4300 m a.s.l. on the mountains of East Africa and Ethiopia. Earlier taxonomic revisions reduced nine previously recognized species to synonyms of C. deltoidea . Two closely related species, C. decipiens and C. atriplicifolia , were also included in the analyses. Thirty-six morphological characters were examined on 111 specimens. Phenograms and scattergrams show partial clusters of specimens of C. deltoidea from individual mountains or geographical regions, but no groups are sufficiently distinct to warrant formal recognition at any rank. The East African specimens from 3000 m a.s.l. and higher tend to cluster together. Growth at high altitude in East Africa is correlated with fewer, larger capitula on longer peduncles, and an absence of a cobwebby indumentum comprising long, narrow-based trichomes. Cineraria deltoidea is thus a highly variable species with geographical and clinal variation evident throughout its range. Cineraria atriplicifolia and C. decipiens are maintained as distinct species, distinguished from C. deltoidea by their growth form, life span and auricle shape.  © 2007 University of the Witwatersrand, Johannesburg. Journal compilation © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 497–521.     

A new species of Exostyles (Leguminosae, Papilionoideae, Swartzieae s.l.), from Paraná State, Brazil

A new species Exostyles godoyensis Soares-Silva & Mansano (Leguminosae, Papilionoideae, Swartzieae), endemic to Paraná state in southern Brazil, is described and illustrated. The new species is distinct from all the other species of Exostyles in having a leaf with up to five leaflets, while the other species have at least seven leaflets per leaf. The new species is distinct from Exostyles amazonica Yakovlev in its longer stamen filaments and ovary stipe and from E. glabra Vogel and E. venusta Schott by its shorter anthers. In addition to these morphological characters E. godoyensis has a disjunct geographical distribution.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 103–106.     

Two approaches for the selection of Relevant Genetic Units for Conservation in the narrow European endemic steppe plant Boleum asperum (Brassicaceae)

Several methods based on population biology, biogeography, ecology, and genetics have been traditionally used for the identification of units for conservation below the species level. We use a combination of two methods based on population genetic structure estimators and on probabilities of loss of rare alleles to identify the Relevant Genetic Units for Conservation (RGUCs). The aims were to assess the genetic diversity and population structure of the endemic steppe plant Boleum asperum (Brassicaceae), and to determine how many and which populations significantly represent the total genetic diversity and the rarest allelic variation. Despite the high amplified fragment length polymorphism genetic diversity values detected in B. asperum ( h _T = 0.744), caused probably by its hexaploidy and allogamy, moderate spatial genetic differentiation was detected among populations (< 20%) and geographical ranges (> 13%), suggesting the existence of an ancestral continuous distribution range that was fragmented into separate 'islands' in more recent historical times. Five RGUCs, accounting for the 85.10% of the total genetic variation and representative of the entire geographical distribution of the species, were selected for in situ conservation. Ex situ conservation is proposed to complement the preservation of B. asperum . This method of objective selection of populations may be applied to other candidate taxa for conservation with prior adjustment of the threshold values of diversity required for effective protection of each particular species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 341–354.     

Morphological variation in house mice from the Robertsonian polymorphism area of Barcelona

Morphometric variation in the Robertsonian polymorphism zone of Barcelona of Mus musculus domesticus was studied by geometric morphometrics. This system is characterized by populations of reduced diploid number (2 n  = 27–39) surrounded by standard populations (2 n  = 40). We investigated the morphological variation in mice from this area, as well as the effect of geographical distance and karyotype on this variation. We also investigated the degree of co-variation between the two functional units of the mandible to explore the origin of this system (primary intergradation or secondary contact). The size and shape of the cranium, mandible and scapula were analysed for 226 specimens grouped by population, chromosome number and structural heterozygosity. Size was estimated as the centroid size, and shape was estimated after Procrustes superimposition. No significant differences in size between populations or chromosomal groups were detected. Diploid number, structural heterozygosity and local geographical isolation contributed to the differentiation in shape. Morphological differentiation between standard mice and Robertsonian specimens was observed, suggesting genetic isolation between these groups. Co-variation between the ascending ramus and alveolar region of the mandible was quantified by the trace correlation between landmark subsets of these modules. The trace values showed an ascending trend, correlated with the distance from the centre of the polymorphism area, a pattern consistent with a primary intergradation scenario.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 555–570.     

Leaf morphometric variation in Quercus affinis and Q. laurina (Fagaceae), two hybridizing Mexican red oaks

Leaf variation was examined in two hybridizing Mexican red oaks, Quercus affinis and Q. laurina . Data of nine traits were obtained for ten randomly chosen leaves in each of 512 individuals from 16 populations sampled along a geographical gradient, including the distribution area of both species and a putative area of secondary contact and hybridization. A cluster analysis recognized two main groups of populations, which were congruent with the taxonomic assignment of the predominant morphological type within the populations and were thus labelled ' Q. affinis -like' and ' Q. laurina -like' population groups. A nested analysis of variance revealed that the largest proportion of the total variation was contained within populations, as among-tree variation (28–54%, depending on the trait), and as intra-individual variation (17–56%). However, differences between the two groups of populations (3–27%) and among populations within groups (5–21%) were also significant for the nine traits. A distinct pattern of change across populations was observed for each trait. Variation was particularly pronounced along the geographical gradient for petiole length and leaf-margin teeth number, possibly implying selection on these two traits. Results suggest that phenotypic plasticity, gene flow, hybridization and natural selection have shaped foliar variation in this oak complex. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 427–435.     

Geographical variation and diversification in the flightless leaf beetles of the Chrysolina angusticollis species complex (Chrysomelidae, Coleoptera) in northern Japan

Geographical variation and differentiation of flightless leaf beetles of the Chrysolina angusticollis species complex were analysed. Nine morphological groups were recognized on the basis of metric characters relating to body size, shape of hind wing and shape of male genitalia, and binary characters relating to females' elytral surface and larval pubescence. The nine groups were essentially parapatric or allopatric to each other. Five groups occupied a continuous geographical range each, whereas another four groups were disjunctively distributed. Morphometric distance and geographical distance between populations were significantly correlated. Analyses based on 13 allozyme loci demonstrated that six out of the nine morphological groups were further divided into more than one genetically distinct group each. Genetic similarity between populations was significantly correlated with morphometric similarity and geographical closeness. Disjunctively distributed populations of a single morphological group more often resembled geographically adjacent populations of different morphological groups than geographically remote populations of the same group. In a morphological group that occupied a wide continuous range, remote populations substantially differed genetically from each other. The results suggest that there exist at least 19 morphologically and/or genetically differentiated units in the C. angusticollis complex in northern Japan. The C. angusticollis complex is likely a superspecies composed of a number of semispecies. We postulate that the diversification of this species complex might have resulted from the repeated range contraction and expansion during the ice ages. Movement of tension zones that separate two adjacent forms might contribute to enhance their geographical differentiation.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 557–578.     

Volatile leaf oil diversity in the narrow range endemic Eucalyptus argutifolia (Myrtaceae) and its widespread congener Eucalyptus obtusiflora

Variation in the qualitative composition of volatile leaf oil was examined in the rare Eucalyptus argutifolia and its widespread congener Eucalyptus obtusiflora . The results revealed that, consistent with the pattern seen with allozymes, E. argutifolia had less variation within populations than E. obtusiflora . Total leaf oil diversity was also significantly lower in the rare species. As found with allozymes, most leaf oil diversity was within populations, but there was also a significant proportion of the variation between populations (25.2% and 27.3% for E. argutifolia and E. obtusiflora , respectively). There were significant associations between phenotypic distance based on leaf oils and geographical distance and between phenotypic and genetic distance across all populations, but these associations were not evident within species. Factors leading to reduced variation in E. argutifolia appear to affect all types of variation, but the relationships between different types of variation within the species are less apparent.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 738–745.     

Astragalus sect. Astragalus (Fabaceae) in Iran

Astragalus rahiminejadii , a new species endemic to Iran, is described and illustrated. This species, which belongs to section Astragalus , is confined to the western part of Iran (Prov. Kermanshah) and is known only from a single population. Seed testa morphology as viewed under the scanning electron microscope is discussed and photomicrographs are provided. In addition, the geographical distribution and ecology of the species belonging to this section are discussed.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 363–368.     

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.     

A new species of Linum (Linaceae) from west Anatolia, Turkey

A new species, Linum vuralianum Yılmaz & Kaynak, is described and illustrated from west Anatolia, Turkey. It is closest to L. flavum L., but differs by its annual or biennial life form, the presence of rosette leaves at anthesis, and the absence of staminodes. The World Conservation Union (IUCN) threat category and observations on the population are noted. The geographical distribution of the new species and its related taxon is given.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 459–462.     

Distinct geographical structure across species units evidenced by chloroplast DNA haplotypes and nuclear ribosomal ITS genotypes of Corylopsis (Hamamelidaceae) in the Japanese islands

The geographical distribution of chloroplast DNA (cpDNA) haplotypes and nuclear ribosomal internal transcribed spacer (nrITS) genotypes of Japanese Corylopsis (Hamamelidaceae), which consists of four species, was investigated. Two hundred and five individuals belonging to four species from 30 populations, covering the entire geographical range, were studied. Based on approximately 1108 bp of the three non-coding regions of cpDNA, nine haplotypes were detected, and each was distinguished from adjacent haplotypes by one substitution. Based on approximately 507-bp nrITS sequences, 47 genotypes were detected, for which three clades were identified in the phylogenetic analysis. There was inconsistency between the cpDNA haplotypes, nrITS genotypes, and classification of Corylopsis taxa, possibly because of incomplete lineage sorting or introgressive hybridization. The distribution of the haplotypes was highly structured geographically, and N _ST (0.893) was significantly greater than G _ST (0.819), implying that the current distribution of Corylopsis species was structured phylogeographically during Quaternary climatic oscillations. The haplotype composition and results of analysis of molecular variance showed that the populations in Hokuriku were highly divergent, suggesting that they are long-term persistent populations arising from refugia during the Quaternary climatic oscillations. Refugial populations in Chugoku and Shikoku may have lost genetic diversity because of a bottleneck resulting from a small population size, followed by post-glacial range expansion. Pre-existing refugia may have been so small that the subsequent range expansion replaced the pre-existing genetic structure.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 501–518.