Patterns of genetic diversity in related taxa of Antirrhinum L. assessed using allozymes

Allozyme diversity was studied within and among populations of five related taxa of Antirrhinum L. endemic to the Iberian Peninsula ( A. graniticum Rothm. ssp. graniticum , ssp. brachycalyx Sutton and ssp. ambiguum (Lange) Mateu & Segarra, A. boissieri Rothm. and A. onubensis (Fdez. Casas) Fdez. Casas). All of the studied taxa are obligate outcrossing endemic perennial herbs which form isolated populations. However, the taxa vary in range and population sizes, and are found on different soil types. The level and distribution of allozyme diversity differed widely between taxa: A. graniticum ssp. brachycalyx had the lowest level of allozyme diversity (H_T = 0.09), whilst the highest level was detected in A. boissieri (H_T = 0.25). Total variation was partitioned into within- and among-population variation. The proportion attributable to variation within populations varied from about 67% up to 84.3% and 89.5% in A. graniticum ssp. brachycalyx and A. graniticum ssp. ambiguum , respectively. Both these subspecies also showed little population divergence (G_ST = 0.10 and 0.09, respectively) and had high levels of estimated gene flow (Nm = 2.18 and 2.62, respectively). These results are discussed in relation to geographical proximity of populations and habitat continuity. Isolation by distance was not detected in any of the studied taxa. This result suggests that divergence among populations is due to random genetic drift.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 79 , 299–307.     

Parnassia palustris: a genetically diverse species in Scandinavia

Patterns of variation at nine enzyme loci were examined in 528 plants representing diploid and tetraploid populations of Parnassia palustris s. l. in Europe to assess genetic variation patterns and migration history. Half of the plants showed a unique multilocus phenotype and 75% of all phenotypes occurred only in Scandinavia. Diploid populations showed similar levels of genetic diversity as other widespread outbreeding species with animal-mediated pollination and F -statistics indicated excessive heterozygosity and low rates of gene flow among them. In spite of dramatic population histories caused by the ice ages, diploid populations have maintained the same genetic diversity in Scandinavia as in central and southern Europe. Northern populations have apparently been established through the gradual advance of genetically variable populations and patterns of variation at individual loci indicate different migration routes, from the south-south-west and the east-north-east, respectively. The data strongly support a repeated autoploid origin of the tetraploid cytotype which has been much more successful than the diploid progenitors in colonizing new land since the last ice age. High genetic diversity in Scandinavia has apparently been obtained by a combination of immigration of plants from different source areas and recurrent formation of autotetraploids from diploid progenitors.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 347−372.     

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.     

Mitochondrial DNA polymorphisms of the house mouse Mus musculus domesticus from Greece, focusing on the Robertsonian chromosomal system of north-west Peloponnese

This study focused on the sequence variation in a mitochondrial region in house mice Mus musculus domesticus from chromosomally variable populations in the north-west Peloponnese, Greece. The mitochondrial molecular variation revealed was among the highest found so far for M. m. domesticus populations. The haplotype distribution pattern was rather complex. There was no clear differentiation between the populations characterized by Robertsonian chromosomes and standard all-acrocentric populations. There is therefore no indication that the Robertsonian populations were formed during a prolonged period of geographical isolation.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 643–651.     

Genetic variation of Saxifraga paniculata Mill. (Saxifragaceae): molecular evidence for glacial relict endemism in central Europe

Saxifraga paniculata Mill. is an amphi-atlantic species that is widely distributed in the Alps but also occurs with disjunct populations in central Europe. These isolated populations are considered to be glacial relicts. In the study presented here, we analysed the genetic variation within and between populations of Saxifraga paniculata in central Europe to test whether the molecular data give evidence for the supposed relict endemism. We used RAPD analysis and the analysis of molecular variance (AMOVA) technique to investigate 30 populations of Saxifraga paniculata from different locations in central Europe and adjacent geographical areas. In the RAPD analysis, 319 fragments could be amplified of which 91.2% were polymorphic. The percentage of polymorphic bands within populations correlated significantly with population size, indicating a higher level of inbreeding in small populations. AMOVA revealed 43.60% of the variation within populations, only 15.45% between populations within locations, and 40.95% between locations. Genetic distance between locations (Φ_ST) correlated significantly with the geographical distance between locations, giving evidence for geographical isolation. In an UPGMA cluster analysis, based on RAPD data, the populations were grouped together according to their location. The results exhibit a strong genetic differentiation which is obviously due to genetic drift in the isolated populations. Our study therefore gives evidence for glacial relict endemism of Saxifraga paniculata in central Europe.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 11–21.     

Occurrence of tetraploid and octoploid cytotypes in Senecio jacobaea ssp. jacobaea (Asteraceae) in Pannonia and the Carpathians

Four different cytotypes have been reported for Senecio jacobaea L. ssp. jacobaea throughout Europe, with the most common occurrence of tetraploids (2 n  = 40). Here we present a survey of previously published chromosome number data on this subspecies and its geographical distribution, and focus on populations from Pannonia and the Carpathians. Two ploidy levels have been determined in the study area, using chromosome counting and flow cytometry: tetraploid (2 n  = 40) and octoploid (2 n  = 80). Fifty-one populations originating from Slovakia, Austria, the Czech Republic, Hungary, Ukraine and Romania have been analysed. Multivariate morphometric analyses have been performed on 39 populations to study morphological differentiation between these two cytotypes. Despite slight morphological tendencies expressed on the level of populations, tetraploid and octoploids cannot be reliably distinguished morphologically and they are not taxonomically classified formally here.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 231–242.     

The Deschampsia cespitosa complex (Poaceae: Aveneae) with special reference to Russia

The Deschampsia cespitosa complex in Russia is revised. Related taxa common to Russia and adjacent regions (i.e. northern Europe, Arctic America and Japan) are also considered. Deschampsia cespitosa s.l. is characterized by open to less commonly rather contracted panicles, spikelets (1) 2 (3)-flowered, 4-toothed lemmas with the lateral teeth larger, and awns straight or bent, inserted from the base to the top of the lemmas. Due to extensive overlapping in diagnostic characters, it is not possible to accept the specific status for D. borealis (Traut.) Roshev., D. brevifolia R.Br., D. macrothyrsa Tatew. et Ohwi and D. obensis Roshev; instead, they are treated as subspecies of D. cespitosa (L.) Beauv. Morphological differences are insufficient for a clear differentiation among subspecies, and geographical distributions also need to be considered. A total of 14 subspecies is accepted. Observations on ecology, problems of specific and subspecific delimitation, a distribution map, and considerations of evolution of the complex are provided. One new combination is made, Deschampsia cespitosa ssp. sukatschewii (Popl.) Chiapella & Prob.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 213–228.     

Cytogeography of the South American Turnera sidoides L. complex (Turneraceae, Leiocarpae)

New chromosome number determinations are presented for 48 populations of five subspecies of Turnera sidoides. Chromosome counts in sspp. carnea, holosericea, integrifolia and sidoides confirm previous data while, in ssp. pinnatifida , the numbers 2n=2x = 14 and 2n = 6x = 42 are reported for the first time. The results show that polyploidy is a very frequent phenomenon. Populations are primarily tetraploids (2n = 4x = 28), while diploids (2n = 2x = 14), hexaploids (2n = 6x =42) and octoploids (2rc = 8x = 56) are less frequent. Diploid, tetraploid and hexaploid cytotypes occur in sspp. carnea and pinnatifida , whereas tetraploids and hexaploids are known in ssp. holosericea. Turnera sidoides ssp. integrifolia shows a polyploid series with ploidy levels from 2x to 8x. The ssp. sidoides appears to be uniformly tetraploid, with the exception of an isolated pentaploid (2rc = 5x = 35). On the basis of these counts and previous reports the geographical distribution of the cytotypes has been determined and related to climatic and ecological factors.     

Determination of ploidy and nuclear DNA content in populations of Atriplex halimus (Chenopodiaceae)

Nuclear DNA contents were determined by flow cytometry for 20 populations of the perennial C₄ shrub Atriplex halimus L. (Chenopodiaceae) originating from the Mediterranean basin and Fuerteventura (Canary Islands). Two populations were also analysed for chromosome number: one (from Ibiza, Spain), with a 2C nuclear DNA content of 2.40 pg, was shown to be diploid (2 n  = 2 x  = 18), whilst the other (from Sicily, Italy), with 5.11 pg, was tetraploid (2 n  = 2 x  = 36). With respect to nuclear DNA content, two groups of populations were detected, diploids with 2.40–2.44 pg and tetraploids with 4.77–5.13 pg. The diploid populations were mainly from the western Mediterranean (Spain and France) and Fuerteventura, whereas tetraploids were generally, but not exclusively, from more arid areas in North Africa and the eastern Mediterranean. In general, the diploid and tetraploid populations corresponded to the subspecies halimus and schweinfurthii , respectively. For certain populations having morphologies intermediate between those considered typical of these two subspecies, nuclear DNA contents showed them to be tetraploid. There was significant variation in nuclear DNA content among the tetraploid populations, with greater values in the more easterly populations.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 147 , 441−448.     

Morphological and genetic analysis of fish of a Carassius complex (Cyprinidae) in Lake Kasumigaura with reference to the taxonomic status of two all-female triploid morphs

The classification of the Carassius complex (Cyprinidae) including all-female triploids, called ginbuna in Japanese, is so confused that three sympatric morphs of crucian carp in Lake Kasumigaura are categorized into two different subspecies within a species. We examined them in order to explain the coexistence of more than one subspecies and determine the founder of the triploid lineages in the crucian carp fauna in the lake. Principal component analysis proved that the three sympatric morphs had a morphometric basis distinguishable from each other. Ploidy was determined by flow cytometry which showed triploids in two morphs and diploids in the other morph. Stepwise discriminant analysis using only meristic characteristics could separate the diploids from the triploids. Phylogenetic analysis using mitochondrial DNA inferred two lineages in which one was composed of a triploid morph and the other was a diploid–triploid mixture. Disagreement between the taxonomic status and the phylogenetic status is explicable by assuming that the triploids in the Carassius complex had independent origins leading to the different subspecies. C. auratus langsdorfii appears to show genetic complexities that traditional taxonomic classification can not unravel.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 351–357.     

Arbuscular mycorrhizal fungi: genetics of multigenomic, clonal networks and its ecological consequences

Arbuscular mycorrhizal fungi are thought to have remained asexual for 400 million years, although recent studies have suggested that considerable genetic and phenotypic variation could potentially exist in populations. A brief discussion of these multigenomic organisms is presented.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 59–60.     

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.     

Complex patterns of morphological and mtDNA variation in Lacerta perspicillata (Reptilia; Lacertidae)

Lacerta perspicillata is a north-west African lacertid lizard that shows considerable intraspecific variation, with three subspecies described on the basis of colour pattern and body size. Recent observations of a population containing two morphological forms and more than one deep genetic lineage, as well as an apparent lack of concordance between forms and genetic lineages, suggest that the complexity is greater than previously thought. To analyse and quantify this variation, we studied the variability within this species at two levels: (1) external morphology (multivariate analysis of scalation, body dimensions, and colour pattern) and (2) mtDNA (sequencing and single-strand confirmation polymorphism analysis). Fifty-two individuals were studied at Taza, northern Morocco. Two morphological groupings (ostensibly representing two previously described subspecies) and two deep mtDNA lineages were detected at this site, with complete correspondence between the two. This, together with an apparent lack of hybrids, would normally support respective full species recognition. However, analysis of 98 individuals from other populations demonstrated that the situation is highly complex with the same genetic lineages having reversed morphotypes in other areas, making such a designation difficult. Across the other studied populations, we found no support for any of the currently recognized subspecies. The lack of congruence between mtDNA lineages and morphometric patterns (in some cases) and the morphological similarity among lizards from different lineages suggest ecophenotypic convergence or multiple introgressive hybridization. The study highlights the tremendous complexity that may exist within a taxon and the inadequacy of older alpha-taxonomy based designations in describing it.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 479–490.     

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.     

Genetic differentiation and gene flow of two sparidae subspecies, Diplodus sargus sargus and Diplodus sargus cadenati in Atlantic and south-west Mediterranean populations

A total of nine enzymes coded by 14 loci were assayed for each of six populations (from the north-eastern Atlantic and the Mediterranean) of two sea bream subspecies ( Diplodus sargus sargus and Diplodus sargus cadenati ). Diagnostic alleles were observed for each subspecies, although there were several common alleles. Estimates of variance in allele frequencies among samples ( F _ST) revealed significant differences ( P  < 0.05) among both subspecies. Genetic divergence was found between Atlantic and Mediterranean samples: values for genetic distances were higher than 0.163. Furthermore, D. sargus cadenati populations displayed a higher mean weight and length than D. sargus sargus populations and significant differences in growth were found among subspecies and populations. These results are discussed in terms of levels of gene flow and its respective relationships with water circulation in the Strait of Gibraltar and geological events.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 705–717.     

Phylogeography of five Polytrichum species within Europe

Using allozymes and microsatellites we have analysed the genetic structure among European populations for several Polytrichum species to infer relevant factors, such as historical events or gene flow, that have shaped their genetic structure. As we observed low levels of genetic differentiation among populations, and no decreasing levels of genetic variation with increasing latitude within most of the examined species, no genetic evidence was obtained for a step-wise recolonization of Europe from southern refugia after the latest glacial period for P. commune , P. uliginosum , P. formosum and P. piliferum . The near absence of population substructuring within these species does indicate that extensive spore dispersal is the most important factor determining the genetic structure among European Polytrichum populations. Gene flow levels have apparently been sufficient to prevent genetic differentiation among populations caused by genetic drift, and to wipe out any genetic structure caused by the postglacial recolonization process. On the other hand, increased genetic differentiation of alpine P. formosum populations suggests that mountain ranges might restrict gene flow significantly among Polytrichum populations. In contrast to most examined Polytrichum species, P. juniperinum showed high levels of genetic differentiation and a profound genetic structure. Assuming that gene flow is not more restricted in P. juniperinum , these findings suggest that this species has recolonized Europe after the latest glacial period from two different refugia, one possibly being the British Isles.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society 2003, 78, 203–213.     

Geographic variation in the long-nosed snake, Rhinocheilus lecontei (Colubridae): beyond the subspecies debate

Scalation, colour pattern, linear and geometric morphometrics were used to quantify geographical differentiation in the long-nosed snake, Rhinocheilus lecontei, and to test the hypothesis that all four subspecies are morphologically distinct. Also investigated were potential associations between morphological (scalation, colour pattern, linear measurements) and environmental variables (climate, vegetation, soil). Sexual dimorphism was weakest for geometric and strongest for linear morphometric variables. Morphological variables differed widely in their ability to differentiate subspecies. Linear morphometric variables achieved the most statistically significant pairwise Mahalanobis distances between subspecies, while geometric morphometrics largely failed to differentiate them. Colour pattern showed the strongest and linear morphometrics the weakest correlation with environment. Several characters varied continuously along latitudinal or longitudinal gradients, such that, in some cases, the clines for closely related traits were discordant. No one subspecies was consistently divergent in all analyses, leading to the conclusion that the three mainland subspecies are not sufficiently distinct to warrant separate subspecies status. The island subspecies, though not always statistically distinct, is geographically separate from other populations and differs in characters related to size. Given the small number of specimens available, a decision regarding its taxonomic status (i.e. elevation to species level) is best deferred until additional specimens can be examined and data on molecular variation can be analysed.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 65–85.     

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.     

Morphological and pollen differentiation in Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii

A multivariate analysis to differentiate morphologically the populations of wild potatoes Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii was carried out. An analysis of the morphology and the viability of pollen of these potato populations was also made. The results of the morphometric analysis indicate that both subspecies are phenetically different. The pollen grain shape and size in ssp. cardiophyllum are different in northern and southern populations. Pollen diameter is significantly different between subspecies. Based on these results we propose that these taxa should be considered as two separate species.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 415–426.