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.     

Infraspecific variation and phylogeography of the high-polyploid Iberian endemic Anthoxanthum amarum Brot. (Poaceae; Pooideae) assessed by random amplified polymorphic DNA markers (RAPDs) and morphology

High morphological and ecological diversity has been observed in the north-west Iberian Peninsula endemic Anthoxanthum amarum Brot., for which two different morphotypes (northern and southern) have been described on the basis of qualitative traits and geographical origin. In the present study, a combined molecular and morphological method was applied to ten populations of this species with the following aims: (1) to assess whether the variation observed was taxonomically meaningful; (2) to assess the influence of the environment on the variation in the morphological characters; and (3) to track the potential phylogeographical information deduced from our random amplified polymorphic DNA marker (RAPD) data in order to draw inferences about the past history of this species in the north-west Iberian Peninsula. To achieve these aims, 26 macromorphological characters were recorded in 279 specimens, and 77 RAPD phenotypes were identified in the 79 plants studied. The association analyses performed using the morphological and molecular data showed that no clear separation existed between the morphs, and a strong correlation between qualitative characters and the environment was detected. Moreover, both the multivariate analyses and the assignment test based on RAPD data revealed that the genetic variation was hierarchically structured, and three genetically distinct groups could be identified. Two of these clusters might correspond to different expansion routes proposed in the literature for different plant species in the north-west Iberian Peninsula.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 179–192.     

Local adaptation in four Iris species tested in a common-garden experiment

Local adaptation is a commonly observed result of natural selection acting in heterogeneous environment. Common-garden experiments are a method of detecting local adaptation, as well as studying phenotypic plasticity and gradients of traits. The present study aimed to analyse reaction norms of four closely-related Iris species of section Oncocyclus and to identify a role of environmentally-specific natural selection in their plastic responses. The plant vegetative and phenological, as well as performance traits were measured in a full factorial common-garden experiment with three levels of water amount and three soil types. We found a significant effect of species identity on all traits measured. Water amount and soil type affected many of the traits, but soil type did not affect the performance. There was no significant difference in the effect of water amount and soil type on performance as reflected by rhizome growth; in other words, there was no significant genotype × environment interaction for performance. Plasticity levels and directions of response were also similar among the species. We conclude that phenotypic differences among species are of genetic origin, although no adaptive value was demonstrated for them at the time and life-stages 'frame' of this experiment.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 98 , 267–277.     

Geography of dental polymorphism in the red fox Vulpes vulpes and its evolutionary implications

Frequency distributions of dental morphotype characters (groups of morphotypes from A to S) in 43 red fox Vulpes vulpes (Linnaeus, 1758) populations from the Holarctic region were analysed. Definite but different geographical gradients in morphotype dental patterns were found both in the Palearctic and Nearctic. In the Nearctic, a great number of characters present geographical gradient related to longitude whereas, in the Palearctic, gradient relates to latitude. Mean annual temperature and mean annual sum of precipitations play a significant role in shaping the geographical variation of morphotype characters. The shape of the teeth in the centre of the morphogenetic field is more heavily geo-climatically conditioned than the morphotype characters in the teeth at the ends of the cheek teeth field. There was an attempt to reconstruct the evolutionary changes in the morphotype dental pattern in the V. vulpes line. The morphotype pattern in the red fox dentition partially follows the phylogeographical evolutionary trends in Canidae in the northern hemisphere. However, the picture of morphotype variation is influenced by both earlier episodes of human interference (reintroductions of foxes in North America) and the recent phenomenon of heavy opportunism of the species.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 61–84.     

Geographical variation in floral traits of the tree tobacco in relation to its hummingbird pollinator fauna

Nicotiana glauca , a hummingbird pollinated plant, exhibits geographical variation in several floral traits. We examined whether geographical differentiation occurred for different flower characters and if this differentiation could be explained, at least in part, by the existence or abundance of different hummingbird species in the respective pollinator assemblages. The comparison between five populations showed significant variation in six floral traits and two female fitness measures. The traits that better discriminated between populations were corolla length and corolla width. There were metric correlations between corolla length and style length in all populations studied and, in four of the five populations, both corolla length and width were also correlated. Among plants in each population, seed weight was correlated positively and significantly with style exertion, suggesting that fruit quality is dependent on the degree of cross pollination. Assemblages of hummingbirds differed between populations in species composition, visitation frequencies, and bill length. Linear regression involving bill length of the more frequent hummingbird pollinators and corolla length yielded positive and significant relationships. Thus, there appears to be an adjustment between pollinators and flowers traits that have high incidence in the among population variation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 657–667.     

A parallel geographical mosaic of morphological and behavioural aposematic traits of the newt, Cynops pyrrhogaster (Urodela: Salamandridae)

Aposematic animals advertise their unprofitability to potential predators with conspicuous coloration, occasionally in combination with other life-history traits. Theory posits that selection on functionally interrelated aposematic characters promotes the unidirectional evolution of these characters, resulting in an increase or decrease in the effectiveness of the signal. To test whether this prediction applies on a microevolutionary scale, the intra- and interpopulational variations in aposematic coloration, behaviour (which enhances the effectiveness of the coloration) and body size of newts, Cynops pyrrhogaster (Urodela: Salamandridae), were investigated. A parallel geographical mosaic of variation in aposematic coloration and behaviour among populations, independent of body size, was found. Newts on islands displayed more conspicuous aposematic traits than those on the mainland, both morphologically and behaviourally. There was no significant relationship between variation in coloration and behaviour within populations. Male newts displayed more conspicuous coloration than females. Surveys of potential predators suggest that variable natural selection at a local scale, such as predation pressure, may primarily be responsible for the microevolution of variable aposematic traits in newts.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 613–622.     

Adaptive spatiotemporal distribution of soil microfungi in 'Evolution Canyon' III, Nahal Shaharut, extreme southern Negev Desert, Israel

We examined adaptive spatiotemporal mycobiota structure in the soil of 'Evolution Canyon' III, Nahal Shaharut, in the extreme southern Negev, Israel. A total of 223 species representing 80 genera were isolated using the soil dilution plate method. The microfungal communities in all localities and seasons were characterized by a superdominance of dark-coloured species with large multicelled conidia:  Ulocladium atrum, U. botrytis, Alternaria alternata, and Al. chlamydospora . Species of the genus Aspergillus (mainly As. fumigatus ) and teleomorphic ascomycetes comprised a basic part of the thermotolerant mycobiota obtained at a temperature of 37 °C. Isolate density displayed high positive dependence on organic matter content. Density was subject to drastic spatiotemporal (especially spatial) fluctuations, with maximum levels found in the shady valley bottom locality. The lowest biodiversity indices were estimated in localities under shrubs and in the most stressful summer and spring. The results demonstrated a clear effect of harsh desert climatic and edaphic selection on the adaptive variation of the mycobiota studied.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 263–277.     

Clinal genetic variation and isolation by distance in the European eel Anguilla anguilla (L.)

The genetic variability and structure of the European eel ( Anguilla anguilla L.) in populations throughout Europe was reassessed using 15 allozymic loci, seven of which were polymorphic. Seven sites were sampled on a latitudinal gradient across the natural continental range, extending from southern France to southern Norway. Heterozygosity ( H _e = 0.05) and level of polymorphism (P = 0.43) were comparable to other marine fish. Populations were poorly differentiated ( G _ST = 0.014, F _ST = 0.002), which is not surprising considering the high dispersal capability of the European eel. However, a significant geographical cline was detected at two alleles ( IDH-1 * 100 and GPI-1 * 110 ), and genetic distances ( D _CE) were concordant with geographical coastal distances. Mantel tests, pairwise F _ST's and multidimensional scaling analyses identify three distinct groups: Northern Europe, Western Europe and the Mediterranean Sea. We propose that the clinal genetic structure in the European eel may be due to (1) isolation by distance (as recently detected with microsatellites), (2) temporal reproductive separation, (3) post-larval selective forces, (4) contact between formerly separated groups or (5) some combination thereof.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 509–521     

Soil microfungal communities of 'Evolution Canyons' in Israel – extreme differences on a regional scale

On a regional scale, we compared the local adaptive patterns of soil mycobiota revealed in four 'Evolution Canyons' located in the northern and southern parts of Israel. These microsites were chosen according to the framework of the Institute of Evolution, University of Haifa programme, focusing on the effect of interslope environmental divergence on biodiversity patterns. The comparative analysis demonstrated remarkable differences in spatiotemporal structure of the microfungal communities and their biodiversity level (species richness, heterogeneity and equitability). In the desert 'Evolution Canyon', stress-selected, slow-reproducing, dark-coloured species with large, multicelled conidia were dominant. At the same time, in the forest localities of the northern canyons, as well as in the agriculturally disturbed locality with soil degradation, ruderal-selected, fast-reproducing Penicillium species predominated. Environmental natural selection appeared to be the major factor affecting adaptive diversity patterns of soil microfungi in the studied area.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 157–163.     

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.     

Morphological differentiation among closely related species with disjunct distributions: a case study of Mediterranean Cyclamen L. subgen. Psilanthum Schwarz (Primulaceae)

In Cyclamen subgenus Psilanthum , previous authors have recognized three species ( C. repandum, C. balearicum and C. creticum ), with C. repandum subdivided into three subspecies (ssp. repandum , ssp. peloponnesiacum and ssp. rhodense ), and two varieties (var. vividum and var. peloponnesiacum ) within C. repandum ssp. peloponnesiacum . To examine the validity of this classification and the degree of morphological differentiation among closely related endemic taxa with disjunct distributions, we undertook analyses of three quantitative and eight qualitative morphological traits of flowers and leaves, most of which have previously been used without statistical comparison in the classification of the different taxa. Cyclamen balearicum showed high levels of morphological differentiation from all other taxa in subgenus Psilanthum consistent with its specific status. In contrast, morphological differentiation between C. creticum and subspecies of C. repandum was similar to that among the three subspecies of C. repandum . When compared with a recent molecular phylogeny of this subgenus, our results suggest that C. creticum may best be described as a geographical subspecies of C. repandum . The two varieties of C. repandum ssp. peloponnesiacum showed few significant differences for individual traits and their overall morphology is very similar to each other. The morphology of plants at three sites on Corsica strongly suggest, consistent with molecular data, the occurrence of hybrid populations between C. balearicum (outside of its previously recognized distribution) and local C. repandum .  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 133–144.     

16S mitochondrial sequences associate morphologically dissimilar males and females of the family Phoridae (Diptera)

The genus Phora (Diptera, Phoridae) includes more than 50 species widely distributed in temperate regions of Eurasia and Africa. Taxonomic identification of most species is based upon the morphology of the male hypopygium: hence for many species, including the type species of the genus, females are unknown. We used mitochondrial 16S ribosomal RNA sequences to match males with previously unidentified females from Phora atra , P. stictica , and P. holosericea . We then identified morphological characters that allow identification of females of P. atra and P. stictica without recourse to DNA sequencing. Our results show that small scale sequencing can aid in the development of taxonomic characters for use in the field to identify previously cryptic females. This iterative method of identifying populations genetically followed by re-examination of morphology should allow development of better keys for rapid identification of heretofore cryptic populations of insects. We also found that sequences from individual of P. holosericea from Cambridge, England and Malakhovka, Russia, were more similar to each other than to sequences from a conspecific fly also collected in Cambridge. This result suggests that there is previously un-suspected population structuring in this species.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 267–273.     

Taxonomic studies on Senecio aquaticus (Asteraceae). A recommendation for the taxonomic status of Aquaticus and Barbareifolius

Aquaticus and Barbareifolius are two subgroups of Senecio aquaticus s. l. (Asteraceae) that have been recognized at various taxonomic levels in European taxonomic literature. In this paper we present the results of morphological studies on S. aquaticus s. l. in order to establish the taxonomic status of these two subgroups. Using multivariate morphometric techniques (discriminant analysis and principal component analysis), we examined the diagnostic potential of 60 macro-morphological characters and studied the delimitation of Aquaticus and Barbareifolius. Seven of the characters studied were included in the discriminant function of the stepwise discriminant analysis and are therefore considered to be the most reliable diagnostic characters of Aquaticus and Barbareifolius. Barbareifolius has, amongst other distinguishing traits, a more divaricate inflorescence than Aquaticus. Because both subgroups are distinct but overlapping clusters in multivariate morphometric space, and because clear differences in their geographical distribution are absent, we recommend the taxonomic status of variety to both groups. Following the rules of nomenclature, Aquaticus and Barbareifolius should therefore be named S. aquaticus Hill var. aquaticus and S. aquaticus Hill var. barbareifolius (Krock.) Wimm. & Grab.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 489–498.     

Molecular and morphological variation of rare endemic oncocyclus irises (Iridaceae) of Lebanon

Oncocyclus irises endemic to Lebanon form a complex of three closely related taxa replacing each other over short geographical distances in a linear habitat. In order to characterize the appropriate taxonomic levels and to assess their conservation status, we investigated patterns of phenotypic variability and the partitioning of genetic variation within and among populations using random amplified polymorphic DNA (RAPD) markers. Multivariate analysis (principal components analysis and multiple correspondence analysis), based on 16 quantitative and six qualitative characters, revealed no separation between populations or taxa. Moreover, no morphological character could be used to define clear boundaries between populations/taxa. The genetic characterization revealed high levels of polymorphism and diversity (H_s). Principal components analysis showed population delimitations, but no groupings reflecting the currently defined taxa could be identified. Both morphological and genetic data showed that Lebanese oncocyclus irises could not be fitted into clear taxonomic boundaries. Consequences for conservation are discussed. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 123–135.     

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.     

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.     

Microsatellite diversity in populations of blind subterranean mole rats (Spalax ehrenbergi superspecies) in Israel: speciation and adaptation

The genetic diversity, structure, and relationships of 12 populations of blind subterranean mole rats belonging to four species of the Spalax ehrenbergi superspecies were examined. Diversity levels and divergence of 19 nuclear microsatellite markers were analysed. Results indicate that the 12 populations fall into two major clusters representing the species pairs Spalax galili (2 n  = 52) and S. golani (2 n  = 54) vs. S. carmeli (2 n  = 58) and S. judaei (2 n  = 60). There is very little gene flow between the species pairs, except for one population (Afiq) of S. carmeli (2 n  = 58), which is located near the hybrid zone between S. golani (2 n  = 54) and S. carmeli (2 n  = 58). Higher values of the mean for alleles per locus, gene diversity, and number of unique alleles were observed for the southern species pair, S. carmeli and S. judaei, than for the northern species pair, S. galili and S. golani . Microsatellite diversity increases southward toward the Negev Desert and is positively correlated with aridity stress. Climatic factors of water availability coupled with temperature account for 76.7% of microsatellite variance. Natural selection appears to adaptively determine microsatellite evolution in Spalax regionally across Israel.  © 2004 The Linnean Society of London , Biological Journal of the Linnean Society , 2004, 83 , 229–241.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

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.     

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.