A study of Draba cacuminum (Brassicaceae)

A morphological analysis of the Scandinavian mountain endemic Draba cacuminum Elis. Ekman is presented, based on Scandinavian herbarium collections and a population sample from the Finse area, S Norway. It is compared with supposedly close relatives, especially D. norvegica Gunnerus, and is found to be a distinct species with infraspecies variation at two levels. The populations of southern and northern Scandinavia have been separated as subspecies, ssp. cacuminum and ssp. angusticarpa Elven ssp. nov., respectively, and there is also variation within the southern subspecies. A lectotype has been chosen for D. cacuminum , a revised distribution map is presented, and the ecology and phytogeography of the species is discussed. It seems to be a weak competitor, and its present very discontinuous area is assumed to be a remnant of one continuous or more probably two areas (in S Norway and N Scandinavia) in the late Weichselian.     

Parallel ecoclinal evolution and taxonomy of F'rankenia (Frankeniaceae) in the Cape Verde Islands, W Africa

One species, the Macaronesian and N African Frankenia ericifolia , with three subspecies is recognized in the Cape Verde Islands: ssp. ericifolia (a coastal, xerophytic ecotype), ssp. caboverdeana ssp. nov. (a coastal, mesophytic ecotype), and ssp. montana ssp. nov. (a montane, hygrophytic ecotype). Subspecies caboverdeana and ssp. montana are endemic to the Cape Verde Islands. In addition, ssp. latifolia comb. et stat. nov. is tentatively recognized as endemic to the Canary Islands. Reproductive data suggest that the Capeverdean plants are self-compatible but mainly outcrossing. Mor-phometric analyses of field-collected material and progeny families cultivated under uniform conditions demonstrate that there is genetically based, clinal morphological variation connecting ssp. ericifolia and ssp. caboverdeana in several coastal areas that vary in humidity conditions because of varying exposures to the trade wind. Clinal variation was observed in leaf-, calyx-, petal-, and seed characters, probably caused by one or a few genetic factors that control organ size and are subjected to differential selection with respect to drought tolerance. The variation in progeny families derived from morphologically intermediate parents was no higher than the variation in progeny families derived from parents belonging to the "pure" subspecies. It is thus likely that the clinal variation is caused by primary evolution along continuous humidity gradients rather than hybridization. The existence of similar ecoclines in different islands and the potentially simple genetic basis of such clinality suggest that ssp. caboverdeana has evolved several times in mesic environments from xerophytic, ericifolia-like ancestors.     

Multiple origins of the octoploid Scandinavian endemic Draba cacu minum: electrophoretic and morphological evidence

The octoploid Draba cacuminum (Brassicaceae) is one of the few endemic species in Scandinavia and has frequently been considered an example supporting the glacial survival theory. Two subspecies, ssp. cacuminum in southern Norway and ssp. angusticarpa in northern Scandinavia, have been described and suggested to be geographically diverged remnants of a more continuously distributed ancestor. To test an alternative hypothesis of independent origins of similar octoploids in different areas, we investigated populations of D. cacuminum and its possible progenitors using enzyme electrophoresis and morphological analysis. Electrophoretic analysis of progeny of D. cacuminum revealed high levels of fixed heterozygosity, suggesting that the species is a genetic alloploid. Fixed electrophoretic differences among the populations and additivity of electrophoretic phenotypes indicate that the octoploid D. cacuminum has originated at least three times by alloploidizations involving different populations of the hexaploid D. norvegica and a diploid species, possibly D. fladnizensis. Electrophoretic and morphometric data suggest that populations of D. cacuminum with broad siliculae have originated from populations of D. norvegica that had broad siliculae, and that populations of D. cacuminum with narrow siliculae have originated from populations of D. norvegica that had narrow siliculae. However, the electrophoretic and morphometric variation within D. cacuminum did not correspond to the geographic origin of the populations, and the previously described subspecies could not be recognized. Draba cacuminum gives no relevant information on the glacial survival theory; the polyploidizations may have occurred in postglacial time as well as in pre-Weichselian periods.     

Allozyme and morphological variation in two subspecies of Dryas octopetala (Rosaceae)in Alaska

The Alaskan endemic shrub Dryas octopetala ssp. alaskensis and its circumpolar conspecific ssp. octopetala are adapted to closely adjacent habitats in alpine areas of Alaska. These alpine areas form geographically disjunct "islands" among which there are limited opportunities for gene flow. Allozyme electrophoresis and a common garden experiment were used to examine genetic variation between subspecies and among disjunct populations of each subspecies. Overall, allozyme variation in D. octopetala is low with little differentiation among populations or between subspecies. Morphological differences, however, are greater between subspecies than among populations within subspecies. Divergence for a few morphological and life-history characters has apparently occurred in response to strong selection, but without divergence at allozyme loci. The ancestors of both subspecies of D. octopetala in Alaska were isolated during the Pleistocene in the glacial refugia of Alaska and Yukon, which may explain low overall variation. Dryas. o. alaskensis is thought to be a Pleistocene derivative of ssp. octopetala, which may account for the low allozyme divergence between subspecies. Recent restriction to alpine areas may explain the low differentiation among disjunct populations.     

Ecological divergence and speciation in common bottlenose dolphins in the western South Atlantic

Coastal and offshore ecotypes of common bottlenose dolphins have been recognized in the western South Atlantic, and it is possible that trophic niche divergence associated with social interactions is leading them to genetic and phenotypic differentiation. The significant morphological differentiation observed between these ecotypes suggests they represent two different subspecies. However, there is still a need to investigate whether there is congruence between morphological and genetic data to rule out the possibility of ecophenotypic variation accompanied by gene flow. Mitochondrial DNA (mtDNA) control region sequence data and 10 microsatellite loci collected from stranded and biopsied dolphins sampled in coastal and offshore waters of Brazil as well as 106 skulls for morphological analyses were used to determine whether the morphological differentiation was supported by genetic differentiation. There was congruence among the data sets, reinforcing the presence of two distinct ecotypes. The divergence may be relatively recent, however, given the moderate values of mtDNA nucleotide divergence (dA = 0.008), presence of one shared mtDNA haplotype and possibly low levels of gene flow (around 1% of migrants per generation). Results suggest the ecotypes may be in the process of speciation and reinforce they are best described as two different subspecies until the degree of nuclear genetic divergence is thoroughly evaluated: Tursiops truncatus gephyreus (coastal ecotype) and T. t. truncatus (offshore ecotype). The endemic distribution of T. t. gephyreus in the western South Atlantic and number of anthropogenic threats in the area reinforces the importance of protecting this ecotype and its habitat.     

Genetic structure and differentiation of <Emphasis Type="Italic">Oryza sativa</Emphasis> L. in China revealed by microsatellites

China is one of the largest centers of genetic diversity of Oryza sativa L. in the world. Using a genetically representative primary core collection of 3,024 rice landraces in China, we analyzed the genetic structure and intraspecific differentiation of O. sativa, and the directional evolution of SSR. The genetic structure was investigated by model-based structure analysis and construction of neighbor-joining phylogenetic tree. Comparison between genetic structure and predefined populations according to Ting’s taxonomic system revealed a hierarchical genetic structure: two distinct subspecies, each with three ecotypes and different numbers of geo-ecogroups within each ecotype. Two subspecies evidently resulted from adaptation to different environments. The different cropping systems imposed on the subspecies led to further differentiation, but the variation within each subspecies resulted from different causes. Indica, under tropical-like or lowland-like environments, exhibited clear differentiation among seasonal ecotypes, but not among soil-watery ecotypes; and japonica showed clear differences between soil water regime ecotypes, but not among seasonal ecotypes. Chinese cultivated rice took on evident directional evolution in microsatellite allele size at several aspects, such as subspecies and geographical populations. Japonica has smaller allele sizes than indica, and this may partly be the result of their different domestication times. Allele size was also negatively correlated with latitude and altitude, and this may be interpreted by different mutation rates, selection pressures, and population size effects under different environments and cropping systems.     

Patterns of allozyme and morphological differentiation in the Carex flava complex (Cyperaceae) in Fennoscandiaxs

Allozyme and morphological differentiation patterns within the Carex flava complex in Fennoscandia were described by means of multivariate analyses. As a result of these studies, and considering also data on ecology, distribution and hybridization, the following taxa were recognized: C. flava with no infraspecific taxa, C. lepidocarpa with two subspecies, ssp. lepidocarpa and ssp. jemtlandica, C. demissa with no infraspecific taxa, and C. oederi with three varieties, var. oederi , var. bergrothii , and var. pulchella. Carex lepidocarpa ssp. jemtlandica was allozymically divergent from ssp. lepidocarpa , indicating a long period of isolation of the two subspecies. A poor morphological separation in some localities is apparently due to secondary contacts in connection with recolonization of Fennoscandia after the last ice age, resulting in the formation of fertile hybrids and back-crosses. The subdivision of C. oederi into three varieties is supported by ecological data and it is suggested that they have originated by parallel ecotype formation under similar selective regimes in different areas. However, the varietal limits are not supported by differentiation in neutral (allozyme) characters. A phylogenetic hypothesis was constructed by means of neighbour-joining based allozyme data and it was indicated that C. demissa may be more closely related to C. lepidocarpa than to C. oederi. Previous treatments that have reduced the C. flava complex to one or two species may have been based on inappropriate species concepts, too coarse methods of analysis of morphological data, ignorance of patterns of co-occurrence and hybridization, or a combination of these factors.     

Differentiation patterns in the Scutellaria albida group (Lamiaceae) in the Aegean area

An experimental investigation of the mainly white-flowered Scutellaria albida group in the Aegean area was carried out. Eight populations representing maximum morphological variation as well as geographic separation were chosen for crosses between populations. Strong sterility barriers were observed in the Cretean endemic species, S. sieberi Benth., which also deviated morphologically and is probably an ancient relic. Strong crossing barriers were also found in plants from Euboea, and, in spite of little morphological differentiation from S. albida L. s. str., they are treated as a separate species, S. goulimyi Rech. fil. Crossing barriers of intermediate strength were present in morphologically distinct material from N Sporades, which is also treated as a separate species, S. sporadum Bothmer sp. nov. All other populations showed high compatibility in crosses among themselves, viz. populations from Bulgaria, Thrace, Andros, Naxos, and Rodhos. These were treated as the same species, S. albida , having large local as well as clinal variation. Three subspecies are retained: ssp. albida in the north; ssp. perhispida (Bornm.) Bothmer in E Macedonia, Thrace and on the islands; and the mainly purple-flowered, ssp. vacillans (Rech. fil.) Bothmer, endemic to the peninsula of Athos. Evolutionary pathways and phytogeographical patterns are discussed.
S. sporadum is mainly inbreeding and, in S. sieberi , indications of a self-incompatibility system were found. The other taxa have a more versatile system.     

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.     

Experimental studies of adaptation in Clarkia xantiana. I. Sources of trait variation across a subspecies border

Abstract Both genetic differentiation and phenotypic plasticity might be expected to affect the location of geographic range limits. Co‐gradient variation (CoGV), plasticity that is congruent with genetic differentiation, may enhance performance at range margins, whereas its opposite, counter‐gradient variation (CnGV) may hinder performance. Here we report findings of reciprocal transplant experiments intended to tease apart the roles of differentiation and plasticity in producing phenotypic variation across a geographic border between two plant subspecies. Clarkia xantiana ssp. xantiana and C. xantiana ssp. parviflora are California‐endemic annuals that replace each other along a west‐east gradient of declining precipitation. We analyzed variation in floral traits, phenological traits, and vegetative morphological and developmental traits by sowing seeds of 18 populations (six of ssp. xantiana and 12 of ssp. parviflora) at three sites (one in each subspecies' exclusive range and one in the subspecies' contact zone), in two growing seasons (an exceptionally wet El Niño winter and a much drier La Niña winter). Significant genetic differences between subspecies appeared in 11 of 12 traits, and differences were of the same sign as in nature. These findings are consistent with the hypothesis that selection is responsible for subspecies differences. Geographic variation within subspecies over part of the spatial gradient mirrored between‐subspecies differences present at a larger scale. All traits showed significant plasticity in response to spatial and temporal environmental variation. Plasticity patterns ranged from spatial and temporal CoGV (e.g., in node of first flower), to spatial CnGV (e.g., in flowering time), to patterns that were neither CoGV nor CnGV (the majority of traits). Instances of CoGV may reflect adaptive plasticity and may serve to increase performance under year‐to‐year environmental variation and at sites near the subspecies border. However, the presence of spatial CnGV in some critical traits suggests that subspecies ranges may also be constrained by patterns of plasticity.     

A comparative study of allozyme variation of peripheral and central populations of Silene nutans L. (Caryophyllaceae) from Western Europe: implications for conservation

In Belgium, at the western border of its continental distribution range, the perennial herb S. nutans has evolved two parapatric ecotypes (calcicolous or silicicolous), which show contrasted allozyme, morphological and adaptive patterns, and isolating reproductive barriers. We examined allozyme variation in 21 populations close to Belgium (from France and Luxemburg) in relation to their peripheral or central geographical position, and investigated their genetic relationships with the Belgian ecotypes. Both peripheral and central populations showed high genetic variation. They were differentiated from each other, but not in relation to the substrate (calcareous or siliceous). The peripheral and central populations were related to the Belgian silicicolous and calcicolous ecotype, respectively. This suggests an ancient differentiation and different past histories for the Belgian ecotypes, and parapatric speciation processes. This study exemplifies the high evolutionary potential of populations at range peripheries and the need for considering them in conservation strategies.     

Population differentiation within Festuca rubra L. with regard to soil salinity and soil water

Summary Seed and transplanted adult plants from populations of Festuca rubra, collected from inland, salt-marsh and sand-dune sites were grown on culture solution with added sodium chloride. The growth of the populations of the three habitats was reduced differentially by salt. The salt marsh ecotype Festuca rubra ssp. litoralis was only slightly affected and the inland ecotype F. rubra ssp. rubra was severely retarded at 60 mM NaCl. The dune ecotype F. rubra ssp. arenaria had an intermediate tolerance. The tolerant ecotypes accumulated less sodium chloride as compared to the sensitive ecotype, suggesting that salt tolerance is caused in part by salt exclusion.In addition, the dune ecotype F.r. arenaria appeared to be more drought tolerant than the salt marsh ecotype. Abscission of salt-saturated leaves does not function as an adaptation to salinity in Festuca rubra.All three ecotypes accumulated proline with increased salinity. The response was most pronounced in the drought tolerant F.r. arenaria, indicating that proline accumulation is a response to osmotic stress rather than to ion-specific effects of salinity. The observed differences in salt tolerance may be explained by differential sensitivity to toxic effects of sodium chloride.The occurrence on a beach plain of closely adjacent populations of F.r. arenaria and F.r. litoralis, differing markedly in salt tolerance, is briefly discussed.     

Notes on the Chenopodiaceae of Ethiopia, Somalia and Southern Arabia

Atriplex griffithii is divided into two subspecies, ssp. griffithii and ssp. stocksii , stat. nov. Atriplex farinosa is typified and divided into two subspecies, ssp. farinosa and ssp. keniensis , stat. nov. A new species of Atriplex, A. nogalensis is described from Somalia. The subtribe Sevadinae is revised and found to consist of the two monotypic genera Sevada and Fadenia and the genus Lagenantha with two species, L. cycloptera , comb. nov. (which includes the types of Gyroptera and Cycloptera) , and L. gillettii , comb. nov.     

Taxonomic Studies on the Nocturnal Flowering Group of Hemerocallis (Liliaceae) from China

The nocturnal-flowering group of Hemerocallis was treated as including three separate species, H．lilioasphodelus,H.citrina and H．minor in the Florae Reipublicae Popularis Sinicae Vol．14(1980)．However it is difficult to distinguish them from one another by a single feature,such as the flower number,length of perianth tube,diameter of tuberous roots,for they are strongly overlapping．In the present paper,numerical analysis was made of nine morphological characters and karyotype divergences which are of taxonomic significance as shown in the study of H． fulva and its two varieties (Xiong and Chen,1992),based on dry and living materials from China．In addition,geographical distribution pattern of the three taxa studied shows that they seem to be different geographical races or allopatric ecotypes in a topocline,since they inhabit a climate gradient from warm and moist to cold and drought condition．As a result,the present authors tend to consider it more reasonable to treat them as a single species,H．lilioasphodelus,with three subspecies,that is, H．lilioasphodelus ssp．lilioasphodelus,H．lilioasphodelus ssp． citrina (stat．nov) and H．lilioasphodelus ssp．minor(stat．nov．)     

Environmental conditions and genetic differentiation: what drives the divergence of coexisting Leymus chinensis ecotypes in a large-scale longitudinal gradient?

Aims The two coexisting Leymus chinensis ecotypes exhibit remarkable divergences in adaptive strategies under drought and salinity in semi-humid meadows and semi-arid steppes. In order to detect the major genetic and environmental factors dominating the intraspecific phenotype variations and ecotype formation, the questions regarding the two distinct phenotypic forms (ecotypes) in L. chinensis were addressed: (i) did environments drive the L. chinensis ecotype formation? (ii) was there a molecular basis for the morphological divergence between the two ecotypes? (iii) which driving force dominated the intraspecies divergence, divergent natural selection, genetic drift or stabilizing selection?Methods We applied a series experiments on demographical, morphological and physiological traits of two Leymus chinensis ecotypes with gray green (GG) and yellow green (YG) leaf color in nine wild sites along a longitudinal gradient from 114° to 124°E in northeast China. The environmental data including mean annual precipitation, mean annual temperature, elevation and soil properties were collected. We compared the differences of morphological, physiological and genetic differentiations between the two ecotypes.Important findings The GG type exhibited stronger fitness than YG type from the population densities, morphological traits (e.g. shoot height, leaf area, leaf and seed weights et al.), leaf mass per area (LMA) and physiological traits [relative water content (RWC), proline, soluble sugar contents]. Most of above phenotypes (e.g. total shoot densities, spike length et al.) were significantly correlated with mean annual precipitation, mean annual temperature and soil water content (SWC), rarely a correlated with soil pH and soil nutrient. Transplanted populations showed convergence trend by their leaf chlorophyll contents and osmotic adjustments (proline and soluble sugar contents) in the greenhouse, but still exhibited their divergences between two ecotypes in the outdoor transplantation, suggesting that whether L. chinensis ecotype differentiated could be largely affected by the environmental conditions. Furthermore, by the comparison result of quantitative genetic variation (Q ST) values from phenotypes with theoretical neutral genetic differentiation (F ST), differentiation in phenotypic traits greatly surpassed neutral predictions, implying that directional natural selection played a crucial role in L. chinensis ecotype differentiation. In addition, microsatellite analysis from Neighbor-joining tree and Bayesian assignment generated into two groups according to ecotypes, indicating molecular genetic differentiation also propelled the two ecotypes divergence. We conclude that L. chinensis population variations were driven by combing divergent natural selection (precipitation, temperature and SWCs) along the large-scale gradient and significantly intrinsic genetic differentiation.     

MULTIVARIATE ANALYSIS OF SELECTION IN EXPERIMENTAL POPULATIONS DERIVED FROM HYBRIDIZATION OF TWO ECOTYPES OF THE ANNUAL PLANT DIODIA TERES W. (RUBIACEAE)

Morphologically variable F₂ genotypes derived from hybridization of coastal and inland ecotypes of the annual plant Diodia teres were used to identify selection on morphological traits in the natural habitat of each ecotype. These ecotypes occur in very different habitats, and have evolved pronounced morphological differentiation. Selection analysis can suggest whether present patterns of selection can explain morphological differences between ecotypes. F₂ genotypes were characterized morphologically, clonally replicated, and transplanted into the habitat of each ecotype. Selection was measured on six morphological traits. Directional and stabilizing selection occurred on many traits; direction and strength of selection varied sharply at different stages of growth, as revealed by a path-analysis approach that divided selection into a set of independent components. Directional selection favored traits of the native population at the coastal habitat, but less so at the inland habitat. Selection was of sufficient strength to create the observed morphological differences between ecotypes in 25–100 generations, given constant selection and sufficient genetic variation. In effects on fitness, most traits were neither independent nor consistently interactive with other traits. Rather, many traits entered into strong but evanescent interactions affecting particular components of fitness. Observed interactions did not support the hypothesis that the morphology of each ecotype was functionally integrated to a high degree.     

Genetic (RAPD) Diversity between Oleria onega agarista and Oleria onega ssp. (Ithomiinae, Nymphalidae, Lepidoptera) in North-Eastern Peru

Oleria onega agarista Felder and Felder and Oleria onega ssp. nov. are two Ithomiinae subspecies from north-eastern Peru, that differ for some morphological and behavioural traits. Two contact zones are known near the town of Tarapoto: Ahuashiyacu, where both subspecies cohabit but do not seem to hybridise, and Estero (near the village of Shapaja), where they apparently hybridise. Genetic differences between the two subspecies and between populations were investigated with random amplified polymorphic DNA (RAPD) markers. Both Cluster and Principal Coordinates Analyses (CCoA and PCoA) performed using these data, provided a clear but weak discrimination between the two subspecies. Genetic diversity is much higher within the populations than between them. Moreover, the geographically more distant populations are grouped together by the genetic data. Morphological traits on the wing patterns of the hybrids are intermediary between the two butterflies subspecies, while RAPDs data place them closer to O. onega agarista than to O. onega ssp. The individuals of the Ahuashiyacu population are clearly separated into two groups, those of O. onega ssp. and O. onega agarista, by both morphology and RAPDs data. Moreover, none of those individuals show RAPD similarity with the hybrids, suggesting that hybridisation has not occurred in this population.     

Mismatch in the distribution of floral ecotypes and pollinators: insights into the evolution of sexually deceptive orchids

Plants are predicted to show floral adaptation to geographic variation in the most effective pollinator, potentially leading to reproductive isolation and genetic divergence. Many sexually deceptive orchids attract just a single pollinator species, limiting opportunities to experimentally investigate pollinator switching. Here, we investigate Drakaea concolor, which attracts two pollinator species. Using pollinator choice tests, we detected two morphologically similar ecotypes within D. concolor. The common ecotype only attracted Zaspilothynnus gilesi, whereas the rare ecotype also attracted an undescribed species of Pogonothynnus. The rare ecotype occurred at populations nested within the distribution of the common ecotype, with no evidence of ecotypes occurring sympatrically. Surveying for pollinators at over 100 sites revealed that ecotype identity was not correlated with wasp availability, with most orchid populations only attracting the rare Z. gilesi. Using microsatellite markers, genetic differentiation among populations was very low (G_ST = 0.011) regardless of ecotype, suggestive of frequent gene flow. Taken together, these results may indicate that the ability to attract Pogonothynnus has evolved recently, but this ecotype is yet to spread. The nested distribution of ecotypes, rather than the more typical formation of ecotypes in allopatry, illustrates that in sexually deceptive orchids, pollinator switching could occur throughout a species' range, resulting from multiple potentially suitable but unexploited pollinators occurring in sympatry. This unusual case of sympatric pollinators highlights D. concolor as a promising study system for further understanding the process of pollinator switching from ecological, chemical and genetic perspectives.     

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.     

Investigating the extent of parallelism in morphological and genomic divergence among lake trout ecotypes in Lake Superior

Understanding the emergence of species through the process of ecological speciation is a central question in evolutionary biology which also has implications for conservation and management. Lake trout (Salvelinus namaycush) is renowned for the occurrence of different ecotypes linked to resource and habitat use throughout North America. We aimed to unravel the fine genetic structure of the four lake trout ecotypes in Lake Superior. A total of 486 individuals from four sites were genotyped at 6822 filtered SNPs using RADseq technology. Our results revealed different extent of morphological and genetic differentiation within the different sites. Overall, genetic differentiation was weak but significant and was on average three times higher between sites (mean F_ST = 0.016) than between ecotypes within sites (mean F_ST = 0.005) indicating higher level of gene flow or a more recent shared ancestor between ecotypes within each site than between populations of the same ecotype. Evidence of divergent selection was also found between ecotypes and/or in association with morphological variation. Outlier loci found in genes related to lipid metabolism and visual acuity were of particular interest in this context of ecotypic divergence. However, we did not find clear indication of parallelism at the genomic level, despite the presence of phenotypic parallelism among some ecotypes from different sampling sites. Overall, the occurrence of different levels of both genomic and phenotypic differentiation between ecotypes within each site with several differentiated loci linked to relevant biological functions supports the presence of a continuum of divergence in lake trout.