Genetic structure of the critically endangered endemic Cochlearia polonica (Brassicaceae): efficiency of the last-chance transplantation

Cochlearia polonica , a narrow endemic of southern Poland, is one of the rarest and most endangered species of the European flora. All natural populations are extinct and the species has survived in only one transplanted population derived from 14 original individuals. Using AFLPs, the genetic variation and spatial structure of this population were analysed approximately 30 years after transplantation. The incidence of polymorphic AFLP bands (30.46%) is low compared with data from a natural population of another Cochlearia species, C. tatrae . Principal co-ordinates and spatial autocorrelation analyses demonstrated the presence of significant genetic structure. It is recommended that conservation efforts on C. polonica should preserve the complete population area, because local extinctions may lead to a loss of genetic information. The presence of genetic structure should also be taken into account during the sampling of material (plants or seeds) for ex situ conservation measures.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 527–532.     

Genetic variation in Femeniasia (Compositae, Cardueae), an endemic and endangered monotypic genus from the Balearic Islands (Spain)

Femeniasia balearica is the only representative of its genus and is endemic to a small area on the northern coast of Minorca (Balearic Islands, Spain). The entire range of the species covers only 18 km of coastline. It is extremely rare and classified as endangered in the National Catalogue of Threatened Species and as a priority species in the EU Habitats Directive. We carried out a census of current population size and estimated genetic diversity based on AFLP markers to facilitate conservation of this unique species. In the 66 individuals analysed (∼10% of population), 225 bands were scored and the level of diversity was relatively high. Three divergent population groups corresponding to geographical areas (Western, Central and Eastern) were identified. In AMOVA and Bayesian analyses, most of the diversity was found within populations but there was strong differentiation between the three population groups. Genetic and geographical distances between the populations were strongly correlated. Our results show that the populations of F. balearica are not genetically depauperate, in spite of their small sizes. Unexpectedly, our results suggest that as many as three MUs (management units) should be recognized in this very small area, because the low levels of gene flow among them indicate contemporary demographic independence.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 97–107.     

Population genetics and the conservation status of the threatened Iberian steppe grass Puccinellia pungens (Pau) Paunero (Poaceae)

Puccinellia pungens (Pau) Paunero is a narrowly endemic grass found in two continental saline lagoons of north-eastern Spain. This rare plant has been classified as 'at risk of extinction' in several national and European catalogues of endangered species. Recent demographic studies indicate that population sizes greatly exceed several million individuals, challenging that threat category. Our genetic analysis, based on allozymes, has shown that in spite of the large population sizes, very low levels of genetic variation were found in P. pungens . Genetic variation was similar in most populations, but the largest, Gallocanta lagoon as a whole, had less variation (35% polymorphic loci, 1.4 alleles/locus, H _T = 0.038) than the more restricted Royuela range (45% polymorphic loci, 1.5 alleles/locus, H _T = 0.056), suggesting a recent population expansion of the Gallocanta populations from few founder lines. The low genetic distances among populations also suggest a recent divergence. The low genetic variation observed cannot be explained fully by eventual clonal spread and rare seedling establishment in the hypersaline environment. This low variation seems to result from extreme recent population bottlenecks as a consequence of habitat conversion to agricultural fields. In the light of our data, it seems unlikely that reinforcement of populations could increase the genetic diversity of the populations. Hence, conservation efforts should focus on avoiding further habitat loss of this endangered steppe grass species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 269–281.     

Genetic diversity and structure of the cycad Zamia loddigesii Miq. (Zamiaceae): implications for evolution and conservation

The genetic diversity and structure of four populations of the cycad Zamia loddigesii were studied throughout its range in Mexico. Allozyme electrophoresis of 15 loci was conducted. The mean number of alleles per locus was 1.80 ± 0.09, the percentage of polymorphic loci was 66.6 ± 5.4, and the expected heterozygosity was 0.266 ± 0.02. The results indicated that the genetic diversity was relatively higher, with respect to tropical tree species and other cycads. The genetic variation explained by differences among populations was 18%. On average, gene flow between paired populations was similar ( Nm  = 1.6) to other tropical forest trees and cycad species. Our results indicated that the geographical isolation among populations of Z. loddigesii generated allele loss, as well as a clinal variation in the frequencies of two loci ( MDH and MNR2 ), in relation to the latitudinal distribution of populations. The populations have become fragmented due to increasingly higher pressure of habitat conversion and disturbance. The importance of the establishment of sanctuaries and protected areas and a reduction in deforestation is highlighted in this research as a way of preserving the high genetic diversity of this and other endemic species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 533–544.     

Conservation genetics of Hebe speciosa (Plantaginaceae) an endangered New Zealand shrub

All extant populations of Hebe speciosa (Plantaginaceae), a threatened endemic New Zealand shrub, were investigated using the amplified fragment length polymorphism technique (AFLP). Genetic diversity indices varied significantly among geographical regions and were positively correlated with population size. Among-population genetic differentiation was high (mean pairwise Φ_ST = 0.47), implying complex historical relationships between disjunct populations and negligible contemporary gene flow. Southern populations exhibited extremely low genetic diversity relative to those found in Northland, suggesting that these populations may be more recent in origin. Patterns of genetic relationship among some populations indicate pre-European Māori dispersal and cultivation. The three northernmost populations were found to contain the majority of the species' remaining genetic diversity and thus, should be a focus for future conservation management. Some southern sites may also be culturally significant as evidence of Māori trade and cultivation of Hebe speciosa .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 229–239.     

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.     

Structure of allozyme variation in Nordic Silene nutans (Caryophyllaceae): population size, geographical position and immigration history

We investigated allozyme variation in 34 populations of the perennial herb Silene nutans from Sweden and northern Finland, areas that were ice-covered during the last (Weichselian) glaciation. The present geographical structure of genetic variation in S. nutans in Sweden and northern Finland appears to have been mainly shaped by ancient historical processes. Patterns of variation in allele frequencies suggest two major postglacial immigration routes into Sweden, with populations entering the area from both the south and the east and forming a contact zone with admixed populations in central Sweden. While estimates of within-population genetic diversity and allelic richness are significantly correlated with present population size and geographical position (latitude), population size is not correlated with latitude. Low genetic diversity in the northern populations is more likely to have resulted from ancient stochastic events during the process of immigration than from recent population fragmentation. F _IS values are high and increase with latitude. Evidence of recent bottlenecks was detected in several southern Swedish populations: these can be interpreted in terms of population fragmentation as a result of anthropogenic disturbance. Soil pH is uncorrelated with population size and position.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 357–371.     

Extremely low levels of genetic diversity in the terrestrial orchid Epipactis thunbergii (Orchidaceae) in South Korea: implications for conservation

Levels of allozyme variation, population genetic structure, and fine-scale genetic structure (FSGS) of the rare, both sexually and clonally reproducing terrestrial orchid Epipactis thunbergii were examined for eight ( N  = 734) populations in a 20 × 20-km area in South Korea. Twenty-three putative allozyme loci resolved from 15 enzyme systems were used. Extremely low levels of allozyme variation were found within populations: the mean frequency of polymorphic loci was 3.8% [isocitrate dehydrogenase ( Idh-2 ) with two alleles was polymorphic across populations], the mean number of alleles per locus was 1.04, and the mean expected heterozygosity was 0.013. The overall fixation index was not significantly different from zero ( F _IS = 0.069), although the species is self-compatible. However, a significantly high degree of population differentiation was found between populations at Idh-2 ( F _ST = 0.388) in the studied area. Furthermore, spatial autocorrelation analyses revealed a significant FSGS (up to 3 m) within populations. These observations suggest that the main explanatory factors for the extremely low levels of genetic diversity and the shaping of the population genetic structure of E. thunbergii are genetic drift as a result of a small effective population size, a restricted gene flow, and the isolation of populations. Considering the current genetic structure of E. thunbergii , three guidelines are suggested for the development of conservation strategies for the species in South Korea: (1) protection of habitats of standing populations; (2) prohibition by law of any collection of E. thunbergii ; and (3) protection of nearby pollinator populations, given the fact that fruit set in natural habitats is very low.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 161–169.     

The genetic structure of the steppe plant Iris aphylla L. at the northern limit of its geographical range

Genetic diversity of Iris aphylla was examined in seven Polish populations located at the northern limit of the geographical range of the species. Using RAPD analysis, 491 DNA fragments were amplified, all of which were polymorphic. The peripheral populations were characterized by a low number of unique bands (one to three per population) and moderate values of genetic diversity parameters (percent of polymorphic loci = 30.6%, Shannon's index = 0.149 and Nei's gene diversity = 0.097, on average), which can be explained as being due to the recent expansion of the European biota and the extremely low frequency or absence of seed set and of juveniles, as well as limited pollen and seed dispersal. There was also no significant correlation between ecological properties (population size; number of flowering ramets and fruits) and genetic diversity parameters. Despite this fact, more fixed loci were noted in the smallest populations than were in the largest ones. All populations were significantly differentiated and geographically structured. Analysis of molecular variance between regions showed a low and nonsignificant value, but relatively high molecular variation was partitioned among and within populations. Only one group of south-eastern populations was structured according to isolation by distance pattern and neighbour-joining cluster analysis. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 245–255.     

Population structure in the catfish Trichogenes longipinnis: drift offset by asymmetrical migration in a tiny geographic range

Based on population genetic theory and empirical studies of small populations, we expect that species with very small ranges (narrow endemics) will exhibit reduced genetic diversity, increasing their susceptibility to the negative effects of genetic homogeneity. Although this pattern of reduced diversity applies to most narrow endemics, conservation biologists have yet to identify a general pattern for the degree of spatial population genetic structure expected in species with very small ranges. In part, this is because the degree of population structure within narrow endemics will be highly variable depending on the equilibrium between the homogenizing effects of dispersal and the diversifying effects of drift and local selection in small populations, thus precluding general predictions about the relative importance of small range, small population sizes, and habitat patchiness for maintaining genetic diversity in narrowly-distributed species. We document a striking example of high population structure in the tiny geographic range of a stream-dwelling catfish, Trichogenes longipinnis , endemic to the Atlantic Forest of Brazil. The maintenance of this diversity results from a combination of asymmetrical and limited dispersal, and drift in small populations. Our results highlight the need to understand population structure, and not only overall genetic diversity, of narrowly-distributed species for their conservation planning.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 259–274.     

Implications of contrasting patterns of genetic variability in two vespertilionid bats from the Indonesian archipelago

Wallacean island populations of two Vespertilioninae bats, Myotis muricola and Scotophilus kuhlii , which have similar geographical ranges, showed marked contrast in the amount and pattern of genetic variation. Within islands, genetic variation was on average much higher in M. muricola but declined from west to east, whereas all populations of S. kuhlii had uniformly low levels of genetic variation by mammalian standards. S. kuhlii showed little genetic differentiation between islands and estimates of gene flow were substantial whereas island populations of M. muricola differed markedly and there was a strong isolation-by-distance effect associated with the extent of the sea crossing between islands. Furthermore, the lower mean heterozygosity and small genetic distances between eastern island populations of M. muricola is evidence that there has been a bottleneck associated with the colonization of this area. The attenuation of genetic diversity to the east is also seen in some other mammalian species and may indicate limits to dispersal and have implications for species management. The patterns of variability in S. kuhlii may be a consequence of its strong dispersal capacity and close association with human activity, which, together with other factors, suggest a panmictic population.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 421–431.     

Population genetics of Dioon edule Lindl. (Zamiaceae, Cycadales): biogeographical and evolutionary implications

Dioon edule Lindl. (Zamiaceae) is a cycad endemic to Mexico, that occurs as one species D. edule and the geographical variety D. edule var. angustifolium (Miq.) Miq. Dioon edule has a north to south distribution in eastern Mexico. In this study, we analysed 14 allozymic loci in eight populations of D. edule from its total distribution range by sampling all known populations. Patterns of diversity and genetic variability, within and among populations, were obtained. The mean number of alleles per locus ( A ) was 1.44 and the percentage of polymorphic loci was relatively high ( P  = 54.78). The mean observed ( H _O ) and expected heterozygosity ( H _E ) were 0.27 and 0.24, respectively. F -statistics revealed an excess of heterozygous genotypes, locally and globally ( F  = −0.17 and f  = −0.27, respectively). The genetic variation explained by differences among populations was only 7.5%. We also detected a negative relationship between genetic diversity and latitude. On average, the gene flow between population pairs was relatively high ( Nm  = 2.98); furthermore, gene flow between population pairs was significantly correlated with geographical distances ( r  = −0.38, P  = 0.025). Therefore, patterns of genetic diversity in D. edule appear to be associated with the post-Pleistocene spread of the species, from its southerly (origin) to its northerly range (derived populations, including its central distribution). The biogeographical and evolutionary aspects of the results of this study are discussed. We recognize Dioon angustifolium Miq. for the northernmost disjunct populations.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 457–467     

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.     

Genetic variation within and between Iliamna corei and I. remota (Malvaceae): implications for species delimitation

Iliamna corei and I. remota are classified as endangered species; however, their designation as separate species has been questioned. In order to address this problem, intersimple sequence repeat (ISSR) data were generated to examine patterns of genetic differentiation within and between these two taxa. ISSRs were used to screen individuals for genetic diversity of I. corei from the single known natural population and two garden populations, and individuals of I. remota from the six natural populations and four garden populations. Using ten primers, 140 informative markers were generated. Ninety-four percent of loci detected revealed polymorphisms. Cluster analysis (neighbour-joining, NJ) revealed that the two species are genetically distinct and that the Illinois populations of I. remota were genetically distinct from the Virginia populations of I. remota . Ordination by Principal Coordinate Analysis (PCoA) supported the findings of the NJ, with a separation of I. corei and I. remota . Analysis of Molecular Variance (AMOVA) revealed that the majority of variation detected was within populations, which is consistent with other self-incompatible plants. The results indicate a correlation between the geographical distributions of the species and gene flow.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 345–354.     

Allozyme diversity and history of distribution expansion in the maritime perennial plant Hedyotis strigulosa (Rubiaceae), distributed over the wide latitudes in the Japanese Archipelago

Allozyme diversity was examined in 30 populations of the maritime perennial plant Hedyotis strigulosa var. parviflora , which is distributed from subtropical islands to the central mainland of Japan. Genetic diversity within populations tended to be larger in southern island populations than in northern mainland populations. In the southern part of the distribution, the population size is generally large and populations are distributed more continuously than in the northern area, resulting in the larger effective size of southern populations as a whole. These factors play a major role in maintaining greater genetic diversity in the southern populations. By contrast, genetic diversity in the northern populations is very low, probably resulting from bottlenecks of population establishments during recolonization from refugial area to the northern areas. Geographically close populations were located near each other in the multidimensional scaling and the phenogram based on genetic distances, suggesting that gene flow among remote populations is rather limited. The pattern of genetic diversity in H. strigulosa var. parviflora is likely caused by the distribution expansion of the species; in the last glacier era, the species was restricted to the southern area; its advance to the northern area is relatively recent. Another variety endemic only to the Daito Islands, H. strigulos a var. luxurians , has lower genetic diversity than H. strigulosa var. parviflora and has genetically diverged from H. strigulosa var. parviflor a.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 679–688.     

Genetic differentiation on multiple spatial scales in an ecotype-forming marine snail with limited dispersal: Littorina saxatilis

The population genetic structure of marine species lacking free-swimming larvae is expected to be strongly affected by random genetic drift among populations, resulting in genetic isolation by geographical distance. At the same time, ecological separation over microhabitats followed by direct selection on those parts of the genome that affect adaptation might also be strong. Here, we address the question of how the relative importance of stochastic vs. selective structuring forces varies at different geographical scales. We use microsatellite DNA and allozyme data from samples of the marine rocky shore snail Littorina saxatilis over distance scales ranging from metres to 1000 km, and we show that genetic drift is the most important structuring evolutionary force at distances > 1 km. On smaller geographical scales (< 1 km), divergent selection between contrasting habitats affects population genetic structure by impeding gene flow over microhabitat borders (microsatellite structure), or by directly favouring specific alleles of selected loci (allozyme structure). The results suggest that evolutionary drivers of population genetic structure cannot a priori be assumed to be equally important at different geographical scales. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 31–40.     

Low population differentiation and high genetic diversity in the invasive species Carduus acanthoides L. (Asteraceae) within its native range in the Czech Republic

Colonizing species are predicted to suffer from reductions in genetic diversity during founding events. Although there is no unique mode of reproduction that is characteristic of successful plant colonizers, many of them are predominantly self-fertilizing or apomictic species, and almost all outcrossing colonizers are self-compatible. Carduus acanthoides comprises a species of disturbed habitats with wind-dispersed seeds that colonizes open spaces of various sizes. Population genetic diversity was expressed by assessing patterns of variation at nine putatively neutral allozyme loci within and among 20 natural populations in its native distribution range in the Czech Republic. Overall, C. acanthoides displayed high levels of genetic diversity compared to other herbaceous plants. The percentage of polymorphic loci was 84.5, with values of 2.37, 0.330, and 0.364 for the mean number of alleles per polymorphic locus ( A ), observed heterozygosity ( H _o), and expected heterozygosity ( H _e), respectively. There was only weak evidence of inbreeding within populations ( f  = 0.097) and very low genetic differentiation among populations ( θ  = 0.085). Analyses of the data provide strong evidence for isolation-by-distance for the whole study area. Even the colonizing species, C. acanthoides , currently supports a substantial amount of allozyme variation at both the species and population levels.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 596–607.     

Reduced genetic variation and strong genetic population structure in the freshwater killifish Valencia letourneuxi (Valenciidae) based on nuclear and mitochondrial markers

The genetic variation of the critically endangered Corfu killifish (Valencia letourneuxi), an endemic freshwater fish species of the western Balkans, was assessed for nine populations sampled in eight water systems in western continental Greece, the Peloponnese and the Ionian Island of Corfu, using mitochondrial and microsatellite markers. The analyses were based on data from three mtDNA regions (D‐loop, COI and 16S rRNA sequences) and 14 microsatellite loci. Samples from the congeneric species Valencia hispanica and the phylogenetically closely related species Aphanius fasciatus were also used in the study as outgroups. Both the mitochondrial and the microsatellite analyses revealed three distinct population groupings associated with the geographical distribution of the populations: one southern group occupying rivers draining to the Patraikos Gulf, the second one including the populations flowing into the Amvrakikos Gulf and the third, more northern group, including the other populations from rivers in Corfu Island and Epirus flowing into the Ionian Sea. Within these groupings there is limited genetic differentiation between populations; in addition, there is reduced intrapopulation genetic variation, evidenced by low heterozygosity values, number of alleles and haplotype diversity. In terms of taxonomic implications and appropriate management actions for conservation, our data suggest that the major population groups should be regarded at least as three distinct conservation units (CUs), with translocation and restocking actions to take place only within the geographical range of the CU concerned. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 334–349.     

Allozyme variation and population structure of the very narrow endemic Seseli farrenyi (Apiaceae)

Analyses have been made of allozyme variation of the narrow endemic species Seseli farrenyi Molero & J. Pujadas (Apiaceae), which has only three known populations in Catalonia with a total of around 2000 individuals. All three populations were sampled and subjected to starch gel electrophoresis. Nine enzymes were resolved and 14 loci were interpreted. We detected high values of polymorphism ( P = 83.3%, A = 3.0, H _e = 0.297), far exceeding those expected for endemic species ( P = 26.3%, A = 1.39, H _e = 0.063). Genetic diversity was greater within populations than among populations, and the value of gene flow was very high ( Nm = 5.85). Most loci showed deviations from Hardy– Weinberg equilibrium, possibly due to the presence of subpopulations. The main threats to this species are human activities (tourism, fires), while natural threats are minimal due to its high genetic diversity. Finally, we propose some conservation measures which include both in situ and ex situ strategies. © 2000 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 305–314.     

Genetic structure and differentiation of Gentiana atuntsiensis W. W. Smith and G. striolata T. N. Ho (Gentianaceae) as revealed by ISSR markers

The Hengduan Mountains are the core region of the Himalaya hotspot, and are renowned for their high levels of endemism. Gentiana atuntsiensis and G. striolata are two closely related and morphologically similar species endemic to this region. In this study, inter-simple sequence repeat (ISSR) markers were used to investigate the genetic structure within these two species, as well as the differentiation between them. An analysis of molecular variance-derived estimate demonstrated only 13.5% of genetic differentiation between the two species. Considering their adjacent distribution patterns, low genetic divergence, and clear clustering into two groups, it is hypothesized that the two species arose from a rapid and recent speciation event induced mainly by geographical isolation. A relatively high level of genetic diversity was revealed in each species (Shannon's index of diversity: H _sp = 0.324 and 0.391; H _pop = 0.225 and 0.274; for G. atuntsiensis and G. striolata , respectively). Most of the genetic variation was partitioned within populations ( Ø _ST = 0.232 and 0.226 in G. atuntsiensis and G. striolata , respectively). The large population sizes, outcrossing breeding system, and small, light seeds that disperse widely may explain the genetic structure in both species.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 225–232.