A comparison of levels of genetic polymorphism and self-compatibility in geographically restricted and widespread plant congeners

Summary Evolutionary theory predicts low levels of genetic polymorphism and high levels of self-compatibility in plant species with small ranges and few individuals. To test these predictions, I compared published data on electrophoretically detectable genetic variation and breeding systems for geographically restricted and widespread congeners in eleven genera. The restricted species exhibit significantly fewer polymorphic loci and alleles per polymorphic locus than do their widespread congeners. Although some rare species are genetically impoverished, others are nearly as polymorphic as their widespread congeners. The restricted species and their widespread congeners do not differ consistently with respect to breeding systems.     

Conservation genetics of the rare and endangered Leucopogon obtectus (Ericaceae)

Leucopogon obtectus Benth. is a declared rare species found in the kwongan vegetation in Western Australia. Plants on a mineral sand mine and the rehabilitation area are subject to disturbance. Genetic diversity was examined within and among all known populations using random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphisms (AFLPs) for conservation. Both molecular markers revealed a high percentage (> 89%) of polymorphic markers and a high mean genetic distance among individuals (D = 0.3). Analysis of molecular variance showed that 86.7% (RAPD) and 89.7% (AFLP) of variability was partitioned among individuals within populations. Exact tests showed no significant population differentiation. The analyses indicated that L. obtectus exhibits high levels of genetic diversity despite small population sizes. The high levels of variability among individuals and the lack of clear population differentiation suggest that this species comprises a single, genetically diverse group. Conservation and management of L. obtectus should concentrate on maintaining the high levels of genetic variability through mixing genotypes and promoting outcrossing.     

安徽羽叶报春与毛茛叶报春的遗传多样性研究与新资源评估

采用RAPD方法对分布于安徽的珍稀濒危植物安徽羽叶报春(Primula merrilliana Schltr.)和毛茛叶报春(P．cicutariifolia Pax)的10个居群进行了DNA基因组多态性分析,从50条10碱基随机引物中筛选出14条能扩增出稳定性和重复性好的DNA片段的引物,共扩增出85条带,其中多态性带67条,占总数的78．8％。安徽羽叶报春与毛茛叶报春的多态位点分别达到71．6％与66．7％,因而推测主要致濒因素可能不是通常认为的低水平的遗传多样性。采用N—J法对遗传距离构建的树状图进行聚类分析,从分子角度证明了安徽羽叶报春与毛茛叶报春是2个自然的物种,同时也说明了它们的遗传多样性与生境多样性的密切相关性。由于这2个物种具有地处黄山风景区的地理优势和花朵多、花期长、花姿优美等观赏性特点以及遗传优势,如加以合理的开发与利用,可以作为观赏花卉新资源。     

Characterization of polymorphic microsatellite loci in Haageocereus (Trichocereeae, Cactaceae)

? Premise of the study: Microsatellites were isolated from two species of the genus Haageocereus (H. tenuis and H. pseudomelanostele) to be applied in studies of genetic diversity and population structure. ? Methods and Results: Five loci were employed in a preliminary study of genetic diversity and population differentiation in two rare (H. tenuis and H. repens) and two widespread (H. acranthus and H. pseudomelanostele) species, yielding between one and 44 alleles per locus. All five loci were polymorphic, with overall levels of observed and expected heterozygosities ranging from 0.478 to 0.871 and from 0.564 to 0.956, respectively. Three additional loci were scored in H. pseudomelanostele. These eight plus the remaining 11 loci were amplified from putative parents of three hybrids involving Haageocereus and Espostoa. ? Conclusions: These markers will facilitate analysis of genetic diversity, hybridization, and population differentiation throughout Haageocereus and Espostoa.     

Genetic Diversity in Cymophyllus fraserianus (Cyperaceae), a Rare Monotypic Genus

Cymophyllus fraserianus (Fraser's sedge) is a rare perennial evergreen herb found in late-successional forest communities in the Appalachian Mountains of the U.S. Genetic diversity was assessed at 19 allozyme loci for 12 populations sampled from the southern and central portions of the range of this primitive monotypic genus. Species variation was comparable to that found for other narrowly distributed species, although mean population genetic diversity was somewhat higher. Sixteen of the 19 loci (84%) were polymorphic, with a mean of 46% polymorphic within populations. On average, 2.7 alleles were detected per polymorphic locus. Genetic diversity within populations (Hep) averaged 0.113, whereas species' genetic diversity (Hes) was 0.131. Although a twofold range of within-population genetic diversity (Hep = 0.075-0.158) was found, less than 13% of the genetic variation was distributed among populations (G(ST) = 0.129). An indirect estimate of historical levels of gene flow (Nm = 1.69) was consistent with the high mean genetic identity (mean I = 0.96) found between populations. Despite the relictual nature of C. fraserianus, its rarity and fragmented distribution, genetic diversity within the genus is typical of herbs with similar life-history characteristics. No association was found between geographic and genetic distances between populations (r = -0.01; p > 0.9), suggesting that present day populations are relatively isolated.     

Population genetic structure of wild Phaseolus lunatus (Fabaceae), with special reference to population sizes

To set up an in situ conservation strategy for Phaseolus lunatus, we analyzed the genetic structure of 29 populations in the Central Valley of Costa Rica. Using 22 enzyme loci, we quantified the proportion of polymorphic loci (P(p)), the mean number of alleles per locus (A), and the mean effective number of alleles per locus (A(e)), which equaled to 10.32%, 1.10, and 1.05, respectively. The total heterozygosity (H(T)), the intrapopulation genetic diversity (H(S)), and the interpopulation genetic diversity (D(ST)) were 0.193, 0.082, and 0.111, respectively. The genotypic composition of the analyzed populations showed a deviation from the Hardy-Weinberg proportions (F(IT) = 0.932). This disequilibrium was due to either genetic differentiation between populations (F(ST) = 0.497) or nonrandom mating within populations (F(IS) = 0.866). From the level of genetic differentiation between populations and the private alleles frequencies estimates, gene flow was calculated: Nm(W) = 0.398 and Nm(S) = 0.023, respectively. The results suggested that wild Lima bean maintains most of its isozyme variation among populations. Significant positive correlation was observed between population size and P(p), A, and H(o) (observed heterozygosity), whereas no correlation was observed with the average fixation index of population (F). The loss of genetic variability in populations was attributed to inbreeding and the bottleneck effects that characterized the target populations. In situ conservation and management procedures for wild Lima bean are discussed.     

Allozyme diversity in natural populations of Viola palmensis Webb & Berth. (Violaceae) from La Palma (Canary Islands): implications for conservation genetics

Genetic diversity was measured by allozyme electrophoresis in eight natural populations of the threatened Canarian endemic Viola palmensis Webb & Berth. (Violaceae). Nineteen alleles corresponding to 11 gene loci were detected. High levels of genetic diversity were found, ranging from 36.3 to 45.4 % for the percentage of polymorphic loci (P), from 1.45 to 1.60 for the average number of alleles per locus (A) and from 0.128 to 0.200 for the expected heterozygosity (H(e)). Between 85.5 and 96.6 % of genetic variability was apportioned within populations. As a whole, populations were not at Hardy-Weinberg equilibrium, with a deficit of heterozygous individuals attributable to the existence of genetic structuring in the populations analysed. The levels of interpopulation genetic differentiation were low (mean F(ST) = 0.100), while genetic identity pair-wise comparisons were high (mean I = 0.973) suggesting considerable levels of gene flow among populations. No relationship was detected between genetic differentiation and geographical distances between populations. An outcrossing insect-mediated breeding system might contribute to pollen dispersion of this species. For conservation genetics we suggest in situ preservation areas are defined that are free of disturbance and that include populations with the highest genetic diversity.     

Microsatellite Analysis of Genetic Variation and Population Subdivision for the Black Muntjac, Muntiacus crinifrons

The black muntjac (Muntiacus crinifrons) is a rare deer found only in a restricted region in east China. Recent studies of mitochondrial DNA diversity have shown a markedly low level of nucleotide diversity for the species, and the Suichang population was genetically differentiated from the two other populations, in Huangshan and Tianmushan mountains. In this study, we extended the analysis of genetic diversity and population subdivision for the black muntjac using data from 11 highly polymorphic nuclear DNA microsatellite loci. Contrary to the results based on mtDNA data, the microsatellite loci revealed that the black muntjac retained a rather high nuclear genetic diversity (overall average H (E) = 0.78). Nevertheless, both types of markers supported the idea that the extant black muntjac population is genetically disrupted (overall phi (ST) = 0.16 for mtDNA and overall F (ST) = 0.053 for microsatellite, both P < 0.001). The correlation between genetic differentiation and geographic distance was not significant (Mantel test; P > 0.05), implying that the patterns of genetic differentiation observed in this study might result from recent habitat fragmentation or loss. Based on the results from the mtDNA and nuclear DNA data sets, two management units were defined for the species, Huangshan/Tianmushan and Suichang. We also recommend that a new captive population be established with individuals from the Suichang region as a founder source.     

Genetic variability of an endangered Bromeliaceae species (Pitcairnia albiflos) from the Brazilian Atlantic rainforest

Pitcairnia albiflos is a Bromeliaceae species endemic to Brazil that has been included as data-deficient in the extinction risk list of Brazilian flora. We analyzed genetic variability in P. albiflos populations using RAPD markers to investigate population structure and reproductive mechanisms and also to evaluate the actual extinction risk level of this species. Leaves of 56 individuals of P. albiflos from three populations were collected: Urca Hill (UH, 20 individuals), Chacrinha State Park (CSP, 24 individuals) and Tijuca National Park (TNP, 12 individuals). The RAPD technique was effective in characterizing the genetic diversity in the P. albiflos populations since it was possible to differentiate the populations and to identify exclusive bands for at least two of them. Even if there is low genetic diversity among them (CSP-UH = 0.463; CSP-TNP = 0.440; UH-TNP = 0.524), the populations seem to be isolated according to the low genetic diversity observed within them (H(pop) CSP = 0.060; H(pop) UH = 0.042; H(pop) TNP = 0.130). This fact might be the result of clonal and self-reproduction predominance and also from environmental degradation around the collection areas. Consequently, it would be important to protect all populations both in situ and ex situ to prevent the decrease of genetic variability. The low genetic variability among individuals of the same population confirms the inclusion of this species as critically endangered in the risk list for Brazilian flora.     

How much genetic variation is stored in the seed bank? A study of Atriplex tatarica (Chenopodiaceae)

We investigated to what extent the soil seed bank differed genetically and spatially in comparison to three consecutive life history stages (seedlings, mature plants, and fruiting plants) in a natural population of Atriplex tatarica. Representatives of particular life history stages from twenty subunits within a large population were randomly collected and subjected to allozyme analysis. Comparison of population polymorphism among various life history stages showed significant differences in observed heterozygosity (H(O)) and F statistics (F(IS) and F(ST)), but nonsignificant ones in the cases of number of alleles per polymorphic locus (A) and gene diversity (H(S)). These results indicate an increasing number of heterozygotes, a decreasing level of inbreeding and an increase of the partitioning genetic diversity among populations with increasing population age. Spatial autocorrelation was used to calculate f, the average co-ancestry coefficient between individuals within distance intervals of two meters along a 39 m long transect. Significant positive fine scale genetic structure was detected in mature and fruiting plants but not in soil seeds and seedlings stages. The results of the presented study on A. tatarica indicated that significant differences exist in genetic differentiation, differentiation in allele frequencies and spatial autocorrelation among early (soil seeds and seedlings) and late (mature and fruiting plants) life history stages but not within early and late ones. This pattern suggests that, rather than storing genetic variability in the soil or germination and establishment success, self-thinning might be the major microselective force in populations of A. tatarica.     

Molecular diversity and population structure of the forage grass Hemarthria compressa (Poaceae) in south China based on SRAP markers

Hemarthria compressa is one of the most important and widely utilized forage crops in south China, owing to its high forage yield and capability of adaptation to hot and humid conditions. We examined the population structure and genetic variation within and among 12 populations of H. compressa in south China using sequence-related amplified polymorphism (SRAP) markers. High genetic diversity was found in these samples [percentage polymorphic bands (PPB) = 82.21%, Shannon's diversity index (I) = 0.352]. However, there was relatively low level of genetic diversity at the population level (PPB = 29.17%, I = 0.155). A high degree of genetic differentiation among populations was detected based on other measures and molecular markers (Nei's genetic diversity analysis: G(ST) = 54.19%; AMOVA analysis: F(ST) = 53.35%). The SRAP markers were found to be more efficient than ISSR markers for evaluating population diversity. Based on these findings, we propose changes in sampling strategies for appraising and utilizing the genetic resources of this species.     

Does habitat fragmentation reduce fitness and adaptability? A case study of the common frog (Rana temporaria)

Studies examining the effects of anthropogenic habitat fragmentation on both neutral and adaptive genetic variability are still scarce. We compared tadpole fitness-related traits (viz. survival probability and body size) among populations of the common frog (Rana temporaria) from fragmented (F) and continuous (C) habitats that differed significantly in population sizes (C > F) and genetic diversity (C > F) in neutral genetic markers. Using data from common garden experiments, we found a significant positive relationship between the mean values of the fitness related traits and the amount of microsatellite variation in a given population. While genetic differentiation in neutral marker loci (F(ST)) tended to be more pronounced in the fragmented than in the continuous habitat, genetic differentiation in quantitative traits (Q(ST)) exceeded that in neutral marker traits in the continuous habitat (i.e. Q(ST) > F(ST)), but not in the fragmented habitat (i.e. Q(ST) approximately F(ST)). These results suggest that the impact of random genetic drift relative to natural selection was higher in the fragmented landscape where populations were small, and had lower genetic diversity and fitness as compared to populations in the more continuous landscape. The findings highlight the potential importance of habitat fragmentation in impairing future adaptive potential of natural populations.     

Allozyme differentiation and genetic structure of the Lady's slipper (Cypripedium calceolus) island populations in north-east Poland

The population genetic structure in the rare and endangered perennial plant, Cypripedium calceolus , from north-east Poland was investigated. A total of 11 loci were analysed, of which five were polymorphic for the species and in every population. The 11 loci resolved gave a total of 19 alleles with the mean number of 1.73 alleles per locus. The proportion of polymorphic loci (P) was the same for the species and in every population, 45.5%. The average observed and expected heterozygosities for the total Lady's slipper data set were 0.156 and 0.184, respectively. The average expected heterozygosity for the populations under study ranged from 0.176 to 0.188 and did not differ significantly among populations. Genetic diversity within C. calceolus populations from the Biebrza Valley is relatively high compared with rare taxa and taxa with similar life histories. The genetic diversity among populations of C. calceolus studied ( F _ST = 0.016) is much smaller than the genetic diversity within populations. In the species studied, there was no significant correlation between genetic and geographical distances ( r = −0.73, Mantel test). This may indicate the relatively recent origin of the populations studied, or a high level of gene flow among populations. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 138 , 433–440.     

Allozyme diversity and genetic structure of the leafy cactus (Pereskia guamacho [Cactaceae])

We examined levels of genetic variation and genetic structure in the leafy cactus (Pereskia guamacho) in arid and semiarid zones in Venezuela. We surveyed genetic diversity within 17 populations using 19 allozyme loci. Genetic diversity was relatively high at both the species (P(s) = 89%, A(s) = 3.26, AP(s) = 3.53, H(es) = 0.24) and population (P(p) = 63%, A(p) = 1.90, AP(p) = 2.42, H(ep) = 0.20) levels. A significant deficit of heterozygote individuals was detected within populations in the Paraguana Peninsula region (F(IS) = 0.301). Relatively low levels of population differentiation were detected at macrogeographic (G(ST) = 0.112) and regional levels (G(ST) = 0.044 for peninsula region and G(ST) = 0.074 for mainland region), suggesting substantial genetic exchange among populations; however, gene flow in this species seems to be regulated by the distance among populations. Overall, estimates of genetic diversity found in P. guamacho are concordant with the pattern observed for other cacti surveyed, namely high levels of polymorphism and genetic diversity with one common allele and several rare alleles per locus. Differences in gene dispersal systems between this species and other cacti studied were not reflected in the patterns of genetic diversity observed. The concentration of the highest estimates of genetic variation in northwestern Venezuela suggests a potential reservoir of plant genetic diversity within xerophilous ecosystems in northern South America.     

Patterns of Genetic Diversity in Rare and Common Orchids Focusing on the Korean Peninsula: Implications for Conservation

To provide basic information for orchid conservation, we surveyed the plant allozyme literature to summarize genetic diversity and structure data for (i) rare orchids native to the Korean Peninsula, and (ii) their congeners irrespective of being common and rare or Korean or not. A total of 68 taxa (32 taxa in Korea and 37 outside Korea; Goodyera repens being included in both datasets) were considered in this study. Overall, rare Korean orchid species had significantly lower levels of genetic diversity than their common congeners and common orchids in general at both population and species levels. However, mean values of G _ST (or F _ST) for rare and common orchids (Korean or not) did not differ significantly from each other. We found patterns of both low and high genetic diversity in rare Korean orchids. Many rare orchids harbored a complete lack of allozyme variation or extremely low within-population variation, perhaps due to rarity associated with random genetic drift and/or, for the case of warm-temperate orchids, to founder effects during post-glacial re-colonization. In contrast, high levels of genetic variation were found for a few orchids that have become recently rare (due to over-collection during the past several decades), probably because there have not been sufficient generations for the initial diversity to be substantially eroded. In addition, several orchids occurring in the main mountain system of the Korean Peninsula (the Baekdudaegan), that served as a glacial refugium, maintained moderate to high levels of within-population genetic diversity. Based on our genetic data, conservation priority should be given to rare orchid species. Particularly, urgent measures should be implemented on Jeju Island, a popular vacation spot, because it also a hotspot for threatened orchids with low levels of genetic diversity.     

Genetic differentiation among Hemarthria compressa populations in south China and its genetic relationship with H. japonica

Within and among populations genetic variance of twelve Hemarthria compressa populations and one Hemarthria japonica population from China were analyzed using inter simple sequence repeat (ISSR). Twelve primers amplified a total of 165 genomic DNA fragments across a total of 148 individuals of which 156 were polymorphic (94.55%). 75.76% of the bands were unique to each species, while the average genetic distance (GD) between one population of H. japonica and twelve populations of H. compressa was 0.44, which suggest that there was distinct differentiation between these two species. In H. compressa, twelve primers produced 145 bands across 145 individuals. High genetic diversity was observed at species level. The percentage of polymorphic loci (P) was 86.21% and Shannon's information index of diversity (I) was 0.357. In contrast, there were relatively low levels of genetic diversity within population (P=32.93%, I=0.174). Analysis of molecular variance (AMOVA) showed that a considerable proportion of genetic variation (48.02%) resided among populations. The coefficient of gene differentiation (G(ST)=48.6%) also suggested that there was strong genetic differentiation among H. compressa populations in southern China. An indirect estimate of the number of migrants per generation (N(m)=0.264) indicated that gene flow was low among populations of this species. Relative high clonal diversity was found, and all local genotypes were found.     

Isozyme variability of the wetland specialist Swertia perennis (Gentianaceae) in relation to habitat size, isolation, and plant fitness

We examined the effects of size and spatial isolation of fens on the isozyme variability of 17 populations of Swertia perennis. This long-lived perennial is a locally abundant fen specialist in Switzerland, where wetlands have been strongly fragmented. Isozyme variability was comparable to other outcrossing plants (A = 1.53, AP(p) = 2.01, P(p) = 42.5, H(o) = 0.113, H(e) = 0.139). F statistics indicated both inbreeding within and differentiation between populations (F(IS) = 0.076, F(IT) = 0.194, F(ST) = 0.128), with moderate gene flow between populations (N(e)m = 1.703). Populations in small, isolated fens had reduced genetic variability and the highest within-population inbreeding coefficients (F(IS)). Isozyme variability was significantly associated with vegetative fitness traits (MANOVA), and the magnitude of leaf herbivory decreased as the percentage of polymorphic loci increased. These data suggest that the reduced genetic variability of S. perennis in small, isolated populations may reduce plant fitness, thereby increasing susceptibility to herbivore damage. Our study also shows that habitat fragmentation can reduce the genetic variability of populations of fairly common habitat specialists, which so far have attracted less conservation attention than rare species.     

Domestication and the distribution of genetic variation in wild and cultivated populations of the Mesoamerican fruit tree Spondias purpurea L. (Anacardiaceae)

Domestication occurs as humans select and cultivate wild plants in agricultural habitats. The amount and structure of variation in contemporary cultivated populations has been shaped, in part, by how genetic material was transferred from one cultivated generation to the next. In some cultivated tree species, domestication involved a shift from sexually reproducing wild populations to vegetatively propagated cultivated populations; however, little is known about how domestication has impacted variation in these species. We employed AFLP data to explore the amount, structure, and distribution of variation in clonally propagated domesticated populations and sexually reproducing wild populations of the Neotropical fruit tree, Spondias purpurea (Anacardiaceae). Cultivated populations from three different agricultural habitats were included: living fences, backyards, and orchards. AFLP data were analysed using measures of genetic diversity (% polymorphic loci, Shannon's diversity index, Nei's gene diversity, panmictic heterozygosity), population structure (F(ST) analogues), and principal components analyses. Levels of genetic variation in cultivated S. purpurea populations are significantly less than variation found in wild populations, although the amount of diversity varies in different agricultural habitats. Cultivated populations have a greater proportion of their genetic variability distributed among populations than wild populations. The genetic structure of backyard populations resembles that of wild populations, but living fence and orchard populations have 1/3 more variability distributed among populations, most likely a reflection of relative levels of vegetative reproduction. Finally, these results suggest that S. purpurea was domesticated in two distinct regions within Mesoamerica.     

Genetic diversity and geographic differentiation in Tacca chantrieri (Taccaceae): an autonomous selfing plant with showy floral display

BACKGROUND AND AIMS: Despite considerable investment in elaborate floral displays, Tacca chantrieri populations are predominantly selfing. It is hypothesized that this species might possess considerable spatial or temporal variation in outcrossing rates among populations. To test this hypothesis, genetic variability and genetic differentiation within and among T. chantrieri populations were investigated to find out if they are in agreement with expectations based on a predominantly inbred mating system. METHODS: Genetic diversity was quantified using inter-simple sequence repeats (ISSR) in 303 individuals from 13 populations taken from known locations of T. chantrieri in China, and from one population in Thailand. KEY RESULTS: Of the 113 primers screened, 24 produced highly reproducible ISSR bands. Using these primers, 160 discernible DNA fragments were generated, of which 145 (90.62 %) were polymorphic. This indicated considerable genetic variation at the species level. However, there were relatively low levels of polymorphism at population levels, with percentages of polymorphic bands (PPB) ranging from 8.75 % to 55 %. A high level of genetic differentiation among populations was detected based on different measures (Nei's genetic diversity analysis: G(ST) = 0.5835; AMOVA analysis: F(ST) = 0.6989). Furthermore, based on levels of genetic differentiation, the 14 populations clustered into two distinct groups separated by the Tanaka Line. CONCLUSIONS: High levels of differentiation among populations and low levels of diversity within populations at large spatial scales are consistent with earlier small-scale studies of mating patterns detected by allozymes which showed that T. chantrieri populations are predominantly selfing. However, it appears that T. chantrieri has a mixed-mating system in which self-fertilization predominates, but there is occasional outcrossing. Significant genetic differences between the two distinct regions might be attributed to vicariance along the Tanaka Line. Finally, possible mechanisms of geographic patterns based on genetic differentiation of T. chantrieri are discussed.     

Population genetics of two rare perennials in isolated wetlands: Sagittaria isoetiformis and S. teres (Alismataceae)

We investigated genetic structure in two closely related perennial plants that occur in isolated wetlands: Sagittaria isoetiformis, restricted to the southeastern Coastal Plain of North America, and S. teres, endemic to the northeastern Coastal Plain. Using horizontal starch-gel electrophoresis, we screened 527 individuals from 11 populations of S. isoetiformis and 367 individuals from seven populations of S. teres. A high proportion of the 16 loci were polymorphic (%P(S) = 93.8% in S. isoetiformis and %P(S) = 75.0% in S. teres), with higher mean numbers of alleles per polymorphic locus and effective alleles per locus in S. isoetiformis (AP = 3.27, A(E) = 1.90) than in S. teres (AP = 2.58, A(E) = 1.30). Species- and population-level expected heterozygosities were higher in S. isoetiformis (H(ES) = 0.399, H(EP) = 0.218) than in S. teres (H(ES) = 0.177, H(EP) = 0.101). Jackknife estimates of F statistics indicated moderate levels of inbreeding in S. teres (F(IS) = 23.1%). Strong differentiation characterized these geographically isolated populations (G(ST) = 39.9% in S. isoetiformis, and G(ST) = 26.1% in S. teres). Genetic identities varied substantially within (ī = 75%, range = 0.558-0.963 in S. isoetiformis; ī = 89%, range = 0.776-0.963 in S. teres) and among species (ī = 81%, range = 0.506-0.882), leading to the discrimination of four regional population clusters using nonmetric multidimensional scaling (NMDS). It appears that S. isoetiformis and S. teres are a progenitor-derivative species pair.