Allozyme diversity in the federally threatened golden paintbrush, <Emphasis Type="Italic">Castilleja levisecta</Emphasis> (Scrophulariaceae)

Castilleja levisecta (Scrophulariaceae), the golden paintbrush, is an insect-pollinated herbaceaous perennial found in the Pacific Northwest. Currently restricted to two island populations off British Columbia and nine populations (eight on islands) in Washington, C. levisecta is a rare species threatened with extinction. Allozymes were used to describe genetic diversity and structure in these eleven populations. Despite its threatened status and small geographic range, exceptionally high levels of genetic diversity are maintained within C. levisecta. All sixteen of the loci resolved were polymorphic within the species (P_s=100%), while the mean percentage of loci polymorphic within populations (P_p) was 65.7%. The mean number of alleles per polymorphic locus (AP_s) was 2.94 within the species and averaged 2.38 within populations (AP_p). Genetic diversity (H_es) was 0.285 for the species, whereas mean population genetic diversity (H_ep) was 0.213. Smaller populations had, on average, fewer observed alleles and less genetic diversity. A significant negative correlation (r = –0.72) was found between genetic identity and geographic distance, indicating reduced gene flow between distant populations. The most geographically isolated population was one of the larger populations, one of the most genetically diverse and the most genetically divergent. A wide range of pairwise population genetic identities (I = 0.771 – 0.992) was found, indicating considerable genetic divergence between some populations. Overall, 19% of the total genetic diversity was distributed among populations. Results of this survey indicate that genetic augmentation of existing populations is unnecessary. The high allelic diversity found for the species and within its populations holds promise for conservation and restoration efforts to save this rare and threatened plant species.     

Genetic diversity within a threatened,endemic North American species, <Emphasis Type="Italic">Euphorbia telephioides</Emphasis> (Euphorbiaceae)

The southeastern United States and Florida support an unusually large number of endemic plant species, many of which are threatened by anthropogenic habitat disturbance. As conservation measures are undertaken and recovery plans designed, a factor that must be taken into consideration is the genetic composition of the species of concern. Here we describe the levels, and partitioning, of genetic diversity in 17 populations of the rare and threatened Florida endemic, Euphorbia telephioides (telephus spurge). Species-wide genetic diversity was high (P_s = 91%, AP_s = 3.81, A_s = 3.57 and H_es = 0.352) as was mean population genetic diversity (P_p = 81%, AP_p = 2.98, A_p = 2.59 and H_ep = 0.320) which ranks it among the highest 10% of plant species surveyed. Partitioning of genetic variation (G_st = 0.106) was low compared to other herbaceous outcrossing perennials indicating high historical gene flow across its limited geographic range. Among population G_st values within the three Florida counties in which it occurs, Gulf (0.084), Franklin (0.059) and Bay Counties (0.033), were also quite low. Peripheral populations did not generally have reduced genetic variation although there was significant isolation by distance. Rarefaction analysis showed a non-significant relationship between allelic richness and actual population sizes. Our data suggest that E. telephioides populations were probably more continuously distributed in Bay, Gulf and Franklin Counties and that their relative contemporary isolation is a recent phenomenon. These results are important for developing a recovery plan for this species.     

A cline of allozyme variation inAbies mariesii

Genetic variation at 22 allozyme loci was examined for 1,003 trees from 11 isolated natural populations ofAbies mariesii covering all except the southernmost region of its geographic range. Genetic diversity within species (H _es=0.063) was low compared to many other long-lived woody species. Most of the genetic variation is found within populations (G _ST=0.144) despite their isolated distribution. Genetic distance between populations was positively correlated with geographic distance. Genetic diversity within populations was generally low (meanH _ep=0.054), but varied across populations in a clinal fashion such that genetic variation decreased with increasing latitude. These genetic characteristics may reflect the distribution history of this species.     

Contrasting genetic structures in sister species of North American scrub-jays

Threatened Florida scrub-jays (Aphelocoma coerulescens breed communally, are restricted to xeric sandy scrub habitat, generally disperse fewer than three territory diameters. Closely related Western scrub-jays (A. californica) do not breed communally, have a broader habitat range, disperse greater distances, and are not threatened. We compared the genetic structure of 445 individuals in 11 populations in Florida with 157 individuals in eight populations of Western scrub-jays. At ten microsatellite loci, Florida had 24 out of 47 total alleles, while Western scrub-jays had 44. The Florida populations were more differentiated (G_ST = 0.048) than were a set of five California populations (G_ST = 0.015). A randomization extension of a Mantel test showed a stronger correlation between geographic and Cavalli-Sforza genetic distances among Western than Florida populations. Neighbour-joining trees clustered Florida populations from the same sandy ridge systems, suggesting that habitat continuity is more important than geographic proximity in allowing gene flow and preventing differentiation. For Western populations, isolation by distance appears to be the major determinant of genetic structure. Our results suggest that contrasting genetic structures may arise between closely related species, as a result of differences in ecology and social system. Conserving extant genetic variation in Florida jays will require maintaining viable populations in each of the major sandy ridge systems.     

Concordance between life history traits, invasion history, and allozyme diversity of the Everglades invader Melaleuca quinquenervia

During the century following its initial introduction in 1886, the Australian tree Melaleuca quinquenervia (Myrtaceae) dispersed from a few introduction points to occupy over 200,000 ha, primarily in historic Everglades wetlands of southern Florida. Cellulose acetate gel electrophoresis (CAGE) was used to investigate the allozyme diversity and population genetic structure of 208 individuals in a dozen populations resulting from this invasion. The analyses showed that these populations have a high (82%) rate of polymorphic loci and an average of 2 alleles/locus. There was substantial heterozygosity (mean H_e = 0.356), which concords well with recent studies reporting a greater number of introduction events and sources than generally recognized. The introduction history and distributional patterns within Florida have led to geographic structuring (G_ST = 0.419) in which the Gulf Coast metapopulation has a greater effective number of alleles and greater heterozygosity than the Atlantic Coast metapopulation. The gene diversity in M. quinquenervia was comparable to other tropical woody species. Its strong population divergence was reminiscent of pioneer species and consistent with its status as a plant invader in Florida.     

Low Levels of Genetic Diversity within Populations of Hosta clausa (Liliaceae)

Abstract Genetic diversity of Korean populations in Hosta clausa was investigated using starch gel electrophoresis. Hosta clausa is widespread, grows only along streamsides, and has both sexual and asexual reproduction. Populations of the species are small and isolated. Thirty-two percent of the loci examined were polymorphic, and mean genetic diversity within populations (He_p=0.082) was lower than mean estimates for species with very similar life history characteristics (0.131), particularly for its congener H. yingeri (0.250). The mean number of multilocus genotypes per population was 8.7, and genotypic diversity index (D_G) was 0.84. Significant differences in allele frequencies among populations were found in all seven polymorphic loci (P < 0.001). About one-fifth of the total allozyme variation was among populations (G_ST=0.192). Indirect estimate of the number of migrants per generation (Nm=0.48, calculated from mean G_ST) and nine private alleles found indicate that gene movement among populations was low. The low levels of genetic diversity within populations and the relatively high levels of genetic diversity among populations suggest that strong moist habitat preferences, clonal reproduction, low level of gene flow among populations, genetic drift, and historical events may have played roles in the genetic structuring of the species.     

Does GST underestimate genetic differentiation from marker data?

The widely applied genetic differentiation statistics F_ST and G_ST have recently been criticized for underestimating differentiation when applied to highly polymorphic markers such as microsatellites. New statistics claimed to be unaffected by marker polymorphisms have been proposed and advocated to replace the traditional F_ST and G_ST. This study shows that G_ST gives accurate estimates and underestimates of differentiation when demographic factors are more and less important than mutations, respectively. In the former case, all markers, regardless of diversity (H_S), have the same G_ST value in expectation and thus give replicated estimates of differentiation. In the latter case, markers of higher H_S have lower G_ST values, resulting in a negative, roughly linear correlation between G_ST and H_S across loci. I propose that the correlation coefficient between G_ST and H_S across loci, r_GH, can be used to distinguish the two cases and to detect mutational effects on G_ST. A highly negative and significant r_GH, when coupled with highly variable G_ST values among loci, would reveal that marker G_ST values are affected substantially by mutations and marker diversity, underestimate population differentiation, and are not comparable among studies, species and markers. Simulated and empirical data sets are used to check the power and statistical behaviour, and to demonstrate the usefulness of the correlation analysis.     

Population Genetic structure ofPotentilla fragariodes var.major (Rosaceae) in Korea

The genetic diversity and population structure of eighteenPotentilla fragariodes var.major (Rosaceae) populations in Korea were determined using genetic variations at 22 allozyme loci. The percent of polymorphic loci within the enzymes was 66.7%. Genetic diversity at the species level and at the population level was high (Hes = 0.203; Hep = 0.185, respectively), whereas the extent of the population divergence was relatively low (G_ST = 0.069). F_IS, a measure of the deviation from random mating within the 18 populations, was 0.075. An indirect estimate of the number of migrants per generation (Nm = 3.36) indicated that gene flow was high among Korean populations of the species. In addition, analysis of fixation indices revealed a slight heterozygote deficiency in some populations and at some loci. Wide geographic ranges, perennial herbaceous nature and the persistence of multiple generations are associated with the high level of genetic variation. AlthoughP. fragariodes var.major usually propagated by asexually-produced ramets, we could not rule out the possibility that sexual reproduction occurred at a low rate because each ramet may produce terminal flowers. Mean genetic identity between populations was 0.983. It is highly probable that directional movement toward genetic uniformity in a relatively homogeneous habitat operates among Korean populations ofP. fragariodes var.major.     

Fire-mediated effects of shrubs, lichens and herbs on the demography of Hypericum cumulicola in patchy Florida scrub

Understanding the effect of disturbance and interspecific interactions on population dynamics and availability of suitable habitats for colonization and growth is critical for conservation and management of endangered species. Hypericum cumulicola is a narrowly endemic, small perennial herb virtually restricted to open areas of well-drained white sand in Florida rosemary scrub, a naturally patchy community that burns about every 20–80 years. Over 1 year (September 1994 to September 1995) we evaluated variation in survival, growth and fecundity among 1214 individuals in 14 rosemary scrub patches of different sizes (0.09–1.85 ha) and fire histories (2, 8–10, and >20 years since the last fire). Fire kills aboveground individuals of H. cumulicola, but new individuals were present a year after fire. Recruitment decreased in patches more than a decade post-fire. Survival, annual height growth rate, and fecundity (number of flowers and fruits) were higher in recently burned patches. Scrub patch size did not affect these demographic variables. Survival was positively associated with the presence of conspecifics and negatively related to proximity to the dominant shrub Florida rosemary (Ceratiola ericoides), prior reproductive output, and ground lichen cover. Since H. cumulicola and other herbaceous species in the rosemary scrub depend on sporadic fires to decrease interference of shrubs and ground lichens, its persistence may be threatened by fire suppression. Received: 4 December 1996 / Accepted: 5 June 1997     

Sequential loss of genetic variation in flea beetle Agasicles hygrophila (Coleoptera: Chrysomelidae) following introduction into China

The flea beetle (Agasicles hygrophila) was imported to Florida, USA and then introduced from Florida into China in 1987 as a biological control agent for the invasive plant alligator weed (Alternanthera philoxeroides). The initial beetle population was subsequently used for sequential introductions in other areas of China, but little is known about the genetic consequences of the introductions. In this study, the genetic diversity and population structure of five beetle populations, the source Florida population, three intentionally introduced China populations and one accidentally dispersed China population, were examined using amplified fragment length polymorphisms. The results showed a clear pattern of decreasing genetic diversity with the sequential introductions. The diversity was highest in the Florida population followed by the first introduction to Chongqing and then in Kunming and Fuzhou. The lowest diversity was found in the accidentally dispersed Guangzhou population that was first recorded in 1996. Both loci parameters and Nei's genetic diversity showed a high variation among these populations. Genetic differentiation among populations was further verified by the G_ST statistic (0.136–0.432). Beetles in Kunming had the highest gene flow with those in Guangzhou, and therefore lowest differentiation and closest genetic distance. These data show that sequential introduction influenced the genetic diversity of populations in China. Genetic diversity should be considered in planning introduction and long‐term maintenance of populations.     

A biogeographical profile of the sand cockroach Arenivaga floridensis and its bearing on origin hypotheses for Florida scrub biota

Florida scrub is a xeric ecosystem associated with the peninsula's sand ridges, whose intermittent Pliocene–Pleistocene isolation is considered key to scrub endemism. One scrub origin hypothesis posits endemics were sourced by the Pliocene dispersal of arid‐adapted taxa from southwestern North America; a second invokes Pleistocene migration within eastern North America. Only one study to date has explicitly tested these competing hypotheses, supporting an eastern origin for certain scrub angiosperms. For further perspective, we conducted a genetic analysis of an endemic arthropod, the Florida sand cockroach (Arenivaga floridensis), with two aims: (1) to reconstruct the peninsular colonization and residence history of A. floridensis and (2) determine whether its biogeographic profile favors either origin hypothesis. We sequenced the cox2 mitochondrial gene for 237 specimens (65 populations) as well as additional loci (cox1, nuclear H3) for a subset of Florida roaches and congeners. Using Network and Bayesian inference methods, we identified three major lineages whose genetic differentiation and phylogeographical structure correspond with late Pliocene peninsula insularization, indicating Arenivaga was present and broadly distributed in Florida at that time. Stem and crown divergence estimates (6.36 Ma; 2.78 Ma) between A. floridensis and western sister taxa span a period of extensive dispersal by western biota along an arid Gulf Coast corridor. These phylogeographical and phylogenetic results yield a biogeographic profile consistent with the western origin hypothesis. Moreover, age estimates for the roach's peninsular residence complement those of several other endemics, favoring a Pliocene (or earlier) inception of the scrub ecosystem. We argue that eastern versus western hypotheses are not mutually exclusive; rather, a composite history of colonization involving disparate biotas better explains the diverse endemism of Florida scrub.     

Genetic diversity within and among populations of Shorea leprosula Miq. and Shorea parvifolia Dyer (Dipterocarpaceae) in Indonesia detected by AFLPs

The genetic diversity within and among populations of Shorea leprosula and Shorea parvifolia from Indonesia was investigated using amplified fragment length polymorphisms (AFLPs). The results indicated that S. leprosula is genetically more variable than S. parvifolia. At the population level, a higher level of genetic diversity was revealed for S. leprosula with a percentage of polymorphic loci (PPL_p) of 53.32% and an expected heterozygosity (H _ep) of 0.16 in comparison with S. parvifolia showing PPL_p of 51.79% and H _ep of 0.14. At the species level, S. leprosula showed PPL_s of 92.86% and H _es of 0.21, while S. parvifolia showed PPL_s of 85.71% and H _es of 0.21. Genetic differentiation (G _st) indicated that 25 and 31% of total genetic diversity in S. leprosula and S. parvifolia, respectively, were attributed to the differences among populations. An analysis of molecular variance (AMOVA) at two hierarchical levels exhibited that most genetic variation resided within populations with proportion of 70.2% for S. leprosula and 66.2% for S. parvifolia. The AMOVA at three hierarchical levels performed for S. leprosula and S. parvifolia together revealed that the genetic difference between the two species was remarkably higher with a proportion of 44.1% than the differences within and among populations (38.1 and 17.8%, respectively). The genetic differentiation between islands was significant for S. leprosula but not for S. parvifolia. The observed genetic diversity agreed with the life history traits of Shorea species. Highly differentiating individual AFLP markers were found for each species, which will serve as diagnostic markers for the identification of wood of different species, from different islands and regions.     

Genetic variation and population structure ofCarex breviculmis (Cyperaceae) in Korea

We determined the genetic diversity and population structures ofCarex breviculmis (Cyperaceae) populations in Korea, using genetic variations at 23 allozyme loci.C. breviculmis is a long-lived herbaceous species that is widely distributed in eastern Asia. A high level of genetic variation was found in 15 populations. Twelve enzymes revealed 23 loci, of which 11 were polymorphic (47.8%). Genetic diversity at the speciesand population levels were 0.174 and 0.146, respectively. Total genetic diversity (H_T = 0.363) and within-population genetic diversity (H_s = 0.346) were high, whereas the extent of the population divergence was relatively low (G_ST = 0.063). Deviation from random mating (Fis) within the 15 populations was 0.206. An indirect estimate of the number of migrants per generation(Nm = 3.69) indicated that gene flow was extensive among Korean populations of this species. Analysis of fixation indices revealed a substantial heterozygote deficiency in some populations and at some loci. Genetic identity between popu-lations was high, exceeding 0.956.     

Genetic diversity and gene flow estimates among three populations of Botryodiplodia theobromae causing die-back and bark canker of pear in Punjab

Three populations of Botryodiplodia theobromae causing die-back and bark canker of pear in Punjab were analysed to know about the genetic diversity, gene flow, heterogeneity and the probable rate of spread of races capable of overcoming the resistance present in elite cultivars. There was no relationship between the morphologically similar isolates and their pathogenic behaviour. Majority of the isolates of B. theobromae within and among the populations produced lesions of 1.9–7.2 × 0.8–3.1 cm size on detached pear twigs cv. Punjab Beauty. Incubation period and per cent infection also varied within and among the three populations of the pathogen. The genetic diversity was calculated on the basis of allele frequencies of nine simple sequence repeat (SSR) markers using Nei's genetic diversity formulae. Allele frequencies (X_ij ) varied significantly among the three populations ranging from 0.0 to 1.0. The mean genetic diversities within population (H _S) also varied in all the locations ranging from 0.18 to 0.36 indicating a high genetic diversity within the population. The total genetic diversity across all the populations (H _T) ranged between 0.1 and 0.5 with a mean of 0.36, and differed significantly from the mean H _S (0.25). Diversity indices of SSR loci varied from low (H _S < 0.1) to high (H _S > 0.4) across all the populations (G _ST) of B. theobromae in Punjab which were consistent over the presumed SSR loci with a mean G _ST of 0.29. The lower G _ST values of 0.13, 0.10, 0.11 and 0.13 were observed at LAS15-16, LAS27-28, BOT17-18 and BOT35-36 loci, respectively, showing high genetic diversity among all the isolates. However, at BOT19-20 locus, the G _ST value observed was 0.72 thereby indicating low diversity in the population at this locus. Out of 25 loci, 8 loci showed the gene flow (Nm) < 1 indicating a high genetic differentiation within the population and the remaining 6 loci showed the Nm > 1 indicating the frequent existence of gene flow across the populations of B. theobromae. Pairwise comparison (G _ST) values among all the loci were ranging from 0.18 to 0.19, which indicate a low genetic differentiation among the populations of B. theobromae. The high genetic diversity (H _S) within the population was observed in the present study suggesting that B. theobromae possess high variability in Punjab. Keeping this in view, new races would be expected soon outside the state of origin, as naturally occurring gene flow is likely to be increased by the human activity for comprehensive cultivation of pear in the near future. Therefore, it is difficult to predict the durability of resistant sources in Punjab, which in turn emphasises the identification and introgression of R genes into commercial cultivars.     

Patterns of genetic diversity and differentiation in the tsetse fly <Emphasis Type="Italic">Glossina morsitans morsitans</Emphasis> Westwood populations in East and southern Africa

Genetic diversity and differentiation within and among nine G. morsitans morsitans populations from East and southern Africa was assessed by examining variation at seven microsatellite loci and a mitochondrial locus, cytochrome oxidase (COI). Mean COI diversity within populations was 0.63 ± 0.33 and 0.81 taken over all populations. Diversities averaged over microsatellite loci were high (mean number of alleles/locus ≥7.4; mean H _E ≥ 65%) in all populations. Diversities averaged across populations were greater in East Africa (mean number of alleles = 22 ± 2.6; mean h _e = 0.773 ± 0.033) than in southern Africa (mean number of alleles = 18.7 ± 4.0; mean h _e = 0.713 ± 0.072). Differentiation among all populations was highly significant (R _ST = 0.25, F _ST = 0.132). Nei’s G _ij statistics were 0.09 and 0.19 within regions for microsatellites and mitochondria, respectively; between regions, G _ij was 0.14 for microsatellites and 0.23 for mitochondria. G _ST among populations was 0.23 for microsatellite loci and 0.40 for mitochondria. The F, G and R statistics indicate highly restricted gene flow among G. m. morsitans populations separated over geographic scales of 12–917 km.     

Allelopathic effects of <Emphasis Type="Italic">Ceratiola ericoides</Emphasis> (Empetraceae) on germination and survival of six Florida scrub species

Allelopathic inhibition of germination by Florida scrub plants has been demonstrated in the greenhouse and lab, but not in the field. We studied the allelopathic effects of Florida rosemary (Ceratiola ericoides) roots, leaves, and litter leachates on field germination and three-month survival of six Florida scrub species, three habitat generalists (Lechea deckertii, Palafoxia feayi, and Polygonella robusta), and three rosemary scrub specialists (Hypericum cumulicola, Lechea cernua, and Polygonella basiramia). We used AIC and model averaging to evaluate support for a series of non-exclusive hypotheses. Species varied in germination (2.7–24.6%) and survival (39.2–71%) percentages, and in their sensitivity to leachates. Germination of scrub species was most negatively affected by leaf > root > litter leachates, although not all species followed the overall trend. Additional germination suppression by leachate combinations (relative to single leachates) was minimal. Sites did not vary in germination, but seedling survival did differ among sites. This study further documents the negative impact of Florida rosemary leachates on the germination of co-occurring plant species. Allelopathy may be partly responsible for bare sand gaps in Florida rosemary scrub, and therefore be one of the forces structuring Florida rosemary scrub ecosystems.     

Population structure of red-cockaded woodpeckers in south Florida: RAPDs revisited

Six south Florida populations of the endangered red-cockaded woodpecker (Picoides borealis) were sampled to examine genetic diversity and population structure in the southernmost portion of the species' range relative to 14 previously sampled populations from throughout the species range. Random amplified polymorphic DNA (RAPD) analyses were used to evaluate the populations (n= 161 individuals, 13 primers, one band/primer). Results suggested that south Florida populations have significant among-population genetic differentiation (F_ST= 0.17, P < 0.000), although gene flow may be adequate to offset drift (N_m= 1.26). Comparison of Florida populations with others sampled indicated differentiation was less in Florida (F_ST for all populations = 0.21). Cluster analyses of all 20 populations did not reflect complete geographical predictions, although clustering of distant populations resulted in a significant correlation between genetic distance and geographical distance. Overall, results suggest populations in south Florida, similar to the remainder of the species, have low genetic diversity and high population fragmentation. Exact clustering of distant populations supports the ability of RAPDs to differentiate populations accurately. Our results further support past management recommendations that translocations of birds among geographically proximate populations is preferable to movement of birds between distant populations.     

Genetic and clonal diversity in Korean populations ofVitex rotundifolia (Verbenaceae)

Vitex rotundifolia L.f. is a woody perennial and has sexual and asexual modes of reproduction. Allozyme study was conducted on 550 plants in 13 Korean populations. The levels of genetic variability and divergence within and among populations, respectively, are considerably lower and higher than the mean values for woody plants with similar life history tralts. Mean percentage of polymorphic loci (P _P), mean number of alleles per locus (A _P), and mean genetic diversity (He _P) within populations ofV. rotundifolia were: 16.7%, 1.21, and 0.047. On average, about 79% of the total variation inV. rotundifolia was common to all populations (meanG _ST=0.208). In addition, significant differences in allele frequencies among populations were found in all polymorphic loci examined (P<0.001). On the other hand, levels of genotypic diversity within and among populations were moderate. About 44% (18/41) of multilocus genotypes were “local genotypes” (genotypes occurring in only one population), whereas only one “widespread genotype” (genotypes occurring in more than 75% of the populations) were detected. The mean number of multilocus genotypes per population (G) and mean genotypic diversity index (D _G) were 8.4 and 0.74, respectively. Most common multilocus genotypes found in populations were homozygous for five polymorphic loci. The abundance of ramets of these genets is responsible for the low levels of expected heterozygosity within populations. The results indicate that clonal reproduction may act as an enhancer of genetic drift by reducing effective size of local populations ofV. rotundifolia.     

Genetic diversity of Poa annua in western Oregon grass seed crops

The genetic diversity of Poa annua L.populations collected from western Oregon grass-seed fields was surveyed using 18 randomly amplified polymorphic DNA (RAPD) markers. Markers from 1357 individual plants from 47 populations collected at three sampling dates (fall, winter, and spring) for 16 sites were used to measure genetic diversity within and among populations. Site histories varied from low to high herbicide selection pressure, and some sites were subdivided by 3 years of differing post-harvest residue management. Gene diversity statistics, simple frequency of haplotype occurrence, and analysis of molecular variance (AMOVA) revealed the presence of significant variability in P. annua among sites, among collection dates within sites, and within collection dates. Nei gene-diversity statistics and population-differentiation parameters indicated that P. annua populations were highly diverse. Mean Nei gene diversity (h) for all 47 populations was 0.241 and total diversity (H_T) was 0.245. A greater proportion of this diversity, however, was within (H_S=0.209) rather than among (G_ST=0.146) populations. When populations were grouped by season of collection, within-group diversity was H_S=0.241, while among-group diversity was G_ST=0.017. When populations were grouped by site, within-group diversity was H_S=0.224, while among-group diversity was G_ST=0.087. The diversity among populations within season for fall, winter, and spring collections was G_ST=0.121, 0.142, and 0.133, respectively. Populations collected from fields with histories of high herbicide selection pressure showed low differentiation among collection dates, with G_ST as low as 0.016, whereas those collected from fields with low herbicide selection pressure showed greater differentiation among collection dates, with G_ST as high as 0.125. At high selection-pressure sites, populations were also lower in gene diversity (as low as h=0.155), while at low selection-pressure sites there was higher gene diversity (as high as h=0.286). The site to site variability was greater for the high selection-pressure sites (G_ST=0.107 or 69% of the total among-population variance), while the season of germination variability was greater at sites of low herbicide-selection pressure (G_ST=0.067, or 70% of the total among-population variance). High initial diversity coupled with a long-term re-supply of genotypes from the seed bank must have been factors in maintaining the genetic diversity of this weed despite the intensive use of herbicides. Knowledge of the genetic diversity of Willamette Valley P. annua should help in formulating more effective strategies for managing this weed. Received: 24 July 1999 / 11 November 1999     

Allozyme variability within and among populations of the epiphytic moss Leucodon (Leucodontaceae: Musci)

Enzyme electrophoresis was used to estimate genetic variability within and among populations of five species of the genus Leucodon (L. atrovirens, L. luteus, L. nipponicus, L. sapporensis, and L. temperatus). All of these perennial, dioecious plants are epiphytic on tree trunks and produce numerous spores that are released in winter. Gene diversity within populations (H_S) was low (0.070) in L. nipponicus, which grows in disturbed habitats, and high (0.169–0.202) in the other four species, which inhabit stable environments. Gene diversity between populations relative to total diversity (G_ST) was low in L. temperatus (0.042) and L. luteus (0.043), suggesting that extensive gene flow occurs among populations. G_ST was moderate in L. sapporensis (0.104) and L. atrovirens (0.114); these values are still less than those previously reported for terrestrial mosses. Since epiphytes could disperse their spores effectively by the wind, low genetic divergence among populations may be related to the epiphytic habit of the species. On the other hand, high G_ST value (0.208) was found in L. nipponicus. It is probably explained by the genetic differentiation of populations by genetic drift, for the species forms geographically and ecologically isolated populations with small number of individuals. We also observed incongruity between morphological and genetic similarity among species.