Chaoborus and fish-mediated influences on Daphnia longispina population structure,dynamics and life history strategies

Summary This study examined the long term effects of predation by larvae of the midge Chaoborus and simulated fish predation on experimental Daphnia longispina populations. Chaoborus predation, relative to fish predation, led to populations composed of larger individuals as a whole, larger egg-bearing individuals, and a larger primiparous instar. Daphnia retained helmets beyond the first instar in response to the presence of Chaoborus. Both types of predation, relative to predator-free controls, reduced prey population size and rates of increase, but increased population death rates. The reduction in population size due to predation led to increased resource availability for individuals remaining in the populations and increased individual fecundity in the predation treatments. The differences noted between the Chaoborus, fish, and control treatments increased with predation intensity.     

Chaoborus predation and delayed reproduction in Daphnia: a demographic modeling approach

I develop a demographic model that examines the impact of Chaoborus predation on the population dynamics and life history of Daphnia. Predation effects are determined through analysis of the various components of the predator-prey interaction (encounter, attack, strike efficiency), and are integrated into a stage-classified matrix population model. The model is parameterized with data from interactions between D. pulex and fourth-instar C. americanus. I test this model with two laboratory experiments that examine population growth of D. pulex under the influence of five different levels of Chaoborus predation. With the exception of a single predation treatment in each experiment, the model accurately predicted the observed reduction in Daphnia numbers with increasing Chaoborus predation. I then use this model to investigate the evolution of delayed reproduction in D. pulex that are exposed to Chaoborus. I ask whether delayed reproduction may evolve in Daphnia that are subjected to Chaoborus predation as a trade-off for the benefits of larger body size. The model predicts that the effectiveness of such a life history trade-off depends on the relative sizes of predator and prey. In some interactions between Chaoborus and Daphnia, increasing Daphnia body length by as little as 5% from base growth trajectories sufficiently increases fitness (by reducing vulnerability to Chaoborus predation) to compensate for the cost of delayed reproduction. In other interactions, however, increased body length provides no benefit to Daphnia (and may even reduce fitness), and selection would act against the evolution of delayed reproduction. This revised version was published online in July 2006 with corrections to the Cover Date.     

Phosphorus availability mediates plasticity in life-history traits and predator–prey interactions in Daphnia

We analysed growth plasticity of two Daphnia pulex clones under low‐phosphorus (LP) and high phosphorus (HP) conditions, in the presence of Chaoborus kairomones to examine how food quality (P‐availability) might impact life‐history responses and vulnerability to predation. Overall, clone 1 grew faster, and was larger at maturity. Under HP, both clones responded to kairomones by increasing growth, age and size at maturity, and decreasing fecundity. Under LP, both clones suffered reduced growth, and fecundity. However, the magnitude of response to kairomones depended on a clone by P‐availability interaction. Chaoborus presented a 1 : 1 clonal mixture under HP or LP, consumed more individuals under LP. Moreover, fewer clone 1 individuals were consumed. Studying the effects of P‐availability on life histories, and predator–prey interactions may help us understand the mechanisms generating and maintaining plasticity, as well as influencing genotypic diversity and microevolutionary processes in natural populations.     

A PHYLOGENETIC ANALYSIS OF BODY SIZE EVOLUTION AND BIOGEOGRAPHY IN CHUCKWALLAS (SAUROMALUS) AND OTHER IGUANINES

The evolution of body size was reconstructed in chuckwallas (genus Sauromalus), large herbivorous lizards of southwest North America, using a phylogeny derived from sequence variation in the mitochondrial cytochrome b gene. The body mass of two endemic island species (S. hispidus and S. varius) is typically fivefold larger than mainland species. We tested the hypothesis that large body size has evolved on these islands in response to local ecological conditions against the alternative hypothesis that large size is simply retained from large iguanine ancestors. The most parsimonious tree topology depicts the insular gigantic Sauromalus as monophyletic, having diverged from a common ancestor on the Baja California peninsula after the radiation of smaller bodied clades. In a robustness analysis of this topology, we found general support for this tree over alternative topologies representing minimum evolution hypotheses that imply large body size is retained from large iguanine ancestors. The most parsimonious reconstruction of body size evolution implies a change from large to small size after the Sauromalus ancestor diverged from Iguana, and one reversal back to large size within Sauromalus. The large size increase in the gigantic clade contrasts with evolutionary stasis of small body size (for an iguanine) in mainland populations. The gigantic species show 3–4% total sequence divergence from S. obesus populations on the nearby Baja California peninsula, and mainland populations of S. obesus obesus show similar levels of divergence from each other. An analysis of character transitions and comparative behavior implicates predation, and its relaxation on isolated islands, as a strong selective force in Sauromalus. Patterns of genetic differentiation in Sauromalus and biogeographic implications are discussed.     

An experimental study of the effects of nutrient supply and Chaoborus predation on zooplankton communities of a shallow tropical reservoir (Lake Brobo, Côte d'Ivoire)

1. Based on two mesocosm experiments and 10 in vitro predation experiments, this work aimed to evaluate the impact of nutrient supply and Chaoborus predation on the structure of the zooplankton community in a small reservoir in Côte d'Ivoire. 2. During the first mesocosm experiment (M1), P enrichment had no effect on phytoplankton biomass (chlorophyll a) but significantly increased the biomass of some herbivorous zooplankton species (Filinia sp, Ceriodaphnia affinis). During the second experiment (M2), N and P enrichment greatly increased phytoplankton biomass, rotifers and cladocerans (C. affinis, C. cornuta, Moina micrura and Diaphanosoma excisum). In both experiments, nutrient addition had a negative impact on cyclopoid copepods. 3. Larger zooplankton, such as cladocerans or copepodites and adults of Thermocyclops sp., were significantly reduced in enclosures with Chaoborus in both mesocosm experiments, whereas there was no significant reduction of rotifers and copepod nauplii. This selective predation by Chaoborus shaped the zooplankton community and modified its size structure. In addition, a significant Chaoborus effect on chlorophyll a was shown in both experiments. 4. The preference of Chaoborus for larger prey was confirmed in the predation experiments. Cladocerans D. excisum and M. micrura were the most selected prey. Rotifer abundance was not significantly reduced in any of the 10 experiments performed. 5. In conclusion, both bottom‐up and top‐down factors may exert a structuring control on the zooplankton community. Nutrients favoured more strictly herbivorous taxa and disadvantaged the cyclopoid copepods. Chaoborus predation had a strong direct negative impact on larger crustaceans, favoured small herbivores (rotifer, nauplii) and seemed to cascade down to phytoplankton.     

A contribution to characterisation of genetic variation in some natural Polish populations of Elymus repens (L.) Gould and Elymus hispidus (Opiz) Melderis (Poaceae) as revealed by RAPD markers*

To determine the relative importance of clonal growth and sexual reproduction, the Randomly Amplified Polymorphic DNA (RAPD) method was used to study genetic diversity and clonal structure of six populations of Elymus repens and four populations of Elymus hispidus from Poland. These outbreeding species are virtually self‐sterile and form widely spreading and long‐lived rhizomes. Using 12 primers, a total of 150 unambiguous RAPD fragments were amplified and scored. Results of AMOVA showed no significant genetic distinction between morphologically distinguished varieties of E. repens and E. hispidus. E. repens had slightly higher intra‐specific genetic polymorphism than E. hispidus; the percentage of polymorphic bands per population ranged from 38 to 49 and from 19 to 38 respectively. Clonal diversity measured using the Simpson diversity index (D) indicated different contributions of clonal reproduction in particular populations of E. repens (D: 0.20–0.72). Populations of E. hispidus were dominated by one or a few clones, which were generally restricted to a single population (D: 0.00–0.22). RAPD revealed that most genetic diversity resided within populations of the two studied species, suggesting that, despite their clonal character, propagation by seeds contributes considerably to reproduction of E. repens and E. hispidus.     

Sexual reproduction, clonal diversity and genetic differentiation in patchily distributed populations of the temperate forest herb Paris quadrifolia (Trilliaceae)

Clonal plant species have been shown to adopt different strategies to persist in heterogeneous environments by changing relative investments in sexual reproduction and clonal propagation. As a result, clonal diversity and genetic variation may be different along environmental gradients. We examined the regional and local population structure of the clonal rhizomatous forest herb Paris quadrifolia in a complex of forest fragments in Voeren (Belgium). Relationships between population size (the number of shoots), shoot density (the number of shoots per m²) and local growth conditions were investigated for 47 populations. Clonal diversity and genetic variation within and among 19 populations were investigated using amplified fragment length polymorphism markers. To assess the importance of sexual reproduction, seed set, seed weight and germination success were determined in 18 populations. As predicted, local growth conditions largely affected population distribution, size and density of P. quadrifolia. Populations occurring in moist and relatively productive sites contained significantly more shoots. Here, shoots were also much more sparsely distributed compared to populations occurring in dry and relatively unproductive sites, where shoots showed a strongly aggregated distribution pattern. Clonal diversity was relatively high, compared with other clonal species (G/N ratio = 0.43 and Simpson’s D=0.81). Clonal diversity significantly (P<0.01) decreased with increasing shoot density while molecular genetic variation was significantly (P<0.01) affected by population size and local environmental conditions. Lack of recruitment and out-competition of less-adapted genotypes may explain the decreased genetic variation in dry sites. Analysis of molecular variance revealed significant genetic variation among populations (Φ _ST=0.42, P<0.001), whereas pairwise genetic distances were not correlated to geographic distances, suggesting that gene flow among populations is limited. Finally, the number of generative shoots, the number of seeds per fruit and seed weight were significantly and positively related to population size and local growth conditions. We conclude that under stressful conditions populations of clonal forest plant species can slowly evolve into remnant populations characterized by low levels of genetic variation and limited sexual reproduction. Conservation of suitable habitat conditions is therefore a prerequisite for effective long-term conservation of clonal forest plant species.     

EVOLUTION OF GIGANTISM IN NINE‐SPINED STICKLEBACKS

The relaxation of predation and interspecific competition are hypothesized to allow evolution toward “optimal” body size in island environments, resulting in the gigantism of small organisms. We tested this hypothesis by studying a small teleost (nine‐spined stickleback, Pungitius pungitius) from four marine and five lake (diverse fish community) and nine pond (impoverished fish community) populations. In line with theory, pond fish tended to be larger than their marine or lake conspecifics, sometimes reaching giant sizes. In two geographically independent cases when predatory fish had been introduced into ponds, fish were smaller than those in nearby ponds lacking predators. Pond fish were also smaller when found in sympatry with three‐spined stickleback (Gasterosteus aculeatus) than those in ponds lacking competitors. Size‐at‐age analyses demonstrated that larger size in ponds was achieved by both increased growth rates and extended longevity of pond fish. Results from a common garden experiment indicate that the growth differences had a genetic basis: pond fish developed two to three times higher body mass than marine fish during 36 weeks of growth under similar conditions. Hence, reduced risk of predation and interspecific competition appear to be chief forces driving insular body size evolution toward gigantism.     

CLONAL STRUCTURE,GENOTYPIC DIVERSITY,AND SEED PRODUCTION IN POPULATIONS OF FILIPENDULA RUBRA (ROSACEAE) FROM THE NORTHCENTRAL UNITED STATES

Queen of the prairie, Filipendula rubra (Rosaceae), is a clonal plant species inhabiting calcareous fens and wet meadows of the northcentral United States. F. rubra reproduces asexually by underground rhizomes and sexually by seed. While many studies have explored genotype diversity in clonal species with limited sexual reproduction, fewer have been conducted on clonal species with the potential for extensive sexual reproduction. We studied the relationship between the extent of sexual reproduction and genotype diversity. Although genotype diversity in F. rubra was double that reported by others for 27 nearly obligate clonal plant species, it was still quite low. For 25 populations studied, the mean number of genotypes was 5.5 (range = 1–15; SE = 0.964) and the average proportion of distinguishable genotypes was 0.38 (range = 0.03–1.00; SE = 0.07). The production of viable seed was quite variable among populations (mean proportion of viable seeds = 0.242; range = 0.002–0.565; SE = 0.04). Considering that some inflorescences can produce over 5,000 seeds, the potential for recruitment of sexually produced individuals is very large. No correlation was found between seed production and genotype diversity as was expected in a self-incompatible species in which one-third of the populations possessed a single genotype. It was hypothesized that the low genotype diversity found in numerous populations may be due to competition limiting recruitment of new seedlings.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

The interaction of Chaoborus size and vertical distribution determines predation effects on Daphnia

1. Larvae of Chaoborus, the phantom midge, are important pelagic planktivores in many freshwater lakes and ponds. The effect of Chaoborus on its prey depends on its size, especially mouth gape diameter, and vertical migration pattern, which affects predator–prey spatial overlap. These two features vary considerably in different Chaoborus species and instars. In this study, the interacting effects of both Chaoborus size and vertical distribution on population growth of Daphnia pulex was analysed with a field enclosure experiment and a matrix population model. 2. In the field experiment, Daphnia were grown in four replicated treatments that included a control (no Chaoborus) and three combinations of instar III and IV Chaoborus of two species (C. trivittatus and C. americanus). Parameters of the matrix model were based on differences between Chaoborus species and instars in capture and ingestion of Daphnia of differing sizes (prey vulnerability) and in vertical overlap with Daphnia in each treatment (density risk). 3. In comparison with the control, the two treatments containing the smaller, migratory C. americanus showed a significant effect on Daphnia population growth rate, while the treatment containing only the larger, non‐migratory C. trivittatus did not. The model accurately simulated these effects. 4. A Daphnia predation risk model, which uses prey vulnerability and density risk parameters, illustrated the individual and combined effect of the different Chaoborus types on Daphnia. Daphnia have a high prey vulnerability to the large C. trivittatus, but overall predation risk was low because of very little overlap. On the contrary, the smaller C. americanus affects only a small range of Daphnia instars, each with a low vulnerability, yet those instars that were vulnerable had a very high density risk because of an increased overlap. 5. This analysis of Daphnia predation risk parameters with coexisting Chaoborus species strongly supports an integrated approach using both size and vertical distribution to determine the ultimate predation effect on Daphnia.     

Effects of clonality on the genetic variability of rare,insular species: the case of Ruta microcarpa from the Canary Islands

Many plant species combine sexual and clonal reproduction. Clonal propagation has ecological costs mainly related to inbreeding depression and pollen discounting; at the same time, species able to reproduce clonally have ecological and evolutionary advantages being able to persist when conditions are not favorable for sexual reproduction. The presence of clonality has profound consequences on the genetic structure of populations, especially when it represents the predominant reproductive strategy in a population. Theoretical studies suggest that high rate of clonal propagation should increase the effective number of alleles and heterozygosity in a population, while an opposite effect is expected on genetic differentiation among populations and on genotypic diversity. In this study, we ask how clonal propagation affects the genetic diversity of rare insular species, which are often characterized by low levels of genetic diversity, hence at risk of extinction. We used eight polymorphic microsatellite markers to study the genetic structure of the critically endangered insular endemic Ruta microcarpa. We found that clonality appears to positively affect the genetic diversity of R. microcarpa by increasing allelic diversity, polymorphism, and heterozygosity. Moreover, clonal propagation seems to be a more successful reproductive strategy in small, isolated population subjected to environmental stress. Our results suggest that clonal propagation may benefit rare species. However, the advantage of clonal growth may be only short‐lived for prolonged clonal growth could ultimately lead to monoclonal populations. Some degree of sexual reproduction may be needed in a predominantly clonal species to ensure long‐term viability.     

Evolution of pleiotropic alleles for maturation and size as a consequence of predation

Understanding life-history evolution requires knowledge about genetic interactions, physiological mechanisms and the nature of selection. For platyfish, Xiphophorus maculatus, extensive information is available about genetic and physiological mechanisms influencing life-history traits. In particular, alleles at the pituitary locus have large and antagonistic effects on age and size at sexual maturation. To examine how predation affects the evolution of these antagonistic traits, I examined pituitary allele evolution in experimental populations differing in predation risk. A smaller size, earlier maturation allele increased in frequency when predators were absent, while a larger size, later maturation allele increased in frequency when predators were present. Thus, predation favours alleles for larger size, at the cost of later maturation and reproduction. These findings are interesting for several reasons. First, predation is often predicted to favour early reproduction at smaller sizes. Second, few studies have shown how selection acts on alleles that affect age and size at sexual maturation. Finally, many studies assume that trade-offs between these life-history traits result from antagonistic pleiotropy, with alleles that positively affect one trait negatively affecting another, yet this is rarely known. This study unequivocally demonstrates that genetically based trade-offs affect life-history evolution in platyfish.     

Genetic variation in Pueraria lobata (Fabaceae), an introduced, clonal, invasive plant of the southeastern United States

Pueraria lobata (kudzu), a clonal, leguminous vine, is invading the southeastern United States at a rate of 50 000 ha per year. Genetic variability and clonal diversity were measured in 20 southeastern U.S. populations using 14 allozyme loci. Within its U.S. range, 92.9% of the loci were polymorphic and overall genetic diversity was 0.290. Such high levels of genetic diversity are consistent with its history of multiple introductions over an extended period of time. The average proportions of polymorphic loci and genetic diversity within populations were 55.7% (range = 28.6–85.7%) and 0.213 (range = 0.114–0.317), respectively. The proportion of total genetic diversity found among populations was similar to species with equivalent life history characters (G_ST = 0.199). No regional patterns of variation were seen. The number of putative genotypes in each population ranged from 2 to 26. Mean genotypic diversity was 0.694, ranging from 0.223 to 0.955. Such high levels of genotypic diversity indicate that local sites are often colonized by several propagules (most likely seeds) and/or that sexual reproduction occurs within populations after establishment. An excess of heterozygosity was observed in populations with few unique genets, implying that selection for highly heterozygous individuals may occur in populations of P. lobata.     

Environmental correlates of the intrinsic rate of natural increase in primates

Morphological and life history traits of two clones of the cladoceran Daphnia pulex were measured in the presence and absence of size-selective insect predators, the midge larva Chaoborus flavicans, which preys on small Daphnia, and the water bug Notonecta glauca, which preys on large Daphnia. The aim was to detect predator-induced phenotypic changes, particularly the effect of simultaneous exposure to both types of predators. Other work has shown that in the presence of Chaoborus americanus, Daphnia pulex produce a socalled neck spine which may carry several teeth. The morphological modifications are supposed to serve as an anti-predator device. Furthermore, females exposed to Chaoborus often delay their maturation; this has been interpreted as a cost that balances the benefits of the neck teeth. In this investigation, females of both clones produced fewer but larger offspring than control animals when reared in the presence of Chaoborus flavicans. The offspring showed the typical neck spine and delayed first reproduction. In the presence of Notonecta glauca, one of the clones produced more and smaller offspring, and maturation occurred at earlier instars. The other clone also produced more offspring than the control but there was no size difference. When both predators were present, in most cases the reactions of the daphnids were similar to those in the Notonecta experiment. The response to Chaoborus appeared to be suppressed. The observed modifications are interpreted as evolved strategies that reduce the impact of size-selective predation. They are consistent with predictions of life-history theory.     

GENOTYPIC VARIATION AND CLONAL STRUCTURE IN CORAL POPULATIONS WITH DIFFERENT DISTURBANCE HISTORIES

Genotypic diversity in six populations of the endemic Hawaiian reef coral, Porites compressa, was directly related to habitat-disturbance history. The highest diversity (lowest amount of clonal proliferation) was found in populations that had been intensely or recently disturbed. In these populations, space was not limited and mean colony size was small (< 500 cm²), suggesting early stages of recolonization. In an undisturbed, protected habitat, lower genotypic diversity was a result of a significant degree of clonal replication of established genotypes. Unoccupied substratum was rare in this habitat, and average colony size was large (> 2500 cm²). Populations in intermediately disturbed habitats showed intermediate levels of diversity and clonal structure as a result of the combined contributions of sexual and asexual reproduction. Individual clones were distributed over small areas (< 4 m²) or distances (< 1 m) in young populations, and more broadly (> 256 m²) and over longer distances (> 90 m) in the older, undisturbed population. Interpretations of life-history parameters and estimates of total genetic variability in species that have the potential to reproduce asexually are dependent upon an assessment of the overall clonal structure of populations. The power of genotypic assays to reliably detect clonal versus unique colonies, as well as the spatial scales over which clonal populations are sampled, are critical to such assessments.     

The phylogeographical and population genetic approach to the investigation of the genetic diversity patterns in self‐incompatible clonal and polyploid Linnaea borealis subsp. borealis

Surveys of genetic diversity patterns of self‐incompatible clonal polyploid plant species are still scarcer than those of diploid plant species. Therefore, I studied the phylogeographical history of Linnaea borealis subsp. borealis to shed light on the colonization history of this clonal self‐incompatible polyploid plant in Eurasia using selected regions of plastid DNA and genetic diversity patterns of 22 populations of this species employing AFLP markers. I also addressed the question of whether the genetic diversity patterns in L. borealis subsp. borealis in Eurasia are similar to those of earlier published studies of clonal self‐incompatible diploid or polyploid plants. This survey revealed that the shallow phylogeographical history (six plastid haplotypes forming one haplogroup, 100% bootstrap support) and moderate genome‐wide diversity estimated using AFLP markers (Frag_poly = 10.8–38.9%, I = 0.060–0.180, F_ST = 0.289) were general characteristics of L. borealis subsp. borealis in its Eurasian range. The sampling strategy, in most cases at 1–2‐m or even 3–5‐m intervals, showed that a balance between vegetative and sexual reproduction and limited pollen dispersal among compatible mates can be important for genetic diversity patterns in populations of this taxon. Despite the fact that one‐half of the investigated populations were strongly isolated, they still preserved similar levels of genetic diversity across the geographical range. I found no support for the hypothesis that a bottleneck and/or inbreeding had accompanied habitat fragmentation as factors shaping genetic diversity. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 64–76.     

Demographic and life history response of the cladoceranBosmina longirostris to variation in predator abundance

Population dynamics, demography and body size of the cladoceranBosmina longirostris were examined in an experimental study in which the abundance of its predator (the cyprind fishPhoxinus eos) was varied in an unproductive lake. Four densities of fish were used, encompassing the biomass of fish in the lake.Bosmina was most abundant at low and medium fish densities (1.06 and 2.12 g fish biomass · m^-3) and less abundant when fish were either absent or present at high density (3.71 g fish biomass · m^-3). The unimodal response to predator abundance resulted from effects on both birth and death rates.Bosmina birth rates increased as fish biomass increased, in response to increasing food (phytoplankton) biomass. Death rates were highest at high fish biomass (because of fish predation) and in the absence of fish (because of predation by the dipteranChaoborus, which was most abundant in the absence of fish). Size-frequency distributions revealed that fish eliminated the larger size classes ofBosmina, and mean carapace length ofBosmina populations was inversely proportional to fish biomass.Bosmina initiated reproduction at smaller size in the presence of fish than in their absence, and size at maturity was inversely proportional to fish biomass. Size at birth also tended to decrease with increasing fish biomass, but this trend was not as strong as that of size at maturity. Decreased size at maturity apparently allowedBosmina individuals to reproduce before becoming vulnerable to fish predation. Flexibility in size at maturity, together with low abundance of invertebrate predators and large herbivores (which were preyed upon by fish), allowedBosmina to become abundant in low and medium fish treatments. In the high fish treatment, mortality due to fish predation was too severe to be offset by decreased size at maturity, andBosmina population density was low. The net response ofBosmina populations to fish predation results from interactive effects of predation on mortality, natality, and life history traits.     

High genotypic diversity and strong spatial structure in populations of Trifolium alpestre with low seed production

Studies on clonal plants indicate that the proportion between clonal and sexual reproduction can be variable, depending on local habitat conditions and the biological characteristics of the species. In the present study, we assessed this question in Trifolium alpestre by assaying genetic diversity and spatial genotypic structure of natural populations with the use of allozyme markers. Populations revealed high genetic diversity as well as strong spatial structure of multilocus genotypes. The values of genetic diversity were moderately high. Spatially aggregated, identical genotypes spread up to 15 m along linear transects and across 4‐m² plots indicate extensive clonal propagation within populations. However, the existence of numerous unique and small‐sized clones reflects significant contribution from sexual reproduction. Spatially and temporarily stochastic soil disturbances have evidently opened new opportunities for the successful sexual recruitment from the permanent soil seed bank and thus counteracted losses of genetic and genotypic diversity. Seed production in all populations during the three study years was low, in average up to 1.5–2.4 seeds per shoot. The almost total lack of seed set for 57 bagged flower heads on genotypes grown in a common garden indicates that T. alpestre needs pollinators for seed production.     

Genetic variation and clonal structure of the rare,riparian shrub <Emphasis Type="Italic">Spiraea virginiana</Emphasis> (Rosaceae)

Genetic diversity is often considered important for species that inhabit highly disturbed environments to allow for adaptation. Many variables affect levels of genetic variation; however, the two most influential variables are population size and type of reproduction. When analyzed separately, both small population size and asexual reproduction can lead to reductions in genetic variation, although the exact nature of which can be contrasting. Genetic variables such as allelic richness, heterozygosity, inbreeding coefficient, and population differentiation have opposite predictions depending upon the trait (rarity or clonality) examined. The goal of this study was to quantify genetic variation and population differentiation in a species that resides in a highly stochastic environment and is both rare and highly clonal, Spiraea virginiana, and to determine if one trait is more influential genetically than the other. From populations sampled throughout the natural range of S. virginiana, we used microsatellite loci to estimate overall genetic variation. We also calculated clonal structure within populations, which included genotypic richness, evenness, and diversity. Gene flow was investigated by quantifying the relationship between genetic and geographic distances, and population differentiation (θ) among populations. Observed heterozygosity, genotypic richness, and inbreeding coefficients were found to be representative of high clonal reproduction (averaging 0.505, 0.1, and –0.356, respectively) and the number of alleles within populations was low (range = 2.0–3.6), being more indicative of rarity. Population differentiation (θ) among populations was high (average = 0.302) and there was no relationship between genetic and geographic distances. By examining a species that exhibits two traits that both can lead to reduced genetic variation, we may find an enhanced urgency for conservation. Accurate demographic counts of clonal species are rarely, if ever, possible and genetic exploration for every species is not feasible. Therefore, the conclusions in this study can be potentially extrapolated to other riparian, clonal shrubs that share similar biology as S. virginiana.