Genetic consequences of the metapopulation biology of a facultatively sexual freshwater invertebrate

Theoretical models of the impact of a metapopulation structure on the genetics of a species have resulted in multiple predictions that have seldom been empirically evaluated. Here we present microsatellite data from 14 populations of a freshwater bryozoan, Cristatella mucedo, collected along a waterfowl migratory route in north‐western Europe. C. mucedo is facultatively sexual and has the unusual tactic of dispersing via asexually generated propagules. These propagules are likely to be dispersed by waterfowl and therefore the populations that we sampled were expected to maintain some degree of connectivity. Our data illustrate a metapopulation comprising well‐differentiated populations connected by low levels of ongoing gene flow, patterns that agree with predictions based on theoretical work. However, contrary to expectations of a metapopulation, particularly one in which asexual reproduction predominates, genetic variation within populations was often high. This diversity seems to be at least partially attributable to the gene flow that results from ongoing dispersal.     

Reproductive strategy, clonal structure and genetic diversity in populations of the aquatic macrophyte Sparganium emersum in river systems

Many aquatic and riparian plant species are characterized by the ability to reproduce both sexually and asexually. Yet, little is known about how spatial variation in sexual and asexual reproduction affects the genotypic diversity within populations of aquatic and riparian plants. We used six polymorphic microsatellites to examine the genetic diversity within and differentiation among 17 populations (606 individuals) of Sparganium emersum, in two Dutch-German rivers. Our study revealed a striking difference between rivers in the mode of reproduction (sexual vs. asexual) within S. emersum populations. The mode of reproduction was strongly related to locally reigning hydrodynamic conditions. Sexually reproducing populations exhibited a greater number of multilocus genotypes compared to asexual populations. The regional population structure suggested higher levels of gene flow among sexually reproducing populations compared to clonal populations. Gene flow was mainly mediated via hydrochoric dispersal of generative propagules (seeds), impeding genetic differentiation among populations even over river distances up to 50 km. Although evidence for hydrochoric dispersal of vegetative propagules (clonal plant fragments) was found, this mechanism appeared to be relatively less important. Bayesian-based assignment procedures revealed a number of immigrants, originating from outside our study area, suggesting intercatchment plant dispersal, possibly the result of waterfowl-mediated seed dispersal. This study demonstrates how variation in local environmental conditions in river systems, resulting in shifting balances of sexual vs. asexual reproduction within populations, will affect the genotypic diversity within populations. This study furthermore cautions against generalizations about dispersal of riparian plant species in river systems.     

Spatial genetic structure of aquatic bryophytes in a connected lake system

Using genetic markers, we investigated the genetic structure of three clonal aquatic moss species, Calliergon megalophyllum Mikut., Fontinalis antipyretica Hedw. and F. hypnoides Hartm. on two scales: among populations in a connected lake system (large‐scale spatial genetic structure) and among individuals within populations (fine‐scale spatial genetic structure). Mean genetic diversities per population were 0.138, 0.247 and 0.271, respectively, and total diversities equalled 0.223, 0.385 and 0.421, respectively. Relative differentiation levels (F_ST values of 0.173, 0.280 and 0.142, respectively) were significant but showed that there is a moderate amount of gene flow taking place within the lake system connected with narrow streams. Bayesian STRUCTURE analysis provided some indication that the direction of water flow influences population genetic structuring in the studied aquatic mosses. We propose that dispersal leading to gene flow in C. megalophyllum, F. antipyretica and F. hypnoides takes place both along water via connecting streams and by animal vectors, such as waterfowl. Nevertheless, the slight genetic structuring pattern along the direction of water flow suggests that dispersal of shoots or their fragments along water is a means of dispersal in these mosses. The absence of sexual reproduction and spores may have caused the observed spatial genetic structure within populations, including aggregations of similar genotypes (clones or closely related genotypes) at short distances in populations otherwise showing an isolation by distance effect. Regardless of the results pointing to the dominance of vegetative propagation, it is impossible to completely rule out the potential role of rare long‐distance spore dispersal from areas where the species are fertile.     

Sexual and asexual reproduction of the holothurian Stichopus chloronotus (Echinodermata): a comparison between La Réunion (Indian Ocean) and east Australia (Pacific Ocean)

Summary

Stichopus chloronotus (Brandt, 1835) is one among nine aspidochirotide holothurian species known to reproduce both sexually by broadcast spawning and asexually by transverse fission. New data on the sexual cycle of this species in La Réunion are presented here and information available on sexual and asexual reproduction in this species is summarised. Sexual reproduction on La Réunion shows a distinct seasonality with a main spawning period in the warm season (November-February). The spawning period the Great Barrier Reef appears to be at the same time. Some intriguing deviations from unity in sex-ratio, usually biased towards more male individuals, have been observed in both geographic regions (sex ratio at La Reunion 31:1). New data on the asexual reproduction of this species in La Réunion confirm the high rates of fission. The peak of asexual reproduction in both the Indian and Pacific Ocean was observed in winter (June-July). Thus, asexual reproduction in this species occurs outside the season for sexual reproduction. The rate of asexual reproduction appears to vary between sample locations. However, results of population genetic studies on S. chloronotus (Uthicke et al., 1999; Uthicke et al., 2001) indicated that in most populations investigated a maximum of about 60% of all individuals may be derived from sexual recruitment. Cluster analyses on genetic distances between populations grouped populations within Oceans together, with the exception of one sample from a nearshore reef of the GBR. Although genetic differences between the two regions exist, these are relatively small regarding the large geographic distance. We conclude that asexual reproduction in S. chloronotus is important to maintain local population sizes, but that larval exchange between populations mediated by sexual reproduction is important for colonisation of new areas and to provide connectivity between populations. Here, we present the first synthesis of these phenomena for a holothurian species.     

Long‐distance dispersal and genetic structure of natural populations: an assessment of the inverse isolation hypothesis in peat mosses

It is well accepted that the shape of the dispersal kernel, especially its tail, has a substantial effect on the genetic structure of species. Theory predicts that dispersal by fat‐tailed kernels reshuffles genetic material, and thus, preserves genetic diversity during colonization. Moreover, if efficient long‐distance dispersal is coupled with random colonization, an inverse isolation effect is predicted to develop in which increasing genetic diversity per colonizer is expected with increasing distance from a genetically variable source. By contrast, increasing isolation leads to decreasing genetic diversity when dispersal is via thin‐tailed kernels. Here, we use a well‐established model group for dispersal biology (peat mosses: genus Sphagnum) with a fat‐tailed dispersal kernel, and the natural laboratory of the Stockholm archipelago to study the validity of the inverse isolation hypothesis in spore‐dispersed plants in island colonization. Population genetic structure of three species (Sphagnum fallax, Sphagnum fimbriatum and Sphagnum palustre) with contrasting life histories and ploidy levels were investigated on a set of islands using microsatellites. Our data show (, amova , IBD) that dispersal of the two most abundant species can be well approximated by a random colonization model. We find that genetic diversity per colonizer on islands increases with distance from the mainland for S. fallax and S. fimbriatum. By contrast, S. palustre deviates from this pattern, owing to its restricted distribution in the region, affecting its source pool strength. Therefore, the inverse isolation effect appears to hold in natural populations of peat mosses and, likely, in other organisms with small diaspores.     

Parentage analyses identify local dispersal events and sibling aggregations in a natural population of Millepora hydrocorals,a free‐spawning marine invertebrate

Dispersal is a critical process for the persistence and productivity of marine populations. For many reef species, there is increasing evidence that local demography and self‐recruitment have major consequences on their genetic diversity and adaptation to environmental change. Yet empirical data of dispersal patterns in reef‐building species remain scarce. Here, we document the first genetic estimates of self‐recruitment and dispersal distances in a free‐spawning marine invertebrate, the hydrocoral Millepora cf. platyphylla. Using twelve microsatellite markers, we gathered genotypic information from 3,160 georeferenced colonies collected over 27,000 m² of a single reef in three adjacent habitats in Moorea, French Polynesia; the mid slope, upper slope, and back reef. Although the adult population was predominantly clonal (85% were clones), our parentage analysis revealed a moderate self‐recruitment rate with a minimum of 8% of sexual propagules produced locally. Assigned offspring often settled at <10 m from their parents and dispersal events decrease with increasing geographic distance. There were no discrepancies between the dispersal distances of offspring assigned to parents belonging to clonal versus nonclonal genotypes. Interhabitat dispersal events via cross‐reef transport were also detected for sexual and asexual propagules. Sibship analysis showed that full siblings recruit nearby on the reef (more than 40% settled at <30 m), resulting in sibling aggregations. Our findings highlight the importance of self‐recruitment together with clonality in stabilizing population dynamics, which may ultimately enhance local sustainability and resilience to disturbance.     

CONTRIBUTIONS OF SEXUAL AND ASEXUAL REPRODUCTION TO POPULATION STRUCTURE IN THE CLONAL SOFT CORAL,ALCYONIUM RUDYI

Numerous studies of population structure in sessile clonal marine invertebrates have demonstrated low genotypic diversity and nonequilibrium genotype frequencies within local populations that are monopolized by relatively few, highly replicated genets. All of the species studied to date produce planktonic sexual propagules capable of dispersing long distances; despite local genotypic disequilibria, populations are often panmictic over large geographic areas. The population structure paradigm these species represent may not be typical of the majority of clonal invertebrate groups, however, which are believed to produce highly philopatric sexual propagules. I used allozyme variation to examine the population structure of the temperate soft coral, Alcyonium rudyi, a typical clonal species whose sexually produced larvae and asexually produced ramets both have very low dispersal capabilities. Like other clonal plants and invertebrates, the local population dynamics of A. rudyi are dominated by asexual reproduction, and recruitment of new sexually produced genets occurs infrequently. As expected from its philopatric larval stage, estimates of genetic differentiation among populations of A. rudyi were highly significant at all spatial scales examined (mean θ = 0.300 among 20 populations spanning a 1100-km range), suggesting that genetic exchange seldom occurs among populations separated by as little as a few hundred meters. Mapping of multilocus allozyme genotypes within a dense aggregation of A. rudyi ramets confirmed that dispersal of asexual propagules is also very limited: members of the same genet usually remain within < 50 cm of one another on the same rock surface. Unlike most previously studied clonal invertebrates, populations of A. rudyi do not appear to be dominated by a few widespread genets: estimates of genotypic diversity (G_o) within 20 geographically distinct populations did not differ from expectations for outcrossing, sexual populations. Despite theoretical suggestions that philopatric dispersal combined with typically small effective population sizes should promote inbreeding in clonal species, inbreeding does not appear to contribute significantly to the population structure of A. rudyi. Genet genotype frequencies conformed to Hardy-Weinberg expectations in all populations, and inbreeding coefficients (f) were close to zero. In general, the population structure of A. rudyi did not differ significantly from that observed among outcrossing sexual species with philopatric larval dispersal. Age estimates suggest, however, that genets of A. rudyi live for many decades. Genet longevity may promote high genotypic diversity within A. rudyi populations and may be the most important evolutionary consequence of clonal reproduction in this species and the many others that share its dispersal characteristics.     

Using genetic evaluation to guide conservation of remnant Juniperus communis (Cupressaceae) populations

• Many critically endangered plant species exist in small, genetically depauperate or inbred populations, making assisted gene flow interventions necessary for long‐term population viability. However, before such interventions are implemented, conservation practitioners must consider the genetic and demographic status of extant populations, which are strongly affected by species’ life‐history traits. In northwestern Europe, Juniperus communis, a dioecious, wind‐pollinated and bird‐dispersed gymnosperm, has been declining for the past century and largely exists in small, isolated and senescent populations.
• To provide useful recommendations for a recovery plan involving translocation of plants, we investigated genetic diversity and structure of populations in Belgium using four microsatellite and five plastid single‐nucleotide polymorphism (SNP) markers.
• We detected no clonality in the populations, suggesting predominantly sexual reproduction. Populations exhibited high genetic diversity (H_e = 0.367–0.563) and low to moderate genetic differentiation (F_ST ≤ 0.133), with no clear geographic structure. Highly positive inbreeding coefficients (F_IS = 0.221–0.507) were explained by null alleles, population substructuring and biparental inbreeding. No isolation by distance was observed among distant populations, but isolation at close geographic proximity was found. Patterns were consistent with high historical gene flow through pollen and seed dispersal at both short and long distances. We also tested four pre‐germination treatments among populations to improve germination rates; however, germination rates remained low and only cold‐stratification treatments induced germination in some populations.
• To bolster population regeneration, introductions of cuttings from several source populations are recommended, in combination with in situ management practices that improve seedling survival and with ex situ propagation.

     

Dispersal potential of spores and asexual propagules in the epixylic hepatic Anastrophyllum hellerianum

Dispersal ability is of great importance for plants, which commonly occupy spatially and temporally limited substrate patches. Mixed reproductive strategies with abundant diaspore production are favoured in a heterogeneous landscape to ensure successful colonisation at different distances. In bryophytes, long-distance dispersal has been thought to take place primarily by spores, while asexual propagules are important in local dispersal and in the maintenance of colonies. In the present study, we investigated the dispersal potential of two equally sized propagules, sexually formed spores and asexually produced gemmae in the dioecious, epixylic hepatic, Anastrophyllum hellerianum, which inhabits spatially and temporally limited substrate patches. We trapped propagules at different distances (0–10 m) and directions from the source colonies in two experiments: one in a natural habitat within a forest and another involving an artificial set-up in an open habitat. Spore dispersal showed only slight distance dependence both in the open and the forest habitats, presumably as a consequence of wind affecting the dispersal pattern. Gemma dispersal was more strongly distance-dependent in the open habitat than in the forest sites. Considerably more gemmae were deposited during rainy than dry periods, possibly because of the effect of rain drops on gemma release. However, weather conditions had no effect on the dispersal patterns of spores or gemmae. In A. hellerianum, the combination of occasional spore production and practically continuous, massive gemma production facilitates dispersal both on local scale and over long distances. Unlike previously assumed, not only spores but also the asexual propagules may contribute to long-distance dispersal, thus allowing considerable gene flow at the landscape level.     

Biogeography of sexual and asexual populations in Bostrychia moritziana (Rhodomelaceae, Rhodophyta)

Bostrychia moritziana (Sonder ex Kützing) J. Agardh is recorded from many regions around the world. Our laboratory culture investigations have verified a sexual life cycle in isolates from Australia, Venezuela, Colombia, South Africa, Fiji, New Zealand and Indonesia. By contrast, asexual isolates producing successive generations of tetrasporophytes in laboratory culture and, presumably, in the field, are known from Australia. New Caledonia and Japan. In Australia, asexual reproduction is absent only in Victoria. In Western Australia, Northern Territory and Queensland, 99% of the isolates have asexual reproduction. In New South Wales (NSW), asexual and sexual populations are often intermixed. Of the 176 worldwide field collections, 58% were vegetative, 39% were tetrasporic, 2% were female and 1% were male. After several years of observations on the asexual isolates in culture, at least 30 successive asexual tetrasporophytic generations have developed. Only two asexual isolates (3558 and 3575) from NSW have formed a single male and female gametophyte in culture. In a self-cross of 3568, the carpospores developed into tetrasporophytes that recycled asexually. All outcrosses done with normal sexual isolates produced normal carposporophytes and the carpospores developed into tetrasporophytes that also recycled asexually. Asexual populations may arise repeatedly by loss of meiosis in tetrasporangia of sexual populations. Asexual reproduction apparently does not diminish the overall dispersal and abundance in the field. Our present bio-geographic data show that sexually reproducing popuiations of B. moritziana occur worldwide, while asexuaily reproducing populations are confined to the western Pacific. Bostrychia bispora West et Zuccarello, initially described on the basis of its asexual reproduction to distinguish it from B, moritziana, is now reduced to synonymy with B. moritziana.     

Microgeographic variation in recruitment under adult trees: arrival of new genotypes or perpetuation of the existing ones?

• Investigating spatial variation in the relative importance of sexual reproduction and clonal propagation is critical to obtain more accurate estimates of future effective population sizes and genetic diversity, as well as to identify ecological correlates of clonality.
• We combined a stratified sampling scheme with microsatellite genetic analyses to estimate variation in the proportion of sexual versus clonal recruits among saplings in five populations of the tree Pyrus bourgaeana. Using a likelihood framework, we identified clones among the genotypes analysed and examined variation among populations regarding the proportion of saplings coming from clonal propagation. We also examined the relationship between the relative abundance of clonal shoots across the studied populations and their herbivory levels.
• Our results revealed that one third of the saplings examined (N = 225 saplings) had a probability above 0.9 of being clones of nearby (<10 m) trees, with the ratio between clonal propagation and sexual recruitment varying up to eight‐fold among populations. A small portion of these putative clonal shoots reached sexual maturity. Relative abundance of clonal shoots did not significantly relate to the herbivory by ungulates.
• Our results call into question optimistic expectations of previous studies reporting sufficient levels of recruitment under parental trees without animal seed dispersal services. Nevertheless, given that some of these clonal shoots reach sexual maturity, clonal propagation can ultimately facilitate the long‐term persistence of populations during adverse periods (e.g. environmental stress, impoverished pollinator communities, seed dispersal limitation).

     

Estimating seed dispersal distance: A comparison of methods using animal movement and plant genetic data on two primate‐dispersed Neotropical plant species

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Molecular evidence for limited dispersal of vegetative propagules in the epiphytic lichen Lobaria pulmonaria

Propagation, whether sexual or asexual, is a fundamental step in the life cycle of every organism. In lichenized fungi, a great variety of vegetative propagules have evolved in order for the symbiotic partners to disperse simultaneously. For lichens with the ability of sexual and asexual reproduction, the relative contribution of vegetative dispersal is unknown but could, nonetheless, be inferred by studying genotype distribution. The genetic structure of three Lobaria pulmonaria (Lobariaceae) populations from Switzerland was investigated based on the observed variation at six microsatellite loci. All three populations had a clustered distribution of identical genotypes at small spatial scales. The maximum distance between identical genotypes was 230 m. At a distance of 350 m from a source tree, seemingly suitable habitat patches were too far apart to be colonized. Some multilocus genotypes were frequent within local populations but no genotypes were shared among populations. The restricted occurrences of common genotypes as well as the clustered distributions are evidence for a limited dispersal of vegetative propagules in L. pulmonaria. Gene flow among isolated populations will ultimately depend on the capacity of long-distance dispersal and thus probably depend on sexual reproduction.     

Species distribution modelling and cpSSR reveal population history of the Neotropical annual herb Tithonia rotundifolia (Asteraceae)

• The impacts of the historical geologic and climatic events on the diversity and genetic structure of Neotropical taxa have recently become a subject of study. However, annual plants associated with tropical dry forests remain under‐studied. The exploration of additional taxa in contrasting environments will improve the current understanding of responses of the Neotropical biota to these events. Here, we explore the species distribution and geographic structure of the annual herb Tithonia rotundifolia.
• We sampled 175 individuals from 19 populations of T. rotundifolia. Species distribution modelling and six microsatellite chloroplast loci were used to infer its population history. We identified areas of historical climate suitability and then tested if there is genetic structuring among these areas.
• Haplotypes showed strong phylogeographic structure. Historical climatic suitability areas were found along the Pacific coast; however, a gap was found at the Isthmus of Tehuantepec (IT). Although Bayesian analysis showed population structuring, amova revealed that the IT is not its main driver. Instead, a subdivision into a higher number of regions had higher F_CT values. Also, populations to the east of the IT showed evidence of recent population expansion and migration in a south–north direction.
• Pleistocene climate fluctuations partially explain the geographic structure of T. rotundifolia. However, life‐history characteristics such as limited seed dispersal and the patchy distribution of suitable habitats explain the high haplotype diversity and population sub‐structuring and diversity. Lastly, the absence of geographic structure of some haplotypes may indicate long‐distance dispersal, or hybridisation with the closely related T. tubaeformis.

     

Genetic differentiation amongst populations of the coral Pocillopora damicornis off southwestern Australia

An electrophoretic study of genetic differentiation amongst local populations of the reef-coral Pocillopora damicornis was used to group coral heads into units defined as the area of effective dispersal of a clone, and termed colonies. For reefs off southwestern Australia, colonies were usually under a few hundred metres in extent. Although most new recruits within a colony were derived asexually, sexually produced propagules acted to connect populations outside the boundaries of a colony. Such connections were weak, and allelic frequencies varied considerably over a few kilometres. The primary agent of genetic differentiation was suggested to be the small effective population size resulting from the asexual proliferation of a few genotypes at any site. The effective number of genotypes per colony was approximately six. Asexual reproduction appears also to limit gene flow and accentuate selection in this species.     

Distance-dependent performance of asexual progeny in Allium vineale (Liliaceae)

Local adaptation within and among populations may have an impact on processes ranging from speciation to the evolution of mixed breeding systems and dispersal strategies. It is also one potential factor that could favor the production of asexual over sexual propagules. This field experiment tested whether asexually produced bulbils of Allium vineale demonstrate local adaptation to the parental microsite at the scale of natural dispersal from the parent (5, 25, 50, 100, and 1000 cm). Both "home' and randomly chosen "away' genotypes were planted at each location to determine the relative performance of the "home' genotype. Overall, bulbil performance declined with distance from the parent. In particular, "home' bulbils outperformed "away' bulbils at a distance of 25 cm from the parent, indicating that local adaptation has occurred at the scale of natural dispersal in this species. The variance in propagule performance also increased at farther distances from the parent, indicating that the predictability of offspring performance decreases with distance. Fine-scale local adaptation within the range of seed dispersal in this population may be one factor favoring asexual reproduction in Allium vineale.     

Crayfish invasion facilitates dispersal of plants and invertebrates by gulls

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Large‐scale,multidirectional larval connectivity among coral reef fish populations in the Great Barrier Reef Marine Park

Larval dispersal is the key process by which populations of most marine fishes and invertebrates are connected and replenished. Advances in larval tagging and genetics have enhanced our capacity to track larval dispersal, assess scales of population connectivity, and quantify larval exchange among no‐take marine reserves and fished areas. Recent studies have found that reserves can be a significant source of recruits for populations up to 40 km away, but the scale and direction of larval connectivity across larger seascapes remain unknown. Here, we apply genetic parentage analysis to investigate larval dispersal patterns for two exploited coral reef groupers (Plectropomus maculatus and Plectropomus leopardus) within and among three clusters of reefs separated by 60–220 km within the Great Barrier Reef Marine Park, Australia. A total of 69 juvenile P. maculatus and 17 juvenile P. leopardus (representing 6% and 9% of the total juveniles sampled, respectively) were genetically assigned to parent individuals on reefs within the study area. We identified both short‐distance larval dispersal within regions (200 m to 50 km) and long‐distance, multidirectional dispersal of up to ~250 km among regions. Dispersal strength declined significantly with distance, with best‐fit dispersal kernels estimating median dispersal distances of ~110 km for P. maculatus and ~190 km for P. leopardus. Larval exchange among reefs demonstrates that established reserves form a highly connected network and contribute larvae for the replenishment of fished reefs at multiple spatial scales. Our findings highlight the potential for long‐distance dispersal in an important group of reef fishes, and provide further evidence that effectively protected reserves can yield recruitment and sustainability benefits for exploited fish populations.     

Managing diversity: Domestication and gene flow in Stenocereus stellatus Riccob. (Cactaceae) in Mexico

Microsatellite markers (N = 5) were developed for analysis of genetic variation in 15 populations of the columnar cactus Stenocereus stellatus, managed under traditional agriculture practices in central Mexico. Microsatellite diversity was analyzed within and among populations, between geographic regions, and among population management types to provide detailed insight into historical gene flow rates and population dynamics associated with domestication. Our results corroborate a greater diversity in populations managed by farmers compared with wild ones (H_E = 0.64 vs. 0.55), but with regional variation between populations among regions. Although farmers propagated S. stellatus vegetatively in home gardens to diversify their stock, asexual recruitment also occurred naturally in populations where more marginal conditions have limited sexual recruitment, resulting in lower genetic diversity. Therefore, a clear‐cut relationship between the occurrence of asexual recruitment and genetic diversity was not evident. Two managed populations adjacent to towns were identified as major sources of gene movement in each sampled region, with significant migration to distant as well as nearby populations. Coupled with the absence of significant bottlenecks, this suggests a mechanism for promoting genetic diversity in managed populations through long distance gene exchange. Cultivation of S. stellatus in close proximity to wild populations has led to complex patterns of genetic variation across the landscape that reflects the interaction of natural and cultural processes. As molecular markers become available for nontraditional crops and novel analysis techniques allow us to detect and evaluate patterns of genetic diversity, genetic studies provide valuable insights into managing crop genetic resources into the future against a backdrop of global change. Traditional agriculture systems play an important role in maintaining genetic diversity for plant species.     

Local population genetic structure of the montane caddisfly Drusus discolor is driven by overland dispersal and spatial scaling

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