Phenotypic differentiation in fitness related traits between populations of an extremely rare sunflower: Conservation management of isolated populations

Knowledge of the genetic and demographic consequences of rarity is crucial when evaluating the effects of habitat loss and fragmentation on population viability, and for creating management plans in rare plant species. Reduction in population size and in the number of populations can lead to decreased genetic diversity and increased inbreeding. Genetic diversity is often correlated with fitness and is frequently used to identify populations of greatest conservation concern, or those that may be good candidates for ex situ conservation programs. However, an association between these factors is not always clear, and crossing studies evaluating whether there is phenotypic differentiation among populations in fitness related traits can inform managers of suffering populations or good sources for ex situ materials. Crossing studies can also evaluate the potential for genetic rescue to boost fitness in suffering populations. To address these questions, we conducted two generations of controlled crosses between populations of the extremely rare and fragmented sunflower, Helianthus verticillatus. We measured achene viability, germination, survival, and pollen viability (F₁ only) in 176 F₁ and 159 F₂ families. The populations were differentiated with respect to phenotypic fitness measures with one population having significantly lower achene viability and germination. Also, the potential for genetic rescue was observed as gene flow into the less fit population resulted in higher fitness measures in both the F₁ and F₂. Results are discussed with respect to the importance of combining genetic marker data with crosses and the implications for conservation in disjunct populations of rare species.     

Population differentiation and genetic variation inform translocation decisions for Liatris scariosa var. novae-angliae, a rare New England grassland perennial

Augmentation of small rare species populations is sometimes suggested on genetic grounds. However, outbreeding depression via dilution of local adaptation or break-up of genomic coadaptation may occur. These effects depend on the causes of population divergence. Here, we compare genetic measures of population divergence in Liatris scariosa var. novae-angliae, a rare New England perennial. We measured G_ST, neutral marker subdivision, and Q_ST, quantitative subdivision of propagule and juvenile plant traits. G_ST was relatively high. Q_ST for leaf shape exceeded G_ST, indicating local adaptation, while Q_ST for other traits fell within or below the G_ST range. Local adaptation appears low for juvenile traits, although the high G_ST cautions against translocation because of potential coadaptation. If translocation is still required, however, donor populations should contain high quantitative genetic diversity. We assess population size and allozyme diversity as predictors of quantitative genetic variation, but find these poor proxies for direct measurement.     

Microsatellite analysis of genetic population structure of the endangered cyprinid Anaecypris hispanica in Portugal: implications for conservation

The endangered fish species Anaecypris hispanica is restricted to eight disjunct populations in the Portuguese Guadiana drainage. The genetic structure of these populations was studied in order to determine levels of genetic variation within and among populations and suggest implications for conservation of the species. Based on five microsatellite loci, the null hypothesis of population homogeneity was tested. Tests for genetic differentiation revealed highly significant differences for pairwise comparisons between all populations, and substantial overall population subdivision (F_ST=0.112). All sampled populations contained unique alleles. Our findings indicate marked genetic structuring and emphasise limited dispersal ability. The high levels of genetic diversity detected within and among A. hispanica populations suggest, however, that the observed fragmentation and reduction in population size of some populations during the last two decades, has impacted little on levels of genetic variability. Data imply that most A. hispanica populations should be managed as distinct units and that each has a high conservation value containing unique genetic variation. It is argued that geographic patterns of genetic structuring indicate the existence of eight management units.     

AFLP fingerprinting of Anchusa (Boraginaceae) in the Corso-Sardinian system: Genetic diversity, population differentiation and conservation priorities in an insular endemic group threatened with extinction

Although several plants endemic to Corsica and Sardinia are included in various redlists, no attempts have been made to analyse their genetic diversity with molecular techniques. Genus Anchusa occurs with seven taxa in either mountain or coastal habitats of the two islands, but the very restricted range and low population size pose these endemisms in a very precarious conservation status. Highly variable markers (AFLP) were therefore used to analyse the patterns and levels of genetic diversity in a sample of 11 populations from the entire range of the group.Results indicate the separation between a mountain genic pool including Anchusa formosa, Anchusa capellii and Anchusa montelinasana, and four groups of coastal accessions. In spite of small size, mountain taxa show low interpopulation differentiation (F_st = 0.02) and relatively high intrapopulation genetic variation (0.365), while coastal accessions showed on average a stronger differentiation (mean F_st = 0.20) and a lower diversity (0.281), possibly due to higher rates of inbreeding. The particularly low levels of variation found in A. sardoa, A. littorea and A. crispa ssp. maritima from the Coghinas bay are likely due to a historical decrease of populations and to bottleneck events caused by loss of habitat and natural stochastic factors on sand dune ecosystems. While habitat maintenance and regulation of grazing by domestic herbivores should be sufficient to ensure the persistence of the mountain endemics, additional actions of in situ and ex situ conservation are needed for the critically endangered coastal species A. sardoa and A. littorea. A. crispa showed a relatively high variation, especially on Corsica. No correlation between population size and genetic variation was found in the latter species, highlighting the importance of the small patches for its conservation. Also, the genetic separation between subspecies crispa and maritima stresses the need of keeping them distinct in redlists and conservation actions on Sardinia.     

Genetic diversity and local population structure of fragmented populations of Trillium camschatcense (Trilliaceae)

Trillium camschatcense, a long-lived common woodland herb, has been experiencing intensive habitat fragmentation over the last century in eastern Hokkaido, Japan. We examined the genetic diversity and population genetic structure of 12 fragmented populations with different population sizes using allozyme electrophoresis. The percentage of polymorphic loci and mean number of alleles per locus were positively related to population size, probably due to the stochastic loss of rare alleles (frequency of q<0.1) in small populations. Populations with 350 flowering plants or fewer had lost almost all of their rare alleles. While the heterozygosity and inbreeding coefficient were not related to population size, some small populations showed relatively high inbreeding coefficients. In spite of the low genetic differentiation among overall populations (F_ST=0.130), local population structuring was recognized between the two geographically discontinuous population groups. Within groups, sufficient historical gene flow was inferred, whereas a low dispersal ability of this species and geographical separation could produce apparent differentiation between groups.     

Habitat fragmentation reduces genetic diversity and connectivity among toad populations in the Brazilian Atlantic Coastal Forest

Tropical rainforests are becoming increasingly fragmented and understanding the genetic consequences of fragmentation is crucial for conservation of their flora and fauna. We examined populations of the toad Rhinella ornata, a species endemic to Atlantic Coastal Forest in Brazil, and compared genetic diversity among small and medium forest fragments that were either isolated or connected to large forest areas by corridors. Genetic differentiation, as measured by F_ST, was not related to geographic distance among study sites and the size of the fragments did not significantly alter patterns of genetic connectivity. However, population genetic diversity was positively related to fragment size, thus haplotype diversity was lowest in the smallest fragments, likely due to decreases in population sizes. Spatial analyses of genetic discontinuities among groups of populations showed a higher proportion of barriers to gene flow among small and medium fragments than between populations in continuous forest. Our results underscore that even species with relatively high dispersal capacities may, over time, suffer the negative genetic effects of fragmentation, possibly leading to reduced fitness of population and cases of localized extinction.     

Spatial demographic and genetic consequences of harvesting within populations of the terrestrial orchid Cymbidium goeringii

Populations of many orchids, especially terrestrial species, have been rapidly decreasing due to mass collection by plant sellers and enthusiasts. Given the presumably negative demographic and genetic consequences, such anthropogenic activity should be taken into consideration for predicting ecological and evolutionary dynamics and for planning conservation strategies. To determine how recent human disturbance alters spatial demographic and population genetic processes, populations of the terrestrial orchid Cymbidium goeringii located in South Korea were examined to quantify the spatial distributions of individuals and genotypes with respect to three levels of disturbance: “disturbed” (four populations), “putatively disturbed” (two), and “undisturbed” (two). Undisturbed and putatively disturbed populations were found to possess significantly positive spatial clustering of individuals over a range of spatial scales. In contrast, disturbed populations exhibited little or no spatial aggregation, consistent with the selective removal of plants by collectors from higher density areas within these populations. Although overall genetic differentiation among populations was moderate and significant (F_ST = 0.082), levels of genetic diversity within populations were similar despite the different disturbance histories (mean H_e = 0.257-0.324). Spatial genetic autocorrelation analyses revealed that the undisturbed populations exhibited significant declines in kinship (F_ij) with distance, that mean kinship at interplant distances of ?4-6 m was significantly greater than zero and between plants ?0.5 m apart was in the range expected for first cousins to half-sibs. In contrast, only one putatively disturbed and one disturbed population exhibited significant declines in kinship with distance. These differences between disturbed versus putatively disturbed and undisturbed populations in the spatial distribution of individuals and genetic variation likely reflect the consequences of mass collections. Since these differences (and reduced population density) have important implications for future ecological and evolutionary trajectories, conservation managers of endangered terrestrial orchids may want to analyze the spatial distribution of individuals and their genotypes to infer whether a population with few individuals represents a natural state or the likely outcome of mass collection.     

Low levels of genetic variation among southern peripheral populations of the threatened herb, Leontice microrhyncha (Berberidaceae) in Korea

Levels of genetic variation and intrapopulation genetic structures of Leontice microrhyncha S. Moore (Berberidaceae) were assessed for six populations in South Korea, representing the southern most range of a species found in Northeast China and the Korean peninsula. Detected genetic diversity (H_es) was very low (0.024) and F_IS values showed large heterozygote deficiencies. The small percentage of polymorphic loci and numbers of alleles per locus suggest that L. microrhyncha has a history of severe or long-lasting population bottlenecks that have eroded genetic diversity. This study suggests that the Korean population appears to consist of two historically isolated and independently evolving populations. It seems likely that these groups have been isolated and unstable for a significant period of time. However, the effects of recent habitat fragmentation on the historically disjunct and fragmented population system found in L. microrhyncha were not those predicted from the lack of significant relationships between population-level patterns of genetic variation and population sizes. Most non-unique genotypes were shared by most individuals and the lower level of diversity, high levels of inbreeding and population differentiation as well as high rate of seed production indicated that this species is autogamous and self-compatible and probably largely selfing. Therefore, to preserve extant genetic variation, all populations must be protected across the small geographic range of the species to retain both allelic and genotypic diversity.     

Genetic structure, population dynamics, and conservation of Black caiman (Melanosuchus niger)

Microsatellite DNA polymorphisms were screened in seven populations of the largest Neotropical predator, the Black caiman Melanosuchus niger (n = 169), originating from Brazil, French Guiana and Ecuador. Eight loci were used, for a total of 62 alleles. The Ecuadorian population had the lowest number of alleles, heterozygosity and gene diversity; populations of the Guianas region exhibited intermediate diversities; highest values were recorded in the two populations of the Amazon and Rio Negro. During the last century Melanosuchus populations have been reduced to 1-10% of their initial levels because of hunting pressure, but no strong loss of genetic diversity was observed. Both the inter-locus g-test and the Pk distribution suggested no recent important recovery and/or expansion of current populations. On a global scale, the inter-population variation of alleles indicated strong differentiation (F_ST = 0.137).Populations were significantly isolated from each other, with rather limited gene flow; however, these gene flow levels are sufficiently high for recolonization processes to effectively act at regional scales. In French Guiana, genetic structuring is observed between populations of two geographically close but ecologically distinct habitats, an estuary and a swamp. Similar divergence is observed in Brazil between geographically proximate “black water” and “white water” populations. As a consequence, the conservation strategy of the Black caiman should include adequate ecosystem management, with strong attention to preservation of habitat integrity. Distribution of genetic diversity suggests that current populations originated from the central Amazonian region. Dispersal of the species may thus have been deeply influenced by major climatic changes during the Holocene/Pleistocene period, when the Amazonian hydrographic networks were altered. Major ecological changes such as glaciations, marine transgressions and a hypothesized presence of an Amazonian Lake could have resulted in extension of Black caiman habitats followed by isolation.     

Unexpected high levels of genetic variability and the population structure of an island endemic rodent (Oryzomys couesi cozumelae)

Oryzomys couesi cozumelae is an endemic, threatened rodent from Cozumel Island, Mexico. We estimated its genetic diversity and structure by analyzing microsatellite loci in 228 samples from 12 sampling sites widely distributed throughout the island. Unexpected high levels of genetic and allelic diversity were found: a total of 54 alleles, an average of 10.8 alleles per locus, and high heterozygosity values (mean H_O = 0.624, H_E = 0.690 and H_Nei = 0.689). These values are higher than those reported for small sized insular mammals, higher than that found in 37 individuals of the mainland O. couesi from southern Mexico (H_O = 0.578) that we analyzed for comparative purposes, and similar to those of other mainland small mammal populations. Despite factors that affect Cozumel’s biota, such as exotic predators and competitors, hurricanes, seasonal population fluctuations and anthropogenic activities, no evidence of genetic bottlenecks was found. A significant population structure was observed and a model of isolation-by-distance was supported. Our findings render O. c. cozumelae a high conservation value, not only for its high genetic diversity and structure, but because available data suggests that its population has declined significantly in recent years. Further habitat fragmentation and population isolation could result in a higher genetic structure and loss of genetic diversity. The protection of habitat, the maintenance of habitat connectivity and the removal of introduced competitors and predators are a conservation priority. Acknowledging that the genetic structure of populations has crucial conservation implications, the present genetic information should be taken into account in management plans for the conservation of O. c. cozumelae.     

Increased genetic differentiation in house sparrows after a strong population decline: From panmixia towards structure in a common bird

A drastic decline in abundance prompts conservation measures, even though a species may still be common, partly because such a decline may be associated with loss of genetic variability. Longitudinal evidence (i.e. repeated measures across time) for loss of genetic diversity is scarce and mostly concerns organisms that have experienced a severe bottleneck. Here, we study the house sparrow in Finland, where a strong (50–86%) reduction in abundance occurred in four decades, starting earlier and resulting steeper decline in the south than in the north. Based on thirteen polymorphic microsatellites, we compared 12 Finnish populations both prior (mid-1980s) and after (2009) the major population decline. There was no evidence of bottlenecks and only little loss of genetic variation, but we found a significant threefold increase in genetic differentiation (F_ST) across the populations. This may reflect a non-equilibrium situation between the rates of change in the genetic diversity and differentiation and indicate future loss of genetic diversity. Our findings indicate that a strong decline in population size in a relatively common species still leaves a noticeable population-genetic imprint and warrants conservation concern.     

Conserving marginal populations of the food plant (Impatiens noli-tangere) of an endangered moth (Eustroma reticulatum) in a changing climate

Impatiens noli-tangere is scarce in the UK and probably only native to the Lake District and Wales. It is the sole food plant for the endangered moth Eustroma reticulatum. Significant annual fluctuations in the size of I. noli-tangere populations endanger the continued presence of E. reticulatum in the UK. In this study, variation in population size was monitored across native populations of I. noli-tangere in the English Lake District and Wales. In 1998, there was a crash in the population size of all metapopulations in the Lake District but not of those found in Wales. A molecular survey of the genetic affinities of samples in 1999 from both regions and a reference population from Switzerland was performed using AFLP and ISSR analyses. The consensus UPGMA dendrogram and a PCO scatter plot revealed clear differentiation between the populations of I. noli-tangere in Wales and those in the Lake District. Most of the genetic variation in the UK (H_T=0.064) was partitioned between (G_ST=0.455) rather than within (H_S=0.034) regions, inferring little gene flow occurs between regions. There was similar bias towards differentiation between metapopulations in Wales, again consistent with low levels of interpopulation gene flow. This contrasts with far lower levels of differentiation in the Lake District which suggests modest rates of gene flow may occur between populations. It is concluded that in the event of local extinction of sites or populations, reintroductions should be restricted to samples collected from the same region. We then surveyed climatic variables to identify those most likely to cause local extinctions. Climatic correlates of population size were sought from two Lake District metapopulations situated close to a meteorological station. A combination of three climatic variables common to both sites explained 81-84% of the variation in plant number between 1990 and 2001. Projected trends for these climatic variables were used in a Monte Carlo simulation which suggested an increased risk of I. noli-tangere population crashes by 2050 at Coniston Water, but not at Derwentwater. Implications of these findings for practical conservation strategies are explored.     

Distinguishing historical fragmentation from a recent population decline - shrinking or pre-shrunk skink from New Zealand?

Species that are rare when first described present a practical management problem because it may be unclear whether the taxon is in the final stages of an anthropogenic decline, or is naturally uncommon, and each scenario dictates a distinct approach to management. We analysed mitochondrial and microsatellite DNA data with population genetic and phylogeographic tools to distinguish between these possibilities in a rare lizard from southern New Zealand. Grand skinks, Oligosoma grande, are large rock-dwelling lizards that have a fragmented distribution consisting of a western and eastern cluster of populations separated by ca. 120 km. This distribution could result from human disturbance, pre-human climatic and vegetation changes, or both. All populations were highly genetically structured (overall F_ST 0.171, R_ST 0.235), indicating that populations were demographically independent and skinks are unlikely to expand their range without human intervention. In addition, the current fragmented distribution is likely to have both historical and recent anthropogenic elements. Two eastern populations showed evidence of being historically large (high θ mtDNA genetic diversity), although they are now small, supporting anecdotal data that grand skinks have declined in historical times. However, eastern and western populations were reciprocally monophyletic for mtDNA lineages, suggesting long independent evolutionary histories that predate the arrival of humans in New Zealand. Eastern and western populations fulfil many criteria to be considered as evolutionarily significant units, but such a classification must be balanced against addressing more immediate threats to the species’ survival, such as introduced predators.     

Genetic Diversity in Tunisian Ceratonia siliqua L. (Caesalpinioideae) Natural Populations

The last two centuries witnessed the human-caused fragmentation of Tunisian Ceratonia siliqua L. (Caesalpinoideae) populations which were often represented by scattered individuals. Seventeen populations growing in four bioclimatic zones: sub-humid, upper semi-arid, mean semi-arid and lower semi-arid zones, were sampled for allozyme diversity to assess their genetic diversity and structuration using eight isozymes revealed by starch gel electrophoresis. The species showed high diversity within populations. The average number of alleles per polymorphic locus was 1.98, the percentage of polymorphic loci was 83.4% and the mean observed heterozygosity (Ho) and expected heterozygosity (He) under Hardy–Weinberg equilibrium were respectively 0.247 and 0.316. A substantial level of inbreeding within populations induced by Wahlund effect, was observed (F_IS = 0.231). High diversity resulted from the great number of genotypes in the ancestral population before fragmentation, favoured by the outbreeding of the species. High differentiation and low gene flow were detected among populations (F_ST = 0.200) and among pairs of ecological zones (0.113< F_ST <0.198). However, the differentiation coefficient of the four zones was low (F_ST = 0.085) and similar to the average F_ST for forest trees. Population structuration depends on geographic distance between sites rather than bioclimate, indicating that structuration has occurred slowly and that climatic conditions have had little effect. Nei's genetic distances (D) between populations were low and ranged from 0.004 to 0.201. Mean D value for all population was 0.087. The UPGMA clustering established for all populations through Nei's genetic distances did not clearly show that, for the majority of populations, grouping had resulted from ecological factors or geographic location. The substantial differentiation and the high genetic similarities between populations indicate that populations have been recently isolated as a result of anthropic pressure. In-situ conservation strategies must first focus on populations with a high level of genetic diversity and rare alleles. Appropriate conservation action should take account of bioclimatic zones. Ex-situ preservation should be based on a maximum number of individuals collected within populations in each ecological group and their propagation in different bioclimates by means of cuttings.     

Genetic diversity and relatedness of boreal caribou populations in western Canada

We studied genetic diversity within and gene flow among six ‘threatened’ populations of boreal caribou (Rangifer tarandus caribou) inhabiting Alberta and British Columbia, Canada. Mean expected heterozygosity (H_E) across 11 loci spanned a narrow range between 0.74 and 0.79. Estimates of H_E were in the mid to high range of those typically observed in natural populations of large mammals, including caribou, and were not suggestive of any immediate threat to survival. We concluded that recent anthropogenic fragmentation of caribou range in western Canada has yet to affect genetic diversity of populations. Analysis of population structure identified a region of relatively low gene flow corresponding with the valley of the Peace River. The highest value of F_ST observed between populations on the same side of the river was 0.025, whereas the lowest value that spanned the river was 0.044. Confirming this result, an assignment test demonstrated that 96.5% of animals could be assigned to the correct side of the Peace River, though only 66.0% of animals could, on average, be assigned to populations of actual origin. Taken as a whole, our results support the existence of two discrete metapopulations bisected by the Peace River, within each of which there exist multiple populations, or at least multiple regions, which experience considerably higher levels of interchange. For caribou inhabiting the boreal plains, large rivers such as the Peace and Mackenzie may serve as biologically meaningful boundaries for managing metapopulations.     

Relationship between population size and genetic diversity in endangered populations of the European bullhead (Cottus gobio): implications for conservation

This study investigated the relationship between the current size of endangered bullhead (Cottus gobio) populations and microsatellite genetic variability. Additionally, the microsatellite data were used to evaluate whether a genetic test for population bottlenecks was able to provide evidence of recent severe population declines. Finally, our results were used to develop conservation priorities and measures. Population size appears to be a crucial parameter in determining the amount of genetic diversity that can be preserved in bullheads, since a significant positive correlation was observed between both variables. Furthermore, in some populations we were able to detect genetic signatures of the documented decline in population size. We suggest that the most immediate goal for bullhead conservation should be to increase the size and the range of the populations, and in doing so minimise or even reverse further genetic erosion. Potential management actions like habitat quality improvement, reduction of river fragmentation and supplementation programmes (translocation, supportive breeding) are discussed.     

Genetic Diversity and Structure of Wild Tunisian Myrtus communis L. (Myrtaceae) Populations

Myrtus communis L. (Myrtaceae), in Tunisia, is closely associated with Quercus suber L. forest which stretched continuously from the North to parts of Cap Bon and Tunisian Dorsal. The destruction of the primary oak forests associated to an over-exploitation of Myrtus for its essential oil quality had led to discontinuous populations exhibiting various levels of degradation. Using starch gel electrophoresis, we analyzed the polymorphism of nine isozymes in order to assess genetic diversity and structuring of 17 natural populations prospected in the three geographical regions and coinciding with subhumid, humid inferior and semi-arid superior bioclimates. The analysis of the level and the distribution of the genetic diversity in this species might help in its conservation. Out of the 18 loci detected for all populations and isozymes analyzed, 12 loci were polymorphic. Allelic frequencies differed according to populations and particular alleles characterized ecological groups. A high level of genetic variation within populations was observed. The mean number of alleles per polymorphic locus was Ap = 1.67, the percentage of polymorphic loci was P = 60.3% and the observed (Ho) and the expected (He) heterozygosities were respectively 0.144 and 0.215. Populations belonging to subhumid (Cap Bon) and semi-arid superior (Tunisian Dorsal) climates, located in degraded sites exhibited the highest level of inbreeding (0.425 < F_IS < 0.450). A high level of differentiation (F_ST = 0.396) and a low gene flow (Nm = 0.337) among populations, as a result of habitat intermediate destruction, were revealed. The differentiation of populations within the same bioclimate (or geographic) group was substantial and relatively higher for semi-arid superior populations (F_ST = 0.262), which were more distant. The three ecological groups exhibited a high level of structuring (F_ST = 0.401). These differentiations might be due to geographic distances and to the variations of ecological factors between sites, including human activities and environmental factors. Nei’s (1972) genetic distances calculated between pairs of populations were globally low (0.006 < D < 0.367) with a mean of 0.15. They indicate a high level of similarity between populations. UPGMA clustering, established through Nei’s genetic distances, showed three population aggregates according to their geographic/bioclimatic appartenances. The high differentiation between populations and the low level of their genetic divergence indicated their recent isolation under anthropic pressures. The species conservation (in situ or ex situ) strategies should take into account the genetic diversity level within populations and its variation between geographic groups.     

Population baseline data for monitoring genetic diversity loss for 2010: A case study for Brassica species in the UK

The Convention on Biological Diversity and the subsequent International Treaty on Plant Genetic Resources for Food and Agriculture have proved a watershed in plant genetic resources (PGR) conservation and consequent initiatives have set various PGR conservation targets to be met by 2010. The aim of our paper is to develop ideas and issues concerning the monitoring of natural genetic resources: particularly in terms of developing a baseline from which to measure levels of genetic diversity. Three species of Brassica (B. nigra, B. oleracea, B. rapa) found wild in the UK were assessed for levels of genetic diversity using AFLP. The relationship between genetic distribution and ecogeographic distribution was considered for each species to determine patterns that may be useful in formulating conservation strategies. Genetic distance between populations of B. nigra and B. rapa were correlated to geographic distance. Levels of genetic polymorphism in B. oleracea were correlated to soil pH while in B. rapa they were correlated to soil coarseness. In terms of PGR conservation these findings may suggest an emphasis toward in situ conservation of a selection of disparate populations would be appropriate where possible as such adaptations may be lost in ex situ collections.     

Restricted gene flow in the clonal hepatic Trichocolea tomentella in fragmented landscapes

We studied the genetic diversity, gene flow and population structure among 18 populations of the clonal bryophyte Trichocolea tomentella located in Finland, Lithuania, the UK and Canada using DNA fingerprinting methods. T. tomentella is a habitat-limited, unisexual hepatic, which occupies spring and mesic habitats in woodland. The relatively small populations are increasingly fragmented with a high risk for extinction for extrinsic reasons. The presence of relatively high levels of genetic diversity regardless of population size highlights the role of even small remnant populations as important sources of genetic diversity in T. tomentella. The long-term accumulation of genotypes and somatic mutations may explain the observed levels of diversity. Gene flow among populations seems to be infrequent indicating dispersal limitation also on the relatively small spatial scale. Colonization within populations is not affected by isolation by distance suggesting the occurrence of random short-range dispersal of detached vegetative fragments. The population structure study confirmed the low mortality rates of shoots indicating a long life span of the clones in favourable conditions. Efficient ramet production by branching is likely to operate against interspecific competition. To conclude, T. tomentella appears to persist well in undisturbed habitats due to clonal regeneration, although restricted dispersal capacity is likely to prevent successful (re-)colonization in the potential habitat patches of recovering forest landscapes. The implications of the results for conservation are introduced.