Population structure and movements of a threatened butterfly (Lopinga achine) in a fragmented landscape in Sweden

The red-listed butterfly Lopinga achine was studied by mark-recapture methods in southern Sweden for three seasons. We examined movement within and between populations and egg production in relation to age. The majority of the movements were small with mean movements between recaptures of 45-54 m for males and 94-116 m for females. There were few movements between sites, 20 of 996 recaptured males moved and 36 of 391 recaptured females, even though the distance to other sites was in many cases <100 m. The distance moved and the number of females moving between sites increased with increasing age. On average, a female that moves does so after laying two-thirds of its eggs in its natal site. It is therefore important to take account of the proportion of reproductive effort involved in dispersal when estimating colonisation ability. The males did not move more with increasing age. Female behaviour can be seen as a “spread-the-risk” strategy, an adaptation to the successional habitat of L. achine, whose natal site sooner or later will deteriorate. Butterflies like L. achine living in successional habitats may exhibit mobility that is intermediate between butterflies living in ephemeral habitats (very mobile) and in long-lived habitats (sedentary).     

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.     

Neutral DNA markers fail to detect genetic divergence in an ecologically important trait

The development of strategies for in situ, ex situ conservation and reforestation of the monkey puzzle tree (Araucaria araucana), a vulnerable tree endemic to southern South America, has led to an interest in the level and distribution of the genetic diversity of the species. Neutral DNA markers (RAPDs) and quantitative genetic techniques were used to characterise genetic heterogeneity within and among populations from throughout the natural range of the species. Both the level and pattern of genetic variation estimated using the different techniques were essentially uncorrelated. An important discrepancy was found with the neutral markers failing to detect an important quantitative genetic divergence across the Andean Range relating to drought tolerance. This study clearly demonstrates the potential problems associated with making recommendations for conserving the genetic resource of threatened species based solely on neutral marker studies. Alternative approaches are discussed, including a stronger focus on ecologically important traits and the potential use of surrogate measures of genetic variability.     

Artificial dispersal of endangered epiphytic lichens: a tool for conservation in boreal forest landscapes

Due to increasing fragmentation of the boreal forests of Fennoscandia, a number of epiphytic lichens are now becoming threatened. Since these species typically are limited by a poor ability of dispersal, one possible but largely unexplored strategy for conservation is to disperse lichen material artificially into suitable habitats. Therefore, the objective of this study was to evaluate survival and vitality in lichen fragments from Evernia divaricata (L.) Ach. and Ramalina dilacerata (Hoffm.) Hoffm. after transplantation into three different stand types situated in northern Sweden, using different fragment sizes and modes of transplantation. After one year, survival ranged between 85% and 97.5%, and new growth occurred at all transplantation sites. The study has shown that transplantations of small fragments might constitute a resource-efficient option for establishment of new populations of endangered lichen species, or for enlarging their populations at the present sites of occurrence. In addition, the mode of transplantation was of importance for fragment vitality.     

Habitat associations of the endangered damselfly, Coenagrion mercuriale, in a water meadow ditch system in southern England

The damselfly Coenagrion mercuriale (Charpentier) (Odonata: Coenagrionidae) is threatened throughout Europe. It is listed on the EC Habitats and Species Directive and is the only odonate currently given priority status in the UK Biodiversity Action Plan. Information about the habitat preferences of this species is required to guide conservation and management programmes. We obtained estimates of density of mature adult C. mercuriale during an intensive mark-release-recapture study over 7.65 km of a water meadow ditch network in the Itchen Valley, Hampshire. Detailed habitat information was also collected, including a variety of physical variables, and data about the in-channel and bankside vegetation. C. mercuriale density and movement were analysed in relation to habitat variables and local population size using Generalized Linear Models. Mean adjacent population density was the single most important factor determining density. However the species was also associated with a number of habitat features, the most important of which were: a channel substrate consisting primarily of silt, wide underwater ledges (berms), in-channel emergent dicots, and bankside monocots. The presence of trees was negatively associated with damselfly density. Mean net lifetime movement was greatest from sections with low density, with smaller than average berms, and with deeper water. The causes and consequences of these findings are discussed in relation to the conservation and management of this rare species.     

Development of a dragonfly awareness trail in an African botanical garden

The IUCN/SSC Status Survey and Conservation Action Plan: Dragonflies calls for an increase in educating the public and increasing awareness of dragonflies (Odonata). Dragonflies are conspicuous and attractive, and can ‘stand in’ for other invertebrates in raising awareness of the necessity to conserve invertebrates. While reserves have been set aside for dragonfly awareness and conservation in the northern hemisphere, no such dragonfly reserves appear to exist in the southern hemisphere, despite its rich dragonfly fauna. This paper describes the development of a dragonfly-awareness trail in an established and well-visited botanical garden (The National Botanical Gardens, Pietermaritzburg, South Africa). Correlations between species and environmental variables were significantly high for six measured environmental variables: pH, percentage shade, vegetation (structural and compositional), ambient and water temperature, and water depth. Multivariate analyses of data, classified 20 a priori selected sampling units into four ecologically meaningful biotope types, each with a characteristic dragonfly assemblage. The four biotopes provided potential viewing points. To these were added a further three duplicate biotopes so as to link the trail into a circuit. Questionnaires assessed public awareness of dragonflies, and helped design a preliminary leaflet. A final trail design was drawn up along with an expanded booklet. The trail has now been implemented. Recommendations are made on the minimal scientific underpinning required for future trail design.     

Genetic isolation and morphological divergence of Black Sea bottlenose dolphins

The Black Sea is a semi-enclosed body of water that differs from the adjacent Mediterranean Sea in terms of its biodiversity, oceanographical and ecological characteristics. There is growing international concern about pollution in the Black Sea and other anthropogenic threats to its fauna. The bottlenose dolphin (Tursiops truncatus) is one of three species of cetaceans living in the Azov-Black Sea basin. Despite considerable research on bottlenose dolphins elsewhere, the extent of human impacts on the Black Sea populations is unknown. Previous attempts to award special conservation status to Black Sea cetaceans have failed specifically because policy makers have viewed their ecological and evolutionary uniqueness as equivocal. This study assessed divergence between Black Sea, Mediterranean Sea and Atlantic Ocean bottlenose dolphins for 26 cranial measurements (n = 75 adult bottlenose dolphin skulls) and mitochondrial DNA (n = 99 individuals). Black Sea bottlenose dolphins are smaller than those in the Mediterranean, and possess a uniquely shaped skull. As in a previous study, we found the Black Sea population to be genetically distinct, with relatively low levels of mtDNA diversity. Population genetic models suggest that Black Sea bottlenose dolphins have so little gene flow with the Mediterranean due to historical isolation that they should be managed separately.     

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.     

Two genetically distinct units of the Chinese sika deer (cervus nippon): analyses of mitochondrial DNA variation

Chinese sika deer, Cervus nippon, are currently threatened by habitat loss, fragmentation and human hunting, which has led to the extinction of three subspecies in the wild. The remaining subspecies subsist in the narrow regions of Jilin, Heilongjiang, Sichuan, Gansu, Jiangxi, Anhui, and Zhejiang provinces. In order to design effective conservation strategies for the Chinese sika deer, we have investigated genetic diversity, population structure and gene flow in the Chinese sika deer populations by analyzing ≈995 base pairs of the mitochondrial DNA (mtDNA) control region in 59 individuals sampled from the northeast of China, Sichuan, Jiangxi and Zhejiang. Chinese sika deer exhibited low mtDNA diversity and high gene flow among the four populations, and showed no strong geographical structure. The analyses of mtDNA variation among individuals of sika deer identified only two main phylogenetic groups even though three subspecies were sampled. These data singles out the Zhejiang population as being highly genetically distinct and worthy of separate conservation consideration. Therefore, it is recommended that a breeding program for the Zhejiang population be established.     

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.     

Geographic isolation, genetic divergence, and ecological non-exchangeability define ESUs in a threatened sky-island rattlesnake

We utilized nine microsatellite loci and 865 bases from two mtDNA genes to estimate demographic parameters and visualize historic/contemporary connectivity among populations of a sky-island rattlesnake (New Mexico ridge-nosed rattlesnake, Crotalus willardi obscurus). This taxon is listed as threatened under the United States Endangered Species Act (ESA) and is distributed patchily within three borderland mountain ranges [Animas (ANM), Peloncillo (PEL), Sierra San Luis (SSL)] of southeastern Arizona, southwestern New Mexico and north-central México. Molecular data support a hypothesis of northward range expansion from México, with subsequent isolation on sky-islands through vicariant desertification that transformed intervening wooded valleys and low elevation passes into inhospitable habitat. Historic and recent movements have been within rather than among mountains. Although populations are linked via ancestral polymorphism, they do not represent a single mtDNA gene pool. All are genetically bottlenecked, with the Peloncillo reflecting deepest/sharpest declines and fewest captures per unit effort. Most recent population declines occurred ∼4 kybp (thousands of years before present). Thus, population reductions are historic and environmental rather than contemporary and anthropogenic. Our data demonstrate that PEL is ecologically non-exchangable with other sky-island populations, and thus comprises one ‘evolutionary significant unit’ (ESU), while SSL and ANM comprise ‘management units’ (MUs) within a second ESU. All three meet legal criteria for recognition as ‘distinct population segments’ (DPS) under the ESA.     

Molecular markers and a sequence deletion in intron 2 of the putative partial homologue of LEAFY reveal geographical structure to genetic diversity in the acutely threatened legume genus Clianthus

Clianthus is an acutely threatened, bird-pollinated genus endemic to New Zealand, represented in the wild by only one population of C. puniceus and 11 populations of C. maximus, each with very few individuals (typically <10 per population). A limited number of named Clianthus cultivars of indeterminate origin are commonly grown as ornamentals. Genomic DNA from individual Clianthus plants was extracted for genetic diversity analysis using a range of molecular markers, including amplified fragment length polymorphism (AFLP). Data were analysed by the unweighted pair-group method with arithmetic averaging (UPGMA), the generation of Neighbor-Joining trees, and analyses of molecular variance (AMOVA). Genetic distance between wild populations of C. maximus was highly correlated with geographical distance between populations. Sequencing of intron 2 of a putative partial homologue of the floral meristem identity gene LEAFY (CmLFY) revealed a 7 bp deletion that was exhibited homozygously in the more northern populations of C. maximus, and in all individuals tested from the sole population of C. puniceus. This deletion was not exhibited in more southern populations of C. maximus. Further, one geographically intermediate population contained some plants that were heterozygous for the deletion. Parallel analyses of cultivated Clianthus genotypes, more than half of which were also homozygous for the 7 bp deletion, showed that these were not representative of the broad, but threatened, diversity remaining in the wild. It is argued that wild populations of C. maximus are unlikely to have arisen from the escape of plants from cultivation. Conservation effort should focus on the protection and study of the extant plants in these wild populations, rather than on the introduction of disturbance regimes to uncover potential seed banks.     

Anthropogenic and natural disturbance lead to differing patterns of gene flow in the Rocky Mountain tailed frog, Ascaphus montanus

Understanding the influence of habitat alteration on population structure and persistence is critical for effective conservation strategies. Timber harvest and wildfire are two of the most prevalent disturbances across temperate forests, yet the long-term effects of these two forces on population connectivity have rarely been studied. We studied populations of the Rocky Mountain tailed frog (Ascaphus montanus) across landscapes that have experienced either timber harvest or broad scale fires. Rocky Mountain tailed frogs generally require forested habitat, and are a species of concern in managed forests. We used landscape genetic techniques to test the explanatory power of alternative paths of connectivity across both burned and harvested forests and identified topographical and climatic variables that significantly influenced gene flow. We found that timber harvest and fire led to different patterns of genetic connectivity. Widespread terrestrial gene flow was maintained across previously burned areas and connectivity was primarily limited by solar radiation. In contrast, gene flow among populations in managed forests followed riparian corridors, presumably to avoid the loss of cover due to timber harvest. Gene flow along riparian corridors in Rocky Mountain tailed frogs differs from a population of the closely related coastal tailed frog (Ascaphus truei), for which gene flow appears to occur primarily over land. These results suggest management of Rocky Mountain tailed frogs should focus on maintaining riparian buffer zones, which may be less effective for the coastal species. Thus, consideration of landscape differences is essential even for conservation of closely related, morphologically similar species.     

Florida panther dispersal and conservation

We studied dispersal in 27 radio-collared Florida panthers Puma concolor coryi in southern Florida from 1986 to 2000. Male dispersal was longer (mean=68.4 km) than that of females (mean=20.3 km), tended to be circular, frustrated, and of insufficient length to ameliorate inbreeding. Females were philopatric and established home ranges that were less than one home range width away from their natal ranges. All females were successful in establishing territories, whereas males were successful 63% of the time. Dispersing panthers avoided moving toward the southeast and into an area of limited forest cover. Independence and the initiation of dispersal occurred at about 14 months of age and lasted for an average of 7.0-9.6 months for females and males, respectively. On average, Florida panthers disperse shorter distances than are typical for western populations of Puma concolor. A recent increase in long distance male dispersal events may be related to an increase in reproduction and population density resulting from the introduction of female cougars P. c. stanleyana into south Florida. Although the population exhibits the behavioral ability to colonize nearby vacant range, females have yet to do so. Successful dispersal to these areas could be facilitated by habitat restoration and translocation of females.     

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.     

Nectar plant sources for the threatened Apollo butterfly (Parnassius apollo L. 1758) in populations of central Spain

The use of flowers by the apollo butterfly at several sites along the Sierra de Guadarrama (central Spain) is documented. The Apollo butterfly uses a narrow range of flowers as nectar sources. Males feed more frequently on Armeria arenaria, Jasione montana and Carduus carpetanus. Females feed on the same species as males, but incorporate other species in their diet (Jurinea humilis, Thymus bracteata and Anthyllis lotoides). These observations also indicate that Parnassius apollo generally direct their foraging activities towards a limited number of the available plants but that the plants used can differ slightly between sites and change within sites through time. These data on resources for the adult butterflies should serve for the management of the species in future conservation programmes.     

Restoration of threatened red gorgonian populations: An experimental and modelling approach

The increasing disturbances affecting marine communities highlight the need to examine restoration measures that can be added to other conservation efforts for threatened populations. The main goal of this study was to examine the usefulness of ecological restoration in the management of gorgonian populations damaged by diving activity in intensively visited marine protected areas (MPAs). We used field experiments as well as simulations from size-structured matrices to assess the utility of transplantation of living fragments from damaged colonies to increase the viability of threatened populations. Despite results showing that technique failure caused the loss of 40% of transplants, well-attached transplants achieved survival rates (80%) similar to those of natural colonies. Surprisingly, environmental conditions (light level and presence of algae) did not have a significant effect on the mortality of the transplants, but did affect methodological failure rates (37% of transplants were lost in the photophilous treatments in contrast to the 25% lost in sciaphilous treatment). The simulations showed a substantial increase in the annual population growth rate (λ) only when transplantation was performed every two years and under the most demanding conditions (recovering 75% of the detached colonies and obtaining 3 fragments from each one). Predictions from the size-structured matrix model suggest severe limitations of this technique at larger spatial scales. However, our study confirms the feasibility of this restoration measure to contribute to the recovery of populations in MPAs affected by local disturbances. The experimental and modelling approaches developed here may provide useful guidelines for future studies on the restoration of marine populations.     

Ecological effects on estimates of effective population size in an annual plant

Effective population size (N_e) is a critical indicator of the vulnerability of a population to allele loss via genetic drift, and it can also be used to assess the evolutionary potential of a population. While some plant conservation plans have focused on outcrossing through cross-pollination as a way to increase estimated N_e, variance in reproductive output determined by ecological factors such as competition can also strongly affect estimated N_e. We examined the effects of intraspecific and interspecific competition, stressful soils, and local adaptation on estimates of N_e in an annual plant species. While ecological influences on plant growth rate variance have been predicted to influence estimates of N_e/N, we found a significant effect on the estimate of N_e/N, but no significant ecological effects on growth rate variance. Lower survivorship on stressful soil was the most important effect reducing estimates of N_e/N. If stochastic mortality is greater in environments that are abiotically stressful, then populations in these stressful environments may be slower to adapt because of lower census sizes and reduction of N_e/N. In populations of conservation concern, increasing survivorship may be of greater benefit for maximizing N_e than the reduction of variability in reproductive output among surviving adults.     

Cryptic genetic bottlenecks during restoration of an endangered tropical conifer

Forest restoration programmes aim to use material for re-planting that is genetically diverse and not inbred. However, restricted seed sampling, high variance in reproductive output, and the production of inbred seeds that survive in the nursery but not in the wild can lead to forest restoration stock being genetically compromised. The aim of this study was to evaluate whether the reproductive biology of the New Caledonian endemic conifer Araucaria nemorosa makes it susceptible to these genetic problems and to assess whether there is evidence for genetic bottlenecks and elevated inbreeding in nursery stock compared to seedlings and adults from wild source populations. Reproductive output was low with high variance among trees (only 14% of adult trees surveyed produced mature cones, >50% of examined cones had <10 viable seeds). Evidence for an extreme genetic bottleneck was detected in a nursery population established from cones collected from adult trees. A second nursery population established with seed collected from the forest floor showed no evidence of a genetic bottleneck, but was inbred compared to its wild source population. In light of these results, we do not recommend collecting cones directly from A. nemorosa as an efficient means of establishing genetically diverse stock for restoration programmes. Collecting seed from the forest floor is likely to be more effective, but the planting stock may contain a high proportion of inbred individuals. Collecting established wild seedlings already subjected to natural selection is suggested as an alternative method of maximising the diversity captured, whilst minimising sampling effort and proportion of inbred individuals.     

Forest fragmentation differentially affects seed dispersal of large and small-seeded tropical trees

The responses of plant-animal interactions to forest fragmentation can vary. We hypothesized that large-seeded plant species would be more susceptible to forest fragmentation than small-seeded species because large-seeded species rely on a few, extinction prone dispersers. We compared seed dispersal of the large-seeded, mammal dispersed Duckeodendron cestroides and the small-seeded, avian dispersed Bocageopsis multiflora. The number, percentage, distance, and distributions of dispersed seeds were all reduced in fragments for Duckeodendron but not for Bocageopsis. Other fragmentation research in tropical communities supports this hypothesis through three lines of evidence: (1) Large-seeded plant species are more prone to extinction, (2) Fragmentation restricts or alters the movement of large animal dispersers more than small dispersers, and (3) Large and small-seeded species seem to be differentially linked to primary and secondary forest habitats. Therefore, small-seeded plants may be more resilient to forest fragmentation while large-seeded species may be more susceptible and should be a priority for conservation.