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.     

Ecological and genetic measurements of dispersal in a threatened dragonfly

Leucorrhinia caudalis is a rare dragonfly, threatened throughout its European distribution. The species was formerly widespread in the Swiss lowlands, but only a single population remained in the 1980s. However, a spread has recently been observed, with additional ponds being colonised, sometimes at considerable distance. Despite this evidence of recent long-distance dispersal, it is unknown whether L. caudalis regularly moves among ponds or whether this is a rather rare event. A combination of an ecological mark-resight and a population genetic study was applied to investigate contemporary dispersal and the genetic footprint of the recent population history of L. caudalis in Switzerland. DNA for genetic microsatellite analysis was extracted from exuviae. The mark-resight study and the genetic analysis gave congruent results. They showed that L. caudalis is mostly a sedentary species, with only a few contemporary dispersal events over distances up to 5 km being observed. The genetic analysis was in agreement with the recent population history of the Swiss populations. The oldest and largest population showed large genetic diversity and acted as source population for the recent spread of L. caudalis in Switzerland. Recurrent gene flow among this source population and close populations caused substantial local genetic variation in the latter, as well as low population differentiation. The two recently founded distant populations (?30 km distance) were genetically less diverse and highly differentiated. These distant populations and another recently colonised population also expressed signatures of genetic bottlenecks.     

The western jewel butterfly (Hypochrysops halyaetus): factors affecting adult butterfly distribution within native Banksia bushland in an urban setting

Adult western jewel butterflies Hypochrysops halyaetus Hewitson were studied in the Koondoola Regional Bushland Reserve on the northern outskirts of the Perth metropolitan area, Australia, in 1999. The butterfly is myrmecophilous, and the southern (Perth) form is considered to be ‘Vulnerable’. Butterfly dispersal, distribution, population size and habitat preferences were estimated using mark-recapture techniques. Butterflies were capable of moving large distances, and were widespread within the reserve but with a low density; however, one small physical area (80 × 20 m) contained a colony with large numbers of individuals. This colony, and an adjacent area of bush, was subject to a detailed microdistribution study: 1158 butterflies were marked within it over a 25-day period. The small size of the microdistribution study area compromised statistical independence assumptions: spatial autocorrelation was tested for and corrected, where appropriate, using a Markov-chain Monte-Carlo approach. Sex differences were evident in spatial autocorrelation: female data were not spatially autocorrelated, whilst male data was. The latter probably reflects the impact of the perching mode of mate location employed by males. Butterflies appeared to prefer rather degraded bush typical of a post fire/disturbance regime with high densities of the host plant Jacksonia sternbergiana. Males were positively correlated with the proportion of bare ground, again probably relating to preferred perching sites for mate location. Females were negatively correlated with Conospremum stoechadis probably indicating dense ground cover in degraded areas and Xanthorrhoea preissii probably an indicator at the site of more mature vegetation. Reserve management to improve vegetation quality may threaten the persistence of the butterfly within the reserve, whilst controlled burning to create even quite small new areas of habitat in an urban situation may be difficult to achieve. The creation of suitable habitat may be possible using mechanical means, provided the action of fire is not an essential component of habitat creation. As the western jewel appears to prefer degraded vegetation, it is possible that there may be opportunities for this species’ expansion within the urban environment provided that its mutualistic ant partner is also present.     

Persistence and conservation of a consumer-resource metapopulation with local overexploitation of resources

The monophagous flightless weevil Hadramphus spinipennis causes the frequent local extinction of its host plant Aciphylla dieffenbachii through overexploitation. Both species are endemic and restricted to the Chatham Islands group (New Zealand). The weevil is classified as endangered and the plant as a threatened species. As this herbivore-plant system is locally unstable long-term persistence only appears possible via a metapopulation structure in which subpopulations are connected by dispersal. This paper investigates this hypothesis using a spatially explicit model of a consumer population whose resource is patchily distributed. The parameters are adapted to the H. spinipennis-A. dieffenbachii system. Our model includes local population dynamics and dispersal of the consumer, the destruction of host plant patches due to foraging and their regeneration. The results show that the key factor for long-term persistence is the short-range dispersal of the consumer with high mortality during dispersal. Only this highly inefficient dispersal prevents the synchronisation of local dynamics while ensuring sufficient colonisation of regenerated patches. We also show that although synchronisation through spatially correlated environmental fluctuations may be critical for long-term persistence, it cannot replace the strong effects of dispersal. Thus, in a consumer-resource metapopulation with deterministic local extinction, the strength and spatial scale of consumer dispersal in relation to the spatial and temporal scales of the resource patch dynamics (patch destruction and regeneration, number, size and distance of patches) play a vital role for long-term persistence. The consequence for conservation management is that such metapopulations could in fact profit rather than suffer from decreasing connectivity of resource patches.     

Predicting short-term extinction risk for the declining Little Bustard (Tetrax tetrax) in intensive agricultural habitats

Populations of Little Bustard (Tetrax tetrax) have shown pronounced declines in European farmland landscapes over the last 25 years due to the intensification of agricultural practices. In France, the number of breeding males in agricultural habitats has declined by 92% since 1980 as a result of decreases in insect abundance and nest destruction during harvesting. We formulate an age- and sex-structured stochastic model for the remaining Little Bustard population in SW France that has been studied since 1997 and, using actual values of demographic rates, we estimate its extinction risk over a time period of 30 years to be of 0.45. At the level of local populations, the extinction risk ranged between 0.66 and 0.90, largely depending on the initial population size and local fecundity of each population. A comprehensive sensitivity analysis evaluated the influence of uncertain demographic rates and of aspects of the spatial dynamics (sex and age dispersing individuals, sex-biased dispersal and different dispersal rates) on the predicted extinction risks and showed that our model was robust to changes of a wide range of combinations assessed. Given the severity of the current decline, and the spatial issues raised by our analysis, implications of our findings for the conservation of this endangered species are suggested.     

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.     

Wing loading and habitat selection in forest beetles: Are red-listed species poorer dispersers or more habitat-specific than common congenerics?

Traits such as poor dispersal ability and high habitat specificity are thought to predispose some species to a greater risk of extinction than others. Habitat preferences and morphological features associated with dispersal ability were compared between red-listed species and common congenerics co-occurring in boreal forests in northern Sweden. Measurements of body size, wing loading and wing aspect ratio were used to compare dispersal abilities, while catches from experimental treatments of dead wood were used to compare habitat preferences. We also compared how restricted red-listed species were to particular sites or habitats relative to common species. The red-listed Epuraea longipennis was longer and wider, while Epuraea. deubeli weighed less than common Epuraea species. In contrast to expectations, these red-listed species had a larger wing area (relative to their body mass) and thus a lower wing-loading than congeneric species, suggesting superior dispersal abilities. The red-listed Tachinus elegans possessed intermediate morphological characteristics, compared with common congenerics. However, the relationships between the risk of extinction in fragmented habitats and size and dispersal ability are likely to be scale-dependent, so intermediate or superior dispersal abilities may increase extinction risk at some scales. Red-listed species were not found in fewer sites or habitat types than congenerics so were not more likely to be habitat specialists. However, some red-listed species preferred deeply shaded and Fomitopsis pinicola-inoculated logs, relative to congenerics, suggesting that specificity to these particular microhabitats may be connected with extinction risk.     

Dynamics of threatened goldenseal populations and implications for recovery

Goldenseal (Hydrastis canadensis) is a North American perennial clonal herb highly prized for its medicinal value. It is threatened at the northern range limit with only 20 populations known in Canada. To assist recovery planning, 13 populations were sampled to model dynamics. The fate of all ramets in one square-meter was monitored from 1998 to 2001. Transition matrices were built for 2000-2001, using three stage classes based on size and reproductive status. A six-stage pooled matrix, separating established ramets from newly produced ramets, was also constructed. Recruitment by seed was not observed and therefore excluded. The average population growth rate (λ) was 1.062 ± 0.053, which did not significantly differ from the equilibrium value (1.0) suggesting that the northern population is stationary. However, growth rates among population samples varied largely and had wide confidence intervals. Populations with λ-values less than or close to 1.0 require environmental change to increase. Recovery of goldenseal, and possibly other woodland perennials at risk, requires intervention aimed at population size augmentation, habitat optimization, and targeted dispersal.     

Does reproductive plasticity in Lumbricus rubellus improve the recovery of populations in frequently inundated river floodplains?

Flooding events often eradicate all of the individuals of the earthworm species Lumbricus rubellus living in river floodplains, although earthworm cocoons usually survive immersion, permitting populations to recover after the flood waters recede. Yet, if the area is flooded again before earthworms hatching from cocoons or migrating from adjacent areas reach reproductive maturity, it is unlikely that their populations will recover. The objective of this study is to determine the importance of the length of the dry period for population recovery in L. rubellus. Earthworms were collected at three floodplain sites along the Rhine River that were frequently, moderately or seldom flooded. Reproductively mature L. rubellus from the frequent flooded site were half the weight and probably younger than those from the other sites. A mechanistic population model was used to estimate the time for earthworm development from hatching to reproductive maturity, and to calculate the probability of population recovery after flooding. The model results show that the probability of extinction increases when the dry period is not long enough for individuals to reach reproductive maturity. When this condition is met population extinction is virtually absent resulting from the high lifetime reproductive output of L. rubellus. Parameterization of the model with site-specific data indicate that population survival on the site with the shortest dry period drastically decreases if worms mature at the weight measured at the other sites. The results therefore strongly suggest that the dry period is critical for population recovery in river floodplains and that earthworm populations have adapted to local (site-specific) conditions.     

Herbicides, weeds and endangered species: management of bitou bush (Chrysanthemoides monilifera ssp. rotundata) with glyphosate and impacts on the endangered shrub, Pimelea spicata

Environmental weed invasion threatens the biodiversity of native species. Unfortunately, managing these weeds may also affect biodiversity adversely. A recent example occurred when glyphosate, a herbicide used to control the highly invasive weed, bitou bush (Chrysanthemoides monilifera ssp. rotundata), accidentally drifted over a small population of an endangered shrub, Pimelea spicata. Following concerns that the affected population would not recover and, thereby, cause the local extinction of P. spicata, we conducted a series of glasshouse and field experiments to explore the impacts of glyphosate on this endangered species. Seedlings and young plants of P. spicata, in which the tap root was undeveloped, were killed by a single application of glyphosate. Older plants with a well developed tap root also died back initially, but about 50% of individuals re-sprouted. This re-growth was associated with a significant decrease in tap root diameter, implying that further disturbance, including repeated treatment with glyphosate, would kill plants by impairing their potential for recovery. Unlike some sclerophyllous native shrubs, the tolerance of P. spicata to glyphosate was limited, even when its growth was slowed artificially by limiting water availability. Winter applications of glyphosate to manage infestations of bitou bush will impact adversely on populations of P. spicata and may also affect the other rare and endangered species whose survival is threatened by this species, even though some natives are unaffected by the herbicide. Protecting native biodiversity from bitou bush will involve sustainable weed management that minimises impacts on non-target native species.     

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.     

Life on the edge - High levels of genetic diversity in a cliff population of Bertya ingramii are attributed to B. rosmarinifolia (Euphorbiaceae)

Wind-facilitated migration of new genotypes into small, geographically disjunct populations should buffer them against local extinction. Bertya ingramii, a monoecious, wind-pollinated shrub, is restricted to three populations in a 4 km² area in eastern Australia. Populations are separated by deeply dissected gorges where it is unlikely that seeds are exchanged but where wind movement may facilitate pollen dispersal. Using 156 highly polymorphic ISSR markers, we found moderate genetic variation within and among populations of B. ingramii and less genetic diversity in a nearby and small population of the widespread Bertya rosmarinifolia. The smallest population of B. ingramii (<30 plants) had the highest genetic variation (65% polymorphic markers, Shannon Information Index = 0.30). AMOVA and a Bayesian analysis showed that molecular variance was equally distributed within and among populations suggesting that gene flow is as limited within as in among populations. Genetic distances between populations were only weakly explained by their relative geographic distances (mantel test, R² = 0.21, P = 0.001) but the distribution of private bands, the departure from Hardy-Weinberg equilibrium, and a UPGMA tree showed that the smallest population of B. ingramii was generally more similar to an upstream population of B. rosmarinifolia. Thirty-eight percent of bands in this small population of B. ingramii were exclusively shared with B. rosmarinifolia. This covert hybridisation may have been an ancient event but may be responsible for contemporary declines in germination and establishment in B. ingramii. The conservation implications are amplified by the endangered status of B. ingramii.     

Genetic structure of soil invertebrate populations: Collembolans,earthworms and isopods

Soil-living animals such as collembolans, earthworms and isopods are considered to be sedentary animals with only limited dispersal capacities. Therefore, gene flow among populations is expected to be low leading to significant population genetic differentiation due to random drift and local adaptation. We reviewed the literature to test this expectation. Our survey reveals a clear signature in the current pattern of genetic variation due to post-glacial colonization events. It also reveals that habitus can be a misleading predictor of dispersal capacity. In some species relatively high gene flow across considerable distances, most likely through passive dispersal (mediated by wind, water flow or animals), might counteract local genetic adaptation and will prevent loss of genetic variation.     

Dynamics of a black bear population within a desert metapopulation

Understanding metapopulation dynamics in large carnivores with naturally fragmented populations is difficult because of the large temporal and spatial context of such dynamics. We coupled a long-term database of visitor sighting records with an intensive 3-year telemetry study to describe population dynamics of recolonization by black bears (Ursus americanus) of Big Bend National Park in Texas during 1988-2002. This population, which occurs within a metapopulation in western Texas and northern Mexico, increased from a single pair of known breeding-age animals in 1988 to 29 bears (including 6 females of breeding age) in March 2000 (λ = 1.25/year). A migration and dispersal event in August-December 2000 reduced the population to 2 adult females and as few as 5-7 individuals. One-way movement distances from the study area during this event averaged 76 km for females (n = 7) and 92 km for males (n = 4), and 3 animals conducted migrations of at least 154, 178, and 214 km, respectively. Our observations exemplify the importance of stochastic events on demographics of small populations and highlight the potential scale of bear movement among montane islands of southwestern North America. They also provide insight into the use of dispersal data in parameterizing metapopulation models for large carnivores.     

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.     

Population viability analysis and fire return intervals for an endemic Florida scrub mint

We use population viability analysis of an endangered Florida scrub mint, Dicerandra frutescens, to specify the optimal fire return intervals for its long-term persistence and for its specific habitat. We derived 83 population projection matrices from 13 years of demographic data from eight populations, 59 matrices from scrub populations and 24 from firelane or yard edges. Seed dormancy and germination transitions were inferred based on experimental data and verified by comparing modeled vs. observed population trajectories. Finite rates of increase in scrub sites were highest shortly after fire and declined steeply through 10 years postfire. The break-even value of λ = 1 was passed quickly, in about six years, suggesting that populations >6 years postfire were already facing decline. The decline is probably related to the rapid growth of competing shrubs in the habitat of D. frutescens. In long-unburned sites, finite rates of increase were nearly always <1 and declined the most in the long-unburned site with no foot trails or treefall gaps. Finite rates of increase in firelane populations also declined with years since fire or last disking. The yard edge population showed λ values both >1 and <1, with no temporal trend. Stochastic simulations in scrub sites suggested an optimal regular fire return interval of about 6-12 years. Regular fires at this interval were more favorable than stochastic fire regimes, but stochasticity reduced extinction percentages at longer fire return intervals. Stochastic fire return intervals implied a wider optimal fire return interval of 6-21 years. We suggest that prescribed fire in Florida scrub on yellow sand has occurred (and needs to occur) more frequently than previously recommended.     

Recovery and maintenance of North Island kokako (Callaeas cinerea wilsoni) populations through pulsed pest control

Kokako (Callaeas cinerea wilsoni) population recovery on the North Island of New Zealand depends primarily on control of key introduced mammal pests, especially ship rats (Rattus rattus) and brushtail possums (Trichosurus vulpecula). Recovery can still occur if pest control is pulsed (x years ‘on’; y years ‘off’) because kokako sub-adults and adults are generally long-lived, although chick production is high only during ‘on’ years. Pulsing effort means that conservation resources can be extended to other sites or problems during ‘off’ years; that toxin input at any one site is reduced; and that project staff do not burn out by repeatedly working at a site. Mathematical modelling supports empirical evidence that pests need not be controlled every year in order to maintain or greatly increase kokako populations. It predicts that the total number of years during which there is pest control is the main factor determining population size. Three years of pest control in each 10 should be sufficient to at least maintain a population with 20 females when mean parameters apply, but pulsed control should still be effective with very pessimistic parameters. In the safest strategies, control should occur in minimum pulses of 2-3 years to avoid single poor years when few breeding attempts are made. Very small populations should first be increased to at least 20 females by translocation or continuous pest control. This will greatly reduce the probability of chance extinction, and increase the efficiency of subsequent pest control. The model will apply best to closed kokako populations below carrying capacity, in which pests are controlled over the entire block. Empirical data on the effects of habitat carrying capacity on kokako dispersal, and on the importance of stoats as predators of adult females are required to further strengthen the model.     

Minimum viable population and conservation status of the Atlantic Forest spiny rat Trinomys eliasi

A population viability analysis (PVA) was conducted to assess the minimum viable population (MVP) of the Atlantic Forest spiny rat Trinomys eliasi, a species threatened by habitat loss and restricted geographical distribution. Objectives were to suggest quasi-extinction thresholds, estimate minimum areas of suitable habitat (MASH) and MVPs, and compare results with the species’ current status. The computer package VORTEX was used. The model predicted sizes of 200 animals to achieve demographic stability, but buffering declines in genetic variability required populations of 2000 animals. Estimated MASHs were approximately 250 and 2500 ha for demographic and genetic stability, respectively. Mortality rate and mean litter size were the most sensitive parameters to changes in model assumptions. The protection of known populations and the search for extant populations are the first steps in conservation. T. eliasi's issue could help protecting the coastal shrubland ecosystem of Rio de Janeiro state. Observing IUCN's criteria for listing threatened species, it is suggested that T. eliasi should be ranked as vulnerable in red lists.     

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.