Modelling the effect of El Niño on the persistence of small populations: The Galápagos penguin as a case study

Small populations are vulnerable to long-term declines, even where short-term censuses indicate increasing trends in numbers. Census data for the Galápagos penguin (Spheniscus mendiculus) collected between 1970 and 2004 provide evidence that despite year-to-year population increases detected in most of the annual censuses, the strong El Niño events of 1982-83 and 1997-98 were followed by population declines of more than 60% from which the species has yet to recover. Such large declines raise concerns about the future viability of the species because the frequency and severity of El Niño events are predicted to increase. We used the simulation software VORTEX to evaluate the potential effects of El Niño on the risk of extinction of the Galápagos penguin population and its four constituent subpopulations. Weak and strong El Niño events were treated as catastrophes, with varying frequencies, which simulated past, current and future effects on the penguin population. The “Current El Niño” scenario, based on the frequency of El Niño events recorded in the Galápagos between 1965 and 2004, indicated an approximately 30% probability of extinction within the next 100 years for the penguin population. More ominously, the species may be at a greater risk if the frequency of strong El Niño episodes increases only marginally. A probability of extinction greater than 80% was predicted when the current frequency (5%) of strong El Niño events was doubled (to 10%). The probabilities of extinctions were higher for each subpopulation treated individually, ranging from 34% for Isabela and Fernandina, 64% for Bartolomé-Santiago to 78% for the smallest subpopulation on Floreana. Sensitivity analyses identified survival of penguins during El Niño events and sex ratio as influential parameters. The estimates of extinction risk may be conservative as other threats associated with increased human activities on the islands may further compromise species persistence.     

Biological effects of El Niño on the Galápagos penguin

Long-term monitoring of physical and biological parameters is essential for understanding the effects of El Niño on bird populations, particularly for small or declining populations. We examined the biological effects of El Niño activity from 1965 to 2004 using instrumental sea-surface temperatures from the Galápagos Islands and 20 years of census counts of the Galápagos penguin. Between 1965 and 2004, nine El Niño events were recorded of which two were strong and seven were weak. The two strong El Niño events of 1982-1983 and 1997-1998 were followed by crashes of 77% and 65% of the penguin population, respectively. The evidence suggests that the increased frequency of weak El Niño events limits population recovery. The 2004 penguin population is estimated to be at less than 50% of that prior to the strong 1982-1983 El Niño event. We discuss the biological effects of increased El Niño intensity and frequency within the context of a 6000-year record of El Niño influence and in the light of increasing anthropogenic threats operating after 1535, when the Archipelago was discovered by Europeans.     

Polygyny and female breeding failure reduce effective population size in the lekking Gunnison sage-grouse

Populations with small effective sizes are at risk for inbreeding depression and loss of adaptive potential. Variance in reproductive success is one of several factors reducing effective population size (N_e) below the actual population size (N). Here, we investigate the effects of polygynous (skewed) mating and variation in female breeding success on the effective size of a small population of the Gunnison sage-grouse (Centrocercus minimus), a ground nesting bird with a lek mating system. During a two-year field study, we recorded attendance of marked birds at leks, male mating success, the reproductive success of radio-tagged females, and annual survival. We developed simulations to estimate the distribution of male reproductive success. Using these data, we estimated population size () and effective population size N_e for the study population. We also simulated the effects of population size, skewed vs. random mating, and female breeding failure on N_e. In our study population, the standardized variance in seasonal reproductive success was almost as high in females as in males, primarily due to a high rate of nest failure (73%). Estimated N_e (42) was 19% of in our population, below the level at which inbreeding depression is observed in captive breeding studies. A high hatching failure rate (28%) was also consistent with ongoing inbreeding depression. In the simulations, N_e was reduced by skewed male mating success, especially at larger population sizes, and by female breeding failure. Extrapolation of our results suggests that six of the seven extant populations of this species may have effective sizes low enough to induce inbreeding depression and hence that translocations may be needed to supplement genetic diversity.     

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.     

Evaluating effects of deforestation, hunting, and El Niño events on a threatened lemur

Madagascar ranks as one of the world’s top extinction hotspots because of its high endemism and high rate of habitat degradation. Global climate phenomena such as El Niño Southern Oscillations may have confounding impacts on the island’s threatened biota but these effects are less well known. We performed a demographic study of Propithecus edwardsi, a lemur inhabiting the eastern rainforest of Madagascar, to evaluate the impact of deforestation, hunting, and El Niño on its population and to re-evaluate present endangerment categorization under the IUCN. Over 18 years of demographic data, including survival and fecundity rates were used to parameterize a stochastic population model structured with three stage classes (yearlings, juveniles, and adults). Results demonstrate that hunting and deforestation are the most significant threats to the population. Analysis of several plausible scenarios and combinations of threat revealed that a 50% population decline within three generations was very likely, supporting current IUCN classification. However, the analysis also suggested that changing global cycles may pose further threat. The average fecundity of lemurs was over 65% lower during El Niño years. While not as severe as deforestation or hunting, if El Niño events remain at the current high frequency there may be negative consequences for the population. We suggest that it is most critical for this species continued survival to create more protected areas, not only to thwart hunting and deforestation, but also to give this endangered lemur a better chance to recover from and adapt to altered climate cycles in the future.     

Reduced effective population size in an overexploited population of the Nile crocodile (Crocodylus niloticus)

Unchecked exploitation of wildlife resources is one of the major factors influencing species persistence throughout the world today. A significant consequence of exploitation is the increasing rate at which genetic diversity is lost as populations decline. Recent studies suggest that life history traits affecting population growth, particularly in long-lived species, may act to moderate the impact of population decline on genetic variation and lead to remnant populations that appear genetically diverse despite having passed through substantial demographic bottlenecks. In this study we show that the retention of genetic variation in a partially recovered population of Nile crocodile is deceptive, as it masks the reality of a significant decline in the population’s effective size (N_e). Repeated episodes of unchecked hunting in the mid to late 20th century have today led to a five-fold decrease in the population’s N_e. Using current census data we estimate the contemporary N_e/N ratio as 0.05 and, in light of quotas that permit the ongoing removal of adults, simulated the likely effects of genetic drift on extant levels of variation. Results indicate that even if the current effective size is maintained, both allelic diversity and heterozygosity will decline. Our findings have complex implications for long-lived species; an emphasis on the retention of genetic variation alone, whilst disregarding the effects of population decline on effective size, may ultimately obscure the continued decline and extinction of exploited populations.     

Conserving genomic variability in large mammals: Effect of population fluctuations and variance in male reproductive success on variability in Yellowstone bison

Loss of genetic variation through genetic drift can reduce population viability. However, relatively little is known about loss of variation caused by the combination of fluctuating population size and variance in reproductive success in age structured populations. We built an individual-based computer simulation model to examine how actual culling and hunting strategies influence the effective population size (N_e) and allelic diversity in Yellowstone bison over 200 years (～28 generations). The N_e for simulated populations ranged from 746 in stable populations of size 2000 up to 1165 in fluctuating populations whose census size fluctuates between 3000 and 3500 individuals. Simulations suggested that ～93% of allelic diversity, for loci with five alleles will be maintained over 200 years if the population census size remains well above ～2000 bison (and if variance in male reproductive success is high). However for loci with 20 alleles, only 83% of allelic diversity will be maintained over 200 years. Removal of only juveniles (calves and yearlings) resulted in longer generation intervals which led to higher maintenance of allelic diversity (96%) after 200 years compared to the culling of adults (94%) when the mean census size was 3250 (for loci with five alleles). These simulations suggest that fluctuations in population census size do not necessarily accelerate the loss of genetic variation, at least for the relatively large census size and growing populations such as in Yellowstone bison. They also suggest that the conservation of high allelic diversity (>95%) at loci with many alleles (e.g., ?5) will require maintenance of a populations size greater than approximately 3250 and removal of mainly or only juveniles.     

Effects of the 1982–1983 El Nino on Humboldt penguin colonies in Peru

The Humboldt penguin Spheniscus humboldti is endemic to the Peruvian Current which flows northward along the coast of Chile and Peru. This species has greatly diminished from its former abundance.The coast of Peru is characterised by high biological productivity which concentrates fish such as the anchovy Engraulis ringens, the main prey item of marine predators including seabirds. In years of the abnormal oceanographic conditions of El Nino, the schools of anchovies become unavailable to the seabirds and they disperse in search of food. Massive mortality, especially of juveniles, results and there is nest desertation and lack of reproduction.This paper describes the effects of the 1982–1983 El Nino on Humboldt penguin colonies in Peru. There has been an overall population decline of 65% and the surviving population in 1984 was estimated to be between 2100 and 3000 adults. Although El Nino is a periodic event and the Humboldt penguin has evolved to adapt to such unpredictable changes, the environment has now been altered by man. Under these circumstances, the 1982–1983 El Nino has contributed to placing this species in a critical position.     

Environmental uncertainty and commercial fishing: Effects on Peruvian guano birds

The roles of the El Niño phenomenon, a cessation of upwelling, and of human fishing in limiting populations of Peruvian guano birds (Guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata, and Peruvian brown pelican Pelecanus occidentalis thagus) were investigated using historical records and the scientific literature. In normal years, the combined populations of the three species showed an 18% increase per year. El Niños occurred at approximately five-year intervals and caused a population decrease of 17% and 35% desertion of nesting areas. In severe El Niños at approximately 12-year intervals, there was a mean decrease of 47% and complete nesting failure. El Niños varied in severity and a possible explanation is presented.In non-El Niño years, commercial fishing did not directly control bird populations through starvation of adults or drowning in nets. Increased commercial fishing apparently led to decreased nesting success: the greater the quantity of fish landed, the lower the percentage increase of the bird population that year.Short-lived, heavily-fished species such as anchoveta are in constant danger of population collapse during years of occasional, unpredictable reproductive failure. Bird populations could be maintained as buffers to prevent terminally-destructive overfishing by humans.     

Genetics of Wild and Managed Populations of <Emphasis Type="Italic">Leucaena esculenta</Emphasis> Subsp. <Emphasis Type="Italic">esculenta</Emphasis> (Fabaceae; Mimosoideae) in La Montaña of Guerrero,Mexico

Genetic consequences of silvicultural management of Leucaena esculenta subsp. esculenta were analyzed from eight allozyme loci in half-sib families of one wild and one managed in situ (selectively cleared) population from La Montaña de Guerrero region, Central Mexico. A reference sample (including wild, feral and cultivated individual plants) from the states of Morelos, Puebla and Guerrero, Mexico was also analyzed. Genetic variation, population structure and mating system were analyzed. All loci showed high variation (75–87.5% polymorphic loci at 95% level; 2.4–2.8 mean number of alleles per locus). All progenies showed heterozygous deficiency, but both wild and managed parental inbreeding coefficients were negative, suggesting heterosis. Progenies of managed populations differed from those of the wild and reference samples (Nei’s unbiased identities 0.874–0.934). Biparental inbreeding is suggested by Wright’s-statistics ( f = 0.313), and by outcrossing rate estimates: t_m = 0.644 (SE 0.094), and 0.645 (SE 0.193); t_s = 0.576 (SE 0.189), and 0.523 (SE 0.182), for managed and wild samples respectively. Population differentiation is significant (Θ = 0.210). The species is self-incompatible and deviations from the mixed mating model were found. Indirect estimates of products of effective population size (N_e) by the proportion of migrants (N_m) were moderate, as were the N _evalues. Variation due to ecotypic differentiation (related to altitude), prolonged artificial selection, and introduction from other areas is supported. A model of domestication of seed-propagated trees is suggested, based on extensive and in situ selection of locally adapted populations, and their diffusion to other areas.     

Phenotype management: a new approach to habitat restoration

The goal of habitat restoration is to provide environmental conditions that promote the maintenance and growth of target populations. But rarely is it considered how the allocation of resources influences the diversity of phenotypes in these populations. Here we present a framework for considering how habitat restoration can shape the development and expression of phenotypes. We call this approach phenotype management as it entails restoring the resources in a habitat to manage phenotypic diversity. Phenotype management is achieved by manipulating the spatial and temporal distribution of resources to alter the degree of competition among individuals. Differences in competition, in turn, lead to changes in phenotypic and life history expression that affect population parameters including demography and effective population size (N_e). To illustrate how phenotype management can be applied, we explore how resource distributions shape variation in phenotypes in two imperiled fishes, Pacific salmon and desert pupfish. In both examples, modulating male reproductive phenotypes changes the allocation of reproductive success among population members to subsequently affect N_e. These examples further demonstrate that whether to increase or decrease phenotypic diversity depends on the primary conservation pressures faced by the species.     

The ecology of extinction: population fluctuation and decline in amphibians

Even among widespread species with high reproductive potentials and significant dispersal abilities, the probability of extinctions should be correlated both with population size variance and with the extent of population isolation. To address how variation in demographic characteristics and habitat requirements may reflect on the comparative risk of species decline, I examined 617 time series of population census data derived from 89 amphibian species using the normalized estimate of the realized rate of increase, ΔN, and its variance. Amphibians are demonstrably in general decline and exhibit a great range of dispersal abilities, demographic characteristics, and population sizes. I compared species according to life-history characteristics and habitat use. Among the populations examined, census declines outnumbered increases yet the average magnitudes for both declines and increases were not demonstrably different, substantiating findings of amphibian decline. This gives no support for the idea that amphibian population sizes are dictated by regimes featuring relatively rare years of high recruitment offset by intervening years of gradual decline such that declines may outnumber increases without negative effect. For any given population size, those populations living in large streams or in ponds had significantly higher variance than did populations of completely terrestrial or other stream-dwelling amphibians. This could not be related to life-history complexity as all the stream-breeding species examined have larvae and all of the wholly terrestrial species have direct development without a larval stage. Variance in ΔN was highest amongst the smallest populations in each comparison group. Estimated local extinction rates averaged 3.1% among pond-breeding frogs, 2.2% for pond-breeding salamanders, and negligible for both stream-breeding and terrestrial direct-developing species. Recoveries slightly exceeded extinctions among European pond-breeding frogs but not among North American pond-breeding frogs. Less common species had greater negative disparities between extinctions and recoveries. Species with highly fluctuating populations and high frequencies of local extinctions living in changeable environments, such pond- and torrent-breeding amphibians, may be especially susceptible to curtailment of dispersal and restriction of habitat.     

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.     

Plasmodium blood parasite found in endangered Galapagos penguins (Spheniscus mendiculus)

This is the first report of a Plasmodium blood parasite found in the Galapagos Archipelago. Phylogenetic analyses place this parasite, recovered from endangered Galapagos penguins (Spheniscus mendiculus), within the genus Plasmodium, and suggest a close relationship to some of the most dangerous lineages of Plasmodium that have been known to cause severe mortality and morbidity in captive penguin populations. Infectious disease is an increasingly important cause of global species extinctions, and extinctions due to avian pox and avian malaria (Plasmodium relictum) have been well documented in Hawaiian avifauna. Plasmodium blood parasites had not been detected in Galapagos birds until now, despite previous microscopic and molecular screening of many of the species, including the Galapagos penguin. While penguin populations now appear healthy, it is unclear whether this parasite will have an obvious impact on their survival and reproduction, particularly during El Niño events, which cause stress due to reduced food availability. It is possible that this parasite arrived with or shortly after the recent arrival of an introduced mosquito, Culex quinquefasciatus, known elsewhere as a competent vector of Plasmodium blood parasites.     

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 population structure of Chinese Cherry revealed by chloroplast DNA trnQ-rps16 intergenic spacers variation

Chinese Cherry, generally referring to Prunus pseudocerasus Lindl. species, is one of the most economically important domestic fruit tree in China. To effectively preserve and sustainably utilize this species, a study in genetic diversity and population structure was carried out by the sequence variation of chloroplast DNA trnQ-rps16 intergenic spacers. Polymorphism was calculated among the 200 individuals from 7 local and 10 wild populations across its geographical range in China. The results showed that (1) Total genetic variation in species level was poor (h = 0.478, π = 0.0018). But in the wild populations, it was much higher than that in the local populations (h = 0.565, π = 0.0021 vs. h = 0.152, π = 0.0006). (2) Low levels of genetic differentiation (F_SC = 0.19595, P = 0.000), genetic distance (0.00000–0.00297), as well as pairwise Fst value among populations were detected. These results, combined with the insignificant gene genealogy pattern and neutrality tests indicate that Chinese Cherry has relatively low levels of genetic diversity and insignificant population structure. Coancestry might be a combined effect of the reduction in effective population size with the recent demographic bottlenecks and mating systems. The pronounced seed dispersal abilities and relatively high rates of gene exchange are supported by the limited genetic differentiation among populations.     

ENSO classification indices and summer crop yields in the Southeastern USA

This research uses a quantitative methodology for directly comparing the responses of observed crop yields in the SE USA to ENSO phenomena classified using dissimilar ENSO indices. ENSO condition has been related to agricultural yields in many parts of the world. It has been generally recognized that the strongest effects on spring and summer crops occur during the boreal winter, and therefore most directly affect spring-summer field crops in the southern hemisphere. However, some ENSO effects on northern hemisphere spring and summer crops have been found, particularly when researchers have used sub-annual indicators of ENSO conditions that, unlike annual ENSO indices, distinguish between continuity and change prior to or during the crop season. To evaluate the utility of such sub-annual ENSO indicators for agriculture in the SE USA, a tercile-based scoring system was devised to compare four distinct ENSO indices: three monthly ENSO indices and the JMA annual ENSO index. Each index was scored in its ability to predict crop yields differing from the historically normal tercile for corn (Zea mays L.), cotton (Gossypiumhirsutum L.), and peanut (Arachis hypogaea L.). Annual crop yield data were used from selected counties in five Southeastern USA states. No geographic differentiation among the data was included in the analysis. This aggregation of county data increased the sample size for each crop, to address the limitation of a short time-series (47 years) distributed among up to 9 ENSO categories. Statistical significance was compared using contingency tables and the Fisher exact test. Peanut and corn yield response matched best to the Multivariate ENSO Index (MEI) and cotton, to the Oceanic Niño Index (ONI). The MEI and ONI are quantitative indices, while the lower-scoring JMA and Modified-JMA indices are categorical. Therefore, future efforts may reduce the focus on categorical (Niño, Niña, Neutral) classification, and explore the response of crop yields to quantitative ENSO data.     

Nitrate removal from drained and reflooded fen soils affected by soil N transformation processes and plant uptake

Knowledge about nitrate transformation processes and how they are affected by different plants is essential in order to reduce the loss of valuable N fertiliser as well as to prevent environmental pollution due to nitrate leaching or N₂O emission after fertilisation or the reflooding of degraded fens with nitrate-containing municipal sewage. Therefore four microcosm ¹⁵N tracer experiments were performed to evaluate the effect of common wetland plants (Phalaris arundinacea, Phragmites australis) combined with different soil moisture conditions (from dry to reflooded) on nitrate turnover processes. At the end of experiment, the total formation of gaseous N compounds was calculated using the ¹⁵N balance method. In two experiments (wet and reflooded soil conditions) the N₂O and N₂ emissions were also directly determined.Our results show that in degraded fen soils, which process mainly takes place—denitrification or transformation into organic N compounds—is determined by the soil moisture conditions. Under dry soil moisture conditions (water filled pore space: 31%) up to 80% of the ¹⁵N nitrate added was transformed into organic N compounds. This transformation process is not affected by plant growth. Under reflooded conditions (water filled pore space: 100%), the total gaseous N losses were highest (77-95% of the ¹⁵N-nitrate added) and the transformation into organic N compounds was very low (1.8% of ¹⁵N nitrate added). Under almost all soil conditions plant growth reduced the N losses by 20-25% of the ¹⁵N nitrate added due to plant uptake. The N₂ emissions exceeded the N₂O emissions by a factor of 10-20 in planted soil, and as much as 30 in unplanted soil. In the treatments planted with Phragmites australis, N₂O emission was about two times higher than in the corresponding unplanted treatment. 15% of the N₂O and N₂ formed was transported via the Phragmites shoots from the soil into the atmosphere. By contrast, Phalaris arundinacea did not affect N₂O emissions and no emission via the shoots was observed.     

Genetic and bioclimatic variation in <Emphasis Type="Italic">Solanum pimpinellifolium</Emphasis>

Solanum pimpinellifolium, due to its close relationship to S. lycopersicum, has been a genetic source for many commercially important tomato traits. It is a wild species found in the coastal areas of Peru and Ecuador. In this study, the genetic variation of S. pimpinellifolium was studied using the diversity found in 10 microsatellites in 248 plants spread throughout its entire distribution area, including Ecuador, which has been underrepresented in previous studies. Peruvian and Ecuadorian accessions are genetically quite differentiated. A possible cause of these differences could be the non-uniform nature of the coastal Ecuadorian and Peruvian climates, seeing as an important correlation between genetic differentiation and climate has been found. In addition, Ecuadorian and south Peruvian accessions have a lower genetic diversity and a higher homozygosity due to their higher autogamy, lower population size, and possible colonization bottlenecks. The Galápagos Islands population is an extreme case, with no diversity, likely caused by a recent colonization from the northern continental Ecuadorian region where genetically identical plants have been found. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Elena Zuriaga and José M. Blanca contributed equally to this work.     

Conservation implications of the distribution of genetic diversity at different scales: a case study using the marsh fritillary butterfly (Euphydryas aurinia)

In the UK, Euphydryas aurinia exists in fragmented habitat patches, and undergoes population fluctuations as a result of a larval parasitoid. Its range is declining in the UK and conservation is thought to require a landscape approach since populations spread over large areas in some years and contract to core breeding patches in others. We examined populations at a range of geographic scales using allozyme electrophoresis to look for evidence of gene flow and differences in genetic diversity among populations. Nationally, our F_ST value was 0.1542 but between population groups within the suspected colonisation range of the butterfly (ca. 20 km), F_ST values were not significantly different from zero. Genetic diversity in terms of number of alleles and heterozygosity was reasonably high in natural populations (H_e=0.267) but low in an introduced, isolated population. We infer that migration between closely spaced subpopulations (in a metapopulation) maintains a high genetic effective population size (large number of individuals in a population that contribute genes to the next generation) which offsets any local reductions in population numbers due to stochastic extinctions or parasitoid effects. We therefore conclude that effective conservation of the species must seek to provide networks of suitable habitat for groups of subpopulations, rather than maintaining habitat for isolated populations.