Mating system in a gopher tortoise population established through multiple translocations: Apparent advantage of prior residence

Population manipulations such as translocation are becoming increasingly important tools in the management of rare and declining species. Evaluating the effectiveness of such manipulations requires comprehensive monitoring of population processes, including dispersal, survivorship, and reproduction. We investigated the mating system of a translocated population of gopher tortoises (Gopherus polyphemus) established through multiple releases, which occurred primarily during 1987–1994. During 2006–2007, we sampled and genotyped 27 candidate males (candidate sires), 34 candidate females (candidate dams), and 121 offspring from 19 clutches at five polymorphic microsatellite loci to determine the relative frequency of multiple paternity and to estimate individual reproductive success. Multiple paternity was detected in 57% of clutches genotyped, and females of single-sire clutches and females of multiple-sire clutches were of similar size. Reproductive success varied among male tortoises, and successful sires were significantly larger than males to which no offspring were attributed. Among successful sires, previously established males sired a disproportionate number of the offspring sampled, despite being significantly smaller than subsequently released males. The high variance in individual reproductive success and the apparent reproductive advantage associated with prior residence observed in this gopher tortoise population has important implications for the design of future translocation projects.     

A nine-year study of successful breeding in a Bonelli’s eagle population in southeast Spain: a basis for conservation

We analyse the factors influencing breeding success in a healthy population of Bonelli’s eagle (Hieraaetus fasciatus) from south-east Spain, a raptor that has suffered a serious decline in Europe. Between 18 and 33 pairs were annually monitored during the period 1994-2002. Several factors that may affect four breeding parameters were studied, namely human presence, vegetation, relief, climatic factors, intra- and inter-specific relationships, diet, prey abundance, nest building, adult mortality and age of reproduction. A consistently high breeding success was registered during the study period (productivity=1.43, SD=0.11), which was probably the result of high adult survival, adequate prey availability and mild weather conditions. However, a certain vulnerability to the presence of golden eagles (Aquila chrysaetos) and to human disturbance was observed. One interesting result was the reduced survival of young chicks on north-facing cliffs owing to colder conditions, which may partially explain the decline of the Bonelli’s eagle populations along its European distribution limits. We discuss the role of this healthy subpopulation in an Iberian metapopulation context, and propose that the potential interference of golden eagles should be taken into account when designing management strategies for Bonelli’s eagles. Finally, we pinpoint some conservation priorities and the importance of reducing the main causes of adult mortality (i.e. shooting and electrocution) to assure successful reproduction and survival of this species.     

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.     

Higher breeding densities of the threatened little bustard Tetrax tetrax occur in larger grassland fields: Implications for conservation

Previous studies have found that densities of little bustard Tetrax tetrax breeding males tend to be higher in areas with smaller agricultural fields, presumably due to increased habitat diversity. However, exceptionally high densities have been found in large grassland fields in Portugal, which suggests that the influence of field size varies geographically, and that the role of this factor is not yet fully understood, despite its importance as a key management issue.We studied how field size, together with vegetation structure, influences the presence and density of breeding little bustards in a region of southern Portugal. Fifty-four grassland fields were sampled in 2007 and another 29 in 2008, with sizes ranging from 23 to 172 ha. A total of 183 breeding males were found in 47 of these fields, reaching densities of up to 37 males/100 ha. A higher probability of occurrence of breeding males was found in larger fields with a vegetation height below 40 cm and field size alone explained 46% of the variability in male density. These results suggest that larger continuous areas of suitable habitat attract many males, most likely as a consequence of their lek mating system. We conclude that conservation efforts, in a landscape context of large farm sizes, should: (1) be channelled to farms with large fields; (2) ensure adequate livestock grazing to create suitable habitat and (3) promote management at a landscape level to ensure the most continuous grassland habitat patches possible.     

Reproduction and pre-dispersal survival of Iberian lynx in a subpopulation of the Doñana National Park

Little is known about the reproduction of the Iberian lynx (Lynx pardinus) even though it is the most endangered felid in the world. We studied during 9 years several reproductive parameters of the Iberian lynx in one of the subpopulations situated in Doñana National Park (south-western Spain), by means of radio-tracking, direct observations and photo-trapping. The potential breeding subpopulation was usually composed of 3 adult females, which bred 83% of the total possible 29 female-reproductive year. The minimum total number of cubs born during the study was 64 (7.7 ± 0.69 per year). There was no correlation between the number of cubs born or number of breeding females and population size of European rabbit (Oryctolagus cuniculus, the main prey of the lynx). All known births (n = 16) occurred in March except one in April and another in June. Mean litter size was 3.0 ± 0.16 (n = 16, range = 2-4). Sex-ratio (n = 59 cubs) was 1.03:1.00 (females:males). Nevertheless, the commonest picture was a female with 2 cubs older than 3 months. Altogether, at age of 3 months, 75% of cubs survived. Number of lynx alive at 10 months old and before dispersing was 69% and 57%, respectively. Sex did not affect survival for any age. Normally, it was not possible to know the causes of the death of cubs younger than 3 months. Survival at this age was not related with mother, mother age, or rabbit abundance. All females that bred were older than 3 years. The age of last reproduction was 9 years. For 3 females that were tracked during almost their complete reproductive life, the life time reproductive output was between 11 and 19 cubs. Iberian lynx reproductive parameters did not respond to wide changes in prey abundance during the study. Conservation plans considering the extraction of cubs with a low survival probability should be considered by managers, for instance, in translocation campaigns.     

Effective population size in a bottlenecked fur seal population

The census population size (N) is usually the only information available for most threatened species. For evolutionary matters, the effective population size (N_e), not the census number, is a prime concern. Factors such as variation in the sex ratio of breeding individuals, variation of population size in different generations and mating system are important. The South American fur seal, Arctocephalus australis, has been exploited in Peru by humans since ca. 2000 BC and now the original population declined 72%, as a result of low food availability during the severe El Niño in 1997-1998. In this sense A. australis is now classified as in danger of extinction in Peru. We present the first estimate of N_e of the Peruvian population of A. australis that takes into account the effects of mating system and variation in population size caused by the 1997-1998 El Niño. The resulting N_e was 2153 specimens. We believe that the estimated N_e for the Peruvian population is a critical value, because it is significantly lower than the mean minimum viable population for vertebrates (7000 breeding age adults). This estimated N_e is of critical importance because combined with the current El Niño events are reasons of great concern for the survival of the species and should be taken into account in future management plans to ensure the conservation and protection of the species in the Peruvian coast.     

Autumn swarming behaviour of Natterer’s bats in the UK: Population size, catchment area and dispersal

Many species of temperate bats visit underground sites in autumn, prior to hibernation, and chase each other in and around the entrances in an event known as autumn swarming. It has been suggested that swarming is a mating event that facilitates gene flow between bats from otherwise isolated summer colonies. We describe swarming behaviour at a cluster of four caves in the North of England and provide direct evidence for seasonal migration of Natterer’s bats (Myotis nattereri ) between these caves and many summer nursery roosts in a catchment area of at least 60 km radius. Mark-recapture analysis suggests that an estimated 2000-6000 Natterer’s bats use the three best-studied caves, together with smaller numbers of four other species: M. daubentonii, M. brandtii, M. mystacinus and Plecotus auritus. Recaptures also show that there is a small interchange of bats between these three caves, which are 0.6-1.5 km apart. Capture rates, automatic logging and video recordings at one cave suggest that an average 310-400 bats visit the cave each night. Mating at swarming sites would be particularly important for gene flow if a species showed no other dispersal pattern. We provide evidence for natal philopatry, with little or no dispersal, in both sexes of Natterer’s bats. The protection of swarming sites is discussed in light of our findings.     

Habitat fragmentation reduces plant fitness by disturbing pollination and modifying response to herbivory

In fragmented landscapes plant species are often confined to remnants of formerly more widespread habitats, with many of their populations being small and isolated. This study experimentally examined the effects of population size and isolation on pollination, herbivory and reproductive success in the forest herb Phyteuma spicatum (Campanulaceae). In an experiment in which population size and isolation were manipulated using plants from the same origin, population size positively affected pollinator visitation, but did not alter the generally high levels of herbivory. As a result, seed production was higher in large populations. Conversely, plants originating from 14 natural populations of varying size and degree of isolation did not differ in reproductive success when grown in the same environment, suggesting similar attractiveness to pollinators and reproductive potential. The intensity of herbivory, however, was higher in progeny of small populations, at least in terms of the proportion of biomass removed. In both experiments, there were no effects of population isolation. The results suggest (1) that small population size decreases reproductive success via direct negative effects on plant-pollinator interactions, (2) that this pattern is not offset by herbivory, but (3) that herbivory enforces fragmentation effects on pollination by further reducing the number of flowering individuals and (4) that habitat fragmentation may influence plant fitness by affecting plant response to herbivory. The effects of habitat fragmentation on plant populations in present-day landscapes are thus complex, illustrating the need for more integrated studies in conservation biology that take into account both mutualistic and antagonistic plant-animal interactions.     

Role of human-modified habitat in protecting specialist species: A case study in the threatened Florida Scrub-Jay

Habitat specialists are especially vulnerable to habitat alterations in vegetation structure and composition, making them difficult to protect and restore amidst agricultural landscapes. Protection strategies in such cases require information on species’ ability to survive and reproduce on marginal and modified habitat. We examined reproductive success, survival, and foraging efficiency of the Florida Scrub-Jay (Aphelocoma coerulescens), an extreme habitat specialist, in a human-modified habitat - regenerating pasture - along a pasture-native scrub interface. From 1985 to 2003, Florida Scrub-Jays were equally successful at producing young in regenerating pasture and native scrub. Production of eggs, nestlings, fledglings, independent young, and yearlings were not statistically different between territories containing pasture and those containing only scrub. Similar trends were observed for nest success and survival. When in pasture, individual jays were significantly more efficient at capturing small prey items, but significantly less efficient at capturing medium and large prey items. Availability of small prey items was significantly higher in pasture than in scrub; availability of medium and large prey items was not significantly different between habitat types. We conclude that regenerating pasture provides suitable supplemental habitat for Florida Scrub-Jays when in close proximity to native scrub, suggesting a novel conservation strategy for maintaining and potentially increasing local populations in fragmented agricultural settings. This case study illustrates how a broadened definition of ‘suitable habitat’ may augment existing conservation strategies for specialist species facing substantial and rapid modification of their native habitat.     

Correlates of abundance and breeding success in the globally threatened Hinde's babbler (Turdoides hindei) and its congener, northern pied babbler (T. hypoleucus)

We investigated the relationship between habitat quality, abundance, group size and breeding success in the threatened Kenyan endemic Hinde's babbler (Turdoides hindei; Family Timaliidae), a co-operative breeder mainly associated with riverine thickets in cultivated areas. At two intensively cultivated sites, significantly more Hinde's babbler groups occurred in areas retaining at least 3% thicket cover, while breeding outcome improved where cover exceeded 9%. Despite previous concerns over low breeding success, the percentage of young birds present (17%, n=424 birds aged) was similar to that of a sympatric congener, the northern pied babbler (T. hypoleucus) (12%, n=162), and comparable with that of two other African congeners. Although breeding outcome was positively correlated with adult group size, there was no evidence that larger groups monopolised more favourable habitat or defended larger territories. Group density showed a positive, non-linear relationship with adjacent cover, levelling out where this exceeded 15-20%. Our findings suggest that even a modest increase in thicket cover may significantly raise the density and productivity of Hinde's babblers at these sites.     

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.     

Landscape occupancy and local population size depends on host plant distribution in the butterfly Cupido minimus

We tested whether the landscape occupancy and local population size of the monophagous butterfly Cupido minimus can be predicted by patch size and isolation of its host plant or by other habitat characteristics. C. minimus and its larval food plant Anthyllis vulneraria are classified as rare and endangered in northern Germany. Adults of C. minimus are ranked as the most sedentary butterfly species in northern Europe.Around the city of Göttingen (Germany), we checked all known locations of A. vulneraria (n=70) in June 2002 for butterfly eggs (in blooming flowerheads) and adult butterflies (within 20-min transects).We found eggs of C. minimus or a high number of adults (>7) in all habitats with A. vulneraria (which are calcareous grasslands) even when isolated up to 2-4 km. In multiple regression analyses, local population size of adult butterflies was positively related to the cover of its larval food plant A. vulneraria explaining 65% of variance. Cover of A. vulneraria increased with increasing habitat area and increasing cover of plant species in flower and decreased with increasing cover of shrub layer. Habitat isolation and further factors describing habitat quality were not related to C. minimus population size or cover of its larval food plant.The results suggest that dispersal ability of C. minimus is greater than expected and that management should focus to increase A. vulneraria patches. For conservation, low impact grazing once a year and removing of excessive shrubs in winter seems to be the most appropriate strategies.     

Genetic and demographic responses of fragmented Acacia dealbata (Mimosaceae) populations in southeastern Australia

Genetic and demographic studies of fragmented populations of common plant species often reveal negative impacts that are likely to constrain persistence. Examining species that are broadly representative of functional groups within fragmented landscapes is one approach to providing a better understanding of how these processes will influence vegetation persistence. Acacias are a significant component of the Australian flora, with Acacia dealbata being a common and representative species of fragmented landscapes across New South Wales. Previous reproductive assessments of fragmented A. dealbata populations indicated significant constraints for small populations through low reproductive output mediated by fertilisation success. This study examined genetic diversity, mating system, and progeny growth parameters of the seed crops produced by these populations to assess whether further constraints to persistence could be detected. Spatially explicit simulation studies were also conducted to assess the persistence likelihood of fragmented populations. Landscape parameters such as population size and plant density were useful predictors for some of the genetic and demographic responses, but a poor response signal was generally observed. Strong evidence for a self-incompatibility mechanism was observed in A. dealbata and is likely to be the major driver of population persistence. Self-incompatibility in small populations limits mate availability and eliminates inbred progeny early in the reproductive cycle leading to poor reproductive output. The simulation data provides further evidence that mate limitation in smaller populations (<200 plants and 40 S alleles) constrains reproductive output and persistence. These data indicate that introducing new germplasm to smaller populations can dramatically improve their persistence likelihood.     

The importance of mating system in translocation programs: reproductive success of released male bridled nailtail wallabies

Translocation is an important tool for the conservation of species that have suffered severe range reductions. The success of a translocation should be measured not only by the survival of released animals, but by the reproductive output of individuals and hence the establishment of a self-sustaining population. The bridled nailtail wallaby is an endangered Australian macropod that suffered an extensive range contraction to a single remaining wild population. A translocated population was established and subsequently monitored over a four year period. The aim of this study was to measure the reproductive success of released males using genetic tools and to determine the factors that predicted reproductive success. Captive-bred and wild-caught animals were released and we found significant variation in male reproductive success among release groups. Variation in reproductive success was best explained by individual male weight, survival and release location rather than origin. Only 26% of candidate males were observed to sire an offspring during the study. The bridled nailtail wallaby is a sexually dimorphic, polygynous macropod and reproductive success is skewed toward large males. Males over 5800 g were six times more likely to sire an offspring than males below this weight. This study highlights the importance of considering mating system when choosing animals for translocation. Translocation programs for polygynous species should release a greater proportion of females, and only release males of high breeding potential. By maximizing the reproductive output of released animals, conservation managers will reduce the costs of translocation and increase the chance of successfully establishing a self-sustaining population.     

Island differences in population size structure and catch per unit effort and their conservation implications for Komodo dragons

Species inhabiting archipelagos are often characterised by high levels of interpopulation divergence (e.g. size related traits). This divergence may, in turn, influence their life-history. To facilitate better management and conservation of the Komodo dragon (Varanus komodoensis), an island endemic, we identified demographic differences between two island populations in Komodo National Park, Indonesia. Comparison of data collected from dragon populations inhabiting Rinca Island and the much smaller Gili Motang Island indicated that between 1994 and 2004, the Komodo dragon population on Gili Motang significantly decreased its: (1) mean body mass, (2) body condition and (3) relative abundance. These results suggest that the numerically small Gili Motang population was oscillating downwards; in contrast, the Rinca Island population had been relatively stable. More importantly these results emphasize the necessity for managers of this priority conservation species to understand further the inherent functional differences among dragon populations to develop island specific management units. Current management practices (e.g. monitoring) instigated by Komodo National Park management ignore small island dragon populations and thus run the risk of being unable to detect adverse effects for populations that are potentially most prone to decline.     

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.     

Differences in the reproductive success and survival of the rufous treecreeper (Climacteris rufa) between a fragmented and unfragmented landscape

The rufous treecreeper (Climacteris rufa) has declined in abundance in the agricultural regions of southwestern Australia. The patterns of decline are well documented, but the processes that threaten population persistence are poorly understood. I compared the reproductive success and survival of the treecreeper between three sites in an unfragmented landscape and four remnant categories (large, small, grazed and ungrazed) in a fragmented, agricultural landscape. Nest success and annual productivity were significantly higher in the unfragmented landscape, but varied between sites and remnant categories within landscapes. Nest success was lowest in grazed remnants and annual productivity was positively associated with territory size in the fragmented landscape. Fledgling survival rates did not differ between landscapes, but there was a trend for juvenile survival rates to be higher in the unfragmented landscape. I used artificial nests to compare relative predation rates between landscapes, and provisioning rates and prey biomass brought to nestlings to assess differences in food availability. There were no landscape differences in predation rates, but provisioning rates to nestlings and total prey biomass were significantly lower in the fragmented landscape. Mean habitat quality was also lower in the fragmented landscape, although it differed between remnant categories. Reduced reproductive success, juvenile survival, food availability and habitat quality may threaten the viability of the rufous treecreeper population living in the fragmented landscape. Limiting the modification of remaining habitat (e.g. removing stock grazing) and improving habitat quality are required to assist in the conservation of this species.     

Effects of supplementary feeding on the offspring sex ratio of kakapo: a dilemma for the conservation of a polygynous parrot

The kakapo (Strigops habroptilus) is a large, flightless, nocturnal parrot, endemic to New Zealand. It is critically endangered, with a world population of ca. 62 individuals and a male-biased adult sex ratio. The species has a polygynous “lek” mating system and adult males typically weigh 30-40% more than females. The kakapo is subject to intensive conservation efforts, including the provision of supplementary food to wild birds to encourage successful nesting. There is mounting evidence that, in polygynous species with large variance of male reproductive success, females in better condition may maximise their fitness by producing more offspring of the larger, more costly sex to be reared. We used data on the sex ratio of progeny of female kakapo that had or had not received supplementary food, to test the hypothesis that supplementary feeding might cause a male-biased offspring sex ratio. There was a significant excess of males in the clutches of females provided with supplementary food, suggesting that changes need to be made to the feeding regime to increase recruitment of females. This is an example of applying evolutionary theory to a practical conservation problem.     

Grassland nature reserves for breeding wading birds in England and the implications for the ESA agri-environment scheme

The two principal mechanisms available to conserve breeding wading birds on lowland wet grassland in England are nature reserves and agri-environment schemes, particularly the Environmentally Sensitive Area (ESA) scheme. ESAs offer payments to landowners in return for maintaining (‘low tiers’) or enhancing (‘high tiers’) the landscape, biodiversity and historic value of the land. Lowland wet grassland nature reserves managed by the Royal Society for the Protection of Birds (RSPB) have been successful at conserving breeding lapwings and redshank but habitat management usually only results in short-term increases in numbers of breeding snipe. Within ESAs, populations of breeding wading birds fare better on land managed under high tiers than on land managed under low tiers. Land managed under high tiers also supports more pairs of breeding wading birds per £ of ESA grant received than land managed under low tiers. Means of increasing the cost-effectiveness of the ESA scheme in conserving breeding wading birds are discussed in the light of these results.     

Reproductive biology and conservation implications of three endangered snapdragon species (Antirrhinum, Plantaginaceae)

About 32% of Antirrhinum species are considered to be endangered; however, no field studies have focused on their reproductive biology. In this work, several aspects of the reproductive biology (flowering phenology, floral biology, breeding system) and potential limits on seed quantity and quality (pollen limitation, inbreeding depression) were studied in natural populations of three endangered species of the genus (Antirrhinum charidemi, Antirrhinum subbaeticum, Antirrhinum valentinum). Results disclose that all three species need insect visitors for seed production since fruit set after autonomous self-pollination was lower than under hand cross-pollination. A. charidemi and A. valentinum were mainly self-incompatible, whereas A. subbaeticum was self-compatible but herkogamous. Supplementary pollination in open-pollinated flowers only increased fruit set and seed set relative to controls in a given population of A. valentinum. Preliminary data on inbreeding depression at early life-cycle stages of the self-compatible A. subbaeticum revealed that the cumulative level was low. Despite the three species being closely related and sharing many ecological characteristics, they show different mating systems, and different factors limit seed quantity and quality. Thus, caution should be taken when making a common conservation plan for a group of closely related taxa.