Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

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.     

Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii

The roosts of many IUCN-listed cave-roosting bat species are under threat from tourist development in SE Europe and other regions of the world. Much-needed conservation strategies require, among other information, an understanding of their roost movements and population dynamics, which can now be obtained relatively quickly using advanced models. We have studied the long-fingered bat, Myotis capaccinii, an obligate cave-dweller, in Dadia National Park, Greece. The species formed colonies of up to a few thousand individuals and was highly mobile, frequently switching summer roosts up to 39 km apart, even during late pregnancy. The bats migrated to distant hibernacula including a cave in Bulgaria 140 km NW of the Park. Adult recapture probabilities varied with season and sex: low female recapture rates in autumn, relative to spring and summer, indicated non-random temporary emigration following nursery colony dispersal. The opposite pattern was seen in males: increasing recapture rates in the autumn suggest that males gather in these roosts to mate with females in transit. Adult survival (0.86-0.94) was similar in females and males, similar in winter and summer, and comparable to recent estimates for other bats based on similar modelling techniques. Sex-based differences in juvenile recapture suggest female philopatry and male-biased dispersal. Our work shows that protection of M. capaccinii roosts must extend beyond the Park’s and indeed the country’s boundaries: its conservation requires large-scale, trans-national integrated conservation plans. Our results will apply to many other warm-temperate species with similar life history cycles.     

Population ecology of the Chinese grouse, Bonasa sewerzowi, in a fragmented landscape

We report the first population ecology study of the endangered Chinese grouse (Bonasa sewerzowi). The study population was stable during 1995-2000. Annual rates of survival were 0.64 for adults and only 0.17 for chicks to 13 weeks of age. Reproductive parameters were 6.1 eggs/clutch, 63% nest success, and overall 3.6 chicks hatched/female/year. The spring sex ratio was 64% males, with 44% of the males unmated in spring. Local people found an estimated 10-29% of the nests and took the eggs for food. Only about 9% of the reserve containing the study area is mature mixed forest, the preferred habitat. Inadequate brood habitat may have influenced chick survival and the skewed adult sex ratio may be a result of habitat fragmentation. We recommend protecting the remaining mixed forest, creating more brood habitat, and stopping the taking of eggs.     

Perceived risk, displacement and refuging in brown bears: positive impacts of ecotourism?

Ecotourism is a rapidly growing industry with unknown impacts on viewed wildlife that may require novel management action. We examined the impact of viewing activities on the behaviour of brown bears (Ursus arctos) in coastal British Columbia.Domination of the best feeding sites and human avoidance by large male bears has consistently been reported. We, however, saw displacement in time rather than space - during the viewing day large males were less active than at other times, while females with cubs tended to be more active.In each year, females with cubs spent similarly high proportions of their time fishing when people were present. In years with large male activity, less time was spent fishing when people were absent. When freed from the potential threat of large male bears, females with cubs showed no measurable impact of controlled human activity.Human presence at a feeding site impacts the behaviour of brown bears, but not as expected. Temporal avoidance of human activity by large males was observed; indications that they departed upon satiation, before the arrival of morning tours, however, suggests that there was little energetic impact. By displacing large males, viewing activities created a temporal refuge, enhancing feeding opportunities for subordinate age/sex classes. With the strong positive relationships between mean female mass and litter size, this may in turn increase population productivity.     

The impact of sport-hunting on the population dynamics of an African lion population in a protected area

Between 1999 and 2004 we undertook an ecological study of African lions (Panthera leo) in Hwange National Park, western Zimbabwe to measure the impact of sport-hunting beyond the park on the lion population within the park, using radio-telemetry and direct observation. 34 of 62 tagged lions died during the study (of which 24 were shot by sport hunters: 13 adult males, 5 adult females, 6 sub-adult males). Sport hunters in the safari areas surrounding the park killed 72% of tagged adult males from the study area. Over 30% of all males shot were sub-adult (<4 years). Hunting off-take of male lions doubled during 2001-2003 compared to levels in the three preceding years, which caused a decline in numbers of adult males in the population (from an adult sex ratio of 1:3 to 1:6 in favour of adult females). Home ranges made vacant by removal of adult males were filled by immigration of males from the park core. Infanticide was observed when new males entered prides. The proportion of male cubs increased between 1999 and 2004, which may have occurred to compensate for high adult male mortality.     

Large-scale noninvasive genetic monitoring of wolverines using scats reveals density dependent adult survival

Noninvasive genetic monitoring has the potential to estimate vital rates essential for conservation and management of many species. In a long-term genetic capture-mark-recapture study using scats we evaluated temporal variation in adult survival in a wolverine (Gulo gulo) population in southern Norway. In contrast to most previous studies of large mammals we found evidence for negative density dependence in adult survival in this large carnivore. Both sexes showed the same pattern of density dependence, with higher annual survival rates in adult females than males. In addition, we also found an additive mortality effect of harvesting in the population, resulting in the lowest adult survival rates at a combination of high population density and high harvest rate. The additive effects of density and harvest on adult survival of wolverines have relevance to the conservation and management of solitary carnivores with strong intrasexual territoriality, especially for species where combats among conspecifics can cause serious injury or even mortality.     

Behavioural response of a trophic specialist, the Iberian lynx, to supplementary food: Patterns of food use and implications for conservation

Prey scarcity compromises population survival, especially for specialist predators. Supplementary feeding is a management tool that can be applied to reverse the decline of food-limited populations. We analyse how a population of Iberian lynx, a threatened food specialist, initially reacted to, and subsequently used, supplementary food. Twenty-seven feeding stations (FS) with domestic rabbits were placed in the Doñana Biological Reserve, SW Spain, between 2002 and 2005. We recorded lynx tracks inside and around stations to analyse spatio-temporal patterns of use, as well as the performance of several station designs. Iberian lynx used 62% of the FS installed, and consumed most of the supplied food. All station designs were used and food provision apparently covered the energetic needs of the lynx inhabiting the reserve. There was spatial aggregation in the use of the FS. Fourteen weeks were needed on average for lynx to become familiar with feeding stations and making a regular use of the supplementary food. Seasonal variations in consumption frequency appeared to be modulated by fluctuations of wild rabbit numbers throughout its annual cycle as well as by variation in energy demand of breeding females. The Iberian lynx responded positively to our supplementary feeding programme. We show that this technique allows the persistence of lynx populations during long periods in areas where wild rabbits are extremely scarce. Supplementary food may be used to sustain lynx whenever rabbit populations need recovery, as well as in the context of lynx restocking or reintroduction programmes.     

Conservation of the red-winged grasshopper, Oedipoda germanica (Latr.): the influence of reproductive behaviour

Reproductive isolation can function as a mechanism to maintain locally adapted gene complexes while decreasing the heterozygosity in distinct populations. As a result, reproductive behaviour should be considered a fundamental factor influencing reproductive isolation. This is of interest to conservation biology when one desires to regulate gene flow between two populations either by creating opportunities for increased dispersal, by relocation of individuals, or by re-introduction of a species to its natural habitat. Reproductive behaviour can also influence the effective population size and the actual population size through the Allee effect. We investigated the reproductive behaviour of individuals from two isolated populations of the red winged grasshopper, Oedipoda germanica, an endangered species in Central Europe. We detail several methods to show how several aspects of the reproductive behaviour of this species interact with the conservation of this species. Foreign males were not disfavoured in mate choice and male body size was also unimportant in mating success. Heterogamic matings were as productive as homogamic matings in terms of total number of eggs per female, egg hatching rate, or nymph survival. Therefore, we suggest that cross-matings of individuals from different populations do not positively influence population size by heterosis effects nor act they negatively in the form of outbreeding depression. We found that female O. germanica were able to store viable sperm for extended periods but egg pods showed a decrease in hatching rate when these females were deprived of additional mating opportunities. Multiple mated females laid more eggs than once-mated females. Hence, females are capable of founding new populations even after only one mating but their reproductive output may be reduced. Present-day gene flow between populations of O. germanica probably does not occur. Consequently, there is an urgent need to pursue habitat management and release programmes that maintain current population sizes of this species. With regard to the reproductive behaviour of the red-winged grasshopper, we suggest that relocation programmes release males and female together and at an early adult stage. Also, because female reproductive output increases with mating activity, the initial release should involve excess females. Because males have a shorter lifespan than females a subsequent release of males at a later time may enhance the overall success of local conservation efforts by resulting in more offspring per female.     

Dispersal of breeding, adult male Phocarctos hookeri: Implications for disease transmission, population management and species recovery

Dispersal impacts on a range of population parameters making it a key piece of information in species conservation. Despite its importance, dispersal is poorly characterized for many species: pinnipeds are no exception. Understanding dispersal patterns of the New Zealand sea lion Phocarctos hookeri is crucial in the conservation management of the species as its recovery to a non-threatened status hinges on range recolonisation. In this study, we examined the movements of breeding adult male New Zealand sea lions within and following the breeding season of the 2002/03 austral summer using a novel multi colony approach. Based on resightings of 202 individually identifiable adult males, we found (1) a previously unappreciated, high level of dispersal by adult territorial males between breeding colonies during the pupping period and (2) that breeding males disperse to the extremes of the species’ range at the end of female oestrous. Our findings are contrary to the current paradigm of otariid breeding behaviour, which is believed to consist of prolonged, uninterrupted male territoriality based on intense male-male competition and sustained fasting. Adult male dispersal between colonies and across the species range has important implications for adult males as vectors of disease in three recent epizootics, species management and species recovery via recolonisation as males are apparently remaining part of a localized, vulnerable breeding population.     

Measuring the cost of risk avoidance in brown bears: Further evidence of positive impacts of ecotourism

The objective of this study was to determine the cost to female brown bears with cubs of avoiding risk, or perceived risk, from both large adult male bears and ecotourists. We measured salmon consumption by brown bears in British Columbia under different risk scenarios (the presence and absence of large male bears and people). Declines occurred despite superabundant food where consumption was never limited by salmon abundance.While there were significant changes in the proportion of time spent foraging and in the consumption of captured fish between periods with and without ecotourists present, there was no change in foraging effectiveness, catch per unit effort.Selection of sub-optimal habitats to reduce predation risk incurs energy costs. Risk avoidance reduced daily salmon consumption by females with cubs by more than one third. By associating with people, females with cubs were able to avoid encounters with large, potentially aggressive males, which avoided people in both time and space. Controlled human activity at feeding sites can provide a temporal feeding refuge for vulnerable age/sex classes. This is the first study to quantify the energetic cost of intra-specific risk-avoidance for a large carnivore.     

Behavioral responses of rats exposed to long-term dietary vinclozolin

Vinclozolin is a fungicide used on food crops with human exposure estimated at approximately 2 microg/kg/day from ingestion; occupational exposure, however, may be greater. The metabolites of vinclozolin have been reported to act as antiandrogens and have adverse effects on reproductive physiology and behavior in animals. Here, pregnant rats were fed soy-free diets containing 0, 10, 150, or 750 ppm of vinclozolin (approximately 0, 0.8, 12, and 60 mg/kg/day for an adult) beginning on gestational day 7, and offspring were continued on these diets through sacrifice at postnatal day 77. Male and female offspring were assessed for changes in several nonreproductive sexually dimorphic behaviors: open field and running wheel locomotor activity, play behavior, and consumption of saccharin- and sodium chloride-flavored solutions. There was a significant interaction of sex with vinclozolin exposure on running wheel activity, which indicated that females in the high-dose exposure group were hypoactive compared to same-sex controls. There was a significant overall effect of vinclozolin exposure on fluid consumption, and high-dose animals showed increased intake of the saccharin solution and decreased intake of plain water while saccharin was available. Effects were more pronounced in females, which drank 40.8% more saccharin than control females, whereas males drank 6.2% more than control males. There were no effects of vinclozolin treatment on play behavior or sodium solution intake. Gestational duration, total and live pups per litter, litter sex ratios, and birth weight were also not significantly affected, nor were body weight and food intake for dams and offspring. These results indicate that long-term dietary exposure to vinclozolin does not have severe toxicological consequences on the nonreproductive behaviors measured here. However, exposure may cause subtle alterations in locomotor activity and consumption of saccharin-flavored solution.     

Survival, interactions with conspecifics and reproduction in 37 chimpanzees released into the wild

We report the results of eight years of post-release monitoring of 37 wild-born, captive chimpanzees released into the Conkouati-Douli National Park, Republic of Congo. Overall survival was high, with 23 (62%) individuals remaining in the release zone, and only 5 (14%) confirmed dead. Released females regularly interacted with wild chimpanzees. Several females appeared to have integrated into wild groups for extended periods of time, and four released females gave birth to a total of five offspring. However, encounters with wild chimpanzees were a major cause of mortality in released males, and 40-50% of released males would have died without veterinary intervention. These sex differences are in accordance with knowledge of chimpanzee behavioural ecology. Our results demonstrate that wild-born, captive chimpanzees can be released into the wild successfully, under certain specific conditions. Most importantly, careful planning and preparation is critical at all stages; a suitable release area must be identified; potential risks to existing wild populations, including the possibility of disease transmission, must be minimised; and post-release monitoring is essential. Adolescent females are the most suitable candidates for release, as they appear to be able to integrate successfully into wild communities. However, males should not be released where wild chimpanzees occur, as they are likely to be attacked and killed. Release into the wild addresses the welfare of certain individual animals, although it clearly cannot address the fate of all captive, wild-born chimpanzees. Knowledge of how to successfully release chimpanzees into the wild also has both current and potential future benefits for the conservation of wild chimpanzee populations.     

Managing supplementary feeding for avian scavengers: Guidelines for optimal design using ecological criteria

Current European sanitary laws are promoting the concentration of domestic livestock carcasses in limited locations called ‘vulture restaurants’ or supplementary feeding stations in order to safeguard human health. However, this type of food concentration could lead to monopolization by a few, large dominant species. Management guidelines are urgently needed to ensure that potentially less competitive and more endangered scavengers can also benefit from this resource, considering that once abundant carcasses are now absent from the field. Here, we assess factors affecting the abundance of six avian scavenger species at 17 feeding stations in northern Spain, considering aspects such as carrion availability, physiographic features, humanization, presence of heterospecifics and densities of scavengers during both winter and summer seasons. Results indicate that conditions for interspecific facilitatory processes are possible at low numbers (<100 individuals) of griffon vultures (Gyps fulvus). Otherwise, the other avian scavengers are excluded. In a few cases, spatio-temporal patterns of segregation become apparent. Differences in the size of carcasses supplied to feeding stations as well as the local characteristics influence the balance of species at these points and, therefore, how species of conservation concern use feeding stations. Future legislation should encourage the opening of numerous feeding stations supplied with low quantities of food to mimic the original condition of temporal and spatial unpredictability of carcasses and to maintain ecological relationships within the scavenger guild.     

Home range size and daytime habitat selection of leopards in Huai Kha Khaeng Wildlife Sanctuary, Thailand

Leopards (Panthera pardus) are endangered in South East Asia yet little is known about which resources need to be secured for their long-term conservation or what numbers of this species this region can support. This study uses radio telemetry to investigate seasonal variation in habitat selection and home range size of Leopards in Huai Kha Khaeng Wildlife Sanctuary, Thailand. Over a five year period, 3690 locations were recorded from nine individuals. The mean ± standard error of fixed kernel home range size for six adult females was 26 ± 8.2 km², for two adult males was 45.7 ± 14.8 and for two sub-adult females was 29 km² ± 5.5. Adult female wet and dry season home range sizes did not differ significantly. One adult male showed an increase in home range size from dry to wet seasons. Estimated density was 7 adult females/100 km², which suggests 195 adult female leopards living in Huai Kha Khaeng alone, thus highlighting the larger Western Forest Complex’s potential contribution to leopard conservation. Compositional analysis of second and third order habitat selection suggested mixed deciduous and dry evergreen forest types, flat slope and areas close to stream channels are important landscape features for leopards. These results can help formulate a much needed conservation strategy for leopards in the region.     

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

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

Hybridization and conservation of Mediterranean orchids: Should we protect the orchid hybrids or the orchid hybrid zones?

Natural hybridization between plant species often occurs in disturbed habitats and it is generally considered a threat for rare and endangered species. A different situation occurs in Mediterranean food deceptive orchids, where hybridization is a common phenomenon, as a natural consequence of their unspecific pollination system.Here, we present molecular and ecological evidence from a hybrid zone between Orchis mascula and O. pauciflora in order to address the consequences of hybridization on local orchid evolution and to assess conservation priorities for hybrids and hybrid zones in orchid conservation programs.We find that, although hybrids among the two target taxa are formed relatively frequently, hybrid zones often consist of F1 individuals, whereas backcrosses and later generation hybrids appear to be either rare or absent, presumably as a consequence of post-zygotic reproductive barriers acting in these food-deceptive orchids. Experimental evidence further indicates that hybrids are less fit than the parental species in attracting pollinators and that parental taxa exhibit higher or similar fitness in sympatry compared to allopatric populations. Since introgression is not frequent as indicated by molecular analyses, fitness and phenotypic trait differences observed among sympatric and allopatric populations of the parental taxa may be a consequence of pollinator-mediated selection.Our experimental data confirm that hybridization is a natural phenomenon in food-deceptive orchids and that it does not pose a threat to their survival. Furthermore, sympatric zones provide the stage for evolutionary processes in orchids, and this peculiarity should be taken into account when devising orchid conservation strategies.     

Female biased mortality caused by anthropogenic nest loss contributes to population decline and adult sex ratio of a meadow bird

Meadow breeding birds such as the whinchat Saxicola rubetra have been declining due to increased farming intensity. In modern grassland management, the first mowing and the bird’s breeding cycle coincide, causing high nest destruction rates and low productivity of grassland bird populations. However, it is virtually unknown whether the mowing process directly affects adult survival by accidentally killing incubating females. We studied adult survival of an Alpine whinchat population during two breeding seasons using either colour-ringing or radio-tracking of 71 adults. Assessing territories, mowing phenology and nest destruction from 1988 to 2007 allowed changes in the factors associated with female mowing mortality to be estimated. Adult survival over 5-day-periods was Φ = 0.986, but during the period of mowing female survival was strongly reduced (Φ = 0.946). As a result, 80.6% of the males, but only 68.4% of the females survived the breeding season. Mowing undoubtedly killed two of 20 radio-tagged females when they were laying or incubating. In the 20-year period, an increasing proportion of nests were destroyed before the chicks hatched and this change was associated with an increased distortion of the adult sex ratio. Modelling the population growth rate showed that including the additional effect of mowing on female mortality resulted in a 1.7 times faster local population decline. These results are consistent with the hypothesis that the extinction of whinchat populations in the lowlands of central Europe was caused not only by habitat degradation and low productivity, but also by increased man-made female mortality.     

Responses of cycads with different life histories to the impact of plant collecting: simulation models to determine important life history stages and population recovery times

Collection of plants and seeds from wild populations threatens a large number of cycad species. We investigated to what extent individual life history stages contribute to population growth (λ) and compared two species with major differences in life histories in the African genus Encephalartos: Encephalartos cycadifolius, a highly persistent grassland species that resprouts after fire, and Encephalartos villosus, a relatively fast growing, non-sprouting forest species. Several harvesting scenarios impacting different sized individuals were simulated to determine the sensitivity of the two functional types to harvesting. In both species λ was most sensitive to changes in abundance of adult plants. The harvesting of seeds had minimal impact on population growth rates, whereas harvesting of adult plants led to rapid population decline. This response from two very different functional types suggests that the conservation of adult plants is critical for all cycad species. Despite similar responses to adult mortality, the two species had substantially different population growth rates. This determined recovery time after harvesting of adult individuals. Encephalartos cycadifolius is typical of highly persistent plant species associated with low levels of recruitment and unable to recover from even small losses of adults within a reasonable conservation time frame (<100 years). Our results suggest that the ability to recover from loss of individuals is an important factor that should be considered when assessing the vulnerability of wild populations to threats.