Animal Conservation

Cover image for Vol. 19 Issue 6

Early View (Online Version of Record published before inclusion in an issue)

Edited By: Res Altwegg, Darren Evans, John Ewen, Iain Gordon, Jeff A. Johnson, Nathalie Pettorelli and Julie Young

Impact Factor: 2.788

ISI Journal Citation Reports © Ranking: 2015: 10/49 (Biodiversity Conservation); 48/150 (Ecology)

Online ISSN: 1469-1795

Associated Title(s): International Zoo Yearbook, Journal of Zoology, Remote Sensing in Ecology and Conservation

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  1. 1 - 34
  1. Original Articles

    1. Costs are key when reintroducing threatened species to multiple release sites

      K. J. Helmstedt and H. P. Possingham

      Version of Record online: 29 NOV 2016 | DOI: 10.1111/acv.12319

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      Multiple sites might be options for threatened species reintroductions; therefore, managers must determine when to prepare sites for new populations, release individuals into those sites, and eventually cease releases. We find optimal release schedules for bridled nailtail wallabies (Onychogalea fraenata) with three different cost scenarios: no ongoing site-management cost, a financial ongoing site-management cost, and a demographic cost of continuous releases. Our general results provide a guide for planning future reintroduction programs and illustrate the importance of categorizing and understanding ongoing costs for reintroduction planning.

    2. Social implications of a colony collapse in a highly structured vertebrate species (long-tailed bat, Chalinolobus tuberculatus)

      J. M. Monks and C. F. J. O'Donnell

      Version of Record online: 28 NOV 2016 | DOI: 10.1111/acv.12324

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      Strong social structuring within a population can confer fitness advantages to group members, but may also impact on the ability of a population to recover from local extinctions. We used a 19-year mark-recapture dataset of long-tailed bats, in which one colony collapsed during a period of high predator numbers, to investigate the social impacts of a colony collapse. The 11 known survivors were accepted into the neighbouring colony, demonstrating resilience of highly structured vertebrate populations to local crashes.

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      Translocation and hand-rearing result in short-tailed albatrosses returning to breed in the Ogasawara Islands 80 years after extirpation

      T. Deguchi, F. Sato, M. Eda, H. Izumi, H. Suzuki, R. M. Suryan, E. W. Lance, H. Hasegawa and K. Ozaki

      Version of Record online: 28 NOV 2016 | DOI: 10.1111/acv.12322

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      We investigated the attendance and breeding attempts of hand-reared short-tailed albatross chicks translocated to a historic breeding island in the Ogasawara Islands, 350 km from the natal island, for 8 consecutive years after the first translocation. Thirty-nine per cent of hand-reared birds returned to the translocation site at least once per breeding season, of which 67% also visited the natal island. Our preliminary results suggest that even though more translocated and hand-reared albatrosses visited and recruited to their natal island compared to the translocation site, the early re-establishment of breeding by short-tailed albatrosses in the Ogasawara Islands 80 years after extirpation would not have occurred without the initial translocation effort.

    4. When the neighbourhood goes bad: can endangered black robins adjust nest-site selection in response to the risk of an invasive predator?

      C. Lawrence, D. Paris, J. V. Briskie and M. Massaro

      Version of Record online: 21 NOV 2016 | DOI: 10.1111/acv.12318

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      Chatham Island black robins are subject to nest predation by invasive European starlings. Robins re-nest in safer sites following predation of an earlier nest, suggesting robins are showing adaptive behavioural responses to this novel nest predator.

    5. Rebuilding beluga stocks in West Greenland

      M. P. Heide-Jørgensen, R. G. Hansen, S. Fossette, N. H. Nielsen, D. L. Borchers, H. Stern and L. Witting

      Version of Record online: 11 NOV 2016 | DOI: 10.1111/acv.12315

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      Beluga whales are hunted in many Arctic communities and it is necessary to promote a scientific basis for establishing sustainable exploitation levels. This study utilizes a long-term time series of abundance estimates and data on sea ice coverage to assess future harvest levels of belugas in West Greenland.

  2. Letters to the Editor

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  3. Original Articles

    1. The vacant planting: limited influence of habitat restoration on patch colonization patterns by arboreal marsupials in south-eastern Australia

      D. B. Lindenmayer, A. Mortelliti, K. Ikin, J. Pierson, M. Crane, D. Michael and S. Okada

      Version of Record online: 31 OCT 2016 | DOI: 10.1111/acv.12316

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      Remnant native vegetation cover in Australian temperate agricultural landscapes includes old growth woodland, regrowth woodland and planted (restored) woodland. We quantified occupancy and colonization probability of these kinds of woodland by arboreal marsupials over an 11-year period. Occupancy levels were highest in old growth, approached zero in plantings, with regrowth intermediate between these two vegetation types. Plantings were not occupied at the outset of our investigation and only rarely colonized throughout the subsequent 10 years, most probably as a result of a lack of shelter sites in large old hollow-bearing trees. The probability of colonization was positively related to the amount of vegetation cover in the landscape surrounding a patch, but not influenced by a temporal increase in woody vegetation cover. Our findings emphasize the value of old growth woodland for arboreal marsupials and suggest that restoration efforts be targeted around old growth and regrowth woodland patches.

    2. Predicting landscape connectivity for the Asian elephant in its largest remaining subpopulation

      J.-P. Puyravaud, S. A. Cushman, P. Davidar and D. Madappa

      Version of Record online: 25 OCT 2016 | DOI: 10.1111/acv.12314

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      To better assess population connectivity for the Asian elephant in the Nilgiri Biosphere Reserve, we evaluated a combination of three resistance layers and three dispersal abilities with a connectivity modeling tool over the landscape. We evaluated the optimality of 24 expert corridors and obtained a much better agreement between the calculated corridors and the expert corridors than the expert teams achieved. Our results provide the first rigorous, spatially synoptic and empirically validated evaluation of the connectivity of the elephant population across the reserve.

    3. Fishers' knowledge as an information source to investigate bycatch of marine mammals in the South China Sea

      M. Liu, M. Lin, S. T. Turvey and S. Li

      Version of Record online: 21 OCT 2016 | DOI: 10.1111/acv.12304

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      Using fishers’ knowledge as an information source, by-catch of marine mammals in the South China Sea were investigated, including which fishing gears caused by-catch, spatial pattern and potential seasonality of by-catch, and possible relation between by-catch and fishing activities, as well as fishers’ attitudes and behaviors towards by-catch, and they perceptions about marine mammals. Based on these findings, we provide new baseline data on regional fishing methods and activities, associated geographic and seasonal patterns of by-catch for different marine mammal species around Hainan, and patterns and drivers of other interactions between marine mammals and fishers, all of which will strengthen the evidence-base for marine mammal conservation and management in the South China Sea.

    4. Mismatch between goals and the scale of actions constrains adaptive carnivore management: the case of the wolverine in Sweden

      M. Aronsson and J. Persson

      Version of Record online: 19 OCT 2016 | DOI: 10.1111/acv.12310

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      We show how wolverines in Sweden colonize areas with low conflicts and poor snow cover where they are not monitored. Instead, management focuses on high-conflict areas and dismisses an opportunity to implement sound population-level management. This illustrates the importance of flexible management that considers the entire range of conditions under which carnivores, people and the environment interact, because the solution may be found outside areas with high management focus.

    5. Forecasting disturbance effects on wildlife: tolerance does not mitigate effects of increased recreation on wildlands

      B. P. Pauli, R. J. Spaul and J. A. Heath

      Version of Record online: 17 OCT 2016 | DOI: 10.1111/acv.12308

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      We created TRAILS (Tolerance in Raptors and the Associated Impacts of Leisure Sports), an individual-based model that simulates the interaction between recreationists and nesting raptors, to assess the effects of human disturbance on raptor populations and test if changes in tolerance to disturbance could mitigate negative consequences of increased recreation activity. We used behavioral and demographic data from golden eagles (Aquila chrysaetos) and recreation activity data to parameterize TRAILS and simulate the likelihood of territory occupancy, egg-laying, and nest survival of eagles in the absence of recreation, with stationary 2014 levels of recreation, and with annual increases in recreation. Recreation activity significantly lowered the population growth rates, population sizes, and territory occupancy of eagles and no simulated method of tolerance acquisition buffered eagle populations from the detrimental effects of recreational disturbance.

    6. Defaunation and biomass collapse of mammals in the largest Atlantic forest remnant

      M. Galetti, C. R. Brocardo, R. A. Begotti, L. Hortenci, F. Rocha-Mendes, C. S. S. Bernardo, R. S. Bueno, R. Nobre, R. S. Bovendorp, R. M. Marques, F. Meirelles, S. K. Gobbo, G. Beca, G. Schmaedecke and T. Siqueira

      Version of Record online: 16 OCT 2016 | DOI: 10.1111/acv.12311

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      The Brazilian Atlantic forest is considered a biodiversity hotspot and although highly fragmented, it still contains large forest patches that may be important for the conservation of mammals that require large areas. Here we estimated species richness, density and biomass of medium and large sized mammals along the largest remnant of the Atlantic rainforest in Brazil. We found that the density and biomass of mammals varied 16 and 70 fold among sites, respectively. Mammalian biomass declined by up to 98% in intensively hunted sites and was 53 fold lower than in other Neotropical non-fragmented forests. Therefore, we show that hunting is depleting the mammalian biomass in the largest Atlantic forest remnant. Photo: the muriqui (Brachyteles arachnoides) (Pedro Jordano).

    7. In aid of (re)discovered species: maximizing conservation insights from minimal data

      T. A. Brichieri-Colombi, J. M. McPherson, D. J. Sheppard and A. Moehrenschlager

      Version of Record online: 16 OCT 2016 | DOI: 10.1111/acv.12306

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      Species (re)discoveries often involve small, isolated populations that require urgent conservation interventions despite limited data. We use a case study of the recent rediscovery in Ghana of sitatunga Tragelaphus spekei, a cryptic wetland antelope, to illustrate how even sparse data can yield considerable, conservation-relevant insights. Our approach first paired observations of occurrence with landscape characteristics derived from open-access remote sensing and map data. The resulting habitat suitability model, a first for sitatunga, then served to: (1) elucidate habitat preferences; (2) assess possible existence and connectivity of remnant populations elsewhere in Ghana; and (3) estimate maximum total and effective population size. Our results demonstrate the conservation challenges associated with the rediscovery of relict populations, and the utility of applying tools such as habitat suitability models to sparse data. Moreover, our research stresses the need to implement immediate conservation action upon species (re)discoveries to prevent (regional) extinction.

    8. You have full text access to this OnlineOpen article
      Toll-like receptor variation in the bottlenecked population of the endangered Seychelles warbler

      D. L. Gilroy, C. van Oosterhout, J. Komdeur and D. S. Richardson

      Version of Record online: 13 OCT 2016 | DOI: 10.1111/acv.12307

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      In small populations, drift can result in a loss of genetic variation which can reduce adaptive evolutionary potential. However, under certain circumstances, balancing selection can counteract drift and maintain variation at functional loci. Identifying these loci can have important conservation implications and give insight into how different evolutionary forces interact in small populations. Here we investigate toll-like receptor (TLR) genes that play a key role in vertebrate innate immune defence, and whether variation has been maintained at TLR loci in a bottlenecked population of the endangered Seychelles warbler. We find depauperate levels of variation that suggests drift is the dominating evolutionary force in this island population and this may well be disadvantageous for the long-term viability of this species. Although the finding of variation of specific TLR loci, in spite of the recent bottleneck event, does warrant further study.

    9. Challenges of using behavior to monitor anthropogenic impacts on wildlife: a case study on illegal killing of African elephants

      S. Z. Goldenberg, I. Douglas-Hamilton, D. Daballen and G. Wittemyer

      Version of Record online: 13 OCT 2016 | DOI: 10.1111/acv.12309

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      Behavioral indicators are increasingly advocated as a way to monitor populations of conservation concern, but the efficacy of this approach in field settings must be investigated. In this study we use a rare combination of movement, behavior, demography, ecology, and poaching pressure from a threatened population of African elephants in northern Kenya to assess the ability of a behavioral indicator to reflect levels of human pressure. Our results suggest that behavioral indicators may not always reflect the anthropogenic impacts they are intended to measure, should be carefully tailored to systems of interest, and should incorporate non-human variables that may influence wildlife behavior.

    10. Interactions between domestic and wild carnivores around the greater Serengeti ecosystem

      M. E. Craft, F. Vial, E. Miguel, S. Cleaveland, A. Ferdinands and C. Packer

      Version of Record online: 6 OCT 2016 | DOI: 10.1111/acv.12305

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      Interactions between carnivore species have important implications for direct interference competition and cross-species transmission of shared pathogens. We asked villagers residing around a conservation area to report on the presence of wild carnivores in their village, the number of domestic dogs and cats in their household and interactions between domestic and wild carnivores. We find that wild carnivores are abundant near households surrounding the National Park, villagers would like to have more domestic dogs, interactions between wild and domestic carnivores are common and that social surveys were a useful tool for obtaining data on carnivore interactions as well as providing a direction for future targeted and in-depth research to reduce interspecific conflict. Photo credit: Andrew Ferdinands.

    11. Modelling different reintroduction strategies for the critically endangered Floreana mockingbird

      C. Bozzuto, P. E. A. Hoeck, H. C. Bagheri and L. F. Keller

      Version of Record online: 6 SEP 2016 | DOI: 10.1111/acv.12299

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      In agreement with one of the main recommendations in reintroduction biology, we used local knowledge of species to develop quantitative advice for the planned reintroduction of the critically endangered Floreana mockingbird Mimus trifasciatus in the Galápagos Archipelago. Using population viability analysis and simulations of genetic diversity, we compared six potential reintroduction strategies that differ, among other aspects, in the use of captive breeding. Although in general our findings favour a captive breeding strategy, we discuss situations where these advantages vanish.

    12. Global representation of threatened amphibians ex situ is bolstered by non-traditional institutions, but gaps remain

      A. Biega, D. A. Greenberg, A. O. Mooers, O. R. Jones and T. E. Martin

      Version of Record online: 18 AUG 2016 | DOI: 10.1111/acv.12297

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      We provide an analysis of amphibian ex-situ holdings by testing how eight ecological and biogeographical variables relating to extinction risk vary between amphibian species held in zoos and their close relatives not held in zoos. We find that species held in zoos are equally as threatened as their close relatives not found in zoos but that range and habitat-restricted species remain under-represented ex-situ. We highlight that the taxon specific conservation organization, Amphibian Ark, and non-traditional zoos are having an important effect on the representation of threatened species held ex-situ but also warn that species most at risk of short-term extinction (range-restricted specialists) require greater attention from the ex-situ conservation community.

    13. Estimates of avian collision with power lines and carcass disappearance across differing environments

      D. Costantini, M. Gustin, A. Ferrarini and G. Dell'Omo

      Version of Record online: 17 AUG 2016 | DOI: 10.1111/acv.12303

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      Our article provides for the first time raw counts and estimates of bird collisions with high-voltage power lines across seven Italian areas, and estimates of carcass disappearance that will help to improve monitoring programmes of fatal collisions of birds with the power lines.

    14. Are tourism and conservation compatible for ‘island tame’ species?

      T. Worrell, R. Admiraal, P. W. Bateman and P. A. Fleming

      Version of Record online: 14 AUG 2016 | DOI: 10.1111/acv.12301

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      Island tameness can make many island species attractive for wildlife tourism. For example, Rottnest Island quokkas allow people to approach and even touch them, which can make these animals more vulnerable to malicious attention or accidental injury. We found that quokkas at tourism sites allowed a person to approach closer before moving away compared with non-tourisms sites, and spent less time being vigilant and feeding on tourism sites. Understanding the impact of ecotourism on animal behavior will help to frame conservation management actions to ensure persistence of threatened wildlife species.

    15. Noise from a phantom road experiment alters the age structure of a community of migrating birds

      C. J. W. McClure, H. E. Ware, J. D. Carlisle and J. R. Barber

      Version of Record online: 2 AUG 2016 | DOI: 10.1111/acv.12302

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      Using a phantom road, we demonstrate experimentally that traffic noise alone can change the age structure of a community of migrating birds. We further show that the ability of both adult and hatch-year birds to gain weight during stopover is negatively affected by anthropogenic noise, although the effect is stronger for young birds. Photo credit: Ashley Jensen.

    16. Behaviour during handling predicts male natal dispersal distances in an establishing reintroduced hihi (Notiomystis cincta) population

      K. M. Richardson, J. G. Ewen, P. Brekke, L. R. Doerr, K. A. Parker and D. P. Armstrong

      Version of Record online: 25 JUL 2016 | DOI: 10.1111/acv.12296

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      Natal dispersal is a key behaviour in reintroduced populations, as it both influences the probability of recruitment, and simultaneously dictates the spatial configuration of the reintroduced population as it establishes. We examined natal dispersal distances (NDD) in a reintroduced passerine population, and found mean NDD to be significantly higher in males than in females. Males that distress-called during handling had greater NDD than those that did not. Our results show that while clusters of individuals established across the release area due to conspecific attraction, there is still movement between these clusters, primarily a consequence of natal dispersal of males of a distinct temperament.

    17. Adaptive space use by baboons (Papio ursinus) in response to management interventions in a human-changed landscape

      G. Fehlmann, M. J. O'Riain, C. Kerr-Smith and A. J. King

      Version of Record online: 12 JUL 2016 | DOI: 10.1111/acv.12293

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      In the Cape Peninsula, South Africa, chacma baboons Papio ursinus are habitual raiders of urban areas, foraging on a variety of human-derived foods. To mitigate this conflict, Cape Town municipality employs field rangers with paintball markers that ‘herd’ baboons away from the urban edge. We find that baboons still use space close to the urban edge, but choose areas close to the refuges in forested habitat and where inter-individual variation in field ranger management strategy is highest, suggesting adaptive space use by the baboons to exploit uncertainty in risk variability.

    18. Ecotourism marketing alternative to charismatic megafauna can also support biodiversity conservation

      A. Hausmann, R. Slotow, I. Fraser and E. Di Minin

      Version of Record online: 5 JUL 2016 | DOI: 10.1111/acv.12292

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      Charismatic species are arguably the main attractor of ecotourists to protected areas, but this narrow interest leads to under-appreciation of other biodiversity as well as cultural values of protected areas. We used a choice experiment and latent class model and found that less charismatic biodiversity, landscape, biodiversity-related activities, such as camping and game drives, the sense of wilderness and accessibility, also affected tourists' preferences in South African national parks. Domestic tourists, as well as more experienced international tourists, were more likely to support initiatives that promote a broader biodiversity experience than charismatic megafauna alone and were prepared to travel longer distances to do so. Our results reveal new opportunities to promote and support biodiversity conservation at sites where only less charismatic biodiversity is present.

    19. Joint effects of rising temperature and the presence of introduced predatory fish on montane amphibian populations

      N. Polo-Cavia, L. Boyero, B. Martín-Beyer, L. A. Barmuta and J. Bosch

      Version of Record online: 28 JUN 2016 | DOI: 10.1111/acv.12294

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      Warmer conditions might exacerbate the impact of exotic predators in montane amphibian populations. A joint effect of increasing temperatures and the presence of fish predators was observed on activity and development of larval amphibian, although predator recognition was not precluded at rising temperatures.

    20. Landscape connectivity losses due to sea level rise and land use change

      P. B. Leonard, R. W. Sutherland, R. F. Baldwin, D. A. Fedak, R. G. Carnes and A. P. Montgomery

      Version of Record online: 19 JUN 2016 | DOI: 10.1111/acv.12289

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      Landscape connectivity will be reduced by increasing land use and climate change, especially where these two phenomena interact. We used multiple species with various life-history requirements to indicate changes to landscape connectivity in the year 2100. We found a future landscape structure that was more homogenized, and thus, lead to broad connectivity decreases for each species and for all of them collectively. However, our results also highlight the spatial variability of these decreases and elucidate focus areas for conservation that may preserve connectivity. (Image credit: P.B. Leonard and E.B. Duffy.)

    21. Testing the demographic effects of divergent immigrants on small populations of Trinidadian guppies

      J. A. Kronenberger, W. C. Funk, J. W. Smith, S. W. Fitzpatrick, L. M. Angeloni, E. D. Broder and E. W. Ruell

      Version of Record online: 14 JUN 2016 | DOI: 10.1111/acv.12286

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      Augmenting threatened populations with individuals from elsewhere is a contentious conservation strategy, in part because immigrants can diminish overall fitness of the target population. This risk is further elevated if immigrants are divergent from recipients. In this study, we demonstrate via a controlled laboratory experiment with Trinidadian guppies that divergent immigrants can have positive demographic effects on small populations.

  4. Letter From the Conservation Front Line

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  5. Original Articles

    1. Colour pattern variation can inform about extinction risk in moths

      P.-E. Betzholtz, M. Franzén and A. Forsman

      Version of Record online: 2 JUN 2016 | DOI: 10.1111/acv.12287

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      In this study of noctuid moths we demonstrate, for the first time for invertebrates, that the degree of among-individual colour pattern variation can inform about extinction risk. Our findings have important direct implications for nature conservation, because insects constitute the vast majority of terrestrial organisms, provide many ecosystem services, are of high economic interest in agriculture and forestry (as pollinators, pests, and invasives), and comprise several endangered species.

    2. Seasonality and human disturbance alter brown bear activity patterns: implications for circumpolar carnivore conservation?

      A. Ordiz, S. Sæbø, J. Kindberg, J. E. Swenson and O.-G. Støen

      Version of Record online: 25 MAY 2016 | DOI: 10.1111/acv.12284

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      Wildlife may adapt activity patterns to daily and seasonal variations in environmental factors and human activity, and diurnal or nocturnal activity can be a response to variations in food availability and/or human avoidance. We analyzed the movement patterns of 133 GPS-collared brown bears in Sweden in spring, when bears prey on the calves of domestic reindeer and moose, and in summer–early fall, when bears rely mostly on berries, in three areas with a gradient of human disturbance. Flexibility in daily movement patterns by large carnivores may help them survive in human-dominated landscapes, but behavioral changes may also reflect environmental degradation, for example human disturbance and diurnal human activity influencing foraging opportunities for carnivores. However, brown bear depredation on reindeer occurs mostly at night, and therefore, carnivores and reindeer should be separated spatially to reduce depredation.

    3. Impacts of grassland management on wader nest predation rates in adjacent nature reserves

      S. G. Leigh, J. Smart and J. A. Gill

      Version of Record online: 24 MAY 2016 | DOI: 10.1111/acv.12283

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      Deploying agri-environment schemes (AES) adjacent to reserves in order to reduce the extent of negative impacts on the reserve has been proposed as a potential management strategy, but its efficacy has yet to be assessed. This study examines how the targeted deployment of AES adjacent to a wet grassland reserve affects wader nest predation rates upon the reserve and found that areas of the reserve adjacent to commercial land suffer lower predation rates than areas of the reserve adjacent to AES land. This suggests that predators concentrate activity away from commercial farmland and that the consequences of this strategy will depend on the extent to which it attracts breeding waders and the strength of any consequent density-dependent effects on predation rates.

    4. National assessment of threatened species using sparse data: IUCN Red List classification of Anatidae in Iran

      E. Nourani, M. Kaboli, M. Farhoodinia and B. Collen

      Version of Record online: 24 MAY 2016 | DOI: 10.1111/acv.12282

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      Using opportunistically collected data for threat assessment of Anatidae in Iran showed that effective use of the IUCN categories and criteria at the national level is hampered in situations where monitoring schemes have a short history. We suggest that in many cases opportunistically collected data need to be complemented with some level of standardized data collection or modeling in order for assessors to reliably determine the threat status of species at the national level.

    5. All roads lead to Iran: Predicting landscape connectivity of the last stronghold for the critically endangered Asiatic cheetah

      E. M. Moqanaki and S. A. Cushman

      Version of Record online: 18 MAY 2016 | DOI: 10.1111/acv.12281

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      With a worldwide population of ≤70 individuals, the critically endangered Asiatic cheetah Acinonyx jubatus venaticus only persists in several fragmented nuclei in Iran. We predicted alternative models for the landscape connectivity of Asiatic cheetahs, considering the combination of relative landscape resistance and different dispersal ability scenarios. We propose that conservation of the Asiatic cheetah urgently requires integrated landscape-level management to reduce mortality risk, and to protect core areas and corridors. (Photo credit: Iranian Cheetah Society/Iran DoE/CACP)

    6. Divergent forecasts for two salt marsh specialists in response to sea level rise

      E. A. Hunter, N. P. Nibbelink and R. J. Cooper

      Version of Record online: 26 APR 2016 | DOI: 10.1111/acv.12280

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      We forecasted the effects of sea level rise (SLR) on two salt marsh specialist bird species: clapper rails Rallus crepitans and seaside sparrows Ammodramus maritimus. Although both of these species are salt marsh obligates, we found striking differences in their habitat preferences, which will lead to divergent responses to SLR. Clapper rails will likely benefit from SLR in the short term; however, seaside sparrows will experience dramatic declines in their preferred habitats of high elevation, high salinity marshes far from forests.

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