Use of monitoring data and population viability analysis to inform reintroduction decisions: Peregrine falcons in the Midwestern United States

The peregrine falcon (Falco peregrinus anatum) has not yet recolonized natural cliff sites in Illinois and much of the lower Midwestern US, and remains restricted to urban areas. We constructed a landscape-linked population viability analysis using RAMAS/GIS software to compare possible reintroduction strategies for the species. Habitat-specific (i.e. cliff and urban) demographic parameters such as survival, fecundity, and dispersal rates were derived from the Midwest Peregrine Society Database for peregrines in the central Mississippi River region during 1982-2006. We simulated a base scenario of no reintroduction and 18 models of reintroduction with varying cohort sizes, supplementation schedules, and number of reintroduction sites, and used the Lake Superior population to test our model. Our analysis indicated that even without reintroductions in Illinois, the peregrine population in the lower Midwestern region is slowly increasing and is not likely to go extinct. Recolonization of cliff sites in southern Illinois likely will occur via dispersal from urban populations, however further research on dispersal rates between urban areas and cliffs is needed. Analysis indicated that the most cost-effective reintroduction strategy would be priced at approximately $280,000 and would result in only two additional breeding pairs compared to the no-action scenario. Thus, funds would be more effectively used in other management efforts such as habitat preservation. This study provides an example of how post-release monitoring can be used to inform future reintroduction plans.     

Relative fitness of wild and captive-reared piping plovers: Does egg salvage contribute to recovery of the endangered Great Lakes population?

Since 1992, efforts to recover the US federally endangered Great Lakes piping plover (Charadrius melodus) have included population supplementation with captive-reared young raised from abandoned eggs. Using banding data collected 1993-2008 and Cormack-Jolly-Seber models in program MARK we estimated resighting rates (ρ) and apparent survival (Φ) of first-year (1) and after-first-year (2+) plovers. Reproductive success was measured by estimating mean number of eggs laid, chicks hatched and chicks fledged per wild-reared or captive-reared parent, and these values were compared with a permutation test. The best-supported mark-resighting model indicated wild-reared plovers had higher Φ₁ and Φ₂₊ than captive-reared plovers. Breeding propensity influenced detection of wild plovers, whereas unique band combinations facilitated detection of yearling captive-reared plovers. Captive-reared (n = 10) and wild-reared (n = 57) plovers laid similar numbers of eggs, but wild-reared plovers hatched 36% more chicks and fledged 56% more young. Reproductive values derived from matrix models suggest captive-reared piping plovers are less fit than similarly aged wild-reared birds upon release and demonstrate reduced fitness in subsequent years. The Great Lakes captive-rearing effort has successfully produced a minimum of 10 breeding adults from 192 eggs that otherwise would have had no reproductive value; these captive-reared individuals now constitute up to 3% of the total population. Small scale salvage and captive-rearing of abandoned eggs should be considered as a technique to supplement existing recovery efforts in highly imperiled populations.     

The Jolly-seber model: More than just abundance

The Jolly-Seber model provides estimates of abundance, survival, and capture rates from capture-recapture experiments. This article will describe recent extensions to the following cases: (a) multiple-cohort studies where recruitment rates are compared among cohorts, (b) age-specific breeding proportions, and (c) population growth rates. Finally, new areas of research needed for this model are proposed.     

Long-term influence of mowing on population dynamics in the rare orchid Dactylorhiza lapponica: The importance of recruitment and seed production

Many orchids are currently red-listed due to changes in land use, and their future persistence will depend on management. Traditional land use like mowing is believed to favour orchids through increased survival and reproduction of established individuals, but the lack of data connecting their complex life cycle presently limits our ability to evaluate effects of management. Here we used data from 16 years of demographic monitoring to study how mowing affects population dynamics in two populations of the rare orchid Dactylorhiza lapponica. Both populations were characterized by long-lived individuals, low adult mortality and high seedling mortality. The traditional regime of mowing every second year strongly increased recruitment and reduced seed production in both populations, but had moderate effect on adult survival, growth rate and flowering probability. Population growth rate was positive for all population × treatment combinations. Traditional mowing significantly increased growth rate in both populations, and LTRE-analyses revealed that this primarily was a result of increased recruitment. The results indicate that demographic rates commonly associated with orchid persistence may be insensitive to traditional management, and underscores the importance of seed production and recruitment to maintain population growth in this long-lived species. The combination of low establishment success and no seed bank makes D. lapponica dependent on high seed input. Our results suggest that D. lapponica would benefit most from traditional mowing performed after seed dispersal in the study areas, but also suggest a high probability of future survival in the absence of mowing.     

Impact of West Nile virus and other mortality factors on American white pelicans at breeding colonies in the northern plains of North America

American white pelicans (Pelecanus erythrorhynchos) are colonial-nesting birds and their breeding sites are concentrated in a few small areas, making this species especially vulnerable to factors that can influence productivity, such as disease, disturbance, predation, weather events and loss of nesting habitat. Nearly half of the American white pelican population breeds at four colonies in the northern plains: Chase Lake National Wildlife Refuge (NWR) in North Dakota, Bitter Lake (Waubay NWR) in South Dakota, Medicine Lake NWR in Montana, and Marsh Lake in Minnesota. Thus, sustained productivity at these colonies is crucial to the health of the entire species. During the latter half of the 2002 and 2003 breeding seasons, unusually high mortality of pelican chicks was observed at these colonies. West Nile virus (WNv) was identified as one source of these losses. In 2004–2007 we monitored three major colonies in the northern plains to assess mortality of chicks during the late breeding season. We documented severe weather events, disturbance, and WNv as factors contributing to chick mortality. Before WNv arrived in the region in 2002, chick mortality after mid-July was 4%, and then jumped to as high as 44% in the years since WNv arrived. WNv kills older chicks that are no longer vulnerable to other common mortality factors (e.g., severe weather, gull predation) and typically would have survived to fledge; thus WNv appears to be an additive mortality factor. Persistence of lower productivity at American white pelican colonies in the northern plains might reduce the adult breeding population of this species in the region.     

Dynamics of a declining amphibian metapopulation: Survival, dispersal and the impact of climate

Climate can interact with population dynamics in complex ways. In this study we describe how climatic factors influenced the dynamics of an amphibian metapopulation over 12 years through interactions with survival, recruitment and dispersal. Low annual survival of great crested newts (Triturus cristatus) was related to mild winters and heavy rainfall, which impacted the metapopulation at the regional level. Consequently, survival varied between years but not between subpopulations. Despite this regional effect, the four subpopulations were largely asynchronous in their dynamics. Three out of the four subpopulations suffered reproductive failure in most years, and recruitment to the metapopulation relied on one source. Variation in recruitment and juvenile dispersal was therefore probably driving asynchrony in population dynamics. At least one subpopulation went extinct over the 12 year period. These trends are consistent with simulations of the system, which predicted that two subpopulations had an extinction risk of >50% if adult survival fell below 30% in combination with low juvenile survival. Intermittent recruitment may therefore only result in population persistence if compensated for by relatively high adult survival. Mild winters may consequently reduce the viability of amphibian metapopulations. In the face of climate change, conservation actions may be needed at the local scale to compensate for reduced adult survival. These would need to include management to enhance recruitment, connectivity and dispersal.     

The importance of chemical communication studies to mammalian conservation biology: A review

The relevance of chemical communication to mammalian conservation is not often the focus of scientific investigation. Our review identifies and discusses ten key areas in which the study of chemical communication aids conservation behaviour. Articles (n = 140) were revealed, most were concerned with population monitoring (22.50%), reducing human-wildlife conflicts (18.93), influencing habitat selection (18.57%), increasing welfare of captive animals (12.86%), encouraging captive breeding (12.86%), reducing predation (5.71%), and increasing the success of release programmes (5.00%). Few articles (<4%) were found relating olfactory studies to health status of wild populations, reducing hybridization or as indication of pollution. A growing number of articles are addressing how olfactory studies may aid conservation, but more rigorous experimental testing and manipulations are required. The vast majority of studies linking olfaction with conservation involved the population monitoring of wild carnivores. We suggest that animal behavioural studies and manipulations of chemical communication can have significant impacts on conservation in these areas, which should be further developed to generate practical applications. Areas of future study include chemical communication of aquatic mammalian species, the transfer of olfactory cues under water, and the identification of genetic markers that may link ‘personality’ with olfactory responses. Linking olfactory studies to fitness, either on an individual or population scale, particularly in a wider ecological context is more likely to increase conservation value. Animal translocations and reintroduction programmes may offer a means to do this and could be an important area to direct future studies.     

Vaccination strategies to conserve the endangered African wild dog (Lycaon pictus)

Pathogens such as rabies virus and canine distemper virus present a significant risk to the long-term survival of some endangered African wild dog (Lycaon pictus) populations. Vaccination of wild dogs is one approach that might reduce population extinction risks; however clear guidelines for how best to do this are lacking. Hence, we developed a mathematical model of wild dog population dynamics that integrates a pathogen transmission model and a vaccination process. Simulations indicated that the most effective vaccination strategy includes all age classes (when pups are 2-months old), prioritizes the breeding pair, and vaccinates at least 4 animals per pack included in the vaccination effort. In addition, populations for which an Allee effect, high rates of pathogen introduction, or low rates of recovery and immunity were simulated, required greater vaccination coverage (dogs/pack), to protect an equivalent number of packs compared to populations for which no Allee effect, low rates of pathogen introduction, or high rates of recovery and immunity were simulated. For populations in which some packs (high-risk) have a greater risk of pathogen exposure than others (low-risk), vaccinating both high- and low-risk packs, or only low-risk packs, is more effective than vaccinating only high-risk packs when pathogen introduction rates are high. Finally, model results suggest that vaccination of wild dog populations against pathogens, such as canine distemper virus, that do not cause 100% mortality and against which some wild dogs develop acquired immunity, may be unnecessary.     

Survival, dispersal and early migration movements of captive-bred juvenile eastern loggerhead shrikes (Lanius ludovicianus migrans)

Avian captive breeding programs pose a particular challenge with migratory birds due to natal dispersal and high mortality during migration. In Canada a captive breeding program for the eastern loggerhead shrike (Lanius ludovicianus migrans) has released over 400 juveniles since 2001, but the fate of almost all these birds is unknown. In 2007 and 2008, we used radio-telemetry of captive-reared juveniles to determine pre-migration survival and dispersal movements away from the release site. Overall, 76% (29 of 38) of shrikes survived from release to the initiation of migration and the daily survival rate was 0.987. Most deaths (78%) occurred in females, suggesting a possible sex-biased pre-migration mortality. Shrikes typically dispersed independently, there was little overlap between dispersal sites, and the average pre-migration dispersal distance from the release site was 4.2 km (SD 2.4). Release date was negatively correlated with time spent at the release site prior to dispersal, but did not have a significant effect on survival, time spent at dispersal sites or distance of dispersal. Migration initiation date ranged over 6 weeks and early-hatched juveniles were the first to begin migration. Using aerial telemetry, we located five of 29 birds that had begun migration at distances up to 180 km from the release site; no birds were found dead after initiating migration despite the extensive search area. High juvenile survival to migration for captive-reared juveniles suggests that the captive breeding and release program has high potential to augment wild populations of the eastern loggerhead shrike.     

Population trends in northern spotted owls: Associations with climate in the Pacific Northwest

We used reverse time capture-mark-recapture models to describe associations between rate of population change (λ) and climate for northern spotted owls (Strix occidentalis caurina) at six long-term study areas in Washington and Oregon, USA. Populations in three of six areas showed strong evidence of declining populations, while populations in two additional areas were likely declining as well. At four areas, λ was positively associated with wetter-than-normal conditions during the growing season, which likely affects prey availability. Lambda was also negatively associated with cold, wet winters and nesting seasons, and the number of hot summer days. The amount of annual variation in λ accounted for by climate varied across study areas (3-85%). Rate of population change was more sensitive to adult survival than to recruitment; however, there was considerable variation among years and across study areas for all demographic rates. While annual survival was more closely related to regional climate conditions, recruitment was often associated with local weather. In addition to climate, declines in recruitment at four of six areas were associated with increased presence of barred owls. Climate change models predict warmer, wetter winters and hotter, drier summers for the Pacific Northwest in the first half of the 21st century. Our results indicate that these conditions have the potential to negatively affect annual survival, recruitment, and consequently population growth rates for northern spotted owls.     

Successful reintroductions of the endangered long-lived Sargent’s cherry palm, Pseudophoenix sargentii, in the Florida Keys

The 1991-1994 reintroductions of Florida endangered Pseudophoenix sargentii to 13 Florida Keys sites represent a rare example of a successful multi-agency long-term effort to conserve a long-lived palm. To assess reintroduction success, we compared population demographics with and without reintroduced plants and conducted population viability analyses. Since 1991, the wild population has increased 6.4-fold. Survival from 2000-2004 was 94%, growth was positive (λ = 1.013), and there was no predicted extinction risk. Recent wild population growth is attributed to good seedling recruitment and removing the greatest threats. After 14 years, reintroductions had 43% survival, increased total plants in the wild by 27%, and expanded the species’ distribution. Reintroduced plants had faster maturation rates, improved population age structure, and enhanced population growth (λ = 1.032). Success varied with transplant year, location, microsite, and original transplant size. Failures in 1991 and at some historic sites emphasize the need for a multi-year, multi-site approach to reintroductions to buffer against stochastic losses. Rockland hammocks and the tops of coastal berms had greatest plant growth and survival. Large transplants had the greatest survival. Because no reintroduced plants are reproductive, transitions between stages are extremely slow, and plants may require >30 years to mature, continued institutional dedication to long-term monitoring will be required to assess whether the populations are self-sustaining. Horticultural expertise and ex situ collections complimented support of land managing agencies for the species’ preservation. These first rare plant reintroductions to Florida State Parks opened avenues for more plant conservation efforts and public interpretation.     

Linking nest histories,remotely sensed land use data and wildlife crime records to explore the impact of grouse moor management on peregrine falcon populations

Wildlife crime can be difficult to quantify, and its true impact on populations can be underestimated if rates are under-recorded. The illegal killing of birds of prey is an important form of wildlife crime, which in the UK, is often associated with land managed for the recreational shooting of red grouse Lagopus lagopus scoticus. In the UK, increases in peregrine falcons Falco peregrinus following recovery from organo-chlorine pesticides have not been uniform, with slow growth and localised declines in some areas, including those managed for red grouse shooting. In this study, we combined 1081 peregrine nest histories across northern England between 1980 and 2006 with a remotely sensed map of grouse moor management, to test whether breeding performance was lower in areas with active management for grouse shooting. Productivity of pairs on grouse moors was 50% lower than pairs breeding on non-grouse moor habitat. However, clutch size and brood size of successful nests did not differ between habitat types, suggesting that food constraints were unlikely to explain this difference. Population models suggested source-sink dynamics, with populations on grouse moors unable to sustain themselves without immigration. Population data confirmed that growth rates were indeed lower on grouse moors than on non-grouse moor sites. Analysis of wildlife crime data confirmed that persecution of the species was more frequent on grouse moors than in other habitat types. This population will be more secure, and better able to function as a barometer of environmental health and climate change, if illegal persecution of the species ceases on areas of land managed for grouse shooting.     

Incidental and intentional catch threatens Galápagos waved albatross

As large, long-lived seabirds with delayed and slow reproduction, albatrosses have low intrinsic mortality rates and are especially vulnerable to extinction from extrinsic sources of mortality such as fishery bycatch. Leg-band recovery information for waved albatrosses revealed mortality from both incidental catch and intentional catch for human consumption. Annual adult survival in 1999-2005, estimated from capture-mark-recapture data, was lower than historical estimates. This recent increase in adult mortality probably contributed to recent and dramatic shrinkage of the breeding population; periodic matrix models confirm that population growth rate is most sensitive to changes in adult survival. Banding data and recovery information also suggest that capture by fisheries is male-biased, which should reduce fecundity in this species with obligate bi-parental care. This new documentation of bycatch, harvesting, and associated demographic consequences provides reason for serious concern about the persistence of the single breeding population of the waved albatross.     

Lessons from a failed translocation program with a seabird species: Determinants of success and conservation value

Conservation diagnosis of ex situ techniques (e.g. releasing chicks by hacking) cannot be evaluated without quantitative assessment of the fate of individuals, mainly their survival and recruitment. Here we use a long-term monitoring at a large spatial scale of a hacking programme (i.e. chick translocations) for Audouin’s gulls in an uninhabited site so as to establish a new breeding patch and reinforce the metapopulation. The success of the programme relied on the assumption that birds tend to recruit to the site where they fledge (i.e. they were philopatric). Multi-state capture-recapture models applied to a large dataset (more than 43,500 resights at 30 colonies during 1999–2010) were used to evaluate the survival of released chicks and the probability of being philopatric. Adult survival was high, but juvenile survival was lower than that of wild gulls, suggesting that there was a cost associated with hacking only during their first year of life. As expected, most released birds returned to the hacking site, but very few (including immigrants from abroad) attempted to breed here. Dispersal was inversely correlated with distance from the hacking site and positively associated with the population size of the patch (i.e. conspecific attraction). Even though most hacking procedures met the ecological conditions to succeed, results clearly showed that the program failed to establish a new breeding site. The ability of prospecting and the attraction made by conspecifics at established sites may be a constraint for the success of hacking programs, particularly for social and nomadic species.     

Wolves in the Casentinesi Forests: insights for wolf conservation in Italy from a protected area with a rich wild prey community

The Casentinesi Forests, in the northern Apennines, harbour a rich community of wild ungulates, with the wolf representing the largest predator in the area. Between 1993 and 2000, wolf pack distribution in the area was monitored and estimates of pack size were obtained by wolf-howling surveys, snow-tracking, and occasional observations. Three to five packs were detected yearly, with sizes averaging 4.2 ± 0.9 wolves (maximum 7). The overall density in the area was 4.7 wolves per 100 km² with an average distance between adjacent packs of 11.1 km. The high wolf density in the Casentinesi Forests is mostly related to abundance and size of wild prey. In this, like in other areas at low latitudes, wolf density depends mainly on the number of packs, as pack size is rather small and recruitment limited by early dispersal and high mortality. Three homesites used in several years by resident packs were discovered. Homesite fidelity and pack reproductive success were higher in fully protected rather than harvested areas. Establishing a network of protected areas with high ungulate diversity and abundance is proposed as the main factor for allowing a full recovery of the wolf population in Italy.     

野燕麦繁殖和抗逆特性及其对小麦的他感效应研究

作为一种分布广泛的农田恶性杂草,野燕麦具有很强的环境胁迫适应能力,并且在种间竞争中常处于优势地位。本试验对野燕麦种群的繁殖与扩散特性、种子与幼苗抗逆特性及其根系分泌物对小麦的异株克生作用进行了研究,以期为找寻治理野燕麦的有效措施提供参考。结果表明:(1)野燕麦具有很强的种子繁殖与扩散能力,开花结实比小麦早,种子产量大、地下储量多、扩散距离远;(2)野燕麦种子对温度的适应范围较广,在室温与变温条件下均可以正常萌发,而冷冻后置于室温环境是其萌发最佳条件,总发芽率可达93.33%;(3)野燕麦种子对盐胁迫具有一定的适应性,在浓度低于1.8%的NaCl溶液中均可正常萌发;(4)野燕麦幼苗对盐胁迫和干旱胁迫具有较强的适应性,随着NaCl浓度和PEG浓度的升高,野燕麦幼苗叶片的脯氨酸含量显著增加(P0.01),过氧化氢酶活性呈现先增加后下降的趋势;(5)野燕麦根系水浸提液对小麦幼苗株高、干重、根长及根系活力均有极显著影响(P0.01),证明野燕麦根系分泌物对小麦幼苗生长具有异株克生作用。以上结果表明,野燕麦的竞争优势主要体现在强大的繁殖扩散能力、较强的抗逆性以及异株克生效应,相关研究可为大田野燕麦防除提供一定理论参考。     

Evaluation of the extinction risk and of conservation alternatives for a very small insular population: the bearded vulture Gypaetus barbatus in Corsica

The bearded vulture Gypaetus barbatus is a large, long-lived osteophagus vulture whose abundance and breeding range have drastically declined during the last century, making it one of the most endangered European bird species. We evaluated the extinction risk of the bearded vulture population in Corsica (a small, isolated breeding population of 8-10 pairs), one of the last extant populations in Western Europe, and estimated its probability of extinction to be 0.165 over the next 50 years. A sensitivity analysis to assess the influence of uncertain demographic rates showed that it is critical to estimate precisely the values of pre-adult survival. Neither the type nor the parameters of density dependence acting on fecundity and survival rates influenced much the extinction risk of the Corsican population. We evaluated the effect of four realistic conservation actions that could be implemented on the Corsican bearded vulture population and rank them in terms of their respective decrease of the current extinction risk faced by this population. We found that the release of two juveniles every other year for 12 years and the increase of fecundity due to selective food provisioning would reduce by more than one-half the current extinction risk of Corsican bearded vulture population. In contrast, even substantial increases in the carrying capacity through large supplemental feeding produced very modest decreases in the extinction risk, thus calling into question the efficacy of one of the main pan-European conservation strategies for this species. Re-establishing a population network within the Mediterranean could be a potentially better strategy, though its efficacy depends on natal dispersal among populations that is currently unknown.     

Assessing the impact of fisheries, climate and disease on the dynamics of the Indian yellow-nosed Albatross

Many seabird populations are currently decreasing, especially albatrosses for which the primary threat is recognised to be mortality in fisheries. Introduced predators, climate change and other factors such as diseases can also have large impacts on seabirds. Here, we assessed the relative effect of three potential threats: climate, fisheries and diseases on the demography of an endangered marine predator and modelled its population dynamics to project its size under different scenarios. We based our study on a long-term monitoring of a colony of individually marked Indian yellow-nosed albatrosses at Amsterdam Island, subtropical Indian Ocean, that has declined during the past twenty years. We found no evidence for an impact of legal tuna longlining on demographic parameters. Hatching success was lower during El Niño years but survival (0.902 ± 0.011) was not affected by climatic factors. Avian cholera caused high chick mortality (0.808 ± 0.181) which in turn probably triggered the high emigration rate (0.038 ± 0.011) through dispersal of failed breeders. This colony has a high risk of extinction. However, the rest of the population at Amsterdam Island seemingly not affected to the same extent, declined but stabilised since 1998. Matrix models indicated that lowered adult survival and the very low breeding success, resulting in low recruitment, have both contributed to the decline of the yellow-nosed albatross colony until the mid-1990s, but that more recent decline was primarily caused by low fledging success. Our results highlight that potential threats such as fisheries, diseases or climate have to be considered simultaneously to disentangle their roles when assessing the conservation status of a marine predator species.     

Large scale risk-assessment of wind-farms on population viability of a globally endangered long-lived raptor

Wind-farms receive public and governmental support as an alternative energy source mitigating air pollution. However, they can have adverse effects on wildlife, particularly through collision with turbines. Research on wind-farm effects has focused on estimating mortality rates, behavioural changes or interspecific differences in vulnerability. Studies dealing with their effects on endangered or rare species populations are notably scarce. We tested the hypothesis that wind-farms increase extinction probability of long-lived species through increments in mortality rates. For this purpose, we evaluate potential consequences of wind-farms on the population dynamics of a globally endangered long-lived raptor in an area where the species maintains its greatest stronghold and wind-farms are rapidly increasing. Nearly one-third of all breeding territories of our model species are in wind-farm risk zones. Our intensive survey shows that wind-farms decrease survival rates of this species differently depending on individual breeding status. Consistent with population monitoring, population projections showed that all subpopulations and the meta-population are decreasing. However, population sizes and, therefore, time to extinction significantly decreased when wind-farm mortality was included in models. Our results represent a qualitative warning exercise showing how very low reductions in survival of territorial and non-territorial birds associated with wind-farms can strongly impact population viability of long-lived species. This highlights the need for examining long-term impacts of wind-farms rather than focusing on short-term mortality, as is often promoted by power companies and some wildlife agencies. Unlike other non-natural causes of mortality difficult to eradicate or control, wind-farm fatalities can be lowered by powering down or removing risky turbines and/or farms, and by placing them outside areas critical for endangered birds.     

Modelling the effects of persecution on the population dynamics of golden eagles in Scotland

We used observations of the age structure and breeding productivity of the Scottish population of golden eagles Aquila chrysaetos together with the classic theory of population dynamics to derive current `unmanipulated' estimates of life history parameters. We then used regional differences in age structure associated with differences in persecution intensity to derive estimates of prospective `persecution-free' life history parameters. The different parameter combinations were entered into a population model to simulate their effects on the number of occupied territories over time. Most simulations suggested that with unmanipulated demographic parameters the population should decline. The disparity between these predictions and the observed apparent stability in occupied territories was ascribed to the buffering effect of a lowering in age of breeding in areas where persecution is most intense and that more favourable parameter estimates within the estimated limits may be more realistic. The results indicated, nevertheless, that currently the population is vulnerable to decline as also suggested by the apparent lack of adults to occupy vacant territories. In the absence of the estimated 3-5% annual adult mortality through persecution, modelling suggested the population would increase. Removing estimated effects of persecution on reproductive rate and preadult survival were on their own insufficient to reverse the declines predicted from unmanipulated parameters, although the effect of persecution on preadult survival may be more severe than we estimated. In the absence of persecution we conclude that the population could expand to fill currently vacant but apparently suitable habitat and have a more secure long-term status.