Competitive Asymmetries in the Use of Supplementary Food by the Endangered Iberian Lynx (Lynx pardinus)

Background

As a conservation tool, supplementary feeding programs may be directed to specific individuals or sectors of the target population whose productivity or survival is thought to be limited by food scarcity. However, the use of supplemental food by different sex and age classes has received little attention. We studied individual variation in the access of the endangered Iberian lynx (Lynx pardinus) to supplementary food.

Methodology/Principal Findings

From 5349 pictures taken with automatic cameras placed in 25 feeding stations, we identified 28 individuals whose sex and age were known. All individuals known to live in areas subjected to supplementation regularly visited feeding stations. Food consumption was not proportional to expected variations in energy demand within sex and age classes. Food consumption by males was higher than by females, and increased with age, in agreement with a despotic distribution. Food consumption also increased with lynx body mass, and this pattern held for individuals sharing the same breeding territories. The access of inferior competitors increased with the number of feeding stations available within lynx territories.

Conclusions/Significance

All lynx exposed to food supplementation made a regular use of extra food but individuals predicted to be competitively dominant visited stations more frequently than subordinates of the same breeding territory. Our results suggest that insufficient provision of supplementary food could restrict the access of juveniles, or even adult females, to feeding stations. Limited consumption by these target individuals may compromise the efficiency of the supplementary feeding programme at the population level, in endangered species that, as the Iberian lynx, exhibit marked sexual dimorphism in body size.     

Behavioural and spatial adaptation of the Eurasian lynx to a decline in prey availability

The distribution and abundance of food resources is a major factor influencing animal populations. I studied the effect of a roe and red deer population decline on diet composition, home range size and foraging pattern in the Eurasian lynxLynx lynx (Linnaeus, 1758) in the Bia?owie?a Primeval Forest (BPF), eastern Poland. The population of cervids in BPF experienced a nearly two-fold reduction in size from 1991 through 2006 due to severe hunting pressure between 1991 and 1996. Comparison of published data on lynx diet during the high abundance of ungulates with new data obtained for the low abundance period showed that despite a significant decline in their availability, cervids (roe and red deer) continued to form the majority of the diet of lynx, with roe deer being most preferred in both periods. Home range sizes of lynx showed a tendency to increase with declining prey densities, as indicated by relative percentage increases in average yearly home range sizes amongst different sex/age groups. In response to lower availability of their main prey, lynx increased their daily straight-line movement distances by 44% and doubled the ranges covered in 5-day periods. This illustrated that, with declining prey abundance, the lynx increased their hunting efforts by either spending more time actively searching for prey or continuing foraging even after a successful hunt. Spatial analysis of the distribution of ungulates and lynx indicated that deer were evenly distributed throughout lynx ranges in BPF and spatial proximity of the predator to prey sites did not play an important role in the efficiency of hunting. Lynx may adapt to changing prey availability by increasing search effort, but this was not sufficient to prevent the negative influences of the prey decline on the lynx population. Prey depletion has an immediate effect on lynx spatial organization and, in consequence, on their density. This information has to be considered in prioritizing lynx conservation measures and management of ungulates.     

The cost of migratory prey: seasonal changes in semi‐domestic reindeer distribution influences breeding success of Eurasian lynx in northern Norway

Migratory prey is a widespread phenomenon that has implications for predator–prey interactions. By creating large temporal variation in resource availability between seasons it becomes challenging for carnivores to secure a regular year‐round supply of food. Some predators may respond by following their migratory prey, however, most predators are sedentary and experience strong seasonal variation in resource availability. Increased predation on alternative prey may dampen such seasonal resource fluctuations, but reduced reproduction rates in predators is a predicted consequence of migratory primary prey behavior that has received little empirical attention. We used data from 23 GPS collared Eurasian lynx Lynx lynx monitored during 2007–2013 in northern Norway, to examine how spatio‐temporal variation in the migratory behavior of semi‐domestic reindeer Rangifer tarandus influences lynx spatial organization and reproductive success using estimates of seasonal home range overlap and breeding success. We found that lynx of both sexes maintained seasonally stable home ranges and exhibited site fidelity across years, independent of whether they had access to reindeer throughout the year or experienced a scarcity of reindeer in winter due to migration. However, lynx without access to reindeer in winter showed a decreased probability of reproducing and a tendency for lowered kitten survival into their first winter, when compared to female lynx with reindeer available year around. This supports the hypothesis that sedentary predators experience demographic costs in systems with migratory primary prey. Changes in the migratory behavior of ungulates, including disrupted migrations, is therefore likely to have bottom–up effects on the population dynamics of sedentary predators as well as the previously documented consequences for ungulate population dynamics.     

Internal structure and patterns of contraction in the geographic range of the Iberian lynx

We use reports obtained in a field survey to characterize the internal structure, and to reconstruct the strong contraction in the geographic range of the Iberian lynx Lynx pardinus during a 35-yr period. Lynx were distributed in one large central population surrounded by smaller peripheral ones. Abundance was autocorrelated, attained high values only in a few scattered sites mostly within the central population, and increased from west to east along major mountain chains. Abundance and occupancy were positively related. The strength of range contraction was similar in both large and small populations. We were able to date two peaks of local extinction, which were accompanied by many events of population fragmentation. We also identified five areas where local extinctions aggregated in space. Lynx relative abundance appeared to be site-specific, and the probability of local extinction decreased with increasing abundance. We suggest that sublethal deterministic factors operating with similar intensity all over the range, rather than in progression from a focal point, best explain the observed trajectory of contraction. Two factors that meet such characteristics may have had an outstanding role in lynx decline. One of them is myxomatosis, an introduced viral disease that decimated rabbits Oryctolagus cuniculus , the staple prey of the lynx. The other is related to the changes in land use prompted by extensive human emigration from the countryside 40 yr ago. Other stochastic or superimposed deterministic factors may have accounted for regional aggregation of local extinctions. The identification of these factors is very important to reverse the lasting decline of this endangered felid.     

Demographic differences in diet breadth of Canada lynx during a fluctuation in prey availability

Population dynamics of specialist carnivores are closely linked to prey availability, but the extent of variability in diet breadth of individual carnivores relative to natural variability in the abundance of their primary prey is not well understood. Canada lynx (Lynx canadensis) specialize on snowshoe hares (Lepus americanus) and exhibit cyclic fluctuations in abundance that lag 1–2 years behind those of snowshoe hares. Declining hare densities spur demographic changes in lynx, but it is unclear whether a corresponding increase in diet breadth occurs: (1) broadly across a lynx population; (2) only among individuals who are able to effectively switch to alternative prey; or (3) only among individuals who cannot capture sufficient primary prey. We measured stable isotope ratios of lynx muscle tissue spanning a cyclic increase and decline in hare density (1998–2001) in Fort Providence, NT, Canada. We found that lynx cohorts responded differently to hare population change, with yearling animals having broader diets at low hare densities, while adults and dependent juveniles maintained a constant diet through the initial decline in hare density. This result was consistent irrespective of lynx sex and indicates that yearling lynx likely are forced to adopt a broader diet when primary prey densities decline. Our results imply that select cohorts of specialist carnivores can exhibit high dietary plasticity in response to changes in primary prey abundance, prompting the need to determine whether increased diet breadth in young lynx is a successful strategy for surviving through periods of snowshoe hare scarcity. In this way, cohort‐specific niche expansion could strongly affect the dynamics of organisms exhibiting population cycles.     

Habitat-related heterogeneity in breeding in a metapopulation of the Iberian lynx

Identifying attributes associated with good breeding habitat is critical for understanding animal population dynamics. However, the association between environmental heterogeneity and breeding probability has been often overlooked in habitat analyses. We evaluated habitat quality in a metapopulation of the endangered Iberian lynx Lynx pardinus by analyzing spatiotemporal patterns in breeding records. Data summarizing successful production of litters after emergence from dens over four years within 13 lynx territories were examined. We designed a set of generalized linear mixed models representing different hypotheses regarding how patterns in breeding records relate to environmental heterogeneity. Environmental heterogeneity was described by two characteristics: 1) a landscape index measured in lynx territories indicative of time‐averaged prey availability and 2) yearly variability in prey abundance not captured with this index. By including the random effect of the lynx territory we also accounted for other territory‐specific effects on reproduction. We found significant differences in yearly prey density dynamics among lynx territories. However, temporal variation in prey density contributed poorly to explaining lynx breeding. The most parsimonious model included the landscape structure as the only effect explaining breeding patterns. A multinomial‐model‐representation of the landscape hypothesis explained nearly 50% of variability in breeding records. Results pointed to the existence of a habitat quality gradient associated with particular landscape structures influencing lynx habitat selection and breeding performance. Underlying this gradient was time‐averaged prey availability. Probably as a result of long‐term fitness strategies in long‐lived territorial species, the short‐term fluctuations in prey availability had a minor influence. Our results illustrate how habitat inferences can be enhanced by incorporating the link between spatiotemporal patterns in reproduction and environmental heterogeneity.     

Large manipulative experiments reveal complex effects of food supplementation on population dynamics of Brandt's voles

Although food supplementation is well known to increase population density,there is still debate on the causative effects of food supplementation on reproduction,survival,and immigration.Large manipulative experiments,which exclude any confounding effects of dispersal and predation,are essential for clarifying the debate.In this study,we investigated the effects of food supplementation on Brandt's vole population dynamics and plant community in eight large enclosures(0.48 ha each) from2010 to 2014.Food supplementation showed significant positive effects on population density due to increases in recruitment;however,it showed a complex effect on survival of voles:positive in non-breeding seasons,but negative in breeding seasons.In addition,food supplementation increased the quality of plants(as reflected by increased crude protein content),but decreased the quantity of less preferred plants in experimental enclosures.Thus,food seems to have direct positive effects on small rodents through improvement of food supply and indirect negative effects through food-induced density-dependent effects,and may have long-term effects on rodents through altering plant community composition and abundance.     

Diet–demography relationships in a long‐lived predator: from territories to populations

Understanding the mechanisms that shape animal population dynamics is of fundamental interest in ecology, evolution and conservation biology. Food supply is an important limiting factor in most animal populations and may have demographic consequences. Optimal foraging theory predicts greater consumption of preferred prey and less diet diversity when food is abundant, which may benefit key fitness parameters such as productivity and survival. Nevertheless, the correspondence between individual resource use and demographic processes in populations of avian predators inhabiting large geographic areas remains largely unexplored, particularly in complex ecosystems such as those of the Mediterranean basin. Based on a long‐term monitoring program of the diet and demography of Bonelli's eagle Aquila fasciata in western Europe, here we test the hypothesis that a predator's diet is correlated to its breeding productivity and survival at both the territorial and population levels, and ultimately to its population growth rate. At the territorial level, we found that productivity increased with greater consumption of European rabbits Oryctolagus cuniculus, the Bonelli's eagle's preferred prey, and pigeons, an important alternative prey for this predator. The survival of territorial pairs was negatively affected by higher diet diversity, which probably reflected the inability to find sufficient high quality prey. Diet effects at the population level were similar but more noticeable than at the territorial level, i.e. a greater consumption of rabbits, together with lesser consumption of small‐to‐medium avian species (‘other birds’; non‐preferred prey), increased productivity, while greater diet diversity and lower consumption of rabbits was associated with reduced survival and population growth rate. Overall, our study illustrates how the diet of a predator species can be closely related to key individual vital rates, which, in turn, leave a measurable fingerprint on population dynamics within and among populations across large spatial scales.     

Dynamics of a small re-introduced population of wild dogs over 25 years: Allee effects and the implications of sociality for endangered species’ recovery

We analysed 25 years (1980–2004) of demographic data on a small re-introduced population of endangered African wild dogs (Lycaon pictus) in Hluhluwe-iMfolozi Park (HiP), South Africa, to describe population and pack dynamics. As small populations of cooperative breeders may be particularly prone to Allee effects, this extensive data set was used to test the prediction that, if Allee effects occur, aspects of reproductive success, individual survival and population growth should increase with pack and population size. The results suggest that behavioural aspects of wild dogs rather than ecological factors (i.e. competitors, prey and rainfall) primarily have been limiting the HiP wild dog population, particularly a low probability of finding suitable mates upon dispersal at low pack number (i.e. a mate-finding Allee effect). Wild dogs in HiP were not subject to component Allee effects at the pack level, most likely due to low interspecific competition and high prey availability. This suggests that aspects of the environment can mediate the strength of Allee effects. There was also no demographic Allee effect in the HiP wild dog population, as the population growth rate was significantly negatively related to population size, despite no apparent ecological resource limitation. Such negative density dependence at low numbers indicates that behavioural studies of the causal mechanisms potentially generating Allee effects in small populations can provide a key to understanding their dynamics. This study demonstrates how aspects of a species’ social behaviour can influence the vulnerability of small populations to extinction and illustrates the profound implications of sociality for endangered species’ recovery. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A pilot study of the performance of captive-reared delta smelt Hypomesus transpacificus in a semi-natural environment

A captive breeding programme was developed in 2008 for delta smelt Hypomesus transpacificus in reaction to dramatic population decline over several decades. We took 526 sub-adult captive-reared delta smelt and cultured them for 200 days without providing artificial food or water quality management to assess their performance once released in the wild. The results indicated captive-reared sub-adult delta smelt could survive in a semi-natural environment with uncontrolled water quality and naturally produced wild prey through spawning and into their post spawning phase.     

Food quantity–quality co‐limitation: Interactive effects of dietary carbon and essential lipid supply on population growth of a freshwater rotifer

相似文献   

Woodland caribou calf mortality in Newfoundland: insights into the role of climate,predation and population density over three decades of study

The rates and causes of juvenile mortality are central features of the dynamics and conservation of large mammals, like woodland caribou (Rangifer tarandus caribou (Gmelin, 1788)), but intrinsic and extrinsic factors may be modified by variations in animal abundance. We tested the influences of population size, climate, calf weight and sex on survival to 6 months of age of 1241 radio-collared caribou calves over three decades, spanning periods of population growth (1979–1997) and decline (2003–2012) in Newfoundland, Canada. Daily survival rates were higher and rose more quickly with calf age during the population growth period compared to the decline. Population size (negatively) and calf weight (positively) affected survival during the decline but neither had a detectable influence during the growth phase. Sex, climate and plant productivity (the latter two derived from the North Atlantic Oscillation and Normalized Difference Vegetation Index, respectively) exerted minimal influence during either phase. Predation was the dominant source of mortality. The mean percentage of calves killed by predators was 30 % higher during the decline compared to the growth phase. Black bears (Ursus americanus) and lynx (Lynx canadensis) were the major predators during the population increase but this changed during the decrease to black bears and coyotes (Canis latrans). Our findings are consistent with the hypothesis that Newfoundland caribou experienced phase-dependent survival mediated proximally by predation and competition for food.     

Predation risk perception and food scarcity induce alterations of life-cycle traits of the mosquito Culex pipiens

Abstract.  1. Predators may affect prey populations by direct consumption, and by inducing defensive reactions of prey to the predation risk. Food scarcity frequently has effects on the inducible defences of prey, but no consistent pattern of food–predation risk interaction is known.
2. In this study the combined effect of food shortage and predation-risk perception in larvae of the mosquito Culex pipiens was investigated. Water exposed to the aquatic predator bug Notonecta glauca was used as a source of predation intimidation. Mosquito larvae were reared in three different media containing either no predator cues or the cues of N. glauca that had been fed on either C. pipiens larvae or on Daphnia magna . Food was provided in favourable or limited amount for these set-ups.
3. The results showed that chemical cues from the predators fed with prey's conspecifics caused a decreased survival, delayed pre-imaginal development, and reduction in body size of emerged mosquitoes, whereas chemical cues from predators fed with D. magna caused only delayed development. Food scarcity significantly exacerbates the negative effect of the predator cues on pre-imaginal development of C. pipiens . Effects of the cues on larval development and body size of imagoes are significantly stronger for females than for males.
4. The present study suggests that when food is limited, predators can affect population dynamics of prey not only by direct predation, but also by inducing lethal and sublethal effects due to perception of risk imposed by chemical cues. To understand the effects of predators on mosquito population dynamics, environmental parameters such as food deficiency should be considered.     

Reproductive biology of captive female Eurasian lynx, Lynx lynx

Studies on wild Eurasian lynx (Lynx lynx) have revealed variation in reproduction between areas, years and individuals. In order to explore potential causes for this variation other than food supply, we analysed data from captive lynx, which provide conditions with minimal environmental variation as all were fed ad libitum. Data from 37 individual female lynx were available from 20 zoos in Norway, Sweden, Finland, Switzerland and the Czech Republic. Data on 177 reproductive events (where a male was available to the female at mating time) are presented. Of these events, 85% resulted in litters being born. Average litter size was 1.95, with a variation from 1 to 4. The mean birth date was 26th May, and sex ratio was not significantly different from parity. The probability of reproduction was related to age, with fewer litters produced by the very young (2–3-year old), and no sign of a senescence effect. However, a clear effect of senescence on litter size was evident. The captive lynx did not have higher reproductive rates than wild lynx, indicating that either factors other than food supply are driving the variation in wild lynx reproduction, or that a factor such as stress may be causing additional variation in the captive population.     

Population cycles and changes in body size of the lynx in Alaska

The lynx Lynx canadensis is a common predator in the boreal forests of North America. Its population fluctuates during a 9- to 11-year cycle in synchrony with the population size of its main prey, the snowshoe hare Lepus americanus. Using adult museum specimens, we studied changes in skull (and hence body) size of the lynx in Alaska during the second half of the 20th century. The population cycle in Alaska averaged 9 years, similar to that reported in the neighbouring Yukon. Using harvest data of lynx as an estimate of population size, we found that skull size was negatively related to population size. This relationship was strongest not for the population density in the year of death (X), but for year X-3, a carry-over effect from the first year (or years) of life, indicating that conditions during the fast-growth years are determining body size. We suggest that the density-dependent effect is probably due to changes in food supply, either resulting from the adverse effects of competition or a possible diminished availability of food. Two skull parameters decreased significantly during the second half of the 20th century. We do not know the cause for the year effect and suggest that it might be due to a long-term change in the availability of prey. Canine size did not change during the study period, probably an indication that snowshoe hares maintained their status as the main prey of the lynx throughout the study period. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The dynamics of top-down and bottom-up effects in food webs of varying prey diversity, composition, and productivity

Prey diversity is thought to mediate the strength of top-down and bottom-up effects, but few experiments directly test this hypothesis. I assembled food webs of bacteria and bacterivorous protist prey in laboratory microcosms with all combinations of five productivity levels, two top predator treatments (present or absent), and three prey compositions. Depauperate food chains contained one of two edible prey species, while more diverse food webs contained both edible prey species plus two additional less-edible/inedible prey. Equilibrium theory predicts that prey diversity should weaken the top-down and bottom-up effects on trophic level biomasses, due to density compensation among prey species. Top-down effects should increase with productivity in food chains, but decrease with productivity in food webs. Results revealed highly dynamic top-down effects, the strength of which varied more over time than among treatments. Further, top-down effects did not merely vary in absolute strength over time, but also in relative strength across different prey compositions and productivity levels. It might be expected that equilibrium models would qualitatively reproduce time-averaged results. However, time-averaged data largely failed to support equilibrium predictions. This failure may reflect strong temporal variability in treatment effects combined with nonlinear density dependence of species' per-capita growth rates. Strong temporal variability in the strength of top-down effects has not previously been demonstrated, but likely is common in nature as well.     

Impact of spatial variation of a crucial prey,the molecricket,on hoopoe territory occupancy and reproduction

Direct benefits accrued from securing a territory of sufficient quality are important determinants of individual fitness and population persistence. Food supply is one of the main factors of animal territory quality, with spatial and temporal variation in prey availability largely dictating reproductive output and thus population dynamics. In a Swiss hoopoe population, molecrickets Gryllotalpa gryllotalpa, the most profitable prey locally, can constitute most of the food biomass delivered to chicks by parents. We first investigated the impact of molecricket prey on hoopoes' fitness‐related traits by quantifying the spatial variation in the food allocation pattern of both male and female parents to chicks across the whole population range; and second, analysed the impact of this prey on current reproduction and, using a 11 yr dataset, on the temporal occupancy rate of each territory. We found considerable but spatially repeatable variation, over the years, of molecricket biomass in the diet provisioned to chicks. This spatial heterogeneity in chicks' diet composition was mirrored both in the history of territory occupancy (2002–2012) and in current reproductive success (2012). Territories with a greater biomass of molecrickets in chicks' diet produced more fledglings in better body condition. Yet, these effects on current reproduction were exclusively demonstrated for male parents, corroborating that paternal provisioning patterns play a predominant role in hoopoe reproductive success. This study demonstrates how a single, very profitable prey species might affect spatial variation in territory settlement and individual reproductive success in a regionally endangered bird species, with potential consequences for its population dynamics and persistence.     

Diet of Eurasian lynx, Lynx lynx, in the boreal forest of southeastern Norway: the relative importance of livestock and hares at low roe deer density

The year-round food habits of lynx were studied using radio-telemetry and snow-tracking in the boreal forest of southeastern Norway. The main objectives of the study were to clarify the importance of domestic sheep and small prey species in the diet of lynx in an area with a very low density of roe deer. During the period 1995–1999, we found 193 scats and 358 kills made by lynx. Our results indicate that roe deer were the most common prey species (contributing to 83 and 34% of the biomass consumed in winter and summer, respectively), although a wide range of other species were also found, including mountain hares, tetranoids, red foxes, domestic sheep, wild reindeer, and even moose. Most of the diet was obtained by predation, although we did document several cases of scavenging. Roe deer were more important in the diet in winter than in summer, perhaps because they were easier to locate in winter as they clustered around feeding sites. In summer, domestic sheep and small prey increased in importance. Despite the very low density of roe deer in this study area, lynx seemed to still specialise on them, although domestic sheep did constitute a significant amount to their diet, especially for males and yearlings. However, the contribution of sheep to summer diet was far from that expected if their relative density was considered.     

Diet of Eurasian lynx <Emphasis Type="Italic">Lynx lynx</Emphasis> in the northern Dinaric Mountains (Slovenia and Croatia)

The Eurasian lynx Lynx lynx (Linnaeus 1758) is an opportunistic predator that usually selects the smallest ungulate available. Its diet varies considerably among different regions; therefore it is important to study lynx diet in different parts of the species’ range. We studied the diet of lynx from the endangered Dinaric population in Slovenia and Croatia by analyzing lynx scats, prey remains, and stomach contents. Dinaric lynx mainly killed European roe deer Capreolus capreolus (0.64 frequency of occurrence, 79% of all consumed biomass), which were used more frequently during winter and spring. Ungulates were killed more often by adult males than by lynx of other age and sex groups. In contrast to studies from other regions, lynx in the northern Dinaric Mountains also frequently fed on the edible dormice Glis glis (0.18 frequency of occurrence, 7% of all consumed biomass). This large rodent appears to be an important alternative prey, especially for females and young lynx, and was the reason for the highest use of rodents reported so far for the Eurasian lynx. Edible dormice in Dinaric forests have highly variable numbers of active animals. Seasonal and possibly annual variation in dormouse availability obviously affects lynx diet. This is a rare example where variability in the availability of the alternative prey and not the preferred prey leads to the dietary shift. This study confirms the opportunistic nature of Eurasian lynx and the regional variability of its diet.     

Could we live with reintroduced large carnivores in the UK?

1. Literature on the wolf Canis lupus, brown bear Ursus arctos and lynx Lynx lynx is reviewed to determine if sufficient semi‐natural habitat exists in the UK for a viable population of any of these species and to assess the potential risks to human safety, livestock and economically valuable wildlife. Public attitudes to the recovery and reintroduction of some other mammals are also briefly reviewed. 2. The large home range sizes and low population densities of large carnivores mean that the Scottish Highlands is the only UK region with the potential to support a viable population. Human population density is also lower in the Highlands and the density of wild ungulate prey higher than in many parts of Europe where large carnivores survive. 3. Attacks on people have been recorded in Europe for healthy bears and for rabid bears and wolves but there are no reports of attacks by lynx. Bears are more carnivorous in the north of their range than in the south and although wild mammals seldom appear to be important prey serious predation of livestock can occur. Livestock predation is also reported for the wolf and the lynx but they appear to prefer wild prey if available. However, mass kills of up to 100 or more sheep are occasionally recorded for wolves. 4. Attitudes to reintroductions and carnivores generally tend to be favourable amongst the general public, but negative amongst those most likely to be adversely affected. Fears for human safety and significant livestock predation with bears and wolves, respectively, suggest that reintroduction of these species is unlikely to be acceptable in the foreseeable future. Reintroduction of the lynx may be feasible but habitat suitability and potential impact on vulnerable native wildlife need to be assessed. Socio‐economic and legal issues also need to be addressed before such a reintroduction is considered.