A note on the effectiveness of incorporating management objectives with ecological variables when modeling red deer abundance

Big game populations are being raised in areas characterized by different land uses and are being managed under a wide range of circumstances which might influence their population densities and structure. However, the consideration of explanatory variables related to management activities is not generally explicitly considered in game population models. This paper focuses on how estate owners’ management objectives and strategies influence red deer (Cervus elaphus L.) hunting yields. We study red deer harvest in 187 hunting estates in Andalusia (Spain) and use both ecological and management predictors to describe capture levels in each hunting estate. We have found that the main land use of the area where red deer are being raised, the type of hunting holder, and the long-term management strategies (such as fencing and the hunting practices implemented) significantly explain red deer hunting bag, which is often used as proxy of population abundance. Therefore, ecological variables alone are not always sufficient to determine big game harvests. We provide empirical support that reinforces the relevance of considering humans and their long-term decisions when trying to interpret wildlife harvests, populations, and trends. We urge scientists and technicians to incorporate human aims in the game species distribution models and resource user decision models, particularly when game populations are being intensively managed.     

Willow grouse bag size is more sensitive to variation in hunter effort than to variation in willow grouse density

Annual variation in harvested animals (hereafter bag size) is often used as an index of population abundance when investigating population dynamics. Few studies have evaluated how well bag size tracks population change despite its widespread use. Two recent studies on grouse harvest statistics have reached contrasting conclusions. Th ere is limited information about the functional response of hunters in relation to varying game densities, and effort is seldom recorded. We investigated how much of the variation in bag size (total number of harvested grouse km^?2) is explained by variation in willow grouse Lagopus lagopus density (adult and young grouse km^?2) and hunting eff ort (total number of hunting days km^?2). We also evaluated catch per unit effort (CPUE) as an index of grouse abundance, and estimated the response in harvest rate (total bag size in relation to total grouse density) to varying hunting effort. We used data from the 88 management areas on state land in Jämtland county, Sweden (1996–2007), where hunting days and bag size are recorded in detail. Willow grouse density was estimated in four of these management areas in August using line transects and distance sampling. The hunting effort and total grouse density explained most of the variation in bag size (R²= 0.89). Bag size was twice as sensitive to changes in hunting effort compared to changes in grouse density. More than a ten times change in the grouse population density was required to one unit change in bag size. The use of CPUE did not provide a better alternative index of grouse density, and variation in density only explained 23% of the variation in CPUE. Harvest rate showed a strong relationship with hunting effort, and we suggest that an upper limit in hunting effort can be used to reduce the risk of high harvest rates. Hunters became more efficient at low densities and controlling hunting effort is most important when there are indications of population lows and/or poor breeding. CPUE may be less sensitive to changes in game abundance than previously assumed, and bag size as a proxy for population density would then depend on the ability of hunters to adjust their effort according to population change. We speculate that this ability will depend on whether or not hunters have long‐term experience of a hunting area where they can return to hunt throughout the hunting season. We propose that recording hunting eff ort should be encouraged and possible correlations with game abundance and other factors such as weather should be investigated for game species.     

Estimating red deer abundance in a wide range of management situations in Mediterranean habitats

When human interventions interfere with the natural regulation of wildlife populations by favouring some species, overabundance can emerge. We evaluated different methods of estimating red deer abundance in a wide range of population densities from southern Spain. Distance sampling estimates were used as the reference method across 22 localities and were compared with two kilometric abundance indices (KAIs), four indices based on pellet group counts and two browsing indices (BWIs). The average red deer density estimated by distance sampling was 19.51±3.19 deer per 100 ha, showing a wide range across the study area (0.04–66.77). Distance sampling estimates correlated with the KAIs, pellet group-based index and the BWI. The agreement with distance sampling improved when groups were used instead of individuals in the KAIs, when the minimum pellet group size was fixed at 20 pellets in the dropping counts, and when only highly palatable species were used in the BWI. Thus, several direct and indirect methods can estimate red deer abundance in Mediterranean habitats from Southern Spain with appropriate modifications.     

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.     

Spatial and temporal habitat use and selection by red deer: The use of direct and indirect methods

Given the importance of red deer Cervus elaphus for hunting and conservation purposes, understanding the interactions between this species and its habitats in the Mediterranean region is a crucial step for the sustainable management of this species. Aiming to compare pellet group counts and direct observations methods to study the habitat use and selection by red deer, the results obtained by both methods were compared, and their advantages and disadvantages were discussed. To understand the temporal patterns of habitat use and selection, the survey was conducted at three different seasons, birth period, rut season and winter. The habitat use and selection were studied in relation to land cover, watercourse, roads, ecotone zones and other topographic features (altitude, slope and aspect), using generalized linear models and selection ratios. The similarity of the results provided by pellet group counts and direct observations indicate that both methods may constitute useful tools to study the habitat use and selection by red deer. Globally, red deer seemed to select habitats that provide simultaneously food and some cover, as shown by its preference for shrublands, independently of the sampling season. The positive selection of ecotone zones embodies the need for open spaces. Males and females showed a similar use of shrubland, but selected patches with different characteristics therein. The spatial and temporal patterns exhibited by our results suggest that red deer balance their habitat requirements in respect to each phase of their reproductive cycle. Pellet group counts and direct observations seem to be useful methods to analyze habitat use and selection, and may provide helpful knowledge to the management and conservation of red deer.     

Red deer hunting—commercializing versus availability

Many deer populations in Europe and North America have increased in abundance over the last decades. The increasing populations potentially entail both ecological and economic challenges and opportunities, but in practice we still know little about the extent to which these opportunities are being exploited in different management systems. The Norwegian red deer population has increased in density and expanded rapidly since the 1950s. Traditionally, red deer hunting has been undertaken by the local landowner and his relatives and friends. The present large population raises the question whether attracting other hunters could provide a higher economic return for the landowners and, if so, if they are interested in providing such hunting opportunities. We designed a survey to learn more about the landowners, both with respect to the present level of hunting income as well as economic costs of, for example, forest and agricultural damage; we also sought to understand their interest in increasing their income from red deer hunting and potential obstacles to realizing such an increased economic benefit. The results indicate that landowners on average think that red deer populations on their land result in higher costs than income but are nevertheless satisfied with the way things are. This highlights that increased numbers of deer need not automatically lead to more income for landowners and that the potential for income may be hindered by cultural factors such as reluctance to allow access to non-local hunters.     

Red oak seedlings as indicators of deer browse pressure: Gauging the outcome of different white‐tailed deer management approaches

After decades of high deer populations, North American forests have lost much of their previous biodiversity. Any landscape‐level recovery requires substantial reductions in deer herds, but modern societies and wildlife management agencies appear unable to devise appropriate solutions to this chronic ecological and human health crisis. We evaluated the effectiveness of fertility control and hunting in reducing deer impacts at Cornell University. We estimated spring deer populations and planted Quercus rubra seedlings to assess deer browse pressure, rodent attack, and other factors compromising seedling performance. Oak seedlings protected in cages grew well, but deer annually browsed ≥60% of unprotected seedlings. Despite female sterilization rates of >90%, the deer population remained stable. Neither sterilization nor recreational hunting reduced deer browse rates and neither appears able to achieve reductions in deer populations or their impacts. We eliminated deer sterilization and recreational hunting in a core management area in favor of allowing volunteer archers to shoot deer over bait, including at night. This resulted in a substantial reduction in the deer population and a linear decline in browse rates as a function of spring deer abundance. Public trust stewardship of North American landscapes will require a fundamental overhaul in deer management to provide for a brighter future, and oak seedlings may be a promising metric to assess success. These changes will require intense public debate and may require new approaches such as regulated commercial hunting, natural dispersal, or intentional release of important deer predators (e.g., wolves and mountain lions). Such drastic changes in deer management will be highly controversial, and at present, likely difficult to implement in North America. However, the future of our forest ecosystems and their associated biodiversity will depend on evidence to guide change in landscape management and stewardship.     

Disentangling demographic effects of red deer on chamois population dynamics

Investigating the impact of ecological factors on sex‐ and age‐specific vital rates is essential to understand animal population dynamics and detect the potential for interactions between sympatric species. We used block count data and autoregressive linear models to investigate variation in birth rate, kid survival, female survival, and male survival in a population of Alpine chamois Rupicapra rupicapra rupicapra monitored over 27 years within the Stelvio National Park, Central Italian Alps, as function of climatic variables, density dependence, and interspecific competition with red deer Cervus elaphus. We also used path analysis to assess the indirect effect of deer abundance on chamois growth rate mediated by each demographic parameter. Based on previous findings, we predicted that birth rate at [t] would negatively relate to red deer abundance at year [t − 1]; survival rates between [t] and [t + 1] would negatively relate to red deer abundance at year [t − 1] and to the interactive effect of winter precipitation at [t + 1] and chamois density at [t]. Our results showed that birth rate was positively related to spring–summer precipitation in the previous year, but this effect was hampered by increasing red deer abundance. Kid and female survival rates were negatively related to the combined effect of chamois abundance and winter precipitation. Male and female survival rates were negatively related to lagged red deer abundance. The path analysis supported a negative indirect effect of red deer abundance on chamois growth rate mediated by birth rate and female survival. Our results suggest that chamois population dynamics was largely explained by the synergistic effect of density dependence and winter harshness, as well as by interspecific competition with red deer, whose effects were seemingly stronger on the kid–female segment of the population.     

Differential Responses to Hunting in Two Sympatric Species of Brocket Deer (Mazama americana and M. nana)

Hunting by humans may affect the abundance and activity patterns of game species. We examined the effect of hunting on the abundance and activity patterns of sympatric red brocket deer Mazama americana and dwarf brocket deer M. nana . We conducted four camera-trap surveys (158 sampling stations, 10,244 trap-days, total area sampled 1200 km²) in three areas within the Atlantic Forest of Misiones, Argentina, that differ in protection and hunting pressure. We used logistic regression and tests of independence to evaluate if protection, hunting pressure, and other independent variables affect the probability of recording each species and their recording rate. We used the Mardia–Watson–Wheeler test to examine if the daily activity pattern differs between species and changes with hunting pressure. Red brocket deer were more frequently recorded (397 records, 58% of stations) than dwarf brocket deer (100 records, 37% of stations). The probability of recording red brockets was higher in areas with better protection and increased with the distance to the main accesses used by poachers. The probability of recording dwarf brockets was higher in areas with low protection. Red brockets were more nocturnal than dwarf brockets, a difference that may reduce interspecific competition. However, red brockets were more diurnal in the best-protected areas, suggesting that they can adjust their activity to local hunting pressure. Hunting has opposite effects on the abundance of these deer and may facilitate their coexistence. Hunting should be carefully controlled or managed to ensure the conservation of these little known species.     

Predation has a greater impact in less productive environments: variation in roe deer, Capreolus capreolus, population density across Europe

Aim We aimed to describe the large‐scale patterns in population density of roe deer Caprelous capreolus in Europe and to determine the factors shaping variation in their abundance. Location Europe. Methods We collated data on roe deer population density from 72 localities spanning 25° latitude and 48° longitude and analysed them in relation to a range of environmental factors: vegetation productivity (approximated by the fraction of photosynthetically active radiation) and forest cover as proxies for food supply, winter severity, summer drought and presence or absence of large predators (wolf, Canis lupus, and Eurasian lynx, Lynx lynx), hunter harvest and a competitor (red deer, Cervus elaphus). Results Roe deer abundance increased with the overall productivity of vegetation cover and with lower forest cover (sparser forest cover means that a higher proportion of overall plant productivity is allocated to ground vegetation and thus is available to roe deer). The effect of large predators was relatively weak in highly productive environments and in regions with mild climate, but increased markedly in regions with low vegetation productivity and harsh winters. Other potentially limiting factors (hunting, summer drought and competition with red deer) had no significant impact on roe deer abundance. Main conclusions The analyses revealed the combined effect of bottom‐up and top‐down control on roe deer: on a biogeographical scale, population abundance of roe deer has been shaped by food‐related factors and large predators, with additive effects of the two species of predators. The results have implications for management of roe deer populations in Europe. First, an increase in roe deer abundance can be expected as environmental productivity increases due to climate change. Secondly, recovery plans for large carnivores should take environmental productivity and winter severity into account when predicting their impact on prey.     

Adaptive management of sika deer populations in Hokkaido, Japan: theory and practice

We investigated the utility of adaptive management (AM) in wildlife management, reviewing our experiences in applying AM to overabundant sika deer (Cervus nippon) populations in Hokkaido, Japan. The management goals of our program were: (1) to maintain the population at moderate density levels preventing population irruption, (2) to reduce damage to crops and forests, and (3) to sustain a moderate yield of hunting without endangering the population. Because of significant uncertainty in biological and environmental parameters, we designed a “feedback” management program based on controlling hunting pressure. Three threshold levels of relative population size and four levels of hunting pressure were configured, with a choice of four corresponding management actions. Under this program, the Hokkaido Government has been promoting aggressive female culling to reduce the sika deer population since 1998. We devised a harvest-based estimation for population size using relative population size and the number of deer harvested, and found that the 1993 population size (originally estimated by extrapolation of aerial surveys) had been underestimated. To reduce observation errors, a harvest-based Bayesian estimation was developed and the 1993 population estimate was again revised. Analyses of population trends and harvest data demonstrate that hunting is an important large-scale experiment to obtain reliable estimation of population size. A serious side effect of hunting on sika deer was inadvertent lead poisoning of large birds of prey. The prohibition of the use of lead bullets by the Hokkaido Government was successful in reducing the lead poisoning, but the problem still remains. Two case studies on sika population irruption show that the densities set by maximum sustainable yield may be too high to prevent damage to agriculture, forestry, and/or ecosystems. Threshold management based on feedback control is better for ecosystem management. Since volunteer hunters favor higher hunting efficiency in resource management (e.g., venison), it is necessary to support the development of professional hunters for culling operations for ecosystem management, where lower densities of deer should be set for target areas. Hunting as resource management and culling for ecosystem management should be synergistically combined under AM.     

An Integrated Population Model for Harvest Management of Atlantic Brant

Atlantic brant (Branta bernicla hrota) are important game birds in the Atlantic Flyway and several long-term monitoring data sets could assist with harvest management, including a count-based survey and demographic data. Considering their relative strengths and weaknesses, integrated analysis to these data would likely improve harvest management, but tools for integration have not yet been developed. Managers currently use an aerial count survey on the wintering grounds, the mid-winter survey, to set harvest regulations. We developed an integrated population model (IPM) for Atlantic brant that uses multiple data sources to simultaneously estimate population abundance, survival, and productivity. The IPM abundance estimates for data from 1975–2018 were less variable than annual mid-winter survey counts or Lincoln estimates, presumably reflecting better accounting for observer error and incorporation of demographic estimates by the IPM. Posterior estimates of adult survival were high (0.77–0.87), and harvest rates of adults and juveniles were positively correlated with more liberal hunting regulations (i.e., hunting days and the daily bag limit). Productivity was variable, with the percent of juveniles in the winter population ranging from 1% to >40%. We found no evidence for environmental relationships with productivity. Using IPM-predicted population abundances rather than mid-winter survey counts alone would have meant fewer annual changes to hunting regulations since 2004. Use of the IPM could improve harvest management for Atlantic brant by providing the ability to predict abundance before annual hunting regulations are set, and by providing more stable hunting regulations, with fewer annual changes. © 2021 The Wildlife Society.     

There is only one winner: The negative impact of red deer density on roe deer numbers and distribution in the Słowiński National Park and its vicinity

Red and roe deer are the most numerous cervids in Europe, and they occur in sympatry in most regions. Roe deer were considered to be an inferior competitor in studies in which they co‐occurred with fallow deer or muntjac. Despite the remarkable overlap of their ranges, there are few studies on the competition between the red and roe deer. Since interspecific interactions among ungulates are often related to their mutual densities, the current study focused on the effects of high red deer density on the roe deer numbers and spatial distribution in the unhunted Słowiński National Park (SNP) in northern Poland and forest districts open to hunting bordering the park. Using fecal pellet group counts, it was found that in the forest districts (where red deer densities were 2–3 times lower than in the SNP), roe deer densities were significantly higher than in the park. The red‐to‐roe deer density ratio was 10.8 and 2.7, in the SNP and the surrounding forest districts, respectively. Moreover, in the SNP, the roe deer distribution was negatively affected by the red deer habitat use, while in the hunting areas, such an effect was not recorded. The negative influence of the red deer on the roe deer population in the park was most probably due to the red deer impact on food availability. The biomass of the plant groups forming the staple food of the roe deer (Rubus spp., forbs, dwarf shrubs) was significantly higher in the fenced plots than in the unfenced ones. Lack of hunting in the protected areas may benefit only some species in ungulate assemblages which, in turn, may contradict one of their objectives—to maintain viable and ecologically functional populations.     

Seasonal variation in the sex and age composition of the woodcock bag in Denmark

The Eurasian woodcock is a highly valued game bird in Western Europe from which c. 2.7 million individuals are harvested annually from an estimated population of 20–26 million birds. The population size and status remains uncertain due to the cryptic behaviour and widespread and solitary occurrence of woodcock, on breeding and wintering areas, making reliable population surveys difficult. Hunting bag records provide age ratios amongst bagged birds, but sex ratios remain poorly known because of the sexually monomorphic nature of this species. We used DNA analysis to determine sex ratios amongst 327 shot woodcocks from two hunting seasons in Denmark (1 October–31 January, 2012/13 and 2013/14). Based on bag totals, age ratios and sex ratios, juvenile females constituted 37%, juvenile males 27%, adult females 16% and adult males 20% of the annual woodcock bag. The female bias was related to a significant deviation from parity in the sex ratio amongst juvenile birds in October, although no such deviation was found at other times or amongst adults. Compared to limited data from other European countries, our data suggest that autumn migration of woodcock involves an initial wave of juvenile females followed by juvenile males and adults, and perhaps that males stay further north in Europe than females during autumn and winter. This migratory pattern would suggest that postponing the opening of the hunting season could reduce the hunting bag on reproductively valuable females in this polygamous species.     

Differential long-term impacts of a management control program of axis deer and wild boar in a protected area of north-eastern Argentina

Exotic ungulates are among the top global invasive mammals and a threat to biodiversity. Axis deer (Axis axis) and wild boar (Sus scrofa) are of increasing concern in multiple regions. A management program reduced wild boar abundance and soil damage below target levels through controlled still shooting from watchtowers and dog-hunting performed by recreational hunters at El Palmar National Park, Argentina. Here we assess program impacts on axis deer over a 10-year period in which 2380 deer were dispatched, and document two largely unexpected outcomes: increasing axis deer abundance toward a plateau, and a strong inverse correlation between deer and wild boar numbers. Unlike the initial steep decline and subsequent stabilization of wild boar, deer abundance indexed by standardized catch-per-unit-effort increased at 37.6% per year over 0–5 years post-intervention (YPI) and stabilized from 7 YPI on when still-shooting effort averaged 948 hunting party-hours per quarter. Deer catch was non-linearly related to still-shooting effort. Timing of deer and boar catches did not differ significantly regardless of sex, season and YPI. Catch-per-unit-effort indices and nightly spotlight deer counts showed similarly increasing trends. The fraction of older adult deer declined over 0–4 YPI and remained stable thereafter. Sex ratios were consistently skewed toward males only among older adults. Failure to reduce deer abundance may be explained by several major processes: protracted exponential growth of the deer population after park invasion; deer regional expansion with increasing immigration; insufficient sex- and stage-biased hunting mortality, and competitor (and perhaps predator) release from wild boar.     

Seasonal migration of male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in southern Sweden and consequences for management

Red deer Cervus elaphus is a highly appreciated and intensively managed game species throughout Europe. A common management objective is a sustainable harvest of large trophies. In southern Sweden, management has mainly aimed at preserving the nominate subspecies C. elaphus elaphus. Seasonal migration of red deer males may, however, complicate both harvest management as well as conservation efforts. I used individually identified male red deer in southern Sweden to observe distance travelled from rutting areas to areas used by males in summer and winter. Adult males were identified by antler shape and photo-documented during rut. Photos from the rut were compared to trophies of deer harvested or found dead, to found cast antlers and to stags photographed during summer. From 1969 to 2007, a distance between rutting ground and summer/winter quarters was established for 96 identified stags. An average distance of 14 km and a maximum distance of 47 km were recorded between rut and summer/winter observations. The seasonal migration of males increases the risk of overexploitation of males with harvest in both rutting areas and wintering areas. Harvest management and conservation efforts may fail if males seasonally migrate outside the management unit. The results suggest that seasonal migration must be considered in harvest management and conservation and that there is a need for a regulation of male harvest. Furthermore, the study stresses that the success in deer management of single hunting units, may be largely dependent on the harvest policies in the near surroundings as well as in areas tenths of kilometres away, suggesting that a successful management must rely on co-operation and co-ordination on a landscape scale.     

Population structure and genetic diversity of red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in forest fragments in north-western France

Red deer have been subjected to anthropogenic interference for many centuries. Most populations are managed according to hunting schedules, some have been kept long-term in enclosures and other populations have been restocked with foreign deer. The red deer in the Brittany region of north-western France only occupy the largest forests in the region, reaching quite high densities in restricted areas. Here, we aimed to assess the extent of the genetic variability of the populations in four forest fragments and investigate their population genetic structure. We show that, despite relatively large expected heterozygosity values, these geographically isolated populations are genetically impoverished relative to individuals from large continuous forests in other parts of Western Europe. We provide evidence for population genetic structure with large genetic differentiation between geographically close populations, suggesting the absence of effective exchange between the forests. Using samples from the most likely source population, we show that at least two populations were non-indigenous. In order to limit further loss of genetic diversity, it should be a management objective to reduce isolation of the different forests, rather than further increase it by fences and hunting practices that could limit free movement of red deer.     

Country,Cover or Protection: What Shapes the Distribution of Red Deer and Roe Deer in the Bohemian Forest Ecosystem?

The Bohemian Forest Ecosystem encompasses various wildlife management systems. Two large, contiguous national parks (one in Germany and one in the Czech Republic) form the centre of the area, are surrounded by private hunting grounds, and hunting regulations in each country differ. Here we aimed at unravelling the influence of management-related and environmental factors on the distribution of red deer (Cervus elaphus) and roe deer (Capreolus capreolus) in this ecosystem. We used the standing crop method based on counts of pellet groups, with point counts every 100 m along 218 randomly distributed transects. Our analysis, which accounted for overdispersion as well as zero inflation and spatial autocorrelation, corroborated the view that both human management and the physical and biological environment drive ungulate distribution in mountainous areas in Central Europe. In contrast to our expectations, protection by national parks was the least important variable for red deer and the third important out of four variables for roe deer; protection negatively influenced roe deer distribution in both parks and positively influenced red deer distribution in Germany. Country was the most influential variable for both red and roe deer, with higher counts of pellet groups in the Czech Republic than in Germany. Elevation, which indicates increasing environmental harshness, was the second most important variable for both species. Forest cover was the least important variable for roe deer and the third important variable for red deer; the relationship for roe deer was positive and linear, and optimal forest cover for red deer was about 70% within a 500 m radius. Our results have direct implications for the future conservation management of deer in protected areas in Central Europe and show in particular that large non-intervention zones may not cause agglomerations of deer that could lead to conflicts along the border of protected, mountainous areas.     

Population dynamics in a guild of four Mediterranean ungulates: density‐dependence,environmental effects and inter‐specific interactions

Population fluctuations in ungulates are driven by both intrinsic and extrinsic factors. Available information, however, mainly refers to arctic, temperate and African ungulate populations, while the dynamics of Mediterranean species, exposed to a milder climate, is known to a much lesser extent. Here we studied the population dynamics of four wild ungulate species in the Castelporziano Preserve near Rome, Italy, as obtained from detailed bag counts from hunting drives during the period 1878–1986: the Italian roe deer Capreolus capreolus italicus, the Maremma wild boar Sus scrofa majori (both endemic to Italy), the native red deer Cervus elaphus, and the alien fallow deer Dama dama. We also considered the effects of the presence of another alien ungulate, the nilgai Boselaphus tragocamelus. This ungulate community experienced an accidental ‘removal experiment’ when, during World War II, red deer and nilgai were exterminated. This event and the length of the time series allowed us to test two main hypotheses: 1) that the complexity level of the ungulate community affects the strength of intra‐ and inter‐specific competition; and 2) that in Mediterranean environments intra‐ and inter‐specific interactions are stronger than climate forcing. Statistical methods ranged from state‐space‐modelling, GLM analysis and structural equation models. The results indicated that direct intra‐specific density dependence played a relevant role for all species, and was stronger after the removal. A complex pattern of species interactions was however revealed; fallow deer had a negative effect on roe deer population, while roe deer had an apparent positive effect on red deer and wild boar, possibly mediated by environmental factors. Nilgai appeared to facilitate all deer species. The results of the analysis also confirmed that at present climate appears to play a minor role with respect to density dependence; however, the increasing aridity of the Mediterranean area could change this picture in coming decades.