Spatiotemporal variation of breeding gull species in a boreal lake complex in Finland: Implications for conservation

Understanding spatial and temporal patterns of species is a prerequisite for successful species and habitat conservation. Spatial variation in breeding sites of four gull species was studied in southern Finland in an oligo-mesotrophic lake complex covering almost 50 km² of water areas and 290 km of shoreline in three census periods in 1986-2004. Two of the species have declined and are regarded as red-listed in Finland (black-headed gull ridibundus and lesser black-backed gull L. f. fuscus) and two have increased (common gull L. canus and herring gull L. argentatus) in numbers during the past decades. The numbers of breeding pairs and the percentage similarity in the spatial distribution of pairs of each species in grid squares were compared between different census periods at resolutions of 0.25, 1 and 4 km². The common gull showed very high percentage similarity between the different census periods and consequently low spatial turnover in nesting sites, whereas the red-listed species, particularly the black-headed gull, had much higher spatial turnover. The spatiotemporal dynamics of gull species should thus be taken into account in conservation planning. If site protection is based only on information of breeding gulls in one year, a large or even major proportion of the breeding red-listed gulls might be outside the protected areas after a few decades. Due to the large spatiotemporal variation of red-listed gulls, areas to be protected should cover a rather large proportion of a boreal lake, not only individual islets or islands.     

Vulnerability of ground-nesting waterbirds to predation by invasive American mink in the Cape Horn Biosphere Reserve, Chile

Biological invasions constitute one of the most important threats to biodiversity. This is especially true for “naïve” birds that have evolved in the absence of terrestrial predators in island ecosystems. The American mink (Mustela vison) has recently established a feral population on Navarino Island (55°S), southern Chile, where it represents a new guild of terrestrial mammal predators. We investigated the impact of mink on ground-nesting coastal waterbirds with the aim of deriving a vulnerability profile for birds as a function of different breeding strategies, habitat, and nest characteristics. We compared rates of nest survival and mink predation on 102 nests of solitary nesting species (Chloephaga picta, Tachyeres pteneres), on 361 nests of colonial birds (Larus dominicanus, Larus scoresbii, Sterna hirundinacea), and on 558 artificial nests. We calculated relative mink and bird densities at all nest sites. Nests of colonial species showed the highest nest survival probabilities (67-84%) and no predation by mink. Nest survival rates for solitary nesting species were lower (5-20%) and mink predation rates higher (10-44%). Discriminant analyses revealed that mink preyed upon artificial nests mainly at shores with rocky outcroppings where mink were abundant. High nest concealment increased the probability for predation by mink. Conservation planning should consider that invasive mink might severely affect the reproduction success of bird species with the following characteristics: solitary nesting, nesting habitat at rocky outcrop shores, and concealed nests. We recommend that work starts immediately to control the mink population with a priority in the nesting habitats of vulnerable endemic waterbirds.     

The evaluation of key sites for breeding waders in lowland Scotland

This study considers the selection of sites based on the densities of breeding waders that would be considered to be distributed widely across common habitats in Scotland: so-called ‘wider countryside’ species. Five methods to provide a hierarchical classification of sites are assessed. Fifty percent of the populations of redshank (Tringa totanus) and snipe (Gallinago gallinago) occur on just 2 and 4.5% of Scottish lowlands, respectively, yet neither are suitably protected by conservation measures and both would benefit from a carefully targeted approach to their conservation. Three species, lapwing (Vanellus vanellus), curlew (Numenius arquata) and oystercatcher (Haematopus ostralegus) are more appropriately considered to be dispersed species. We propose that, if the density of a breeding wader on a site exceeds the threshold level predicted for the top 1% of Scottish lowland for that species, and if the area of the site exceeds 1 km² then that site be considered to be a key site for farmland waders. The minimum density (in pairs km⁻²) must exceed 16.8 for lapwing, 10.1 for oystercatcher, 6.1 for snipe, 7.5 for curlew or 3.6 for redshank.     

Estimating the density of a jaguar population in the Brazilian Pantanal using camera-traps and capture-recapture sampling in combination with GPS radio-telemetry

The jaguar (Panthera onca) is the largest feline in the Americas and third largest world-wide, smaller in size only to the tiger (P. tigris) and lion (P. leo). Yet, in comparison, relatively few studies on jaguar population densities have been conducted and baseline data for management purposes are needed. Camera trapping and capture-recapture sampling methods were used to estimate the size of a jaguar population in the Pantanal’s open wet grassland habitat, an important area for the long-term survival of the species. This study is the first jaguar population estimate conducted in co-operation with a GPS-telemetry study providing an important opportunity for comparing different methods of density estimation. An accessible area within a 460 km² privately-owned ranch was sampled with equal effort during the dry seasons of 2003 and 2004. Thirty-one and twenty-five individual jaguars were identified in 2003 and 2004, respectively. Estimates of jaguar abundance were generated by program CAPTURE. Density estimates were produced according to different methods used to calculate the effectively sampled areas which ranged from 274 to 568 km². For 2003, the currently-used mean maximum distance moved (MMDM) method produced a density of 10.3 jaguars/100 km², while GPS-telemetry-based calculations produced a mean density of 6.6 jaguars/100 km². For 2004, the MMDM method produced an estimate of 11.7 jaguars/100 km² while GPS-telemetry calculations produced a density of 6.7 jaguars/100 km². Our results suggest that the widely-used MMDM method used to calculate effectively sampled areas is significantly under-reflecting maximum distances moved by jaguars and their range-use and, thereby, considerably inflating cat density estimates. This overestimation could place a population in a difficult situation by lengthening the time taken to initiate protection measures because of underestimating the risk to that population.     

Movements and activities of snow leopards in Southwestern Mongolia

Four adult (2M:2F) snow leopards (Uncia uncia) were radio-monitored (VHF; one also via satellite) year-round during 1994-1997 in the Altai Mountains of southwestern Mongolia where prey densities (i.e., ibex, Capra siberica) were relatively low (∼0.9/km²). Marked animals were more active at night (51%) than during the day (35%). Within the study area, marked leopards showed strong affinity for steep and rugged terrain, high use of areas rich in ungulate prey, and affinity for habitat edges. The satellite-monitored leopard moved more than 12 km on 14% of consecutive days monitored. Home ranges determined by standard telemetry techniques overlapped substantially and were at least 13-141 km²in size. However, the satellite-monitored individual apparently ranged over an area of at least 1590 km², and perhaps over as much as 4500 km². Since telemetry attempts from the ground were frequently unsuccessful , we suspect all marked animals likely had large home ranges. Relatively low prey abundance in the area also suggested that home ranges of >500 km²were not unreasonable to expect, though these are >10-fold larger than measured in any other part of snow leopard range. Home ranges of snow leopards may be larger than we suspect in many areas, and thus estimation of snow leopard conservation status must rigorously consider logistical constraints inherent in telemetry studies, and the relative abundance of prey.     

Predator-prey relationships and responses of ungulates and their predators to the establishment of protected areas: A case study of tigers, leopards and their prey in Bardia National Park, Nepal

Interactions among sympatric large predators and their prey and how they respond to conservation measures are poorly known. This study examines predictions concerning the effects of establishing a protected area in Nepal on tigers (Panthera tigris), leopards (Panthera pardus), and their ungulate prey. Within a part of the park, after 22 years the total density of wild ungulates had increased fourfold, to ca. 200 animals/km², almost exclusively due to a remarkable increase in chital deer (Axis axis). Tiger density also increased markedly to nearly 20 animals/100 km², whereas leopard density did not and was ca. 5 animals/100 km². The prediction that grazers should increase more than browsers was only partially supported. The prediction of positive density dependence in prey selection was not supported. Instead, the most abundant species (chital and hog deer, Axis porcinus) were killed less frequently than expected, whereas the lower-density wild boar (Sus scrofa) was preferred. Predictions that (i) initially rare species suffer highest predation was partially supported, that (ii) predation is highest among the most abundant prey was not supported, and that (iii) predation is highest among the most preferred prey independently of their densities was supported. Clearly, the conservation efforts adopted in Bardia were successful, as both tigers and their natural prey base increased. However, the positive numerical response of tigers limited and depressed the abundance of some prey species. Thus, conservation activities aimed at restoring large predators are likely to change in the composition of the overall mammal community, potentially eliminating rare but preferred prey species.     

Human interference and gull predation in cormorant colonies

The vulnerability of cormorant eggs and nestlings to gull predation is greatly increased by human intrusion into breeding colonies. Observations of the behaviour of double-crested cormorants (Phalacrocorax auritus) and king shag (P. albiventer) vis-à-vis the behaviour of great black-backed and herring gulls (Larus marinus and L. argentatus) on the one hand and dolphin gulls (Leucophaeus scoresbii) on the other indicate that certain predictable offensive and defensive interactions occur when breeding birds are disturbed. Knowledge of these behavioural patterns suggests that interference with normal breeding activities and the resulting reduction in productivity of cormorants can be minimised in colonies where human access is carefully regulated.     

Numerical and behavioral responses of waterfowl to the invasive American mink: A conservation paradox

A behavioral response is usually the first adaptation of animals to environmental changes such as the introduction of an alien invasive species, and it seems to be the most important factor in reducing the negative impact of new predators on naive prey. This study examined changes in the abundance and behavior of breeding coots (Fulica atra) and great crested grebes (Podiceps cristatus) in response to invasion of the Mazurian Lakeland (NE Poland) by American mink (Neovison vison), first recorded here in 1984. Feral mink densities varied from 1.0 to 14.6 individuals per 10 km of shoreline. Mink were responsible for the losses of 17–58% of experimental clutches (artificial nests containing chicken eggs) that were deployed. Mink destroyed more experimental clutches than other predators; the effects of mink predation were similar in reeds and on the bank. Radio-tracking showed that mink avoided lake banks near human settlements. Mink have markedly affected the breeding distribution of both species and caused a greater than 20-fold decline in coot numbers. For coots and grebes breeding distribution and nesting success were positively associated with built-up areas along lake shores and also with whether the birds were breeding in colonies. Increased brood survival of coots and grebes near human settlements is a paradox in the sense that a man-made habitat change has provided refuge from an alien predator.     

Evidence-based culling of a facultative predator: Efficacy and efficiency components

Human activities have greatly modified predator-prey dynamics within seabird communities by favouring a rapid increase in density of large predatory gulls. To counteract such a subsidized growth, conservation agencies perform massive random culling programs, which generally fail to restore the original predator-prey relationship. We used long-term individual-based information to evaluate the effects of a selective culling of a top seabird predator, the yellow-legged gull (Larus michahellis), on the predatory pressure, survival and reproductive success probabilities of a secondary prey, the vulnerable European storm-petrel (Hydrobates pelagicus). The selective removal of only 16 gulls in 3 years led to a reduction of ca. 65% in the number of petrels killed, and to a relative increase in their survival and breeding success probabilities of 16% and 23%, respectively. Our results show that only a few specialised predators were responsible for the bulk of the impact on a secondary prey and that the removal of those specialised individuals was an effective and efficient way to improve prey demographic parameters.     

Restoring forest herb communities through landscape-level deer herd reductions: Is recovery limited by legacy effects?

White-tailed deer (Odocoileus virginianus) overbrowsing has altered plant species diversity throughout deciduous forest understories in eastern North America. Here we report on a landscape-level (306 km²) project in Pennsylvania, USA that tracked the herbaceous community response to deer herd reductions. From 2001 to 2007, we estimated deer densities, browse impact on woody seedlings, and censused the herbaceous flora in permanent plots throughout the area. We assessed herb layer species richness, abundance, and dominance and measured three known phytoindicators of deer impact: Trillium spp., Maianthemum canadense, and Medeola virginiana. We predicted that browse-sensitive taxa would increase in abundance, size, and flowering as would overall species diversity following deer culls and browse impact that declined by an order of magnitude by 2007. Following intensified deer harvests, we observed a limited recovery of the herbaceous community. Trillium spp. abundance, height, and flowering; M. canadense cover; and M. virginiana height all increased following herd reductions. Similarly, forb and shrub cover increased by 130% and 300%, respectively. Nevertheless, species diversity (i.e., richness and dominance) did not vary. Our work demonstrates that reducing deer densities can provide rapid morphological and population-level benefits to palatable species without a concomitant recovery in diversity. We suggest that decreasing deer populations alone may not promote plant diversity in overbrowsed, depauperate forests without additional restoration strategies to mitigate a browse-legacy layer dominated by browse-resistant species.     

How does human exploitation affect impala populations in protected and partially protected areas? - A case study from the Serengeti Ecosystem, Tanzania

Human exploitation can have severe conservation implications for wildlife populations. In the Serengeti ecosystem, Tanzania, illegal hunting is a serious concern for wildlife management, and in this study we investigated if density, demography and behaviour can be used as indicators of human exploitation. We used impala (Aepycerus melampus) as a model species to study human exploitation inside and outside a strictly protected area. Over a six month period, a total of 2050 km of transects were driven in the different protected areas (National Park, Game Reserve, Open Area). Densities were estimated by using distance sampling and the partially protected areas were found to have significantly lower densities (4.3 ind/km²) than the National Park (15.3 ind/km²). A variation in density between different sections within the National Park was also found. However, we found no differences in group sizes. Moreover, the sex-ratio was more skewed towards females in the partially protected areas and in sections within the National Park close to villages. In addition, impalas showed higher alertness levels, and longer flight initiation distance to an approaching human in the partially protected areas compared to the National Park. The present harvest levels by illegal hunting in the study area are most likely the cause of the observed differences. Our results suggest that density, demography and behaviour can be used as indicators of human exploitation, but that this probably varies according to local hunting pressure. Furthermore, it could be expected that the results obtained in this study might reflect the state of other ungulates in the area, which raises concern whether management objectives for the buffer zones of Serengeti National Park are met.     

Degradation of tropical forest in Hainan, China, 1991-2008: Conservation implications for Hainan Gibbon (Nomascus hainanus)

The Hainan gibbon (Nomascus hainanus) is one of the most endangered primates in the world, confined to mature natural forest in Hainan Island, China. We assessed changes in habitat condition on the island between 1991 and 2008, using vegetation maps generated by remote-sensing images. We defined forest suitable for gibbons based on composition, tree size and canopy cover. During the 17-year period, the area of suitable gibbon forest decreased by 540 km² (35%) across the whole island, and by 6.3 km² (7%) in the locality of the sole remaining gibbon population at Bawangling National Nature Reserve. The forest patches large enough (>1 km²) to support a gibbon group decreased from 754 km² to 316 km² in total area, and from 92 to 64 in number. Suitable natural forest was mainly replaced by plantations below 760 m, or degraded by logging, grazing and planting of pines above 760 m. Meanwhile, forests in former confirmed gibbon areas became more fragmented: mean area of patches decreased by 53%. We mapped the patches of natural forest in good condition which could potentially support gibbons. We recommend a freeze on further expansion of plantations between core patches at Bawangling, Jiaxi-Houmiling and Yinggeling Nature Reserves in accordance with forest protection regulations; establishment of nature reserves in currently unprotected natural forest patches elsewhere in line with the local government’s nature reserve expansion policy; and active natural-forest restoration between remaining fragments at Bawangling.     

Interference competition in a threatened seabird community: A paradox for a successful conservation

It is often assumed that conservation actions targeting a threatened community (e.g. habitat protection) will result in similar benefits for all species. However, complex interactions between species, such as interference competition, may result in displacement of subordinate, vulnerable species. We analysed here the spatio-temporal population dynamics of a threatened seabird community since the protection in the 1980s of several breeding sites at the Ebro Delta, western Mediterranean, Spain during 1980-2007. Competition for the most suitable patches was governed by body size, with smaller species avoiding associations with larger, dominant species. We tested whether the density increase of the larger species (yellow-legged gull Larus michahellis and Audouin’s gull Larus audouinii) at La Banya (the highest quality patch within the Delta) affected species diversity at the local level. As expected, such diversity decreased, resulting also from the colonisation of other sites by smaller species, some abandoning the former area. The conservation paradox appeared because the interference competition was dominated not only by the largest species of the community, the yellow-legged gull, which is sometimes considered a pest species, but also by the vulnerable Audouin’s gull, a flagship species which has ca. 65% of the total world population at La Banya. Nevertheless, the availability of alternative patches within the Ebro Delta ensured a high biodiversity at regional scale. Results highlight the role of turnover of competing species within a community in ecosystem function and stability, and the importance of alternative sites and dispersal abilities for the conservation of vulnerable communities.     

Building large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species

We developed large-scale spatially explicit models to predict the distribution of suitable habitat patches for the Greater rhea (Rhea americana), a near-threatened species, in two areas of central Argentina with different land use: a grassland area (ca. 4943 km²) mainly devoted to cattle grazing and an agro-ecosystem area (ca. 4006 km²) mostly used for crop production. The models were developed using logistic regression and were based on current records of Greater rhea occurrence coupled with remote sensing data, including land cover and human presence variables. The habitat suitability maps generated were used to predict the suitable habitat patch structure for wild rhea populations in each area. Fifty-one percent of the total grassland area was suitable for the species, being represented by a single large patch that included 62% of the individual locations. In the agro-ecosystem, only 28% of the total area was suitable, which was distributed among four patches. Seventy percent of rhea observations were in suitable habitat, with all rheas grouped in the largest patch. Conservation efforts for preserving wild rhea populations should be focused on maintaining habitats similar to grasslands, which are less profitable for landowners at present. Consequently, the protection of the pampas grasslands, a key habitat for this species as well as for others with similar habitat requirements, will demand strong conservation actions through the reconciliation of interests between producers and conservationists, since the proportion of croplands is increasing.     

The use of camera traps for estimating tiger and leopard populations in the high altitude mountains of Bhutan

We used camera traps in combination with capture-recapture data analysis to provide the first reliable density estimates for tigers and leopards from the high altitude and rugged terrain in Bhutan’s Jigme Singye Wangchuck National Park. Fifty days of camera trapping in each of five study zones collapsed into two trapping blocks, resulted in a sampling effort of 4050 trap days. Camera trapping yielded 17 tiger photos (14 left flanked and 3 right flanked) and 48 leopard photos (25 left flanked and 23 right flanked). Using photos of these left flank, the closed heterogeneous Jackknife Model M_h was the best fit for the capture history data. A capture probability () of 0.04 was obtained for both tigers and leopards, thus generating population size (N) of 8 tigers (SE = 2.12) and 16 leopards (SE = 2.91) with densities of 0.52 tiger 100 km⁻² and 1.04 leopard 100 km⁻². Photographic evidence indicated that tigers and leopards did not overlap in their spatial use of space. Tigers preferred less disturbed areas located further away from settlements, while leopards appeared to be more resilient to disturbances in so far as they were found nearer to human settlements. Camera trapping using a capture-recapture framework was an effective tool for assessing population sizes for tiger and leopard in low density areas such as Bhutan.     

The impact of trophy hunting on lions (Panthera leo) and other large carnivores in the Bénoué Complex, northern Cameroon

In West and Central Africa large carnivores have become increasingly rare as a consequence of rapid habitat destruction and lack of resources for protected area management. The Bénoué Complex (23,394 km²) in northern Cameroon is a regionally critical area for large mammal conservation. In the complex lions (Panthera leo), leopards (Panthera pardus) and spotted hyenas (Crocuta crocuta) are formally protected in three national parks and 28 hunting zones. Over-hunting may be having a strong additive effect precipitating declines in large carnivore numbers across the complex. We used a coarse level track index method to estimate the relative abundance of these three species both in hunting zones and national parks. The results were interpreted with respect to ungulate abundance, and hunting impact. There was no significant difference between the densities of medium to larger species of ungulates in the hunting zones and the national parks, and no difference in leopard and spotted hyena densities in the respective areas. However, lions occurred at significantly lower densities in the hunting zones, and even in the national parks occurred at significantly lower densities than prey biomass would predict.     

Monitoring rare or elusive large mammals using effort-corrected voluntary observers

Populations of rare or elusive large mammals are difficult to monitor, because they usually are secretive, solitary, occur at low densities, and have large home ranges. The global trend of generally decreasing large carnivore populations necessitates new, feasible, reliable, and cost-effective monitoring methods. We evaluate an index method developed for monitoring populations of moose (Alces alces) based on voluntarily and systematically collected observations from hunters, corrected for effort, for use in monitoring populations of large carnivores in Sweden. For our evaluation, we used independent estimates of minimum brown bear (Ursus arctos) densities from DNA-based scat surveys and brown bear distribution from mandatory reports from successful bear hunters. We verified that the index correctly reflected bear distribution. We also found strong linear relationships between the indices and the independent density estimates for bears at the scale of local management units (about 1000-2000 km²) in all three regional study areas (adjusted R² = 0.88-0.60). Our results suggest that systematic, effort-corrected reports of observed animals can be an alternative and accurate monitoring method for the conservation and management of large mammals occurring over large areas when large numbers of willing volunteers are available (effort >30,000 h).     

Turning back the tide of American mink invasion at an unprecedented scale through community participation and adaptive management

Successful eradications of harmful invasive species have been mostly confined to islands while control programs in mainland areas remain small, uncoordinated and vulnerable to recolonisation. To allow the recovery of threatened native species, innovative management strategies are required to remove invasives from large areas. We took an adaptive approach to achieve large scale eradication of invasive American mink in North East Scotland. The project was centred on the Cairngorms National Park (Scotland), with the primary aim of protecting endangered water vole populations. The project was initiated by scientists and supported and implemented through a partnership comprising a government agency, national park authority and local fisheries boards. Capitalising on the convergent interests of a diverse range of local stakeholders, we created a coordinated coalition of trained volunteers to detect and trap mink. Starting in montane headwaters, we systematically moved down river catchments, deploying mink rafts, an effective detection and trapping platform. Volunteers took increasing responsibility for raft monitoring and mink trapping as the project progressed. Within 3 years, the project removed 376 mink from 10570 km² with the involvement of 186 volunteers. Capture rate within sub-catchments increased with greater connectivity to mink in other sub-catchments and with proximity to the coast where there is more productive habitat. The main factor underpinning the success of this project was functional volunteer participation. The project is a reason for optimism that the tide of invasion can be rolled back on a large scale where the convergent interest of local communities can be harnessed.     

黄土高原不同侵蚀类型区侵蚀产沙强度变化及其治理目标

为了确定黄土高原不同侵蚀类型区的治理目标,采取"水文—地貌法",利用98个水文站控制区和234个侵蚀产沙单元,在分析其不同治理阶段土壤侵蚀产沙变化特征与减沙幅度,不同侵蚀强度面积的变化及其空间分布的基础上,提出了未来20a黄土高原主要流失区的区域治理目标:土壤流失量控制在3.60×108 t左右,土壤侵蚀模数1 300 t/(km2.a)左右。其中,黄土峁状丘陵沟壑区为3 000t/(km2.a),黄土梁状丘陵沟壑区为2 000t/(km2.a),干旱黄土丘陵沟壑区为2 000t/(km2.a),黄土平岗丘陵沟壑区为1 000t/(km2.a),风沙黄土丘陵沟壑区为1 000t/(km2.a),黄土山麓丘陵沟壑区为1 000t/(km2.a),森林黄土丘陵沟壑区为300t/(km2.a),黄土高塬沟壑区为1 500t/(km2.a),黄土残塬沟壑区为3 000t/(km2.a),黄土阶地区为500t/(km2.a),风沙草原区为500t/(km2.a),高原土石山区为100t/(km2.a)。未来20a黄土高原的治理重点区域为黄土峁状丘陵沟壑区(2.20×104 km2)、干旱黄土丘陵沟壑区(1.50×104 km2)、黄土高塬沟壑区(8 600km2)、黄土梁状丘陵沟壑区(4 600km2)。     

Ecology and use of arctic fox Alopex lagopus dens in Norway: tradition overtaken by interspecific competition?

Arctic fox Alopex lagopus dens in Norway are mostly large and well-defined structures of great age. Examinations of these dens have been used in surveys of the breeding and population status of the arctic fox in Fennoscandia. In this study, dens all over north Norway and in some parts of south Norway were surveyed (n=214), and several aspects of den characteristics, localisation and use were recorded. Red fox Vulpes vulpes dens were also recorded whenever found, including red fox use of arctic fox dens (total n=211). Most of the arctic fox dens were of the “classic” type dug out in sandy soil (n=210). The average number (±1 SD) of open entrances (at least half open) was 27.4±19.0, and the average size was 363.1±288.7 m². Regional variation was found in both these measurements, as well as in two variables classifying the vegetation at the den. However, no consistent south-north gradient was found, although the most southern and the most northern regions were extremes in several respects. A positive correlation was found between den size and the number of entrances, as well as between these two variables and the two vegetation variables. For each arctic fox den, height above the tree line, distance to the tree line, distance between arctic fox dens, distance to the nearest red fox den, distance to a main road and distance to other human activity were estimated. Regional variation was found in all these variables, but no consistent south-north gradient. Again, the most southern region deviated most. There were large differences in the proportion of dens used by arctic foxes between the nine regions, with one region outstanding in its production of arctic fox litters and paucity of red foxes. The proportion sum of arctic fox entrances to sum of red fox entrances reliably, but not perfectly, identified the best arctic fox regions. Arctic fox dens that were known to have produced litters at least once during 1980-2001 had about twice as many entrances as dens without pups, were 32% larger and had a more lush vegetation. They were also situated further from a red fox den. Distance to red fox dens and height above the tree line were the most important factors regarding arctic fox use of dens. Thus, the red fox was identified as a threat to the arctic fox population.