Demography and conservation of the Tristan albatross Diomedea [exulans] dabbenena

The Tristan albatross Diomedea [exulans] dabbenena is the third rarest albatross species, with a breeding population of around 1500 pairs almost totally restricted to Gough Island in the Tristan da Cunha group, central South Atlantic. During January 2000, the entire breeding population of Gough was surveyed for the first time since 1956, and 2400 incubating pairs were counted. An analysis of the areas that are likely to have been surveyed most accurately in the past suggests that the population has decreased by around 28% over 46 years. The number of large chicks counted over three successive seasons (1999-2001) was highly variable (range 318-1129). The average count over this period (705 chicks) is less than counts made in 1979 (792) and 1982 (798). A total of 656 chicks were counted in September 2001, giving an island breeding success of just 27.3%. However, breeding success varied considerably in different areas of the island, ranging from 17.6 to 68.0%. During the 2001 season most breeding failures were of large chicks, and over 4 years where data were available, 75% of breeding failures occurred during the chick period. Predation by introduced house mice Mus musculus is the most likely cause of chick mortality. In a small study population, birds began breeding at an average age of 9.7 years and annual adult survival from 1985 to 2001 was 92.6% (SE=1.6%). Both breeding success and adult survival estimates are low in comparison with other Diomedea species and population modelling predicts a population decreasing at an annual rate of 2.9-5.3%. Further research is needed urgently to assess whether breeding success is typical, and to confirm that mouse predation is the cause of chick mortality. The low productivity of this species will compound the negative impacts of longline fishing mortality, which are likely to be reducing adult and juvenile survival.     

Conservation of bats in suburban landscapes: roost selection by Myotis yumanensis in a residential area in California

Protection of roosting habitat is essential to the conservation of bats in human-dominated landscapes. To help define bat roosting needs in suburban settings, we used radio telemetry to locate day roosts of a common North American species (Myotis yumanensis) within a residential area in California. Between June and August 2000, we tracked 16 bats to 20 roosts in two buildings and 18 trees. We used multiple logistic regression to assess roost selectivity at multiple spatial scales. Of 15 tree, plot, and site characteristics considered, only three helped distinguish roosts from random comparisons: tree diameter, distance to water, and forest cover. Myotis yumanensis preferred large trees (mean diameter 115 cm), and roosted only in the five species of largest mean diameter (Sequoia sempervirens, Pseudotsuga menziesii, Quercus agrifolia, Quercus lobata, and Acer macrophyllum). At the site level, these bats selected roosts that were close to water and had substantial forest cover in the surrounding 100-m radius. Unlike other North American bats, they often roosted in live trees (89% live). Relatively high roost fidelity (mean 4.8 days) and large travel distances between consecutive roosts (mean 1130 m) and between capture sites and roosts (mean 2007 m) may indicate a greater degree of roost limitation at this site versus other research sites. We recommend the preservation of large trees and forested parkland, particularly along stream corridors, to help maintain bat populations in urbanizing landscapes.     

Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii

The roosts of many IUCN-listed cave-roosting bat species are under threat from tourist development in SE Europe and other regions of the world. Much-needed conservation strategies require, among other information, an understanding of their roost movements and population dynamics, which can now be obtained relatively quickly using advanced models. We have studied the long-fingered bat, Myotis capaccinii, an obligate cave-dweller, in Dadia National Park, Greece. The species formed colonies of up to a few thousand individuals and was highly mobile, frequently switching summer roosts up to 39 km apart, even during late pregnancy. The bats migrated to distant hibernacula including a cave in Bulgaria 140 km NW of the Park. Adult recapture probabilities varied with season and sex: low female recapture rates in autumn, relative to spring and summer, indicated non-random temporary emigration following nursery colony dispersal. The opposite pattern was seen in males: increasing recapture rates in the autumn suggest that males gather in these roosts to mate with females in transit. Adult survival (0.86-0.94) was similar in females and males, similar in winter and summer, and comparable to recent estimates for other bats based on similar modelling techniques. Sex-based differences in juvenile recapture suggest female philopatry and male-biased dispersal. Our work shows that protection of M. capaccinii roosts must extend beyond the Park’s and indeed the country’s boundaries: its conservation requires large-scale, trans-national integrated conservation plans. Our results will apply to many other warm-temperate species with similar life history cycles.     

Patterns of roost use by bats in a neotropical savanna: implications for conservation

Neotropic bats comprise a highly species-rich group and are considered important regulators of complex ecological processes. Resource partitioning of roosts and food is considered a key element in shaping bat communities. In this paper we examine the roosting patterns of 10 bat species belonging to a neotropical bat community in a Bolivian savanna. Roosts were more common in open woodlands than in forest islands and human settlements. In open woodlands, Tabebuia heptaphylla comprised the largest proportion of roosting trees, whereas in forest islands, Gallesia integrifolia was the most important roosting tree for phyllostomid bats. By comparing active roosts with control holes in Tabebuia heptaphylla, we found that roost site selection by Noctilio albiventris was mainly based on tree diameter and inner width of the cavities. The earlier results are discussed in view of resource partitioning within savanna bat communities and their practical conservation.     

Location of roosts of the lesser long-eared bat Nyctophilus geoffroyi and Gould's wattled bat Chalinolobus gouldii in a fragmented landscape in south-eastern Australia

The location of roosts of two species of vespertilionid bats, the lesser long-eared bat (Nyctophilus geoffroyi) and Gould’s wattled bat (Chalinolobus gouldii), was investigated in a remnant vegetation-farmland mosaic and adjacent floodplain forest in south-eastern Australia. A total of 45 individuals of N. geoffroyi and 27 C. gouldii were fitted with radio transmitters, which resulted in the location of 139 and 89 roosts respectively. Selection of roosting habitat showed both inter- and intra-specific differences. At the landscape level, locations of roosts used by male and female N. geoffroyi differed markedly. Most males roosted in the farmland mosaic within 3 km (mean 1.9±2.9 km) of where they were captured while foraging in remnant vegetation. In contrast, roost sites of females were predominantly in the floodplain forest, 6-12 km from their capture site in farmland (mean for all females, 6.7±2.9 km). All maternity roosts were in the extensive floodplain forest, 4-10 km from foraging areas. Distances moved by this species were greater than previously recorded and further than predicted by flight morphology. Most C. gouldii roosted in the floodplain forest, 4-10 km from their capture site (6.9±1.6 km). Within the floodplain forest, both species roosted in areas that had higher densities of hollow-bearing trees than generally available; dead hollow-bearing trees for N. geoffroyi and large, live trees for C. gouldii. The scale of movements undertaken by these species means that they can obtain resources from multiple landscape elements. Both species used different habitats for roosting and foraging despite the energetic costs of commuting relatively large distances. Conservation of bat populations in rural environments needs to be considered at the landscape scale, with particular attention to identifying landscape elements that provide key resources.     

Influence of temperature in roost selection by Pipistrellus pygmaeus (Chiroptera): relevance for the design of bat boxes

The soprano pipistrelle (Pipistrellus pygmaeus) is a house-dwelling bat, whose maternity colonies are often affected by exclusion from buildings in southern Europe. Bat boxes with a suitable thermal behaviour can provide alternative roosts. However, this is particularly difficult to achieve in hot climates, where boxes may overheat. We studied roost temperatures and roost thermal preferences of this bat in attics and roofs in a Mediterranean region. Roosts reached high temperatures, experienced great thermal fluctuations and had wide spatial thermal ranges. Bats preferred the warmest locations within roosts, indicating that they are a heat tolerant species. However, they avoided temperatures of 40 °C and above, suggesting that this is the upper limit of the species thermally neutral zone. The study of experimental sets of white, grey and black boxes demonstrated that the latter were the ones that most often reached the temperatures observed in roosts in buildings. They were also the ones occupied by larger numbers of bats. These results suggest that, even in hot Mediterranean climates, black boxes are the best option as alternative roosts for evicted colonies. However, on particularly hot days, bats left them, suggesting that black boxes could be improved by creating a wider thermal range.     

Long-term monitoring of tropical bats for anthropogenic impact assessment: Gauging the statistical power to detect population change

Bats are ecologically important mammals in tropical ecosystems; however, their populations face numerous environmental threats related to climate change, habitat loss, fragmentation, hunting, and emerging diseases. Thus, there is a pressing need to develop and implement large-scale networks to monitor trends in bat populations over extended time periods. Using data from a range of Neotropical and Paleotropical bat assemblages, we assessed the ability for long-term monitoring programs to reliably detect temporal trends in species abundance. We explored the magnitude of within-site temporal variation in abundance and evaluated the statistical power of a suite of different sampling designs for several different bat species and ensembles. Despite pronounced temporal variation in abundance of most tropical bat species, power simulations suggest that long-term monitoring programs (?20 years) can detect population trends of 5% per year or more with adequate statistical power (?0.9). However, shorter monitoring programs (?10 years) have insufficient power for trend detection. Overall, our analyses demonstrate that a monitoring program extending over 20 years with four surveys conducted biennially on five plots per monitoring site would have the potential for detecting a 5% annual change in abundance for a suite of bat species from different ensembles. The likelihood of reaching adequate statistical power was sensitive to initial species abundance and the magnitude of count variation, stressing that only the most abundant species in an assemblage and those with generally low variation in abundance should be considered for detailed population monitoring.     

Population recovery following differential harvesting of Allium tricoccum Ait. in the southern Appalachians

Over 100 native herbaceous forest perennials are harvested in the United States. Management of these populations for continued survival requires information on the long-term effects of varied harvesting levels on populations of forest perennials. Allium tricoccum Ait. is a species that is harvested throughout its range for its edible bulb. We examined the recovery of A. tricoccum populations over a 5-year period following experimental harvests of different intensity to predict the number of years required to return to pre-harvest levels and to establish guidelines for sustainable harvesting. Our harvest treatments included removal of 0, 25, 50, 75, and 100% of plants within experimental plots. Treatments were repeated at three sites. Population recovery was monitored for 4 years at two sites and 7 years at one site. We found a relationship between the level of harvest and the negative impacts on populations of A. tricoccum, with limited population recovery in the 4 years following harvesting. We projected recovery times as a function of harvest level using a population projection model. Deterministic projections of recovery time ranged from 148 years for a 95% harvest to 2.5 years for a 5% harvest. Incorporating stochasticity into our projections increased the uncertainty in our estimates of recovery time. For example the recovery time from a 5% harvest ranged from 1 to 89 years (95% confidence limit). Based on our results, a 10% harvest once every 10 years would, on average, be a sustainable level of harvest for A. tricoccum in southern Appalachian forests.     

Autumn swarming behaviour of Natterer’s bats in the UK: Population size, catchment area and dispersal

Many species of temperate bats visit underground sites in autumn, prior to hibernation, and chase each other in and around the entrances in an event known as autumn swarming. It has been suggested that swarming is a mating event that facilitates gene flow between bats from otherwise isolated summer colonies. We describe swarming behaviour at a cluster of four caves in the North of England and provide direct evidence for seasonal migration of Natterer’s bats (Myotis nattereri ) between these caves and many summer nursery roosts in a catchment area of at least 60 km radius. Mark-recapture analysis suggests that an estimated 2000-6000 Natterer’s bats use the three best-studied caves, together with smaller numbers of four other species: M. daubentonii, M. brandtii, M. mystacinus and Plecotus auritus. Recaptures also show that there is a small interchange of bats between these three caves, which are 0.6-1.5 km apart. Capture rates, automatic logging and video recordings at one cave suggest that an average 310-400 bats visit the cave each night. Mating at swarming sites would be particularly important for gene flow if a species showed no other dispersal pattern. We provide evidence for natal philopatry, with little or no dispersal, in both sexes of Natterer’s bats. The protection of swarming sites is discussed in light of our findings.     

Factors influencing survival and long-term population viability of New Zealand long-tailed bats (Chalinolobus tuberculatus): Implications for conservation

A population viability analysis is important for the management of endangered populations and requires the estimation of survival parameters. The long-tailed bat (Chalinolobus tuberculatus) is one of only two native terrestrial mammals currently found in New Zealand and is classed as vulnerable. Its viability in temperate beech (Nothofagus) forest, Eglinton Valley, Fiordland, New Zealand was estimated using mark-recapture data collected between 1993 and 2003 using the Program MARK. Survival was estimated based on a total of 5286 captures representing 1026 individuals. Overall annual survival varied between 0.34 and 0.83 but varied significantly among three sub-populations and with sex and age. Females generally had a higher survival rate compared to males; and adults had higher survival relative to juveniles. Survival of all bats was lower in years when the number of introduced mammalian predators was high and when the winter temperature was warmer than average. High numbers of introduced predators occurred during three of the 10 years in the study. Climate change may mean that the conditions that promote high predator numbers may occur more frequently. A preliminary population viability analysis using a projection matrix on the overall adult female population showed an average 5% decline per year (λ = 0.95). Increased predator control targeting a range of predators is required in years when their numbers are high in order to halt the decline of this population of long-tailed bats. Population estimates using minimum number alive estimates supported the population estimates derived from Program MARK and a population viability analysis using matrices.     

Statistical power for detecting trends with applications to seabird monitoring

Power analysis is helpful in defining goals for ecological monitoring and evaluating the performance of ongoing efforts. I examined detection standards proposed for population monitoring of seabirds using two programs (MONITOR and TRENDS) specially designed for power analysis of trend data. Neither program models within- and among-years components of variance explicitly and independently, thus an error term that incorporates both components is an essential input. Residual variation in seabird counts consisted of day-to-day variation within years and unexplained variation among years in approximately equal parts. The appropriate measure of error for power analysis is the standard error of estimation (S.E._est) from a regression of annual means against year. Replicate counts within years are helpful in minimizing S.E._est but should not be treated as independent samples for estimating power to detect trends. Other issues include a choice of assumptions about variance structure and selection of an exponential or linear model of population change. Seabird count data are characterized by strong correlations between S.D. and mean, thus a constant CV model is appropriate for power calculations. Time series were fit about equally well with exponential or linear models, but log transformation ensures equal variances over time, a basic assumption of regression analysis. Using sample data from seabird monitoring in Alaska, I computed the number of years required (with annual censusing) to detect trends of −1.4% per year (50% decline in 50 years) and −2.7% per year (50% decline in 25 years). At α=0.05 and a desired power of 0.9, estimated study intervals ranged from 11 to 69 years depending on species, trend, software, and study design. Power to detect a negative trend of 6.7% per year (50% decline in 10 years) is suggested as an alternative standard for seabird monitoring that achieves a reasonable match between statistical and biological significance.     

Species-specific barrier effects of a motorway on the habitat use of two threatened forest-living bat species

It is generally recognized that roads can adversely affect local animal populations but little is known how roads affect bats. In particular, no study compared the response of bats that differ in foraging ecology to motorways that cut through the breeding habitat. As bats are key species in conservation, such data are urgently needed for designing management plans. Using radio-telemetry, mist netting, and mark-recapture data we investigated the effects of a motorway with heavy traffic on the habitat use of two threatened forest-living bats. We compared barbastelle bats (Barbastella barbastellus), which forage in open space, to Bechstein’s bats (Myotis bechsteinii), which glean prey from the vegetation. Five of six radio-tracked barbastelle bats crossed the motorway during foraging and roost switching, flying through underpasses and directly over the motorway. In contrast, only three of 34 radio-tracked Bechstein’s bats crossed the motorway during foraging, all three using an underpass. Bechstein’s bats, unlike barbastelle bats, never crossed the motorway during roost switching. Moreover, only in Bechstein’s bats individuals foraging close to the motorway had smaller foraging areas than individuals foraging further away, whereas other forest edges had no such effect. Our data show that motorways can restrict habitat accessibility for bats but the effect seems to depend on the species’ foraging ecology and wing morphology. We suggest that motorways have stronger barrier effects on bats that forage close to surfaces than on bats that forage in open space, and discuss the implications of our findings for bat conservation during road construction.     

Sampling design and its effect on population monitoring: How much monitoring do turtles really need?

Marine turtles are a taxon of world-wide conservation concern. Effective long-term monitoring is hampered by the fact that populations are widely dispersed except during the breeding season. Thus most monitoring programmes focus on nesting beaches, necessitating resource intensive studies, often over months, that could conceivably, be less comprehensive and focus on more parsimonious sampling. We analyse 11 years of exhaustive monitoring data for two species of Mediterranean marine turtles (Chelonia mydas and Caretta caretta). We resample using a variety of plausible sub-sampling regimens to estimate the total annual nesting population. We project our dataset into the future applying a range of population change rates to explore how adopting a monitoring programme based on sub-sampling would, for example, affect our ability to detect population decline. We show that accurate annual population estimates can be achieved with as few as 14 days of survey effort providing monitoring spans the peak of nesting. Furthermore, the modelled impact of sampling-based monitoring suggests that the duration to detect population change is not increased greatly. Our findings have implications for all marine turtle monitoring and have applicability to other animal groups. It is often considered desirable to perform exhaustive monitoring, with aversion of basing policy recommendations on partial data. However, comprehensive long-term monitoring programmes, particularly in developing nations, although presenting a number of advantages, are often impossible. Accurate total annual censuses can be achieved through a variety of sub-sampling regimens without sacrificing the ability to detect changes in the population trends over time. In this example, a solid block of at least 3 weeks sampling that encompasses the peak of the nesting season is advised.     

Swarming of bats at underground sites in Britain—implications for conservation

We investigated the extent of visitation of underground sites during late summer and autumn (‘swarming’) by bats. Bats were captured at eight cave, mine and tunnel sites in southern England over 6 years (1995-2000). A total of 3077 bats representing 11 of Britain's 16 species was caught. Myotis bats predominated in autumn. At some sites these are rarely seen during winter hibernation counts. Myotis nattereri and M. daubentonii were most common. However, species composition changed seasonally. At one site, M. brandtii dominated early in the season and at all sites peak activity of M. daubentonii was earlier than in M. nattereri. Relatively high numbers of internationally vulnerable species such as M. bechsteinii and Barbastella barbastellus were recorded. There was a strong male bias in captures of swarming species. Swarming probably has an important reproductive function, most males are reproductively active at this time and such sites may be important for outbreeding. Due to this and the large number of individuals and species concerned, the conservation of ‘swarming sites’ is vitally important. Clearly swarming sites merit special consideration in conservation management strategies for cave-dwelling bats.     

The distribution of cave-dwelling bats and conservation status of underground habitats in Northwestern Turkey

The distribution and abundance of cave-dwelling bats were investigated in the Thrace region of northwest Turkey. Data were collected in two periods, January-March and April-May 2001, from 32 underground sites, 25 of which had not been surveyed previously. Approximately 76,000 bats representing 13 species were recorded. The most abundant species were Miniopterus schreibersii, Rhinolophus euryale, Myotis myotis/blythii, Myotis capaccinii, and Rhinolophus ferrumequinum. The roosts were evaluated for their conservation importance. The most important sites in Turkish Thrace are Dupnisa and Koyunbaba. The Dupnisa Cave serves as a hibernaculum to approximately 28,000 bats representing five species. The Koyunbaba Cave is a nursery roost to approximately 23,000 bats of six species. Presently, none of the caves in the region has adequate protection and some bat populations are under serious threat.     

The proportion of immature breeders as a reliable early warning signal of population decline: evidence from the Spanish imperial eagle in Doñana

Methods to evaluate population trends have recently received particular attention because of perceived declines in several species during the 20th century. We investigated whether age at first breeding could be used as an “early warning signal” to detect possible changes in population trends in long-lived species with deferred maturity using data from the Spanish imperial eagle (Aquila adalberti) population in Doñana National Park (Spain). This bird of prey is an endangered species that has suffered a rapid decline in this population during the last 10 years. As a result of our 27-year monitoring (1976-2002) study, we detected that an increase in immature breeding birds occurred before population decline became evident. The proportion of immature-plumaged breeders in the population was significantly higher during the period of decline than during the period of stability. In our case, more than 10% of immature breeders can be considered as an “early warning signal” that anticipates population decline. Owing to the ignorance of this warning signal, urgent actions for the recovery of this eagle population started 10 years later than necessary, and when population size had been reduced.     

The recent decline of a New Zealand endemic: how and why did populations of Archey's frog Leiopelma archeyi crash over 1996-2001?

Dramatic changes have been documented in New Zealand's vertebrate faunas since human settlement, involving major declines and extinctions, but over recent years few species have declined in numbers so rapidly as the terrestrial Archey's frog Leiopelma archeyi (Anura: Leiopelmatidae). Long-term monitoring over more than 20 years revealed a major population reduction of the species over 1996-2001 and L. archeyi is now classified as Nationally Critical under the New Zealand threat classification system. The decline progressed northwards in the Coromandel ranges, and mostly larger (female) frogs survived. On a 100 m² study plot at Tapu Ridge, annual population estimates averaged 433 frogs (SE ±32) over 1984-1994, declining by 88% to average 53 frogs (SE ±8) over 1996-2002. A mean annual survival rate of 82% for most years declined to 33% over 1994-1997. There is mounting evidence to suggest that disease is the major agent of decline, supported by (1) the rapidity and severity of decline, (2) the progressive (south to north) nature of decline, and (3) finding frogs with chytriodiomycosis from Batrachochytrium dendrobatidis at the time of decline. Surprisingly, sympatric populations of the semi-aquatic Leiopelma hochstetteri have not declined dramatically, nor has a western population of L. archeyi at Whareorino, despite chytridiomycosis occurring in some frogs there. Sustaining and restoring populations of L. archeyi in New Zealand raises major challenges for conservation management.     

Removal of nonnative fish results in population expansion of a declining amphibian (mountain yellow-legged frog, Rana muscosa)

The mountain yellow-legged frog (Rana muscosa) was once a common inhabitant of the Sierra Nevada (California, USA), but has declined precipitously during the past century due in part to the introduction of nonnative fish into naturally fishless habitats. The objectives of the current study were to describe (1) the effect of fish removal from three lakes (located in two watersheds) on the small, remnant R. muscosa populations inhabiting those lakes, and (2) the initial development of metapopulation structure in each watershed as R. muscosa from expanding populations in fish-removal lakes dispersed to adjacent habitats. At all three fish-removal lakes, R. muscosa population densities increased significantly following the removal of predatory fish. The magnitude of these increases was significantly greater than that observed over the same time period in R. muscosa populations inhabiting control lakes that remained in their natural fishless condition. Following these population increases, R. muscosa dispersed to adjacent suitable (but unoccupied) sites, moving between 200 and 900 m along streams or across dry land. Together, these results suggest that large-scale removal of introduced fish could result in at least partial reversal of the decline of R. muscosa. Continued monitoring of R. muscosa at the fish-removal sites will be necessary to determine whether the positive effects of fish eradication are sustained over the long-term, especially in light of the increasingly important role played by an emerging infectious disease (chytridiomycosis, caused by Batrachochytrium dendrobatidis) in influencing R. muscosa populations.     

Fishery by-catches of marine mammals in Galician waters: results from on-board observations and an interview survey of fishermen

Rates of cetacean by-catch were investigated in Galician waters (NW Spain) using a combination of observer trips on fishing vessels, a carcase recovery scheme and an interview survey of fishermen, carried out over two years (1998-1999). All these data sources are suspected of underestimating by-catch due to the sample of co-operating fishermen being, necessarily, self-selecting. No by-catches were seen during observer trips, although not all sectors of the fishery could be covered. The carcass recovery scheme yielded seventeen cetacean carcases over two years, which compares to around 35 by-caught cetaceans recorded annually by the Galician strandings network. Analysis of interview data suggested that around 200 cetaceans might be caught annually in inshore waters and around 1500 in offshore waters. Confidence limits were wide for all estimates. The highest by-catch rates were estimated for gillnets and offshore trawling. The majority of by-catches are small dolphins, probably mainly Delphinus delphis. Smaller numbers of Tursiops truncatus and Globicephala melas are also reported. Comparing the interview estimates of by-catch rates with minimum estimates of population size, it is suggested that by-catches of D. delphis and T. truncatus may be unsustainably high and that routine monitoring of fishery by-catches in Galician fisheries is required.     

Foraging site fidelity shapes the spatial organisation of a population of female western barbastelle bats

Information about the spatial distribution of individual foraging habitats, which determines the space required by a population to be viable, is vitally important for the conservation of bats. Detailed knowledge of this kind is crucial for the design of nature reserves and management plans. Recent field studies that examined habitat use and home range distribution of bats largely ignored factors like traditional range use vs. intra- and interspecific competition, which may be responsible for the spatial organisation of a population home range. We investigated the home range sizes and distribution of a maternity colony of the western barbastelle bat via radio telemetry in four consecutive tracking sessions (2004-2007). Based on 19 data sets with a total of 2737 fixes obtained from 12 females, we examined (i) how colony members partition the population home range (home range overlap analysis), and (ii) if individuals tracked over several years exhibit site fidelity. Home range sizes ranged from 125 to 2551 ha (median: 403 ha), with a median number of 2 core areas (range: 1-5 core areas per individual per year). The core area sizes ranged from 5 to 285 ha (median: 67 ha). A home range and core area overlap analysis showed that site fidelity across years seems to be more important for home range distribution than competition among colony members. This allows researchers to combine information from several years to get a deeper insight into the population’s spatial requirements.