A conservation framework for the golden eagle in Scotland: Refining condition targets and assessment of constraint influences

Conservation strategies have three elements: species protection, site protection and, the most challenging component, conservation in the wider environment. Watson and Whitfield [Watson, J., Whitfield, P., 2002. A conservation framework for the golden eagle Aquila chrysaetos in Scotland. Journal of Raptor Research 36(1 Suppl.), 41-49.] proposed a conservation framework for the golden eagle in Scotland whose main innovation, taking a lead from EU conservation Directives, was to incorporate the concept of favourable condition and its maintenance by implementing conservation policies that are regionally targeted at known constraints in the wider environment. Three criteria were suggested to judge favourable condition: a national abundance target of breeding pairs, national and regional targets for breeding productivity, and regional targets for the extent of suitable habitat which is occupied by breeding pairs. Here we refine these criteria, first to take the national abundance target and use it to set regional targets. Distribution targets were implicitly incorporated by this process because abundance targets were set regionally, with regard to the proportion of known territories which should be occupied. This allowed us to dispense with the distribution criterion as originally proposed. Our next refinement was to base regional demographic targets not only on breeding productivity, but also on indirect measures of survival (which are likely to have more influence on population dynamics). Despite apparent overall population stability over the last 20 years, the national golden eagle population failed to meet the abundance target and only 3 of 16 regions where eagles have occupied territories since 1982 were considered to be in favourable condition. The key constraint preventing favourable condition being met was persecution, predominantly in some areas managed for grouse shooting. The advantages of a conservation framework over similar approaches, such as species action plans, are highlighted.     

Variations in the age of mates as an early warning signal of changes in population trends? The case of Bonelli's eagle in Andalusia

Methods to evaluate population trends have recently received particular attention because of the decreasing patterns shown by several species since the nineteenth century. We have studied demographic traits of the Bonelli's eagle (Hieraaetus fasciatus) population of Andalusia (southern Spain) for 20 years (1980-2000). This species is endangered and is suffering a rapid decline in most of its distribution range. Although our study population has remained stable for the last 10 years, both age at first breeding and productivity have decreased during the last 5 years. We propose that age at first breeding could be used as a potential early warning signal to detect possible future changes in population trends of long-lived species with deferred maturation age. In particular, in this species spatial variation in demographic traits could help to detect whether unnatural events such as shooting are decreasing age at first breeding and productivity. If occupation of territories by non-adults is caused by human persecution, conservation efforts in the particular case of Bonelli's eagle have to be directed to protect those breeding territories where an increase in the percentage of pairs containing at least one non-adult individual would be detected.     

The effects of persecution on age of breeding and territory occupation in golden eagles in Scotland

Several studies have documented negative effects of persecution by humans on the population dynamics of large birds of prey. Several areas have seen a reduction in persecution in modern times, but the demography of large raptors still makes these birds vulnerable to such effects. In a GIS analysis, utilising data collected during two national censuses of the golden eagle Aquila chrysaetos in Scotland and contemporary data on the distribution of poisoning incidents (a common method of raptor persecution in Scotland), we show that persecution was associated with a reduction in the age of first breeding, territory vacancies, and the use of territories by non-breeding immatures. Persecution was therefore reducing adult survival and distribution and was probably creating ecological traps that attract mobile immatures, increasing subadult mortality in birds that may originate from persecution-free areas. Mapping a form of land management unique to moors managed for red grouse Lagopus lagopus scoticus shooting, where most poisoning occurs, we also show similar associations between golden eagle demography and grouse moor management. As such, persecution may be having a major impact on the golden eagle population of Scotland, since it occurs over a wide area and at least in some areas has continued unchecked over many decades.     

Spatial association as an indicator of the potential for future interactions between wind energy developments and golden eagles Aquila chrysaetos in Scotland

Despite their environmental benefits in generating electricity without emission of ‘greenhouse’ gases, wind farms have attracted controversy with regard to their impacts on birds, especially golden eagles Aquila chrysaetos. Evidence from USA studies suggest eagle fatalities through collision with turbines may be the main potential impact whereas for breeding eagles in Scotland, displacement from wind farm areas (indirect habitat loss) may be the primary impact. In this study, we examined the co-occurrence potential for golden eagles and wind farms in Scotland by documenting the spatial association between wind farm proposals and breeding eagle territories and areas potentially suitable for non-breeding eagles. Although there were records for over 500 wind farm proposals at various stages of development, relatively few coincided with eagle territories (ca. 4% of territories had a proposal within 3 km of territory centre). Similarly, only 2% of habitat predicted to be suitable for non-breeding eagles overlapped with proposed or installed wind farm areas. Moreover, estimates of the potential for electricity generation from all wind farm proposals, with respect to government targets for renewable energy supplies, suggested most proposals were unlikely to be constructed. We conclude that in comparison with other constraints on Scotland’s golden eagles, notably persecution, wind farms should not represent a serious concern if best practice in planning their location and minimising their impact are maintained. Potential future regional pressures on breeding eagles from wind farms are highlighted, however, and uncertainty of impact with respect to displacement or collision fatalities requires continued scrutiny.     

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.     

An assessment of the olive ridley turtle (Lepidochelys olivacea) nesting population in Orissa, India

Olive ridley mass nesting events or ‘arribadas’ have been documented in Orissa, India since 1974. However, since standardised techniques have not been used to census turtles, actual population trends remain unknown. Herein, we summarise information on nesting populations in Orissa, using data from multiple sources to arrive at consensus estimates and to derive trends. We conducted a quantitative estimate of an arribada in March 1999, where nesting was estimated as ∼180,000 turtles by the strip transect method. Non-linear (quadratic) fits for arribada data from 1976-1999 and a recent decrease in the size of adults suggest a potential or imminent decline, consistent with fishery-related mortality of at least 90,000 turtles since 1994. Though statistical support for the recent decline is equivocal, efforts to reduce mortality and close monitoring of the population would be prudent. The absence of reliable data on which to base conservation action highlights an urgent need to train management personnel in data collection and estimation techniques for effective monitoring of status, threats and trends.     

Conservation status and limiting factors in the endangered population of Egyptian vulture (Neophron percnopterus) in the Canary Islands

Egyptian vulture populations have decreased sharply in the Western Palearctic; island populations are almost extinct in the Mediterranean and the Macaronesian regions. In the Canary archipelago, the species only survives in the islands of Fuerteventura and Lanzarote. During 1998-2001 we examined population parameters and evaluated some potential limiting factors for this isolated and sedentary population. The total population (breeding and non-breeding birds) was monitored annually. In addition, 26 fledglings and 33 immatures (<6 years old) and adult birds were captured for individual marking with plastic rings. Twenty-three/twenty-four occupied territories were located in the island and the total population estimated at around 130 birds. Breeding success was lower than recorded elsewhere in the species' distribution area: only 0.43 fledglings/pair/year were produced. Adult (>6 years old birds) and immature annual survival rates were similar, around 90%. Adult survival was lower than expected as territorial birds seem more susceptible to poisoning. Immature survival could be favoured by the existence of regular feeding places. Casualties from power lines was the main cause of mortality (12 cases during the study period). Blood sampling revealed high frequencies of lead poisoning: 13.5 and 2.7% of individuals showed sub-clinical and clinical intoxication levels, respectively, probably caused by the ingestion of lead shot. Priority conservation measures should be directed to reduce electrocution risks, illegal poisoning, and lead contamination. Population reinforcement with birds coming from other populations is not recommended as previous information reveals morphological and genetic differentiation of Canarian Egyptian vultures compared with continental populations.     

Decline in the distribution and abundance of flesh-footed shearwaters (Puffinus carneipes) on Lord Howe Island, Australia

The flesh-footed shearwater (Puffinus carneipes) is a migratory seabird that ranges widely across the Pacific and Indian Oceans. The principal breeding populations are in Australia and New Zealand. The only breeding site in eastern Australia is on Lord Howe Island. Despite it being afforded a high level of legislative protection, the population on Lord Howe Island has declined substantially during the last few decades. The total extent of nesting habitat in 2002 was 24.3 ha, a reduction of 13.4 ha (35.6%) since 1978. Loss of nesting habitat was associated with increased urbanisation, the adverse impact of which extended beyond the footprint of buildings and gardens. In 2002, overall burrow density was 0.123 per m² and the total number of burrows was estimated to be 29,853 ± 5867, a decline of about 19.0% since 1978. A substantial decline in burrow density was evident in the colony where loss of habitat to urbanisation had been greatest. In 2002, 58% of burrows were occupied by breeding birds, and the total population size was estimated to be 17,462 breeding pairs. Breeding success (the proportion of eggs that produced fledglings) was 50%, but was lowest in the most urbanised colony. To avert further declines in the population of flesh-footed shearwaters on Lord Howe Island major changes in land use practices, enforced through appropriate legislation, are needed, together with reductions in the level of seabird bycatch in fisheries operations and in the amount of plastics that litter the world’s oceans.     

The size and status of the population of southern sea lions Otaria flavescens in the Falkland Islands

Pup production of southern sea lions in the Falkland Islands was estimated to be 80,550 (total population ca. 380,000) in 1937, but by 1965 it had fallen to around 6000; a 93% reduction in under 30 years. We describe the results of an aerial survey of part of the breeding population in 1990 and comprehensive ground counts of the entire population in 1995 and 2003. Results indicate that the decline continued. In 1995, 63 breeding and 42 non-breeding groups were found. Pup production was estimated at 2034 pups; less than 2.7% of the 1930s estimate. All known and potential sites were revisited in 2003. 2747 pups were counted at 68 breeding sites, seven of which were new since 1995. Results indicate that between 1965 and 1990 the population reached a minimum of less than 1.5% of the 1937 population. Since then, pup production has increased at a rate of 8.5% p.a. between 1990 and 1995 and at 3.8% p.a. between 1995 and 2003.The Falklands' trajectory is similar to that of the adjacent Argentinian population. The causes of these declines are not clear. Around 44,000 sea lions were killed in the Falklands between 1935 and 1962, more than 500,000 were taken in Argentina in the same period. We present the results of a simple population model which suggests that, if sea lions migrated between the two areas, the combined hunt may explain the initial decline in the Falklands population. However, the continued decline after 1965 is as yet unexplained.     

A sensitivity analysis to guide research and management for Hector's dolphin

Hector's dolphin (Cephalorhynchus hectori) is an uncommon endemic of New Zealand which is suspected to be in decline due to entanglement mortality. However, uncertainty in available data has led to a dispute between the New Zealand Ministry of Fisheries and the New Zealand Department of Conservation over the status of this species. We use a density-dependent deterministic model to predict the future abundance and geographic distribution of Hector's dolphin under different scenarios of fisheries management. We then examine the sensitivity of this model to a number of parameters for which few or no data are available. We find that two populations of Hector's dolphins are predicted to decline in the future even when the most optimistic parameter estimates are used. The status of the third population is dependent upon the estimate of maximum annual population growth rate. Because of the dependence of final abundance estimates on the estimates of entanglement mortality rates and maximum population growth rate, research efforts should be concentrated on estimating these parameters.     

Using occupancy and population models to assess habitat conservation opportunities for an isolated carnivore population

An isolated population of the fisher (Martes pennanti) in the southern Sierra Nevada, California, is threatened by small size and habitat alteration from wildfires, fuels management, and other factors. We assessed the population’s status and conservation options for its habitat using a spatially explicit population model coupled with a fisher probability of occurrence model. The fisher occurrence model was selected from a family of generalized additive models (GAM) generated using numerous environmental variables and fisher detection–nondetection data collected at 228 survey arrays sampled repeatedly during 2002–2006. The selected GAM accounted for 69% of the Akaike weight using total above-ground biomass of trees, latitude-adjusted elevation, and annual precipitation averaged over a 5 km² moving window. We estimated equilibrium population sizes (or carrying capacities) within currently occupied areas, and identified likely population source, sink, and expansion areas, by simulating population processes for 20 years using different demographic rates, dispersal distances, and territory sizes. The population model assumed that demographic parameters of fishers scale in proportion to habitat quality as indexed by the calculated probability of fisher occurrence. Based on the most defensible range of parameter values, we estimate fisher carrying capacity at ∼125–250 adults in currently occupied areas. Population expansion into potential habitat in and north of Yosemite National Park has potential to increase population size, but this potential for expansion is predicted to be highly sensitive to mortality rates, which may be elevated in the northern portion of the occupied range by human influences, including roadkill and diseases carried by domestic cats and dogs.     

Potential biological removal of albatrosses and petrels with minimal demographic information

Seabirds such as albatrosses and petrels are frequently caught in longline and trawl fisheries, but limited demographic data for many species creates management challenges. A method for estimating the potential biological removal (the PBR method) for birds requires knowledge of adult survival, age at first breeding, a conservation goal, and the lower limit of a 60% confidence interval for the population size. For seabirds, usually only the number of breeding pairs is known, rather than the actual population size. This requires estimating the population size from the number of breeding pairs when important demographic variables, such as breeding success, juvenile survival, and the proportion of the adult population that engages in breeding, are unknown. In order to do this, a simple population model was built where some demographic parameters were known while others were constrained by considering plausible asymptotic estimates of the growth rate. While the median posterior population estimates are sensitive to the assumed population growth rate, the 20th percentile estimates are not. This allows the calculation of a modified PBR value that is based on the number of breeding pairs instead of the population size. For threatened albatross species, this suggests that human-caused mortalities should not exceed 1.5% of the number of breeding pairs, while for threatened petrel species, mortalities should be kept below 1.2% of the number of breeding pairs. The method is applied to 22 species and sub-species of albatrosses and petrels in New Zealand that are of management concern, of which at least 10 have suffered mortalities near or above these levels.     

Adaptive selection of foraging and nesting habitat by black kites (Milvus migrans) and its implications for conservation: a multi-scale approach

Black kites (Milvus migrans) are vulnerable and in decline within Europe. Here, we investigate selection of foraging and breeding habitat in a high-priority population in the Italian pre-Alps. Compared to a random distribution, kites foraged preferentially near water, over extensively managed grassland and within 1 km of nest-sites. Urban areas were positively selected near lakes but otherwise avoided. Foraging performance was higher over water than over terrestrial habitats. Kites nested on cliffs and trees and preferentially near water, far from paths and villages and in rugged and steep micro-sites. Tree-nests were located in the most mature tree in the stand. Productivity was positively related to the availability of water bodies. Therefore, food availability and human disturbance/persecution limited foraging and breeding performance. Guidelines to maintain or enhance current population levels include: (1) setting up reserves covering 10% of the areas within 1 km of large lakes; (2) converting current derelict coppice-woodland to high forest; and (3) enhancing subsidies for extensively managed grassland. Our results highlight the importance of cross-scale models integrating selection of foraging and breeding habitat and reinforce the importance of the spatial configuration of key resources for more realistic conservation management in mosaic landscapes.     

Evidence for food limitation in the declining hen harrier population on the Orkney Islands, Scotland

The hen harrier (Circus cyaneus) population on the Scottish Orkney Islands has declined dramatically since the end of the 1970s. We postulate that the cause of this decline was due to a reduction in the amount of available prey and predict that if this was the case the population would currently be limited by food. The evidence for this hypothesis is explored by examining the rates at which males deliver prey to their females in relation to breeding performance both among individuals within the declining population and also between this declining population and another, non-declining population in southern Scotland. Breeding performance within the Orkney population was related to male provisioning rates: males that provided more food to their females were more likely to initiate a breeding attempt and there was a tendency for males with the highest provisioning rates to breed with more females. Comparisons between the two populations revealed that harriers on Orkney had a lower breeding performance and also a lower rate of food provision. Changes in agriculture, in particular decreases in rough grazing and increases in sheep densities are thought to be the most likely cause for a reduction in food supply. Conservation measures should be aimed toward increasing the areas of rough grass habitat.     

Buzzards Buteo buteo and land use in the new forest,Hampshire, England

The population ecology of buzzards Buteo buteo in the New Forest, southern England, was studied from 1961 to 1982. Until 1973 the population was relatively constant (32–37 pairs), though density and productivity were low compared with elsewhere in Britain, probably as a result of low densities of mammalian prey resulting from intensive grazing by free-ranging cattle and ponies.In 1973 there was a dramatic fall in productivity: only two pairs fledged young. Comparing 1962–1971 with 1973–1982, the average number of breeding attempts declined by 35%, the average number of succesful attempts by 44%, and the average number of young fledged by 40%. The number of occupied territories fell from 33 in 1974 to 21 in 1982 (a 36% decline). It is hypothesised that this was the result of further declines in numbers of small rodents caused by greatly increased grazing pressure by large domestic herbivores, and their penetration into hitherto ungrazed areas.The number of successful breeding attempts varied widely from year to year after 1973. From 1968 the number of successful breeding pairs of buzzards fluctuated in relation to the amount of tree and shrub seed produced the previous autumn, and we suggest that from the early 1970s buzzards responded positively to early spring numbers of small rodents, which were in turn related to seed abundance in the previous autumn. It seems likely that before 1973 population levels of small rodents were sufficiently high even after a poor seed year, to form the basis of the buzzards' food in early spring, which is believed to be a critical period, and that only when population levels were reduced further by habitat deterioration caused by increased grazing pressure did fluctuations become critical to buzzards. The implications of intensive grazing for other predators of small rodents, and the implications for the management of the New Forest are briefly reviewed.     

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.     

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

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

Testing a mechanistic model. XI. The effects of time and of level of application on isotopically exchangeable phosphate

A model for the reaction of ions with soil was improved to permit time trends to be followed at a given level of phosphate addition. Difference equations were also developed to describe the rate of reaction of ions with both vacant sites and occupied sites, while diffusive penetration of the surface was occurring. The model was applied to data for the effects of time and of level of application on exchangeable phosphate. Many of the observed values for isotopically exchangeable phosphate could be well-described if it was assumed that equilibration of ³²P with surface sites was very rapid and this was followed by a diffusive penetration into the adsorbing particles. However, for short periods of contact between soil and ³²P, it was necessary to also take into account the rate of the reaction between ³²P and surface sites. This reaction was largely with vacant sites. Reaction with occupied sites–that is, true exchange–was unimportant. It is suggested that the electric potential of the surface may determine whether reaction is with occupied or vacant sites. In contrast to reaction of ³²P with occupied sites, reaction with vacant sites involves a net transfer of charge. Reaction with vacant sites would be slow if the potential was large and negative. It is shown that when reaction with vacant sites is slow, the proportion of previously added ³¹P recorded as exchangeable increases with level of addition of ³¹P. This may explain published observations of slow and non-linear exchange in some soils.     

Estimating the rate of quasi-extinction of the Australian grey nurse shark (Carcharias taurus) population using deterministic age- and stage-classified models

Grey nurse sharks off the east coast of Australia are listed nationally as “critically endangered” under Schedule 1 of the Environmental Protection and Biodiversity Conservation Act (1999) and may number no more than 300 in New South Wales and southern Queensland waters. They are an inshore, coastal dwelling species and were severely depleted by spearfishing in the 1960s. The population has continued to decline despite protection since 1984. Their life history (long-lived to 25+ years), late maturation (6-8 years), low fecundity (maximum 2 live young biennially), specific habitat requirements, limited inshore distribution, and small population size render them particularly vulnerable to extinction. We estimated the time to quasi-extinction (years elapsed for the population to consist of ?50 females) for the grey nurse shark population off the east coast of Australia based on current estimates of abundance and known anthropogenic rates of mortality. Estimated minimum population size was 300 as of 2002, and minimum anthropogenic mortality assessed from recovered carcasses was 12/year of which 75% were females. We modelled time to quasi-extinction using deterministic age- and stage-classified models for worst-, likely and best-case scenarios. Population size was estimated at 300 (worst), 1000 (likely) and 3000 (best). Anthropogenic mortality was added to the model assuming either all carcasses are being recovered (best), or conservatively, that only 50% are reported (realistic). Depending on model structure, if all carcasses are being reported, quasi-extinction times for worst-, likely and best-case scenarios range from 13 to 16 years, 84-98 years and 289-324 years, respectively. If under-reporting is occurring, time to quasi-extinction ranges from 6 to 8 years, 45-53 years and 173-200 years, respectively. In all scenarios modelled the grey nurse shark population will decline if no further steps are taken to remove anthropogenic sources of mortality. Because estimates of quasi-extinction rate depend on initial population size, and sensitivity analysis revealed that population rate of change was most sensitive to changes in the survival probability of the smallest length classes, obtaining precise estimates of abundance and annual survival of young females is critical.     

The conservation status of the spectacled petrel Procellaria conspicillata

The spectacled petrel Procellaria conspicillata is listed as critically endangered due to its small population size and ongoing mortality on long-lines. Spectacled petrels were counted in 2004, repeating a census made in 1999 at their sole breeding locality, Inaccessible Island. The 2004 survey took place earlier in the breeding season than the previous count, allowing for more robust estimates of burrow occupancy. During early incubation, birds responded to call playback at 69% of burrow entrances, but birds in at least 8% of burrows remained silent. Birds in shallow burrows were less likely to respond to playback than were those in deep burrows. Two repeat trials at 100 marked nests showed that at least 61% of apparently ‘unoccupied’ burrows were occupied on subsequent checks, resulting in an overall occupancy estimate of 91%. Occupancy was equally high in peripheral colonies. The apparent spatial extent of colonies increased slightly from 1999, and the estimate of total burrow numbers increased by 50%, from 5900 burrows in 1999 to 8900 in 2004. Validation surveys indicated that burrow numbers were underestimated (84 ± 3%) to the same extent as that in 1999 (85 ± 4%), and repeat checks of one colony where all nests were marked showed that even careful counts underestimated actual numbers of burrows by up to 10%. This suggests there are some 11-12,000 burrows, and assuming 90% occupancy, the adult population is likely to be at least 20,000 birds. The population has increased over the last five years, continuing the apparent recovery from a very small population size in the early 20th century. Despite this increase, demographic models indicate that the population remains at risk from relatively small increases in mortality, if mortality is determined primarily by fishing effort. Mitigation of long-line mortality remains the key conservation goal for this species.