Insular pest control within a metapopulation context

Introduced pests threaten many species and their control is generally beneficial for conservation, particularly on islands where complete eradication is possible. Unfortunately on ‘nearshore’ islands neighbouring source populations exist and unaided reinvasion is likely. Pest control programmes at these sites thus require a metapopulation context to adequately manage movements between source and sink populations. We investigated the ecology of introduced ship rats (Rattus rattus) on a nearshore island, and gene flow with adjacent mainland populations, in order to understand the metapopulation dynamics and relative levels of pest control required within the landscape. We sampled the entire population by trapping (n = 30), achieving eradication, and found a low rat density (3.2 ha⁻¹) indicative of a sink population. Seed and other plant material constituted the major dietary component of rats. Despite its proximity to mainland source populations, the island population was genetically distinct with reduced allelic diversity caused by a recent reinvasion founder effect. Genetic analyses also detected recent migrants between the populations. In contrast, two mainland populations separated by a similar distance displayed complete genetic mixing. The small water gap therefore provides a sufficient barrier to lower the migration rate to the island and delay reinvasion, which nonetheless eventually happens. In order to maintain nearshore islands pest-free, conservation management will require a metapopulation approach simultaneously focusing on both island and source population pest control.     

Rapid eradication of feral pigs (Sus scrofa) from Santa Cruz Island, California

Eradication of invasive alien species from islands is often necessary to protect native biota. The rapidity in which eradication projects are implemented can help reduce risk they will fail. We describe the eradication of feral pigs (Sus scrofa) from Santa Cruz Island, California, highlighting those control techniques that removed the most pigs and those that removed the last pigs. In 411 days, a total of 5036 pigs were removed from the 25,000-ha island. Before the eradication began, the island was fenced into five zones. Within each zone, the same general sequence of control methods was used: trapping (16% of dispatches in 1660 trap-nights); aerial shooting from a helicopter (77% of dispatches in 13,822 km of flight path); and then ground-based hunting with trained dogs (5% of dispatches in 1111 hunter-days). Sterilized adult pigs fitted with radio collars were subsequently used to aid in the location of surviving wild pigs and to monitor the success of the project. Female telemetered pigs were more effective than males at locating remaining wild pigs. Only 10% of the last 102 pigs (the last 20 or so present in each fenced zone) were dispatched as a result of being found with a telemetered pig, but telemetered animals were responsible for finding 43% of the very last pigs once normal hunting had ceased. The time taken to eradicate pigs on Santa Cruz Island was about half that taken on a neighbouring island of similar size (Santa Rosa Island) and 12 times as fast as that on Santiago Island (58,465 ha), Galapagos Islands. The deliberate sequencing of control methods, using first those that taught surviving pigs the least, and the intensive implementation of those methods, represent important advances in the practice of eradication and so biodiversity conservation.     

Is the Yelkouan shearwater Puffinus yelkouan threatened by low adult survival probabilities?

Many seabird species are experiencing population declines, with key factors being high adult mortality caused by fishery by-catch and predation by introduced predators on nesting islands. In the Mediterranean, both of these pressures are intensive and widespread. We studied the adult survival of an endemic Mediterranean seabird, the Yelkouan shearwater (Puffinus yelkouan), between 1969–1994 and 2007–2010 in Malta and between 2004–2010 in France using mark–recapture methods. Mean annual survival probabilities for breeding adults were below 0.9 for all colonies and periods. Between 1969–1994, annual survival for adults of unknown breeding status was on average 0.74 (95% confidence interval: 0.69–0.80) in Malta, possibly as a result of various human disturbances (including illegal shooting), light pollution and fisheries by-catch. Over the period 2004–2010, we found strong support for variation in adult survival probabilities between breeders and non-breeders, and islands with and without introduced predators in France. Survival probabilities for non-breeders (0.95, 0.81–1.0) appeared to be higher than for breeders (0.82, 0.70–0.94), but were imprecise partly due to low recapture probabilities. In Malta, we found evidence for heterogeneity in survival probabilities between two unknown groups (probably breeders and non-breeders), and seasonal variation in survival probability. Birds were more likely to survive the period including the peak breeding season than an equally long period during which they roam widely at sea. Although annual adult survival probability was still low (0.85, 0.58–1.0), colony protection measures appear to have reduced mortality at nesting cliffs. A population model indicated that colonies in France and Malta would currently require continuous immigration of 5–12 pairs per year to maintain stable populations. Our estimates of adult survival probabilities over the past four decades are consistent with overall population declines. Threats to Yelkouan shearwaters require immediate management actions to avoid ongoing population declines in the western Mediterranean.     

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.     

Populations of inshore serranids across the Canarian Archipelago: Relationships with human pressure and implications for conservation

We investigated spatio-temporal variability in the abundances and biomasses of four species of inshore serranids (the dusky grouper Ephinephelus marginatus, the island grouper Mycteroperca fusca, the painted comber Serranus scriba, and the blacktail comber S. atricauda) throughout the Canarian Archipelago (central-east Atlantic Ocean) with underwater visual transects. By means of a multiscaled sampling design spanning three orders of magnitude of spatial variability (from 10 s of meters among replicated 100 m² transects to 100 s of kilometres among islands) and four sampling periods, we related differences in the distributions of serranids to differences in the degree of human pressure, such as fishing intensity and human population. Differences in human pressure among islands provide the most parsimonious explanation for many of the consistent inter-island differences in the abundance and biomass of the analyzed species. Larger-bodied serranids (E. marginatus and M. fusca) are more vulnerable than the smaller species (S. scriba and S. atricauda). In fact, the larger, more vulnerable species have been almost completely extirpated from the most intensely fished islands. Our results show that the larger groupers have been overexploited throughout the Canary Islands, and highlight the urgent need for stringent management measures and better control of littoral reef fish resources.     

The impact of introduced predators, light-induced mortality of fledglings and poaching on the dynamics of the Cory’s shearwater (Calonectris diomedea) population from the Azores, northeastern subtropical Atlantic

Cory’s shearwater (Calonectris diomedea) is a procellariiform seabird which breeds in the Mediterranean and the north-eastern subtropical Atlantic, and which is considered “Vulnerable” in Europe due to recent declines at some localities. In the Azores archipelago (Atlantic), the introduction of mammalian predators by man has led to petrels being extirpated from the main islands, except for Cory’s shearwater. Currently, the Azorean population of Cory’s shearwaters represents 65% of the species’ world population. However, its dynamics remains unknown, although: (1) numbers might have declined by 43% between 1996 and 2001, (2) on the main islands, the young suffer mortality from introduced mammals, poachers, and urban lights upon fledging, and (3) at sea, the level of fishery mortality remains unknown.To fill this gap, we conducted a 7-year demographic survey on a mammal-free islet in the Azores to determine adult survival rate using capture-mark-recapture of the breeders and to estimate fecundity. We also assessed urban mortality using the data from the rescue campaigns annually conducted in the archipelago. Urban mortality concerned about 6% of fledglings, but its importance greatly varied among islands. When rescue campaigns occur, the rate might drop below 0.5%. Overall, our simulations concerning the next 100 years do not allow excluding a decrease in Cory’s shearwater numbers in the Azores without rescue campaigns (by 87% under the least favourable scenario). Rescue campaigns should only slow down the decline unless, simultaneously, lower competition for nests compared to Vila islet allows adults to breed almost every year on the main islands and juvenile survival exceeds a threshold value. Since adult survival rate was high (>0.93), an eventual decline of the Azorean population of Cory’s shearwaters would probably not result from fishery mortality of adults, but rather from poor fledgling productivity and perhaps also from low survival during the first year at sea.     

Seasonal fluctuation of invasive flatworm predation pressure on land snails: Implications for the range expansion and impacts of invasive species

Introduction of the snail-eating flatworm Platydemus manokwari has caused extinction and decline of native land snails on tropical and subtropical islands. As the factors influencing flatworm predation pressure on land snails remain unclear, I examined the effects of seasonal variation in flatworm predation pressure on land snail survival in the wild on a subtropical island. I also examined the feeding activities of P. manokwari under laboratory conditions. Survival rates of land snails experimentally placed on the forest floor for 7 days ranged from 0% to 100% among seasons on the oceanic island Chichijima [Ogasawara (Bonin) Islands]. More than 90% of snails were killed by P. manokwari within 7 days in the period from July to November, while less than 40% of snails were killed in other months. Snail mortality rate (0-100%) attributable to P. manokwari was positively correlated with mean temperature (17.1-27.3 °C) in the study area. Laboratory experiments showed that low temperature influenced snail survival and regulated feeding activity of P. manokwari. Laboratory experiments also suggested that high densities of P. manokwari may cause a rapid decline in snail survival. Therefore, ambient temperature and density of P. manokwari may regulate seasonal variations in predation pressure on land snails. Recent global warming may increase the probability of invasion and population establishment, and elevate the impacts of P. manokwari in temperate regions.     

Translocations of eight species of burrow-nesting seabirds (genera Pterodroma, Pelecanoides, Pachyptila and Puffinus: Family Procellariidae)

Development of seabird translocation techniques is required to meet species recovery objectives, to improve conservation status, and to restore ecological processes. During 1997-2008 we undertook translocation trials on eight petrel species of four genera within the New Zealand region: common diving petrel (Pelecanoides urinatrix), fairy prion (Pachyptila turtur), grey-faced petrel (Pterodroma macroptera gouldi), Pycroft’s petrel (Pterodroma pycrofti), Chatham petrel (Pterodroma axillaris), Chatham Island taiko (Magenta petrel; Pterodroma magentae), fluttering shearwater (Puffinus gavia), and Hutton’s shearwater (Puffinus huttoni). A total of 1791 chicks within 5 weeks of fledging were moved up to 240 km, placed in artificial burrows and hand-fed until they fledged. Of these, 1546 fledged, and so far at least 68 have returned as adults to the translocation sites. Most birds were crop-fed a puree based on tinned sardines and fresh water. This diet worked well for all species regardless of their typical natural diet (planktonic crustaceans, squid, or fish) with all species fledging above or close to mean natural fledging weights.     

Edge effects on nesting success of cavity-nesting birds in fragmented forests

Forest fragmentation leads to the creation of isolated forest patches and habitat edges with subsequent impact on forest-interior bird species. Although the effects of fragmentation and edge on avian nesting success are well documented for open cup-nesting species in eastern deciduous forests in North America, it is unclear whether these effects are common for all birds nesting in predominantly forested landscapes. In particular, edge effects on nesting success of cavity-nesting birds are poorly understood. Using natural cavity nests, we examined nesting success of four species of cavity-nesting birds (two nonexcavators and two excavators), the yellow-rumped flycatcher (Ficedula zanthopygia), the great tit (Parus major), the great spotted woodpecker (Picoides major), and the grey-faced woodpecker (Picus canus) in relation to forest edges in Zuojia Nature Reserve, Jilin province, northeastern China. Our primary objective was to assess whether distance to the edge of agricultural lands was related to nesting success for cavity-nesting birds in fragmented forests. A total of 439 natural cavity nests of the four species were located and monitored during four breeding seasons. Probability of nest success was influenced by distance to forest edge for nonexcavators, but not for excavators. The rate of nesting success of the two nonexcavators, yellow-rumped flycatcher and the great tit, increased with distance from the edges. For all cavity nests, nesting success was 0.43 at 0-100 m, 0.56 at 101-200 m, 0.68 at 201-300 m, 0.61 at 301-400 m, 0.77 at 401-500 m from the edges. Nesting success ranged from 0.57 for the yellow-rumped flycatcher to 0.89 for the Grey-faced Woodpecker. Failed nests were often occupied by nest-site competitors (accounting for 68%). However, predation only accounted for 20% of all nest failures. Our results suggest that negative edge effects do exist for some cavity-nesting birds, especially for nonexcavator species.     

Protecting islands from pest invasion: optimal allocation of biosecurity resources between quarantine and surveillance

Removing pests from islands, and then keeping them pest free, is a common management goal. Given that goal we face a decision: how much effort should we invest in quarantine to reduce the risk of a pest arriving vs. surveillance, looking for the pest on the island with the view of eradicating it before it gets out of control. We use models of an island under threat of invasion by a pest (animal, plant or disease) and a cost minimisation approach to optimally allocate management resources between quarantine and surveillance. In the optimal allocations joint investment in both quarantine and surveillance is uncommon. Investment in quarantine is optimal if quarantine is more effective than surveillance or if large costs associated with pest impact and eradication are incurred at low pest density. Investment in quarantine is also favoured as our ability to eradicate a pest declines. Surveillance is optimal if it is considerably more cost-effective than quarantine and we can generate significant savings through early detection of the pest population. We illustrate how theses models are useful ways to examine these trade-offs by applying the model to the prevention of black rat (Rattus rattus) invasion on Barrow Island, Western Australia. Our model predicts an optimal strategy different to the management strategy currently being used on the island. We suggest that this is due to a risk-averse tendency in managers and the difficulty of estimating costs that combine management, environmental and social factors.     

Maximum foraging ranges in solitary bees: only few individuals have the capability to cover long foraging distances

To preserve populations of endangered bee species, sound knowledge of their maximum foraging distance between nest and host plants is crucial. Previous investigations predicted maximum foraging distances of 100-200 m for small bee species and up to 1100 m for very large species based on mainly indirect methods. The present study applied a new and direct approach to experimentally investigate maximum foraging distances in solitary bees. One endangered and two common species of different body sizes, all of which restrict pollen foraging to a single plant genus, were established in a landscape lacking their specific host plants. Females were forced to collect pollen on potted host plants that were successively placed in increasing distance from fixed nesting stands. The maximum foraging distance recorded for the small Hylaeus punctulatissimus was 1100 m, for the medium sized Chelostoma rapunculi 1275 m and for the large Hoplitis adunca 1400 m, indicating that maximum foraging distances at species level have been underestimated. However, the capability to use resources on such a large spatial scale applied only to a small percentage of individuals as 50% of the females of H. punctulatissimus and H. adunca did not forage at distances longer than 100-225 m and 300 m, respectively. This finding suggests that a close neighbourhood of nesting and foraging habitat within few hundred meters is crucial to maintain populations of these species, and that threshold distances at which half of the population discontinues foraging are a more meaningful parameter for conservation practice than the species specific maximum foraging distances.     

Evaluating alternative rodenticides for island conservation: roof rat eradication from the San Jorge Islands, Mexico

Introduced commensal rats (Rattus spp.) are a major contributor to the extinction and endangerment of island plants and animals. The use of the toxin brodifacoum to completely eradicate rats from islands is a powerful conservation tool. However, brodifacoum is toxic to animals other than rats and on some islands its use may not be feasible without prohibitively expensive mitigation. As part of a regional conservation program, we experimentally tested brodifacoum and two less toxic rodenticides, diphacinone and cholecalciferol, in eradicating Rattus rattus from three small islands in the northern Gulf of California, Mexico. All three rodenticides were successful in eradicating rats, suggesting that the less toxic diphacinone and cholecalciferol may be useful alternatives to brodifacoum for some island eradication programs. However, the choice of rodenticide must be balanced between efficacy and the risks to non-target species. Applied field research is needed on less toxic rodenticides, as well as improving palatability of baits. This may prove invaluable in preventing extinctions and in restoring larger and more diverse island ecosystems.     

Eradication of rock pigeons,Columba livia,from the Galápagos Islands

Alien species can negatively affect global biodiversity, especially on islands. Significant advances in methods for eradicating mammals from islands have been achieved. In contrast, development of methods for eradicating birds from islands has lagged and few islands have had alien birds successfully eradicated. We report on a 7 year campaign to remove rock pigeons from the Galapagos Islands. To date this is the largest successful eradication of an alien bird from an island system and the only eradication of rock pigeons from an island. Multiple methods were tested and used, including alpha-chloralose, however, shooting with a high-powered air rifle was the most efficient removal technique. Incorporating the support of the community and local agencies into the campaign was critical to the success of the project.     

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.     

The conservation status of Townsend's shearwater Puffinus auricularis auricularis

Townsend's shearwater (Puffinus auricularis auricularis) is an endangered seabird endemic to the Revillagigedo Archipelago. It nested on Socorro, Clarion, and San Benedicto Islands. It was extirpated by the Barcena volcano on San Benedicto in 1952, and there are no recent indications of nesting. Introduced mammals—pigs and rabbits—preyed on them and destroyed habitat at Clarion; shearwaters were extirpated by 1988, and no breeding attempts have been reported since. Our results confirm that Socorro holds the last breeding grounds. We found breeding colonies above 800 m and a minimum population of 1100 individuals. This represents a significant reduction in distribution and population size. Intensive cat predation at Socorro could potentially kill ca. 350 females per season, and sheep progressively destroy nesting areas. Population projections suggest that demographic instability could occur in less than 100 years under severe predation and habitat degradation. Only low predation rates would allow population persistence for more than 150 years in spite of a declining population. Thus, the immediate eradication of all introduced mammals is necessary to prevent the extinction of this seabird.     

Aerial surveying of the world’s largest leatherback turtle rookery: A more effective methodology for large-scale monitoring

For many marine megavertebrate species it is challenging to derive population estimates and knowledge on habitat use needed to inform conservation planning. For marine turtles, the logistics required to undertake comprehensive ground-based censuses, across wide spatial and temporal scales, are often insurmountable. This frequently leads to an approach where a limited number of index nesting beaches are monitored in great detail by foot. In this study we use nationwide aerial surveying interfaced with ground assessments across three seasons of leatherback turtle nesting in Gabon (Equatorial West Africa), highlighting the importance of a synoptic approach to marine turtle monitoring. These surveys allow the first complete population assessment of this nesting aggregation to be made, identifying it as the world’s largest for the species (36,185-126,480 clutches, approximating to 5865-20,499 breeding females per annum and a total estimate of 15,730 to 41,373 breeding females). Our approach also serendipitously provides insights into the spatial appropriateness of Gabon’s protected areas network, for example (mean ± 1SD) 79 ± 6% (range 67-86%) of leatherback turtle activities recorded during aerial surveys (n = 8) occurred within protected areas (345 km, 58%, of surveyed coastline). We identify and discuss sources of potential error in estimating total nesting effort from aerial surveying techniques and show that interannual variation in nesting is considerable, which has implications for the detection of statistically significant changes in population size. Despite its relative costliness per day, aerial surveying can play an important role in providing estimates of relative population abundance of large vertebrates dispersed over extensive areas. Furthermore, it can provide data on habitat use and deliver real-time information on the spatial efficacy of protected area networks.     

Disperser communities and legacies of goat grazing determine forest succession on the remote Three Kings Islands, New Zealand

Many remote islands are degraded as a result of deforestation and browsing of vegetation by introduced goats. Goat eradication is therefore a focus for island restoration, but there are few long-term records of change on islands after eradications. In 1946, three permanent plots were established immediately after goats were eradicated from Great Island (Manawa Tawhi), 60 km from northern New Zealand, and provide a 57-year record of change across a sequence of forest succession. Since 1946, the native and non-native bird communities that disperse 75% of the woody flora have increased from six to eight species and bird-dispersed woody plants in plots have increased from 7 to 11 species. After 1946, palatable trees were recruited in the plots. Unpalatable understorey sedges, present when goats were abundant, have persisted and may impede tree seedling establishment. Of the bird-dispersed woody plant species, 41% occur in the plots compared with 67% of the non-bird-dispersed species. Large-seeded species were unable to germinate away from parents until native pigeons Hemiphaga novaeseelandiae were present during the last decade. Forest succession is a consequence of interactions between the legacy of goat grazing and current disperser communities. Survival of seed-limited rare plants is not guaranteed in these circumstances. Although non-native goats no longer influence succession directly, non-native birds have been and remain important components of the disperser community. Our study supports the view that a whole-ecosystem understanding of the interactions between native and non-native species is needed to predict the consequences of eradications on islands worldwide.     

MtDNA genetic diversity and population history of a dwindling raptorial bird, the red kite (Milvus milvus)

The red kite (Milvus milvus) occurs in a relatively small area in the southwestern Palearctic region, with population strongholds in Central Europe. Following strong human persecutions at the beginning of the 20th century, populations have receded, particularly in peripheral areas and islands. In order to describe and compare levels of genetic diversity and phylogeographic patterns throughout its entire distribution in Europe, sequence variation of a 357 bps part of the mitochondrial DNA control region was assessed in eight populations and 105 individuals. Overall, results indicate that population declines have affected red kite mtDNA variation. We found low levels of genetic diversity (values of nucleotide diversity ranging from 0 in Majorca island to 0.0062 in Central Europe), with only 10 distinct haplotypes, separated by low levels of genetic divergence (mean sequence divergence = 0.75%). Highest haplotype and nucleotide diversities match with demographic expectations, and were found in Central European and Central Spanish samples, where present strongholds occur, and lowest values in the declining southern Spanish and insular samples. Φst estimates indicated moderate gene flow between populations. Phylogeographic patterns and mismatch distributions analyses suggest central European regions may have been colonized from southern glacial refugia (in the Italian or Iberian peninsulas). Interspecific phylogenetic comparisons and divergence date estimates indicated the genetic split between the red kite and its closely related species, the black kite (Milvus migrans), might be relatively recent. The low level of genetic variation found in the red kite mitochondrial control region, compared to the black kite, is likely the result of relatively recent divergence (associated with founder events), successive bottlenecks and small population sizes. As there are several ongoing projects aimed at reinforcing populations in countries such as the United Kingdom, Italy or Spain, our results may prove useful for the genetic management of the species.     

The effects of heat stress, predation risk and parental investment on Malaysian plover nest return times following a human disturbance

Waders leave nests and conduct distractive displays when approached by people. The time taken for waders to return to nests depends on numerous factors that affect the costs and benefits of incubation and anti-predator behavior. Understanding this trade-off may help assess the reproductive consequences of different nest return times and identify variables to consider in breeding disturbance studies. We subjected 73 Malaysian plover (Charadrius peronii) nests to standardized human disturbances and an analysis of covariance was used to determine how weather, time of day, embryonic age, weeks into breeding season and nest attendance (proportion of time adults incubated nests) influence nest return times. Egg temperatures were estimated using a regression model that predicted the temperature inside unshaded eggs from air temperature, cloud cover and time of day (r² = 0.88). We assessed the relationship between nest return times and hatch success. Plovers returned to nests faster at higher modeled egg temperature (P = 0.010), in the morning (P = 0.003), if they had younger clutches (P = 0.038), and if they had high nest attendance prior to the disturbance (P = 0.015). Pairs that returned to nests faster had lower hatch success (P = 0.021). This may be because pairs that spend more time distracting humans may also do so for predators. These results suggest that short nest return times may not indicate low fitness costs of disturbance. The thermal and predation environment in addition to nest return times should be taken into account when assessing the deleterious effects of human disturbance.