The impacts of introduced House Mice on the breeding success of nesting seabirds on Gough Island

Invasive species are the main threat to island biodiversity; seabirds are particularly vulnerable and are one of the most threatened groups of birds. Gough Island, a UNESCO World Heritage Site in the South Atlantic Ocean, is an Important Bird and Biodiversity Area, and one of the most important seabird colonies globally. Invasive House Mice Mus musculus depredate eggs and chicks of most seabird species on the island, but the extent of their impact has not been quantified. We used field data and bootstrapped normal distributions to estimate breeding success and the number of surviving chicks for 10 seabird species on Gough Island, and compared estimates with those of analogous species from predator‐free islands. We examined the effects of season and nest‐site location on the breeding success of populations on Gough Island, predicting that the breeding success of Gough birds would be lower than that of analogues, particularly among small burrow‐nesting species. We also predicted that winter‐breeding species would exhibit lower breeding success than summer‐breeding species, because mice have fewer alternative food sources in winter; and below‐ground nesters would have lower breeding success than surface nesters, as below‐ground species are smaller so their chicks are easier prey for mice. We did indeed find that seabirds on Gough Island had low breeding success compared with analogues, losing an estimated 1 739 000 (1 467 000–2 116 000) eggs/chicks annually. Seven of the 10 focal species on Gough Island had particularly high chick mortality and may have been subject to intense mouse predation. Below‐ground and winter breeders had lower breeding success than surface‐ and summer‐breeders. MacGillivray's Prion Pachyptila macgillivrayi, Atlantic Petrel Pterodroma incerta and Tristan Albatross Diomedea dabbenena are endemic or near‐endemic to Gough Island and are likely to be driven to extinction if invasive mice are not removed.     

Disturbance and demography of Phylica arborea (Rhamnaceae) on the Tristan-Gough group of islands

Populations of Phylica arborea Thouars, an evergreen tree endemic to six southern oceanic islands, have been depleted by wood gathering and browsing on Amsterdam Island and Tristan da Cunha, and it was feared that introduced rodents were destroying seed and reducing regeneration on uninhabited Gough Island. A comparison of the demography of this species on four islands in the Tristan-Gough group suggests that rodents have little effect on seedling numberS. Regeneration seldom occurs in undisturbed P. Arborea groves: germination and seedling survival appears to be dependent on periodical removal of established trees and understorey vegetation by soil-slips and occasional fireS. Climatic and vegetation differences problably account for the less frequent regeneration at Gough Island relative to that at the Tristan islands.     

Estimating the home range and carrying capacity for takahe (

Predator-free offshore islands play an important role in the conservation of many of New Zealand's endemic species. Takahe (Porphyrio mantelli) have small populations established on four offshore islands and although hatching success is lower than that of the wild mainland population in Fiordland, juvenile and adult survival is high and populations are growing exponentially. Accurate estimates of home range size and potential carrying capacities are therefore essential for the future management of the population as a whole. The mean home range size of takahe pairs in one study population on Mana Island (217 ha) was 2.8 ± 1.9 ha. The island was assessed for current and maximum available area for takahe and the potential carrying capacity was estimated at 22—53 pairs. Current and maximum available areas were also used to calculate carrying capacities on each of three other islands using two different estimates of mean home range size for Maud Island (7—34 pairs) and Kapiti Island (5—33 pairs) and one estimate of home range size for Tiritiri Matangi Island (25 pairs). A model of the population growth of takahe on islands predicted that estimated carrying capacities would be reached between 1997 and 2009. The urgency of planning to make use of the considerable potential of island populations of takahe is stressed.     

Demographic history of a recent invasion of house mice on the isolated Island of Gough

Island populations provide natural laboratories for studying key contributors to evolutionary change, including natural selection, population size and the colonization of new environments. The demographic histories of island populations can be reconstructed from patterns of genetic diversity. House mice (Mus musculus) inhabit islands throughout the globe, making them an attractive system for studying island colonization from a genetic perspective. Gough Island, in the central South Atlantic Ocean, is one of the remotest islands in the world. House mice were introduced to Gough Island by sealers during the 19th century and display unusual phenotypes, including exceptionally large body size and carnivorous feeding behaviour. We describe genetic variation in Gough Island mice using mitochondrial sequences, nuclear sequences and microsatellites. Phylogenetic analysis of mitochondrial sequences suggested that Gough Island mice belong to Mus musculus domesticus, with the maternal lineage possibly originating in England or France. Cluster analyses of microsatellites revealed genetic membership for Gough Island mice in multiple coastal populations in Western Europe, suggesting admixed ancestry. Gough Island mice showed substantial reductions in mitochondrial and nuclear sequence variation and weak reductions in microsatellite diversity compared with Western European populations, consistent with a population bottleneck. Approximate Bayesian computation (ABC) estimated that mice recently colonized Gough Island (~100 years ago) and experienced a 98% reduction in population size followed by a rapid expansion. Our results indicate that the unusual phenotypes of Gough Island mice evolved rapidly, positioning these mice as useful models for understanding rapid phenotypic evolution.     

A global comparison of plant invasions on oceanic islands

Oceanic islands have long been considered to be particularly vulnerable to biotic invasions, and much research has focused on invasive plants on oceanic islands. However, findings from individual islands have rarely been compared between islands within or between biogeographic regions. We present in this study the most comprehensive, standardized dataset to date on the global distribution of invasive plant species in natural areas of oceanic islands. We compiled lists of moderate (5–25% cover) and dominant (>25% cover) invasive plant species for 30 island groups from four oceanic regions (Atlantic, Caribbean, Pacific, and Western Indian Ocean). To assess consistency of plant behaviour across island groups, we also recorded present but not invasive species in each island group.We tested the importance of different factors discussed in the literature in predicting the number of invasive plant species per island group, including island area and isolation, habitat diversity, native species diversity, and human development. Further we investigated whether particular invasive species are consistently and predictably invasive across island archipelagos or whether island-specific factors are more important than species traits in explaining the invasion success of particular species.We found in total 383 non-native spermatophyte plants that were invasive in natural areas on at least one of the 30 studied island groups, with between 3 and 74 invaders per island group. Of these invaders about 50% (181 species) were dominants or co-dominants of a habitat in at least one island group. An extrapolation from species accumulation curves across the 30 island groups indicates that the total current flora of invasive plants on oceanic islands at latitudes between c. 35°N and 35°S may eventually consist of 500–800 spermatophyte species, with 250–350 of these being dominant invaders in at least one island group. The number of invaders per island group was well predicted by a combination of human development (measured by the gross domestic product (GDP) per capita), habitat diversity (number of habitat types), island age, and oceanic region (87% of variation explained). Island area, latitude, isolation from continents, number of present, non-native species with a known invasion history, and native species richness were not retained as significant factors in the multivariate models.Among 259 invaders present in at least five island groups, only 9 species were dominant invaders in at least 50% of island groups where they were present. Most species were invasive only in one to a few island groups although they were typically present in many more island groups. Consequently, similarity between island groups was low for invader floras but considerably higher for introduced (but not necessarily invasive) species – especially in pairs of island groups that are spatially close or similar in latitude. Hence, for invasive plants of natural areas, biotic homogenization among oceanic islands may be driven by the recurrent deliberate human introduction of the same species to different islands, while post-introduction processes during establishment and spread in natural areas tend to reduce similarity in invader composition between oceanic islands. We discuss a number of possible mechanisms, including time lags, propagule pressure, local biotic and abiotic factors, invader community assembly history, and genotypic differences that may explain the inconsistent performance of particular invasive species in different island groups.     

Spatial scale and evolutionary history determine the degree of taxonomic homogenization across island bird assemblages

It is widely documented that human activities have elevated the extirpation of natural populations as well as the successful introduction to new areas of non-native species. These dual processes of introduction and extirpation can change the similarity of communities, but the direction and magnitude these changes take are likely to depend on the manner in which introductions and extirpations occur, the spatial scale at which the changes are measured, and the initial similarity of the communities before the human-induced drivers occurred. Here, we explore patterns of extirpation and introduction and their influence on the similarity of global oceanic island bird assemblages from four different Oceans (Atlantic, Caribbean, Indian, Pacific). We show that different historical patterns of introduction and extirpation have produced varying trends in compositional similarity both between islands within archipelagos and between islands across different archipelagos within the same ocean. Patterns of bird assemblage convergence (i.e. taxonomic homogenization) or divergence (i.e. taxonomic differentiation) among islands depended on the scale of examination, the evolutionary associations among species of the region, and the cultural history of human colonization. These factors are all likely to be leading to a series of multiple interacting processes that are shaping the complex compositional changes observed among global island bird faunas over time.     

Across island and continents, mammals are more successful invaders than birds (Reply)

Aim  The establishment success of exotic species is calculated as the fraction of introduced species that have become established, and invasion success is estimated as the fraction of established species that have spread significantly from their points of introduction. Records on species introductions are highly incomplete, so strong conclusions about the tens rule and invasibility of island and continents cannot be drawn.
Location  Global.
Methods  Using Jeschke (2008) as an example, we explain the issue of the inadequacy of data to draw conclusions about the tens rule and invasibility of island and continents.
Results  Lack of adequate data.
Main Conclusion  Jeschke (2008) probably overestimates the establishment and invasion success rates, so his conclusions about violation of the tens rule and that islands are not more susceptible to invasion than continents are misleading.     

Ant communities on small tropical islands: effects of island size and isolation are obscured by habitat disturbance and ‘tramp’ ant species

Aim Comparisons among islands offer an opportunity to study the effects of biotic and abiotic factors on small, replicated biological communities. Smaller population sizes on islands accelerate some ecological processes, which may decrease the time needed for perturbations to affect community composition. We surveyed ants on 18 small tropical islands to determine the effects of island size, isolation from the mainland, and habitat disturbance on ant community composition. Location Thousand Islands Archipelago (Indonesian name: Kepulauan Seribu) off Jakarta, West Java, Indonesia. Methods Ants were sampled from the soil surface, leaf litter and vegetation in all habitat types on each island. Island size, isolation from the mainland, and land‐use patterns were quantified using GIS software. The presence of settlements and of boat docks were used as indicators of anthropogenic disturbance. The richness of ant communities and non‐tramp ant species on each island were analysed in relation to the islands’ physical characteristics and indicators of human disturbance. Results Forty‐eight ant species from 5 subfamilies and 28 genera were recorded from the archipelago, and approximately 20% of the ant species were well‐known human‐commensal ‘tramp’ species. Islands with boat docks or human settlements had significantly more tramp species than did islands lacking these indicators of anthropogenic disturbance, and the diversity of non‐tramp species decreased with habitat disturbance. Main conclusions Human disturbance on islands in the Thousand Islands Archipelago promotes the introduction and/or establishment of tramp species. Tramp species affect the composition of insular ant communities, and expected biogeographical patterns of ant richness are masked. The island with the greatest estimated species richness and the greatest number of unique ant species, Rambut Island, is a forested bird sanctuary, highlighting the importance of protected areas in preserving the diversity of species‐rich invertebrate faunas.     

The role of introduced species in shaping the distribution and abundance of island reptiles

Summary Species interactions, as revealed by historical introductions of predators and competitors, affect population densities and sometimes result in extinctions of island reptiles. Mongoose introductions to Pacific islands have diminished the abundance of diurnal lizards and in some cases have led to extinctions. Through these population level effects, biogeographic patterns are produced, such as the reciprocal co-occurrence pattern seen with the tuatara and its predator, the Polynesian rat, and with the tropical gecko competitorsHemidactylus frenatus andLepidodactylus lugubris in urban habitats in the Pacific. Although competition has led to changes in abundance and has caused habitat displacement and reduced colonization success, extinctions of established reptile populations usually occur only as a result of predation.These introductions, along with many manipulative experiments, demonstrate that present day competition and predation are potent forces shaping community structure and geographic distributions. The human introduction of species to islands can be viewed as an acceleration of the natural processes of range expansion and colonization. The immediate biotic consequences of these natural processes should be of the same intensity as those of the human introductions. Coevolution may subsequently act to ameliorate these interactions and reduce the dynamical response of one species to the other. The role played by coevolution in mediating interactions between competitors and predator and prey is highlighted by the susceptibility of predator-naive endemic species to introduced predators and the invalidity of species-poor communities.     

A Characterization of the Non-indigenous Flora of the Azores Archipelago

The 9 Azores islands are located in the North Atlantic Ocean, about 1500 km from Europe. The vascular plant flora, consisting of 1002 taxa, was characterized using a database. Biogeographic and historic criteria were used to categorize the taxa as indigenous (31%) and non-indigenous (69%). The proportion of non-indigenous vascular plant taxa is higher than in other island ecosystems. This might have resulted from the removal of native vegetation and the introduction of many cultivated and ornamental plants, and from the relatively large extension of the agricultural landscape. The introduced plants were largely subcosmopolitan therophytes and hemicryptophytes, mainly Dicotyledoneae and Monocotyledoneae. The families with highest absolute contributions were similar to those in other areas of the world (Poaceae, Asteraceae, Fabaceae, Brassicaceae, Scrophulariaceae), but not all contributed as expected by their species richness worldwide. Many were introduced as ornamental or crop plants, but there were also many accidental introductions of weeds (about 55% of non-indigenous taxa). Considering the 9 islands, the percentage of introductions was positively correlated with human population density and island surface below 300 m, and negatively correlated with island surface allocated to natural areas.     

The island biogeography of exotic bird species

Aim   A recent upsurge of interest in the island biogeography of exotic species has followed from the argument that they may provide valuable information on the natural processes structuring island biotas. Here, we use data on the occurrence of exotic bird species across oceanic islands worldwide to demonstrate an alternative and previously untested hypothesis that these distributional patterns are a simple consequence of where humans have released such species, and hence of the number of species released.
Location   Islands around the world.
Methods   Statistical analysis of published information on the numbers of exotic bird species introduced to, and established on, islands around the world.
Results   Established exotic birds showed very similar species–area relationships to native species, but different species–isolation relationships. However, in both cases the relationship for established exotics simply mimicked that for the number of exotic bird species introduced. Exotic bird introductions scaled positively with human population size and island isolation, and islands that had seen more native species extinctions had had more exotic species released.
Main conclusion   The island biogeography of exotic birds is primarily a consequence of human, rather than natural, processes.     

Systematic review of the land snails of the Pitcairn Islands

The land snails (and semi-terrestrial molluscs) of the four islands that comprise the Pitcairn group are reviewed and the indigenous species illustrated. The strictly terrestrial molluscan faunas from the two atolls (Oeno and Ducie) are poor, like many other atolls in the Pacific. Each supports less than six species with wide geographical ranges. In contrast, the terrestrial molluscan fauna from Henderson Island, an uplifted atoll, is more diverse with at least 16 species belonging to seven families. Over half these taxa appear to be endemic, at least at the level of sub-species. Two species of semi-terrestrial molluscs have also been found on Henderson. Analyses of archaeological deposits in caves near the North Beach have revealed that at least a further six species of land snail formerly occurred on Henderson. The volcanic island of Pitcairn, the only island in the group still inhabited, supports the greatest number of terrestrial molluscs. Twenty-six species of land snail (and one semi-terrestrial pulmonate) were found living there during the recent expedition and a further three taxa were recognized amongst museum material. At least seven of these species are thought to be recent adventives and a further three are likely to have been prehistoric introductions. One Henderson (Georissa hendersoni) and three Pitcairn endemics (Pacificella filica, Sinployea pitcaimensis and Diastole tenuistriata) are formally described as new species. Some of the Pitcairn endemics occur in very restricted areas (less than a hectare) and it is important that measures should be taken to prevent the spread of invasive plants, such as rose-apple, that would threaten their survival.     

Introduced house mice as a conservation concern on Gough Island

House mice have previously been identified as a significant threat to both species and ecosystems on Southern Ocean islands. To date, these impacts have been quantified on several sub-Antarctic islands, but the role of house mice on more temperate islands is poorly known. On South Atlantic Gough Island, non-commensal house mice (Mus musculus L.) were probably introduced in the early 19th century and are now extremely abundant. To assess the likely impacts of mice on the fauna and flora at Gough Island we examined the diet of this population from September 1999 to July 2000 using conventional snap trapping techniques. The population has a single breeding season from September to March and mean body mass is notable in being amongst the largest reported for non-laboratory M. musculus. At low elevations (<250 m above sea level [a.s.l.]), avian carrion was the most prevalent dietary item during September and October. From November to February, plant material constituted the bulk of stomach contents and from March to July lumbricid worms were the most common food item. Indigenous invertebrate matter contributed little to mouse diet, independent of season. At altitudes greater than 500 m a.s.l., larvae of endemic brachypterous moths made up a significant proportion of stomach contents. In light of studies elsewhere, these data suggest that mouse predation may pose a significant threat to these species. However, it is not clear whether conservation action, such as an eradication attempt, is warranted. Further assessments of the impacts of mice are required, and in the interim every effort should be made to prevent introductions of other potentially harmful invasive species.     

Reassembling island ecosystems: the case of Lord Howe Island

Exotic species that invade remote islands, usually following human settlement, have had catastrophic effects on native biota. However, on islands it is increasingly feasible to eradicate key exotic species allowing extant native species to recover in situ or to return naturally. The practice of marooning threatened species on islands where the threat is absent, irrespective of whether the threatened species once occurred on the island, is well established. However, less focus has been given to the 'island' as the management unit on which to return extirpated species or related surrogates for extinct species. We use the example of Lord Howe Island as a case study to explore options for island restoration should the remaining critical exotic pests (rodents and perhaps owls in this case) be eradicated as planned. Lord Howe Island, in the south-west Pacific Ocean, is remote, biologically diverse, has a high degree of endemism, and was only discovered by humans in 1778. Consequently, the original and exotic biota and their interactions are all better known than for many islands with a more ancient human history. Two species of plants, nine terrestrial birds, one bat and at least four invertebrates have been lost from the island since 1778. One plant and two invertebrates could be returned as conspecifics. One plant and all the terrestrial birds that are extinct could be replaced by closely related species from elsewhere in the Pacific Ocean. Decisions on replacing extinct species with surrogates should be based on the taxonomic relatedness of the candidates for reintroduction: the same species before subspecies before genera, with functional replacement being a further filter on candidates that are not the same species. In our opinion, taxa with functional equivalence but without taxonomic relatedness would not be acceptable candidates for reintroduction.     

Genetic differentiation within and among island populations of the endangered plant Aster miyagii (Asteraceae), an endemic to the Ryukyu Islands

Genetic diversity was examined at 17 putative allozyme loci in 18 populations of the insular endemic plant Aster miyagii (Asteraceae). This species is geographically restricted to only three islands of the Ryukyu Islands and is on the federal list of threatened plants. Genetic differentiation within an island is small, suggesting that gene flow among populations on the same island is sufficiently large to prevent divergence. By contrast, genetic differentiation among islands is large, especially between Amamioshima Island and the other two islands, suggesting that gene flow between the islands is highly restricted. Two unique alleles are nearly fixed in populations on Amamioshima Island, which is the southernmost island of the three. Comparatively, genetic diversity is the smallest on Amamioshima Island. This genetic paucity on Amamioshima Island is probably a result of a population bottleneck at colonization or the small effective population size on this island. Genetic diversity at the species level of A. miyagii is larger than those of the species with a similar life history and of the congeneric widespread species, suggesting that the species has an old origin as an insular endemic species.     

Historical bird and terrestrial mammal extinction rates and causes

Aim Conservation of species is an ongoing concern. Location Worldwide. Methods We examined historical extinction rates for birds and mammals and contrasted island and continental extinctions. Australia was included as an island because of its isolation. Results Only six continental birds and three continental mammals were recorded in standard databases as going extinct since 1500 compared to 123 bird species and 58 mammal species on islands. Of the extinctions, 95% were on islands. On a per unit area basis, the extinction rate on islands was 177 times higher for mammals and 187 times higher for birds than on continents. The continental mammal extinction rate was between 0.89 and 7.4 times the background rate, whereas the island mammal extinction rate was between 82 and 702 times background. The continental bird extinction rate was between 0.69 and 5.9 times the background rate, whereas for islands it was between 98 and 844 times the background rate. Undocumented prehistoric extinctions, particularly on islands, amplify these trends. Island extinction rates are much higher than continental rates largely because of introductions of alien predators (including man) and diseases. Main conclusions Our analysis suggests that conservation strategies for birds and mammals on continents should not be based on island extinction rates and that on islands the key factor to enhance conservation is to alleviate pressures from uncontrolled hunting and predation.     

The island syndrome and population dynamics of introduced rats

The island syndrome predicts directional changes in the morphology and demography of insular vertebrates, due to changes in trophic complexity and migration rates caused by island size and isolation. However, the high rate of human-mediated species introductions to some islands also increases trophic complexity, and this will reduce the perceived insularity on any such island. We test four hypotheses on the role of increased trophic complexity on the island syndrome, using introduced black rats (Rattus rattus) on two isolated coral atolls in the Mozambique Channel. Europa Island has remained relatively pristine and insular, with few species introductions, whereas Juan de Nova Island has had many species introductions, including predators and competitors of rats, anthropogenically increasing its trophic complexity. In the most insular environments, the island syndrome is expected to generate increases in body size and densities of rodents but decreases in the rates of reproduction and population cycling. Morphology and reproduction were compared using linear regression and canonical discriminant analysis, while density and population cycling were compared using spatially explicit capture–recapture analysis. Results were compared to other insular black rat populations in the Mozambique Channel and were consistent with predictions from the island syndrome. The manifestation of an island syndrome in rodents depends upon the trophic composition of a community, and may not relate to island size alone when many species additions, such as invasions, have occurred. The differing patterns of rodent population dynamics on each island provide information for future rodent eradication operations.     

Long-term effects of feral goats (<Emphasis Type="Italic">Capra hircus</Emphasis>) on Mediterranean island communities: results from whole island manipulations

Islands exhibit disproportionally high biodiversity, however high levels of endemism and simplified food webs make their communities susceptible to invasive species. Introduced goats (Capra hircus), a generalist herbivore, are among the most harmful invasive species on islands. Concern about goat impacts on island communities have resulted in eradication programs, which have been generally implemented without comprehensive evaluation and monitoring. Unintended consequences may follow eradication, as grazing can have complex effects on island food webs. Using whole island manipulations, we evaluate the long-term, community-wide effects of goat herbivory, as well as their subsequent removal, in a system of 16 islands in the Aegean Sea (Greece) located within the Mediterranean biodiversity hotspot. Goat grazing on these islands is a major conservation concern, as these support endemic plant communities that have evolved in low herbivory conditions and lack appropriate defenses. We show that goat introductions lead to significant decreases in vegetation height, percent cover, and biomass but not to immediate plant species loss, as native island endemics are replaced with widespread generalist taxa carried in by the livestock. Additionally, goats contribute to the desertification of islands by initiating a long-term soil loss cycle that continues even after goats are removed; however, remaining soil structure and chemistry are not affected. Island arthropod populations do not appear to be significantly impacted by goat introduction or removal, except for a distinct increase in the order Diptera with goat presence. This study also reaffirms the role of seabirds in providing important marine subsidies, rich in nitrogen and phosphorus, to island food webs. Plant species diversity declines following goat removal, and vegetation cover returns only partially, as further recovery is being prevented by the long-term loss of soil. This suggests that following goat removal, island communities may require additional restoration efforts, including seabird reestablishment and reintroduction of extirpated plant populations, to promote island recovery.     

Can predation by invasive mice drive seabird extinctions?

The house mouse, Mus musculus, is one of the most widespread and well-studied invasive mammals on islands. It was thought to pose little risk to seabirds, but video evidence from Gough Island, South Atlantic Ocean shows house mice killing chicks of two IUCN-listed seabird species. Mouse-induced mortality in 2004 was a significant cause of extremely poor breeding success for Tristan albatrosses, Diomedea dabbenena (0.27 fledglings/pair), and Atlantic petrels, Pterodroma incerta (0.33). Population models show that these levels of predation are sufficient to cause population decreases. Unlike many other islands, mice are the only introduced mammals on Gough Island. However, restoration programmes to eradicate rats and other introduced mammals from islands are increasing the number of islands where mice are the sole alien mammals. If these mouse populations are released from the ecological effects of predators and competitors, they too may become predatory on seabird chicks.