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.     

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.     

Review of impacts of the introduced house mouse on islands in the Southern Ocean: are mice equivalent to rats?

Research on the impacts of house mice Mus musculus introduced to islands is patchy across most of the species’ global range, except on islands of the Southern Ocean. Here we review mouse impacts on Southern Ocean islands’ plants, invertebrates, land birds and seabirds, and describe the kinds of effects that can be expected elsewhere. A key finding is that where mice occur as part of a complex of invasive mammals, especially other rodents, their densities appear to be suppressed and rat-like impacts have not been reported. Where mice are the only introduced mammal, a greater range of native biota is impacted and the impacts are most severe, and include the only examples of predation on seabird eggs and chicks. Thus mice can have devastating, irreversible and ecosystem-changing effects on islands, impacts typically associated with introduced rats Rattus spp. Island restoration projects should routinely include mouse eradication or manage mouse impacts.     

Rats are not the only introduced rodents producing ecosystem impacts on islands

In addition to rats, nutria (Myocastor coypus) and the North American beaver (Castor canadensis) have certainly caused damage at an ecosystem level when introduced to islands, in both cases primarily by ecosystem engineering. Of other introduced rodents successfully established on islands, the gray squirrel (Sciurus carolinensis) may be in the process of damaging entire forest ecosystems, particularly by bark-stripping. Though introduced muskrats (Ondatra zibethicus) have had ecosystem-level impacts in continental Europe, their impact on islands worldwide to which they have been introduced has been very limited. The North American red squirrel (Tamiasciurus hudsonicus) and Barbary ground squirrel (Atlantoxerus getulus) have each had substantial impacts when introduced to particular islands, but for neither species have these impacts yet been demonstrated to spread through an entire ecosystem. Introduced house mice (Mus musculus) may well generate ecosystem impacts on remote islands lacking rats, and it is possible that explosions of house mice on islands after rat eradication, a common occurrence, will lead in some instances to ecosystem impacts.     

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.     

Review article: The catastrophic impact of invasive mammalian predators on birds of the UK Overseas Territories: a review and synthesis

The UK has sovereignty over 16 Overseas Territories, which hold some of the world’s great seabird colonies and collectively support more endemic and globally threatened bird species than the whole of mainland Europe. Invasive alien mammalian predators have spread throughout most of the Territories, primarily since European expansion in the 16th century. Here we review and synthesize the scale of their impacts, historical and current, actions to reduce and reverse these impacts, and priorities for conservation. Mammalian predators have caused a catastrophic wave of extinctions and reductions in seabird colony size that mark the UKOTs as a major centre of global extinction. Mammal‐induced declines of threatened endemics and seabird colonies continue, with four Critically Endangered endemics on Gough Island (Tristan da Cunha), St Helena and Montserrat directly threatened by invasive alien House Mice Mus musculus, Feral Cats Felis catus and rats Rattus spp. Action to reduce these threats and restore islands has been modest in comparison with other developed countries, although some notable successes have occurred and a large number of ambitious eradication and conservation plans are in preparation. Priority islands for conservation action against mammalian predators include Gough (which according to one published prioritization scheme is the highest‐ranked island in the world for mammal eradication), St Helena and Montserrat, but also on Tristan da Cunha, Pitcairn and the Falkland Islands. Technical, financial and political will is required to push forward and fund the eradication of invasive mammalian predators on these islands, which would significantly reduce extinction risk for a number of globally threatened species.     

The effects of mice on stoats in southern beech forests

Introduced stoats (Mustela erminea) are important invasive predators in southern beech (Nothofagus sp.) forests in New Zealand. In these forests, one of their primary prey species – introduced house mice (Mus musculus), fluctuate dramatically between years, driven by the irregular heavy seed‐fall (masting) of the beech trees. We examined the effects of mice on stoats in this system by comparing the weights, age structure and population densities of stoats caught on two large islands in Fiordland, New Zealand – one that has mice (Resolution Island) and one that does not (Secretary Island). On Resolution Island, the stoat population showed a history of recruitment spikes and troughs linked to beech masting, whereas the Secretary Island population had more constant recruitment, indicating that rodents are probably the primary cause for the ‘boom and bust’ population cycle of stoats in beech forests. Resolutions Island stoats were 10% heavier on average than Secretary Island stoats, supporting the hypothesis that the availability of larger prey (mice verses wētā) leads to larger stoats. Beech masting years on this island were also correlated with a higher weight for stoats born in the year of the masting event. The detailed demographic information on the stoat populations of these two islands supports previously suggested interactions among mice, stoats and beech masting. These interactions may have important consequences for the endemic species that interact with fluctuating populations of mice and stoats.     

Vagrant Antarctic fur seals at the Tristan da Cunha Islands

Antarctic fur seals Arctocephalus gazella mainly breed at islands south of the Antarctic Polar Front, but stragglers occasionally occur farther north, with records from Gough Island (40°S, 10°W) in the central South Atlantic Ocean in October/November 2005 and September/October 2009. We report the first record from Tristan da Cunha (37°S, 12°W) in September 2013, and another individual that was observed at Gough Island. Both individuals were lean, lethargic subadult males that were present before the onset of the breeding (pupping) season of the resident populations of subantarctic fur seals Arctocephalus tropicalis.     

The diet of feral cats at New Island, Falkland Islands, and impact on breeding seabirds

We studied the diet of feral cats (Felis catus) on New Island, Falkland Islands, through the analysis of 373 scats collected during the austral summers of 2004/2005 and 2005/2006. The most frequent prey were three introduced mammals (house mice Mus musculus, ship rats Rattus rattus and rabbits Sylvilagus sp.) and the thin-billed prion Pachyptila belcheri (each season present on ca. 21% of the analysed scats). These represent the first systematic data on feral cat diet for the Falklands. A simple bioenergetics model suggests that cats could be eating in the region of 1,500–11,000 prions per season, representing <1% of the local adult and subadult population. Predation on other seabirds nesting on New Island (several penguin species, albatrosses and cormorants) was unimportant, with the possible exception of white-chinned petrels Procellaria aequinoctialis, which nest locally in very small numbers. For each prion eaten, cats were estimated to have killed 1.1–1.9 ship rats during the summer season, and probably more in autumn and winter. Knowing that ship rats are prion predators, it is conceivable that, on the whole, cats are having a positive impact on the prion population, a scenario predicted by general theoretical models. Thus, considering the available information, we would not recommend the implementation of any eradication programme on New Island that would target cats in isolation. Nevertheless, it would be prudent to consider some local action targeting cats and rats around the small New Island white-chinned petrel colony.     

Diet of the house mouse (Mus musculus) on Guillou Island, Kerguelen archipelago, Subantarctic

The diet of the house mouse (Mus musculus domesticus), introduced to the Kerguelen archipelago in the 1800s, was studied at monthly intervals from August 1997 to July 1998 in the closed communities of Acaena magellanica, the main habitat of mice on Guillou Island. The analysis of 291 stomach contents showed that this opportunistic rodent included a variety of items in its diet: earthworms (Dendrodrilus rubidus tenuis, Microscolex kerguelensis), caterpillars of a flightless moth (Pringleophaga kerguelensis), weevil adults and larvae (Ectemnorrhinus spp.), seeds of Acaena magellanica, and floral parts of dandelion (Taraxacum officinale). The animal prey were dominant in its diet all year round, except in summer. Based on the presence of chaetae in the stomach contents, our results show that earthworms are an important prey for the house mouse at Kerguelen. The consequences of these food habits for the invertebrate communities of the subantarctic islands are discussed.     

Evolution of an invasive rodent on an archipelago as revealed by molar shape analysis: the house mouse in the Canary Islands

Aim The aim of this paper is to identify the patterns in the morphological differentiation in Canary Island mice, based on fossil and modern samples. In order to achieve this, the mouse species present on the archipelago were first compared with a set of continental mice. The differences between the continental and Canary Island samples, and among the Canary Island samples, provide insights into the processes of colonization and the subsequent insular evolution. Location Canary archipelago. Methods An outline analysis based on Fourier transformation was used to quantify shape differences between lower molars. Together with the fossil and modern Canary Island samples, a reference set of genotyped continental populations of the commensal Mus musculus and the wild Mus spretus was used for comparison. Results The morphometric analysis showed that all the mouse specimens from the Canary Islands and Cape Verde belonged to Mus musculus domesticus. Lower molars of extant mice from La Gomera, El Hierro, Gran Canaria, Tenerife, and to a lesser degree from Lanzarote, were similar to those of genotyped M. m. domesticus from the continent, while teeth of extant mice from Fuerteventura were more divergent. Fossil mice from Fuerteventura were very similar to the extant representatives on this island, and similar to the fossil mice on the nearby islands of Lobos and La Graciosa. Main conclusions The mouse present on the Canary archipelago has been identified as the house mouse M. m. domesticus. Based on the shape of the lower molar, the Canary Island mice are divergent from the continental ones, but the degree of divergence varies with the geography of the archipelago. Overall, populations from eastern islands are more divergent from the continental mice than populations from western ones. Fossil populations indicate that this situation was established several centuries ago. Two main factors may have contributed to this pattern: the appearance of different types of environment on the islands since the successful settlement of the mouse, and/or the number of subsequent introductions of continental individuals via shipping.     

The diet of the King Penguin Aptenodytes patagonicus at Macquarie Island

The diet of King Penguins Aptenodytes patagonicus at Macquarie Island was studied between November 1984 and November 1985 based on stomach flushed samples (obtaining 93% of the total stomach content) from ten birds each month. The mean stomach content mass of the 118 samples was 923 0 g. Percentage by number, percentage by weight and dietary coefficient analysis all showed the main prey of the penguins to be myctophid lantern fish of the species Electrona carlsbergi and Krefftichthys anderssoni. Juvenile fish of both species were eaten from December to July, and adults in August and September. Cephalopods were relatively unimportant in contrast to previous indications. The amount of food brought ashore and the composition of the diet varied over the year, with K. anderssoni the dominant food in all but the winter months when E. carlsbergi replaced it as the principal food item.     

Introduced mammals coexist with seabirds at New Island,Falkland Islands: abundance,habitat preferences,and stable isotope analysis of diet

The largest known colony of Thin-billed prions Pachyptila belcheri has been coexisting with introduced mammals for more than 100 years. Three of the introduced mammals are potential predators of adults, eggs and chicks, namely ship rats Rattus rattus, house mice Mus musculus and feral cats Felis catus. We here determine habitat preferences over three seasons and dietary patterns of the unique set of introduced predators at New Island, Falkland Islands, with emphasis on the ship rats. Our study highlights spatial and temporal differences in the levels of interaction between predators and native seabirds. Rats and mice had a preference for areas providing cover in the form of the native tussac grass Parodiochloa flabellata or introduced gorse Ulex europaeus. Their diet differed markedly between areas, over the season and between age groups in rats. During the incubation period of the prions in November–December, ship rats had mixed diets, composed mainly of plants and mammals, while only 3% of rats had ingested birds. The proportion of ingested birds, including scavenged, increased in the prion chick-rearing period, when 60% of the rats consumed prions. We used δ¹³C and δ¹⁵N to compare the importance of marine-derived food between mammal species and individuals, and found that rats in all but one area took diet of partly marine origin, prions being the most frequently encountered marine food. Most house mice at New Island mainly had terrestrial diet. The stable isotope analysis of tissues with different turnover times indicated that individual rats and mice were consistent in their diet over weeks, but opportunistic in the short term. Some individuals (12% of rats and 7% of mice) were highly specialized in marine-derived food. According to the isotope ratios in a small sample of cat faeces, rodents and rabbits were the chief prey of cats at New Island. Although some individuals of all three predators supplement their terrestrial diet with marine-derived food, the current impact of predation by mammals on the large population of Thin-billed prions at New Island appears small due to a number of factors, including the small size of rodent populations and restriction mainly to small areas providing cover. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Ingrid Schenk: deceased     

A molecular characterization of the charismatic Faroe house mouse

Faroe house mice are a ‘classic’ system of rapid and dramatic morphological divergence highlighted by J. S. Huxley during the development of the Modern Synthesis. In the present study, we characterize these charismatic mice using modern molecular techniques, examining specimens from all Faroe islands occupied by mice. The aims were to classify the mice within the modern house mouse taxonomy (i.e. as either Mus musculus domesticus or Mus musculus musculus) using four molecular markers and a morphological feature, and to examine the genetic diversity and possible routes of colonization using mitochondrial (mt) control region DNA sequences and microsatellite data (15 loci). Mice on the most remote islands were characterized as M. m. domesticus and exhibited exceptionally low genetic diversity, whereas those on better connected islands were more genetically diverse and had both M. m. musculus and M. m. domesticus genetic elements, including one population which was morphologically M. m. musculus‐like. The mtDNA data indicate that the majority of the mice had their origins in south‐western Norway (or possibly southern Denmark/northern Germany), and probably arrived with the Vikings, earlier than suggested by Huxley. The M. m. musculus genetic component appears to derive from recent mouse immigration from Denmark. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 471–482.     

Recombination rate variation in mice from an isolated island

Recombination rate is a heritable trait that varies among individuals. Despite the major impact of recombination rate on patterns of genetic diversity and the efficacy of selection, natural variation in this phenotype remains poorly characterized. We present a comparison of genetic maps, sampling 1212 meioses, from a unique population of wild house mice (Mus musculus domesticus) that recently colonized remote Gough Island. Crosses to a mainland reference strain (WSB/EiJ) reveal pervasive variation in recombination rate among Gough Island mice, including subchromosomal intervals spanning up to 28% of the genome. In spite of this high level of polymorphism, the genomewide recombination rate does not significantly vary. In general, we find that recombination rate varies more when measured in smaller genomic intervals. Using the current standard genetic map of the laboratory mouse to polarize intervals with divergent recombination rates, we infer that the majority of evolutionary change occurred in one of the two tested lines of Gough Island mice. Our results confirm that natural populations harbour a high level of recombination rate polymorphism and highlight the disparities in recombination rate evolution across genomic scales.     

Impact of introduced house mice (Mus musculus) on burrowing seabirds on Steeple Jason and Grand Jason Islands,Falklands, South Atlantic

Whilst there is good evidence for negative impacts of introduced rat species on island ecosystems, the effects of house mice (Mus musculus) are generally less well documented. In some situations, introduced house mice can exert severe impacts, particularly where this is the only introduced mammal. Here, we examine the distribution, relative abundance and breeding success of small burrowing seabirds on Steeple Jason Island, Falklands, in relation to habitat types and the distribution of house mice which is the sole introduced mammal species, and we make comparisons with seabird distribution and densities on the neighbouring island of Grand Jason where mice are absent. Grey-backed storm-petrel (Garrodia nereis) and Wilson’s storm-petrel (Oceanites oceanicus), which due to their extremely small size are likely to be the most vulnerable to mouse predation, were considerably more abundant on mouse-free Grand Jason than on Steeple Jason. Grey-backed storm-petrel, which are typically associated with tussac grass, avoided this habitat on Steeple Jason where it is associated with high levels of house mouse activity (assessed from the proportion of wax baits gnawed overnight), whereas on mouse-free Grand Jason, there was no such avoidance. Wilson’s storm-petrel nesting on Steeple Jason suffered high rates of egg and chick loss. Whilst we found evidence for detrimental impacts of house mice on the two small storm-petrel species, there was no relationship between relative mouse activity levels and the distribution or abundance of the larger thin-billed Prion (Pachyptila belcheri).     

Feeding habits of the mochokid Synodontis punctifer Daget, 1964, in the Bandama River,Côte d'Ivoire

The feeding habits of Synodontis punctifer were investigated in the basin of the Bandama River. Fish specimens were collected using gill nets in the rainy season (May–July 2009) and in the dry season (November 2008 to March 2009 and August 2009). In this article, the importance of the different prey types was assessed using the corrected occurrence and weight percentage. These values were used to calculate the index of preponderance for each taxonomic category. One hundred and ten specimens of S. punctifer were examined, nine stomachs were found empty (8.18%). A total of 39 different prey species belonging to seven major groups Insects, Crustacea, Rotifera, Molluscs, Other invertebrates, Other organisms and Macrophytes were identified in the stomach contents. The results showed that Macrophytes were the primary food item and Insects the secondary prey item. Other preys were recorded occasionally. At the rainy season, the main foods of S. punctifer were Macrophytes and during the dry season this species ate Insects and Macrophytes. The food composition varied between localities. In the riverine localities, specimen fed on 38 prey types. Twenty‐seven prey categories were consumed in the lacustrine. In terms of diet overlap, Schoener coefficient values were above the 0.6 limit (α = 0.99), indicating some high overlap.     

PREY OF NORTHERN ROCKHOPPER PENGUINS AT GOUGH ISLAND,SOUTH ATLANTIC OCEAN

Klages, N. T., Brooke, M. de L. & Watkins, B. P. 1988. Prey of Northern Rockhopper Penguins at Gough Island, south Atlantic Ocean. Ostrich 59:162-165.

The diet of Northern Rockhopper Penguins Eudyptes chrysocome moseleyi breeding on Gough Island, south Atlantic Ocean was studied, during November 1984, 1985 and 1986 by stomach content analysis. Rockhopper Penguins fed chiefly on the euphausiids Thysanoessa gregaria, Euphausia lucens and E. similis. Fish and squid were of minor importance by mass but constituted the largest individual prey items.     

Spatio-temporal patterns of introduced mice and invertebrates on Antipodes Island

House mice (Mus musculus) are a widespread introduced species with major but often overlooked impacts on ecosystems, proportionally greater when they are the only introduced mammal present. Studies conducted on the ecology of mice on Antipodes Island, where they are the only introduced mammal, are presented and compared to previous work over the past four decades. Mice live-trapped on grids were more abundant in dense coastal tussock (147 mice/ha) compared to inland plateau grasslands (59 mice/ha), with a significant effect of age, but not sex, on both capture probability and range size. Body-size of mice has not changed over four decades, providing no evidence of gigantism, which on other Southern Ocean islands has been speculated to increase the predation risk to birds. Over 2,405 invertebrates from fourteen Orders were identified from pitfall traps and litter samples across five sites. Differences in invertebrate communities and taxonomic units attributable to habitat and altitude were detected among sites in both pitfall and litter samples on Antipodes Island. Differences in invertebrate communities were detected from litter samples on a neighbouring mouse-free island, with significantly greater abundance of large Amphipods and Collembola, but fewer Spiders. These data on introduced mouse ecology and invertebrate distribution on Antipodes Island contribute to the body of knowledge on Southern Ocean islands.     

Eradication of the House Mouse (Mus musculus) from Montague Island,New South Wales,Australia

Summary Of all the introduced mammalian pests, rodents are the most common invaders of islands and have triggered numerous extinctions around the globe. They have a short gestation period and large litter size, giving them the ability to colonise new areas rapidly. The same biological traits make them difficult to eradicate. Although the biodiversity benefits of removing exotic rodents from islands are increasingly being recognised, there are few published analyses of eradication attempts that critically evaluate eradication tools. This study examined the response of House Mouse (Mus musculus) populations to an eradication operation on Montague Island, Australia. While the specific impacts of mice have not been studied on this particular island, elsewhere they have been shown to have negative impacts on a range of species including plants, invertebrates, lizards and seabirds. On Montague Island, mouse abundance across different habitats was examined using mark–recapture data collected before and after the deployment of brodifacoum baits. Data obtained before baiting showed significantly lower numbers of mice in sites dominated by exotic grass compared with those dominated by native vegetation. Mouse numbers overall were declining during winter and the population was not breeding, making this an optimal time to undertake the eradication. Trapping immediately after the initial bait drop failed to capture any survivors, and no individuals have been detected during the 3 years since the deployment of baits. This study demonstrated that both small and large baits (0.6 and 2 g pellets) were effective in eradicating the House Mouse from Montague Island. While present, mice were probably slowing the re‐establishment of native vegetation by grazing seedlings and consuming seeds. With this pest now gone, the process of natural regeneration is expected to accelerate. The eradication of the House Mouse from Montague is likely to have other positive effects on the island’s biodiversity. The operation itself contributed to enhancing local capacity to eradicate exotic rodents from larger or more complex islands.