Invasive rats alter assemblage characteristics of land snails in the Ogasawara Islands

Although a large number of studies have shown that rats have a serious effect on island ecosystems, questions remain regarding how their invasion alters native species diversity and assemblage patterns on islands. In particular, a long history of invasions makes it difficult to understand the impact of rat invasion. In the present study, I investigate how an increase in predation by black rats (Rattus rattus) alters the assemblage characteristics of land snails on the Ogasawara Islands in the West Pacific. Radical declines in many land snail species have occurred since 2006 due to increasing predation by black rats. The frequency of shells with predation scars was greater on larger species and on those living on the trees and in a shallower position in the litter, suggesting greater predation pressure on these species. As a result of this selective predation, large species living on trees and in shallower parts of the litter declined. Accordingly, when black rats increased on the island, small species and species with a burrowing microhabitat became dominant in the land snail assemblage. The present findings can be used to aid in predicting the effects of rat eradication and in estimating the assemblage patterns and diversity prior to rat invasion.     

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.     

Prioritizing rat eradication on islands by cost and effectiveness to protect nesting seabirds

To prioritize conservation actions on Italian islands we used the case study of the eradication of the Black rat Rattus rattus to protect Cory’s shearwater Calonectris diomedea and Yelkouan shearwater Puffinus yelkouan. We evaluated for each island the effectiveness of rat eradication by means of two different indices, both based on the relative importance of the island’s nesting population of the two species at the national and regional scale, but differing in the parameters set at the divisor, i.e., respectively, the number of nesting pairs in rat-free islands and the number of islands occupied by shearwaters. We estimated analytically the monetary costs of rat eradication on each island. Islands at high risk of recolonization were excluded from further analyses, while costs and effectiveness of rat eradication were compared for the remaining islands. Rat eradication was most cost-effectively carried out on the island hosting the largest colony of P. yelkouan. Eradicating rats from all the islands in the ranking provided benefits to 63.9% of the Italian population of P. yelkouan, but only to 7.1% of that of C. diomedea. Comparing costs and effectiveness of all possible island combinations, ranging from a minimum budget of 50,000 € and a maximum of 1600,000 € (i.e. the cost for eradicating rats from all the listed islands), the maximum increase in effectiveness (marginal effectiveness) fell around a relatively small budget (200,000 €). For both species, when adopting the cost/effectiveness rankings, the number of pairs protected for 1000 € of investment was significantly higher than adopting rankings of effectiveness alone, demonstrating that conservation priorities are more efficiently identified by including monetary costs in the analysis.     

Browsing down our natural heritage: Deer impacts on vegetation structure and songbird populations across an island archipelago

Declines in forest and woodland birds have largely been attributed to habitat loss and fragmentation. In the past decade, however, the potential for herbivores to influence bird species abundance and community composition via their direct impact on vegetation structure has also been recognised. We tested the hypothesis that deer influence vegetation structure and bird assemblages in a large island archipelago in western North America using surveys of 18 islands with deer densities ranging from 0 to over 1 deer/ha. Amongst these islands, reduced predation and hunting pressure has allowed deer populations to increase above those likely to have existed in pre-European times. Our results support a growing body of evidence that deer regulate both the cover and architecture of understory vegetation which in turn profoundly affects island bird assemblages. Deer-free islands supported the most abundant and diverse bird fauna. Iconic songbirds such as the rufous hummingbird, song and fox sparrow were abundant on islands with no deer but substantially reduced on islands with high deer densities. Only one bird species, the dark-eyed junco, preferred moderate and high density deer islands. Our observations suggest that current cohorts of palatable shrubs on islands with high deer densities are relatively old and potentially represent an impending extinction debt, where the full effects of high deer density on island biota may take decades to fully unfold. Our results suggest that deer densities below a threshold of 0.1 deer/ha should allow native vegetation to recover and a rich and diverse bird species assemblage to persist. We suggest that adaptive management be used to test the validity of this threshold, and that without active management of deer abundance, local extinctions of native flora and fauna appear likely to accelerate.     

Understanding the decline and extinction of monarchs (Aves) in Polynesian Islands

Understanding the decline and extinction of species has become critical to conservation biology. The five monarch species of the genus Pomarea, endemic to the southeastern Pacific, are all listed as threatened. Introduced mammals and birds are believed to be responsible for their rarefaction. We analyzed the historical and current distribution of monarchs and introduced animals and found no relation between presence of Polynesian rats (Rattus exculans) and monarch distribution. There was a highly significant correlation between the arrival of the black rat (Rattus rattus) and the decline and extinction of monarch populations. The extinction of monarch populations after colonization by black rats tended to take longer on larger islands than on smaller ones. On islands without black rats, monarchs persisted even where forests have been reduced by more than 75%. After an island was colonized by black rats the number of monarch pairs with young decreased dramatically. Eggs in artificial nests placed in sites used by monarchs were only preyed upon by black rats. No eggs were preyed upon by Polynesian rats (Rattus exulans) or introduced birds. Monarch nests were mainly placed on horizontal branches inside the canopy and were more accessible than nests of Polynesian warbler (Acrocephalus caffer), a species still locally abundant. Warbler nests were placed higher up on vertical branches near the top of trees. These studies suggest that nest predation by black rats has been the main cause of monarch decline. However observations of direct aggression of adult monarchs by introduced red-vented bulbuls (Pycnonotus cafer), especially when monarchs raise their young, suggest that introduced birds could aggravate the decline of monarch populations already weakened by black rats. We discuss the practical implications of these findings for monarch conservation.     

The exploitation of sooty tern eggs in the seychelles

In the Seychelles, populations of sooty terns Sterna fuscata have declined this century, but the decline has not been as great as previously thought, and factors apart from over-exploitation of the eggs have been involved. A recent study of sooty tern ecology in the Seychelles has indicated that the present close season regulation serves no useful biological purpose. A system of quotas of eggs that may be taken from each island has not been effective due to chaning tern populations and to the voluntary cessation of collecting on some islands.This paper recommends that the eggs should in future be collected from two islands only, and that all other sooty tern colonies should be protected as far as is practicable. Exploitation should be restricted to the eggs of about 20% of the Seychelles/Amirantes population of sooty terns, but further research on the movements of the birds is urgently needed to determine the extent of inter-colony movements.     

The trade in fruit bats Pteropus spp. on Guam and other Pacific islands

Residents of Guam, Mariana Islands, have imported large numbers of fruit bats Pteropus spp. from other Pacific islands during the past 10 years as a delicacy. The trade in bats began in the 1960s or early 1970s after overhunting had greatly reduced the populations of Guam's native fruit bats. The quantity of bats shipped to the island peaked in 1979 and 1980, with an estimated total of more than 24 000 animals brought in each year. From 1981 to 1984, imports fell to an estimated mean of 14 475 bats per year. During the 1970s, Guam's importers purchased bats mainly from the Caroline Islands (Palau and Yap) and other islands in the Marianas (Saipan, Rota and Tinian). Changes in availability and laws during the 1980s have caused the fruit bat trade to expand geographically, making islands outside Micronesia (Western Samoa, Tonga, American Samoa and Papua New Guinea) important suppliers of bats. Because large-scale commercial exploitation may severely deplete an island's population of fruit bats, perhaps beyond the point of recovery, we recommend that island governments be conservative in the number of bats allowed for harvest until studies on abundance and life history have been completed.     

The Galápagos giant tortoises (Geochelone elephantopus) Part II: Conservation methods

Eight of the 11 surviving races of Galápagos tortoises are threatened due to decreased population sizes and predation and/or competition by introduced mammals. Systematic hunting has been effective for controlling and even eliminating goats on the less elevated, more sparsely vegetated, smaller islands and for controlling pigs, even on some of the larger islands. However, the method has been ineffective against dogs and cats and it is doubtful that it could be used to eliminate goats or pigs on the more elevated, larger islands which have a diversity of vegetation zones. Alternative control or extermination methods are being sought and tested for these feral mammals and black rats. In an attempt to increase yearly recruitment of hatchlings, lava corrals have been constructed around nests, resulting in almost 100 per cent success in preventing nest destruction by pigs; but the method was ineffective against dogs. While other control methods are being sought, young of the endangered races are being hatched and raised in captivity for restocking of endemic populations. During the past seven years, improved and highly successful techniques have been found for (1) establishment of breeding colonies and construction of artificial nesting sites, (2) transport of eggs from wild nests, (3) incubation of eggs, and (4) raising of young in captivity. By August 1972, 231 young of six races were being raised, and 71 4·5–6·5 year old captive raised tortoises released on Pinzón in 1970 and 1971 were in good condition and growing rapidly. The advantages of conducting the breeding/raising programme in the Galápagos rather than foreign locations are discussed.     

Urban conservation genetics: Study of a terrestrial salamander in the city

Urbanization is a global process that negatively affects ecosystems in various ways. It is namely an important cause of habitat loss, and animal species inhabiting cities often exhibit highly fragmented populations. Those populations may be prone to problems associated with reduced abundance and isolation such as genetic drift and inbreeding. However, few studies have dealt with animal populations living in cities. In this paper, we assessed the genetic structure of urban red-backed salamander populations located in remnant forested areas within a major city (Montréal) and on two nearby islands. Microsatellite analysis revealed a high degree of genetic differentiation among populations from Montréal and from neighboring islands. Within Montréal island, no clear genetic structure was detected, with some populations as far as 35 km apart not being differentiated. On the other hand, genetic differentiation was observed at a small spatial scale (0.7-1.7 km) for three populations located near downtown area where anthropogenic perturbations are older. We believe these populations had very low effective population size for a long time, allowing for fast genetic drift and subsequent population differentiation. Although no inbreeding was detected in any population, genetic variation was relatively low. These results illustrate the importance of preserving large forested areas within cities, especially given that these patches may also harbor rarer and threatened species which may be even more negatively affected by urbanization.     

The impact of feral cats and dogs on populations of the West Indian rock iguana, Cyclura carinata

A population of rock iguanas, Cyclura carinata, inhabiting Pine Cay in the Caicos Islands was nearly extirpated during the three years following construction of a hotel and tourist facility. The decline, from an estimated adult lizard population of nearly 5500, was due primarily to predation by domestic dogs and cats introduced to the island simultaneously with hotel construction. Population declines on other nearby islands were also attributed to predation by these feral mammals.     

Long-term trends in the demography of the Allen Cays Rock Iguana (Cyclura cychlura inornata): Human disturbance and density-dependent effects

Allen Cays Rock Iguanas (Cyclura cychlura inornata) are native to two small islets (Leaf and U Cay) in the north-central Bahamas. These populations were nearly extirpated in the early 1900s because of heavy hunting pressure (for food), but increased to a total of ca. 150 lizards in 1970, and now number over 500 (not including juveniles). Over the past several decades poaching has declined, but tourist visitation (including nearly daily supplemental feeding of iguanas) has increased. To examine human impacts on the demography of these iguanas, survival, population growth rates, and population sizes for subadult and adult (>25 cm snout-vent length) males and females on the two cays were estimated based on mark-recapture data collected over a 25-year period (1980-2004). As predicted, annual survival probability was higher on U Cay (with less human visitation) than on Leaf Cay, was higher in females than in males (which are bolder), and exhibited a declining trend. Both populations more than doubled during this study, but population growth rates declined to near zero in recent years. These data reflect the importance of human impacts, but also suggest that the populations may be nearing carrying capacity. The rapid population growth observed on these cays, and that seen for several other translocated iguana populations, suggest that if unnatural causes of mortality are reduced or eliminated, island populations of iguanas are capable of rapid recovery. The inexpensive establishment of assurance colonies on undisturbed “islands” should be considered for any comprehensive management plan for endangered species of iguanas.     

The Prince Edward Islands: A sanctuary for seabirds in the Southern Ocean

This paper reviews the breeding distribution and numerical status of selected species of seabirds at 23 oceanic island groups in the Southern Ocean (between latitudes 35 and 70°S), based on census data and population estimates which have become available through increased scientific endeavour in the region during the last 15 years or so. The paper focuses on the avifauna of the Prince Edward islands, in the southern Indian Ocean, and points to the importance of these islands as a breeding station and sanctuary for seabirds. Only one oceanic island group, the Crozets, contains more species of breeding seabirds than the Prince Edward islands. Substantial segements (about 10% and greater) of the world breeding populations of the king and macaroni penguins, and the wandering, grey-headed, yellow-nosed and sooty albatrosses are based at the Prince Edward group. We recommend that Prince Edward Island (sensu stricto) should be a wilderness area, and that development at Marion Island should be kept to an absolute minimum.     

Connectivity between island Marine Protected Areas and the mainland

In addition to protecting important species or ecosystems, ideally Marine Protected Areas (MPAs) should also act as propagule sources and be located where they can receive larval influxes so as to buffer species against local extinctions and allow genetic mixing. There are many examples of island MPAs across the world because of their advantages compared to mainland sites, and although evidence suggests they effectively protect biodiversity and may have localized fisheries benefits, their contribution to larval export and connectivity to mainland areas remains largely unknown; these are important aspects of MPA sustainability and functioning. Here I examined the genetic differentiation between four island MPAs in northern Europe and their respective mainlands, relative to differentiation of populations along the coast at similar spatial scales for two marine invertebrates (Semibalanus balanoides, planktonic larvae; Nucella lapillus, direct development). These data were used to infer past patterns of larval exchange between populations. Gene flow was restricted at all four locations between the islands and the mainland, compared to gene flow along the coast (for both species) and more distant islands had greater differentiation than those closer to the mainland for S. balanoides, but not for N. lapillus. My results of genetic analysis suggest that island MPAs may not provide as much larval export or receive as much buffering against local extinctions, compared with mainland populations (for a similar sized protected area). My findings are especially relevant to the creation of MPA networks and in understanding the importance of connectivity in achieving the greatest fisheries benefits and in ensuring inter-MPA linkages. Low levels of larval exchange may limit the success of any protected area and may prevent multiple conservation objectives from being achieved.     

Single species island biogeography and Montana mule deer

The theory of island biogeography supplemented with dynamic concepts from other areas of biology can be applied to a single species. Data series from mule deer Odocoileus hemonius hemonius populations inhabiting diverse mountain as well as prairie habitats were compared. The fawn-doe ratios obtained at 6 months of age were inversely correlated (r = ?0·97; p < 0·01) with ‘island’ size as indicated by the percentage of cover. Populations inhabiting small islands of cover in the prairie consistently had higher winter fawn-doe ratios than those of the relatively secure mountain habitats. Cover fragmentation and low security level of the prairi populations relates to a high annual turnover rate compared with that of the mountain populations. The impact of cover fragmentation is consistent with this interpretation of theory which pictures reproduction as annual recolonisation. This approach may be useful in predicting hunting and ecological impact effects.     

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.     

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.     

High levels of hatching failure in an insular population of the South Island robin: a consequence of food limitation?

Endangered birds in insular environments like New Zealand are often translocated to predator-free offshore islands for conservation purposes. Some translocated populations however exhibit reduced fecundity, and it has been suggested that either inbreeding (due to small number of founders) or resource shortages (due to high population density) may limit reproductive success. For example, the South Island robin (Petroica a. australis) population on Motuara Island (59 ha) was founded by only five birds but has increased to ∼600 individuals, or a density >10 times that of the mainland. Despite this apparent success, the rate of hatching failure in this population is three times higher than in mainland populations, with more than a third of eggs failing to hatch. To test if elevated hatching failure is the result of food limitation, we carried out a food supplementation experiment by providing females with mealworms equivalent to ∼50% of their daily energy requirements. Food supplementation had no effect on hatching success, clutch size, incubation attentiveness or nest size. Egg volume increased with food supplementation in one of the two years of this study, but both egg volume and incubation attentiveness were unrelated to rate of hatching failure. As previous genetic analyses confirmed that the bottlenecked population of robins on Motuara Island have significantly less genetic variation than their source population, we conclude that inbreeding depression, and not food limitation, is the most likely explanation for their high rate of hatching failure. We suggest that the experimental translocation of genetically dissimilar individuals be considered as a possible remedy for low productivity in island populations that were initiated with few founders.     

Current status of rodents on islands in the Gulf of California

On islands in the Gulf of California, many species and subspecies of rodents are relict species, restricted to the islands, and now considered extinct or verging on extinction. From 1991 to 1999, each island was surveyed and an average of 15 work-nights were spent on each island, for a total of 15,000 trap-nights. A Fourier series was used to estimate population density. Five native taxa are now considered extinct (Chaetodipus baileyi fornicatus, Peromyscus guardia mejiae, P. g. harbinsoni, P. pembertoni, and Neotoma bunkeri). Two species are at risk of extinction (Dipodomys insularis and Neotoma albigula varia). The most probable cause for extinction is the introduction of nonnative species, specifically cats. We believe that the most vulnerable rodent populations of the islands in the Gulf of California to non-native species are Neotoma lepida latirostra on Danzante island and Neotoma lepida abbreviata on San Francisco island, both in the state of Baja California Sur.     

Pathogen exposure in endangered island fox (Urocyon littoralis) populations: Implications for conservation management

Island fox (Urocyon littoralis) populations on four California Channel Islands have declined severely since 1994. Canine distemper (CDV) was suspected to be responsible for the decline of the Santa Catalina Island fox, so knowledge of infectious disease exposure in the remaining island fox populations was urgently needed. This study reviewed previous pathogen exposure in island foxes and investigated the current threat by conducting a serologic survey of foxes on all islands and sympatric feral cats on three islands from 2001 to 2003 for antibodies against canid pathogens. Before the decline, foxes had evidence of exposure to CDV, canine adenovirus (CAV), canine parvovirus (CPV), and Toxoplasma, with exposure to these five pathogens differing greatly by island. Exposure to canine coronavirus (CCV), canine herpesvirus (CHV), and Leptospira was rare. In 2001-2003, wild-born foxes had evidence of exposure to CDV (5.2-32.8%) on 5 of 6 islands, CPV (28-100%) and CAV (4.7-100%) on five islands, and Toxoplasma gondii (2.3-15.4%) on four islands. Exposure to CCV, CHV and Leptospira was less common. Sharing of infectious agents between sympatric foxes and feral cats appeared minimal, but CDV exposure was detected in two cats on Santa Catalina Island. Domestic dogs have historically been present on the islands, but it is not known if canine diseases can be maintained in fox populations without the continual presence of dogs. Targeted vaccination programs against the most virulent pathogens and continued intensive disease surveillance may help protect the critically small remaining fox populations from disease outbreaks that could threaten the success of ongoing conservation efforts.     

Husking stations provide evidence of seed predation by introduced rodents in Tongan rain forests

The islands of Tonga, in Western Polynesia, lacked rodents prior to human colonization. During the past 3000 years, people have introduced three species of rats (Rattus). Seed predation by rodents structures plant communities in many parts of the world. Therefore, there is the potential for rats to have a significant impact on the regeneration of plant communities in Tonga. We found 53 husking stations—sheltered sites used by rats to strip inedible protective structures (husks) from the seeds they eat—on eight islands in Tonga. The stations contained 13,555 empty husks from at least 18 plant species, but only 165 viable seeds and seedlings. These husking stations provide evidence of seed predation by rats. Determining the extent to which seed predation threatens plant regeneration will be important for conservation of native forest remnants.