Control of invasive rats on islands and priorities for future action

Invasive rats are one of the world's most successful animal groups that cause native species extinctions and ecosystem change, particularly on islands. On large islands, rat eradication is often impossible and population control, defined as the local limitation of rat abundance, is now routinely performed on many of the world's islands as an alternative management tool. However, a synthesis of the motivations, techniques, costs, and outcomes of such rat‐control projects is lacking. We reviewed the literature, searched relevant websites, and conducted a survey via a questionnaire to synthesize the available information on rat‐control projects in island natural areas worldwide to improve rat management and native species conservation. Data were collected from 136 projects conducted over the last 40 years; most were located in Australasia (46%) and the tropical Pacific (25%) in forest ecosystems (65%) and coastal strands (22%). Most of the projects targeted Rattus rattus and most (82%) were aimed at protecting birds and endangered ecosystems. Poisoning (35%) and a combination of trapping and poisoning (42%) were the most common methods. Poisoning allows for treatment of larger areas, and poison projects generally last longer than trapping projects. Second‐generation anticoagulants (mainly brodifacoum and bromadiolone) were used most often. The median annual cost for rat‐control projects was US$17,262 or US$227/ha. Median project duration was 4 years. For 58% of the projects, rat population reduction was reported, and 51% of projects showed evidence of positive effects on biodiversity. Our data were from few countries, revealing the need to expand rat‐control distribution especially in some biodiversity hotspots. Improvement in control methods is needed as is regular monitoring to assess short‐ and long‐term effectiveness of rat‐control.     

Feral Goat Eradications on Islands

Abstract:  Introduced mammals are major drivers of extinction. Feral goats ( Capra hircus ) are particularly devastating to island ecosystems, causing direct and indirect impacts through overgrazing, which often results in ecosystem degradation and biodiversity loss. Removing goat populations from islands is a powerful conservation tool to prevent extinctions and restore ecosystems. Goats have been eradicated successfully from 120 islands worldwide. With newly developed technology and techniques, island size is perhaps no longer a limiting factor in the successful removal of introduced goat populations. Furthermore, the use of global positioning systems, geographic information systems, aerial hunting by helicopter, specialized hunting dogs, and Judas goats has dramatically increased efficiency and significantly reduced the duration of eradication campaigns. Intensive monitoring programs are also critical for successful eradications. Because of the presence of humans with domestic goat populations on large islands, future island conservation actions will require eradication programs that involve local island inhabitants in a collaborative approach with biologists, sociologists, and educators. Given the clear biodiversity benefits, introduced goat populations should be routinely removed from islands.     

Importance of Assessing Population Genetic Structure before Eradication of Invasive Species: Examples from Insular Norway Rat Populations

Abstract:  Determining the inter-island migration abilities of pest species and delimiting eradication units enable more viable long-term eradication campaigns because recurrent colonization from neighboring islands is avoided. We examined the genetic structure of the invasive Norway rat ( Rattus norvegicus ) to identify gene flow between islands and delimit population units at different geographical scales. We investigated variation in eight microsatellite loci in rat populations from 18 islands, representing five archipelagos off the Brittany coast (France). Although most of the islands are isolated from each other, short genetic distances, weak F_ST values between close islands, and a high level of cross-assignment showed that individuals collected on different islands could represent a single population unit. A Bayesian clustering method also supported the existence of high levels of gene flow between some neighboring islands. Thus, the statement "one island equals one population" can be false when inter-island distances are less than a few hundred meters. Genetic studies enable the definition of island clusters among which migration may occur that should be considered eradication units. To avoid reinvasion and to minimize ecological and economic costs, rats on all islands in an eradication unit should be eradicated simultaneously. We suggest that the genetic monitoring we performed here can be applied for management of any pest.     

Eradication of Invasive Mammals on Islands Inhabited by Humans and Domestic Animals

Abstract: Non‐native invasive mammal species have caused major ecological change on many islands. To conserve native species diversity, invasive mammals have been eradicated from several islands not inhabited by humans. We reviewed the challenges associated with campaigns to eradicate invasive mammals from islands inhabited by humans and domestic animals. On these islands, detailed analyses of the social, cultural, and economic costs and benefits of eradication are required to increase the probability of local communities supporting the eradication campaign. The ecological benefits of eradication (e.g., improvement of endemic species’ probability of survival) are difficult to trade‐off against social and economic costs due to the lack of a common currency. Local communities may oppose an eradication campaign because of perceived health hazards, inconvenience, financial burdens, religious beliefs, or other cultural reasons. Besides these social challenges, the presence of humans and domestic animals also complicates eradication and biosecurity procedures (measures taken to reduce the probability of unwanted organisms colonizing an island to near zero). For example, houses, garbage‐disposal areas, and livestock‐feeding areas can provide refuges for certain mammals and therefore can decrease the probability of a successful eradication. Transport of humans and goods to an island increases the probability of inadvertent reintroduction of invasive mammals, and the establishment of permanent quarantine measures is required to minimize the probability of unwanted recolonization after eradication. We recommend a close collaboration between island communities, managers, and social scientists from the inception of an eradication campaign to increase the probability of achieving and maintaining an island permanently free of invasive mammals.     

Cost-Effective Suppression and Eradication of Invasive Predators

Abstract: Introduced predators can have pronounced effects on naïve prey species; thus, predator control is often essential for conservation of threatened native species. Complete eradication of the predator, although desirable, may be elusive in budget‐limited situations, whereas predator suppression is more feasible and may still achieve conservation goals. We used a stochastic predator–prey model based on a Lotka‐Volterra system to investigate the cost‐effectiveness of predator control to achieve prey conservation. We compared five control strategies: immediate eradication, removal of a constant number of predators (fixed‐number control), removal of a constant proportion of predators (fixed‐rate control), removal of predators that exceed a predetermined threshold (upper‐trigger harvest), and removal of predators whenever their population falls below a lower predetermined threshold (lower‐trigger harvest). We looked at the performance of these strategies when managers could always remove the full number of predators targeted by each strategy, subject to budget availability. Under this assumption immediate eradication reduced the threat to the prey population the most. We then examined the effect of reduced management success in meeting removal targets, assuming removal is more difficult at low predator densities. In this case there was a pronounced reduction in performance of the immediate eradication, fixed‐number, and lower‐trigger strategies. Although immediate eradication still yielded the highest expected minimum prey population size, upper‐trigger harvest yielded the lowest probability of prey extinction and the greatest return on investment (as measured by improvement in expected minimum population size per amount spent). Upper‐trigger harvest was relatively successful because it operated when predator density was highest, which is when predator removal targets can be more easily met and the effect of predators on the prey is most damaging. This suggests that controlling predators only when they are most abundant is the “best” strategy when financial resources are limited and eradication is unlikely.     

Invasive Non‐Native Species’ Provision of Refugia for Endangered Native Species

Abstract: The influence of non‐native species on native ecosystems is not predicted easily when interspecific interactions are complex. Species removal can result in unexpected and undesired changes to other ecosystem components. I examined whether invasive non‐native species may both harm and provide refugia for endangered native species. The invasive non‐native plant Casuarina stricta has damaged the native flora and caused decline of the snail fauna on the Ogasawara Islands, Japan. On Anijima in 2006 and 2009, I examined endemic land snails in the genus Ogasawarana. I compared the density of live specimens and frequency of predation scars (from black rats[Rattus rattus]) on empty shells in native vegetation and Casuarina forests. The density of land snails was greater in native vegetation than in Casuarina forests in 2006. Nevertheless, radical declines in the density of land snails occurred in native vegetation since 2006 in association with increasing predation by black rats. In contrast, abundance of Ogasawarana did not decline in the Casuarina forest, where shells with predation scars from rats were rare. As a result, the density of snails was greater in the Casuarina forest than in native vegetation. Removal of Casuarina was associated with an increased proportion of shells with predation scars from rats and a decrease in the density of Ogasawarana. The thick and dense litter of Casuarina appears to provide refugia for native land snails by protecting them from predation by rats; thus, eradication of rats should precede eradication of Casuarina. Adaptive strategies, particularly those that consider the removal order of non‐native species, are crucial to minimizing the unintended effects of eradication on native species. In addition, my results suggested that in some cases a given non‐native species can be used to mitigate the impacts of other non‐native species on native species.     

Remote Forest Refugia for Fijian Wildlife

Abstract:  On Pacific islands non-native rats and mongooses threaten many native species. In Fiji we compared visitation rates of rats and mongooses at bait stations and measured biomass of leaf-litter invertebrates to assess the relative predation pressure from these species in forest areas at different distances from the forest edge. Forest areas over 5 km from the forest edge had significantly fewer baits encountered by rats or mongooses than did natural forest areas nearer agricultural and urban habitats. Remote forest areas may function as a last refuge for island species threatened by predation from non-native rats and mongooses. The biomass of leaf-litter invertebrates in remote forest areas was higher, indicating a refuge effect for some taxa targeted by rats and mongooses. Protection of the few remaining large blocks of natural forests on Pacific islands may be the most cost-effective approach for conserving many island endemics threatened by rats and mongooses. Logging roads can compromise this refuge effect by acting as dispersal routes for rats into natural forests.     

Responses of Tuatara (Sphenodon punctatus) to Removal of Introduced Pacific Rats from Islands

Abstract:  Invasive mammalian predators such as rats are now widespread on islands, but hypotheses about their effects have rarely been tested. Circumstantial evidence from New Zealand indicates that, when introduced to islands, Pacific rats ( Rattus exulans ) have negative effects on endemic plants, invertebrates, birds, and reptiles, including the tuatara ( Sphenodon punctatus ). We tested the effects of Pacific rats on tuatara by comparing the demographic structure and body condition of tuatara populations on three islands before and after removal of rats and on a fourth island where rats remained. In the presence of rats, juvenile tuatara constituted on average 0–5% of the sample tuatara populations. When Pacific rats were removed after at least 200 years' occupancy, the proportion of juvenile tuatara increased 3.5- to 17-fold and body condition of adult males and females also improved (sometimes dramatically). We predict that, unless Pacific rats are removed from Taranga Island, the tuatara population will collapse because of low population density and the lack of juvenile recruitment. Our results demonstrate that when invasive species exert subtle effects on recruitment and body condition, the effects on populations of long-lived endemic species may only become apparent long after the invasion.     

Current Practices and Future Opportunities for Policy on Climate Change and Invasive Species

Abstract:  Climate change and invasive species are often treated as important, but independent, issues. Nevertheless, they have strong connections: changes in climate and societal responses to climate change may exacerbate the impacts of invasive species, whereas invasive species may affect the magnitude, rate, and impact of climate change. We argue that the design and implementation of climate-change policy in the United States should specifically consider the implications for invasive species; conversely, invasive-species policy should address consequences for climate change. The development of such policies should be based on (1) characterization of interactions between invasive species and climate change, (2) identification of areas where climate-change policies could negatively affect invasive-species management, and (3) identification of areas where policies could benefit from synergies between climate change and invasive-species management.     

Managing Aquatic Species of Conservation Concern in the Face of Climate Change and Invasive Species

Abstract:  The difficult task of managing species of conservation concern is likely to become even more challenging due to the interaction of climate change and invasive species. In addition to direct effects on habitat quality, climate change will foster the expansion of invasive species into new areas and magnify the effects of invasive species already present by altering competitive dominance, increasing predation rates, and enhancing the virulence of diseases. In some cases parapatric species may expand into new habitats and have detrimental effects that are similar to those of invading non-native species. The traditional strategy of isolating imperiled species in reserves may not be adequate if habitat conditions change beyond historic ranges or in ways that favor invasive species. The consequences of climate change will require a more active management paradigm that includes implementing habitat improvements that reduce the effects of climate change and creating migration barriers that prevent an influx of invasive species. Other management actions that should be considered include providing dispersal corridors that allow species to track environmental changes, translocating species to newly suitable habitats where migration is not possible, and developing action plans for the early detection and eradication of new invasive species.     

Effect of Invasive Plant Species on Temperate Wetland Plant Diversity

Abstract:  Invasive species are a major threat to global biodiversity and an important cause of biotic homogenization of ecosystems. Exotic plants have been identified as a particular concern because of the widely held belief that they competitively exclude native plant species. We examined the correlation between native and invasive species richness in 58 Ontario inland wetlands. The relationship between exotic and native species richness was positive even when we controlled for important covarying factors. In addition, we examined the relationship between the abundance of four native species (  Typha latifolia, T. angustifolia, Salix petiolaris, Nuphar variegatum ) and four invasive species (  Lythrum salicaria, Hydrocharis morsus-ranae, Phalaris arundinacea, Rhamnus frangula ) that often dominate temperate wetlands and native and rare native species richness. Exotic species were no more likely to dominate a wetland than native species, and the proportion of dominant exotic species that had a significant negative effect on the native plant community was the same as the proportion of native species with a significant negative effect. We conclude that the key to conservation of inland wetland biodiversity is to discourage the spread of community dominants, regardless of geographical origin.     

Drivers of Seabird Population Recovery on New Zealand Islands after Predator Eradication

Eradication of introduced mammalian predators from islands has become increasingly common, with over 800 successful projects around the world. Historically, introduced predators extirpated or reduced the size of many seabird populations, changing the dynamics of entire island ecosystems. Although the primary outcome of many eradication projects is the restoration of affected seabird populations, natural population responses are rarely documented and mechanisms are poorly understood. We used a generic model of seabird colony growth to identify key predictor variables relevant to recovery or recolonization. We used generalized linear mixed models to test the importance of these variables in driving seabird population responses after predator eradication on islands around New Zealand. The most influential variable affecting recolonization of seabirds around New Zealand was the distance to a source population, with few cases of recolonization without a source population ≤25 km away. Colony growth was most affected by metapopulation status; there was little colony growth in species with a declining status. These characteristics may facilitate the prioritization of newly predator‐free islands for active management. Although we found some evidence documenting natural recovery, generally this topic was understudied. Our results suggest that in order to guide management strategies, more effort should be allocated to monitoring wildlife response after eradication. Conductores de la Recuperación de Poblaciones de Aves Marinas en Islas de Nueva Zelanda después de la Erradicación de Depredadores     

Assessing the Effects of Climate Change on Aquatic Invasive Species

Abstract:  Different components of global environmental change are typically studied and managed independently, although there is a growing recognition that multiple drivers often interact in complex and nonadditive ways. We present a conceptual framework and empirical review of the interactive effects of climate change and invasive species in freshwater ecosystems. Climate change is expected to result in warmer water temperatures, shorter duration of ice cover, altered streamflow patterns, increased salinization, and increased demand for water storage and conveyance structures. These changes will alter the pathways by which non-native species enter aquatic systems by expanding fish-culture facilities and water gardens to new areas and by facilitating the spread of species during floods. Climate change will influence the likelihood of new species becoming established by eliminating cold temperatures or winter hypoxia that currently prevent survival and by increasing the construction of reservoirs that serve as hotspots for invasive species. Climate change will modify the ecological impacts of invasive species by enhancing their competitive and predatory effects on native species and by increasing the virulence of some diseases. As a result of climate change, new prevention and control strategies such as barrier construction or removal efforts may be needed to control invasive species that currently have only moderate effects or that are limited by seasonally unfavorable conditions. Although most researchers focus on how climate change will increase the number and severity of invasions, some invasive coldwater species may be unable to persist under the new climate conditions. Our findings highlight the complex interactions between climate change and invasive species that will influence how aquatic ecosystems and their biota will respond to novel environmental conditions.     

Effects of Spatially Extensive Control of Invasive Rats on Abundance of Native Invertebrates in Mainland New Zealand Forests

Predation on native fauna by non‐native invasive mammals is widely documented, but effects of predation at the population level are rarely measured. Eradication of invasive mammals from islands has led to recovery of native biota, but the benefits of controlling invasive mammal populations in settings where eradication is not feasible are less understood. We used various combinations of aerially delivered toxic bait and control measures on the ground to reduce abundances of invasive rats (Rattus rattus) to low levels over large areas on mainland New Zealand and then monitored the abundance of invertebrates on replicated treatment sites to compare with abundances on similar nontreatment sites. We also assessed rat diet by examining stomach contents. Abundance of the rats’ most‐consumed invertebrate prey item, the large‐bodied Auckland tree weta (Hemideina thoracica), increased 3‐fold on treatment sites where we maintained rats at <4/ha for approximately 3 years, compared with the nontreatment sites. Auckland tree weta also increased in abundance on sites where rats were controlled with a single aerial‐poisoning operation, but rat abundance subsequently increased on these sites and tree weta abundance then declined. Nevertheless, our data suggest that biennial reduction of rat abundances may be sufficient to allow increases in tree weta populations. Other invertebrates that were consumed less often (cave weta [Rhaphidophoridae], spiders [Araneae], and cockroaches [Blattodea]) showed no systematic changes in abundance following rat control. Our results suggest that the significant threat to recruitment and individual survival that predation by rats poses for tree weta can be mitigated by wide‐scale aerial pest control. Efectos del Control Extensivo Espacial de Ratas Invasoras sobre la Abundancia de Invertebrados Nativos en Bosques de Nueva Zelanda     

Impact of Invasive Cane Toads on Australian Birds

Abstract: The cane toad (Bufo marinus), a large, toxic, American anuran, was introduced to Australia in 1935. Populations of many of Australia's reptiles (snakes, varanid lizards, crocodiles) and carnivorous mammals (dasyurid marsupials) have declined because these predators are killed by the toad's powerful toxins. In contrast to these well‐studied species, little is known about the cane toads impacts on Australian birds. We reviewed published and unpublished data on behavioral interactions between Australian avian predators and cane toads and collated distributional and dietary information to identify avian taxa potentially at risk from cane toad invasion. Cane toads are sympatric with 172 frog‐eating bird species in Australia, and an additional 8 bird species overlap with the predicted future range of the toad. Although many bird species thus are potentially at risk, behavioral observations suggest the risk level is generally low. Despite occasional reports of Australian birds being killed when they ingest cane toads, most birds either ignore toads or survive the predation event. The apparently higher tolerance of Australian birds to toad toxins, compared with Australian reptiles and marsupials, may reflect genetic exchange between Australian birds and Asian populations that encounter other bufonid species regularly and hence have evolved the capacity to recognize or tolerate this toxic prey.     

Prioritizing islands for the eradication of invasive vertebrates in the United Kingdom overseas territories

Invasive alien species are one of the primary threats to native biodiversity on islands worldwide. Consequently, eradicating invasive species from islands has become a mainstream conservation practice. Deciding which islands have the highest priority for eradication is of strategic importance to allocate limited resources to achieve maximum conservation benefit. Previous island prioritizations focused either on a narrow set of native species or on a small geographic area. We devised a prioritization approach that incorporates all threatened native terrestrial vertebrates and all invasive terrestrial vertebrates occurring on 11 U.K. overseas territories, which comprise over 2000 islands ranging from the sub‐Antarctic to the tropics. Our approach includes eradication feasibility and distinguishes between the potential and realistic conservation value of an eradication, which reflects the benefit that would accrue following eradication of either all invasive species or only those species for which eradication techniques currently exist. We identified the top 25 priority islands for invasive species eradication that together would benefit extant populations of 155 native species including 45 globally threatened species. The 5 most valuable islands included the 2 World Heritage islands Gough (South Atlantic) and Henderson (South Pacific) that feature unique seabird colonies, and Anegada, Little Cayman, and Guana Island in the Caribbean that feature a unique reptile fauna. This prioritization can be rapidly repeated if new information or techniques become available, and the approach could be replicated elsewhere in the world. Priorización de Islas para la Erradicación de Vertebrados Invasores en los Territorios Exteriores del Reino Unido     

Effects of an invasive ant on land snails in the Ogasawara Islands

We investigated how Pheidole megacephala has affected endemic achatinellid snails because these snails are excellent indicators of the impact of ants and they have high conservation value in Ogasawara. In 2015 we surveyed the Minamizaki area of Hahajima Island of Ogasawara, designated a core zone of the World Heritage Site, for P. megacephala. In Minamizaki, we determined the distribution and density of achatinellid snails in 2015 and compared these data with their distribution and density in 2005. Land cover in the survey area was entirely forest. We also tested whether P. megacephala preyed on achatinellid snails in the laboratory. P. megacephala was present in the forested areas of Minamizaki. Achatinellid snails were absent in 19 of 39 sites where P. megacephala was present, whereas in other areas densities of the snails ranged from 2 to 228 individuals/site. In the laboratory, P. megacephala carried 6 of 7 achatinellid snails and a broken shell was found. Snail distribution and density comparisons and results of the feeding experiments suggest that the presence of P. megacephala has contributed to the decline of achatinellid snails in forests in the survey area. Yet, P. megacephala is not on the official list of invasive non‐native species. Stakeholders using the list of invasive species to develop conservation programs should recognize that invasiveness of non‐native species differs depending on the ecosystem and that official lists may not be complete.