The population ecology of Rattus exulans on Tiritiri Matangi Island,and a model of comparative population dynamics in New Zealand

Abstract

On Tiritiri, a small predator-free island in northern New Zealand, kiore (Rattus exulans) were live and snap trapped in grassland and forest. In both habitats, kiore abundance peaked in late summer/autumn. The increase followed a 3 month breeding season during which females produced two to three litters, each averaging 7 young. During the population decline in autumn and winter, animals lost weight. Few bred in the breeding season of their birth and none lived to breed in a second breeding season, so the population consisted of distinct age cohorts. These patterns may relate to a highly seasonal food supply.

Kiore elsewhere in New Zealand show seasonal breeding, but the length of breeding, sexual maturation, and litter size vary. Other studies of kiore in the Pacific show less marked seasonal fluctuations, longer breeding seasons, and smaller litters. We propose a model to explain the variation in rodent demography in New Zealand. The model is based on the seasonal availability of food, along with the modifying influences of predation and dispersal.     

Modelling the distribution and interaction of introduced rodents on New Zealand offshore islands

Aim To establish the factors that correlate with the distribution of the four most commonly introduced rodent species on New Zealand offshore islands — ship rat (Rattus rattus), Norway rat (R. norvegicus), Pacific rat or kiore (R. exulans) and house mouse (Mus musculus) — and examine if these distributions are interactive at the archipelago scale. Location The 297 offshore islands of the New Zealand archipelago (latitude: 34° S to 47° S; longitude: 166° E to 179° E). Methods Data on the distribution of all four introduced rodent species and the characteristics of New Zealand offshore islands were collated from published surveys and maps. The distribution of individual rodent species was regressed on island characteristics using a logistic generalized linear model. Interactions were examined by including the distributions of other rodent species as predictors in models. Results All four rodent species appear to be limited by a variety of factors, which differ between species in both number and type. The distribution of ship rats is limited by the most factors, reflecting the extent of its distribution across the archipelago. The distribution of mice is the least explicative. Only the three rat species interacted in their distribution. The distribution of kiore on offshore islands is significantly negatively related to that of ship rats and to a lesser extent Norway rats. The distribution of mice did not appear affected in any way by the number of other rodent species on an island. Main conclusions Differences in competitive ability and dispersal allow all four species to inhabit the New Zealand archipelago. Kiore distribution appears to be most limited by ship rat (and to a lesser extent Norway rat) distribution. The distribution of kiore was not found to interact with the distribution of mice on offshore islands, as has been suggested by others. The distribution of mice on offshore islands was difficult to model, which highlights the difficulties in managing this species. Overall the results offer valuable insights for management methods to assist preventing the invasion of offshore islands.     

A histochemical study of the swimming musculature of Antarctic fish

The kiore, once common throughout New Zealand, had disappeared from most of the country by the end of the 19th century, and is now found only on certain offshore islands and in areas of Fiordland where at least one of the three introduced European rodent species is absent. It is usually accepted that the kiore was displaced by ship rats (Rattus r. rattus) and Norway rats (R. norvegicus). However, recent investigations on Stewart Island have revealed kiore, ship rats, and Norway rats living in close association, but in the absence of mice (Mus musculus). In the area studied the kiore seemed to inhabit mainly grassland. Re‐examination of possible reasons for the decline of the kiore strongly suggests that competition from mice has been a major contributing factor. It seems that in New Zealand a niche no longer exists for kiore once mice, ship rats, and Norway rats have all become established.     

Heavy rimu (Dacrydium cupressinum) mast seeding and rat (Rattus spp.) population eruptions on Stewart Island/Rakiura

Abstract

This study aimed to quantify changes in rat abundance and population structure before, during, and after a rimu (Dacrydium cupressinum) mast seed event in lowland forest on Stewart Island, New Zealand. Rats, primarily ship rats (Rattus rattus), were trapped in low numbers throughout the study period (March 2000‐March 2003), except when they erupted to very high abundance in spring 2002, shortly after heavy rimu seed fall. In the immediate post‐peak phase, scavenging of trapped rats increased substantially; rats were seen and trapped in daylight; and weights of adult female rats were low in relation to their size, which suggests that food shortage was the cause of the subsequent steep decline in abundance. Rat eruptions have been observed on Stewart Island after heavy rimu seed fall several times over the past 40 years. Eruptions of rats caused by heavy rimu seed‐fall may have triggered the invasion of nearby islands by rats, and caused the extinction of several native species on Stewart Island.     

Eradications as reverse invasions: lessons from Pacific rat (Rattus exulans) removals on New Zealand islands

Eradications of kiore or Pacific rats (Rattus exulans) from islands around New Zealand have been followed by responses from resident species of coastal plants, invertebrates, reptiles and seabirds. These responses are compared with an invasion by ship rats (Rattus rattus), which devastated populations of invertebrates, birds and bats. Post-eradication responses only approximate the effects of invasions because recovery is limited to the residual pool of native species. Greater effects of kiore are indicated by adding incompatible species confined to rat-free locations. The extended list includes at least 15 species of invertebrates, two species of frogs, tuatara (Sphenodon punctatus), 11 species of lizards and 9 species of seabirds. The analyses indicate direct and indirect effects of kiore similar to those reported after ship rat invasions. This is despite indications from the literature that kiore are the least damaging of the three commensal rat species.     

The end of an 80-million year experiment: a review of evidence describing the impact of introduced rodents on New Zealand’s ‘mammal-free’ invertebrate fauna

Since separating from its super-continental origin 80 million years ago, New Zealand has effectively been isolated from the impacts of terrestrial mammals. The arrival of Polynesians in 13th C heralded the end of this era, with the introduction of kiore, (Rattus exulans, or Pacific rat), which had far-reaching effects on plant regeneration, survival of small ground vertebrates, larger invertebrates, and seabird breeding colonies. This paper reviews the evidence available from raptor nest sites and Quaternary beetle fossils to summarise extinctions thought to be caused by kiore in New Zealand. It also utilises invertebrate comparisons between islands with and without rats, or where rats have been eradicated, in order to document the impacts of rats (R. exulans, R. norvegicus) on invertebrate abundance, body mass, and the behavioural responses of some large New Zealand insects to the presence of rats. The role of a ‘mammal-free’ evolutionary history is discussed.     

Habitat selection of feral cats (Felis catus) on a temperate,forested island

Abstract Habitat selection of mammalian predators is known to be influenced by availability and distribution of prey. The habitat selection of feral cats on Stewart Island, southern New Zealand, was investigated using telemetry of radio‐tagged cats. Compositional analysis of the habitat selection of radio‐tagged cats showed they were using the available habitats non‐randomly. Feral cats avoided subalpine shrubland and preferentially selected podocarp‐broadleaf forest. The avoidance of subalpine shrubland by cats was probably due to a combination of the presence of a large aggressive prey species, Norway rats Rattus norvegicus, and the lack of rain‐impervious shelter there. Most cats also used subalpine shrubland more often in dry weather than in wet weather. Cats did not preferentially select all the other habitats with only smaller rat species, Rattus rattus and Rattus exulans, present however. Cats were probably further influenced by the availability of large trees, in podocarp‐broadleaf forest, that can provide shelter. Cats were also more active in dry rather than wet weather which supports this conclusion. Home ranges of feral cats on Stewart Island were some of the largest recorded, probably because of limited primary and alternative prey.     

Competition, coexistence, and adaptation amongst rodent invaders to Pacific and New Zealand islands

Variation in skull size was investigated for three species of rats (kiore –Rattus exulans Peale; ship rat –R. rattus L.; Norway rat –R. norvegicus Berkenhout) which were introduced by humans to various islands in New Zealand and other Pacific islands. Data from seventy-one islands and 882 specimens are examined for evidence of the effects of latitude, island size and interspecific competition among rats and the house mouse (Mus musculus L.) on skull size, using multiple regressions. For R. exulans, skull size increases with latitude as predicted by Bergmann's rule, but no such effect occurs for the other two rats. There was a positive relationship between island size and the number of species inhabiting it, and some species combinations were more likely to occur than others. For example, R. exulans and R. norvegicus were more likely to occur together, while R. rattus and R. exulans were rarely sympatric. R. exulans and R. rattus skull size was negatively correlated with the number of other rodents on the same island. R. exulans skull size increased on smaller islands in some island groups, perhaps because increased density and consequent increased intraspecific competition on smaller islands favours increased body size. This effect is more pronounced in tropical islands (Solomon islands), than in subtropical ones (Hawaiian islands) and less so in temperate New Zealand. Collectively the data demonstrate that rapid evolution of body size in predictable directions can follow within 150 years of the introduction of species to new receiving communities.     

From small Maria to massive Campbell: Forty years of rat eradications from New Zealand islands

Abstract

Over the last four decades the eradication of rats from islands around New Zealand has moved from accidental eradication following the exploratory use of baits for rat control to carefully planned complex eradications of rats and cats (Felis catus) on large islands. Introduced rodents have now been eradicated from more than 90 islands. Of these successful campaigns, those on Breaksea Island, the Mercury Islands, Kapiti Island, and Tuhua Island are used here as case studies because they represent milestones for techniques used or results achieved. Successful methods used on islands range from bait stations and silos serviced on foot to aerial spread by helicopters using satellite navigation systems. The development of these methods has benefited from adaptive management. By applying lessons learned from previous operations the size, complexity, and cost effectiveness of the campaigns has gradually increased. The islands now permanently cleared of introduced rodents are being used for restoration of island‐seabird systems and recovery of threatened species such as large flightless invertebrates, lizards, tuatara, forest birds, and some species of plants. The most ambitious campaigns have been on remote subantarctic Campbell Island (11 300 ha) and warm temperate Raoul Island (2938 ha), aimed to provide long‐term benefits for endemic plant and animal species including land and seabirds. Other islands that could benefit from rat removal are close inshore and within the natural dispersal range of rats and stoats (Mustela erminea). Priorities for future development therefore include more effective methods for detecting rodent invasions, especially ship rats (Rattus rattus) and mice (Mus musculus), broader community involvement in invasion prevention, and improved understanding of reinvasion risk management.     

Rodents and their predators in the eastern Bay of Islands

Abstract

Traps were set for rodents and mustelids on five islands (Motukiekie, Moturua, Okahu, Urupukapuka, and Waewaetorea) in the eastern Bay of Islands in March 1984. Kiore (Rattus exulans) were caught on Moturua Island and Norway rats (R. norvegicus) on all five islands, but no mustelids were caught or seen. Kiore on Moturua Island were very scarce compared with other northern offshore islands, perhaps because of competition from Norway rats and the presence of stoats and cats. Kiore were breeding and young matured in the season of their birth. Norway rats were scarce and found mainly near the shoreline on four of the islands. On Waewaetorea Island they were plentiful and widespread despite the possible presence of stoats. About a third of the mature females were visibly pregnant. Average litter size was 6.9 embryos, and 44% of the parous females had borne two or three litters. Females first ovulated at 180 ± 5 g weight and 356 ± 5 mm total length on average. Males first produced sperm at 189 ± 7 g weight and 364 ± 4 mm total length. Most rats matured before reaching a tooth-wear age index of 5.     

Grey-faced petrel (Pterodroma gouldi) productivity unaffected by kiore (Pacific rats,Rattus exulans) on a New Zealand offshore island

Burrow-nesting seabird populations are vulnerable to predation by introduced rats, because of their nesting habits and slow life histories. We investigated whether control of kiore (Pacific rats, Rattus exulans) by removal trapping, and during an unsuccessful community-led island-wide eradication attempt, had any effects on nest survival of grey-faced petrels (Pterodroma gouldi) on Ririwha (Stephenson Island), northeastern New Zealand. We compared nest survival between two plots at which rats were trapped and six un-trapped plots in 2010, as well as at all plots during and after the poisoning programme in 2011–2012. Neither mean rates of breeding burrow occupancy nor nest survival differed between trapped and un-trapped plots in 2010. We found no significant differences between years or between plots throughout the poisoning programme. Extrapolation of daily nest survival rates to the full 172-day combined egg and chick period gave an estimate of mean annual productivity for all plots combined of 0.285 (95% confidence interval 0.252–0.318), which is higher than on comparable predator-free islands. Although the absence of a detectable effect of kiore on breeding grey-faced petrels on Ririwha is reassuring, we can be less sure that smaller burrow-nesting seabirds on the island are secure.     

Invertebrate survey of coastal habitats and podocarp forest on Ulva Island,Rakiura National Park,New Zealand

Abstract

Inventory of the invertebrate fauna is important to establish taxonomic diversity, abundance and distribution, and hence the conservation of indigenous biodiversity. Invertebrate assemblages have been documented in some broadleaf‐podocarp forests and grassland habitats in New Zealand, but not in dense stands of coastal forest or in mature podocarp forest. This survey aimed to provide a taxonomic inventory of terrestrial invertebrates and their habitat associations on Ulva Island (Rakiura National Park, Stewart Island), an off‐shore sanctuary of significant conservation value in New Zealand. We systematically documented the invertebrate assemblages collected in ground litter and on tree trunks on the island. The invertebrate specimens identified represented 4 phyla, 6 classes, 25 orders and 62 species. The invertebrate fauna reported in this survey was distinct from those of lowland shrubland and broadleaved‐Nothofagus forests on the mainland, but shared species with that reported from another similar off‐shore island, Codfish Island (Whenua Hou).     

An inventory of mammalian pests in a New Zealand city

Abstract

Urban ecosystems include many habitat types supporting native flora and fauna. These habitats may also sustain populations of introduced mammalian pests, although relatively little is known about the composition or distributions of these species in urban environments. We made a preliminary survey of the distribution and relative abundance of pest mammals across three urban habitat types (gully, amenity park, residential) in Hamilton, New Zealand. Tracking tunnel and WaxTag® surveys showed that: (1) rats (Rattus rattus or R. norvegicus), mice (Mus musculus) and brushtail possums (Tricho‐surus vulpecula) were detected most often and in highest abundances in gullies, were relatively rare in amenity parks, and were not detected in residential areas; (2) hedgehogs (Erinaceus europaeus) were regularly detected in all habitats; and (3) mustelids were not detected in any habitat. Live trapping in areas where rats were detected captured 21 rats, of which 19 were R. rattus. Scat and animal sign surveys found evidence of lagomorphs in amenity parks and gullies but not in residential areas. Cats (Felis catus) were detected in all habitats. These data suggest that important mammalian pests other than cats are either absent (mustelids) or comparatively rare (rats) in urban Hamilton and largely confined to gully habitats. Further research is needed to determine whether detection probabilities of target species vary significantly between the species and habitats we sampled, and to quantify the impact that mammalian pests might have in limiting populations of other animals, especially avifauna.     

Eradication of Norway Rats for Recovery of Seabird Habitat on Langara Island, British Columbia

Introduced Rattus norvegicus (Norway rats) caused the decline of Synthliboramphus antiquus (ancient murrelets) and other seabirds breeding on Langara Island (approximately 3,100 ha), British Columbia. Using funds from the litigation settlement following the Nestucca oil spill, Environment Canada eradicated Norway rats using a technique developed in New Zealand which involved dispensing wax baits containing the anticoagulant brodifacoum at 50 ppm from fixed bait stations. Bait stations were placed every 75 to 100 m on a grid over the entire island (1 station/ha). Rats removed bait for 26 days, after which crews placed baits in protective plastic bags in each bait station. Stations loaded with baits were left on the island and rechecked four times over 2 years, after which bait stations and remaining bait were removed. The eradication succeeded. No signs of rats have been detected on Langara Island and its associated islands since January 1996. No rats were trapped during 1,700 trap‐nights following the poison campaign. Incisor marks of rats were not found on apples or oil‐dipped chew‐sticks. Corvus corax (common ravens) likely suffered greater than 50% mortality from the eradication after apparently gaining access to the poison directly from bait stations and from scavenging rat carcasses. A monitoring and response system is being developed in conjunction with current users of the islands. The success on Langara Island demonstrates how the technique proven on small New Zealand islands of less than 300 ha can be effectively extrapolated to much larger islands.     

Have the Harmful Effects of Introduced Rats on Islands been Exaggerated?

Introduced rats are now being eradicated from many islands. Increasingly, these eradications are contested by activists claiming moral, legal, cultural, historic or scientific reasons and poorly documented evidence of effects. We reviewed the global literature on the effects of rats on island flora and fauna. We then used New Zealand as a case study because of its four-decade history of rat eradications and many detailed and innovative studies of how rats affect native species. These include use of exclosures, local manipulations of rat populations, video surveillance, and measurements of responses following eradications. The most intensive studies have been on the Pacific rat (Rattus exulans), a small South-East Asian species spread by Polynesians throughout the Pacific. These and the more recently introduced Norway rat (R. norvegicus) and ship (roof) rat (R. rattus) suppress some forest plants, and are associated with extinctions or declines of flightless invertebrates, ground-dwelling reptiles, land birds, and burrowing seabirds. On islands off France, Norway rats are also implicated in declines of shrews. Globally, ship rats were associated with declines or extinctions of the largest number of indigenous vertebrate species (60), including small mammals such as deer mice and bats. Effects of rats on forest trees and seabird populations are sufficiently pervasive to affect ecosystem structure and function. However, the data are patchy. Deficiencies in our knowledge would be reduced by documenting distribution and abundance of indigenous species before and after eradications. Comprehensive measurements of the responses of indigenous species to rat eradications would enable the development of testable models of rat invasion effects.     

Response of a Tree Weta Population (Hemideina crassidens) After Eradication of the Polynesian Rat from a New Zealand Island

Abstract A population of Hemideina crassidens (the Wellington tree weta) was monitored over a 4‐year period after the eradication of Rattus exulans (the Polynesian rat kiore) and Gallirallus australis australis (the South Island weka) from Nukuwaiata (Chetwode Islands), Pelorus Sound, New Zealand. A novel survey technique (entrance scores) was used in combination with a conventional technique (random searches for active weta) to measure changes in weta population parameters after the removal of predation pressure and to investigate impacts of exotic predators on tree weta. Tree weta density did not increase markedly over the 4‐year period, but the proportion of active adults did increase. Weta were observed to move into larger and more crowded galleries (refuges), to occupy galleries closer to the ground, and to spend less time sitting in gallery entrances. It was concluded that endemic tree weta are well adapted to withstand some introduced vertebrate predators but are able to live a more “relaxed” lifestyle in the absence of this predation. The most significant change detected was in weta age structure, with adults increasing their proportion of the population.     

Habitat use by three rat species (

Ship rats (Rattus rattus) were removed from sites on Pearl Island, southern Stewart Island, in 2004 and 2005, to test whether they excluded Pacific rats (R. exulans) or Norway rats (R. norvegicus) or both from podocarp-broadleaf forest. As predators can influence habitat use in rodents, Pearl Island was selected because no mammalian predators of rodents are present. Rats were trapped in two other habitats to clarify rat distribution on the island and to obtain samples for stable isotope investigation of food partitioning within habitats. The experimental removal of ship rats failed, as Pacific rats were found to share forest and shrubland with ship rats. This result contrasted with the restricted distribution of Pacific rats on Stewart Island. Ship rats were ubiquitous, and appear to have been the dominant species in podocarp-broadleaf forest on Pearl Island. The largest species, the Norway rat, was trapped only on the foreshore of Pearl Island, but on Stewart Island it is more widespread. Ship rats and Norway rats were partitioning the coastal habitat by exploiting different food sources. Stable isotopic ratios (δ¹⁵N and δ¹³C) in muscle samples from Norway rats revealed a strong marine signature, suggesting intensive foraging in the intertidal zone. Ship rats trapped in the same habitat exhibited mixed terrestrial and marine sources in their diet. There was little obvious partitioning between ship rats and Pacific rats in forest, except a possible delay in breeding in Pacific rats relative to ship rats. Whether Norway rats select the intertidal zone to forage, or were excluded from forest by ship rats is unknown, but competitive exclusion is likely. Estimated densities of rats were low (2.1–5.1 rats ha^-1 in forest, 1.42 rats ha^-1 in shrubland) and similar to other New Zealand sites with low soil fertility. Further research will be required to elucidate the roles of food quality, habitat structure and predation in facilitating habitat selection in these species.     

Population dynamics of Rattus argentiventer, Rattus losea, and Rattus rattus inhabiting a mixed-farming system in the Red River Delta, Vietnam

Rodent pests cause significant damage to lowland irrigated rice crops in the Red River Delta of northern Vietnam. Data from a 4-year study were examined to look at the population dynamics of the ricefield rat, Rattus argentiventer (representing 50% of captures), the lesser ricefield rat, Rattus losea (30% of captures), and the black rat, Rattus rattus complex (9% of captures) that inhabit the irrigated mixed-cropping system. We tested the hypothesis that these rodent species were breeding in response to the availability of high-quality food provided by crops rather than in response to rainfall. The abundance of rodents fluctuated annually, with a main peak following the spring rice crop, and a secondary peak following the summer rice crop. There was a strong relationship between the monthly abundance of rats and rainfall, but a weak relationship between monthly rates of increase and rainfall. There were distinct peaks in breeding activity during the reproductive stages of the rice crops suggesting that changes in crop stages were more important than rainfall in this seasonal, but irrigated agroecosystem. The modal litter size for R. argentiventer was 8 (mean of 8.67±0.20 SE, range 2–16), where the mode for R. losea, was 7 (mean of 7.32±0.15 SE, range 3–14). Management of these species needs to be conducted prior to the onset of the main breeding seasons.     

The Diet of Feral Cats (

Feral cats became established on Raoul Island some time between 1836 and 1872; the prey available to them included a great variety of nesting seabirds, few of which are present now, landbirds and kiore (Rattus exulans). Norway rats reached the island in 1921, providing additional prey for cats, but also another potential predator of seabirds. The diet of cats is described from guts and scats collected between 1972 and 1980. Rats are the main food, with land birds second in importance, and seabirds are now a minor item. More than 90% of the rats eaten by cats are kiore although more Norway rats than kiore are trapped. Eradicating cats from Raoul Island is feasible but because Norway rats too are important predators of birds on islands, it is likely that eradicating cats without also eradicating Norway rats will do little to restore the diversity of bird species on Raoul Island, although the densities of a few species now present might be increased.     

New Zealand's giant Collembola: New information on distribution and morphology for Holacanthella Börner, 1906 (Neanuridae: Uchidanurinae)

Abstract

The New Zealand collembolan fauna currently includes five described species of the striking endemic genus Holacanthella (springtails). Holacanthella species are saproxylic decomposers of cool temperate forest ecosystems, and they contribute to nutrient cycling of coarse woody debris. All species of the genus have the dorsal and lateral surfaces furnished with conspicuous red, orange, yellow or white digitations. They are among the largest Collembola known, with some individuals reaching 17 mm in length. We examined new material of the five species from throughout New Zealand, as well as existing museum material, to understand more fully their distributions. We provide an updated key to the five Holacanthella species and discuss the conservation of rare/range restricted species, and propose hypotheses to explain their distributions. One species, H. laterospinosa, is known only from Cuvier Island and the Coromandel Peninsula, North Island, but the distributions of the remaining four species appears to reflect the turbulent geological history of New Zealand during the Pliocene. Intraspecific variation in several gross morphological characters was observed in all species. We provide a comparison of characters with the other genera within the Uchidanurinae in order to characterise the genus more fully with respect to allied genera. Holacanthella species are particularly vulnerable to human‐mediated disturbance by forest modification because of their strict habitat requirements and low mobility, and we stress that conservation efforts should focus on protecting critical habitats for each species.