Kiore (Rattus exulans) distribution and relative abundance on a small highly modified island

ABSTRACT

Kiore (Pacific rat; Rattus exulans) is both a target for eradication and a taonga or highly valued species in New Zealand, and its abundance and distribution vary considerably throughout the country. We investigated reports of an abundant kiore population on Slipper Island (Whakahau), off the east coast of New Zealand’s North Island, in March 2017. We trapped kiore to examine their distribution across a range of habitats with varying degrees of human activity. Kiore were captured in all habitats, with particularly high abundance at a campground with a fruiting fig tree (50 kiore per 100 trap nights corrected for sprung traps). We found no evidence of other rat species; Slipper Island appears to remain one of few New Zealand islands with kiore but without ship rats (Rattus rattus) and Norway rats (R. norvegicus), the two other rat species present in New Zealand. Slipper Island potentially provides opportunities to research kiore behaviour and population dynamics in a New Zealand commensal environment, and genetics of an isolated island population.     

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.     

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.     

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.     

Small mammals as potential seed dispersers in New Zealand

Weed invasion success is strongly influenced by availability of seed dispersal vectors, which may include animals. We examined the potential of several small introduced mammals (mice, kiore, ship rats and possums) to disperse germinable seeds in New Zealand. Captive animals were fed fleshy fruit of weeds (Berberis glaucocarpa, Cotoneaster spp., Crataegus monogyna, Ilex aquifolium, Leycesteria formosa, Ligustrum sinense, Lonicera japonica, Passiflora mollissima, Pyracantha angustifolia, Sorbus hupehensis) and native species (Coprosma spp., Prumnopitys ferruginea and Solanum aviculare). We recorded the percentage of fruit consumed, seed ingested and gut passage time. Faeces were collected and the seeds extracted and tested for germination potential in an unheated glasshouse (two weed species) or under controlled conditions (11 species). The smallest rodents (mice and kiore) generally destroyed all seeds eaten. Large numbers of viable seeds of the small‐seeded (<1 mg) species, L. formosa and S. aviculare, passed through ship rats. Possums consumed the seeds of all adventive and native fruits except P. ferruginea. The proportion of seeds recovered intact from possum faeces varied with plant species and ranged from 6 to 83%. The time required for 50% of all seeds to be passed by possums ranged from 2.5 to 5.5 days with an average of 3.7 days, and was generally unrelated to simple fruit parameters such as percentage pulp and moisture content. For seeds where germination also occurred in the uneaten controls, the germination of seed from possums ranged from 3 to 78%. Germination was mostly lower in seeds from possums than in the controls, where differences were significant. Possums have major potential to disperse a wide range of fleshy fruit‐producing native and introduced plant species. Ship rats have the potential to disperse those with very small seeds.     

Diphacinone with cholecalciferol for controlling possums and ship rats

ABSTRACT

Potent second-generation anticoagulant rodenticides such as brodifacoum have been used as more effective alternatives to first-generation anticoagulants, such as warfarin. A combination of diphacinone at 0.005% and cholecalciferol at 0.06% produces a slow-acting bait that is effective at killing possums (Trichosurus vulpecula) and rodents. Cage trials with groups of possums and ship rats (Rattus rattus) achieved a mortality of 87% and 86% for possums and ship rats, respectively. Two field trials, each 200 hectares in size, targeting possums, ship rats and mice achieved an average reduction in the abundance of 94% for possums, 94% for ship rats and 80% for mice. The combination of diphacinone and cholecalciferol appears effective and has a favourable risk profile compared with second-generation anticoagulant rodenticides, such as brodifacoum. Approval of this new bait by the New Zealand Environmental Protection Agency was granted in 2018 and final registration obtained from the Ministry of Primary Industries in 2019.     

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.     

Hoarding behavior by ship rats (Rattus rattus) in captivity and its relevance to the effectiveness of pest control operations

Hoarding of food items is well known among muroid rodents, but evidence for hoarding behavior among ship rats (Rattus rattus) is scant. Here, we characterize hoarding behavior in ship rats maintained in captivity after capture from the wild. After acclimatization to captivity, 40 ship rats (21 females, 19 males) were presented with baits in experiments designed to emulate a typical poison control operation for vertebrate pests in New Zealand: this involved first offering rats nontoxic cereal baits (of 2- or 6-g size) as a prefeed for three nights consecutively, followed by 6- or 12-g cereal baits laden with 0.15% 1080 on the fourth night. Seventy-eight percent of rats (31/40) hoarded food in distinct cache sites when presented with nontoxic baits although there was no significant effect of bait size or type on hoarding behavior and nor did hoarding behavior vary according to rat gender. When rats were presented with 1080-laden baits, the incidence of hoarding was reduced to 40%, due to the onset of toxicosis. This study indicates that R. rattus will show hoarding behavior analogous to other rat species when presented with an excess of cereal-based baits, at least under conditions of captivity and free from competition. This finding may have practical relevance: since 1080 is the principal toxin used against the major vertebrate pest species in New Zealand (the brushtail possum, Trichosurus vulpecula), ship rats have the potential to deplete supplies of prefeed and/or toxic baits intended for possum control. However, based on typical rat densities recorded in New Zealand native forest (c. 5 rats/ha), the degree of removal and manipulation of toxic baits observed by ship rats here is unlikely to impact adversely on the efficacy of possum control operations.     

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.     

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.     

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.     

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.     

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.     

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.     

The home range of ship rats (Rattus rattus) in beech forest in the Eglinton Valley,Fiordland, New Zealand: A pilot study

Abstract

Three male and two female ship rats (Rattus rattus) were radio‐tagged and tracked in beech (Nothofagus) forest in the Eglinton Valley, Fiordland, New Zealand over two field periods in 1996 and 2000. The home range of each animal was calculated using the minimum convex polygon method. Ranges of three male rats were 7.5, 9.1, and 11.4 ha whereas those of the female rats were 0.89 and 0.27 ha. The home ranges recorded for male rats were considerably larger than those reported from other studies in non‐beech forest. Ship rats are important predators of forest birds, and home range information could be used to provide a guide for trap or bait station spacing in beech forests. To carry out rat control in beech forests effectively, further studies are needed to determine if the results of this pilot study are typical, and if home ranges of ship rats change with season, or at various stages of the beech mast cycle.     

Denning behaviour of ship rats (Rattus rattus) on Taukihepa,a seabird breeding island

Abstract

Den sites of 14 ship rats (Rattus rattus) were located daily during the rat breeding season on Taukihepa (Big South Cape), a seabird island southwest of Rakiura (Stewart Island). In contrast to other New Zealand studies, no arboreal dens were found. Den sites on Taukihepa were in ferns, under logs, in woodpiles, or underground in sooty shearwater (Puffinusgriseus) breeding burrows. The number of times known den sites used was positively related to the amount of leaf litter and woodpiles near the den sites. Overall, 24% of radio‐tagged rats were sharing den sites on any given day. While there was considerable individual variation in the number of times den sites were used, female rats tended to reuse den sites more than males. Many rats were found in dens alone, but frequently males and females shared. Occasionally two females and one male denned together, as did two females, whereas males never denned with another male.     

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.     

Distribution and movements of Buller's albatross (Diomedea bulleri) in Australasian seas

Abstract

The distribution and movements of Buller's albatross in Australasian seas are analysed using results of shipborne surveys (13 238 10‐min counts), counts from trawlers, banding data, recoveries on beaches and fishing vessels, and records from the literature. Patterns of marine distribution are documented by monthly accounts and maps. During the breeding season, highest abundances are recorded over shelves and slopes off southern New Zealand (The Snares shelf to 41–43°S off the South Island, D. b. bulleri), around the Chatham Islands and over oceanic subtropical waters east of New Zealand (probably D. b. platei), with marked seasonal variations observed off southern New Zealand. Both subspecies disperse mostly outside Australasian waters during the non‐breeding season. Birds banded on The Snares were recovered off south‐eastern New Zealand (Stewart Island to Cook Strait) and in the eastern tropical Pacific. Immatures accounted for only 0.25% of birds censused during the ship‐borne surveys; they are recorded around the New Zealand mainland in August‐October and February‐May, off south‐eastern Australia and in the Tasman Sea in November‐December, February, and June‐July. Around New Zealand, males predominate among birds recovered along the eastern seaboard, whereas the sex ratio in south‐western waters tends to vary according to water depth and season. Distribution patterns and movements in New Zealand and Australian seas are discussed in relation to breeding events and breeding status.     

Polytrichum commune Hedw. and Polytrichadelphus magellanicus (Hedw.) Mitt. used as decorative material on New Zealand Maori cloaks

Abstract

Confirmation of the occurrence of the moss Ptychomnion densifolium on the main islands of New Zealand is provided and the features which separate it from the more common P. aciculare are outlined. A brief account of the history of P. densifolium in New Zealand is given and its morphology there is discussed. A key to the two species in New Zealand is provided and habitat differences between them outlined. New Zealand material differs from type locality (Tristan d' Acunha) material in having longer and more twisted acumens, and in this respect is similar to New Zealand material of P. aciculare. New Zealand populations of P. densifolium are recognisable in having stem leaves reflexed from an oblong sheathing base, and well developed basal leaf plications. It is a relatively widespread species in New Zealand, occurring in mostly upper elevation, open sites.