Movements, behaviour and survival of adult Cook Strait giant weta (Deinacrida rugosa; Anostostomatidae: Orthoptera) immediately after translocation as revealed by radiotracking

Giant weta (Orthoptera: Anostostomatidae) are large flightless New Zealand insects vulnerable to predation from introduced mammals. Some species have been transferred to islands or mammal-free mainland sanctuaries to establish additional populations. Radiotelemetry was used to investigate behaviour, movements and survival of adult Cook Strait giant weta (Deinacrida rugosa) immediately after translocation into Karori Sanctuary, New Zealand, to describe their initial movements, and to assess the importance of this establishment phase in relation to the long-term viability of the population. The average distance moved between consecutive daytime refuges for translocated male D. rugosa within Karori Sanctuary was significantly further than for resident weta on Matiu-Somes Island. In contrast, translocated female weta moved significantly smaller distances between consecutive daytime refuges within Karori Sanctuary than those on Matiu-Somes Island. Translocated D. rugosa travelled significantly further between consecutive daytime refuges between 19 and 45?days after release than during the first 19?days and more than 45?days of radiotracking. Deinacrida rugosa survived well following translocation and there was only limited evidence of predation despite an increased abundance of indigenous avian and reptilian predators being present, and the presence of low numbers of mice. The establishment potential of this population was not adversely affected by movements and survival of the weta immediately after translocation. It still remains to be seen if a self-sustaining population of D. rugosa develops in Karori Sanctuary but the indications are that the species is present because progeny of the translocated weta are regularly seen within Karori Sanctuary. Radiotelemetry provided valuable insights into the behaviour of adult D. rugosa and it could be appropriate for monitoring other large bodied invertebrates.     

Saved from extinction? Establishment and dispersal of Mercury Islands tusked weta,Motuweta isolata,following translocation onto mammal-free islands

The Mercury Islands tusked weta, Motuweta isolata (Orthoptera: Anostostomatidae), survived only on 13 ha Ahu or Middle Island, a mammal-free island in the Mercury Group, New Zealand. Between 2000 and 2009, 567 individuals were translocated in nine releases to six nearby islands from which mammals had been removed. These translocations occurred to reduce the chance of accidental extinction of the Middle Island population of only a few hundred adults and to contribute to the restoration of the other islands. All translocated insects originated from the captive-bred progeny of one male and two females collected from Middle Island between 1998 and 2001. Their establishment on Double and Red Mercury Islands, after their releases in 2000 and 2001 respectively, was confirmed by searching plots, and by using footprint tracking tunnels on Red Mercury Island between 2008 and 2012. Tracking tunnels provided better data and proved more cost effective than searching plots for detecting large tusked weta. Tracking tunnels demonstrated that the population on Red Mercury expanded outwards from the release sites by 50–100 m each year between 2009 and 2012. These weta are now estimated to be present over more than half the Island. Tusked weta have also survived on Stanley, Korapuki and Ohinau Islands after releases in 2007, but they remain within 100 m of the release sites. No confirmed progeny of the weta released on Cuvier Island in 2008 and 2011 were detected. No tusked weta were detected on Middle Island using tracking tunnels on eight occasions between 2009 and 2012, suggesting this species is likely to be locally extinct. Despite possible failure on one island, these translocations have resulted in a significant conservation success outcome.     

Male tree weta are attracted to cuticular scent cues but do not discriminate according to sex or among two closely related species

Recognition of conspecifics is an essential precursor of successful mating. Where related species coexist, species discrimination might be important, but because related species are similar, species signal recognition may actually be low. Chemical cues such as cuticular hydrocarbons (CHCs) are frequently used by insects to identify suitable sexual partners. We predicted that New Zealand tree weta (Hemideina spp.), a genus of nocturnal ensiferan Orthoptera that live both allopatrically and sympatrically, use chemical signals from either frass or CHCs to find mates. In a series of six laboratory trials using both H. thoracica and H. crassidens, we found that male tree weta, but not female tree weta, occupied cavities primed with female cuticular cues more often than cavities without. However, males did not discriminate between chemical cues of male and female conspecifics, or between conspecifics and heterospecifics. In field trials, tree weta did not occupy artificial cavities primed with either female frass or female cuticular cues more often than unscented cavities. However, in both trials weta preferentially returned to cavities that had already been occupied earlier in the trials. A final field trial confirmed the presence of mixed species harems during the mating season in one region of sympatry. Our results suggest that selection on sex and species specific chemical cues that could be used to find conspecific mates is weak. Mixed species aggregations suggest that identification of conspecific mating cues has not evolved to be species specific. We infer that for male tree weta, the cost of mating with heterospecifics is likely less than not mating at all.     

Mark-recapture study of mountain stone weta

Mountain stone weta (Hemideina maori) on the Rock and Pillar Range in the South Island, New Zealand, are found primarily in cavities under flat rocks on isolated outcrops or 'tors'. We marked 66 adult weta on one tor and 30 adults on an adjacent tor and recorded their location during the summer and for the following three years to obtain baseline data on survival, longevity, dispersal, and movement within tors. It was not uncommon for adult weta to live for two to three years. Most marked weta were resighted at least once, usually under the same rock. Few weta moved further than the rock adjacent to where they were first captured and only one dispersed to a neighbouring tor. On one tor, a relatively stable group of 6-8 females and 2 males was resighted during most of the summer period under one large rock. An analysis of our mark-recapture methodology and results indicates that H. maori may be an appropriate species for investigating population structure in a metapopulation context as well as local mate competition.     

Life cycle, survival rates and longevity of an alpine weta

Although reproductive and behavioural studies have been conducted on captive tree weta, there have been very few ecological field studies of any of the weta species involving free-ranging, marked individuals. The mountain stone weta (Hemideina maori) is a tree weta that lives on rock tors in the alpine region of the South Island of New Zealand. Over three seasons each of 480 adults and 789 juveniles was individually marked on four large and 14 small tors to gather baseline information on aspects of H. maori’s life cycle and life history. Seasonal patterns were seen in the appearance of the smallest nymphs, moulting, and in the survival and recruitment of adults. Some marked juveniles were recaptured after 10 or more months, with a maximum interval of 14.1 months, indicating that instar intervals can be exceptionally long. Adult males and females had similar survival rates and often lived for 2 or 3 breeding seasons. Relatively high recapture rates (~60–70%) and long life spans make adult H. maori amenable to modern mark-recapture analyses using the programme MARK. We believe this research will be a useful template for further mark-recapture studies such as those to verify life history characteristics of endangered species of weta or the effects of secondary poisoning on invertebrates.     

Seed dispersal effectiveness increases with body size in New Zealand alpine scree weta (Deinacrida connectens)

Weta are giant, flightless orthopterans that are endemic to New Zealand. Although they are known to consume fleshy fruits and disperse seeds after gut passage, which is unusual among insects, their effectiveness as seed dispersal mutualists is debated. We conducted a series of laboratory experiments on alpine scree weta (Deinacrida connectens) and mountain snowberries (Gaultheria depressa) to investigate how fruit consumption rates, the proportion of ingested seeds dispersed intact and weta movement patterns vary with weta body sizes. On average weta dispersed 252 snowberry seeds nightly and travelled at a rate of 4 m min^?1. However, seed dispersal effectiveness varied over three orders of magnitude and was strongly associated with body sizes. Smaller weta consumed few snowberry seeds and acted primarily as seed predators. On the other hand, the largest weta consumed and dispersed thousands of seeds each night and appear to be capable of transporting seeds over large distances. Overall results indicate that scree weta shift from being weakly interacting seed predators to strongly interacting, effective seed dispersers as they increase in size.     

Mutualism or opportunism? Tree fuchsia (Fuchsia excorticata) and tree weta (Hemideina) interactions

Mutualisms or interspecific interactions involving net mutual benefits, are an important component of ecological theory, although effectively demonstrating mutualism is notoriously difficult. Among two New Zealand endemics, a slightly elevated germination rate of Fuchsia excorticata (Onagraceae) seeds after passage through tree weta (Orthoptera: Anostostomatidae) compared with seeds manually extracted from fruit, led to the proposal that a mutualistic relationship exists between this plant and animal. An improved germination rate, or any other single trait, however, does not alone constitute evidence for mutualism; the relative costs and benefits of numerous components of the interaction need to be accounted for. We considered the costs and benefits to F. excorticata of the putative seed dispersal mutualism with tree weta. Tree weta provided with F. excorticata fruits destroyed 78% of the seeds they consumed, did not move fruit; and faeces containing seeds were deposited near their roost holes (which are naturally in trees). The seeds remaining after fruit consumption and those that are ingested but survive gut passage are unlikely to be deposited in suitable habitat for seedling survival. Plant food preferences of captive tree weta assessed using pairwise leaf choice tests showed that the leaves of F. excorticata were the least preferred of six commonly encountered plants. In addition, we found that tree weta did not show a preference for F. excorticata fruit over a standard leafy diet, indicating they are unlikely to be actively seeking fruit in preference to other sources of food. These observations indicate that any interaction between tree weta and F. excorticata is likely to be opportunistic rather than mutualistic, and highlight the difficulty of characterizing such interactions.     

Brodifacoum residue analysis in water, soil, invertebrates, and birds after rat eradication on Lady Alice Island

Rats were eradicated from Lady Alice Island (Northland, New Zealand) in October 1994, using aerially applied cereal-based bait containing brodifacoum. To determine the fate and non- target impact of brodifacoum, streams, soil, invertebrates, and birds were monitored for 7 months after the baits were applied. No brodifacoum was detected in any of the stream or soil samples. Brodifacoum was detected in cave weta found on baits, and in morepork and red-crowned parakeet liver tissue. Significant contamination of water and soil is unlikely after a single aerial application of brodifacoum baits. Invertebrates such as cave weta may be at risk of consuming brodifacoum bait, morepork and red-crowned parakeet may be at risk of secondary poisoning.     

Assessment of non-target impact of 1080-poisoning for vertebrate pest control on weta (Orthoptera: Anostostomatidae and Rhaphidophoridae) and other invertebrates in artificial refuges

Artificial refuges and mark-recapture techniques were used to monitor the non-target impacts of handbroadcast application (simulating aerial application) of Wanganui No.7 cereal-based baits containing 0.15% (1500 µg g-1) 1080 on populations of weta and other invertebrates in Tararua Forest Park, North Island, New Zealand. Wellington tree weta (Hemideina crassidens) and a cave weta (Isoplectron sp.) were the only species of weta that occupied the refuges. Flatworms, slugs, spiders, harvestmen, amphipods, millipedes, centipedes, cockroaches, and beetles also occupied the refuges. Invertebrate numbers in the refuges were monitored for 12 months before and 4 months after bait application on 22 August 2000. Bait application had no significant impact on the numbers of either species of weta, or on slugs, spiders, and cockroaches, the most numerous other invertebrates occupying the refuges. Bait application also had no effect on the number of individually marked tree weta resighted in the refuges. Few weta or other invertebrates were observed on baits at night. The concentration of 1080 in a cave weta collected alive from a bait, and in a tree weta collected alive from outside an artificial refuge, was less than 10% of the average lethal dose. The results indicate that 1080-poisoning for vertebrate pest control is unlikely to have any negative impact on populations of weta or the other invertebrates monitored.     

History of weta (Orthoptera: Anostostomatidae) translocation in New Zealand: lessons learned, islands as sanctuaries and the future

Establishing new populations by transferring founder individuals from source populations has been effective for managing the recovery of many threatened species including some weta (Orthoptera: Anostostomatidae) in New Zealand. These large-bodied flightless insects are ‘flagship species’ for insect conservation in New Zealand and many are rare or threatened. The declining abundance of most weta species, particularly giant weta, can be attributed to the introduction of mammalian predators, habitat destruction, and habitat modification by introduced mammalian browsers. New populations of some weta have been established in locations, particularly on islands, where these threats have been eliminated or severely reduced in order to reduce the risk of extinction. Some populations were established to provide food for endemic vertebrates, ecosystem restoration and ready access for the general public. We illustrate how methods for both transferring weta and monitoring them have become more sophisticated by using a series of case studies. Other transfers of weta not included in the case studies are also summarised. We conclude by re-iterating the importance of documenting the transfer and post-release monitoring for all insect transfers, both for biogeographical reasons and to provide information to improve future transfers.     

Artificial weta roosts: A technique for ecological study and population monitoring of Tree Weta (

Tree weta (Hemideina) are an important component of New Zealand forest ecosystems and have been identified as possible invertebrate indicator species in restoration programmes. We present designs for artificial weta roosts that have been used to monitor tree weta in Hawke's Bay for five years. A variety of invertebrates use the roosts including two species of Hemideina. Our data suggest that occupation of roosts may take a number of years, each roost monitors a very limited area, and that occupation by invertebrates fluctuates seasonally. We recommend that data from weta roosts be used primarily for temporal rather than spatial comparisons, and that installation of roosts takes place as far ahead of changes in management as feasible.     

Dramatic increases in weta (Orthoptera) following mammal eradication on Maungatautari – evidence from pitfalls and tracking tunnels

Opportunities now exist to establish pest-free areas on the mainland of New?Zealand by eradicating introduced mammals from within predator-proof-fenced areas. This has increased opportunities to investigate how the native insect fauna responds to the eradication of introduced mammals. We examined the response of weta populations to mammal eradication in a before-after-control-impact (BACI) experiment within the southern exclosure on Maungatautari. A novel monitoring technique (footprint tracking tunnels) was used in combination with a conventional technique (lethal pitfall traps) to monitor weta populations. Within 2 years after mammal eradication, there were dramatic increases in weta pitfall captures, weta tracking rates and the incidence of weta footprinting per tracking card (proportion squares tracked out of 100). The mean number of weta per pitfall increased 12-fold (95% credible interval 7?20) after mammal eradication for adult Hemideina thoracica and 52-fold (95% credible interval 30?89) for other weta. Before and immediately after mammal eradication approximately equal proportions of juveniles, subadult and adult weta were caught in the pitfall traps. The age structure of weta caught in pitfall traps changed after mammal eradication, with the percentage of adult weta in the samples increasing markedly from 30% in summer 2004/05 to 66% in summer 2008/09. The sex ratio of weta caught also changed, the percentage of females in pitfall traps increasing from 25% in summer 2004/05 to 55% in summer 2008/09. The two monitoring techniques have different advantages and disadvantages, which are discussed, but key advantages of tracking tunnels are that they are non-lethal and time-efficient. Counting squares tracked per card gave a more sensitive and precise indicator of response to mammal eradication than simply recording presence and absence of tracks, but required much more processing time.     

Effects of an aerial 1080 possum poison operation using carrot baits on invertebrates in artificial refuges at Whirinaki Forest Park, 1999– 2002

The effects of an aerial 1080 possum poison operation using carrot baits on invertebrates in Whirinaki Forest Park are described from an un-replicated study of artificial refuges attached to tree trunks. Auckland tree weta (Hemideina thoracica), cave weta (Pharmacus sp. and Isoplectron sp.), cockroaches, spiders and harvestmen, and leaf-veined slugs (Athoracophorus bitentaculatus) were the most frequent occupants, but snails, millipedes, centipedes, flatworms, lepidopteran larvae, glowworm larvae (Arachnocampa luminosa), peripatus, slaters and beetles were also present occasionally. Invertebrate numbers were monitored every second or third month for a year before the poison operation, and for two years afterwards. Numbers of tree weta, cave weta, cockroaches, spiders and harvestmen, and leaf-veined slugs did not decline substantially in refuges in the treatment area relative to those in the non-treatment area immediately after the poison operation. Our results, and those from two other similar studies, suggest that aerial 1080 poison operations are unlikely to have a detrimental effect on invertebrates that occupy cavities above ground.     

An appraisal of simple tree-mounted shelters for non-lethal monitoring of weta (Orthoptera: Anostostomatidae and Rhaphidophoridae) in New Zealand nature reserves

When monitoring rare insect species, or when surveying faunas within nature reserves, it is desirable not to use indiscriminate lethal sampling techniques. In this investigation we assessed the usefulness of simple tree-mounted wooden shelters to monitor endemic weta (Orthoptera) in nature reserves in Canterbury, New Zealand. Fifty shelters were placed out at six sites and examined at three-monthly intervals for a year. A wide variety of invertebrates were found utilizing the shelters, with Arachnida, Blattodea and Collembola being the most common occupants. After three months over 80% of the shelters exhibited signs of use by invertebrates, increasing to 96% after 12 months. Only seven tree weta (Anostostomatidae) and one (dead) ground weta (Hemiandrus sp.) were observed in the shelters over the full 12 month period. There were 52 observations of cave weta (Rhaphidophoridae) in the shelters, 36 of which occurred at one site, Orton Bradley Park. Occupation of the shelters by cave weta was not affected by soil conditions, light intensity or aspect of the shelter. However, cave weta exhibited a preference for shelters less than 50 cm above the ground and for shelters attached to kanuka and vines. Although weta were found in only a small proportion (9%) of the shelters, this method proved useful in confirming the presence of weta without risk of harming vulnerable populations. These shelters are inexpensive and easy to manufacture and have potential for long-term non-lethal monitoring of weta and as a collection/carriage device for live specimens used in conservation translocations.     

Tolerance for Nutrient Imbalance in an Intermittently Feeding Herbivorous Cricket,the Wellington Tree Weta

Organisms that regulate nutrient intake have an advantage over those that do not, given that the nutrient composition of any one resource rarely matches optimal nutrient requirements. We used nutritional geometry to model protein and carbohydrate intake and identify an intake target for a sexually dimorphic species, the Wellington tree weta (Hemideina crassidens). Despite pronounced sexual dimorphism in this large generalist herbivorous insect, intake targets did not differ by sex. In a series of laboratory experiments, we then investigated whether tree weta demonstrate compensatory responses for enforced periods of imbalanced nutrient intake. Weta pre-fed high or low carbohydrate: protein diets showed large variation in compensatory nutrient intake over short (<48 h) time periods when provided with a choice. Individuals did not strongly defend nutrient targets, although there was some evidence for weak regulation. Many weta tended to select high and low protein foods in a ratio similar to their previously identified nutrient optimum. These results suggest that weta have a wide tolerance to nutritional imbalance, and that the time scale of weta nutrient balancing could lie outside of the short time span tested here. A wide tolerance to imbalance is consistent with the intermittent feeding displayed in the wild by weta and may be important in understanding weta foraging patterns in New Zealand forests.     

Identification of weta foraging on brodifacoum bait and the risk of secondary poisoning for birds on Quail Island, Canterbury, New Zealand

Brodifacoum is a second-generation anticoagulant used for rodent control in New Zealand. Concerns about the poisoning of non-target species have resulted in restrictions being imposed on the mainland. It is, however, still commonly employed on offshore islands. Previous research investigating the poisoning risks of brodifacoum has generally focused on birds eating brodifacoum bait (primary poisoning) or through depredation of live rodents or carrion containing brodifacoum residues (secondary poisoning). Other research has highlighted the potential for secondary poisoning of birds via the consumption of contaminated invertebrates. An inspection of rodent bait stations undertaken on Quail Island revealed that both cave and ground weta were feeding on brodifacoum bait. A sample of ground weta (Hemiandrus n. sp.) and cave weta (Pleioplectron simplex) was removed from Quail Island and exposed to toxic bait for 60 days. These weta were then assayed for brodifacoum residues and the values used to quantify the secondary poisoning risk for bird species found around Quail Island. We also calculated the risk to birds of secondary poisoning from the tree weta (Hemideina ricta) and the risk of primary poisoning via direct consumption of brodifacoum bait. The LD50 estimates indicated a low risk of secondary poisoning from contaminated ground weta and cave weta. By contrast, the estimates indicated a higher risk from larger-bodied tree weta; however, our calculations were based on a single residue concentration value and should be treated with caution. Of most concern was the primary poisoning risk from the brodifacoum bait. The results indicated that all the 17 bird species assessed are more susceptible to primary poisoning than secondary poisoning and access to brodifacoum bait by non-target bird species needs to be minimised.     

Fission or fusion? Mitochondrial DNA phylogenetics of the chromosome races of Hemideina crassidens (Orthoptera: Anostostomatidae)

The weta Hemideina crassidens has two chromosomal races that differ by two centric fusions or fissions. The mitochondrial DNA of weta from both chromosomal races and a sister species were sequenced for a 750-bp region of the gene coding for cytochrome oxidase I. The average pairwise genetic distance among the 15 (XO)-chromosome race weta was almost four times greater than the average distance among the 19 (XO)-chromosome race weta. The weta from the 19-chromosome race formed a well-supported monophyletic clade in all shortest maximum parsimony trees. Maximum likelihood and neighbor-joining trees suggested that the 15-chromosome karyotype was paraphyletic with respect to the 19-chromosome karyotype, but this was not supported by maximum parsimony analyses. Although phylogenetic analysis could not exclude chromosome fusion as the rearrangement responsible for the karyotype differentiation, the level of sequence variation and pattern of distribution appear to implicate fission as the more likely event.     

What weta want: colour preferences of a frugivorous insect

Plants use colours as signals to attract mutualists and repel antagonists. Fleshy-fruits are often conspicuously coloured to signal different types of information including fruit maturity and spatial location. Previous work on fruit colour selection focus on large diurnal vertebrates, yet fruit colours are perceived differently by frugivores with different types of visual systems. Here, we tested whether a nocturnal, frugivorous, seed-dispersing insect selects fruits based on their pigmentation and whether different lighting conditions affect fruit colour selection. We captured 20 Wellington tree weta (Hemideina crassidens) from a forest reserve on the North Island of New Zealand and brought them into laboratory conditions to test their fruit colour preferences. The fruits of Coprosma acerosa, a native shrub species that naturally produces translucent, blue-streaked fruits, were dyed either red or blue. Fruits were then offered to weta in a binary (y-maze) choice test in two light conditions, either at night during a full moon or under artificial light conditions in the lab. Weta preferred unmanipulated, naturally blue-streaked fruits and artificially-blue coloured fruits over those dyed red. Furthermore, their colour preferences were unaffected by light environment. Our results therefore suggest that weta can discriminate between colours (using colour vision) in both light and dark light environments. Their consistent preferences for colours other than red indicate that weta might be responsible for the unusual colours of fleshy-fruits in New Zealand.     

Body size determines rates of seed dispersal by giant king crickets

New Zealand is home to giant king crickets called weta, which are the only insects known to consume fleshy-fruits and disperse seeds after gut passage. Although they disperse seeds in viable condition after consumption, their importance as seed dispersers is unknown. We conducted a series of field observations and laboratory experiments to investigate intraspecific variation in the capacity of Wellington tree weta (Hemidenina crassidens) to disperse seeds of tree fuchsia (Fuchsia excorticata). We asked three questions. How frequently do weta disperse fuchsia seeds? Do seed passage rates differ between sexes and different-sized weta? Might weta select for particular seed sizes via differential seed mortality after ingestion? A total of 2,272 F. excorticata seedlings germinated from 241 scats (i.e., faecal pellets) that were collected from the field. Experimental results showed that, on average, 15% of seeds ingested by weta successfully germinated, whereas 75% germinated in control trials. Larger weta dispersed greater numbers of seeds in experimental trials, while no differences in dispersal rates were observed between sexes. Regardless of sex and size, weta preferentially dispersed larger seeds. When interpreted collectively, results indicate that (1) weta are frequent seed dispersers of F. excorticata, although many seeds are destroyed during ingestion, (2) larger-bodied weta consistently disperse greater quantities of seeds, which is unusual in seed dispersal mutualisms, and (3) weta preferentially disperse larger seeds, suggesting that they might interact evolutionarily with New Zealand plants.