Are footprint tracking tunnels suitable for monitoring giant weta (Orthoptera: Anostostomatidae)? Abundance,distribution and movement in relation to tracking rates

Appropriate monitoring tools are essential for assessing the effectiveness of management for all threatened insect taxa. In New Zealand the large-bodied flightless orthopterans in the genus Deinacrida have mostly been monitored by searching through habitat during the day or spotlighting at night but this is time consuming and the results depend on the skill of the searcher. Recently, footprint tracking tunnels, similar to those used for monitoring small mammals in New Zealand, were found to be effective for detecting adults of various giant weta species. In this study, we compared the abundance of Cook Strait giant weta (CSGW) in the vicinity of the tunnels, estimated by mark-recapture, with the number of tracking tunnels tracked by weta. We found strong indications that both baited and unbaited tracking tunnels can be used to estimate the number of adult weta present but that this probably depends on their responses to meteorological conditions which are not yet understood. Our results also show that footprint tracking tunnels are more effective for detecting adult CSGW than searching for these insects at night and that baiting tracking tunnels with peanut butter increases their effectiveness for detecting adult CSGW. We confirmed how far Cook Strait giant weta moved each night on Matiu-Somes Island by attaching transmitters to them and found that day roosts of three adult males were on average 8.6 m apart each day and those of adult females were on average 21.3 m apart. Both the low recapture rates of marked adult CSGW and the nightly displacements of those with transmitters suggest that adult CSGW show no site fidelity and are clearly capable of moving large distances each night. However, an individual weta is unlikely to track more than one tunnel per night if tunnels are 30 m apart. Tracking tunnels have the potential to be used with some other insects, provided their footprints are diagnostic. An advantage of using tracking tunnels is that they are non-lethal and would therefore be particularly suitable for monitoring other large threatened insect taxa.     

Where have all the weta gone? Results after two decades of transferring a threatened New Zealand giant weta, Deinacrida mahoenui

There have been numerous transfers of the large-bodied orthopteran, the Mahoenui giant weta (Family Anostostomatidae: Deinacrida mahoenui), over the past 19 years but there has been limited follow-up monitoring to ascertain establishment and breeding of transferred populations. Recent surveys carried out at all the locations where this weta were transferred, found weta at four of the seven transferred sites. The most important factor determining the success of past transfers is the absence of introduced mammalian predators, particularly rats, at a site. At two sites, Mahurangi Island Scenic Reserve and Warrenheip, weta appeared to be flourishing and have successfully established new populations in the absence of rats. If mammals are present at a site, the occurrence of dense prickly gorse to protect giant weta from predation is another important factor in their survival. Weta were found at very low densities (only single specimens were captured) at Mangaokewa Scenic Reserve and Tikikaru (private land) and it is likely their populations are not viable in the long term. Further efforts to establish Mahoenui giant weta populations should be in mammal-free sanctuaries containing native forest. More intensive post-transfer monitoring using radio-transmitters would enable better understanding of their behaviour after transfer. In addition, long-term regular monitoring of transferred populations is required, particularly those where the likelihood of rat re-invasion is high.     

Does predation result in adult sex ratio skew in a sexually dimorphic insect genus?

Theory proposes that sexually dimorphic, polygynous species are at particularly high risk of sex-biased predation, because conspicuous males are more often preyed upon compared to females. We tested the effects of predation on population sex ratio in a highly sexually dimorphic insect genus (Hemideina). In addition, introduction of a suite of novel mammalian predators to New Zealand during the last 800 years is likely to have modified selection pressures on native tree weta. We predicted that the balance between natural and sexual selection would be disrupted by the new predator species. We expected to see a sex ratio skew resulting from higher mortality in males with expensive secondary sexual weaponry; combat occurs outside refuge cavities between male tree weta. We took a meta-analytic approach using generalized linear mixed models to compare sex ratio variation in 58 populations for six of the seven species in Hemideina. We investigated adult sex ratio across these populations to determine how much variation in sex ratio can be attributed to sex-biased predation in populations with either low or high number of invasive mammalian predators. Surprisingly, we did not detect any significant deviation from 1 : 1 parity for adult sex ratio and found little difference between populations or species. We conclude that there is little evidence of sex-biased predation by either native or mammalian predators and observed sex ratio skew in individual populations of tree weta is probably an artefact of sampling error. We argue that sex-biased predation may be less prevalent in sexually dimorphic species than previously suspected and emphasize the usefulness of a meta-analytic approach to robustly analyse disparate and heterogeneous data.     

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.     

New Zealand endemic open-habitat specialist,the Black-fronted Tern (Chlidonias albostriatus), experienced population expansion during Pleistocene glaciation and recent decline

Understanding how climatic and environmental changes, as well as human activities, induce changes in the distribution and population size of avian species refines our ability to predict future impacts on threatened species. Using multilocus genetic data, we show that the population of a threatened New Zealand endemic open-habitat specialist, the Black-fronted Tern Chlidonias albostriatus – in contrast to forest specialists – expanded during the last glacial period. The population has decreased subsequently despite the availability of extensive open habitat after human arrival to New Zealand. We conclude that population changes for open habitat specialists such as Black-fronted Terns in pre-human New Zealand were habitat-dependent, similar to Northern Hemisphere cold-adapted species, whereas post-human settlement populations were constrained by predators independent of habitat availability, similar to other island endemic species.     

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.     

Diversification of New Zealand weta (Orthoptera: Ensifera: Anostostomatidae) and their relationships in Australasia

New Zealand taxa from the Orthopteran family Anostostomatidae have been shown to consist of three broad groups, Hemiandrus (ground weta), Anisoura/Motuweta (tusked weta) and Hemideina-Deinacrida (tree-giant weta). The family is also present in Australia and New Caledonia, the nearest large land masses to New Zealand. All genera are endemic to their respective countries except Hemiandrus that occurs in New Zealand and Australia. We used nuclear and mitochondrial DNA sequence data to study within genera and among species-level genetic diversity within New Zealand and to examine phylogenetic relationships of taxa in Australasia. We found the Anostostomatidae to be monophyletic within Ensifera, and justifiably distinguished from the Stenopelmatidae among which they were formerly placed. However, the New Zealand Anostostomatidae are not monophyletic with respect to Australian and New Caledonian species in our analyses. Two of the New Zealand groups have closer allies in Australia and one in New Caledonia. We carried out maximum-likelihood and Bayesian analyses to reveal several well supported subgroupings. Our analysis included the most extensive sampling to date of Hemiandrus species and indicate that Australian and New Zealand Hemiandrus are not monophyletic. We used molecular dating approaches to test the plausibility of alternative biogeographic hypotheses for the origin of the New Zealand anostostomatid fauna and found support for divergence of the main clades at, or shortly after, Gondwanan break-up, and dispersal across the Tasman much more recently.     

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.     

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.     

Tracking tunnels: a novel method for detecting a threatened New Zealand giant weta (Orthoptera: Anostostomatidae)

Several species of giant weta, including wetapunga (Deinacrida heteracantha, Orthoptera: Anostostomatidae), New Zealand?s largest terrestrial invertebrate, have high conservation value, yet no methods for detecting and monitoring them have been developed. Here we show that rodent tracking tunnels set over three consecutive nights can be used to detect wetapunga on Little Barrier Island. Adult wetapunga footprints had significantly longer tarsal pad lengths than subadult wetapunga and Auckland tree weta, but tarsal pad length does not distinguish between subadult wetapunga and adult Auckland tree weta. Results suggest that setting tunnels on the ground is more effective than setting them on tree branches and that peanut butter as an attractant bait increases the detection rate of adult wetapunga. Future research is required to determine if the footprints of ground weta, tree weta and giant weta of different ages can be distinguished and to determine how tracking rates of giant weta relate to the population density.     

Life histories, dispersal, invasions, and global change: progress and prospects in New Zealand ecology, 1989?2029

We highlight three areas of significant progress in ecology since 1989 which are particularly relevant to New Zealand, and three major challenges for the next two decades. Progress: (1) The unusual life histories of New Zealand organisms, including extreme longevity and low reproductive rates, are now seen as efficient responses to the low-disturbance environment present before the arrival of large mammals, including humans. (2) Recent data show that long distance dispersal has been far more common than previously supposed, changing our image of New Zealand from a Gondwanan ark to the ?flypaper of the Pacific?. (3) Greatly improved techniques for pest control, and innovative species management, have stabilised numbers of many of the most charismatic of New Zealand?s threatened species. Problems: (1) Native species continue to decline, including many previously thought to be stable, and improved phylogenetics and new discoveries have added threatened species. (2) Despite increased emphasis on biosecurity, biological invasions are continuing, driven by increased trade and lags in naturalisation. (3) Conservation efforts risk being overwhelmed by the direct effects of increasing human population, resource use, invasions, and global climate change at a time when human food supplies and economies are coming under increasing pressure from environmental constraints. Conclusions: (1) We need improved ecological understanding and more management tools for invasive and threatened species, especially for species other than birds. (2) In these decades of rapid climate change and habitat conversion, there is an urgent need for more widespread and sustainable integration of native species into New Zealand?s rural and urban lowland landscapes.     

Managing genetic diversity in threatened populations: a New Zealand perspective

Genetic diversity allows a population to adapt genetically to a changing environment or to buffer it against stochastic events such as harsh weather or disease outbreaks. Genetic diversity is therefore an important consideration in the development of management strategies for threatened populations around the world, with the possible exception of New Zealand, where species recovery programmes tend to focus on increasing population size while neglecting the maintenance of genetic diversity. Many of New Zealand?s threatened species have relatively low genetic variation and consequently may still be at risk in the long-term due to reduced resilience even if the effects of introduced predators were eliminated. The three main factors affecting genetic diversity – genetic drift, inbreeding and population subdivision – are processes that potentially impact on many of our locally threatened species, but their effects tend to occur over a considerably broader timescale than ecological effects, and as such are much more difficult to detect and ultimately to justify additional resource spending towards. Our message is that genetic management of New Zealand threatened species should not take priority over other management concerns such as controlling predators or improving habitat quality, but it needs more attention than it currently receives. We recommend that genetic diversity be a fundamental component in long-term management strategies for threatened species, and that such strategies are made explicit within the New Zealand Department of Conservation?s current species recovery plans so that the persistence of biodiversity becomes of key importance, as opposed to current approaches that seek solely to maximise representation.     

The impact of exotic weed competition on a rare New Zealand outcrop herb,

Nearly one quarter of New Zealand’s unique vascular plant flora is threatened, and weed invasion is implicated in the decline of more than half of these threatened species. However, there is little experimental evidence showing that invasive weeds have a direct impact on threatened native plants. This study experimentally tested the hypothesis that competition with invasive weeds threatens the rare outcrop plant Pachycladon cheesemanii (Brassicaceae). Pachycladon cheesemanii is a threatened South Island, New Zealand endemic with a distribution nearly confined to rock outcrops. It has disappeared from historical record sites throughout its range. The effects of weed competition and habitat on P. cheesemanii establishment, growth and survival were investigated by sowing seed into replicated plots subject to three treatments: weed removal, soil disturbed and unweeded control, in three habitat types: forested and open rock outcrops and open tussock grassland. The experiments were carried out at three locations: Mt Somers (Canterbury), Wye Creek and Diamond Lake (Otago). Within weedy rock outcrop habitat, weed removal significantly increased the rate of P. cheesemanii germination, and appeared to increase seedling growth rates, implying that weeds can negatively impact populations. Relative to rock outcrop habitat, P. cheesemanii germination was very low in adjacent open grassland habitat regardless of weeding treatment. Demographic monitoring of four natural populations of P. cheesemanii revealed that seed production is highly variable among populations and may be limited by browse and mechanical damage to inflorescences. Pachycladon cheesemanii does produce a persistent seed bank but most seed is found close to parent populations. Our results suggest that competition with invading weeds threatens current P. cheesemanii populations, that plant establishment can be enhanced by weed removal, and that considerable potential exists for artificially expanding populations by sowing seed into appropriate weed-free habitat.     

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.     

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.     

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.     

Distribution and forage use of exotic bumblebees in South Island, New Zealand

The rapid decline in bumblebee populations within Europe has been linked to habitat loss through agricultural intensification, and a consequential reduction in the availability of preferred forage plants. The successful introduction of four European Bombus species to the South Island of New Zealand from England (in 1885 and 1906) provides an opportunity to determine how important different forage plants (also introduced from the U.K.) are to two severely threatened European bumblebee species (Bombus ruderatus and B. subterraneus). In January 2003 we conducted a survey of bumblebee populations across 70 sites in the central and southern South Island, recording which plant species were being used as pollen and nectar sources for each Bombus species. All four bumblebee species showed a clear preference for plants of European origin. Only B. terrestris, the most polylectic species, was recorded feeding on native plant species. The longer-tongued bumblebees, B. hortorum, B. ruderatus, and B. subterraneus, foraged predominantly on just two plant species; Trifolium pratense for both nectar and pollen, and Echium vulgare for nectar. These plant species are now declining in abundance in the U.K. Our results provide support for the hypothesis that the loss of flower-rich meadows, particularly those containing populations of Fabaceae species with long corollae, is responsible for the decline of bumblebee species across Europe. Comparison with earlier bumblebee surveys suggests that long-tongued bumblebees may also be in decline in New Zealand, particularly B. subterraneus which is now very localised and scarce.     

Protected species and predator detection dog use in New Zealand wildlife conservation projects

In many New Zealand threatened species (birds, lizards, frogs, invertebrates) recovery programmes dogs are a significant conservation management tool. Protected species detection dogs have regularly been used to locate rare birds since the 1970’s and more recently lizards. They have been used to detect, but not capture, kakapo, takahe, kiwi, whio, pateke, taiko, skinks and geckos, for monitoring and translocation purposes. Kakapo would possibly be extinct today if dogs had not been used to locate birds on Stewart Island and Fiordland for translocation to predator-free islands before predators and old age eliminated these original remnant populations. Use of these dogs on the kiwi recovery projects has advanced the progress of these projects by about ten years. Predator detection dogs support threatened species recovery programmes and their role is to enhance other predator control methods. They have been regularly used since 2002 following a successful three-year pilot project to prove their usefulness. These dogs are used to detect the presence of mammalian predators including rodents, mustelids, cat and hedgehog. They are particularly useful when predator numbers are low when other predator detection methods (tracking tunnels, traps, gnaw sticks) have difficulty confirming their presence. Once detected by the dogs the predators are killed using toxins, traps or shooting. Dogs are used for: surveillance of supposedly predator-free sites; as a tool to focus predator a process of handler application and interview, and a two-step assessment and certification system for dogs and handlers guide these Department of Conservation dog programmes. There are currently 11 predator dog handlers and 26 protected species dog handlers in the programmes.     

After the deluge: mitochondrial DNA indicates Miocene radiation and Pliocene adaptation of tree and giant weta (Orthoptera: Anostostomatidae)

Aim New Zealand broke away from the margins of Gondwana c. 75 Ma. Since then, New Zealand taxa derived from the Gondwanan biota are thought to have been exposed first to a subtropical climate on a low lying terrain, then severe land reduction during the Oligocene marine transgression, followed by much cooler climates of the Pliocene and Pleistocene, at which time mountain ranges emerged. The biological consequence of New Zealand's geological and climatic history is not well understood, in particular the extent to which the Oligocene acted as a biological bottleneck remains unresolved. Methods We used mitochondrial cytochrome oxidase I and 12S DNA sequences to examine the extent of diversity and inferred timing of speciation of New Zealand weta (Anostostomatidae), a group of Orthoptera with a Gondwanan distribution generally thought to be ancient inhabitants of New Zealand. Main conclusions We hypothesize that at least three distinct groups of weta survived the Oligocene marine transgression and radiated subsequently. Speciation followed during the Miocene and radiation into new habitats occurred during the Pliocene when mountain building created novel environments. Patterns of genetic diversity within species reflect, in some instances, geographical subdivision in the Pliocene, and in other cases, Pleistocene range changes resulting from climate change.