Wellington Tree Weta Research Articles

Alternative mating strategies in the Wellington tree weta expressed via genetic polymorphism for precocial male maturation.

The expression of alternative mating strategies is generally hypothesized to be a plastic response of the genome to the environment or a genetic polymorphism. The Wellington tree weta (Hemideina crassidens) is an orthopteran insect in which males express three alternative mating morphotypes related to the size of mandibular weaponry. My common garden experiment in the laboratory shows that male weapon size at maturity in this species is nether condition- nor environment-dependent; therefore, supporting the hypothesis that it is genetically polymorphic. Males that matured at the 8th instar had the shortest development time and the smallest weaponry, males that matured at the 10th instar had the longest development time and the largest weaponry, and males that matured at the 9th instar were intermediate to the other two morphs. These morph-related differences in development time suggest that male alternative mating strategies in this species are expressed via precocial male maturation, a rare mode of strategy expression in insects. Additionally, adult lifespans differed significantly among morphs with 8th instar males having the shortest, and 9th instar males having the longest lifespans. I discuss how these differences in lifespan align with morph-specific investment in pre- and post-copulatory processes to equalize fitnesses among morphs.

Sub-lethal exposure to a mammalian pesticide bait alters behaviour in an orthopteran

Pesticides are a crucial tool to control introduced pest species, facilitating restoration of native ecosystems and reducing mortality risk in endemic species. However, non-target species, including many invertebrates, consume pesticides. Although lethal effects on these non-target species are often documented, sub-lethal effects are poorly understood. New Zealand presents a unique case study in this field, with a range of introduced mammalian pest species that threaten a large proportion of endemic fauna. Current research suggests that vertebrate pesticide baits do not cause mortality in invertebrate consumers, but few studies investigate sub-lethal behavioural effects. In this study, we tested whether consumption of a widely dispersed anticoagulant rodenticide, brodifacoum, influenced the expression and repeatability of behaviour in a known consumer of the bait, Wellington tree weta (Hemideina crassidens) over a 40-day treatment period. We compared refuge emergence, activity, refuge-seeking tendencies and defensive responses of brodifacoum-exposed and untreated control weta. Bait-exposed individuals showed increased emergence but decreased activity, boldness and aggression compared to untreated controls. These results show that brodifacoum, although not directly causing mortality, may alter behaviours directly related to foraging, mating, competitive interactions and anti-predator responses, which could be detrimental to fitness and survival. More research should be undertaken to investigate the capacity for H. crassidens to tolerate exposure, and to determine any age- or duration-related responses. These results should also be a catalyst for further research on non-target behavioural effects in other pesticide-exposed insect species.

Do alternative reproductive strategies in the Wellington tree weta represent different behavioural types?

AbstractIn many animal species, variation in reproductive success among individuals has led to the evolution of alternative mating strategies, which in the case of insects can often be correlated with developmental trajectories. In the Wellington tree weta, Hemideina crassidens, males can mature at the 8th, 9th or 10th instar, while females mature at the 10th instar only. A number of morphological attributes including male head and mandible size correlate with final instar number, and as these attributes represent a form of weaponry, they are often used in mate/site guarding and male–male competition. Tenth instar males have larger head/mandible/body sizes and show a conventional (guarder) reproductive strategy, whereas smaller 8th instar males typically show an unconventional (sneaker) strategy. In contrast, 9th instar males are predicted to adopt a “jack‐of‐all‐trades” strategy whereby they can fight or sneak depending context. Here, we tested whether alternative reproductive morphs exhibit strategy‐specific differences in risk‐taking associated with refuge emergence, activity and antipredator behaviour and further, whether these traits correlate to form a behavioural syndrome. We found that tree weta show consistent and repeatable differences in activity and refuge use at the individual level; however, behavioural covariances suggest that only 8th instar males exhibit a behavioural syndrome. That 9th instar males show high plasticity and variance in their gallery‐related behaviours supports the hypothesis that these males are a “jack‐of‐all‐trades.” Contrary to our predictions, antipredator behaviour was not correlated with other traits, and differences in behaviour overall were consistently more pronounced between individuals rather than between male morphs or sexes.

Refuge size variation and potential for sperm competition in Wellington tree weta.

Ecological variation in resources can influence the distribution and encounter rates of potential mates and competitors and, consequently, the opportunity for sexual selection. Factors that influence the likelihood that females mate multiply could also affect the potential for sperm competition. In Wellington tree weta (Hemideina crassidens, plural “weta”), the size of tree cavities (called galleries) used as refuges affects weta distribution and thus the opportunity for sexual selection and selection on male weaponry size. We examined the predicted effects of gallery size and male weaponry size on the potential for sperm competition. We asked if gallery size influenced the potential for multiple mating by females and potential for sperm competition, if male weaponry size was associated with relative expected sperm competition intensity (SCI), and if estimated male mating success was correlated with potential SCI. To quantify relative competitive environments of males, we created and analyzed networks of potential competitors based on which males could have mated with the same females. We found that small galleries had higher potential for female multiple mating and higher potential for sperm competition. Size of male weaponry was not associated with expected relative SCI. Regardless of gallery size, males with more potential mates were expected to face lower expected relative sperm competition. Thus, in this system, variation in the size of available refuges is likely to influence the potential for sperm competition, in a way that we might expect to increase variation in overall reproductive success.

Sexually dimorphic effect of mating on the melanotic encapsulation response in the harem‐defending Wellington tree weta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae)

AbstractReproductive activities are generally costly to immune responsiveness because limited resources required by reproduction are diverted away from immunity (and vice versa). Reproduction, however, is not expected to affect the immune response in males and females similarly as mating is expected to negatively affect male immunity more so than female immunity. Here, I test the phenotypic plasticity hypothesis in the Wellington tree weta (Hemideina crassidens), a sexually dimorphic orthopteran insect that is endemic to New Zealand. My laboratory experiment showed that although males had higher rates of melanotic encapsulation than females, contrary to prediction, females were the only sex significantly affected by mating and the effect was positive. In addition to immunity differing between the sexes, immune function can differ intrasexually, particularly when males are polymorphic and different investment strategies are used to maximize fitness. Male H. crassidens exhibit alternative mating strategies that are represented by three different morphotypes. I therefore explored whether the morphs differed in their melanotic encapsulation response and whether mating affected the morphs differently. I found no difference among morphs or an effect of mating on male immune response.

Melanin waterproofs mountain-dwelling weta

It can be hard to get your head around just how big some weta are if you're used to working on the Drosophila scale, but giant weta routinely tip the scales at up to 70 g and even the more diminutive tree weta tower over other more familiar insects. While some offshoots of the family make their homes in low-lying areas, others have pushed the boundaries into the mountains, where they are exposed to high winds, low oxygen, the searing sun and dry air. ‘Together, these environmental challenges…mean montane insects are at a significantly higher risk of desiccation’, say Keith King from the University of Otago, New Zealand, and Brent Sinclair from the University of Western Ontario, Canada. So how do these high-altitude insects cope with these conditions? Suspecting that they might reduce their water-loss rates, King and Sinclair set about measuring water-loss across the insect's cuticle in weta ranging from sea-level species (the Wellington tree weta, the tokoriro and the Canterbury tree weta) to the mountain dwellers (the alpine tree weta and the Banks Peninsula tree weta).The duo was impressed to see that the low-level dwellers’ cuticular water loss rate was 0.444 μmol–1 g–1 h–1 while the mountain species’ cuticular water loss rate was 45% lower at 0.245 μmol–1 g–1 h–1. They also found that the mountain dwellers had cut their respiratory water loss by 55%. And when they compared how much water the species lost relative to their metabolic rate, the mountain species were much more conservative than their low-altitude cousins, losing 64% less water.King and Sinclair then explain that the amount of melanin pigment in the cuticle of Drosophila is known to affect insect desiccation rates, with lighter flies losing water across the cuticle faster than darker species. Knowing that there are two forms of the alpine tree weta – one with a melanin-tinted cuticle and a lighter version – the duo decided to test whether the degree of melanisation affected the insects’ cuticular water loss rates. Through a direct comparison of cuticular water loss rates King and Sinclair could see that the darker form lost 46% less water than the lighter form. And when they stripped the waterproofing hydrocarbons from the surface of the cuticles of both insects and measured their cuticular water loss rates, the duo found that although the insects’ water loss rates rocketed, the darker weta still lost water 57% more slowly than the lighter insects. The high melanin content of the cuticle in the darker weta offers more protection from desiccation than the cuticle of the lighter weta.So, mountain-dwelling weta have developed various strategies to protect themselves from desiccation in their harsh mountain homes and some have probably benefited from the waterproofing effects of melanin as an unintended consequence when they became darker for camouflage.

Elevational variation in adult body size and growth rate but not in metabolic rate in the tree weta Hemideina crassidens

Populations of the same species inhabiting distinct localities experience different ecological and climatic pressures that might result in differentiation in traits, particularly those related to temperature. We compared metabolic rate (and its thermal sensitivity), growth rate, and body size among nine high- and low-elevation populations of the Wellington tree weta, Hemideina crassidens, distributed from 9 to 1171ma.s.l across New Zealand. Our results did not indicate elevational compensation in metabolic rates (metabolic cold adaptation). Cold acclimation decreased metabolic rate compared to warm-acclimated individuals from both high- and low-elevation populations. However, we did find countergradient variation in growth rates, with individuals from high-elevation populations growing faster and to a larger final size than individuals from low-elevation populations. Females grew faster to a larger size than males, although as adults their metabolic rates did not differ significantly. The combined physiological and morphological data suggest that high-elevation individuals grow quickly and achieve larger size while maintaining metabolic rates at levels not significantly different from low-elevation individuals. Thus, morphological differentiation among tree weta populations, in concert with genetic variation, might provide the material required for adaptation to changing conditions.

Effect of an immune challenge on the functional performance of male weaponry

Theories of parasite-mediated sexual selection predict a positive association between immune function and the expression of sexually selected ornaments. Few studies, however, have investigated how an immune challenge affects the performance of sexually selected weaponry. Male Wellington tree weta (Hemideina crassidens) (Orthoptera: Anostostomatidae) possess enlarged mandibles that are used as weapons in fights for access to females residing in tree galleries. Intense sexual competition appears to have favoured the evolution of alternative male mating strategies in this species as males have a trimorphic phenotype in which weapon size varies across morphotype: 8th instar males have the smallest jaws, 10th instar males have the largest and 9th instar males being intermediate to the other two. After injecting males and females with either lipopolysaccharide (LPS; immune challenge) or saline (control) I measured over a 24h period each weta's body mass to assess whether they responded immunologically to the LPS and their bite force to assess the functional performance of their jaws. Both sexes responded immunologically to the immune-challenge as LPS-injected individuals lost significantly more body mass than saline-injected controls with females losing more mass than males. Female bite force was significantly reduced 8h after LPS-injection whereas male bite force did not significantly decline. Both sexes regained pre-injection functional performance of their jaws 24h after the immune challenge. My results suggest that females trade-off bite force for immune function whereas males do not.This article is part of a Special Issue entitled: insert SI title.

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.

Reproductive and physiological costs of repeated immune challenges in female Wellington tree weta (Orthoptera: Anostostomatidae)

Fitness-related traits, such as immunity and reproduction, are typically condition dependent, and are predicted to trade off against each other because they share a pool of energy and resources. It is generally assumed that the resources and energy required for immune processes and reproduction is contained in the body fat; however, few studies quantify the effect of an immune insult on fat load. I tested the hypothesis that if mounting an immune response is physiologically costly in terms of consuming limited resources from body fat, and reproduction also relies on these resources, then an immune challenge should cause a reduction in fat load and an associated decline in reproduction. I tested this hypothesis by assaying the fat content of female Wellington tree weta (Hemideina crassidens) after repeated challenges with lipopolysaccharide (LPS), and counting the number of oviposited eggs in addition to assaying the egg protein content, a possible measure of egg quality. In line with my prediction, immune-challenged females had significantly smaller fat loads and laid significantly fewer eggs than saline-injected controls. Moreover, the protein content of the eggs laid by immune-challenged females was significantly lower than that of controls. My experimental results show that an immune challenge with a non-pathogenic stimulant increases fat consumption, which in turn limits the resources available for reproduction. These findings support the hypothesis that reproduction and immunity are condition dependent, and are traded off against each other. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 38–46.

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.

Sexual Selection, Ontogenetic Acceleration, and Hypermorphosis Generates Male Trimorphism in Wellington Tree Weta

Strong sex-specific selection on traits common to both sexes typically results in sexual dimorphism. Here we find that Wellington tree weta (Hemideina crassidens) are sexually dimorphic in both head shape and size due to differential selection pressures on the sexes: males use their heads in male-male combat and feeding whereas females use theirs for feeding only. Remarkably, the sexes share a common ontogenetic trajectory with respect to head growth. Male head shape allometry is an extension of the female’s trajectory despite maturing two instars earlier, a feat achieved through ontogenetic acceleration and hypermorphosis. Strong sexual selection also favours the evolution of alternative reproductive strategies in which some males produce morphologically different weapons. Wild-caught male H. crassidens are trimorphic with regard to weapon size, a rare phenomenon in nature, and weapon shape is related to each morph’s putative mating strategy.

Body size determines rates of seed dispersal by giant king crickets

AbstractNew 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.

Sexually dimorphic immune response in the harem polygynous Wellington tree weta Hemideina crassidens

Abstract Adult males are often less immunocompetent than females. One explanation for this is that intense sexual selection causes males to trade‐off investment in immunity with traits that increase mating success. This hypothesis is tested in the Wellington tree weta (Hemideina crassidens), a large, sexually dimorphic orthopteran insect in which males possess enormous mandibular weaponry used during fights for access to female mates. Field‐collected males have a significantly greater immune response (greater melanotic encapsulation) than females, suggesting that body condition, longevity or an allied trait is important to male fitness, or that females require materials for egg production that would otherwise be used to boost immunity. Although immunity is expected to trade‐off against reproductive traits in both sexes, there is no significant relationship between immune response and weapon or testes size in males, nor fecundity in females.

Why Do Male Tree Weta Aggressively Evict Females from Galleries After Mating?

AbstractThe sexually dimorphic Wellington tree weta (Hemideina crassidens Blanchard) (Orthoptera: Anostostomatidae) is a harem‐defending polygynous insect in which males use their enormous mandibles to fight conspecifics for access to harems residing in tree cavities (gallery). Tree weta ejaculates (spermatophores) do not include a nuptial meal (spermatophylax) to delay removal by females as in some ensiferan Orthoptera. Consequently, male tree weta are predicted to remain with their mates post‐copula to ensure complete sperm transfer. Contrary to prediction, however, males forcibly eject females from the mating site (gallery) soon after copulation. Results suggest that forcible ejection occurs apparently because females consume the previous spermatophore if either the resident male or a rival initiates a subsequent copulation attempt. Thus, ejection appears to reduce the resident male’s risk that his ejaculate will be eaten and his mate will copulate with another male.

Sperm investment in relation to weapon size in a male trimorphic insect?

In the harem polygynous Wellington tree weta (Hemideina crassidens), early maturation at the eighth instar by males is associated with smaller mandibular weaponry. Because these males, compared with larger males (i.e., matured at 10th instar), are less successful at holding harems, they appear to have decreased mating success. Therefore, smaller males may acquire mates using tactics different than those of males with larger weaponry. A source of empirical support for this hypothesis is to show that smaller males offset their mating disadvantage by investing more in spermatogenesis to achieve fertilization success. Contrary to expectation, relative investment in testes (controlling for body size) is similar for smaller and larger males. I suggest that this lack of difference in testes size may be due to both morphs experiencing similar risks of sperm competition on Maud Island. Despite having testes similar in size to larger males, smaller males produce significantly larger ejaculates than males with bigger weaponry. This difference in ejaculate size may stem from larger males investing submaximally in current ejaculates in anticipation of their higher future mating success. Key words: accessory gland size, allometry, ejaculate size, sexual selection, sperm competition, sperm number, strategic ejaculation, testis size. [Behav Ecol]

Identifying a causal agent of sexual selection on weaponry in an insect

In many animal species, males do not seek females directly but instead locate and defend sites that contain spatially or temporally limited resources essential to female survival and reproduction. Resident males that successfully repel conspecific rivals can mate with females attracted to these resources. In theory, increasing resource value increases harem size and thus increases the opportunity (Imates) for and strength of sexual selection on traits crucial to male resource-holding potential and mating success. I experimentally tested this hypothesis in the field using the Wellington tree weta, Hemideina crassidens (Orthoptera: Tettigonioidea: Anostostomatidae), a sexually dimorphic insect in which males use their enlarged mandibles as weapons in male‐male contests over access to females sheltering in tree cavities (galleries). By manipulating gallery size, I showed that, compared with smaller galleries, larger galleries housed larger harems. Variation in gallery size was an important determinant of Imates, but contrary to expectation, greater opportunity existed in small galleries compared with large galleries. As predicted, male weapon size was under stronger directional selection in large galleries because the fitness benefits were greater under these conditions compared with small galleries. My results help explain the positive association between average weapon size and average gallery size observed within and among tree weta populations in New Zealand. Key words: conservation, Hemideina, longitudinal study, resource defense, selection gradient, sexual dimorphism. [Behav Ecol]

Fighting for harems: assessment strategies during male–male contests in the sexually dimorphic Wellington tree weta

Females often aggregate at particular sites for feeding or shelter, thus giving adult males the opportunity to defend harems and increase male reproductive success. Rival males compete for control of harems via ritualized displays or direct combat using weaponry. Contests for harems or the resources required by females can be settled based on asymmetries in fighting ability or resource ownership. Males that accurately assess a rival's fighting ability prior to engaging in potentially costly combat should be favoured by selection. Game theory and optimality models provide three models to explain how individuals decide to persist in or flee from a fight. These models are the energetic war of attrition, the sequential assessment model and the cumulative assessment model. Using staged contests in the laboratory, I tested predictions of these models using the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect native to New Zealand. Male H. crassidens use their enlarged mandibles as weapons in fights for access to adult females that reside in cavities in trees. My results supported a prediction common to each assessment model: contest duration was negatively correlated with the asymmetry in opponent's weapon size. The sequential assessment model of contest settlement was partially supported but the strongest support was for the cumulative assessment model. Predictions of the latter model were supported because: (1) fights are apparently settled based on own-size assessment; (2) fights occur in a single phase and escalate; and (3) contests involve physical combat and injury. I suggest that, in nocturnal species, cumulative assessment will generally be most applicable.

The Relationship Between Resource Control, Association with Females and Male Weapon Size in a Male Dominance Insect

AbstractIn species with a resource‐defence (male dominance) mating system, males are expected to maximize fitness by controlling resources deemed more valuable by sexually receptive females because these sites attract more mates. Furthermore, males, which control more valuable resources should themselves be of high quality. I experimentally tested these predictions in the laboratory using the sexually dimorphic Wellington tree weta, Hemideina crassidens (Blanchard) (Orthoptera: Tettigonioidea: Anostostomatidae). Male H. crassidens use their mandibular weaponry to fight for control of harems (groups of adult females) that seek shelter in trees cavities (galleries). As predicted, larger galleries housed significantly larger groups of females and males with larger weaponry controlled large galleries significantly more often. Therefore, galleries with a larger volume are likely considered more valuable by males because they house larger harems. However, contrary to prediction, males with larger weaponry did not reside with significantly more females overall because females did not always form the largest possible groups in galleries and males with smaller weaponry were able to reside with single females in small galleries. The latter observation suggests a possible alternative mating strategy by disadvantaged males.

Resource quality or harem size: what influences male tenure at refuge sites in tree weta (Orthoptera: Anostostomatidae)?

Rather than seeking females directly, males in many animal species locate and defend sites that contain spatially limited resources essential for female survival and reproduction. In these cases, resident males successfully repelling conspecific rivals will mate with sexually receptive females that seek to use the resident’s resources. Theory predicts that if resources are limiting in nature, are costly to procure, and if residency at the resource site increases male reproductive success, each site should be monopolised by a single adult male. Moreover, if females aggregate at these sites, males should be sedentary and monopolise harems. This predicts that males should reside at resource sites longer than females and male tenure should be positively correlated with harem size. I address these hypotheses using a wild population of the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect in which males use mandibular weapons in fights to control resource sites (galleries in trees) required by females for diurnal refuge. In a longitudinal study using artificial galleries, I show that male tenure in a gallery is positively related to harem size and, contrary to prediction, not gallery size per se. Moreover, contrary to prediction, females are more sedentary than males: males appear to move frequently between galleries to search for novel mates. I show experimentally that galleries represent mating sites to males: males prefer galleries housing adult females whereas females prefer unoccupied galleries. Females likely avoid male-occupied galleries because they incur injuries when interacting with males.