Neuromodulatory unpaired median neurons in the New Zealand tree weta, Hemideina femorata

Wetas are ancient Gondwanan orthopterans (Anostostomatidae) with many species endemic to New Zealand. Like all Orthoptera they possess efferent neuromodulatory dorsal unpaired median (DUM) neurons, with bilaterally symmetrical axons, that are important components of motor networks. These neurons produce overshooting action potentials and are easily stimulated by a variety of external mechanosensory stimuli delivered to the body and appendages. In particular, stimulation of the antennae, mouth parts, tarsi and femora of the legs, abdomen, cerci and ovipositor is very effective in activating DUM neurons in the metathoracic ganglion of wetas. In addition, looming visual stimuli or light on-, light off-stimuli excite many metathoracic DUM neurons. These DUM sensory reflex pathways remain viable after the prothoracic to subesophageal connective is cut, whereas in locusts such reflex pathways are interrupted by the ablation. This suggests that, in wetas, sensory reflex pathways for DUM activation are organized in a less centralized fashion than in locusts, and may therefore reflect a plesiomorphic evolutionary state in the weta. In addition, many weta DUM neurons exhibit slow rhythmic bursting which also persists following the connective ablation.     

Avoidance of intracellular freezing by the freezing-tolerant New Zealand Alpine weta Hemideina maori (Orthoptera: Stenopelmatidae)

Calorimetric analysis indicates that 82% of the body water of Hemideina maori is converted into ice at 10 degrees C. This is a high proportion and led us to investigate whether intracellular freezing occurs in H. maori tissue. Malpighian tubules and fat bodies were frozen in haemolymph on a microscope cold stage. No fat body cells, and 2% of Malpighian tubule cells froze during cooling to -8 degrees C. Unfrozen cells appeared shrunken after ice formed in the extracellular medium. There was no difference between the survival of control tissues and those frozen to -8 degrees C. At temperatures below -15 degrees C (lethal temperatures for weta), there was a decline in survival, which was strongly correlated with temperature, but no change in the appearance of tissue. It is concluded that intracellular freezing is avoided by Hemideina maori through osmotic dehydration and freeze concentration effects, but the reasons for low temperature mortality remain unclear. The freezing process in H. maori appears to rely on extracellular ice nucleation, possibly with the aid of an ice nucleating protein, to osmotically dehydrate the cells and avoid intracellular freezing. The lower lethal temperature of H. maori (-10 degrees C) is high compared to organisms that survive intracellular freezing. This suggests that the category of 'freezing tolerance' is an oversimplification, and that it may encompass at least two strategies: intracellular freezing tolerance and avoidance.     

Control systems models for the circadian clock of the New Zealand weta, Hemideina thoracica (Orthoptera: Stenopelmatidae)

The New Zealand weta, Hemideina thoracica, is a nocturnal orthopteran insect which emerges from holes in trees or from under bark soon after sunset to forage for several hours on plant and animal material before returning to its refuge before dawn. In tests of the internal clock hypothesis it exhibits clear circadian locomotor rhythms in which the period is initially somewhat less than 24 h, but frequently spontaneously increases to over 25 h. The rhythms are entrainable by light and temperature cycles, obey Aschoff's Law and are temperature compensated. A single oscillator feedback model accounts for these basic properties of the weta clock, but does not explain a variety of examples of rhythm lability, such as day skipping, spontaneous change in period, scalloping and desynchrony typically found in the real data. To account for these characteristics the model is expanded into two linked populations of oscillators, which retain the basic properties of the simple model and in addition interact through their coupling to show the various types of free-run lability. To make these control systems models compatible with the molecular interpretation of circadian biology, each of the components in the feedback loop is matched with molecular function and structure.     

Intraspecific karyotype variation is not concordant with allozyme variation in the Auckland tree weta of New Zealand, Hemideina thoracica (Orthoptera: Stenopelmatidae)

Nine karyotypes are described within a single species of common New Zealand tree weta. Their diploid numbers range from 11 to 25. The distribution of the karyotypes suggests that each had a single origin except the 17-karyotype which was the most common karyotype and had a disjunct distribution. The overall level of allozyme diversity observed is similar to that seen within many widespread taxa. The distribution of allozyme alleles did not coincide with the distribution of karyotypes within this species and the Neighbour-Joining tree was not concordant with the chromosome based sub-divisions of the species. Thus, no evidence was found to suggest that chromosomal differentiation has been acting as a barrier to the flow of alleles within H. thoracica. The lack of concordance of genetic markers is thought to result from rapid chromosome radiation and reticulate evolution. Northland peninsula of North Island, New Zealand is a region of high chromosomal and allozymic diversity in H. thoracica. This may have resulted from geographic isolation during the Pliocene when Northland formed an archipelago of many small low-lying islands.     

Microclimate and variations in haemolymph composition in the freezing-tolerant New Zealand alpine weta Hemideina maori Hutton (Orthoptera: Stenopelmatidae)

The microclimate in the habitat of the New Zealand alpine weta Hemideina maori is very variable with winter temperatures down to −6 °C under the rocks where the insects are found. Subfreezing temperatures may in winter prevail for up to 17 days but diurnal cycles of freezing and thawing are common, as is also the case in summer. Rates of temperature change can be very high and up to −7.20 °C/h. During winter, humidity was high for extended periods ranging from 70% to 100% relative humidity (RH). In the summer, humidity ranged from 30% RH during the day to 100% RH at night. The supercooling point of the haemolymph was approximately −8 °C year round, caused by a heat labile substance. The supercooling point of the haemolymph of an insect of the same genus, Hemideina femorata not regularly exposed to subfreezing temperatures, was ca. −16.5 °C. Thermal hysteresis was not detected in the haemolymph of H. maori. Haemolymph osmolality varied from 380 mOsm (summer) to 700 mOsm (winter). Body water content was ca. 75% all year round. Total concentrations of sodium, potassium and chloride in haemolymph varied from 170 mM (winter) to 250 mM (summer). The total concentration of free amino acids varied from 58 mM (summer) to 263 mM (winter). This variation was mostly due to proline which varied from ca. 15 mM (summer) to ca. 100 mM (winter). The freeze-tolerant weta H. maori is exposed to a highly variable and cold environment all year round and several properties of its haemolymph composition can be attributed to these climatic conditions, e.g. the presence of ice-nucleating agents and an increase in the concentration of proline during cold hardening in the autumn. Accepted: 22 February 1999     

Ice nucleation and freezing tolerance in New Zealand alpine and lowland weta, Hemideina spp. (Orthoptera; Stenopelmatidae)

Abstract.The alpine tree weta Hemidiena maori Pictet et Saussure (Orthoptera: Stenopelmatidae) is a large, flightless insect found above the treeline on many of the mountain ranges of the South Island of New Zealand. The population found on the Rock and Pillar Range, Central Otago has been identified as freezing tolerant with a haemolymph ice nucleating agent. The ability of H. maori to survive freezing is compared to the lowland weta Hemideina thoracica Walker and H. crassidens Blanchard, both of which are able to survive the formation of some ice in their bodies. Mortality is associated with time spent frozen in H. thoracica , and it is hypothesized that this species is killed when a critical proportion of its body water is frozen. All five subalpine and alpine populations of H. maori surveyed were found to be freezing tolerant.
Comparison of temperatures of first nucleation and mean supercooling point of haemolymph droplets suggest that haemolymph ice nucleating activity varies between populations of H. maori. Hemideina maori collected from the Mt Cook region appear to lack a haemolymph ice nucleator. This population is nevertheless freezing tolerant, suggesting that the haemolymph ice nucleating agent described in H. maori is not essential for freezing tolerance. Hemideina crassidens and H. ricta Hutton, both of which are found in lowland habitats, also had high mean supercooling point and temperatures of first nucleation of haemolymph droplets, suggesting that these species also have a haemolymph ice nucleator.
Comparison of ice nucleation characteristics of haemolymph and faecal material (representing gut contents) suggests that gut nucleators in H. maori may be at least as efficient as the haemolymph nucleator. It is concluded that freezing tolerance is probably not an adaptation to the alpine environment. This highlights the need for inter- and intraspecific comparative studies if physiological data are to be used to draw evolutionary conclusions.     

Structure of the cuticle of the alpine weta,Hemideina maori (Saussure) (Orthoptera: Stenopelmatidae)

Sclerotized cuticle segments from the thorax, dorsal abdomen, and ventral abdomen of the alpine, weta Hemideina maori (Saussure) (Orthoptera: Stenopelmatidae) were examined by light microscopy and by scanning and transmission electron microscopy. An epicuticle, exocuticle (outer and inner), mesocuticle, endocuticle, and deposition layer are present in transverse sections. The epicuticle is further composed of a cuticulin layer and inner epicuticle, the latter being finely laminated and containing narrow wax canals that terminate below the cuticle surface. Openings to dermal gland ducts are visible on the surface as are large setae and smaller sensory pegs. Frozen fractured cuticle reveals the presence of horizontal ducts or channels that run laterally within the cuticle. The structure of weta cuticle is compared with that of the common house cricket and arthropods in general.     

Sexual selection and infection by ectoparasites in Wellington tree weta,Hemideina crassidens (Orthoptera: Anostostomatidae)

Abstract Sexual selection can affect the prevalence and intensity of infection of individuals by ectoparasitic mites. According to this theory, males should exhibit greater infection by parasites than females and juveniles should be less infected than adults. In the wild, I investigated whether prevalence and intensity of the chyzeriid mite, Nothotrombicula deinacridae (Dumbleton) differed between the sexes and between developmental stages in Wellington tree weta, Hemideina crassidens. Despite being under strong sexual selection, male tree weta did not exhibit greater parasitism and there was some evidence that adults and juveniles differed in prevalence. The sexual selection hypothesis was not supported in this study.     

Robertsonian translocations and B chromosomes in the Wellington tree weta, Hemideina crassidens (Orthoptera: Anostostomatidae)

Two karyotypes within the species Hemideina crassidens are described, 2n = 15 (XO) and 2n = 19 (XO). These two karyotypes have a NF of 28. The 19-karyotype was found exclusively in the southern part of the species range and the 15-karyotype was found in the north. The differences between the two karyotypes are interpreted as arising from two Robertsonian translocations (fission/fusion). Laboratory matings between weta with the two karyotypes produced viable offspring. During meiosis in F1 intraspecific hybrids metacentric and acrocentric autosomes aligned to form two trivalents, confirming homologies predicted by Robertsonian translocations. The subspecies H. c. crassicruris, (confined to Stephens Island) was found to be polymorphic for a metacentric B chromosome. An unusual association of sex and presence of B chromosome was observed in this island population with Bs found only in male weta.     

Structure of the auditory system of the weta Hemideina crassidens (Blanchard, 1851) (Orthoptera,Ensifera, Gryllacridoidea,Stenopelmatidae)

Summary This study of the ultrastructure of the auditory sensilla of the New Zealand weta, Hemideina crassidens, is the first such study on a member of the orthopteran Superfamily Gryllacridoidea. Ultrastructure of the auditory sensilla is similar in all of the tibial mechanosensory organs, here called subgenual organ, intermediate organ and crista acoustica by analogy with comparable structures in Tettigoniidae.Distal to each sensory soma is a dendrite containing multiple ciliary rootlets that fuse into a single ciliary root. This splits into nine root processes that pass around the outside of the proximal basal body and then rejoin at the level of the distal basal body, distal to which the dendrite has a modified ciliary structure with a circlet of nine peripheral paired tubes and rods as it passes through the proximal extracellular space. It is then enclosed by a zone of scolopale cell cytoplasm before expanding into a dilatation within the distal extracellular space. In some sensilla this space is partially occluded by electron dense material which is part of the scolopale cell. Distal to the dilatation the cilium shrinks and ends surrounded by the scolopale cap.Accessory cells consist of glia enwrapping the sensory neuron in the region of its soma, the scolopale cell surrounding the ciliary portion of the dendrite, and the attachment cell surrounding the scolopale cell and scolopale cap and connected to them by desmosomes. The attachment cells are filled with microtubules in differing densities and orientations. Lamellae are present in the acellular matrix surrounding the attachment cells. Banded fibres, presumably of collagen, are also present in the matrix.     

Identification of a hybrid zone between distinctive colour variants of the alpine weta Hemideina maori (Orthoptera: Stenopelmatidae) on the Rock and Pillar range, southern New Zealand

The weta Hemideina maori occurs as yellow (to the north), black (to the south) and intermediate colour variants on the Rock and Pillar range in New Zealand. Isozyme electrophoresis revealed little genetic variation, whereas RFLP analysis of an amplified mtDNA sequence uncovered two haplotypes correlating completely with colour in allopatry and nearly so in sympatry. Intermediates had one or other haplotype. The observed distribution of colour variation and mtDNA genotypes is characteristic of a hybrid zone, perhaps formed by secondary contact. Work is continuing to locate nuclear DNA markers and to study the genetic interactions of the colour variants.     

Diet of the ground weta Zealandosandrus gracilis (Orthoptera: Stenopelmatidae)

Abstract

The diet of the ground weta Zealandosandrus gracilis Salmon was investigated by examining the crop contents of 68 individuals collected throughout the year from the Cass area, Canterbury. Specimens representing all instars of both sexes, as well as adults, were obtained. Z. gracilis is carnivorous, preying on immature and adult invertebrates (mostly insects) of the forest litter. Similar prey was taken by early instars of both sexes; late instars captured a wider range of prey, including larger and more active species. Late-instar and adult females preyed more on adult insects such as mycetophilid flies than did males, which took more insect larvae.     

Stridulatory behaviour in a New Zealand weta, Hemideina crassidens

Using a femoral-abdominal stridulatory mechanism, wetas produce the following stridulatory behaviours: aggression, mating, calling, defence and disturbance. Syllable period, rather than number of syllables/echeme or syllable duration, was the most stereotyped temporal parameter for aggression, mating and calling stridulation. Coefficients of variation of the above parameters were large and overlapped considerably for aggression and mating stridulation. We concluded that, for these two behaviours, a basic sound pattern is used to convey different messages to female and male receiver wetas, respectively, but the syllable period of the pattern decreases with increased excitation of males in aggressive encounters. In adult male combat, winners stridulated last in a bout, and produced significantly more aggression sounds than losers.     

Localization of the circadian pacemakers of Hemideina thoracica (Orthoptera; Stenopelmatidae)

The location of the circadian pacemakers of the orthopteran Hemideina thoracica (White) has been investigated through observation of the effects of surgical removal of brain tissues (principally optic lobes and tracts) on free-running and entrained locomotor rhythms. Bilobectomy and severance of optic tracts invariably resulted in arrhythmicity, whereas rhythmicity was sustained following unilateral lobectomy, generally with increases in the free-running period (FRP) and decreases in both the active-phase lengths and activity-to-rest ratios of the rhythm. Bilobectomized subjects could be entrained by temperature cycles, but exhibited no transients or residual rhythmicity, indicating that temperature brought about a direct response or masking effect. These results support the hypothesis that the circadian locomotor pacemakers of Hemideina are located within each optic lobe, and that there are no extraoptic centers for the control of the timing of locomotor activity. Although confirmation of the pacemaker role of the optic lobes requires transplantation of the tissues, the conclusion may be drawn by inference from other studies (e.g., Leucophaea maderae--Page, 1983; Gryllus bimaculatus--Tomioka and Chiba, 1986). Light entrainment continued after surgical binding and blackening of the compound eyes and ocelli, supporting the view that direct illumination of neural tissue through the cuticle may be one possible pathway for light entrainment.     

Tree wetas (Hemideina crassicruris) (Orthoptera: Stenopelmatidae) of Stephens Island,Cook Strait

Abstract

Tree wetas (Hemideina crassicruris) were studied in kohekohe (Dysoxylum spectabile) forest on Stephens Island, Cook Strait, between 5 and 17 January 1978. Mark and recapture observations suggested a population density of at least 5300 wetas ha^?1 and a biomass of 42 kg ha^?1. Activity of wetas increased with temperature and humidity. Tuataras are important predators on Stephens Island and probably deter wetas from moving onto the forest floor. The enlarged head and jaws of male wetas are associated with head-on fights for occupancy of galleries. Single, adult males occupy galleries with a harem of up to 9 females. Stridulation was associated with agonistic encounters and was most frequent shortly before dawn. Seventeen percent of males and 42% of females were damaged. Moulting, copulation, and oviposition were observed, and females carried up to 96 mature eggs. Stephens Island tree wetas reach larger sizes overall, and males have relatively larger heads, than in other populations of the Cook Strait region. The number and size of galleries and the presence or absence of predators may be important determinants of weta population density, sex ratio, harem formation, and activity on the ground.     

Structure and evolution of stridulatory mechanisms in New Zealand wetas (Orthoptera : Stenopelmatidae)

Four types of stridulatory mechanisms were found in 35 species (7 genera) of New Zealand wetas (Orthoptera : Stenopelmatidae). These include: (1) tergo-femoral; (2) tergo-tergal; (3) mandbbuao-mandibular; and (4) pleuro-coxal. These fell into 12 groups, based upon shape, density and pattern of tergal and femoral pegs, number of terga bearing pegs and presence of mandibular tusks bearing stridulatory tubercles. The plesiomorphic condition, consisting of spinose peg patches on 3 or more abdominal tergites rubbed by bands of pegs on the femur, is found in Hemiandrus spp. Apomorphically-derived tergal files have arisen separately in 3 other genera. In Hemideina, (2 species groups) well-formed files are very similar among species. In Deinacrida, (4 species groups), the trend is toward file reduction into one or 2 massive tergal ridges, and embellished femoral pegs. In the tusked wetas (2 gen., 2 spp.), one species has a crude file of many broken ridges. Stridulatory structures on mandibular tusks of both New Zealand species are unique, although the tusks appear to have their origin in African stenopelmatids. The plesiomorphic condition appears to have been deployed originally for defense stridulation (inter-specific communication). Additional intea-specific stridulatory communication developed in Hemideina and Deinacrida, using the apomorphic file and peg mechanisms. Here, the stridulation is associated with defense, calling, mating and disturbance behaviors.     

Electrochemical characteristics of ion secretion in malpighian tubules of the New Zealand alpine weta (Hemideina maori)

Characteristics of ion and fluid secretion were investigated in isolated Malpighian tubules of the New Zealand Alpine Weta (Hemideina maori). Fluid secretion by tubules in iso-osmotic saline (500mOsm) occurred at a rate of 15+/-3nlh(-1) and was enriched in K(+) (approx. 125mmoll(-1)) relative to the saline (10mmoll(-1)). Maximal fluid secretion (112nlh(-1)) during simultaneous exposure to hypo-osmolality and dibutyryl cAMP resulted in an 8.8x increase in the quantity of K(+) secreted, compared to only a 2.4x increase in Na(+) secretion. Measurements of intracellular ion activities and membrane potentials indicated that Na(+) and K(+) were transported against a strong electrochemical gradient across the apical surface, regardless of saline osmolality. On the basolateral surface, there was a large driving force for Na(+) entry, while K(+) was distributed near its equilibrium potential. Neither bumetanide nor ouabain in the bathing saline had a significant effect on fluid secretion, but Ba(2+) and amiloride decreased fluid secretion by 79 and 57%, respectively. The effect of Ba(2+) on fluid secretion was consistent with a high basolateral permeability to K(+), relative to Na(+) and Cl(-). These results indicate that the characteristics of fluid secretion in this primitive insect are largely conserved with characteristics reported for other insects.     

Properties of a feedback model for the circadian clock of Hemideina thoracica (Orthoptera; Stenopelmatidae)

The single oscillator feedback model describing the circadian system of the nocturnal insect, Hemideina thoracica, (Gander and Lewis, 1979) has been developed and refined by restricting the range of parameter values which successfully simulate the known behaviour of this insect's circadian system. The model accounts for Aschoffs Law for nocturnal animals, and makes new predictions on the combined effects of temperature and constant light on the period of the free-running rhythm, which were verified experimentally. These simulations also indicate that the same general feedback model can be used to describe the circadian systems of other organisms.