In situ measurement of calling metabolic rate in an Australian mole cricket, Gryllotalpa monanka.

Examination of the energetics of sound production usually requires measurement of species that will produce normal calls under unnatural circumstances. Such measurements are potentially compromised by stress-related changes in calling input (through a reduction in calling effort) or output (through forced use of sub-optimal singing burrows). To determine if such measurements are indeed affected by abstraction from a natural setting, we measured the energetics of song production in undisturbed mole crickets Gryllotalpa monanka and employed a new approach where the animal's singing chamber replaces the respirometry chamber normally used in studies of this type. It was therefore possible to measure metabolic rate (MR) of calling crickets in situ for animals within self-constructed burrows under natural conditions. Calling MR measured under these conditions averaged 13.5-fold higher than standard MR and 2.2-fold higher than MR measured during burrowing in the lab. The calling MR of G. monanka was similar to that measured for other calling insects, and to endothermic insects, but was only 10% of that allometrically predicted for a similarly sized insect (0.89 g) during flight. A male mole cricket is estimated to consume 5.9 ml of oxygen during construction of a calling burrow and a 1-h calling bout; by comparison, a flying female would consume a similar volume in less than 6 min.     

Gecko phonotaxis to cricket calling song: A case of satellite predation

Insectivorous Mediterranean house geckos, Hemidactylus tursicus, are found in the field close to the burrows of calling male decorated crickets, Gryllodes supplicans. In playback experiments where adult geckos were presented with calls of these crickets or calls of frogs as controls, geckos exhibited positive phonotaxis to broadcast cricket calling song. This indicates that geckos orient to and approach male cricket calls, even though the crickets call from burrows where they are protected from the geckos. However, this behaviour enables the geckos to intercept and consume female crickets that also respond phonotactically to the cricket calls. Thus geckos act as ‘satellite predators’, a situation which may impose sex-biased mortality on female crickets.     

The underground life of the oriental mole cricket: an analysis of burrow morphology

The underground life of the oriental mole cricket Gryllotalpa orientalis has been investigated by studying the structure of its burrows under different environmental situations and in different seasons. The different uses of different burrow types and their advantages and disadvantages have been examined. The total length, number of tunnels and combination of burrow types varied from a simple tunnel to a more complex one with branches at various angles to the surface, burrow types being divided roughly into shallow horizontal or deep vertical ones. In horizontal burrows, the branching structure was well developed in various directions. It is notable that the vertical burrows of G. orientalis were occupied by only one individual. Both vertical and horizontal burrows were used for foraging: vertical burrows for plants with subterranean stems and horizontal burrows for creeping plants. Vertical burrows were also used for hiding from predators, resting, moulting and overwintering, whereas horizontal burrows were used for escaping from predators and as mating routes. Egg chambers were constructed beside horizontal burrows, and calling burrows were constructed as part of horizontal burrows. Based on their current requirements, mole crickets continuously modify their burrow structures or change burrowing sites.     

A Study of the Behaviour of the Australian Field Cricket Teleogryllus commodus (Walker) (Orthoptera: Gryllidae) in the Field and in Habitat Simulations

The behaviour of T. commodus (Walker) in the field and in simulated natural habitats is described. Regular predictable sequences of behaviour are represented by an ethogram. Most behaviour was centred around burrows or sheltered sites. Burrows appeared to be a limiting resource and there may be behavioural dimorphism in ♂ ♂ (burrower and non-burrower strategies). Individuals were found to form gregarious clusters and appeared to be attracted to areas already occupied by crickets. Territorial spacing and other social interactions occurred within these clusters. ♂ ♂ and ♀ ♀ were found to be mobile, and moved continually between sites. The movement of ♂ ♂ and ♀ ♀ between burrows (calling sites) was measured. Adult ♂ ♂ never remained at a burrow for longer than two days. In simulated natural habitats ♂ ♂ moved in response to two factors: first, and most common, as a result of eviction by other ♂ ♂ second, as a response to the absence of ♀ ♀. When cricket density was increased in habitat simulations crickets clustered at calling sites. The adaptive significance of T. commodus behaviour is discussed.     

Strategies of mate finding in the European field cricket (Gryllus campestris) at different population densities: a field study

Abstract. 1. Members of a field population of Gryllus campestris L. varied in their walking and calling activity. In both sexes, some individuals occupied burrows whereas others walked around in the observation area. Males at burrows could be either silent or calling.
2. In the course of one summer, population density decreased and the initial balanced sex ratio changed to a large surplus of males.
3. At high population density, there were equal numbers of non-calling males at burrows, calling males at burrows and walking males, while walking females predominated over females at burrows. Non-calling males at burrows achieved more encounters with females than did calling and walking males. Females met males by walking through the population and by waiting at burrows. Thus, calling and phonotaxis were not essential for mate finding and calling was less effective than previously thought.
4. At low population density calling males predominated. Calling males at burrows achieved the most encounters with females. Females met males only by walking around in the population area. Calling was thus more important in mate finding than at high population density.
5. Changes in sex ratio and population density may cause the flexibility in mate finding behaviour of individual crickets.     

Power output and efficiency of sound production by crickets

I estimated the sound power of male crickets calling outdoorsusing a microphone array to measure their sound fields. Averagepower output for four species, Scapteriscus borellii (N = 18),S. vicinus (N = 7), Oecanthus quadripunctatus (N = 6), and Anurogryllusarboreus (N = 6), was 6.5, 4.5, 10.2, and 58.6 µW, respectively.Power output for male mole crickets (Scapteriscus spp.) wasvariable (range 2–22 nW), and estimated efficiency wasless than 0.1%. Efficiencies of sound production for tree crickets{Oecanthus) and short-tailed crickets (Anurogryllus) were between1% and 3%. Power output of S. vicinus males was significantlycorrelated with soil moisture near the male's burrow. LargerOecanthus males produced significantly more power than smallermales. Male Oecanthus that used leaves as baffles increasedpower output by a factor of 2 (3 dB). Radiation patterns ofScapteriscus sound fields were hemispherical. Those of Oecanthusand Anurogryllus were dumbbell shaped. Radiation patterns ofScapteriscus and Oecanthus are predicted by simple physicalsystems. The costs and benefits to male reproduction due o increasedpower output relate to changes in efficiency of sound productionand preferences used by females, competitors, or natural enemiesin locating calling males. [Behav Ecol 1991; 2:327–338]     

The Energetics of Acoustic Signaling in Anurans and Insects

SYNOPSIS. This review focuses on the energetics of advertisementcalls in frogs and insects (mostly ensiferans). I also reviewa number of methodological questions relating to the most appropriateways to normalize metabolic rates for calling animals and tocalculate efficiency of sound production Although the mechanism of sound production is very differentin these groups (vocalization vs. stridulation), net metaboliccosts normalized to mass are similar among species producingthe most conspicuous calls. Features of the call that interactto determine energetic cost include repetition rate, call duration,and intensity (sound pressure level). Anurans tend to producelouder (more intense) calls while ensiferans tend to producesound during a greater proportion of the calling bout. All evidencesuggests that advertisement calls are produced aerobically andthe aerobic costs are similar or exceed the aerobic (but notnecessarily total costs) of terrestrial locomotion The pattern of radiated sound tends to be constant within aspecies and can be predicted to some degree from characteristicsof an animal's acoustic radiator. Efficiency of sound production(acoustic power/net metabolic power) is low (0.05 to 6%) andvariable when compared to locomotion ({small tilde} 10–20%).From the present sample it appears that frogs are more efficientthan ensiferans, but as more katydids are studied this trendmay not hold. Of the factors that have been identified as determiningefficiency the most important are the match between the sizeof the radiator and wavelength radiated, the absorption propertiesof the environment immediately around the animal, and the presenceor absence of structures such as baffles and acoustic burrows     

Signaling or Not-Signaling: Variation in Vulnerability and Defense Tactics of Armored Ground Crickets (Acanthoplus Speiseri: Orthoptera, Tettigoniidae, Hetrodinae)

Male Orthoptera singing from exposed perches are at risk from acoustically- and visually-hunting predators. The defensive reactions of armored ground crickets (Acanthoplus speiseri) include falling silent, dropping from their perch, alarm stridulation and autohaemorrhaging. Male and female ground crickets show different reactivity (i.e. the number or intensity of defense tactics used) to predation, depending on level of exposure: calling males were more reactive when approached during daylight, compared with in the dark. During daylight, calling males were more reactive than silent, cryptic, males and females. The level of response presumably reflected the riskiness of the individual’s behavior and situation at that time. Plasticity of response to predation allows individuals to balance risky behavior (i.e. acoustic signaling from exposed perches) by being more reactive to potential threats.     

The energetic cost of calling in the variable field cricket, Gryllus lineaticeps

Abstract. Female preferences for conspicuous male calls have been documented in many groups. However, relatively few studies have examined the metabolic costs associated with the production of call types preferred by females. We measured the oxygen consumption of calling male Gryllus lineaticeps Stål crickets using closed chamber respirometry. Calling song was recorded concurrently. The average increase in mass-specific oxygen consumption during calling was 2.7 times basal rates of oxygen consumption, and calling males consumed approximately 1.2ml O₂g^-lh^-1. Oxygen consumption increased with increasing chirp rate and pulse duration, but not with increasing chirp duration. Females of this species prefer higher chirp rates, thus some call types that increase the male's attractiveness to females require more metabolic energy to produce.     

The Effect of Age on a Sexually Selected Acoustic Display

Models of sexual selection suggest that females should prefer to mate with older males because old age is evidence of heritable high viability. In a longitudinal analysis, we demonstrate that male field crickets (Gryllus campestris) alter their calling song with age. Carrier frequency, a calling song character related to growing condition and the main song component under female preference, changed towards higher sexual attractiveness with age. Body mass decreased slightly with age, while chirp rate, an indicator of current condition, remained stable. By choosing males singing at a low frequency, female field crickets would base their mate choice decision on a sexual trait that indicates superior growing conditions as juvenile and on viability, i.e. enhanced current condition as adult.     

Acoustic communication and sexual selection in Orthoptera (Insecta)

Comparison of calling and courtship songs and mating strategies in different groups of Orthoptera shows that acoustic signals in bush crickets (Tettigonioidea) are used for searching conspecific mates at a distance, and song evolution is primarily driven by the acoustic surroundings and simultaneously singing other species. The role of sexual selection in the evolution of acoustic signals in bush crickets is much less important than in the evolution of other nonacoustic signals employed during direct contact. Acoustic signals of crickets (Grylloidea) are involved in both distant and short-range communication. Acoustic courtship signals, along with other courtship components, may offer the material for sexual selection, although, essentially, evolution of acoustic signals in crickets is determined by the acoustic surroundings. Acoustic communication in the grasshoppers of the subfamily Gomphocerinae is mainly a short-distance communication. Acoustic signals of Gomphocerinae are highly variable and elaborate in temporal parameters, and therefore can serve for evaluation of the mate "quality". Song evolution in this group is to a greater extent driven by sexual selection than by the acoustic surroundings and, therefore, could proceed faster than in other groups of Orthoptera and play a major role in speciation.     

Landscape and weather determinants of prey availability: implications for the Lesser Kestrel Falco naumanni

Spatial and temporal variation in prey abundance have been shown to impact the time of breeding and breeding success of birds. Understanding the ecological requirements of preferred prey can help develop management measures to improve food supply for target species. For the colonial Lesser Kestrel Falco naumanni, mole crickets Gryllotalpa spp. are one of the most important prey items during the mate‐feeding period. Lesser Kestrel colonies with higher mole cricket consumption had earlier egg‐laying dates, suggesting that differences between individuals in the time of breeding could be caused by differences in the diet. Moreover, the mean number of mole crickets in pellets was significantly correlated with clutch size (in one of the studied years) and egg volume. Thus, the impact of environmental variables and land use on mole crickets is likely to be relevant to Lesser Kestrel conservation. Weekly consumption of mole crickets was higher following an increase in either precipitation or minimum temperature values. Furthermore, mole cricket consumption was higher in colonies surrounded by higher quality soils and in wetter areas and years. Predicted probability of mole cricket occurrence in surveyed watercourse margins suggested a positive relationship between soil penetrability and mole cricket occurrence. Among variables that might be the target of management, the presence of riparian vegetation positively influenced the occurrence of mole crickets, whilst tillage and sowing of streambeds were revealed as the most important threats. We suggest that the maintenance of native vegetation in the margins of watercourses could improve soil resilience to erosion, increase water retention, soil penetrability and fertility, and provide a food supply and shelter for mole crickets. Overall, the implementation of such recommendations is likely to benefit other farmland species known to consume mole crickets, including several endangered species.     

Males make poor meals: a comparison of nutrient extraction during sexual cannibalism and predation

Cannibalism is hypothesized to have evolved as a way to obtain a high-quality meal. We examined the extraction of lipid and protein by female wolf spiders, Hogna helluo, during sexual cannibalism of males and predation of crickets. Most food-limited females did not cannibalize males but immediately consumed a size-matched cricket. When consuming male H. helluo and crickets, female H. helluo only consumed 51% of the male body while they consumed 72% of the cricket body. While males had higher protein content in their bodies than crickets and other insects, female H. helluo ingested similar amounts of protein from male H. helluo and crickets. Female H. helluo extracted 47% of the protein present in male H. helluo and 67% of the protein present in crickets. Females were able to extract nearly all of the lipid present in male H. helluo and crickets. However, crickets and other insects had almost 4 times higher lipid content than male H. helluo. The ratio of lipid to protein consumed from crickets appeared more similar to the nutritional requirements of egg production than that of males. Taken together, female hesitancy to engage in cannibalism, low extraction of nutrients from males and a low ratio of lipid to protein in the food extracted from males suggest that males may be poor-quality prey items compared to common insects such as crickets.     

Sex differences in metabolic rates in field crickets and their dipteran parasitoids

Sex differences in metabolic rate (MR) can result from dimorphism in the performance of energetically demanding activities. Male crickets (Teleogryllus oceanicus) engage in costly calling and aggressive activity not performed by females. Consistent with this difference, we found higher maximal MR, factorial scope, and fat content in males than females. T. oceanicus song is also costly because it attracts the parasitoid fly Ormia ochracea. Parasitized crickets had reduced maximal MR consistent with a metabolic cost to harboring larvae. This cost was greater for females, either because females invest more heavily into reproduction at the expense of metabolic capacity, or because males are under stronger selection to respond to infection. Little is known about O. ochracea outside of its auditory system and parasitic lifestyle. We observed greater resting MR in male flies, possibly reflecting a sex difference in the requirement for metabolic power output, because male flies perform potentially costly mating behavior not seen in females. We found a positive relationship between larval density within a cricket and pupal resting MR, suggesting that crickets in good condition are able to both harbor more larvae and produce larvae with higher resting MR. These results reveal a complex interplay between the metabolism of crickets and their fly parasitoids.Communicated by G. Heldmaier     

Sex,death, and genetic variation: natural and sexual selection on cricket song

Male field crickets, Gryllus integer, in Texas, USA, produce a trilled calling song that attracts female crickets, resulting in enhanced mating success. Gravid female parasitoid flies, Ormia ochracea, are also attracted to male cricket calling song, resulting in the death of the male within about seven days. Using playbacks of field-cricket calling song in the natural habitat, we show that both female crickets and female parasitoid flies prefer male calling song with average numbers of pulses per trill. Thus female crickets exert stabilizing sexual selection, whereas flies exert disruptive natural selection on male song. Disruptive natural selection will promote genetic variation and population divergence. Stabilizing sexual selection will reduce genetic variation and maintain population cohesiveness. These forces may balance and together maintain the observed high levels of genetic variation (ca. 40%) in male calling song.     

Biology of cave crickets,Hadenoecus subterraneus,and camel crickets,Ceuthophilus stygius (Insecta: Orthoptera): Metabolism and water economies related to size and temperature

• 1.1. Metabolic rate (MR) and water budget (WB) components of cave and camel crickets are directly related to size and temperature.
• 2.2. MR increases most rapidly with size for insects in general followed by cave crickets (females > males), and lastly, camel crickets (no sex differences).
• 3.3. Metabolic thermal sensitivity of cave crickets (males > females) is much greater than camel crickets.
• 4.4. WB components parallel MR relations.

     

Heat transfer and the energetic cost of singing by canaries <Emphasis Type="Italic">Serinus canaria</Emphasis>

Sexually selected displays, such as male passerine bird song, are predicted to be costly. However, most measurements calculated the rate of oxygen consumption during singing using respirometry have shown that bird song has a low energetic cost. Since birds are reluctant to sing when enclosed inside a respirometry chamber, the energetic cost of singing could differ from that under more normal circumstances. We used heat transfer modelling, based on thermal images, to estimate the energetic cost of singing by canaries (Serinus canaria) that were not enclosed in respirometry chambers. Metabolic rate calculated from heat transfer modelling was 0.70±0.02 W (N=10 birds) during singing, which was 14±5% greater than during standing (0.62±0.02 W). The energetic cost of singing did not differ significantly from that measured previously using respirometry when we took into account that birds sang for a greater proportion of the time during the current experiments. These conclusions were not sensitive to potential errors in the heat transfer model. Heat transfer modelling would be especially useful to obtain measurements of the energetic cost of activities that animals do not perform readily inside respirometry chambers, such as singing in birds.     

Conservation and the cricket culture in China

Several species of Tettigoniidae and Grylloidea are collected and traded as singing or fighting insects in China. There is anecdotal evidence that this trade is causing a reduction in the ranges and populations of some species. However, before any conservation status can be assigned to any species reliably, it will be necessary to reconcile the long-established traditional classification system of singing and fighting insects with scientific classification, determine the ranges and monitor the populations of selected species of commonly traded species, and initiate an educational programme for invertebrate conservation using these crickets as flagship taxa.     

Influence of age and environmental factors on burrow-making behavior of the short-tailed cricket,Anurogryllus muticus (De Geer) (Orthoptera: Gryllidae)

For the short-tailed cricket, Anurogryllus muticus, burrow-making behavior is essential. All nymphal instars construct burrows, but in the adult stage the rate of burrowing behavior is age dependent. Increases in photophase and light intensity stimulate burrowing, and the explicit negative phototaxis is correlated with the cricket's inability to exist under dry conditions. Ingestion of substrate during burrow construction may serve to acquire additional moisture. There is no evidence of burrow recognition, and crickets can construct a burrow when needed. The natural distribution of burrows at the plot investigated on Moorea supports the notion thatA. muticus builds burrows where the preferred food plantAlysicarpus vaginalis is most abundant. By minimizing the traveling distance to food sources when foraging they can retreat to their burrow again.     

A comparison of light and temperature entrainment: evidence for a multioscillator circadian system

ABSTRACT. Both photoperiodic and thermoperiodic cycles synchronize circadian calling activity of male crickets, Teleogryllus commodus (Walker). Because the phase relationships between these cycles and the entrained singing activity are clearly distinguishable, we studied the relative power of these factors in affecting the circadian clock. Upon resumption of constant conditions after exposure to a thermoperiodic cycle, singing activity sometimes splits into two daily bouts, each with a distinctive period. This observation, together with results derived from simultaneous fluctuation of both factors in and out of phase, suggests that the system integrating environmental input is composed of multiple oscillators.