Diel Calling Activity and Field Survival of the Bushcricket,Sciarasaga quadrata (Orthoptera: Tettigoniidae): A Role for Sound-locating Parasitic Flies?

In this study I examine the variability in calling activity of Sciarasaga quadrata over both the short (4 d) and long term (life span), and the temporal pattern to host searching by its most significant natural enemy, the ormiine fly Homotrixa alleni. Indeed, few male S. quadrata survive the entire calling season as this acoustically orienting parasitoid fly continually ‘culls’ calling males out of the population. I found that male S. quadrata commence calling over 1 h before sunset, cease around midnight and call, on average, for over 3 h within this period (range 0–11 h 09 min). There was significant concordance and repeatability in calling activity over both the short and long term, enabling me to conclude that relative differences in calling activity among males persist throughout a male's life span. There was a distinct peak to host searching by gravid female flies, with 82 % of all flies collected at acoustic traps between sunset and midnight. A positive association between call duration (time spent calling) and fly attraction was evident from multiple collections of flies at single traps. Counter to expectation, there was no significant decline in average call duration across successive collections of males, indicating that long-call-duration males were not being selectively culled by flies from the calling population. Males at the end of the calling season, when their remaining reproductive potential is low, allocate the majority of their calling time between sunset and midnight despite the high risk of fly attack. The distribution of call durations within the population was not skewed, which if heritable, indicates a lack of history of directional selection acting on call duration. Limited evidence suggests that encounters with females are rare in S. quadrata so that males may be selected, irrespective of natural selection pressure, to call for long periods of time to optimize their chance of attracting mates. The above findings are further discussed in relation to parasitoid foraging and acoustic reproductive strategies.     

The biology of the phonotactic parasitoid, Homotrixa sp. (Diptera: Tachinidae), and its impact on the survival of male Sciarasaga quadrata (Orthoptera: Tettigoniidae) in the field

Abstract.

• 1 Unlike most parasitoids, tachinid flies of the tribe Ormiini use sound to locate their hosts. Although thought to exert selection pressure on their host's calling behaviour, little is known about the biology of ormiines. Accordingly, this study reports the biology and impact of the ormiine Hornotrixa sp. upon calling males of the univoltine bushcricket Sciarasaga quadrata Rentz in south-western Australia.
• 2 Populations of adult S.quadrata were monitored in the field over two successive calling seasons. Females, which do not call, were not parasitized by Hornotrixa sp., but the risk of parasitism for males increased as the 3-month calling season progressed. Parasitism did not commence until c. 2 weeks into the calling season, but by the end of the season up to 87% of surviving males were parasitized.
• 3 Parasitized males lived for 14 days and were found singing until their penultimate evening before death in the field. Unparasitized males lived on average 69 days and a maximum of 119 days.
• 4 Multiparasitism of hosts was common, with up to sixteen fly larvae found within parasitized males. The number of fly larvae within hosts significantly increased at the end of the season. However, successful emergence of fly larvae from hosts, as well as pupal size, significantly decreased as more than one fly larva developed within the host.
• 5 Hornotrixa sp. has a long pupal duration of 30–31 days at 20°C. As a consequence, only one complete fly generation, which overwinters in the pupal stage, is likely within each host generation.
• 6 No evidence for differential (size-bias) mortality by Hornotrixa sp. on male S.quadrata was found. The size of parasitized and unparasitized males collected in the field was not significantly different.
• 7 It is concluded that Hornotrixa sp. is a significant mortality factor acting on the survival of adult male S.quadrata.

     

Fluctuating asymmetry, call structure and the risk of attack from phonotactic parasitoids in the bushcricket Sciarasaga quadrata (Orthoptera: Tettigoniidae)

The acoustically orienting parasitoid Homotrixa alleni (Diptera: Tachinidae) is a significant mortality factor of calling male Sciarasaga quadrata (Orthoptera: Tettigoniidae), with less than 10% of males escaping parasitism within the duration of the calling season. This study examined fluctuating asymmetry (FA) (small, random deviations from perfect symmetry) in five morphological traits in S. quadrata and its relation to the survival of calling males in the field. A significant relationship was demonstrated in only a single trait; across three consecutive field seasons and among each of the collections per season, parasitised males were shown to have significantly more asymmetric hind tibia than unparasitised males. Since the morphological development of males is complete prior to parasitoid attack, this relationship is not caused by the activities of the parasitoid. Further experimentation revealed that there was no relationship between the level of FA in the hind tibia and the success of parasitism, with males artificially parasitised in the laboratory showing no difference in FA from those where parasitism was unsuccessful. Instead, FA was shown to be negatively related to chirp length, an aspect of call structure that significantly affects the survival of calling males under parasitism risk in the field. We explore the possible reasons why more asymmetric males produce calls of shorter chirp length and hypothesise that the difference in hind tibia asymmetry seen between parasitised and unparasitised males in the field is related to host location in H. alleni rather than the inability of S. quadrata to defend itself against parasitoid attack. Received: 14 November 1997 / Accepted: 20 April 1998     

Impacts of defoliation by tussock moths (Orgyia vetusta) on the growth and reproduction of bush lupine (Lupinus arboreus)

Abstract.

• 1 Outbreaks of tussock moths (Orygia verusta Bdv.) on bush lupines (Lupinus arboreus Sims) may be very intense, although spatially localized, and may last >10 years. To understand better how such outbreaks may persist, we defoliated lupine bushes using variable realistic numbers of tussock moth larvae, and measured the immediate and delayed impacts of defoliation on the growth and seed production of lupine bushes.
• 2 Immediate effects of defoliation included the production of new leaves that were significantly smaller in diameter and biomass, and lower in water content, than the new leaves of undamaged bushes. Also, seed output was reduced by up to 80% on heavily damaged bushes.
• 3 In the longer term, surviving bushes recovered remarkably well, despite having received the full range of possible (survivable) insect densities. After 1 or 2 years’rapid growth, surviving bushes were not affected in terms of height, basal stem diameter or volume by their previous defoliation. However, there was a significant tendency for juvenile bushes to produce more seeds the more heavily they had been attacked the previous year.
• 4 The ability of the host plant to recover, in terms of its biomass, helps to explain the sustained nature of the insect outbreak.

     

Effect of trap height and within-planting location on captures of cranberry fruitworm (Lepidoptera: Pyralidae) in highbush blueberries

Abstract 1 Winged traps baited with synthetic sex pheromone lures [(E,Z)‐8,10‐pentadecadien‐1‐ol and (E)‐9‐pentadecen‐1‐ol acetate] were evaluated for their effectiveness in monitoring cranberry fruitworm, Acrobasis vaccinii Riley, in highbush blueberry, Vaccinium corymbosum L., plantings. Trap effectiveness was compared at different heights within the bush canopy and different locations within plantings. 2 In our trap height study, three positions were evaluated: (i) at the top of bush canopy (15 cm below the uppermost branch); (ii) centrally within bush canopy (60 cm below the uppermost branch); and (iii) at the bottom of the bush, 20 cm above ground level. Traps placed 15 and 60 cm below the uppermost branch captured significantly more male moths compared with traps placed 20 cm above ground level at two organic sites. 3 In our trap location study, four treatments were evaluated based on trap location relative to adjacent woodlands: (i) in trees within 1 m of the woodland boundary; (ii) in blueberry bushes adjacent to woodlands, 15 m from the woodland boundary; (iii) in blueberry bushes in the centre of the planting, 75 m from the woodland boundary; and (iv) in blueberry bushes furthest away from woodlands, 150 m from the woodland boundary. Traps located within 1 m of woodland boundary captured significantly more male moths compared with traps located centrally (15 and 75 m) within plantings.     

Bush selection along foraging pathways by sympatric impala and greater kudu

In order to identify the selection mechanism of two sympatric African browsers, we analysed encounter rates and selection of bushes along foraging pathways. We monitored the tracks, left overnight, by kudu and impala on an experimental plot of natural Acacia nilotica and Dichrostachys cinerea in the highveld of Zimbabwe, and recorded the number of bushes attacked in each category. Both ungulates were selective for the bush categories, but kudu were consistently more selective than impala, and showed a higher preference for the larger A. nilotica and D. cinerea bushes, which had a significantly greater number of bites which were not reachable by impala. For both kudu and impala, the probability of attacking larger bushes increased significantly with the proportion of large bushes encountered along the foraging pathways, whereas the consumption of smaller bushes was apparently unpredictable. For the most abundant food item (medium D. cinerea), the probability of attack by impala along a pathway decreased with increasing proportions of larger bushes in the experimental area, but was also dependent on impala group size and season. In addition, we found that encounter rates with larger bushes were significantly higher for kudu than for impala. Experimentally reducing the availability of the larger bushes had little effect on both impala and kudu during the following rainy season. However, during the following cool dry season, kudu showed an increased selectivity with a strong preference for the remaining large bushes (large A. nilotica), followed by a sharp decrease in selectivity in the hot dry season when they also fed from significant numbers of medium trees. Impala had little reaction to the experimental changes in the availability of bush categories in either season. We suggest that both kudu and impala selected bushes on the basis of the potential number of bites they can provide, and this resulted in different search strategies. Kudu focussed on the larger bushes which have a larger number of twigs which are out of reach of impala and kudu also probably directed their path preferentially towards the few larger bushes to maximize encounter rates with this favoured bush category. These differences in bush selection process lead to a low overlap in resource use between the two browsers in this type of savanna.     

Susceptibility of Scaevola taccada (Gaertn.) Roxb. bushes to attack by the coccid Icerya seychellarum Westwood: the effects of leaf loss

Abstract.

• 1 The distribution of infestation levels of the coccid Iceryu seychellarum on bushes of Scuevola taccudu and its population fluctuations over a 10 month period on Aldabra atoll show that individual plants of S. taccada exhibit differential susceptibility to attack.
• 2 Coccids on heavily and lightly infested bushes show no important differences in growth rate, fecundity or plant induced death rate. There were also no demonstrable differences in bush exposure, food quality (total and soluble nitrogen levels) or settlement behaviour of first instar larvae (crawlers).
• 3 Detailed examination of Iceryu populations on individual leaf clusters showed that mortality caused by leaf fall is density dependent and can be avoided only if a large proportion of the population frequently moves from senescing to mature leaves. The proportion of the population required to move to younger leaves in order to complete development is greater in low density Zceryu populations.
• 4 The effects of leaf fall are discussed with reference to the maintenance of differential susceptibility within the plant population.

     

Resource distribution and its effect on the mating system of a longhorned beetle,Perarthrus linsleyi (Coleoptera: Cerambycidae)

Summary Adults of Perarthrus linsleyi feed on flowers of creosote bush, Larrea tridentata. Mating also occurs on the flowers and foliage of this plant. This food resource is widely and evenly distributed in space, and is usually abundant. The spatial distribution of the beetles bore no relationship to the spatial distribution of flowers among creosote bushes, nor were female beetles distributed predictably with respect to flower distribution. Males moved widely over the area under study, and moved much more frequently than females. Males actively searched for females, and mounted and began copulation attempts without preliminary courtship. Males did not engage in aggressive defense of females, creosote flowers, or creosote bushes. This species exhibits a scramble competition mating system. The spatial distribution of the food resource is a primary factor in the evolution of the mating system of Perarthrus linsleyi.     

Acoustically-orienting parasitoids in calling and silent males of the field cricket Teleogryllus oceanicus

Abstract.

• 1 On three Hawaiian Islands, the introduced Australasian field cricket Teleogryllus oceanicus Le Guillou (Orthoptera: Gryllidae) was found to be attacked by the phonotactic parasitoid tachinid fly, Ormia ochracea Bigot.
• 2 Noncalling males occurred with callers in all locations, but silent males were more heavily parasitized than callers.
• 3 Body size was unrelated to both calling status and the likelihood of harbouring parasitoid larvae.
• 4 An experiment examining the likelihood of calling in the laboratory by males collected as silent or calling individuals showed no difference between the two classes of males, after accounting for parasitoid levels; males harbouring larvae were less likely to call.

     

Transmission and perception of acoustic signals in the desert clicker,Ligurotettix coquilletti (Orthoptera: Acrididae)

Males of the desert clicker, Ligurotettix coquilletti (Acrididae: Orthoptera) defend a femalerequired resource, the creosote bush Larrea tridentata, in desert habitats of the southwestern United States. Males signal acoustically to each other as well as to searching females. The call is produced by tegminal/femoral stridulation where one or both legs are used for sound production. Sound pressure levels, measured laterally, are influenced by the intervening tegmen between the stridulating leg and the microphone. Differences in measured sound pressure levels between sides can vary up to 7 dB. When clicks are produced multiply,these multiple clicks may be 4 dB louder than single clicks. We examine the structure of the call and the effective broadcast area of single males by monitoring acoustic ascending neurons of the ventral nerve cord in the neck. By taking the neurophysiological preparation into the field, we were able to map the broadcast area of isolated males and also of males calling within aggregations. The distance over which the signal of isolated males could be detected was 8–14 m, whereas neural representation of the calls of males within aggregation were detectable within 4–6 m. The sound spectrum of the song, although having a major lower-frequency component around 10 kHz, has extensive power in the ultrasonic range. The tuning characteristics of the ascending auditory neuron matched the overall structure of the male call. The importance of the acoustic cue, as compared to visual cues, is discussed in relation to female attraction.     

Assessment of the range of attraction of pheromone traps to Agriotes lineatus and Agriotes obscurus

• 1 The range of attraction of YATLOR pheromone traps was studied to gain information on the number of traps needed for mass trapping of males of two Agriotes species.
• 2 Male click beetles of the species Agriotes lineatus (L.) and Agriotes obscurus (L.) (25–30 individuals per release point) were marked and released at a distance of 2, 5, 10, 15, 20 and 60 m from a pheromone trap both along and opposite to the known prevailing wind direction. Traps were regularly inspected over approximately 1 month. The percentage of recaptured beetles was calculated and analyzed using analysis of variance. Maximum sampling ranges and effective sampling areas were calculated.
• 3 Averaged over all five trials and distances, approximately 40% of the released beetles (A. lineatus and A. obscurus) were recaptured. The percentage recapture of male adults was significantly affected by release distance, whereas no differences were found for species and release direction.
• 4 Males were recaptured from all release points and the percentage recapture decreased (in part significantly) with increasing distance from 76% (2 m) to 35% (15 m) and 9% (60 m), respectively. Most of the beetles were recaptured within the first 3 days after release, independent of the distance, except 60 m. The effective sampling area for A. lineatus was 1089 m² after 12 days and increased to 1735 m² after 30 days. Corresponding values for A. obscurus were considerably higher: 1518 m² for 12 days and 2633 m² for 30 days.
• 5 We conclude that the range of attraction of the pheromone traps for A. lineatus and A. obscurus is comparatively low, providing high percentage recapture only for release distances up to 10 m. Accordingly, any approach targeted on preventing mating by male mass trapping would require a dense network of pheromone traps.

     

Hilltopping, leks and female choice in the Carpenter bee Xylocopa (Neoxylocopa) varipuncta

The mating system of the Carpenter bee, Xylocopa (Neoxylocopa) varipuncta Patton, closely resembles the leks of certain vertebrates. Males defend hovering stations at small creosote bushes on ridgetops in the Sonoran Desert, as well as at large ironwood trees in dry washes. Ridgetops, like washes, may provide orientation guides for dispersing females, channelling potential mates to waiting males. During the height of the two month flight season, both males and females were more concentrated on the ridge than in the wash. Males were particularly densely aggregated at or near two prominent peaks on the study ridge. At times, two or more males occupied the same creosote bush. Females visited hovering males infrequently, and exhibited a high degree of freedom in mate choice, usually rejecting males after a close inspection.
The reduction in territorial intolerance exhibited by some males at ridgetop sites may occur: (1) because of the costs of repelling many intruders and (2) because females may choose among males through direct analysis of male phenotype, rather than by selecting a male on the basis of territory position.     

The effects of male body size on territorial and mating success in the landmark-defending fly Hermetia comstocki (Stratiomyidae)

Abstract.

• 1 Males of Hermetia comstocki Williston compete for territorial control of certain agaves and yuccas. Winners copulate with females that visit these plants solely to acquire a mate.
• 2 Males vary in body weight by more than an order of magnitude and larger flies almost always defeat smaller ones in aerial contests for control of landmark territories.
• 3 The mean body size (as measured by wing-length) was significantly greater for males retaining residency at a site for at least one hour compared to males unable to do so. Likewise, males able to return to a perch site in the study area on more than one day were larger on average than males unable to do so.
• 4 Male preferences for landmark territories remained similar across years. Large males dominated the perch landmarks most likely to be occupied by males and most likely to be visited by females.
• 5 Despite the fighting and territorial advantages enjoyed by large males, the mean size of males found mating with females was not significantly larger than that of the general population.
• 6 The apparent failure of large males to secure a statistically significant mating advantage may be a statistical consequence of the small sample size of males observed mating. On the other hand, any mating advantage of large males may be reduced because (a) receptive females visit many different landmarks, (b) females mate with the first male they encounter at a landmark, regardless of his size, (c) there are usually many vacant landmarks available for smaller males, and (d) even popular territories are often open to small males, thanks to the low site-tenacity of territory owners.

     

Burrowing behavior and burrowing energetics of a bioindicator under human disturbance

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Inter- and intraspecific variation in nightly calling duration in field crickets,Gryllus integer andG. rubens (Orthoptera: Gryllidae)

The time that male field crickets spend calling was measured electronically in the laboratory to examine nightly calling duration from different species and populations. Male Gryllus integerfrom Davis, CA, and Las Cruces, NM, called approximately 7 h per 24 h, whereas male G. integerfrom San Antonio and Austin, TX, and Norman, OK, and male G. rubensfrom McAlester, OK, and Arkadelphia, AR, called approximately 3 h per 24 h. Variation in duration of calling is discussed in terms of the energetic costs of producing song and selection against calling by acoustically orienting parasites and predators. Duration of calling was also studied infield-collected male G. integer.Males were first observed in field populations and then collected. Those males observed calling in the field later called significantly more in the laboratory than males observed showing non-calling, satellite behavior. Flying G. integercollected under lights were intermediate in their calling duration to calling and satellite males. These data are discussed in terms of the heritability of nightly calling song duration infield crickets.     

Sexual behavior and male volatile compounds in wild and mass‐reared strains of the Mexican fruit fly Anastrepha ludens (Diptera: Tephritidae) held under different colony management regimes

We compared the calling and mating behavior and volatile release of wild males Anastrepha ludens (Loew) with males from 4 mass‐reared strains: (i) a standard mass‐reared colony (control), (ii) a genetic sexing strain (Tap‐7), (iii) a colony started from males selected on their survival and mating competitiveness abilities (selected), and (iv) a hybrid colony started by crossing wild males with control females. Selected and wild males were more competitive, achieving more matings under field cage conditions. Mass‐reared strains showed higher percentages of pheromone calling males under field conditions except for Tap‐7 males, which showed the highest percentages of pheromone calling males under laboratory cage conditions. For mature males of all strains, field‐cage calling behavior increased during the last hour before sunset, with almost a 2 fold increase exhibited by wild males during the last half hour. The highest peak mating activity of the 4 mass‐reared strains occurred 30 min earlier than for wild males. By means of solid phase microextraction (SPME) plus gas chromatography‐mass spectrometry (GC‐MS), the composition of volatiles released by males was analyzed and quantified. Wild males emitted significantly less amounts of (E,E)‐α‐farnesene but emitted significantly more amounts of (E,E)‐suspensolide as they aged than mass‐reared males. Within the 4 mass‐reared strains, Tap‐7 released significantly more amounts of (E,E)‐α‐farnesene and hybrid more of (E,E)‐suspensolide. Differences in chemical composition could be explained by the intrinsic characteristics of the strains and the colony management regimes. Characterization of calling behavior and age changes of volatile composition between wild and mass‐reared strains could explain the differences in mating competitiveness and may be useful for optimizing the sterile insect technique in A. ludens.     

Large scale Agriotes spp. click beetle (Coleoptera: Elateridae) invasion of crop land from field margin reservoirs

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The visual perceptual range of a lizard, <Emphasis Type="Italic">Tiliqua rugosa</Emphasis>

The fragmentation of landscapes produces habitat gaps where the distance between visual landmarks may exceed the perceptual range of a species and impose navigational constraints. We estimated the visual perceptual range of the Australian sleepy lizard, Tiliqua rugosa, by releasing individuals in the centre of a cleared arena in high temperature conditions, with a 0.5-m-high bush placed either 10, 20 or 30 m from the release site. Lizards were more likely to locate those bushes and shelter under them when they were closer, and no lizards found a bush at 30 m. In addition, lizards were less likely to move from the release point when bushes were at 30 m than when they were at the two closer distances. These data suggest that for sleepy lizards the perceptual range for a 0.5-m-high bush is about 20 m. In the uncleared chenopod shrub-land where these lizards live, suitable shelter bushes are an average of 10.5 m from any point in their home range, well within their perceptual range.     

Male Corncrakes Crex crex extend their home ranges by visiting the territories of neighbouring males

Capsule Radiotracked male Corncrake often intruded on the territories of neighbouring males.

Aims To test that intruders' visits are goal-directed, not just a by-product of extended spatial activity during daylight hours.

Methods Using radiotelemetry, we sampled a total of 20 three-day home ranges from 11 tagged males. We recorded daily vocal activity and used a permutation test to see if the movements of tracked males were independent of the position of neighbouring males.

Results The majority of males who had a neighbouring male, up to approximately 600 m from their night calling site, undertook goal-directed visits to the neighbour's territory. Males undertook these visits every day, or every other day, when the neighbours were close. Males undertook visits approximately once every three days when they were more distant. The time spent in the neighbour's territory was longest where the distance between night calling sites was about 200 m. Males tended to be silent in neighbour's territory, apparently to prevent confrontation. Otherwise the distance of neighbouring males did not significantly affect daytime vocal activity. Visiting males tended to sing more often in their home territories.

Conclusions Daily movement of the majority of males was towards the neighbouring male's calling site. We suggest that the purpose of these visits was to seek females. These males may try to drive a female into their territory or gain extra-pair copulation.     

Territoriality and Male Mating Success in the Dart-poison Frog,Epipedobates femoralis (Dendrobatidae,Anura)

The social and reproductive behaviour of the dart-poison frog, Epipedobates femoralis, was studied in Amazonian Peru for 14 months. Males defended territories with advertisement calls and, ultimately, fighting. Territory size ranged from 0.25 to 26 m² and was positively correlated with duration of residence and calling activity of the owner. Females were not territorial and were never attacked when approaching calling males. Males and females only mated once and females sampled calling males before mating. Male mating success was closely correlated with territory size and calling activity. No correlation was found between male body size and mating success. Territories provide residents with sufficient space for mate attraction and reproduction without interference from rivals. Since territory size is dependent on calling activity which involves high energetic costs, it is suggested that territory size reveals male quality.