Mate selection by female katydids (Orthoptera: Tettigoniidae, Conocephalus nigropleurum)

Female katydids receive a large spermatophore at mating which they subsequently eat. Available evidence indicates that spermatophore nutrients are important to female reproduction. Heavier males produce larger spermatophores. When given the choice between two singing males of different weights, females always mated with the larger individual.     

Mate feeding, offspring investment, and sexual differences in katydids (Orthoptera: Tettigoniidae)

Food stress in the katydid Requena veriicalis (Orthoptera: Tettigoniidae)decreases the relative availability of males able to supplynutritious spermatophores to females and increases the valueof the male courtship meal (i.e., relative male parental investment).These changes cause female sexual competition in katydid populations.Here we examine the effect of food stress on male and femaleinvestment in single offspring and test the prediction thatmale-derived nutrients in eggs increase relative to nutrientsfrom the female's reserves. We varied the diet of female R.veriicalis and determined the fate of nutrients from male andfemale sources using I4C and 3H radiolabeled amino acids. Low-dietfemales retained more nutrients from male and female sourcesin somatic tissues and invested less in reproduction both becausethey produced fewer eggs and because they invested less peroffspring (egg) than females maintained on a high-quality diet.Moreover, opposite to our prediction, relative male investmentin individual eggs decreased in foodstressed females; femalesretained more nutrients in somatic tissues from the male sourcethan the female source. Food-stressed females may retain nutrientreserves, particularly those from the male, as an adaptive trategyfor immediate survival needs and future reproduction. Such afemale strategy is unlikely to compromise male reproductivesuccess; first-male sperm precedence means that males matingwith virgin females are likely to father most eggs laid, evenin future reproductive bouts. The decrease in male investmentin eggs of low-diet females does not conflict with the contentionthat relative parental investment controls male intrasexualcompetition because, in mate-feeding species, male investmentinfluencing this competition includes more than investment incurrent offspring; females should compete for males if courtshipgifts aid survival and later reproduction.     

Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago

The extant endemic katydids (Orthoptera: Tettigoniidae) of the Hawaiian Archipelago include one to three species per high island and a single species on Nihoa, all currently placed in the genus Banza. These acoustic insects provide an excellent opportunity for investigating the evolution of reproductive isolation and speciation, but such studies require an understanding of phylogenetic relationships within the group. We use maximum parsimony, likelihood-based Bayesian inference, and maximum likelihood to infer phylogenetic relationships among these taxa, based on approximately 2kb of mitochondrial cytochrome oxidase I and cytochrome b. Our results strongly support two distinct high island clades: one clade ("Clade I") composed of species from Kauai, Oahu, Molokai, and Lanai and another clade ("Clade II") composed of species from Maui and Hawaii (Banza unica, from Oahu, may be basal to both these clades, but its placement is not well resolved). Within these clades, some inferred relationships are strongly supported, such as the sister status of B. kauaiensis (Kauai) and B. parvula (Oahu) within Clade I, but other relationships remain more ambiguous, such as the relative position of B. brunnea (Maui) within Clade II. Although a detailed reconstruction of the historical biogeography of the Hawaiian katydids is difficult, we use our genetic data combined with the known geological history of the Hawaiian Islands to set limits on plausible historical scenarios for diversification of this group. Beyond these historical biogeographic inferences, our results indicate possible cryptic speciation on both Oahu and Hawaii, as well as what may be unusually high average rates of nucleotide substitution. The present work sets the stage for future genetic and experimental investigations of this group.     

Increase in song frequency decreases spermatophore size: correlative evidence of a macroevolutionary trade-off in katydids (Orthoptera: Tettigoniidae)

In many katydids, the male feeds his mate with a large gelatinous spermatophore. Males of most species also produce elaborate calling songs. We predicted a negative relationship between spermatophore size and call frequency because of trade-offs between these two costly traits. Our comparative analysis controlling phylogeny and body size supported this prediction. Although call frequency is expected to decrease with increasing body size, after controlling for phylogeny, both variables were not related. Finally, given that song frequency and spermatophore size are likely targets of sexual selection, we examined the relationship between these variables and sexual size dimorphism (SSD) which can be influenced by sexual selection on body size. We found that only female body size was positively related to SSD, suggesting that natural and/or sexual selection on female body size may be stronger than sexual selection on male and spermatophore size.     

Reverse stridulatory wing motion produces highly resonant calls in a neotropical katydid (Orthoptera: Tettigoniidae: Pseudophyllinae)

This paper describes the biomechanics of an unusual form of wing stridulation in katydids, termed here 'reverse stridulation'. Male crickets and katydids produced sound to attract females by rubbing their forewings together. One of the wings bears a vein ventrally modified with teeth (a file), while the other harbours a scraper on its anal edge. The wings open and close in rhythmic cycles, but sound is usually produced during the closing phase as the scraper moves along the file. Scraper-tooth strikes create vibrations that are subsequently amplified by wing cells specialised in sound radiation. The sound produced is either resonant (pure tone) or non-resonant (broadband); these two forms vary across species, but resonant requires complex wing mechanics. Using a sensitive optical diode and high-speed video to examine wing motion, and Laser Doppler Vibrometry (LDV) to study wing resonances, I describe the mechanics of stridulation used by males of the neotropical katydid Ischnomela gracilis (Pseudophyllinae). Males sing with a pure tone at ca.15 kHz and, in contrast to most Ensifera using wing stridulation, produce sound during the opening phase of the wings. The stridulatory file exhibits evident adaptations for such reverse scraper motion. LDV recordings show that the wing cells resonate sharply at ca. 15 kHz. Recordings of wing motion suggest that during the opening phase, the scraper strikes nearly 15,000 teeth/s. Therefore, the song of this species is produced by resonance. The implications of such adaptations (reverse motion, file morphology, and wing resonance) are discussed.     

Scale effects and constraints for sound production in katydids (Orthoptera: Tettigoniidae): correlated evolution between morphology and signal parameters

Male katydids (Orthoptera: Tettigoniidae) produce mating calls by rubbing the wings together, using specialized structures in their forewings (stridulatory file, scraper and mirror). A large proportion of species (ca. 66%) reported in the literature produces ultrasonic signals as principal output. Relationships among body size, generator structures and the acoustic parameters carrier frequency (f_c) and pulse duration (p_d), were studied in 58 tropical species that use pure‐tone signals. A comparative analysis, based on the only available katydid phylogeny, shows how changes in sound generator form are related to changes in f_c and p_d. Anatomical changes of the sound generator that might have been selected via f_c and p_d are mirror size, file length and number of file teeth. Selection for structures of the stridulatory apparatus that enhance wing mechanics via file‐teeth and scraper morphology was crucial in the evolution of ultrasonic signals in the family Tettigoniidae.     

Structure of the stridulation ofOrchelimum (Orthoptera: Tettigoniidae)

Summary The structure of the stridulation was investigated by re-playing tape-recordings at very slow speed. The findings were corrobrated by sonograms and mingograms.The central part of the song is the ripple, a fast succession of syllables around which isolated syllables (clicks) are distributed according to species and circumstances. The rate of syllables in the ripple is a linear function of temperature.A quantitative expression for the stridulatory activity is the actual number of syllables per time unit, including pauses. By changes in the combination of elements, at leastO. agile is able to increase the output of syllables four to six times.This work was made possible by a grant from the Carlsberg Foundation to whom my most sincere thanks are due. —As I do not have advanced equipment for sound analysis, I am very much indebted to Dr. Bondesen and cand. sci. Poul Hansen, Bioakustisk Laboratorium, Naturhistorisk Museum, Aarhus, Denmark, and Mr. W.B. Broughton and Dr. M. Samways, Animal Acoustic Unit, City of London Polytechnic, London, for analysing part of the material by sonograms and mingograms. The very valuable help of Dr. Th.J. Walker, University of Florida, Gainesville, Florida, in identifying the species is gratefully acknowledged. For friendly discussions and linguistic corrections my best thanks are due to H.T. Evans and F.D.S. Evans.     

Colour polymorphism in Ephippiger ephippiger (Orthoptera, Tettigoniidae)

Colour in the Ephippiger ephippiger complex varies with the geographical distribution and with rearing density. In the W European subspecies E. e. vitium , wild insects from northern and eastern parts of the range tend to have green tergites with relatively inconspicuous banding, whereas those from S France have conspicuously pale banded dark-green or brown tergites. These forms represent the ends of a continuum and there is no zone of distinction. However, the E Pyrenean subspecies, E. e. cunii , lacks the broad banding of its neighbouring E. e. vitium. The tergites of laboratory cultured insects of these stocks when reared in isolation are distinctly green but when reared together become progressively brown. The darkness of the brown is density dependant, so that at high densities the tergites may become almost black. These colour changes are irreversible. Adult insects also become darker with sexual maturity and after contact on mating. Wild-caught E. e. ephippiger from Italy and Yugoslavia have green tergites without obvious banding. Colour changes have not been found or induced in this subspecies. The significance of the colour change is considered and although firm data are wanting, circumstantial evidence suggests a deliberate shift to an aposematic or pseudoaposematic colour pattern.     

Ultrasonic startle behavior in bushcrickets (Orthoptera; Tettigoniidae)

1. In the present work, we show that in flight, bushcrickets not previously known to respond to ultrasound alter their flight course in response to ultrasonic stimuli. Such stimuli elicit in flying Neoconocephalus ensiger an extension of the front and middle legs along the body and a rapid closure of all 4 wings (Fig. 1). This is a short latency acoustic startle response to ultrasound, consistent with acoustic startle responses of other insects. 2. The percentage of trials on which acoustic startle responses were elicited was maximum (90%) for sound frequencies ranging from 25 to at least 60 kHz. No acoustic startle response was observed at frequencies of 5 or 10 kHz (Fig. 2). The threshold for the response was roughly 76 dB between 25 to 60 kHz (Fig. 2) and the behavioral latency was 45 ms (Fig. 3). Recordings from flight muscles show that they cease discharging during the acoustic startle response (Fig. 4). 3. The characteristics of the acoustic startle response match those of an auditory interneuron called the T-neuron. The frequency sensitivity of this neuron is greatest for sound frequencies ranging from 13 to 60 kHz (Fig. 6). Moreover, we found that the neuron produces many more spikes to ultrasound (30 kHz) of increasing intensities than to a conspecific communication sound, whose dominant frequency is 14 kHz (Fig. 7).     

Systematics and phylogeny of Parapholidoptera (Orthoptera: Tettigoniidae: Tettigoniinae)

Genus Parapholidoptera is revised and three species, P. yoruka sp.n., P. bolkarensis sp.n. and P. salmani sp.n., are described. Parapholidoptera delineata Stolyarov is placed in synonymy with P. ziganensis Karaba?. Parapholidoptera bodenheimeri Karaba? and P. distincta bodenheimeri Karaba? are placed in synonymy with P. distincta (Uvarov). Parapholidoptera flexuosa Karaba?, previously a subspecies of P. castaneoviridis (Brunner von Wattenwyl), is recognized as a separate species. A key to world species is provided. Cladistic analysis revealed the monophyly of the genus with two major clades. A short account of distribution is presented.     

Variation in nuptial gift quality in bush crickets (Orthoptera: Tettigoniidae)

Males of many insect species provide the female, during courtshipand copulation, with a nuptial gift consisting of a prey itemor synthesized material (e.g., spermatophores). The studiesperformed so far have mainly focused on effects of nuptial giftsize on male and female reproductive success. However, the qualityof the nuptial gift can differ substantially between taxa andmay potentially have a large impact on male and female reproductiveperformance. In this study the effects of. variation in dieton nuptial gift quality is investigated in several bush cricketspecies with different diets. The effect of diet on nuptialgift quality (e.g., protein in the spermatophylax) and femalereproductive output and, in turn, die effect of variation inspermatophylax quality on female reproductive output are investigated.Female reproductive output and male spermatophore size weremainly found to be influenced by differences in diet betweenspecies. Spermatophylax quality (i.e., protein concentration)was also correlated with differences in diet. There was a largevariation in protein content of the spermatophylax widiin aswell as between species. Moreover, larger spermatophylaxes hada lower protein concentration, indicating a possible trade-offbetween spermatophylax size and quality. Consequendy, productionof larger spermatophylaxes, required for protection of the male'ssperm carrying ampulla during insemination, can lead to a reducedprotein concentration, because the total amount of protein maybe limited. This pattern is also consistent with die idea diatthe spermatophylax functions primarily to ensure sperm transfer.Finally, there was no correlation between the amount of proteinin the spermatophylax and female reproductive output eitheracross diets or within each diet category, further supportingthe finding that female reproductive output is mainly affectedby differences in diet.     

Diet influences male-female interactions in the bushcricketRequena verticalis (Orthoptera: Tettigoniidae)

The influence of diet on the courtship roles of male and female Requena verticaliswas investigated in the laboratory. The protein content of available food was found to affect the frequency of mating attempts. Pairs which were fed on a low-protein diet were involved in fewer mating attempts than pairs which were fed on a high-protein diet. Diet also influenced the relative frequencies of male and female rejections. Males rejected their virgin female partners more often than females rejected their male partners when the pairs were kept on a low-protein diet. The opposite was found when the female had mated once before. No difference in the frequency of male and female rejections was found when the pair was kept on a high-protein diet irrespective of the mating status of the female.     

Dynamics of F-actin during spermiogenesis in Gampsocleis gratiosa ( Orthoptera: Tettigoniidae)

为阐明F-肌动蛋白在优雅蝈螽Gampsocleis gratiosa Brunner von Wattenwyl精子形成过程中的动态变化,本研究利用微分干涉相衬技术和免疫荧光技术首次对优雅蝈螽精子形成过程中的F-肌动蛋白进行了细胞定位,利用透射电镜技术从超微水平观察了优雅蝈螽精子顶体复合体的结构.结果显示:精子形成早期,F-肌动蛋白富集于亚顶体区域,形态由“球状”转变为“棒锥状”;精子形成中期,F-肌动蛋白呈“倒Y型”分布于亚顶体区域和细胞核前端两侧;精子形成后期,亚顶体区域的F-肌动蛋白解聚消失,F-肌动蛋白呈“箭头状”,仅分布于顶体复合体扩张的两翼中.F-肌动蛋白动态变化伴随着细胞核和精子头部的形态改变,F-肌动蛋白的动态装配在精子顶体复合体形态构建和细胞核的形变中起着重要的作用.本研究还发现未成熟的精子尾部有一些富含F-肌动蛋白的细胞质微滴,与精子形成过程中多余细胞质和细胞器的外排有关.F-肌动蛋白的动态变化研究为进一步阐明细胞骨架蛋白在昆虫精子形成过程中的功能和作用机制奠定了基础.     

Determinants of Male Spacing Behaviour in Panacanthus pallicornis (Orthoptera: Tettigoniidae)

Panacanthus pallicornis is a neotropical species of katydid endemic to Colombia that inhabits premontane forests, and individuals are found in regenerating (RF) and dense forest (DF) habits. Members of this species are made conspicuous by their colouration, remarkable defensive body thorns and the loud mating calls of the males. Through capture, marking, and recapture, we determined singing‐site characteristics in two different natural habitats: an RF and a DF. Using nearest neighbour analysis, we found male distribution tends to uniformity in forest habitats, but in the RF, male distribution is more random. Males also showed preference for high singing sites and these were correlated with host plant height. Although host plants in DF were taller than those in the RF, male’s perch preference was independent from the habitat; and in RF, males select lower perches. To investigate if male spacing was a function of the calling song, we manipulated two groups of males from a different population, one group deafened (tympanic membranes torn), and another control group with tympanic membranes intact. Insects were released from a single location in the two different zones of similar area. After a number of days we measured male displacement, and found that the deafened group distribution tended to aggregation, while the control group spread throughout the experimental zone with a random distribution. These results suggest that male spacing behaviour is one of the functions of the calling song. Based on a pre‐established phylogenetic framework of the genus Panacanthus, we discuss some implications for the evolution of the calling song of P. pallicornis in both RF and DF.     

The egg-laying habits of Euthypoda acutipennis Karsch (Orthoptera:Tettigoniidae)

The egg-laying habits of the West African bush-cricket (Euthypora acutipennis Karsch) are described. In addition, from a population of females bred in the laboratory, the longevity of the species has been determined.