Arne W. Lehmann

Spermatophore characteristics in bushcrickets vary with parasitism and remating interval

Abstract Male bushcrickets provide females with a nuptial gift, a spermatophore, which is transferred to females at mating. The spermatophore consists of a gelatinous mass, the spermatophylax, and the sperm-containing ampulla. Male spermatophore size is positively correlated with insemination rate and female refractory period and therefore with male reproductive success. In this study, we examined spermatophylax weight, ampulla weight and sperm number in males of Poecilimon mariannae parasitized by the acoustically orienting fly Therobia leonidei. We show that in parasitized males, spermatophylax weight decreases with the level of parasitism. In line with the hypothesis that parasitism is a cost to reproduction, we found that spermatophylax weight was reduced at remating. In contrast, the replenishment of the spermatophylax in unparasitized males was complete after 2 days and was increased no further after 3 days. Both sperm number and ampulla weight showed an increase over time since last mating and sperm production was estimated at a constant rate of 500,000 per day in all individuals, regardless of parasitism. The allocation of investment in components of the spermatophore varies greatly with parasitism and remating. Both factors had rather independent effects on spermatophore constitution, revealing functional constraints acting on spermatophore characteristics in bushcrickets, which are important for understanding the selection pressures working on its components.

Bushcricket song as a clue for spermatophore size

Bushcricket males of Poecilimon zimmeri transfer large and protein-rich spermatophores during mating, which females directly ingest. There is correlational evidence that heavier males transfer larger nuptial gifts. In no-choice mating trials, females mated randomly with respect to male’s body weight. In contrast, in two-choice mating trials, female bushcrickets exhibit clear choice for the heavier male. This heavier male advantage was also found in pre-mating choice during phonotaxis. With manipulated mute males, females mated at random with regard to body weight of the competitors. The number of physical encounters between a female and males was low in all tests with a single male (no choice) and greater in choice-tests with two competing males. The latencies to mate also differed significantly between treatments. The time mating pairs spent in precopula was short in experiments where the males could hear rivals and significantly longer in all other tests using either a single male or mute males. Thus, acoustic signalling in male bushcrickets seems to signal male body weight. A preference for heavier males may reflect a female’s preference for a larger spermatophore and therefore a greater direct benefit.

Variation in body size among populations of the bushcricket Poecilimon thessalicus (Orthoptera: Phaneropteridae): an ecological adaptation

Abstract We analyzed geographical variation in body size in males and females of nine Greek populations of the bushcricket Poecilimon thessalicus Brunner von Wattenwyl, 1891. We found significant geographic variability in P. thessalicus, with all three morphometric body-size characters (hind femur, front tibia, and pronotum) highly correlated within populations. Populations differed in mean size between three mountain ranges, and were larger on moist eastern, than on dry western slopes. We suggest that the observed smaller size of the bushcrickets on western slopes is mainly the consequence of a shorter growing season due to summer drying on western slopes. Sex-specific growth rates might contribute to the observed smaller body size in males in all populations. Renschs rule that sexual size dimorphism (SSD) decreases as body size increases is supported, with males growing relatively larger compared to females in populations with larger body sizes. This pattern might be sexually selected, as males that produce larger nuptial gifts are favored as mates in Poecilimon bushcrickets.

Female bushcrickets mated with parasitized males show rapid remating and reduced fecundity (Orthoptera: Phaneropteridae: Poecilimon mariannae)

Abstract Following mating, female bushcrickets undergo a refractory period during which they are sexually unreceptive. The length of the refractory period correlates with the size of the spermatophylax. However, the size of the nuptial gift of acoustically signalling bushcrickets is often reduced as a result of infections by parasitoid flies. We examined the effect of male parasitoid infection on the induction of the refractory period and fecundity of females. We found a drastically reduced refractory period in females if the first mating partner was infected. During this shortened period fewer eggs were deposited, as an effect of the shorter refractory period, whereas the daily egg-laying rate remained the same regardless of whether the females were mated with a parasitized or an unparasitized male.

Spatial organization of tettigoniid auditory receptors: Insights from neuronal tracing

The auditory sense organ of Tettigoniidae (Insecta, Orthoptera) is located in the foreleg tibia and consists of scolopidial sensilla which form a row termed crista acustica. The crista acustica is associated with the tympana and the auditory trachea. This ear is a highly ordered, tonotopic sensory system. As the neuroanatomy of the crista acustica has been documented for several species, the most distal somata and dendrites of receptor neurons have occasionally been described as forming an alternating or double row. We investigate the spatial arrangement of receptor cell bodies and dendrites by retrograde tracing with cobalt chloride solution. In six tettigoniid species studied, distal receptor neurons are consistently arranged in double‐rows of somata rather than a linear sequence. This arrangement of neurons is shown to affect 30–50% of the overall auditory receptors. No strict correlation of somata positions between the anterio‐posterior and dorso‐ventral axis was evident within the distal crista acustica. Dendrites of distal receptors occasionally also occur in a double row or are even massed without clear order. Thus, a substantial part of auditory receptors can deviate from a strictly straight organization into a more complex morphology. The linear organization of dendrites is not a morphological criterion that allows hearing organs to be distinguished from nonhearing sense organs serially homologous to ears in all species. Both the crowded arrangement of receptor somata and dendrites may result from functional constraints relating to frequency discrimination, or from developmental constraints of auditory morphogenesis in postembryonic development. J. Morphol.

Potential lifetime reproductive success of male bushcrickets parasitized by a phonotactic fly

Males of the bushcricket Poecilimon mariannae are parasitized by the acoustically orienting fly Therobia leonidei. Developing fly larvae reduce male bushcricket survival and reproductive effort. We estimated potential lifetime reproductive success (PLRS) of male bushcrickets to investigate the likely costs of this parasitism. Our analysis explored the relative reduction in PLRS caused by parasitoid infection by examining effects on survival, calling attractiveness and spermatophylax size. We also incorporated nonparasitoid-related mortality. Parasitized males lost 42% of their PLRS compared to nonparasitized males. The shortened life span after parasitism accounted for half of the PLRS reduction. Decreased spermatophylax production also had a major effect on PLRS, while decreased calling attractiveness had a smaller effect. This fitness loss was context dependent and changed with extrinsic nonparasitoid-related mortality. If nonparasitoid-related mortality was high, the selection on host males to avoid parasitism was weak; if it was low, parasitized males lost a considerable amount of reproductive opportunities compared with unparasitized males.

Increased copulation duration before ejaculate transfer is associated with larger spermatophores, and male genital titillators, across bushcricket taxa

Copulation duration varies considerably across species, but few comparative studies have examined factors that might underlie such variation. We examined the relationship between copulation duration (prior to spermatophore transfer), the complexity of titillators (sclerotized male genital contact structures), spermatophore mass and male body mass across 54 species of bushcricket. Using phylogenetic comparative analyses, we found that copulation duration was much longer in species with titillators than those without, but it was not longer in species with complex compared with simple titillators. A positive relationship was found between spermatophore size and copulation duration prior to ejaculate transfer, which supports the hypothesis that this represents a period of mate assessment. The slope of this relationship was steeper in species with simple rather than complex titillators. Although the data suggest that the presence of titillators is necessary to maintain long copulation prior to ejaculate transfer, mechanisms underlying this association remain unclear.

Changes in the numbers of chromosomes and sex determination system in bushcrickets of the genus Odontura (Orthoptera: Tettigoniidae: Phaneropterinae)

Chromosomes of the males of five species of Odontura, belonging to the subgenera Odontura and Odonturella, were ana- lyzed. Intensive evolution of the karyotype was recorded, both in terms of changes in the numbers of chromosomes (from 2n = 31 to 27) and the sex chromosome system (from X0 to neo-XY and X0 to neo-X1X2Y). Karyotype evolution was accompanied by tandem autosome fusions and interspecific autosomal and sex chromosome differentiation involving changes in the locations of nucleolar organizer regions, NORs, which were revealed by silver impregnation and confirmed by FISH using an 18S rDNA probe. O. (Odon- turella) aspericauda is a polytypic species with X0 and neo-X1X2Y sex determination. The latter system is not common in tettigo- niids. It possibly originated by a translocation of a distal segment of the original X chromosome onto a medium sized autosome, resulting in a shortened neo-X1 and a metacentric neo-Y. The remaining autosome homologue became the neo-X2 chromosome. This shift from X0 to neo-X1X2Y is supported by the length of the X chromosome and location of the NOR/rDNA.

Life-history trade-off between macroptery and reproduction in the wing-dimorphic pygmy grasshopper Tetrix subulata (Orthoptera Tetrigidae)

Dimorphism in wing length is well known in many insect species. In those species a trade-off exists between dispersal capability and reproduction, with the long-winged (LW) morph being a better disperser due to superior flight capability. The short-winged (SW) morph is less mobile and it is hypothesised that females of this morph can invest more energy reserves into offspring production. Morphological examination supports the occurrence of two clearly distinct wing morphs in both sexes of the pygmy grasshopper Tetrix subulata (Orthoptera Tetrigidae). SW females produced larger eggs, in accordance with the reproductive compensation hypothesis. Male wing morph strongly influenced mating duration; mating of LW males with LW females lasted less than 1 min while those of SW males with SW females took around 19 min. Female wing length had little influence on the mating duration. Our results support the trade-off between wing development and reproduction.

Sensory evolution of hearing in tettigoniids with differing communication systems.

In Tettigoniidae (Orthoptera: Ensifera), hearing organs are essential in mate detection. Male tettigoniids usually produce calling songs by tegminal stridulation, whereas females approach the males phonotactically. This unidirectional communication system is the most common one among tettigoniids. In several tettigoniid lineages, females have evolved acoustic replies to the male calling song which constitutes a bidirectional communication system. The genus Poecilimon (Tettigoniidae: Phaneropterinae) is of special interest because the ancestral state of bidirectional communication, with calling males and responding females, has been reversed repeatedly to unidirectional communication. Acoustic communication is mediated by hearing organs that are adapted to the conspecific signals. Therefore, we analyse the auditory system in the Tettigoniidae genus Poecilimon for functional adaptations in three characteristics: (i) dimension of sound‐receiving structures (tympanum and acoustic spiracle), (ii) number of auditory sensilla and (iii) hearing sensitivity. Profound differences in the auditory system correlate with uni‐ or bidirectional communication. Among the sound‐receiving structures, the tympana scale with body size, whereas the acoustic spiracle, the major sound input structure, was drastically reduced in unidirectional communicating species. In the unidirectional P. ampliatus group, auditory sensilla are severely reduced in numbers, but not in the unidirectional P. propinquus group. Within the P. ampliatus group, the number of auditory sensilla is further reduced in P. intermedius which lost acoustic signalling due to parthenogenesis. The auditory sensitivity correlated with the size of the acoustic spiracle, as hearing sensitivity was better with larger spiracles, especially in the ultrasonic range. Our results show a significant reduction in auditory structures, shaped by the differing sex roles during mate detection.