Calling song and selective phonotaxis in the field crickets,Gryllus firmus andG. pennsylvanicus (Orthoptera: Gryllidae)

The field cricket species, Gryllus firmusand G. pennsylvanicus,occur in a mosaic hybrid zone that roughly parallels the eastern slope of the Appalachian mountains in the northeastern United States. It is important to know what role, if any, the calling song plays in mate choice in sympatric and allopatric populations. In this report, we present results on the variability of calling song properties along transects across this hybrid zone. We also present the results of experiments on phonotactic selectivity of females from an allopatric population of G. firmus.The male calling song of allopatric G. firmuswas significantly slower in temporal rhythm (i. e., chirp and pulse repetition rates) and lower in pitch (i.e., dominant frequency) than that of allopatric G. pennsylvanicus.Calling song properties of males recorded in the hybrid zone varied considerably in temporal and spectral properties. In two-stimulus (choice) phonotaxis experiments, allopatric females of G. firmuspreferred synthetic calling songs with conspecific pulse repetition rates over songs that had lower and higher pulse rates. This preference persisted even when the sound pressure levels of alternative stimuli were unequal. Therefore, allopatric females of G. firmuscan discriminate between conspecific and heterospecific calling songs. Whether or not this same selectivity is present in sympatric populations remains unclear. Investigations of phonotactic selectivity in other allopatric and sympatric populations of both species are currently under way.     

Seasonal and individual variation in gregarine parasite levels in the field crickets Gryllus veletis and G.pennsylvanicus

ABSTRACT. 1. Gregarines, debilitating protozoan gut parasites, were monitored in two species of field cricket, Gryllus veletis (Alexander and Bigelow) and G.pennsylvanicus Burmeister, in northern Michigan in 1983 and 1984.
2. Gregarines were found in 31–51% of G.veletis individuals and 50–70% of G.pennsylvanicus.
3. Males were more heavily infected than females in G. veleris , but G.pennsylvanicus males and females had nearly equal infection levels; this disparity resulted in an overall lower infection rate for G. veletis.
4. Both species had fewer gregarines in 1983 than in 1984, possibly due to the adverse effect of drier conditions in 1983 on development of parasite cysts in the soil.
5. Levels of infection were not constant throughout the breeding season for either cricket species.
6. Body size was not correlated with parasite load in either G. veletis or G.pennsylvanicus.
7. Adult age was unrelated to gregarine level in G. veferis , whereas G.pennsylvanicus showed a positive correlation between adult age and parasite load during two non-consecutive weeks of the breeding season.     

An analysis of G matrix variation in two closely related cricket species,Gryllus firmus and G. pennsylvanicus

An important issue in evolutionary biology is understanding the pattern of G matrix variation in natural populations. We estimated four G matrices based on the morphological traits of two cricket species, Gryllus firmus and G. pennsylvanicus, each reared in two environments. We used three matrix comparison approaches, including the Flury hierarchy, to improve our ability to perceive all aspects of matrix variation. Our results demonstrate that different methods perceive different aspects of the matrices, which suggests that, until more is known about these methods, future studies should use several different statistical approaches. We also found that the differences in G matrices within a species can be larger than the differences between species. We conclude that the expression of the genetic architecture can vary with the environment and that future studies should compare G matrices across several environments. We also conclude that G matrices can be conserved at the level of closely related species.     

Biosynthesis of 2-methylalkanes in the crickets Nemobius fasciatus and Gryllus pennsylvanicus.

The biosynthesis of 2-methylalkanes was studied in the crickets $\text{Nemobius}\text{fasciatus}$ and $\text{Gryllus}\text{pennsylvanicus}$. Labelled acetate, valine, and isobutyric acid were incorporated into the cuticular hydrocarbon of $\text{N.}\text{fasciatus}$ at levels of 6.0 ± 1, 6.5 ± 2, and 1.5 ± 0.7 percent respectively. The hydrocarbons of this insect are 20 percent 2-methylalkanes, primarily of even numbered carbon chain lengths, and 80% n-alkanes. Of the label incorporated into the hydrocarbon fraction, 28 ± 2 percent of sodium [1-¹⁴C] acetate, 98 ± 1 percent of L-[G-³H] valine, and 75 ± 10 percent of [1-¹⁴C] isobutyric acid were incorporated into the 2-methylalkanes. This suggests that valine is converted to isobutyric acid and is incorporated into the even numbered carbon chain length 2-methylalkanes during the initial stages of chain elongation. Similar data obtained in $\text{G.}\text{pennsylvanicus}$ suggests that leucine is converted to isovaleric acid which is then incorporated into the odd numbered carbon chain length 2-methylalkanes.     

Sexual selection at varying population densities in male field crickets,Gryllus veletis andG. pennsylvanicus

Male field crickets call and attract females or they silently search for females in the vicinity of calling males. At high population densities, fewer calling sites are available, defense of calling sites is costly, and an increased proportion of matings should result from searching behavior. To test these predictions, individually marked field crickets, Gryllus veletisand G. pennsylvanicus,were observed for 10 h nightly in large outdoor arenas at relatively high and low densities (2020 and 55, males and females). Data were gathered on body weight, calling duration, movement, and mating frequency for individual crickets. These observations showed that variance in male mating success was significantly greater at a low density in G. pennsylvanicus,and calling duration correlated with mating success at this density. Direct selection on a trait was estimated as the partial regression coefficient (selection gradient, ) and the total selection was estimated (direct and indirect selection on correlated traits) as the covariance (standardized intensity of selection, s) of the trait on the relative mating success. Direct selection favored increased movement at a high density in G. veletis,and direct and total selection favored increased calling duration at a low density in G. pennsylvanicus.Most other comparisons were not statistically significant. The data are discussed in terms of density-dependent fluctuations in sexual selection on correlated male traits.     

The effects of gregarine parasites on longevity, weight loss, fecundity and developmental time in the field crickets Gryllus veletis and G.pennsylvanicus

ABSTRACT. 1. Gregarines, debilitating gut parasites, significantly reduced longevity and increased weight loss in the field cricket Gryllus pennsylvanicus Burmeiser when adults were kept on suboptimal diets, but in G.pennsylvanicus and G.veletis (Alexander and Bigelow) fed ad libitum , weight change, longevity, and fecundity were not significantly different in infected and control individuals.
2. Adults harbouring gregarines at eclosion took significantly longer to develop from the second and third instars than did uninfected adults.
3. Longevity of nymphs kept in the laboratory without food was decreased in infected individuals compared with uninfected ones, but not significantly so.
4. The evolutionary and ecological implications of nonlethal parasites are discussed.     

Flight polymorphism in the field cricket Gryllus pennsylvanicus

Summary Wing length polymorphisms are common in many groups of insects. In the field cricket Gryllus pennsylvanicus the long-winged morph occurs in low frequency in most populations but seems not to occur at all in certain areas and to become relatively abundant in others. Laboratory rearing experiments and single-pair crosses suggest that both inter-and intrapopulation variation can be explained at least in part by genetic differences. A model of genetic variation in threshold response is consistent with all available data. One possible mechanism for maintenance of the flight polymorphism is discussed.     

The contribution of multiple mating and spermatophore consumption to the lifetime reproductive success of female field crickets (Gryllus bimaculatus)

ABSTRACT.

• 1 The adaptive significance of multiple mating by female Gryllus bimaculatus (De Geer) was investigated.
• 2 Multiple mating prevented the depletion of sperm stores and, therefore, maintained high hatching success. This may not, however, explain the high frequency of remating in this species.
• 3 Male-derived egg stimulants known to be passed with sperm at mating increased the number of eggs produced only when females mated throughout their lifespans.
• 4 Spermatophore consumption appeared to provide nutrients which, while they did not increase the quantity of eggs, increased egg quality as indicated by weight. Females who consumed spermatophores had a greater hatching success.
• 5 While females may derive non-genetic benefits from mating, these are apparently long-term benefits; females must mate throughout their lives in order to accrue them.
• 6 Since the benefits of mating may not be derived from individual males, the spermatophores and their contents in this species are best considered as mating effort.

     

Reproductive Success and Body Size in the Cricket Gryllus firmus

Male body size influences mate choice and sexual selection in many animal species. Here we investigate the role of male body size in the reproductive success of the field cricket Gryllus firmus. This species hybridizes with a close smaller relative, G. pennsylvanicus, and it is thought that this size difference may affect reproductive isolation between these species. We paired large and small G. firmus males with a single G. firmus female and genotyped the resulting offspring. Overall, larger males sired a greater proportion of offspring and in a majority of the crosses the larger male sired all of the offspring. For crosses in which both males sired offspring, there was no difference in the proportion of offspring sired by small and large males. Intrasexual competition, female choice, and differences in ejaculates between males could all influence the patterns we observe. We discuss the implications of our findings within the context of reproductive isolation between G. firmus and G. pennsylvanicus.     

Hormones in the field: evolutionary endocrinology of juvenile hormone and ecdysteroids in field populations of the wing-dimorphic cricket Gryllus firmus

Virtually no published information exists on insect endocrine traits in natural populations, which limits our understanding of endocrine microevolution. We characterized the hemolymph titers of juvenile hormone (JH) and ecdysteroids (ECDs), two key insect hormones, in field-collected short-winged, flightless (SW) and long-winged, flight-capable (LW(f)) morphs of the cricket Gryllus firmus. The JH titer exhibited a dramatic circadian rhythm in the LW(f) morph but was temporally constant in the flightless SW morph. This pattern was consistent in each of three years; in young, middle-aged, and older G. firmus; and in three other cricket species. The ECD titer was considerably higher in SW than in LW(f) females but did not exhibit temporal variation in any morph and did not differ between male morphs. JH and ECD may control different aspects of the morph-specific trade-off between nocturnal dispersal and reproduction. Results confirm and extend laboratory studies on young female G. firmus; most, but not all, important aspects of morph-specific differences in JH and ECD titers can be extrapolated from field to laboratory environments and vice versa. Hormone titers in Gryllus are more complex than those proposed in evolutionary endocrine models. Directly measuring hormone titer variation remains a fundamentally important task of insect evolutionary endocrinology.     

Extra-nuclear effects on growth and development in the sand cricket Gryllus firmus

Although largely ignored until recently, parental effects on the phenotypes of their offspring are both ubiquitous in nature and of a potentially great importance to evolution. Our study examines the presence of extra-nuclear (maternal and paternal) effects in growth traits, development time and adult size in the sand cricket Gryllus firmus using a diallel cross of inbred lines. Sex linkage was shown to be nonsignificant for development time but the other traits could not be tested. We assume that they are nonsignificant but use the term 'reciprocal' effects to include this effect. We show that reciprocal effects are present in the growth traits and development time, where they account for 10-30% of the phenotypic variance. They are not present in adult size as indexed by head width. We demonstrate that reciprocal effects are due, at least in part, to maternal effects by an analysis of the positive correlation between egg size, a maternal trait, and the growth traits. The growth rate traits show no significant decline with age either with respect to extra-nuclear contributions to variance or difference between phenotypic means of reciprocal pairs. This study demonstrates that extra-nuclear effects are important contributors to the phenotypic variation in life history traits of G. firmus.     

Juvenile hormone titer and morph-specific reproduction in the wing-polymorphic cricket,Gryllus firmus

Juvenile hormone titers and reproductive characteristics were measured in adult wing and flight-muscle morphs of the wing-polymorphic cricket, Gryllus firmus, during the first week of adulthood. This species has three morphs: one flight capable morph with fully-developed wings and fully-developed flight muscles [LW(F)], one flightless morph with fully-developed wings and histolyzed (non-functional) flight muscles [LW(H)], and another flightless morph with underdeveloped (short) wings and underdeveloped flight muscles (SW). Both flightless morphs [LW(H) and SW] had larger ovaries which contained a greater number of postvitellogenic eggs compared with the flight capable [LW(F)] morph. The juvenile hormone titer was significantly higher in SW compared with LW(F) females on days 3-7 of adulthood. On these days, the JH titer also was significantly higher in the other flightless morph, LW(H), compared with flight-capable [LW(F)] females as determined by one statistical test, but did not differ significantly by another test. The JH titer was positively correlated with ovarian mass or terminal oocyte length, but not with the number of post-vitellogenic eggs. This study is the first direct comparison of juvenile hormone titers in adult wing morphs of a wing-polymorphic insect. Results indicate that an elevated juvenile hormone titer may be at least partly responsible for one of the most distinctive features of wing-polymorphic species, the increased early fecundity of flightless females.     

Development of sex and protective behavior of female crickets Gryllus argentinus Sauss. at preproductive and reproductive periods of adult ontogenesis

The work presents data on reproductive and protective behavior of non-copulative female crickets Gryllus argentinus Sauss. at the beginning of adult ontogenesis. There is studied character of interaction of females with "virtual" males (the method of "trackball". The forms of female motor reactions are described and characterized in the model of male intraspecies signals--the call and aggressive ones. The character of development of female acoustic behavior is described by a change in the motor reaction forms in the course of adult ontogenesis. It is shown that at once after moulting the reproductive behavior on adults has not yet been formed, but is maturing gradually for several days. The passive protective behavior was established to be the most expressed at the initial stages of adult ontogenesis.     

Temperature affects the tripartite interactions between bacteriophage WO, Wolbachia, and cytoplasmic incompatibility

Wolbachia infections are a model for understanding intracellular, bacterial symbioses. While the symbiosis is often studied from a binary perspective of host and bacteria, it is increasingly apparent that additional trophic levels can influence the symbiosis. For example, Wolbachia in arthropods harbor a widespread temperate bacteriophage, termed WO, that forms virions and rampantly transfers between coinfections. Here we test the hypothesis that temperatures at the extreme edges of an insect's habitable range alter bacteriophage WO inducibility and in turn, Wolbachia densities and the penetrance of cytoplasmic incompatibility. We report four key findings using the model wasp, Nasonia vitripennis: First, both cold treatment at 18 C and heat treatment at 30 C reduce Wolbachia densities by as much as 74% relative to wasps reared at 25 C. Second, in all cases where Wolbachia densities decline due to temperature changes, phage WO densities increase and inversely associate with Wolbachia densities. Heat has a marked effect on phage WO, yielding phage densities that are 552% higher than the room temperature control. Third, there is a significant affect of insect family on phage WO and endoysmbiont densities. Fourth, at extreme temperatures, there was a temperature-mediated adjustment to the density threshold at which Wolbachia cause complete cytoplasmic incompatibility. Taken together, these results demonstrate that temperature simultaneously affects phage WO densities, endosymbiont densities, and the penetrance of cytoplasmic incompatibility. While temperature shock enhances bacteriophage inducibility and the ensuing bacterial mortality in a wide range of medically and industrially-important bacteria, this is the first investigation of the associations in an obligate intracellular bacteria. Implications to a SOS global sensing feedback mechanism in Wolbachia are discussed.     

Wolbachia and nuclear-nuclear interactions contribute to reproductive incompatibility in the spider mite Panonychus mori (Acari: Tetranychidae)

Maternally transmitted bacteria of the genus Wolbachia are obligate, intracellular symbionts that are responsible for cytoplasmic incompatibility in a wide range of arthropods such as insects and mites. Spider mites often show uni- and bidirectional incompatibilities among populations with and without Wolbachia. Therefore, we surveyed the presence of Wolbachia by PCR and then conducted crossing experiments among 25 populations of Panonychus mori to determine how Wolbachia are related to the incompatibility in this species. Five out of the 25 populations were infected with Wolbachia. These five populations were treated with an antibiotic (rifampicin) to eliminate Wolbachia. We carried out round-robin crosses among 20 Wolbachia-uninfected populations, five infected populations and five rifampicin-treated populations (30 x 30=900 crosses in total). Incompatibility among P. mori populations was caused by Wolbachia infection, nuclear-cytoplasmic interactions or nuclear-nuclear interactions. Wolbachia-mediated incompatibility was observed in crosses between uninfected females and infected males or between females and males harboring different Wolbachia strains. Nuclear-cytoplasmic interactions may be responsible for the unidirectional incompatibility in crosses between the two northern populations and one of the southern populations. Bidirectional incompatibility caused by nuclear-nuclear interactions was observed in 99 combinations of interpopulation crosses (99/300=0.33). Although no geographical trends were detected in the distribution of bidirectionally compatible populations, the results reveal a genetic divergence among P. mori populations.     

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.     

Female mating behavior in the field cricket,Gryllus pennsylvanicus (Orthoptera: Gryllidae) at different operational sex ratios

The influence of operational sex ratio on the mating behavior of female field crickets,Gryllus pennsylvanicus, was investigated. Females were predicted to be more discriminating under conditions of high mate availability and show less selectivity when males were rare. Such selectivity was indicated in this study with the proportion of courtships leading to a mating changing with sex ratio. Females accepted almost 70% of all courtships at the female-biased sex ratio, but only about half of all courtships were successful at even or male-biased sex ratios. Females moved least at the female-biased sex ratio. There was also a trend for females to be guarded more under male-biased conditions. Female weight did not influence any of the behaviors examined.     

Courtship and copulation, but not ejaculates, reduce the longevity of female field crickets (Gryllus bimaculatus)

Females of many taxa incur fitness costs from male sexual coercion and harassment leading to mating. Although male crickets cannot force copulations on females, female Gryllus bimaculatus in this study incurred significant reductions in longevity through being exposed to different levels of male courtship. Virgin females kept in isolation had the longest life spans. Reductions in longevity applied to females in sensory contact with males (without the opportunity to mate), females that courted and mated and females that mated but with fertilization being prevented. Females also incurred significant reductions in longevity when kept with other females, which may have been due to high levels of cannibalism. Consistent with previous studies, females appeared to incur no cost to longevity from receiving sperm or seminal fluid. It is known that female G. bimaculatus benefit genetically from multiple mating. However, this benefit could possibly be offset by the negative effect that male courtship and mating behaviour has on female longevity.