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.     

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.     

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.     

Structure of a mosaic hybrid zone between the field crickets Gryllus firmus and G. pennsylvanicus

Hybrid zones provide insight into the nature of species boundaries and the evolution of barriers to gene exchange. Characterizing multiple regions within hybrid zones is essential for understanding both their history and current dynamics. Here, we describe a previously uncharacterized region of a well‐studied hybrid zone between two species of field crickets, Gryllus pennsylvanicus and G. firmus. We use a combination of mitochondrial DNA sequencing, morphological data, and modeling of environmental variables to identify the ecological factors structuring the hybrid zone and define patterns of hybridization and introgression. We find an association between species distribution and natural habitat; Gryllus pennsylvanicus occupies natural habitat along forest edges and natural clearings, whereas G. firmus occupies more disturbed areas in agricultural and suburban environments. Hybridization and introgression occur across patch boundaries; there is evidence of substantial admixture both in morphological characters and mtDNA, over a broad geographic area. Nonetheless, the distribution of morphological types is bimodal. Given that F₁ hybrids are viable and fertile in the lab, this suggests that strong pre‐zygotic barriers are operating in this portion of the hybrid zone.     

Wolbachia plays no role in the one-way reproductive incompatibility between the hybridizing field crickets Gryllus firmus and G. pennsylvanicus

Wolbachia are cytoplasmically inherited alpha-proteobacteria that can cause cytoplasmic incompatibility (CI) in insects. This incompatibility between sperm and egg is evident when uninfected females mate with infected males. Wolbachia-driven reproductive incompatibilities are of special interest because they may play a role in speciation. However, the presence of Wolbachia does not always imply incompatibility. The field crickets Gryllus firmus and G. pennsylvanicus exhibit a very clear unidirectional incompatibility and have been cited as a possible example of Wolbachia-induced CI. Here, we conduct curing experiments, intra- and interspecific crosses, cytological examination of Wolbachia in testes and Wolbachia quantifications through real-time PCR. All of our data strongly suggest that Wolbachia are not involved in the reproductive incompatibility between G. firmus and G. pennsylvanicus.     

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.     

A fine-scale spatial analysis of the mosaic hybrid zone between Gryllus firmus and Gryllus pennsylvanicus

Abstract.— The pattern of character variation within a hybrid zone, the hybrid zone structure, has been used to infer the processes that maintain hybrid zones. Unfortunately it is difficult to infer process from structure alone because many different processes can produce the same pattern of character variation. Mosaic hybrid zones may be maintained by exogenous selection in a heterogeneous environment and/or endogenous selection against hybrid individuals; habitat preference, premating isolating barriers and/or fertility selection can also contribute. The spatial scale at which a hybrid zone is sampled affects its apparent structure; a hybrid zone may appear clinal at one scale and mosaic at another. Here, we sample the mosaic hybrid zone between two field crickets, Gryllus firmus and G. pennsylvanicus , at a scale that spans the boundaries between individual soil-habitat patches. From our analysis, we find that at fine scales, the mosaic hybrid zone resolves into a set of steep clines across patch boundaries. Both morphological and molecular traits exhibit sharp and generally concordant clines. However, clines for mitochondrial DNA and one anonymous nuclear marker are clearly displaced as a result of current hybridization or past introgression (the "ghost of hybridization past"). Thus, scale is important for the structure of this and probably other hybrid zones. The extremely sharp, concordant clines across patch boundaries indicate that the cricket hybrid zone is undoubtedly structured by selection. However, the detailed mechanisms responsible for the maintenance of the hybrid zone–whether endogenous selection against hybrids, exogenous selection by the environment, and/or behavioral preferences for mates or habitats– remain to be elucidated. Determining these mechanisms will depend on closer inspection of the organisms themselves and their interactions, as is the case for all hybrid zones.     

Molecular evidence of a peripatric origin for two sympatric species of field crickets (Gryllus rubens and G. texensis) revealed from coalescent simulations and population genetic tests

Species pairs that differ primarily in characters involved in mating interactions and are largely sympatric raise intriguing questions about the mode of speciation. When species divergence is relatively recent, the footprint of the demographic history during speciation might be preserved and used to reconstruct the biogeography of species divergence. In this study, patterns of genetic variation were examined throughout the geographical range of two cryptic sister taxa of field crickets, Gryllus texensis and G. rubens; mitochondrial cytochrome oxidase I (COI) was sequenced in 365 individuals sampled from 48 localities. Despite significant molecular divergence between the species, they were not reciprocally monophyletic. We devised several analyses to statistically explore what historical processes might have given rise to this genealogical structure. The analyses indicated that the biogeographical pattern of genetic variation does not support a model of recent gene flow between species. Instead, coalescent simulations suggested that the genealogical structure within G. texensis, namely a deep split between two geographically overlapping clades, reflects historical substructure within G. texensis. Additional tests that consider the concentration of G. rubens haplotypes in one of the two G. texensis genetic clusters suggest a model of speciation in which G. rubens was derived from one lineage of a geographically subdivided ancestor. These results indicate that, despite the contemporary sympatry of G. texensis and G. rubens, the data are indicative of an peripatric origin in which G. rubens was derived from one of the two historical partitions in the species currently recognized as G. texensis. This proposed model of species divergence suggests how the interplay of geography and selection may give rise to new species, although this requires testing with multilocus data. Specifically, the model highlights how that geographical partitioning of ancestral variation in the past may augment the selectively driven divergence of characters involved in the reproductive isolation of the species today.     

Condition-dependent signaling affects male sexual attractiveness in field crickets, Gryllus campestris

The condition dependence of sexually selected traits is an importantassumption of sexual selection theory. Several laboratory studieshave documented a positive relationship between food availability,body condition, and sexual display. However, these studies mightnot reflect the resource allocation between body maintenance,reserves, and the sexually selected trait under natural conditions.Further, the effect of condition-dependent signaling on femalemate choice has hardly been investigated experimentally in thefield. We therefore investigated the effect of food availabilityon body condition, calling behavior, and sexual attractivenessof male field crickets, Gryllus campestris, under field conditions.Food availability was manipulated for individual males by supplementingfood in a confined area close to the burrow. Food-supplementedmales showed a significant increase in body condition, whereasthe opposite was found in the control males. Males receivingextra food called more frequently, whereas the calling-songcharacteristics were not affected by the treatment. Further,food-supplemented males attracted more females than did controlmales, and their higher attractiveness was partly explainedby their superior calling rate. Our study thus indicates condition-dependentsignaling as an important determinant of the sexual attractivenessof males to females under natural condition.     

Weak relationships of parasite infection with sexual and life‐history traits in wild‐caught Texas field crickets (Gryllus texensis)

1. Sex differences in immune investment and infection rate are predicted due to the divergent life histories of males and females, where females invest more toward immunity due to the fitness consequences of a reduced lifespan and males allocate less toward immunity due to increased resource investment in traits critical to sexual selection. Consequently, males are expected to fight infection less adeptly, resulting in higher parasite loads relative to females across all taxa.
2. Wild animals rarely face a single parasite within their given environment, yet nearly all studies on sex‐biased infection rates have focused on a single host–parasite relationship. Here, we investigate how simultaneous natural infections of ecto‐ and endosymbionts (i.e. both parasitic and phoretic taxa) correlate with sex biases in host immune response and reproductive investment in a field‐caught cricket, Gryllus texensis.
3. Our comprehensive analysis found no significant sex differences in two measures of immune response (melanization and nodulation), and found no strong evidence of a sex bias in the prevalence or intensity of parasitism by the three most common parasites infecting wild G. texensis field crickets (Eutrombidiidae, gregarines, and nematodes).
4. Two traits related to female fitness, egg number and egg size, showed no relation to parasitic infection; however, males having wider heads and poorer body condition were significantly more infected by eutrombidiid mites, gregarines, and nematodes.
5. Despite frequent predictions of male‐biased parasitism in the literature, our results concur with many other studies indicating that the divergent life histories of males and females alone are not sufficient to explain natural infection rates in wild insects.

     

Production of chitin and chitosan from the exoskeleton of adult two‐spotted field crickets (Gryllus bimaculatus)

Chitin and chitosan were extracted from all specimens of Type I and II two‐spotted field crickets (Gryllus bimaculatus) following chemical treatment with an acid and alkali. For chitin extraction, 2 N HCl and 1.25 N NaOH solutions were used to achieve demineralization and deproteinization, respectively. For chitosan extraction, 50 % NaOH (w/v) and 50 % NaOH (w/w) solutions were used to achieve deacetylation. Chitosan yielded from adult exoskeletons of G. bimaculatus in Test A of Type I was 1.76 and 8.40 % on a fresh weight (FW) and dry weight (DW) basis, respectively, after treatment with 50 % NaOH (w/v) at 95°C for 3 h. Furthermore, the chitosan yielded in Test D of Type II was 1.79 and 7.06 % on FW and DW basis, respectively, after treatment with 50 % NaOH (w/w) at 105°C for 3 h. The average yield of chitin and chitosan was 2.42 and 1.65 % on a FW basis, and 10.91 and 7.50 % on DW basis, respectively. The deacetylation (%) of chitosan extracted from adult exoskeletons in Tests A, B, C1, C2, D1, and D2 were 81.2 %, 14.5 %, 19.6 %, 90.7 %, 17.1 %, and 95.5 %, respectively. The viscosities of the chitosans extracted from adult exoskeletons in Tests A, C2, and D2 were 32.0, 21.6, and 62.4 cP (centi Poise), respectively. The molecular weight of chitosan from adult exoskeletons of G. bimaculatus was 308.3 kDa. Our results indicate that adult exoskeletons of G. bimaculatus could be used as a source of chitin and chitosan for use as functional additives in industrial animal feeds.     

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.     

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.     

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.

     

Seasonal and interindividual variation in field water metabolism of female meadow voles Microtus pennsylvanicus.

We analyzed variations in water flux rates on a large sample of meadow voles (Microtus pennsylvanicus) to quantify the effect of season on water metabolism of individuals and to examine patterns of intra- and interindividual variability. Voles were nonreproductive females maintained in outdoor enclosures where they fed on natural vegetation. They were injected one to three times with doubly labeled water, which resulted in one to six measures of daily water flux rate per individual. Summer water flux rates of voles were 258% of the predicted values for herbivorous eutherian mammals of similar size. To date, very few studies have focused on mammals with such high water flux rates. Body water volume of individuals was higher in summer than in winter (75.6% vs. 72.5%), and water flux rate of animals was 12.5% higher in the winter season (0.99 vs. 0.88 mL H2O g-1 d-1). Between-season differences in water fluxes were proportional to differences in energy expenditures, hence the water economy index remained constant across seasons (0.30 mL H2O kJ-1). Intraindividual variability of water flux rate was high compared to interindividual variability (repeatability, r<0.30), which will make it difficult to study natural selection of water metabolism in a microevolutionary framework, at least in meadow voles.     

Death comes suddenly to the unprepared: singing crickets, call fragmentation, and parasitoid flies

Male field crickets are subject to a delicate dilemma becausetheir songs simultaneously attract mates and acoustic predators.It has been suggested that in response, crickets have modifiedvarious temporal song parameters to become less attractiveto acoustic predators. We investigated whether crickets withchirping (versus trilling) song structures are less likely toattract acoustically orienting parasitoid flies. Experimentally,we evaluated the phonotactic quest of the parasitoid fly Ormiaochracea in response to broadcast cricket calls, presentedboth simultaneously (choice paradigm) and sequentially (no-choiceparadigm). Flight trajectories were recorded in darkness usingthree-dimensional active infrared video tracking. The flies showed remarkable phonotactic accuracy by landing directly onthe loudspeaker. The introduction of acoustic fragmentationthat resembles calls of many chirping crickets altered theflies' phonotactic accuracy only slightly. Our results documentdifferential attraction between trilling and chirping cricket songs and quantitatively demonstrate that chirping songs, ifpresented alone, do not impair the efficiency (temporal investmentand landing accuracy) of the flies' phonotactic quest. Thisstudy shows that song fragmentation is no safeguard againstacoustic parasitism. We conclude that, in general, a cricket may reduce predation only if its neighbors are acousticallymore conspicuous, chiefly by amplitude.     

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.     

Karyotypes of two American field crickets: Gryllus rubens and Gryllus sp. (Orthoptera: Gryllidae)

The karyotypes of Gryllus rubens (Scudder) and Gryllus sp. collected in the US were studied using conventional Giemsa staining and two differential staining methods. Both species had a chromosome complement of 2n = 28 + XX/XO, and the X chromosome was large and metacentric. In addition, nucleolus organizer regions (NOR) were detected in the short arm of one pair in the two species, and the NOR showed variation in size. The two species had species‐specific chromosome configuration and C‐banding patterns. In Gryllus sp., the chromosome configuration showed polymorphism in size and type among individuals, and the distal C‐bands of Gryllus sp. were larger than those of G. rubens.