Acoustic communication in the trilling field cricket,Gryllus rubens (Orthoptera: Gryllidae)

Male field crickets produce calling songs that are used for attracting conspecific females for mating. Acoustic communication was studied in the trilling field cricket, Gryllus rubens.A trilled calling song consists of a continuous train of sound pulses, each of which was repeated at a stereotyped rate. Singing males were recorded at different temperatures. The pulse period of the calling song decreased with increasing temperature; there was less effect of temperature on pulse duration. Female phonotaxis was studied on a noncompensating, spherical treadmill. In two-stimulus (choice) playback experiments, females preferred the conspecific trilled song over the chirped calling song of a sympatric species, Gryllus fultoni.This preference persisted even when the song of G. fultoniwas 6 dB louder. Females also discriminated between synthetic trills having different pulse periods; females chose trills with the conspecific pulse period over trills having lower and higher pulse periods.     

The characteristics of karyotype and telomeric satellite DNA sequences in the cricket, Gryllus bimaculatus (Orthoptera, Gryllidae)

The chromosomes derived from the Japanese population of Gryllus bimaculatus were characterized by C-banding and Ag-NOR staining. The chromosome number, 2n = 28 + XX (female)/XO (male), corresponded with that of other populations of G. bimaculatus, but the chromosome configuration in idiograms varied between the populations. NORs were carried on one pair of autosomes and appeared polymorphous. The positive C-bands located at the centromere of all chromosomes and the distal regions of many chromosome pairs, and the size and the distribution pattern of the distal C-heterochromatin showed differences among the chromosomes. In addition, this paper reports on the characteristics of HindIII satellite DNA isolated from the genome of G. bimaculatus. The HindIII repetitive fragments were about 0.54 kb long, and localized at the distal C-bands of the autosomes and the interstitial C-bands of the X chromosome. Molecular analysis showed two distinct satellite DNA sequences, named the GBH535 and GBH542 families, with high AT contents of about 67 and 66%, respectively. The two repetitive families seem to be derived from a common ancestral sequence, and both families possessed the same 13-bp palindrome sequence. The results of Southern blot hybridization suggest that the sequence of the GBH535 family is conserved in the genomic DNAs of Gryllus species, whereas the GBH542 family is a species-specific sequence.     

Male responses to conspecific advertisement signals in the field cricket Gryllus rubens (Orthoptera: Gryllidae)

In many species males aggregate and produce long-range advertisement signals to attract conspecific females. The majority of the receivers of these signals are probably other males most of the time, and male responses to competitors' signals can structure the spatial and temporal organization of the breeding aggregation and affect male mating tactics. I quantified male responses to a conspecific advertisement stimulus repeatedly over three age classes in Gryllus rubens (Orthoptera: Gryllidae) in order to estimate the type and frequency of male responses to the broadcast stimulus and to determine the factors affecting them. Factors tested included body size, wing dimorphism, age, and intensity of the broadcast stimulus. Overall, males employed acoustic response more often than positive phonotactic response. As males aged, the frequency of positive phonotactic response decreased but that of the acoustic response increased. That is, males may use positive phonotaxis in the early stages of their adult lives, possibly to find suitable calling sites or parasitize calling males, and then later in life switch to acoustic responses in response to conspecific advertisement signals. Males with smaller body size more frequently exhibited acoustic responses. This study suggests that individual variation, more than any factors measured, is critical for age-dependent male responses to conspecific advertisement signals.     

Ontogeny of the cricket Phaeophilacris bredoides Kaltenbach (Orthoptera, Gryllidae)

Development of Phaeophilacris bredoides Kalt. was studied under stable laboratory conditions: temperature of 26°C, air humidity 60%, and 12L: 12D photoperiod (Knyazev, 1985). The life cycle of Ph. bredoides includes four stages: egg, pronymph, nymph, and adult. The duration of embryonic development is 28 days. The nymphal ontogeny lasts 230 days and consists of 25 instars. The duration of nymphal instars (days) is: 1st—16, 2nd—6, 3rd—8, 4th—10, 5th—10, 6th—15, 7th—10, 8th—8, 9th—9, 10th—11, 11th—11, 12th—8, 13th—9, 14th—10, 15th—7, 16th—9, 17th—7, 18th—7, 19th—7, 20th—10, 21st—11, 22nd—7, 23rd—9, 24th—6, and 25th—9. The duration of adult life is 126 days in males and 125 days in females. Three periods were distinguished in the imaginal ontogeny of males and females: pre-reproductive, reproductive, and post-reproductive. The pre-reproductive period begins with the molt to the adult and ends with the onset of reproductive behavior in males and with the first copulation in females. Its duration is 4 (3–6) days in males and 5 (2–7) days in females. The reproductive period in males starts with the onset of reproductive behavior on the 4th (3rd–6th) day and lasts 119 (98–135) days. In females it begins when they start responding to males’ courtship behavior and lasts 116 (97–133) days. The female reproductive period includes two alternating phases: copulation and egg-laying. The egg-laying phase is initiated by successful copulation. The post-reproductive period in females starts when oviposition ceases and in males, when their reproductive behavior disappears. This period lasts about 3 (2–3) days in males and 4 (2–7) days in females, until the insect dies.     

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.     

Detailed developmental morphology of the spermatophore of the Mediterranean field cricket, Gryllus bimaculatus (De Geer) (Orthoptera: Gryllidae)

The detailed developmental morphology of the spermatophore of a cricket (Gryllus bimaculatus, De Geer) is described for the first time. A specific behavioural reference point is identified from which time developmental processes proceed in a more or less robotic manner. The structures previously identified as forming part of the fully formed spermatophore, that is, the ampulla, the sperm tube and the attachment plate are all identified in detail during their development. Interestingly the attachment plate material is extruded as an amorphous mass from the ejaculatory duct prior to its insertion into the dorsal pouch, the sperm do not enter the sperm sac until the end of spermatophore formation, and the sperm tube itself is formed from a flattened layer of material which rolls up into a tube and involves the use of a die mechanism.     

Reproductive Isolation in the Wood Cricket Gryllus vernalis (Orthoptera: Gryllidae)

Two species of closely related wood crickets, Gryllus vernalis (Orthoptera: Gryllidae) and G. fultoni, occur together in the eastern USA and have a similar calling‐song structure, consisting of three‐pulse chirps. Previous studies revealed that male calling song and female selectivity were divergent between sympatric and far allopatric populations of G. fultoni, consistent with the pattern expected of reproductive character displacement. We studied the reproductive isolation of G. vernalis in relation to G. fultoni by investigating the geographic variation in calling songs of G. vernalis and by examining close‐range mating behaviors. Neither field nor laboratory studies revealed differences in any of the calling‐song characters between sympatric and allopatric populations, but this could reflect the limited sampling within the relatively small region of allopatry for G. vernalis. Although close‐range mating trials revealed that females of both species discriminated against heterospecific mating partners, the strength of discrimination was especially strong in females of sympatric G. vernalis populations. Our studies of long‐range and close‐range mating behaviors suggest that selection pressures for reproductive isolation are exerted primarily on close‐range mating behaviors in G. vernalis but on long‐range mating behaviors in G. fultoni.     

the effect of pair formation on diel calling patterns in two cricket species,Gryllus veletis andGryllodes sigillatus (Orthoptera: Gryllidae)

After pair formation, male crickets should reduce calling to minimize the risks to males of attracting predators and/or rivals. We tested this hypothesis in two cricket species, one in which males exhibit a high mating propensity (Gryllus veletis)and another in which the mating propensity of males is constrained by the manufacture of elaborate, bipartite spermatophores (Gryllodes sigillatus).Calling durations of male G. veletisdeclined precipitously after the introduction of females but remained unchanged in G. sigillatus.We attribute the asymmetric effect of female proximity on male calling to differences in the mating propensity of males of the two species. Male G. veletisabstain from further calling in favor of repeated matings with the same female. Male G. sigillatusprobably lack the opportunity to mate repeatedly with the same female and, thus, resume calling shortly after mating to increase their probability of attracting additional mates.     

Divergence in the calling songs between sympatric and allopatric populations of the southern wood cricket Gryllus fultoni (Orthoptera: Gryllidae)

In the eastern United States the wood cricket Gryllus fultoni (Orthoptera: Gryllidae) occurs in sympatry with G. vernalis in an area between eastern Kansas and west of the Appalachian Mountains. Calling songs were recorded from 13 sympatric and allopatric localities. Both field and laboratory recordings showed that chirp rate (CR) and pulse rate (PR) overlapped extensively between allopatric populations of G. fultoni and sympatric populations of G. vernalis; by contrast, there was little or no overlap in these variables between sympatric populations of these two species. Divergence in PR and CR between the two species was thus greater in areas of sympatry than in areas of allopatry. Our field and laboratory studies of G. fultoni calling songs thus demonstrate the pattern expected of character displacement and support the genetic assumptions of this hypothesis. Other possible explanations for the sympatric divergence such as ecological character displacement and clinal variation are discussed.     

Development of microsatellite markers for the wood cricket, Nemobius sylvestris (Orthoptera: Gryllidae)

Thirty-four novel microsatellite markers developed for wood cricket (Nemobius sylvestris) were tested and optimized. Twenty-five microsatellite loci were polymorphic, exhibiting between two and nine alleles. Observed heterozygosities ranged from 0.038 to 0.925. The microsatellites were also tested in a species belonging to another genus of the Gryllidae family (Gryllus bimaculatus). Two markers produced clear banding patterns with the expected product size. These markers will be used to study the effects of forest fragmentation on genetic connectivity using wood cricket as a model species.     

Expression of allatostatins in the Mediterranean field cricket, Gryllus bimaculatus de Geer (Ensifera, Gryllidae)

The allatostatin (AST) type A gene of the cricket Gryllus bimaculatus encodes a hormone precursor including at least 14 putative peptides with a common C-terminus Y/FXFGL/Iamide. By RT-PCR we have analyzed the expression of the allatostatin precursor in various tissues of 0-21 days old adult virgin and mated females. In 3-day-old virgin females, the gene is strongly expressed in the brain (oesophageal ganglion), the suboesophageal ganglion and the caecum, but to a lower extent in other parts of the digestive tract (ileum, midgut, colon), and in various other tissues such as the fat body, ovaries and female accessory reproductive glands. In the brain and ovaries of virgin females, the AST expression is rather constant throughout adult life, whereas in brains of mated animals, expression is low until day 7, but increases sharply from day 8 onwards to reach values triple those before day 7. In ovaries of mated animals AST gene expression is also age-dependent, with high expression rates during the first 4 days after imaginable moult, a second but smaller peak from day 15 to 21, and very low values in between. In the fat body of virgin crickets allatostatin expression is high during the first 9 days after ecdysis and declines thereafter, whereas in mated animals two peak values, day 1 and day 6, are observed, and a third peak in older animals.     

Heterogonic development of the head, thorax and abdomen in Gryllus domesticus (Orthoptera:Gryllidae)

The heterogonic development of the three basic divisions of the body (head, thorax and abdomen) of Gryllus domesticus , was studied. The relationships in the linear dimensions of each of these parts with the standard parameter (absolute body length) were established through regression equations and the cephalosomatic, thoraco-somatic and abdomeno-somatic ratios. Of the body parts investigated the head was found to be the most heterogonic component followed by the abdomen and the thorax. The heterogonic development in the insect body seemed to be the result of varying intensities of the growth potential of different anatomical locations.     

The influence of female age on phonotaxis during single and multiple song presentations in the field cricket,Gryllus integer (Orthoptera: Gryllidae)

The influence of age on variation in female phonontaxis in the field cricket,Gryllus integer, was investigated using a Kugel treadmill-type device. Synthetic male calling songs, with different pulse rates, were presented in both single-stimulus and three-stimulus designs. Females were either 11–14 or 25–28 days postecdysis. Females varied in motivation, or the degree of reproductive effort they exhibited, but only in single-stimulus trials: older females achieved higher scores than younger females. Females varied in selectivity, or the extent to which they discriminated among potential mates, in both presentation designs. All females discriminated against atypical pulse rates. In multiple-stimulus trials with normal range pulse rates, younger females were discriminatory; older females were not. Mated females showed reduced phonotaxis and selectivity irrespective of mating interval.     

Characterization of an iridescent virus isolated from Gryllus bimaculatus (Orthoptera: Gryllidae)

We have isolated an iridescent virus from commercially produced colonies of Gryllus bimaculatus in Germany, which showed apparent mortality. Transmission electron microscopy studies on adult cricket specimens revealed the paracrystalline assembly of icosahedral virus particles in the cytoplasm of hypertrophied abdominal fat body cells. The infecting agent could be cultivated in the lepidopteran cell line sf-9, where it caused cytopathogenic effects such as cell hypertrophy, cytoplasmic vacuolization, and cell death within 8 days postinfection. Infection titers of the first virus passage reached 10(7.5) TCID(50)/ml. Negatively stained virus particles (n = 100) had dimensions of 172 +/- 6 nm (apex to apex) and 148 +/- 5 nm (side to side). SDS-polyacrylamide gel electrophoresis of virus proteins showed more than 20 distinct polypeptides with a major species of approximately 50 kDa. Analysis of the restriction fragment length profiles from digestion of purified viral DNA with the endonucleases EcoRI, BamHI, and HindIII showed marked differences from the profiles of iridoviruses of lower vertebrates (genus Ranavirus), e.g., Rana esculenta Iridovirus and Frog virus 3. Restriction enzyme digests with the endonucleases MspI and HpaII indicated the lack of methylation of viral DNA. Polymerase chain reaction led to the amplification of a 420-bp gene fragment with 97% sequence homology to the major capsid protein gene of Chilo iridescent virus. These data indicate that this new isolate, which we propose to be termed Gryllus bimaculatus iridescent virus, belongs to the genus Iridovirus of the family Iridoviridae.     

Tracheation of abdominal ganglia and cerci in the house cricket Acheta domesticus (Orthoptera,Gryllidae)

Patterns of tracheation in the abdominal central nervous system and the cerci of Acheta domesticus are described from whole mounts, and light and electron microscopy. The tracheal supply of the ganglia is derived from ventral longitudinal tracheal trunks which have segmental connections to the spiracels. Each abdominal ganglion is served by a single pair of tracheal trunks, except the terminal ganglion, which has two pairs. Within the ganglia, tracheoles occur principally in association with glia-rich areas of the neuropile. We suggest that the respiratory exchange may be concentrated in the cell bodies of neurons and glia. Each cercus has a tracheal supply in paralle with a large air sac which, it is suggested, serves to lighten the cercus, functions as a resonator for sound reception, or facilitates tidal flow of hemolymph and postecdysial expansion of the cercus. No tracheae run continuously between ganglia or between the terminal ganglion and the cerci, and they do not appear to have a potential role as a contact guidance pathway for cercal nerve growth.     

Learning and solar orientation in the mole cricket Gryllotalpa gryllotalpa (Orthoptera: Gryllidae)

The ecologically efficient escape direction assumed by Gryllotalpa gryllotalpais learned by the larvae after they leave the maternal nest. Nonetheless, larvae hatched in the laboratory are innately capable of using the sun to maintain a set escape direction. The species can also modify the set point of its sun compass to conform to topographical changes encountered during its wanderings.     

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.