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Pulse Rates in the Songs of Trilling Field Crickets (Orthoptera: Gryllidae: Gryllus)

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Pulse Rates in the Songs of Trilling Field Crickets (Orthoptera: Gryllidae: Gryllus) ARTHOPOD BIOLOGY Pulse Rates in the Songs of Trilling Field Crickets (Orthoptera: Gryllidae: Gryllus) THOMAS J. WALKER Department of Entomology and Nematology, University of Florida, Gainesville, FL 32611Ð0620 Ann. Entomol. Soc. Am. 93(3): 565Ð572 (2000) ABSTRACT In cricket calling songs, pulse rate is often the feature that allows females to home on the songs of conspeciÞc males while ignoring those of other species. A previous study of the sibling species Gryllus rubens Scudder and G. texensis Cade & Otte from western Florida showed unexpected effects of developmental conditions and parental generation on the pulse rates of their songs. The current study tested for these and other effects in G. rubens from Gainesville, FL, and G. texensis from Dallas, TX. In both species, sibships from different Þeld-fertilized females differed signiÞcantly in the effects of rearing environment on the pulse rates of their songs. Tests for effects of nymphal diapause and adult age were negative. Hybrids between the two species produced pulse rates intermediate to the rates of the control crosses. Males of G. rubens reared at 32ЊC produced faster pulse rates (at a standard temperature) than those reared at 25ЊC. Furthermore, pulse rates of Þeld-collected males in this study and of Þeld-collected and laboratory-reared males in the previous study increased in rough proportion to the known and estimated temperatures during nymphal development. KEY WORDS crickets, calling songs, developmental effects, age effects, hybrids THE RESEARCH REPORTED here was prompted by an ear- ing on their generation, and males reared in the lab- lier study in which I concluded that two Þeld cricket oratory under similar conditions differed in pulse rate species distinguished only by the pulse rates of their depending on the generation of their parents. No calling songs occur in mixed populations in western spring generation G. texensis were collected, but fall Florida (Walker 1998). A continued dichotomy in the males of G. texensis and the sons of fall-collected fe- pulse rates of callings songs during a 6-yr study sup- males had mean pulse rates of 77.6 and 70.6, respec- ported the hypothesis that introgression between the tively, the same relationship as exhibited by the cor- two species was little or lacking. Also supporting this responding classes of G. rubens (58.6 and 51.8) hypothesis were the songs of laboratory-reared sons of In crickets, pulse rate of a given species at a given Þeld-fertilized females. Males of each sibship had sim- temperature generally varies little and is an important ilar pulse rates, and the mean pulse rates of sibships determinant of female response to the song (Walker were clearly dichotomous. Of 21 sibships from females 1957, 1962; Doherty and Callos 1991). In fact, the songs collected in a vacant lot in Milton, FL, 25 September of trilling species that sing together frequently differ 1982, 11 were Gryllus rubens Scudder and 10 were G. only in their pulse rates. Therefore, developmental texensis Cade & Otte. effects on pulse rate, which would make the calls less The most surprising conclusion of the western Flor- species-speciÞc, should have reproductive disadvan- ida study was that developmental conditions and pa- tages and be minimized by natural selection. In the rental generation affected the pulse rates of calling only case where pulse rate differences attributable to males. Both G. rubens and G. texensis are bivoltine as developmental environment have been sought, revealed by a spring peak of adult abundance, when Walker (1962) found the songs of spring and fall gen- overwintering nymphs mature, and a fall peak, when eration Oecanthus argentinus Saussure indistinguish- the progeny of the spring generation mature. G. rubens able, though the songsters developed under contrast- males collected in the spring had a mean pulse rate of ing temperatures and photoperiods. 48.6, whereas males reared from fall-collected females In the current study, I used G. rubens from Gaines- (which in the Þeld would have been spring males) had ville, FL, and G. texensis from Dallas, TX, to test a mean pulse rate of 51.8. G. rubens males collected in whether the following factors inßuence pulse rate: the fall had a mean pulse rate of 58.6 and males reared sibship (mother), parental generation, temperature from spring-collected females had a mean pulse rate of and photoperiod during development, nymphal dia- 55.3. Thus, both spring and fall generation G. rubens pause, adult age, and interspeciÞc hybridization. males differed in pulse rate depending on whether In articles published before 2000, G. texensis was they developed under Þeld or laboratory conditions. called G. integer Scudder, even though the former Field-reared G. rubens differed in pulse rate depend- species was judged distinct from G. integer as early as 0013-8746/00/0565Ð0572$02.00/0 ᭧ 2000 Entomological Society of America 566 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 93, no. 3 1980 (Weissman et al. 1980). Smith and Cade (1987) 3 June 1997. Females to be used in the crosses were showed that G. integer from California (its type local- isolated from males as nymphs to ensure virginity. For ity) would not produce offspring in laboratory crosses each cross, Þve pairs of crickets were established in with “integer” from Austin, TX, or with G. rubens from individual wide-mouth, 1-liter jars one-fourth Þlled McAlester, OK. Walker (1998) conÞrmed the distinct- with damp sand. Parents were removed when dead or ness of G. rubens and G. “integer” where they overlap, when their progeny began to hatch. Hatchlings were and Cade and Otte (2000) described “integer” as G. transferred to 3.8-liter rearing jars corresponding to texensis. their cross and replicate. When numerous F2 hatch- lings were noted in the rearing jar of a TxR sibship, they were reared and the songs of males recorded. All Materials and Methods crosses were made and reared in a room held at 25 Ϯ Specimens. Robert L. Crocker collected fall gener- 1ЊC and a photoperiod of 16:8 (L:D) h. ation G. texensis by broadcasting synthetic G. texensis Tape Recording. As sons of Þeld-collected females calling song at Dallas, TX, 10 October 1996. On 20 May or laboratory-crossed pairs matured, they were trans- 1997, he collected spring generation G. texensis at ferred to individual 120-ml glass or plastic containers Dallas, and at a nearby site in Ft. Worth, by broad- with screen lids, assigned an identiÞcation number, casting synthetic calling song and by inspection. At and placed in an array on a table in a windowed room Gainesville, FL, I collected G. rubens using broadcasts kept at 25 Ϯ 2ЊC. They were fed oat cereal and small of synthetic G. rubens song (Walker 1996). Fall gen- pieces of apple, which also provided moisture. I ini- eration females were collected 21 and 24 September tially monitored the array Ͼ4 h a day and recorded 1996 and spring generation females were collected 17 songs whenever I could. When I found that most May 1997. Males were collected in 11 of the 14 mo calling occurred at dawn or when lights were turned between September 1996 and October 1997. on before or during dawn, I made that a regular time Rearing. Crickets were reared as described by Win- for recording. Recordings were made with a reel-to- eriter and Walker (1988). The 3.8-liter rearing jars reel tape recorder (Nagra IV, Kudelski, New York, were serviced weekly. Six to eight Þeld-collected fe- NY) and a dynamic microphone (D33, American, males representing each generation and species were New York, NY) afÞxed to one end of a 0.4-m shaft. In placed individually in jars with damp sand for ovipo- each taping session, I tried to make a 20-s recording of sition and held in a rearing room at 25 Ϯ 1ЊC and a each calling male that had not been recorded within long-day photoperiod of 16:8 (L:D) h. Females were 24 h. Temperature was measured after each recording removed no later than the week after Þrst hatch. No with an electric thermometer (BAT-12, PhysiTemp, later than 4 wk after Þrst hatch, most sibships that had Clifton, NJ) from a copper-constantan thermocouple Ͼ120 juveniles were divided evenly among three fresh at a container near the center of the array. A cylinder jars. For each of these sibships, one jar was left in the made from a 13 by 18-cm card surrounded each con- rearing room under 25ЊC and long days (16:8 [L:D] h); tainer to keep callers from seeing the approach of the another was placed in a Florida Reach-In environ- microphone. Twice-recorded crickets were usually mental chamber (Walker et al. 1993) that maintained killed, pinned, labeled, and placed in the Florida State a short day (11:13 [L:D] h) and 25.0 Ϯ 0.2ЊC; and a Collection of Arthropods. To determine whether third was placed in a similar chamber at 32.0 Ϯ 0.2ЊC pulse rate changes with adult age, eight males, from and a photophase of either 16 or 11 h. For spring- and fall-collected G. rubens females, were taped again 25 fall-collected female G. rubens and for spring-col- or more days after their second taping. lected G. texensis, three sibships each were divided as Analysis of Taped Songs. The calls of trilling Þeld described above. For fall-collected G. texensis, no fe- crickets often have frequent, brief, irregularly occur- male produced Ͼ35 progeny; therefore, 5 wk after ring delays in what would otherwise be a uniform initial hatch, the 65 progeny in a jar that had held three series of pulses.
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