Temperature dependence of cicada songs (Homoptera, Cicadoidea)

The songs of male Portuguese cicadas Tettigetta argentata, T. josei and Tympanistalna gastrica were recorded at five to seven temperatures within the range 24-38.5 degrees C. To investigate the temperature dependence of the neuromuscular apparatus involved in song production, different temporal elements of the calling songs were measured. We report a strong temperature dependence for the syllable and the echeme rates in T. josei and Ty. gastrica. This suggests that in these species the neuromuscular structures involved in the timbal cycle and in generating the echeme succession of the song are strongly temperature dependent. In T. argentata, the syllable rate was again significantly temperature dependent; the echeme rate, however, increased between 25.5 degrees C and 33.5 degrees C but decreased with the highest temperature. This indicates that at least in T. argentata two separate neuronal networks control both song parameters. Other temporal elements of the song with potential behavioural significance were also measured and found to be temperature dependent (e.g. echeme duration and interval). The possible implications for intraspecific communication are discussed. We also demonstrate that the temperature of these small cicadas is not significantly influenced by the muscle activity involved in song production. On the other hand, exposure to sunlight can be used by these cicadas to elevate their body temperature by more than 10 degrees C.     

Dynamics of periodical cicada chorus centers (Homoptera: Cicadidae:Magicicada)

Mating aggregations of three species of periodical cicadas were monitored during the emergence of Brood XIX at a 16-ha study site in northwest Arkansas, May–June 1985. Magicicada tredecassiniappeared first and formed the most choruses. M. tredecimand M. tredeculachoruses formed next, and M. tredeculachoruses outnumbered those of M. tredecim.Of the 268 choruses seen, 84% were composed of M. tredecassini. M. tredecassiniwere often found chorusing in the same trees with the other two species. Such multispecies mating aggregations apparently are unique to periodical cicadas. Choruses were dynamic with respect to their locations and durations. Initially, choruses were located near areas of high cicada emergence densities. One week later, cicadas chorused in trees throughout the forest and at the forest edge. Many choruses were seen only once at a location. Although cicadas chorused for almost 4 weeks, individual choruses persisted only approximately 8 days, on average. Sound intensities under chorus centers ranged from 50 to 80 dh and were correlated with arena sizes during times of peak chorus activity. No distinct habitat preferences of the three species were observed, however, the tree species used by chorusing cicadas differed among the species.     

Postembryonic development of the auditory system of the cicada Okanagana rimosa (Say) (Homoptera: Auchenorrhyncha: Cicadidae)

Cicadas (Homoptera: Auchenorrhyncha: Cicadidae) use acoustic signalling for mate attraction and perceive auditory signals by a tympanal organ in the second abdominal segment. The main structural features of the ear are the tympanum, the sensory organ consisting of numerous scolopidial cells, and the cuticular link between sensory neurones and tympanum (tympanal ridge and apodeme). Here, a first investigation of the postembryonic development of the auditory system is presented. In insects, sensory neurones usually differentiate during embryogenesis, and sound-perceiving structures form during postembryogenesis. Cicadas have an elongated and subterranian postembryogenesis which can take several years until the final moult. The neuroanatomy and functional morphology of the auditory system of the cicada Okanagana rimosa (Say) are documented for the adult and the three last larval stages. The sensory organ and the projection of sensory afferents to the CNS are present in the earliest stages investigated. The cuticular structures of the tympanum, the tympanal frame holding the tympanum, and the tympanal ridge differentiate in the later stages of postembryogenesis. Thus, despite the different life styles of larvae and adults, the neuronal components of the cicada auditory system develop already during embryogenesis or early postembryogenesis, and sound-perceiving structures like tympana are elaborated later in postembryogenesis. The life cycle allows comparison of cicada development to other hemimetabolous insects with respect to the influence of specially adapted life cycle stages on auditory maturation. The neuronal development of the auditory system conforms to the timing in other hemimetabolous insects.     

Eicosanoids mediate control of thermoregulatory sweating in the cicada,Tibicen dealbatus (Insecta: Homoptera)

Cicadas prevent body temperature from exceeding tolerable levels by a combination of behavioral responses and sweating. Sweating is activated when body temperature reaches a critical set-point temperature. We investigated control of sweating in the cicada, Tibicen dealbatus, by chemically manipulating biosynthesis of prostaglandins and other eicosanoids. Injecting prostaglandins in amounts equal to those that induce behavioral fever in scorpions and crustaceans resulted in only a small increase in set-point temperature. Blocking prostaglandin biosynthesis with cyclo-oxygenase inhibitors such as aspirin produced significant changes in set-point temperature, confirming that prostaglandins are involved in control of sweating. However, the effect of cyclo-oxygenase inhibitors was not the opposite of the effect of prostaglandins. Instead, the effect of cyclo-oxygenase inhibitors depended strongly on the value of setpoint temperature prior to treatment. Results of biochemical manipulations of other steps in eicosanoid biosynthetic pathways corroborated the results of cyclo-oxygenase inhibition and indicated that eicosanoids other than prostaglandins may be involved in control of body temperature in normothermic T. dealbatus. The effect of cyclo-oxygenase inhibitors on a given set-point temperature depended on the ambient temperature experienced by cicadas during the experiment. Surprisingly, cicadas exposed to ambient temperatures 40°C delayed activation of sweating until body temperature exceeded values normally recorded from T. dealbatus in the field. Control of body temperature in normothermic cicadas is thus complex, involving inputs from body temperature sensors, ambient temperature sensors, and at least two cyclo-oxygenase-dependent regulatory pathways.Abbreviations PUFA polyunsatured fatty acid(s) - T _a ambient temperature - T _b body temperature - T _set set-point of body temperature for activation of sweating     

Timbal muscle physiology in the endothermic cicada Tibicen winnemanna (Homoptera: Cicadidae).

The operative muscle temperature and contraction kinetics are described for the endothermic cicada Tibicen winnemanna (Davis). Measurements of timbal muscle temperature in the field demonstrate that timbal muscle temperature is elevated above ambient temperature during activity. Timbal muscle temperature increases as the acoustic output progresses from a 'warm-up' buzz (27.9-29.7 degrees C) to full song production (36.3-39.5 degrees C). Mean muscle-ambient temperature difference increases from 5.8 degrees C in buzzing animals to 13.1 degrees C in animals producing a full calling song. Twitch rise time and onset to 50% relaxation time decrease while tension production increases with increasing temperature to 40 degrees C. Mean force production at 20 degrees C was determined to be 0.22+/-0.06 N/cm(2). The characteristics of T. winnemanna muscle are similar to those reported in other cicada species. Analysis of the songs produced by animals with one timbal destroyed and the sounds produced by mechanical manipulation suggest that the timbals alternately contract and relax in the sequence IN(1)-IN(2)-OUT(1)-OUT(2). Measurements of the inferred period of muscle contraction from song recordings and the contraction kinetics of isolated timbal muscles measured in the laboratory show that the temperature of the timbal muscle must be elevated in order for the cicada to be able to produce the calling song.     

Vitellogenin of Blattella germanica (L.) (Dictyoptera, blattellidae): nucleotide sequence of the cDNA and analysis of the protein primary structure

The cloning and sequencing of a cDNA of the vitellogenin gene from the cockroach Blattella germanica is reported. It is 5,749 nucleotides long and encodes an amino acid sequence of 1,862 residues (including a putative signal peptide of 17 residues). The vitellogenin sequence includes a long serine-rich stretch between amino acids 322 and 349, and two other stretches between amino acids 1691 and 1740. The vitellogenin of B. germanica shows a notable similarity (between 32 and 42%) to those described in other insects, and its alignment shows a high number of motifs conserved in all species, especially in the subdomains I-V. Non-parsimony methods (Neighbor Joining) of phylogenetic analysis of the insect vitellogenin sequences gave a tree showing a topology that is, in general, congruent with the currently accepted insect phylogenetic schemes. Arch.     

Evaporative cooling and endothermy in the 13-year periodical cicada,Magicicada tredecem (Homoptera: Cicadidae)

Summary The thermobiology of a cicada, Magicicada tredecem, from a warm, high humidity environment was investigated. Thoracic temperature (T_th) of M. tredecem in the field was strongly dependent on, and consistently higher than, ambient temperature (T_am), averaging 33.0±0.19°C on warm sunny days (T_am=28–29°C, rh=60–75%). Laboratory studies documented cuticle water fluxes high enough ( 5mg · cm^–2 · h^–1 in dry air at 40°C) to result in a significant degree of passive evaporative cooling, but the ability of M. tredecem to actively facilitate evaporative water loss during thermal stress is comparatively limited: water loss rates (WLR) of live M. tredecem at 40°C (dry air) were only 35–45% greater than those of dead cicadas. The limited ability of M. tredecem to facilitate transcuticular WLR is associated with limited surface distribution of the cuticular ducts through which water is actively extruded during evaporative cooling. In the laboratory, active extrusion of water had no appreciable effect on T_th, demonstrating that evaporative cooling was due largely to passive water flux through the highly permeable cuticle. The location of the abdominal pore tracts is such that extrusion of water through the ducts may preferentially cool the heart and perhaps other abdominal tissues. Long-term climatological data indicate that M. tredecem rarely encounters T_am levels high enough (i.e., above its apparent T_th setpoint of 34–35°C) to require evaporative cooling. Inactive M. tredecem can endothermically increase T_th. An hypothesis is proposed to account for the diversity of body temperature setpoints in cicadas.Abbreviations rh relative humidity - SOT standard operating temperature - T _am ambient temperature - T _b body temperature - T _sp body temperature setpoint - T _th thoracic temperature - TWF transcuticular water flux - WLR water loss rate     

Thermoregulation in the cicada Platypedia putnami variety lutea (Homoptera: Tibicinidae) with a test of a crepitation hypothesis

• 1.Measurements of body temperature (T_b) in the field demonstrate that Platypedia putnami var. lutea Davis regulates T_b through behavioral mechanisms.
• 2.Thermal responses (minimum flight temperature 17.3°C, maximum voluntary tolerance-temperature 32.5°C, and heat torpor temperature 44.4°C) of P. putnami var. lutea are related to the altitude of their habitat.
• 3.Water loss rates increase with ambient temperature (T_a). Water loss rates are not significantly different at the extremes of the active T_b range but increase significantly when exposed to elevated T_a.
• 4.Acoustic activity was restricted at 6.7°C T_b range. This is similar to the lower end of the T_b range for singing measured in cicada species that produce sound with a timbal mechanism.
• 5.The use of the wing musculature to produce acoustic signals in P. putnami var. lutea does not increase the T_b range over which the species can call compared to timbal calls produced by other cicada species.

     

Thermoregulation and the effect of body temperature on call temporal parameters in the cicada Diceroprocta olympusa (Homoptera: Cicadidae)

We investigated the thermoregulatory behavior, thermal responses (minimum flight, maximum voluntary tolerance and heat torpor temperatures) and the effect of body temperature (T(b)) on call parameters in the cicada Diceroprocta olympusa (Walker). Regression of T(b) as a function of ambient (T(a)) or perch temperatures (T(p)) suggests thermoregulation is occurring. Thermoregulation occurs through behavioral changes that alter the uptake of solar radiation. T(p) is a better predictor of T(b) than is T(a). Thermal responses (minimum flight temperature 20.4 degrees C, maximum voluntary tolerance temperature 37 degrees C, and heat torpor temperature 46.7 degrees C) may be related to the humid, grassland habitat of the species. In contrast to other acoustic insects, no significant relationship was found between the temporal parameters of the calling song and T(b) within the population of D. olympusa.     

A study of the abundance and patchiness of cicada nymphs (Homoptera: Tibicinidae) in a New Zealand subalpine shrub-grassland

Abstract

Cicada emergence skins in a subalpine shrub grassland have been sampled during 1969–75 to determine the abundance and spatial distributions of nymphs feeding on plant roots. A guild of six cicada species is primarily associated with two forms of vegetation: shrubs (Dracophyllum and Cassinia) and tall tussock (Chionochloa). Skin locations were mapped relative to dominant vegetation species, litter zones, and soil and rock pavements over a range of aspects, altitudes, and vegetation types, and sampling methods were scaled at four levels: the locality, plot, quadrat, and individual plant. There were significant differences in skin counts over four years, and different measures of mean skin densities are given for the four sampling scales. The two primary vegetation types had cumulative 1969–72 mean densities of 5.2 ± 4.0 and 12.9 ± 10.0 skins/quadrat (2.3 m²) , and the 1969–72 mean productivities of the upper 25% of quadrats (adjusted for percent ground cover) were, respectively, 5.5 and 35.5 skins/m². These productivities are believed to be conservative estimates of the maturing nymph numbers per individual host plant over the span of one cicada generation. Over a 17-year span, such productivities lie within the upper range of mean densities recorded for 17-year periodical cicadas in the United States. As the dominant subalpine vegetation species are very slow-growing,it is suggested that high densities of nymphs feeding on root sap may affect plant vitality, although 1971/1987 comparisons of vitality in 52 Chionochloa tussocks could not positively demonstrate a correlation across all data. Skin dispersion analyses indicated significant levels of patchiness, in agreement with other nymphal studies and with known cicada oviposition behaviour. No single dispersion model fitted the data comprehensively, and it is suggested that a gradual shifting of the centres of cicada aggregation may occur over a cumulative period of several generations.     

Isolation and functional analysis of a 24-residue linear alpha-helical antimicrobial peptide from Korean blackish cicada, Cryptotympana dubia (Homoptera)

A new antimicrobial peptide, cryptonin, was isolated and characterized from the adult Korean blackish cicada, Cryptotympana dubia. It consists of 24 amino acid residues and has a molecular weight of 2,704 Da on mass spectroscopy. The predicted alpha-helical structure analysis and increased helix percent in 40% trifloroethanol of cryptonin suggests that it belongs to the typical linear alpha-helix forming peptide. Binding of the biotin-labeled cryptonin at the surface of E. coli cells and increased influx of propidium iodide in E. coli after cryptonin treatment indicates that it kills microbial cells by binding bacterial cell surfaces and disrupting the cell permeability. Cryptonin showed strong antibacterial (MIC 1.56-25 microg/ml) and antifungal (MIC 3.12-50 microg/ml) activities against tested bacteria and fungi including two antibiotic-resistant bacterial strains; methicilin-resistant S. aureus and vancomycin-resistant Enterococci (MIC 25 microg/ml, each).     

Development of soybean aphid (Homoptera: Aphididae) on its primary overwintering host, Rhamnus cathartica

Thermally dependent development of soybean aphid (Aphis glycines Matsumura) and common buckthorn (Rhamnus cathartica L.) were examined in growth chambers in spring 2005. Models based on ambient air temperatures for all development events were developed. Adjusted models were developed to account for heat units acquired because of solar radiation. These models were tested at field sites in Guelph and Ridgetown, Ontario, Canada. It was found that egg hatch of aphids and bud swell of buckthorn coincided at low temperatures in growth chambers and in the field. Development thresholds of 9 and 10 degrees C were acquired for bud swell and egg hatch, respectively. Models based on ambient air temperatures were poor predictors of bud swell and egg hatch in the field, but models adjusted for solar radiation predicted these events just 1-4 d before they were observed at both sites. The results obtained have broad application for predicting aphid hatch on a regional basis.     

Entomophthora echinospora (Phycomycetes: Entomophthoraceae), a fungus pathogenic on the neotropical cicada, Procollina biolleyi (Homoptera: Cicadidae)

Entomophthora echinospora (Phycomycetes: Entomophthoraceae) has been identified as a major pathogen of the neotropical cicada Procollina biolleyi (Homoptera: Cicadidae). Mortality levels in teneral adults ranged from 40 to 70% during the peak period of adult eclosion at Bajo la Hondura, Costa Rica, in 1971.     

Molecular analysis of sorghum resistance to the greenbug (Homoptera: Aphididae)

Chromosomal regions of sorghum, Sorghum bicolor (L.) Moench, conferring resistance to greenbug, Schizaphis graminum (Rondani), biotypes C, E, I, and K from four resistance sources were evaluated by restriction fragment-length polymorphism (RFLP) analysis. At least nine loci, dispersed on eight linkage groups, were implicated in affecting sorghum resistance to greenbug. The nine loci were named according to the genus of the host plant (Sorghum) and greenbug (Schizaphis graminum). Most resistance loci were additive or incompletely dominant. Several digenic interactions were identified, and in each case, these nonadditive interactions accounted for a greater portion of the resistance phenotype than did independently acting loci. One locus in three of the four sorghum crosses appeared responsible for a large portion of resistance to greenbug biotypes C and E. None of the loci identified were effective against all biotypes studied. Correspondingly, the RFLP results indicated resistance from disparate sorghums may be a consequence of allelic variation at particular loci. To prove this, it will be necessary to fine map and clone genes for resistance to greenbug from various sorghum sources.     

Metapopulation structure of the specialized herbivore Macrosiphoniella tanacetaria (Homoptera,Aphididae)

We investigated population dynamics, genetic diversity and spatial structure in the aphid species Macrosiphoniella tanacetaria, a specialist herbivore feeding on tansy, Tanacetum vulgare. Tansy plants (genets) consist of many shoots (ramets), and genets are grouped in sites. Thus, aphids feeding on tansy can cluster at the level of ramets, genets and sites. We studied aphid population dynamics in 1997 and 2001 and found that within sites: (i). at any time, aphids used only a fraction of the available ramets and genets; (ii). at the level of ramets, most aphid colonies survived only one week; (iii). at the level of genets, mean survival time was less than 4 weeks; and (iv). colonization and extinction events occurred throughout the season. We sampled aphids in seven sites in the Alsace region, France (4-45 km apart) and two sites in Germany in 1999 to study genetic structure within and between populations. Genetic analyses using nine microsatellite loci showed that: (i). genotypic variability was high, (ii). none of the populations was in Hardy-Weinberg equilibrium, (iii). heterozygote deficits and linkage disequilibria were frequent, and (iv). all populations were genetically differentiated, even at a small geographical scale. Renewed sampling of the Alsace sites in 2001 showed that three populations had become extinct and significant genetic changes had occurred in the remaining four populations. The frequencies of extinction and colonization events at several spatial scales suggest a hierarchical metapopulation structure for M. tanacetaria. Frequent population turnover and drift are likely causes for the genetic differentiation of M. tanacetaria populations.     

Heterorhabditis sonorensis n. sp. (Nematoda: Heterorhabditidae), a natural pathogen of the seasonal cicada Diceroprocta ornea (Walker) (Homoptera: Cicadidae) in the Sonoran desert

A new Heterorhabditis species was isolated from nymphal stages of the seasonal cicada Diceroprocta ornea (Walker) in an asparagus field in the state of Sonora, Mexico. Concomitantly, another isolate of the same nematode species was also collected from an oak woodland habitat in the Chiricahua mountain range in southeastern Arizona. Morphological and molecular studies together with cross-hybridization tests indicate these two isolates are conspecific and represent a new undescribed Heterorhabditis sp. This new species is distinguished from other species in this genus by a combination of several qualitative and quantitative morphological traits. Key diagnostic features include: presence of a pronounced post-anal swelling in the hermaphrodite; male with nine pairs of bursal rays, with pairs 4 and 7 bent outwards and one pair of papillae placed on the cloacal opening, value of D% (average: 79); infective juveniles with a well developed cuticular tooth, long tail (average: 105 μm) and values of D% (average: 90) and E% (average: 99). In addition to these diagnostic characters, cross-hybridization tests between the new species with H. bacteriophora and H. mexicana yielded no fertile progeny. Comparison of ITS rDNA sequences with other available sequences of described species depicted the two isolates as a new species. Phylogenetic analysis of these sequence data placed H. sonorensis n. sp. as a member of the indica-group.