Utilization of energy substrates during calling activity in tropical frogs

Calling activity in frogs is energetically demanding to males because they usually perform at or near their physiological capacities. Metabolic fuel for muscle contractions during bouts of aerobic calling activity comes from carbohydrates and lipids that are stored in the trunk muscles. I monitored nightly calling performance in males of seven tropical frog species from two families, Hylidae and Leptodactylidae, and compared levels of glycogen and lipid in the trunk muscles from males collected before and after a three-hour period of calling activity. Trunk muscles from late-evening males in five species had up to 63% less glycogen than the trunk muscles from early-evening males; relatively little depletion was observed in two other species. Overall, glycogen reserves and rates of depletion were highest in species with very high calling rates. It was not possible to measure changes in the relatively large stores of lipid in the trunk muscles after only 3 h of calling. Nevertheless, intramuscular lipid stores probably provide a greater percentage of the energy needed for sound production than glycogen stores, and are largest in species with high calling rates. Received: 7 January 1997 / Accepted after revision: 20 July 1997     

Size assessment in simulated territorial encounters between male green frogs (Rana clamitans)

We examined the ability of male green frogs to assess the size of an opponent based on the dominant frequency of their advertisement call, which is negatively correlated with size, using synthetic stimuli to simulate intruders of different sizes. In one field playback experiment, we broadcast a pair of stimuli representing a small and a large male; in a second experiment, we broadcast calls of a medium and a large male. In both experiments, males produced calls with significantly lower dominant frequencies in response to each stimulus. Contrast analyses revealed that males lowered the dominant frequency of their calls more in response to the large-male stimulus than in responses to the small- and medium-male stimuli. In the second experiment, males also responded to the large-male stimulus by calling at higher rates. There were no differences in mean note duration or the number of moves made toward or around the playback speaker in response to any stimulus. Thus, the frequency of an opponent's calls elicits a differential modification of calling behavior, primarily in the form of differential dominant frequency alteration, suggesting that males use dominant frequency to assess the size of opponents during aggressive encounters. Received: 17 April 1998 / Accepted after revision: 7 October 1998     

The effect of male calling on female locomotor activity of Teleogryllus commodus

Summary The calling song of the -cricket, Teleogryllus commodus, attracts for mating. Both calling and -locomotor activity are under circadian control and the behaviors occur approximately at the same time. Daily playback of calling songs for 3–12 h in constant light failed to entrain -running. Instead, calling acts as an external stimulus which can release -locomotion at any time of the 24-h period, while the clock-induced activity continues to run freely. The stimulatory effect of the calling song causes flexible extension of -locomotor activity, but a subsequent restorative period is necessary. The ecological consequences of acoustical stimulation are discussed.     

Longevity,calling effort,and metabolic rate in two populations of cricket

Intraspecific variation in a resting metabolic rate (RMR) is likely to be an important determinant of energetic-resource use and may influence the resources subsequently available for allocation to traits not directly associated with somatic maintenance. The influence of RMR on resource availability could be especially important for condition-dependent sexual traits, such as cricket calls, that are themselves energetically costly to produce. RMR may also be associated with longevity, either negatively because individuals with a high RMR burn resources faster and die young, or positively as individuals with high RMR are more able to accrue resources to fuel survival. Additionally, the associations between RMR and other characters may vary across populations if differential selection or drift shapes these traits. Here we tested for differences in RMR, body mass, calling effort, and longevity in two populations of cricket Gryllodes sigillatus and then evaluated the potential influence of RMR on calling and longevity. We find that RMR, calling effort, and longevity varied across populations, but mass did not. Controlling for population and mass, RMR was not significantly associated with calling effort, but was negatively associated with longevity. These findings suggest that male crickets that live fast die young.     

Female encounter rate and the calling behavior and mating propensity of male Requena verticalis (Orthoptera: Tettigoniidae)

Mating effort, the energy exerted in finding and persuading a member of the opposite sex to mate, may be influenced by how frequently potential mates are encountered. Specifically, males that frequently encounter females may reduce calling effort and be less eager to mate than males that infrequently encounter females. An experiment was set up to test this hypothesis, using the tettigoniid Requena verticalis. We examined the song structure, calling activity and mating propensity of individual males exposed to one of five different encounter rates with virgin females. Song structure and calling effort were significantly altered by an encounter with a female. After an encounter, males significantly increased chirp rate and decreased variability in interchirp interval. Encounters also stimulated a male to call and to continue to call for up to two hours. The elapsed time since mating affected mating propensity but not calling activity. Mating propensity asymptotically increased to reach a maximum by day 17 since last mating. However, neither the frequency of encounters, nor the number of previous encounters experienced by a male, influenced calling activity or the propensity of a male to mate. The significance of changes in song structure and calling activity following an encounter, and of increasing male mating propensity over time, are discussed. Correspondence to: G.R. Allen     

The effect of perceived intruder proximity and resident body size on the aggressive responses of male green frogs, Rana clamitans (Anura: Ranidae)

We examined variation in aggressive responses within and among individuals in the green frog, Rana clamitans. We tested the hypothesis that resident males respond in a graded fashion to changes in perceived intruder proximity (stimulus intensity). We also investigated how response level varied with responder body size. We found that green frogs differentially alter aggressive responses (increasing movement and calling rates while decreasing the duration and dominant frequency of their calls) with an increase in stimulus intensity. Body size did not appear to be a significant influence on most response levels. We suggest that aggressive responses are graded, and advertisement and aggressive calls represent opposite ends of a continuum. Green frogs do not exhibit clearly defined thresholds for responses like those reported for some other frogs. By using graded responses rather than discrete all-or-nothing responses, males may reduce the potential costs of aggressive encounters.     

Mating effort or paternal investment? Incorporation rate and cost of male donations in the wartbiter

Summary Provision of nuptial gifts and incorporation of male-derived substances occurs in several insects. There has been some discussion as to whether these gifts should be regarded as a form of paternal investment or part of mating effort. One assumption of the paternal investment hypothesis is that the donating male is expected to father all, or the majority of, the offspring. The rate of incorporation into developing eggs is therefore assumed to be rapid enough to assure the donating male of paternity of the offspring in which he is investing. Spermatophore production in a paternal investment system is also expected to be costly, since males are providing females with highly nutritious gifts during copulation, thereby increasing female fecundity. Therefore, under conditions of food limitation males are expected to alter either mating frequency or spermatophore size. Experiments on the wartbiter (Decticus verrucivorus), using radioactive isotopes, show that the incorporation rate of male-derived substances is so slow that females become sexually receptive before any material received from the last male have been incorporated into eggs laid by the female. Since sperm mixing occurs in this species and since the label is present in eggs laid up to more than 30 days after mating, males might potentially benefit another male's offspring. The nuptial gift (the spermatophylax) was found, regardless of mating frequency and diet, to have a low protein content. These findings coincide with previous results showing that consumption of the gift had no effect on female fecundity. Instead, wartbiter males keep mating frequency high, are capable of mating every day regardless of food availability, invest the same proportion of the body weight in spermatophore production regardless of size, and produce a spermatophylax that correlates with ampulla size. These results indicate that the nuptial gift mainly functions to increase fertilization success in the wartbiter.     

Factors affecting male yearly mating success in the common frog,Rana temporaria

Summary The mating behavior of the European common frog, Rana temporaria, was studied experimentally. Female body length was correlated with body mass as well as with fecundity. However, males showed no mating preference with regard to either female body length, body mass, or fecundity. In successive multiple matings, male readiness to re-mate as well as fertilization success did not vary among the first four matings. Further, fertilization success was not correlated with either the number of days since the previous fertilization, water volume in the experimental container, testes mass, female/male body length ratio, or female fecundity. However, there was a positive correlation between fertilization success and male fat reserve status. Sexual competition and mating patterns were studied in tanks in which operational sex ratio (OSR) and male density were manipulated, and time for sexual competition was allowed to vary. Successful take-overs and nonrandom mating (large male advantage) were observed only at a combination of a four-fold male bias in OSR and an unnaturally high male density (30–50/m²). I argue that in natural populations of Rana temporaria: (1) There is considerable intraspecific variation in the opportunity for sexual competition, (2) OSR influences mating pattern more than male density and time (duration of the prespawning period), and (3) nonrandom mating should be rare.     

Significance of spectral and temporal call parameters in the auditory communication of male grass frogs

Summary During the spawning period, male grass frogs (Rana temporaria L.) frequently produce short and long territorial calls in addition to mating calls. The calls differ in mean pulse number, duration, and pattern of amplitude modulation. Experiments in which recorded natural calls are played back reveal that male grass frogs are capable of discriminating the different conspecific calls. A male frog stimulated by mating calls always responds by producing mating calls in greater numbers (Fig. 3). Territorial calls presented at low intensity also cause an increase in the mating-call rate (Fig. 4), but at high intensity they clicit territorial calls and turning toward the loudspeaker. A combination of short and long territorial calls was especially effective in eliciting the phonotaxis response. As play-back experiments with simulated calls show, the carrier frequency and the pulse repetition rate are particularly important cues for recognition of conspecific calls (Fig. 5). A simulated call with a 400-Hz carrier frequency (the dominant frquency of the mating call) is just as effective as the natural call with the complete frequency spectrum (Fig. 3), whereas a 1100-Hz simulated call is ineffective (Fig. 5). The chief factors in discrimination among the conspecific calls are the call repetition rate and probably the amplitude and frequency modulation. Changes in the duration of the calls had little effect (Fig. 6). The available evidence suggests that the mating call has a reciprocally stimulating action on males in a chorus, whereas the territorial calls experess aggressiveness and give warning to other males.     

Oligogyny by unrelated queens in the carpenter ant, Camponotus ligniperdus

Multilocus DNA fingerprinting and microsatellite analysis were used to determine the number of queens and their mating frequencies in colonies of the carpenter ant, Camponotus ligniperdus (Hymenoptera: Formicidae). Only 1 of 61 analyzed queens was found to be double-mated and the population-wide effective mating frequency was therefore 1.02. In the studied population, 8 of 21 mature field colonies (38%) contained worker, male, or virgin queen genotypes which were not compatible with presumed monogyny and therefore suggested oligogyny, i.e., the cooccurrence of several mutually intolerant queens within one colony. Estimated queen numbers in oligogynous colonies ranged between two and five. According to the results of the genetic analysis, most of the queens coexisting in oligogynous colonies were not closely related. Pleometrosis is very rare and queenless colonies adopt mated queens both in the laboratory and field. Therefore, the most plausible explanation for the origin of oligogynous colonies in C. ligniperdus is the adoption of unrelated queens by orphaned mature colonies. The coexistence of unrelated, but mutually intolerant queens in C. ligniperdus colonies demonstrates that oligogyny should be considered as a phenomenon distinct from polygyny. Received: 18 December 1997 / Accepted after revision: 20 June 1998     

Selective phonotaxis by male wood frogs (<Emphasis Type="Italic">Rana sylvatica</Emphasis>) to the sound of a chorus

Frogs and toads commonly form large choruses around suitable breeding habitat during the mating season. Although often regarded as a constraint on the acoustic behavior of signalers and receivers, the sounds of a chorus could also serve as an acoustic beacon that allows some frogs to locate the breeding aggregation. Attraction to chorus sounds might be particularly important for explosively breeding frogs. In these species, which often mate just one or a few days during the year, the timing and location of breeding aggregations can be unpredictable because their formation often depends on local climatic factors, such as rainfall or a change in temperature. I used laboratory playback experiments to test the hypothesis that male wood frogs (Rana sylvatica), an explosively breeding frog, exhibit positive phonotaxis toward the sound of a conspecific chorus. Males were released at the center of a rectangular arena with a speaker positioned in each corner facing toward the release point. In a single-stimulus experiment, more males approached a speaker broadcasting a conspecific chorus than the three silent speakers in the arena. In a two-stimulus experiment, more males approached a speaker broadcasting a conspecific chorus compared to the two silent speakers or a fourth speaker simultaneously broadcasting the spectrally overlapping sound of a heterospecific (R. septentrionalis) chorus. These results are consistent with the hypothesis that male wood frogs could use the sound of a chorus as a beacon to locate a short-lived breeding aggregation.     

Habituation in the green frog, Rana clamitans

Habituation and its role in the dear enemy effect was investigated in a population of green frogs, Rana clamitans. Green frogs have a prolonged breeding season, and males defend territories centered around suitable oviposition sites. We tested the prediction that male green frogs will habituate to broadcasts of synthetic conspecific stimuli. Our results indicate that male green frogs can discriminate familiar from unfamiliar stimuli. We suggest that habituation helps to mediate the territorial interactions between male green frogs. Strangers present a greater threat than familiar neighbors. By habituating to the advertisement vocalizations of their near neighbors, males avoid costly interactions with individuals that are not a major threat to their territories. Received: 20 July 1998 / Accepted after revision: 6 September 1998     

Defense by exploitation in the Florida carpenter ant, Camponotus floridanus, at an extrafloral nectar resource

When resources in a territory have a patchy distribution, intruders may successfully exploit unguarded patches. In such cases, territory owners may use exploitative strategies to reduce the gains of the intruders. The territorial ant Camponotus floridanus attends the leaf nectaries of Urena lobata, which are also visited by the ant Pseudomyrmex mexicanus and other intruders. Residents visited the nectaries at a high rate and in a systematic way, and thereby depressed the mean standing crop per nectary. This reduces the gains of randomly visiting intruders which obtain the mean standing crop per nectary visit. Two or three residents were present on large plants and kept the mean standing crop at the same low level as at small plants with a single ant. This is an ideal free distribution of the ants. The resident ants visited the nectaries at a rate which increased in proportion to the nectar production per nectary. This is the expected systematic visitation when nectar production varies between nectaries. It is suggested that systematic visitation and maximization of the visitation rate are evolutionarily stable strategies in both residents and intruders. However, the intruders are constrained by the residents, so that they visit the nectaries less frequently and in a random manner, and thus have a lower gain rate. When the resident was temporarily absent, the intruders visited the nectaries at a high rate and systematically. Received: 31 May 1999 / Received in revised form: 15 November 1999 / Accepted: 6 December 1999     

Alarm calling behavior of the thirteen-lined ground squirrel,Spermophilus tridecemlineatus

Summary Alarm calling in a population of thirteenlined ground squirrels, Spermophilus tridecemlineatus, was studied over a three-year period. Data on ground squirrel reactions to human and canine approaches and to the approach or presence of avian predators were used to quantify alarm calling behavior.The results support the hypothesis that alarm calling in this species functions to warn genetic relatives. Human and canine approach-elicited calls were most frequently given by mothers and their recently emerged young; adult males and nonparous females rarely called. The onset of maternal calling coincided with the aboveground appearance of a mother's own litter, and both juvenile and maternal calling were terminated at approximately three weeks post-emergence. Alarm calls were rarely emitted during encounters with avian predators.Alarm calling behavior in S. tridecemlineatus thus appeared to be dependent upon the presence of newly emerged juvenile relatives. To investigate whether the population structure of S. tridecemlineatus was perhaps incompatible with the evolution of alarm calling directed toward adult relatives, the distance between the home ranges of adult relatives and the distance over which the alarm vocalization is audible to ground squirrels were measured. The results revealed that females were likely to have adult relatives relatives residing within audible range of the call.     

辛基酚对中国林蛙蝌蚪生长发育的毒性效应

为评价辛基酚(OP)对中国林蛙(Rana chensinensis)蝌蚪的急性毒性,将26期蝌蚪暴露在浓度为5.0×10-7～5.0×10-6mol·L-1OP的水体中进行急性毒性实验.结果显示,蝌蚪的死亡率随着OP浓度的升高和暴露时间的延长而增加,24、48、72、96h的半数致死浓度(LC50)分别为3.55×10-6、2.96×10-6、1.90×10-6、1.52×10-6mol·L-1;96h零致死浓度(96hLC0)为9.70×10-7mol·L-1;安全浓度(SC)为1.52×10-7mol·L-1.另外,为探讨SC以下OP对蝌蚪生长发育的影响,将26期蝌蚪连续暴露在1.0×10-7、5.0×10-8、1.0×10-8、1.0×10-9mol·L-1OP的水体中,并设1.0×10-7、1.0×10-8mol·L-1雌二醇(E2)阳性对照及空白对照,直至70%蝌蚪完全变态.在暴露20d、40d和70%蝌蚪完全变态时共3次测量蝌蚪及幼蛙的体长和体重,分别统计70%蝌蚪发育至跗蹠部伸长期、前肢伸出期和完全变态期所需的时间.结果表明,SC以下OP和E2对蝌蚪死亡率的影响不明显,但1.0×10-9～1.0×10-7mol·L-1OP和1.0×10-7、1.0×10-8mol·L-1E2可不同程度地延缓蝌蚪发育时间,降低蝌蚪体长和体重,并导致少数蝌蚪发育畸形.结果说明低浓度OP与E2相似,对蝌蚪生长发育具有抑制作用.