Signature whistle variations in a bottlenosed dolphin,Tursiops truncatus

To examine whether context-specific information is superimposed upon the individual cues present in the whistling of the bottlenosed dolphin, Tursiops truncatus, parameter variations within the two most frequently emitted whistle types of a captive individual were investigated in three different behavioural contexts. The study concentrated on comparing signal features of spontaneously occurring vocalizations in two possible phases following the performance of a trained discrimination task and those occurring during isolation. Phases of the discrimination task differed according to whether the animal showed correct (reward given) or incorrect performance (no reward). Signature whistles were most common in isolation, but also represented just over half of the whistles following a choice task. Of 14 signature whistle frequency and time parameters measured 9 differed significantly between isolation and at least one of the phases following a choice task (Table 1). Three parameters also varied according to whether performance was correct or incorrect. In contrast, only one out of four parameters (start frequency) measured from the second most frequent whistle type varied significantly between contexts (isolation vs. phase following correct choice). The results indicate that not only identity but also context-related information is available in the whistles of a bottlenosed dolphin.     

Signature whistles of free-ranging bottlenose dolphins Tursiops truncatus: stability and mother-offspring comparisons

Summary Mother-calf whistle exchanges were recorded from temporarily captured free-ranging bottlenose dolphins from 1975 to 1989. This is part of a long-term research project studying social structure and behavior of a community of approximately 100 dolphins in waters near Sarasota, Florida. Analysis of whistle exchanges from 12 mothercalf pairs shows that signature whistles can remain stable for periods up to at least 12 years. We looked for effects of vocal learning on the development of the signature whistle by comparing whistles of calves to those of their mothers. Eight female calves produced whistles distinct from those of their mothers, while four male calves produced whistles similar to those of their mothers. Male calves appeared to produce a greater proportion of whistles other than the signature whistle (termed variants). We hypothesize that these sex differences in whistle vocalizations may reflect differences in the roles males and females play in the social structure of the community.     

The emergence of unshared consensus decisions in bottlenose dolphins

Unshared consensus decision-making processes, in which one or a small number of individuals make the decision for the rest of a group, are rarely documented. However, this mechanism can be beneficial for all group members when one individual has greater knowledge about the benefits of the decision than other group members. Such decisions are reached during certain activity shifts within the population of bottlenose dolphins residing in Doubtful Sound, New Zealand. Behavioral signals are performed by one individual and seem to precipitate shifts in the behavior of the entire group: males perform side flops and initiate traveling bouts while females perform upside-down lobtails and terminate traveling bouts. However, these signals are not observed at all activity shifts. We find that, while side flops were performed by males that have greater knowledge than other male group members, this was not the case for females performing upside-down lobtails. The reason for this could have been that a generally high knowledge about the optimal timing of travel terminations rendered it less important which individual female made the decision. This contribution is part of the special issue “Social Networks: new perspectives” (Guest Editors: J. Krause, D. Lusseau and R. James).     

Association patterns and kinship in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) of southeastern Australia

Kinship has been shown to be an important correlate of group membership and associations among many female mammals. In this study, we investigate association patterns in female Indo-Pacific bottlenose dolphins (Tursiops aduncus) inhabiting an embayment in southeastern Australia. We combine the behavioral data with microsatellite DNA and mitochondrial DNA data to test the hypotheses that genetic relatedness and maternal kinship correlate with associations and social clusters. Mean association between females was not significantly different from a random mean, but the standard deviation was significantly higher than a random standard deviation, indicating the presence of nonrandom associates in the dataset. A neighbor-joining tree, based on the distance of associations between females, identified four main social clusters in the area. Mean genetic relatedness between pairs of frequent female associates was significantly higher than that between pairs of infrequent associates. There was also a significant correlation between mtDNA haplotype sharing and the degree of female association. However, the mean genetic relatedness of female pairs within and between social clusters and the proportion of female pairs with the same and different mtDNA haplotypes within and between clusters were not significantly different. This study demonstrates that kinship correlates with associations among female bottlenose dolphins, but that kinship relations are not necessarily a prerequisite for membership in social clusters. We hypothesize that different forces acting on female bottlenose dolphin sociality appear to promote the formation of flexible groups which include both kin and nonkin.     

Whistle sharing in paired male bottlenose dolphins,<Emphasis Type="Italic"> Tursiops truncatus</Emphasis>

The signature whistle hypothesis states that dolphins produce highly stereotyped, individually distinctive whistles when in isolation. The presence of signature whistles has been called into question by recent studies proposing that dolphins produce a shared, simple upsweep whistle when in isolation, and that whistles produced by socializing dolphins are shared across individuals and social groups. This shared repertoire hypothesis suggests that when two animals produce the same whistle type, it is due to sharing the same common repertoire rather than one animal learning to produce the whistle of another. One difference between studies supporting or denying the existence of signature whistles is the method used to classify whistle types. We examined whistle production by 17 free-ranging bottlenose dolphins while temporarily restrained. We used both a quantitative comparison technique similar to that used to support the shared repertoire hypothesis and human judges to classify whistle types and quantify similarity between types. Contrary to recent studies that emphasize shared whistles, overall whistle sharing between isolated individuals was low (25%) and a simple upsweep did not account for the most common whistle type in half of the animals. Some species of birds, bats, and primates with stable social groups use vocal learning to converge over time to one common group distinctive call type. We examined whistle similarity between adult male dolphins that are partners in a close social alliance in order to test whether vocal learning may enable a similar vocal convergence. Whistle similarity was rated very high between partners and low between non-partners by both the quantitative technique and human observers. This suggests that as in songbirds and some other mammals, adult male bottlenose dolphins may use vocal learning to converge on similar whistles as they develop affiliative social relationships.Communicated by G. Wilkinson     

Whistle repertoires of two bottlenosed dolphins,Tursiops truncatus: mimicry of signature whistles?

Summary The whistle vocalizations of two bottlenosed dolphins, Tursiops truncatus, were recorded at the Sealand Aquarium in Brewster, Massachusetts. The identification of which dolphin within the group produced a vocalization was made possible by a telemetry device attached to the dolphin's head with a suction cup. 77% of the identified whistles (219 our of 284) fell into two primary categories, type 1 and type 2 (Table 1). The remaining 23% of whistles fell into five secondary categories. Of the primary whistles produced by one dolphin, 78% were of type 1 (22% type 2), while 69% of primary whistles from the other dolphin were of type 2 (31% type 1). The result that each of the dolphins favored a different primary whistle supports the findings of Caldwell and Caldwell (1965), that each dolphin produces an individually distinctive whistle. But in the present study, both dolphins produced both primary whistle types. This may represent mimicry of signature whistles.     

Erratum to: Prey selection by resident common bottlenose dolphins (Tursiops truncatus) in Sarasota Bay, Florida

Prey selection was investigated in wild, resident common bottlenose dolphins, Tursiops truncatus, during the summer months in Sarasota Bay, Florida, USA. Stomach content analyses of 15 dolphins with extensive sighting histories and well-documented distributions were used to determine prey use. Prey availability was assessed by purse seine surveys. We compared the relative abundances of prey available to estimates of prey use at closely matching spatial and temporal scales. G-tests determined that dolphins in this study significantly selected for prey at the species, family, and soniferous/non-soniferous prey levels (G _adj  = 753.98–1,775.93, df = 1–21, p ≤ 0.01). While comprising only 6.3% of the total available prey, soniferous fishes accounted for 51.9% of the total prey consumed. Manly’s standardized forage ratios and 95% Bonferroni confidence intervals determined significant positive selection for soniferous prey and against non-soniferous prey (β^S = 0.9461 vs. β^NS = 0.0539). Dolphins selected against Gerridae, Clupeidae, and Sparidae (β ≤ 0.0014), as well as against all the species within those families (β ≤ 0.0190). It is likely that passive listening for soniferous prey provides an ecological or energetic advantage to cetaceans utilizing this specific foraging technique.     

Sex differences in survival costs of reproduction in a promiscuous primate

In sexually promiscuous mammals, female reproductive effort is mainly expressed through gestation, lactation, and maternal care, whereas male reproductive effort is mainly manifested as mating effort. In this study, we investigated whether reproduction has significant survival costs for a seasonally breeding, sexually promiscuous species, the rhesus macaque, and whether these costs occur at different times of the year for females and males, namely in the birth and the mating season, respectively. The study was conducted with the rhesus macaque population on Cayo Santiago, Puerto Rico. Data on 7,402 births and 922 deaths over a 45-year period were analyzed. Births were concentrated between November and April, while conceptions occurred between May and October. As predicted, female mortality probability peaked in the birth season whereas male mortality probability peaked in the mating season. Furthermore, as the onset of the birth season gradually shifted over the years in relation to climatic changes, there was a concomitant shift in the seasonal peaks of male and female mortality. Taken together, our findings provide the first evidence of sex differences in the survival costs of reproduction in nonhuman primates and suggest that reproduction has significant fitness costs even in environments with abundant food and absence of predation.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Sex pheromone of the large aspen tortrix, Choristoneura conflictana(Lepidoptera: Tortricidae)

Summary. Three components that elicited antennal response from male Choristoneura conflictana were found from female gland extracts analyzed using a coupled gas chromatographic-electroantennographic detector system. The main component in gland extracts was (Z)-11-tetradecenal (Z11-14:Ald). Two minor components also elicited antennal response: (E)11-tetradecenal (E11-14:Ald) and (Z)-11- tetradecen-1-ol (Z11-14:OH). Analysis of effluvia indicated that calling virgin females release mostly Z11-14:Ald and trace amounts of Z11-14:OH. Field and wind tunnel behavioral studies showed that Z11-14:Ald alone attracted male moths in a dose response pattern. Tests comparing male response to blends of components detected in gland extracts showed that addition of 1.8% of E11-14:Ald to Z11-14:Ald did not influence male moths in the wind tunnel, but resulted in significantly lower trap captures in the field. The threecomponent blend [Z11-14:Ald (100), E11-14:Ald (1.8), Z11-14:OH (11)], was less attractive than Z11-14:Ald alone in both field and wind tunnel studies. Traps baited with two virgin female moths were equally attractive to males as traps baited with the three-component synthetic blend but less attractive than traps baited with Z11-14:Ald alone. Field tests of various blends of the two components (Z11-14:Ald, Z11-14:OH) detected in the females’ effluvia showed that the addition of 1–10% Z11-14:OH to Z11-14:Ald did not affect the males’ response to Z11-14:Ald. Our data demonstrate that female C. conflictana release sex pheromone components in a different ratio than they are stored in the pheromone gland. The sex pheromone is comprised of a single component, Z11-14:Ald, that can be used to monitor mated and virgin male C. conflictana throughout their flight period.     

Sex ratio dynamics and fluctuating selection pressures in natural populations of the Trinidadian guppy,<Emphasis Type="Italic"> Poecilia reticulata</Emphasis>

In many species, population sex ratios have far-reaching consequences for a wide variety of population-level and behavioural processes and can directly influence sexual selection through differential effects on male and female mating behaviour. Although sex ratios are often treated as more or less stable population characteristics, recent theoretical evidence suggests that sex ratios fluctuate under many conditions, and that the amplitude of these fluctuations can be considerable. Few studies have attempted to quantify this variation in systems with prominent, sex ratio-dependent sexual conflict. One of the species with the greatest potential to integrate these factors in the wild is the Trinidadian guppy, Poecilia reticulata. In this study, we quantified natural sex ratio variation both as detailed longitudinal studies of focal guppy populations and as snapshot estimates across a range of freshwater habitats. In line with theoretical predictions, we expected to detect significant sex ratio variation over time. We also investigated the association between juvenile and adult sex ratios to quantify a possible compensatory feedback implied in standard models of sex ratio evolution. Our results confirm that population-level sex ratios in wild guppy populations have a range of dynamic features, with all four focal populations showing significant variation in sex ratio over time. The survey showed that juveniles were generally close to equal (50:50) sex ratios whereas 7 out of 11 adult sex ratios differed significantly from equality. We found no evidence that a surplus of juveniles of the locally rarer sex had been produced. The results indicate that sex ratios and hence the balance between sexual selection and sexual coercion is normally fluctuating in nature, despite juvenile ratios being close to equality.Communicated by J. Krause     

Sex pheromone components of pyralid moths Terastia subjectalis and Agathodes ostentalis feeding on the coral tree, Erithrina variegata: Two sympatric species share common components in different ratios

Summary. Two common components, identified as (E)-11-hexadecenal (E11-16:Ald) and (E,E)-10,12-hexadecadienal (E10E12-16:Ald), were found in the extract of virgin females of two sympatric pyralid moths, Terastia subjectalis Lederer, and Agathodes ostentalis (Geyer). The amount of E11-16:Ald and E10E12-16:Ald was 0.12 and 2.8 ng/female for T. subjectalis and 2.0 and 1.8 ng/female for A. ostentalis, respectively. Hexadecanal (16:Ald) was also found at 0.7 ng/female in the extract of A. ostentalis, but EAG activity was unclear. In the field, T. subjectalis and A. ostentalis males were captured with species specific blends in the proportions of 5:95 and 50:50 blends. Blends attractive to one species did not attract the opposite one. Rubber septa loaded with 1 mg each of 5:95 and 50:50 blends of E11-16:Ald and E10E12-16:Ald were as attractive as two virgin females of T. subjectalis and A. ostentalis , respectively. This demonstrated that different ratios of shared pheromone components could provide species specific cues for critical mate location in the two sympatric pyralid moths. It is suggested that directional selection probably occurs in the pheromone blends in opposite directions in both species.     

Sex allocation in a monomorphic seabird with a single-egg clutch: test of the environment, mate quality, and female condition hypotheses

Sex allocation theory posits that mothers should preferentially invest in sons when environmental conditions are favorable for breeding, their mates are of high quality, or they are in good body condition. We tested these three hypotheses in rhinoceros auklets (Cerorhinca monocerata), monomorphic seabirds that lay a single-egg clutch, in 2 years that differed in environmental conditions for breeding. Results supported the environment and mate quality hypotheses, but these effects were interactive: offspring sex was independent of paternal traits in the poor year for breeding, while females mated to larger and more ornamented males reared more sons in the better year. Conversely, offspring sex was unrelated to female condition, as indexed by hatching date. We propose that good rearing conditions enable females to rear sons possessing the desirable phenotypic attributes of their mates. Results also supported two critical assumptions of sex allocation theory: (1) dimorphism in offspring condition at independence: daughters fledged with higher baseline levels of corticosterone than sons and (2) differential costs of rearing sons versus daughters: mothers rearing sons when environmental conditions were poor completed parental care in poorer condition than mothers rearing daughters in the same year and mothers rearing either sex when conditions were better. These novel results may help to explain the disparate results of previous studies of avian sex allocation.     

Experimental test of socially mediated sex change in a protandric simultaneous hermaphrodite,the marine shrimp<Emphasis Type="Italic"> Lysmata wurdemanni</Emphasis> (Caridea: Hippolytidae)

In Lysmata wurdemanni, individuals begin benthic life in a male phase (MP) but later change to a female phase (FP) with female external morphology, but with both male and female reproductive capacity (protandric simultaneous hermaphroditism). Previous studies have demonstrated that the size (timing) of sex change varies considerably in natural populations. We experimentally tested for social mediation of sex change by rearing male-phase individuals (MPs) in both large and small social groups with different sexual and size composition. In the large group experiment, speed of sex change was inversely related to the abundance of female-phase individuals (FPs) in the group (sex-ratio induction). Increased allocation to female function (more rapid change to FP) may occur when male mating opportunities are lower because the simultaneous-hermaphrodite FP can immediately reproduce as a female while maintaining male mating capacity. When FPs are abundant, delayed sex change might be adaptive because the costs of female reproduction are considerable. An MP may gain reproductively by increased growth before changing to FP at a larger size (fewer but much larger broods). Size-ratio induction of sex change by small MPs was suggested but not confirmed. Experimental results from small groups (1–2 individuals) were qualitatively similar but not as conclusive as those from large groups. The number and complexity of social interactions in large groups may be necessary to stimulate labile sex change in this species. In L. wurdemanni, sex change may be influenced not only by abiotic factors related to breeding [Bauer (2002) Biol Bull 203:347–357] but also by social factors in certain demographic situations.Communicated by K. Lindström     

Pup production,sex ratios,and survivorship in African wild dogs, <Emphasis Type="Italic">Lycaon pictus</Emphasis>

The local resource enhancement (LRE) model predicts that in cooperatively breeding species, sex ratios will be biased in favor of the more helpful sex. In this study, we assess the assumptions underlying the LRE model in a population of cooperatively breeding wild dogs (Lycaon pictus) in Northern Botswana monitored over a 15-year period. In this population, litter size and pup survival to 1 year are strongly affected by pack size and the breeding female’s age, but adult males have a stronger and more linear effect on females’ reproductive performance than do adult females. This asymmetry in the benefits derived from male and female helpers is reflected in male-biased sex ratios in litters at the time pups emerge from the den. Sex ratio biases are most pronounced in the litters of the youngest mothers who live in significantly smaller packs than older females. The presence of potential rivals for the dominant female’s position depresses pup production at the time of emergence, suggesting that competition among females for breeding positions may also contribute to the selective forces affecting birth sex ratios.     

Operational sex ratio versus gender density as determinants of copulation duration in the walnut fly,Rhagoletis juglandis (Diptera: Tephritidae)

In laboratory and field studies of the walnut fly, Rhagoletis juglandis Cresson (Diptera: Tephritidae), we assessed the effect of operational sex ratio on copulation duration and partitioned the sex ratio effect into component effects due to male density and female density. In our first laboratory experiment, results were clearly consistent with theoretical expectation: increases in male density were associated with significant increases in copulation duration while increases in female density were associated with significant decreases in copulation duration. These component effects yielded a striking composite effect of operational sex ratio (OSR) on copulation duration in which male-biased ratios were associated with low frequencies of short copulations and female-biased ratios were associated with high frequencies of short copulations. Consistent with a priori expectations concerning costs of territorial behavior, the effect of male density on copulation duration was stronger than that of female density. There was no significant interaction between the effects of gender density on copulation duration: each gender density contributed additively to the composite OSR effect on copulation duration. In contrast to the effect of OSR, overall density had little effect. Field data corroborated these findings fully and showed additionally that OSR in the vicinity of fruit tended in nature to be male-biased. In a second laboratory experiment, we measured copulation duration for individuals exposed alternately to male-biased and female-biased ratios. Individual flies consistently copulated for longer in male-biased environments than in female-biased ones. We propose that this plasticity permits individuals to track changes in local sex ratio over space and time and respond appropriately. Received: 15 November 1995/Accepted after revision: 27 April 1996     

Maternal characteristics and the production and recruitment of sons in the eastern kingbird (<Emphasis Type="BoldItalic">Tyrannus tyrannus</Emphasis>)

Sex allocation theory predicts that if variance in reproductive success differs between the sexes, females who are able to produce high-quality young should bias offspring sex ratio towards the sex with the higher potential reproductive success. We tested the hypothesis that high-quality (i.e., heavy) female eastern kingbirds (Tyrannus tyrannus) that bred early in the breeding season would produce male-biased clutches. A significant opportunity for sexual selection also exists in this socially monogamous but cryptically polygamous species, and we predicted that successful extra-pair (EP) sires would be associated with an excess of male offspring. Although population brood sex ratio did not differ from parity, it increased significantly with female body mass and declined with female breeding date, but was independent of the morphology and display (song) behavior (correlates of reproductive success) of social males and EP sires. Male offspring were significantly heavier than female offspring at fledging. Moreover, the probability that male offspring were resighted in subsequent years declined with breeding date, and was greater in replacement clutches, but lower when clutch size was large. Probability of resighting female offspring varied annually, but was independent of all other variables. Given that variance in reproductive success of male kingbirds is much greater than that of females, and that male offspring are more expensive to produce and have a higher probability of recruitment if fledged early in the season, our results support predictions of sex allocation theory: high-quality (heavy) females breeding when conditions were optimal for male recruitment produced an excess of sons.     

The effects of castration, sex ratio and population density on social segregation and habitat use in Soay sheep

We analysed 16 years of census data gathered on the island of Hirta (archipelago of St. Kilda) to investigate the effects of castration, population density, sex ratio, season and group type on habitat use and social segregation of Soay sheep. From 1978 to 1980, 72 male lambs were castrated. We used this experiment to study how a change in reproductive status could affect sociality and habitat choice of these males. Males, females and castrates were all segregated outside the rutting season in autumn. Castrates were the least segregated from females in spring and summer but were most segregated from them during the pre-rut. The more equal the sex ratios, the higher was the degree of social segregation. The three sex classes used similar habitat types, namely, Holcus agrostis, Agrostis festuca and Calluna habitats. Holcus agrostis and Agrostis festuca were top- and second-ranked in female and castrate habitat use, while Holcus agrostis and Calluna were the two top habitat types used by rams. It is unclear why males included Calluna heath habitats, but it cannot be excluded that they might have shifted their use depending on forage availability. A lack in differences in habitat use between castrates and females suggests that body size differences alone cannot be the driving factor for habitat segregation in male and female Soay sheep and that there are reasons other than body size that could motivate reproductive males to use additional habitat types, such as Calluna heath. Although habitat use shifted from one habitat type to the next between low- and high-population-density years and between seasons, there was no clear link between population density and how different groups (male, female or castrate) used these areas. We discuss effects of reproductive status, population density and sex ratio on social segregation and habitat use and suggest that these factors need to be taken into account when investigating causes of sexual segregation in ungulates.     

Breeding Periodicity for Male Sea Turtles,Operational Sex Ratios,and Implications in the Face of Climate Change

Abstract: Species that have temperature‐dependent sex determination (TSD) often produce highly skewed offspring sex ratios contrary to long‐standing theoretical predictions. This ecological enigma has provoked concern that climate change may induce the production of single‐sex generations and hence lead to population extirpation. All species of sea turtles exhibit TSD, many are already endangered, and most already produce sex ratios skewed to the sex produced at warmer temperatures (females). We tracked male loggerhead turtles (Caretta caretta) from Zakynthos, Greece, throughout the entire interval between successive breeding seasons and identified individuals on their breeding grounds, using photoidentification, to determine breeding periodicity and operational sex ratios. Males returned to breed at least twice as frequently as females. We estimated that the hatchling sex ratio of 70:30 female to male for this rookery will translate into an overall operational sex ratio (OSR) (i.e., ratio of total number of males vs females breeding each year) of close to 50:50 female to male. We followed three male turtles for between 10 and 12 months during which time they all traveled back to the breeding grounds. Flipper tagging revealed the proportion of females returning to nest after intervals of 1, 2, 3, and 4 years were 0.21, 0.38, 0.29, and 0.12, respectively (mean interval 2.3 years). A further nine male turtles were tracked for short periods to determine their departure date from the breeding grounds. These departure dates were combined with a photoidentification data set of 165 individuals identified on in‐water transect surveys at the start of the breeding season to develop a statistical model of the population dynamics. This model produced a maximum likelihood estimate that males visit the breeding site 2.6 times more often than females (95%CI 2.1, 3.1), which was consistent with the data from satellite tracking and flipper tagging. Increased frequency of male breeding will help ameliorate female‐biased hatchling sex ratios. Combined with the ability of males to fertilize the eggs of many females and for females to store sperm to fertilize many clutches, our results imply that effects of climate change on the viability of sea turtle populations are likely to be less acute than previously suspected.