Information transfer and gain in flocks: the effects of quality and quantity of social information at different neighbour distances

We assessed experimentally how the quality and quantity of social information affected foraging decisions of starlings (Sturnus vulgaris) at different neighbour distances, and how individuals gained social information as a function of head position. Our experimental set up comprised three bottomless enclosures, each housing one individual placed on a line at different distances. The birds in the extreme enclosures were labelled senders and the one in the centre receiver. We manipulated the foraging opportunities of senders (enhanced, natural, no-foraging), and recorded the behaviour of the receiver. In the first experiment, receivers responded to the condition of senders. Their searching rate and food intake increased when senders foraged in enhanced conditions, and decreased in no-foraging conditions, in relation to natural conditions. Scanning was oriented more in the direction of conspecifics when senders behaviour departed from normal. In the second experiment, responses were dose dependent: receivers increased their searching rate and orientated their gaze more towards conspecifics with the number of senders foraging in enhanced food conditions. In no-foraging conditions, receivers decreased their searching and intake rates with the number of senders, but no variation was found in scanning towards conspecifics. Differences in foraging and scanning behaviour between enhanced and no-foraging conditions were much lower when neighbours were separated farther. Overall, information transfer within starling flocks affects individual foraging and scanning behaviour, with receivers monitoring and copying senders behaviour mainly when neighbours are close. Information transfer may be related to predation information (responding to the vigilance of conspecifics) and foraging information (responding to the feeding success of conspecifics). Both sources of information, balanced by neighbour distance, may simultaneously affect the behaviour of individuals in natural conditions.Communicated by H. Kokko     

Dispersal of phytoplanktotrophic shipworm larvae (Bivalvia: Teredinidae) over long distances by ocean currents

Shipworms or Teredinidae may be dispersed either as adults in floating wooden objects or as pelagic larvae drifting near the sea surface. Five shipworm species, i.e., half of those having an amphi-Atlantic geographical distribution, are known also to have pelagic phytoplanktotrophic larvae which can be carried by ocean currents. From a series of 742 plankton samples taken from throughout the temperate and tropical North Atlantic Ocean, it can be shown that shipworm larvae are not uncommon in the open sea. Teredinid veligers were found in 19% of all samples taken. One species of larvae, identical in all details to that described by Rancurel (1965), is particularly prevalent and is tentatively identified as Teredora malleolus (Turton). A definitive identification will be possible only after the pelagic larvae of the other Atlantic species are known. The larvae of Teredora malleolus are found throughout the North Atlantic Gyre and the adjacent temperate and tropical seas, and from scattered records in the South Equatorial Current. Larvae of other unidentified Teredinidae species were also found. The distance that larvae may be transported depends upon the length of pelagic larval development and the velocity of the currents. From the known current velocities it can be shown that, even in a few weeks, larvae may be dispersed many hundreds of kilometers. The geographical distribution of shipworm larvae suggests that they are carried along the coasts of continents and even across ocean basins, and that this dispersal must be an important factor in the geographical distribution of the adult forms and in the maintenance of genetic continuity between populations otherwise isolated from one another.Contribution No. 2555 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA     

Female choice of multiple male criteria in guppies: interacting effects of dominance,coloration and courtship

Summary I experimentally examined the relative importance of social dominance, color patterns, and courtship behavior in male mating and reproductive success in the guppy Poecilia reticulata. Female choice of males is based on a complex set of behavioral and morphological traits. The results of 59 paired-male one-female visual choice and mating trials showed that male mating success was positively correlated with dominance, courtship intensity, and male coloration. Only dominant males engaged in full copoulations, and they sired two-thirds of the broods. An analysis of the paternity of broods and results of mating trials showed that a female's visual response when the sexes are separated by a glass partition is a good predictor of a male's reproductive success when the partition is removed and they are allowed to mate. A canonical correlation analysis of male behavioral and morphological traits indicated that female visual response and male mating success were positively correlated with male courtship and with agonistic behavior. However, the relative importance of color varied. Carotenoid and iridescent spots were important both in attracting the female's attention and in enhancing male mating success. Melanins were not correlated with either mating success or female response. There was a relatively low correlation (48%) between male behavioral and morphological variables and female response variables (full copulation and female visual response). These results suggest that female choice is subtle, and is based on a complex suite of male behavioral and morphological traits as well as on competitive interactions among males.     

Female choice in sage grouse: the roles of attraction and active comparison

Previous studies of female choice in sage grouse Centrocercus urophasianus have implicated both the acoustic quality and repetition rate of the stereotyped strut display as putative cues for female choice. Stages in the choice process at which specific components of male courtship display influence female decisions were investigated using field observations of female pre-mating behavior. Females visited a subset of territorial males and then actively chose one of these as a mate. The order in which males were visited suggested that females searched until an acceptable mate was found, rather than employing a “best-of-n” tactic. Numbers of females visiting a male were related to differences in an acoustical component of display (inter-pop interval) whereas the probability that a visiting female mated was related to display rate (Table 3), indicating that initial attraction and active choice are influenced by different components of display. In addition, inter-pop interval and display rate tended to covary inversely (Fig. 1), suggesting that attraction and active choice may impose conflicting selection pressures on display performance. Received: 11 November 1995/Accepted after revision: 16 March 1996     

The short and long run effects of tied foreign aid on pollution abatement, pollution and employment: A pilot model

There exists in the literature a presumption that tied foreign aid can be used effectively to reduce cross-border pollution. Focusing, in contrast to the received literature, on the interaction between the public and private provisions of pollution abatement in the recipient country, we question the effectiveness of tied foreign aid in reducing pollution. In this context, we obtain many novel and policy relevant insights. Allowing for changes in labour employment and distinguishing between short and long run effects, tied foreign aid is shown to crowd out the private provision for pollution abatement in the short run. In the long run, tied foreign aid raises employment and therefore may be desirable for the recipient but undesirable for the donor country because it also raises pollution. The results change drastically if only the government provides pollution abatement.     

Female choice of large males in the treefrog Hyla chrysoscelis: the importance of identifying the scale of choice

Summary In this study, I detected the presence of female choice for larger males in the treefrog Hyla chrysoscelis. Mated males were larger than unmated males at the scales females could have assessed males: local groups (males found within 2 meters of a mated pair) and the nightly chorus. In addition, females were observed to initiate amplexus with the larger males in local groups. Mated males were larger than unmated males for two of the four seasons studied. Seasonal mating success was also analyzed with a multiple regression model that included size and chorus attendance (nights spent at the breeding site). The partial selection coefficients, which represent the relative magnitude of directional selection, were significant for size in only one season and for chorus attendance in all four seasons. Therefore, there are two possible advantages to being a large male: increased likelihood of obtaining a mate due to female preference, and increased likelihood of mating even if no female preference because of a greater number of opportunities to obtain a mating if matings occurred at random.     

Female choice for complex song in the European starling: a field experiment

Male European starlings Sturnus vulgaris sing long complex songs that appear to be important in the courtship of females but which also influence competitive interactions between males. We tested the hypothesis that females choose mates on the basis of the complexity of their songs, rather than on the quality of the territories the males defended. In order to determine whether certain territories were preferred over others, the first set of birds to settle in the experimental nest-boxes was removed and a second set allowed to settle. Consistent preferences for certain nest-boxes were indicated by correlations between the settlement patterns of the first and second sets of birds. However, males with the most complex song did not necessarily occupy the most preferred nest sites. Males with more complex song acquired mates faster. This relationship remained significant when nest-site preference was statistically controlled, indicating that female starlings chose males with complex song rather than those that defended preferred nest sites. A number of morphological variables were also found to be uncorrelated with female choice. Song complexity in European starlings increases with age, and the evolution of song complexity in this species is consistent with an age-indicator model of sexual selection. Males with larger repertoires were also in better condition, indicating that females obtain high-quality mates by choosing on the basis of male song. Received: 29 April 1995/Accepted after revision: 9 September 1995     

Female mate selection in the natterjack toad: active choice or passive atraction?

Summary Variation in seasonal mating success among male natterjack toads (Bufo calamita) was influenced by the number of nights that males spent at the pond and by male body size. Large males produced louder and lower frequency calls than small males, and maintained larger acoustic territories. After arriving at the pond, one half of all observed females mated with the first male they encountered. The remainder visited several males before initiating amplexus, but no criteria could be identified that females might have used when deciding whether to accept or reject a male. Movements between several males seem to be best explained by low female responsiveness to male advertisement calls on cold nights which were nonoptimal for oviposition. Females attempted to reject non-calling males both before and after amplexus, but this may be a mechanism to avoid mismating with males of the common toad (Bufo bufo), an explosive breeder that utilised the same pond. In two-choice playback experiments using synthetic advertisement calls, females showed no preference for calls based on their frequency. Females preferred calls of intermediate pulse repetition rate equivalent to those produced by a male at the same body temperature. Pulse rate is thus potentially subject to stabilishing selection and may be an important character for species-recognition. Females preferred fast rather than slow call rates, but only when the alternative rates were extreme. They also preferred calls which they perceived at the highest sound pressure level, but did not discriminate between absolute sound pressure levels of alternative stimuli at different distances. Since females that delay mating and oviposition may suffer predation, it is suggested that female preference for loud, rapidly repeated calls may be adaptive in the sense of minimizing the costs of locating conspecific males, rather than maximizing the probability of obtaining a high quality mate. Competition between males to maintain large acoustic territories and produce calls that can be easily detected by females would seem to be a sufficient mechanism to explain the evolution of the striking calls produced by male natterjacks.     

Female choice in the guppy (Poecilia reticulata): the interaction between male color and display

The effect of two components of male courtship, color and display behavior, on female choice of mates was investigated in the guppy (Poecilia reticulata). Computer-modified videos were constructed to determine the relative importance of a static trait, the presence or absence of carotenoid pigment (C and NC), and a dynamic trait, high and low display rate (HD and LD), on female response. Females were given a choice between all combinations of male display and color in a binary choice design. Preference was determined by the time females spent visually inspecting the animation. Females preferred animations with high display rates when both animations displayed color (CHD vs CLD), but not in the absence of color (NCHD vs NCLD). Equal numbers of females chose the color/low-display animation and the no-color/high-display animation when the two were paired. Conversely, color became a criterion of choice when both animations showed a low display rate (CLD vs NCLD), but not when both displayed at a high rate (CHD vs NCHD). These results suggest that females use both static and dynamic traits to evaluate males, but their rankings are affected by the choices available. Results of these experiments provide insights into how females use multiple traits to assess males.     

Innate movement rules in foraging bees: flight distances are affected by recent rewards and are correlated with choice of flower type

The non-random movement patterns of foraging bees are believed to increase their search efficiency. These patterns may be innate, or they may be learned through the bees’ early foraging experience. To identify the innate components of foraging rules, we characterized the flight of naive bumblebees, foraging on a non-patchy “field” of randomly scattered artificial flowers with three color displays. The flowers were randomly mixed and all three flower types offered equal nectar volumes. Visited flowers were refilled with probability 0.5. Flight distances, flight durations and nectar probing durations were determined and related to the bees’ recent experiences. The naive bees exhibited area-restricted search behavior, i.e., flew shorter distances following visits to rewarding flowers than after visits to empty flowers. Additionally, flight distances during flower-type transitions were longer than flight distances between flowers of the same type. The two movement rules operated together: flight distances were longest for flights between flower types following non-rewarding visits, shortest for within-type flights following rewarding visits. An increase in flight displacement during flower-type shifts was also observed in a second experiment, in which all three types were always rewarding. In this experiment, flower-type shifts were also accompanied by an increase in flight duration. Possible relationships between flight distances, flight durations and flower-type choice are discussed. Received: 20 November 1995/Accepted after revision: 10 May 1996     

Suitability of short or long conservation contracts under ecological and socio-economic uncertainty

Acquisition of land rights has become a primary tool used to protect terrestrial biodiversity. Fixed length contracts are often used when trying to secure conservation benefits on private land in agri-environment schemes and payment for environmental services schemes, but the duration of the conservation contracts used in different programmes varies. To date, very little research has been undertaken to determine the situations in which contracts of differing lengths are optimal or when conservation agencies or groups should use a portfolio of different contract lengths rather than relying on a single type. Using stochastic dynamic programming and related heuristic methods, we investigate how the choice between short or long conservation contracts is affected by uncertainty regarding the future availability of sites and their ecological condition. We also examine the benefits offered by using a portfolio of different contract lengths. Conservation agencies must pay private landowners a premium to secure longer agreements and because of this, shorter contracts are advantageous if sites are likely to remain available for conservation in the future. Long contracts are preferred when future site availability becomes more unlikely. In contrast to uncertainty over site availability, uncertainty over future ecological conditions has little effect on contract selection and only markedly influences the choice between short and long contracts when there is heterogeneity across sites in expected conservation outcomes and future availability of sites is also uncertain. Finally, when future site availability is unlikely, the use of a portfolio of short and long contracts would offer greater conservation gains than using either type in isolation, even though this option is not yet one that is commonly found in conservation practice.     

Individual- and condition-dependent effects on habitat choice and choosiness

Research on consistent individual differences in behavior, or “behavioral syndromes”, continues to grow rapidly, and yet, the aspects of behavior under consideration have remained remarkably limited. Here, we consider individual variation in consistency of choice (termed here “choosiness”), as expressed during habitat choice. We repeatedly tested the responses of female Western Black Widows, Latrodectus hesperus, to two cues of habitat quality: prey chemical cues and variation in web site illuminance. We estimated females’ response by the distance they positioned themselves from (1) the source of prey chemical cues and (2) the darkest edge of our test arena. Individuals with low variance in their responses are deemed more “choosy”, whereas individuals with high variance are deemed less “choosy”. Generally, most females initiated web construction near the source of the prey chemical cues and tended to place themselves in low-light conditions. However, we detected strong, repeatable differences in females’ intensity of response, and within-individual variance of response (i.e., choosiness) was correlated across situations: females with highly consistent responses towards cricket chemical cues also exhibited highly consistent responses towards variation in light conditions. When deprived of food for extended periods, females were indistinguishable in their responses towards prey chemical cues, but tended to initiate web construction in brighter lighting conditions. Food-deprived females universally exhibited higher variance and diminished consistency in their responses (i.e., they were less choosy). Additionally, higher choosiness was associated with greater mass loss during choice trials, suggesting choosiness is energetically costly. Our results demonstrate that consistency of response to environmental cues is yet another element of behavior that varies among individuals and variation in choosiness could beget speed/quality trade-offs during animal decision making.     

Female choice for parasite-free male satin bowerbirds and the evolution of bright male plumage

Summary Hamilton and Zuk proposed that bright male plumage may have evolved in males of polygynous species as a result of female preferences for males that are able to demonstrate their resistance to disease. They predicted an inverse correlation between female mating preferences and the level of parasitic infection of males. We found such a correlation between the level of infection by a common ectoparasite (Myrsidea ptilonorhynchi: Menoponidae) and mating success of male satin bowerbirds (Ptilonorhynchus violaceus). In addition, we tested and were able to confirm three other predictions derived from their model: that (1) older males had fewer parasites than their younger counterparts, (2) levels of individual parasitic infection are highly correlated between years, and (3) that individuals resighted in successive years are less parasitized than those that fail to return. These results support the bright male model, but they are also consistent with two other hypotheses that may explain plumage dimorphism based on the level of parasitic infection. The correlated infection model suggests that females choose males with few ectoparasites because of a correlation between the level of ectoparasitic infection and heritable resistance to internal infections. In the parasite avoidance model, females favor parasitefree males because it lowers their own prospects for parasitic infection. Our data did not show the predicted relationship between parasite numbers with plumage quality that is needed to support the bright male hypothesis, nor did it show the inverse correlation between male condition and parasite numbers that is predicted by both the bright male and correlated infection hypotheses. Our results are most consistent with the parasite avoidance hypothesis.     

Female choice and the quality of parental care in the great tit Parus major

Summary Previous work on great tits suggested that female mate choice was based on the characteristics of the male rather than the quality of his territory. The aim of this paper was to see whether females were deriving any direct benefits as a result of such choice by comparing male plumage colouration (the size of the central black breast-stripe) with the quality of parental care provided by both members of the pair. Males with large stripes were more likely to defend their broods because their nest attentiveness was higher increasing the chances of predator detection. Males with large stripes produced heavier fledglings primarily as a result of faster growth early in the nestling period. Previous work on great tits has shown that heavier fledglings have a higher survival probability than lighter ones. The results of this study can more easily be explained by differences in parental quality between males rather than as a result of variation in female or territory quality. Therefore, female great tits could increase their reproductive success by pairing with males with a large stripe because such males appear to be better quality parents. This suggests that female choice in the great tit may concern, at least in part, the quality of male parental care.     

Species-specific antipredatory behaviours: effects on prey choice in different habitats

Summary Prey species may use many different behaviours to avoid predation. In this study, the antipredator behaviours of juvenile roach (Rutilus rutilus) and juvenile perch (Perca fluviatilis) were studied in wading pools with three kinds of structural complexity: no structure, structure simulating vegetation and structure simulating bottom crevices. Predation experiments with piscivorous perch and habitat choice experiments with the prey were performed, and the foraging success and prey choice of the predators were related to the type of structure. Predator foraging success was lower in the vegetation than in the other treatments. In the absence of structure and with vegetation structure, predators preferred perch over roach, while the preference was reversed in the crevice treatment. Roach and perch differed in their antipredatory behaviours. Roach responded to the presence of predators by schooling, moving fast and remaining at the surface, and escaped from attacks by jumping out of the water. In contrast, perch moved more slowly, dispersed after attacks and tried to hide at the bottom. Perch always preferred the vegetation structure to the non-structured part of the pool, while roach showed preference for the vegetation structure only when predators were present. Roach never occurred in crevices, whereas perch used crevices when predators where present. Predator pursuit speed was lower in the vegetation structure than in the non-structured treatment, but prey escape speed was unaffected. The results suggest that both the quantity and quality of structural complexity interacting with species-specific antipredator behaviours are important for predator-prey dynamics. It is also suggested that the presence of structure can have substantial effects on the structure of North Eurasian fish communities, by affecting relative and absolute predation pressures from piscivorous perch on prey species. Correspondence to: B. Christensen     

Female choice of a non-bodily ornament: an experimental study of cichlid sand craters in Cyathopharynx furcifer

In many species of East African cichlids, males build sand craters or “bowers” to attract females for spawning. It is commonly assumed that these exaggerated sexual traits have a function similar to bodily ornaments. In non-bodily ornaments, however, the behaviour creating the structure may serve as an additional source of information to potential partners, providing multiple signals for mate choice. We tested whether and to what extent females use these signals for choosing males by observing the individual sampling behaviour of female Cyathopharynx furcifer in the field. In addition, we experimentally manipulated crater characteristics in the field and laboratory. We found that females spawn preferably with owners of large and well-maintained craters, but when crater size was enlarged or reduced, the resulting building activity of crater owners affected female choice more strongly than the manipulated crater size per se. We discuss the importance of multiple signals in species constructing extended phenotypes.     

Female mate choice,male migration,and the threat of infanticide in ringtailed lemurs

Summary Paternity determination by DNA fingerprinting is reported for a long-term study group of semi-free-ranging ringtailed lemurs (Lemur catta), together with behavioral data collected independently. In 1985, fraternal twin males unfamiliar and unrelated to the resident ringtailed lemurs were introduced to the forest enclosure. Every mature resident male attacked the immigrants frequently across the next 5 months, whereas no female ever did. All observed estrous females showed sexual proceptivity toward the' immigrant males; three solicited copulation exclusively from them. Each female repelled sons, matrilineal brothers, and other resident males from attempting to copulate. Over a 5-year period, four of five females always reproduced with distantly related or unrelated males (Fig. 3). Despite low dominance status throughout the case study, an immigrant sired the off-spring of each female that was proceptive toward only the immigrants, demonstrating that female choice can override male dominance relations to determine reproductive success among male ringtailed lemurs. In the birth season following the 1985–1986 immigration, each of four females targeted one or two particular adult males for consistent attack across the period of infant dependency, beginning days after parturition. Paternity determinations, colony records, and subsequent study of two groups allowed 66 cases of this mode of maternal aggression to be documented. In each, the targeted male had not fathered the protected infant, and almost invariably, he was unrelated to the infant's mother. New mothers attacked every male that immigrated following their infants' conceptions and a few familiar males with whom they had not been seen to copulate during the previous breeding season. Recent attempts by immigrant males to kill infants confirmed the anti-infanticidal function of maternal targeting of males. All results were interpreted together to advance a prospective model of the mating system of ringtailed lemurs. Female avoidance of incest has led to the evolution of natal male dispersal. Subsequently, males should prefer to transfer into groups containing few and/or status-vulnerable males. We predict that, by killing others' infants, males simultaneously increase chances for success in females' next reproductive efforts and terminate current fathers' reproductive eligibility in a group. Basic hypotheses that await testing are that (a) raising an infant through weaning reduces a female's chances for reproductive success the following year and (b) males that demonstrate the capacity to promote the survival of infant offspring are most attractive to females as mates.     

Female choice and the evolution of the conspicuous plumage coloration of monogamous male great tits

Summary The conspicuous male plumage coloration of many avian species is often regarded as the result of sexual selection through female choice. In general terms such plumage characters may evolve in monogamous species if males bearing them pair with high quality females and so reproduce more successfully than males lacking the character. Male great tits have a conspicuous, central black breast stripe which varies in size between individuals. The stripe is also present in the female although it is smaller in size. Male great tits with large stripes paired with females which laid large clutches. Furthermore, in one of three years, females paired with such males commenced breeding earlier in the season than other females. Individual females were significantly more consistent in their clutch size and laying date between years than were nesting boxes. Males with large stripes paired with females which had previously laid a large clutch. Although there was evidence that territory quality may affect female reproductive success by influencing nesting success and nestling quality, there was no significant relationship between the stripe size of a male and the quality of his territory. Therefore, the results suggest that female great tits are choosing the characteristics of the male rather than the quality of his territory. The evidence thus suggests that female choice may be important in the evolution of male secondary sexual characteristics in great tits.     

Male choice and competition in Tetraopes tetraophthalmus: effects of local sex ratio variation

Summary Male milkweed beetles (Tetraopes tetraophthalmus) are shown to exhibit choice between potential mates. Males, however, also exhibit direct competition for access to potential mates. These differences in male behavior are dependent on the local sex ratio, with males becoming increasingly choosy when the sex ratio is female-biased, and more competitive when the sex ratio is malebiased. Females show neither choice nor competition. In both the laboratory and in the field, body size is important in determining patterns of competition and choice. Larger males win most aggressive encounters, and males generally prefer large females as mates. The combination in a single sex, of competition for mates and mate choice is not predicted by classical sexual selection theory. Male choice and competition may have evolved in T. tetraophthalmus due to the combination of (1) the mating system (2) the pattern of variation in female fecundity, and (3) the spatially and temporally variable operational sex ratios. This explanation is not dependent on high heritability of body size.