What do pairing patterns in common tern, Sterna hirundo, recruits reveal about the significance of sex and breeding experience?

In general, reproductive output in long-lived bird species increases in older compared to younger individuals. Therefore, experienced mates should be advantageous for first-time breeders. To examine requirements and consequences of experienced pair mates we investigated the first pair bonds of common terns, Sterna hirundo, recruiting to their natal colony. We found that male recruits were usually the same age as their mates, whereas female recruits were usually the younger member of the pair. In order to acquire experienced mates, it was necessary for males to arrive early in the year of recruitment. Mates with 2 or more years of breeding experience were only attainable by male recruits characterised by greater body mass and age. Female recruits arrived more than 1 week later than their experienced mates and significantly later in the season than male recruits paired with experienced females. In general, females first bred at a younger age than males, and neither the female recruits body mass nor their age was related to the level of experience of their first mate. These sex-specific differences in obtaining an experienced mate did not result in different levels of reproductive success between the sexes. Male and female recruits with mates with 2 or more years breeding experience benefited from having experienced mates: they had greater reproductive success. First-time breeders paired with mates with only 1 year of breeding experience did not differ from pairs where both members were breeding for the first time in terms of reproductive success, but clutches were larger. Our results illustrate not only different prerequisites for males and females, but also males need for experienced mates. Delayed male breeding (postponing breeding for another year), supposed to be a negative trait, and high body mass, supposed to be a trait of superior individual quality, can be combined in some individuals, improving reproductive success and showing that breeding common terns use a range of tactics to begin reproduction.Communicated by F. Trillmich     

The “dawn chorus” and incubation in the coot (Fulica atra L.)

Summary It has been proposed that both acoustic and non-acoustic factors could explain the dawn peak of territorial advertisement in birds. The coot is a territorial waterbird which does not use vocal advertisement either to attract mates or defend territory boundaries. In spite of the absence of acoustic factors affecting the time budgeting of advertisement, this species shows a dawn peak of display and aggressive behaviours. This trend occurs both in the presence and absence of non-territorial prospecting birds, but the trends are quantitatively different in these two cases (Fig. 1). Males participate in display and fighting more often than females, probably because of their greater size and effectiveness in defence. This asymmetry in defence abilities governs the time budgeting of incubation, the male tending to take time off the nest when territorial defence is most necessary (Fig. 2).     

Kin-preferential cooperation,dominance-dependent reproductive skew,and competition for mates in communally nesting female house mice

Little is known about the behavioural mechanisms facilitating kin-preferential communal breeding in wild house mice (Mus domesticus). We evaluated the effect of kinship and male availability on aggression, social structure and reproductive skew in groups of female mice freely interacting and reproducing in semi-natural indoor enclosures. Triplets of either sisters or non-sisters were established in enclosures provided with either one or three littermate males, which were unrelated and unfamiliar to the females. Sisters were more spatially associated and less aggressive than non-sisters, leading to higher incidences of communal breeding and reproduction. This is in agreement with theoretical considerations on kin selection in house mice. Reproductive success was highly skewed in favour of dominant females due to subordinate infertility or complete loss of first litters, which might have been caused by dominant females. In spite of this, subordinates only rarely dispersed from the enclosures, suggesting that perceived dispersal risk generally outweighed relatively reduced reproductive potentials. Aggression levels among females were significantly higher when one male was available, compared to when three males were available. We suggest that this might result from higher female-female competition for mates, due to the risk of missing fertilisation when synchronously oestrous females encounter limited numbers of males in a deme. Our results indicate that, first, communal nursing in house mice might have evolved to make the best out of a bad job rather than to enhance offspring fitness; and, second, that female-female mate-competition might play an important role in shaping female social structure in this polygynous mammal.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by F. Trillmich     

The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

Notes on some factors affecting aggressive behaviour in Carcinus mediterraneus

The relationships between aggressive behaviour and animal density and between aggressive behaviour and metabolic excretion have been investigated in Carcinus mediterraneus Czerniavsky. Increasing the crab density by increasing the number of crabs or reducing the available area revealed that the aggressive interactions perindividual did not increase. The frequency of interactions reached a maximum in several minutes following the introduction of crabs. Individuals isolated between 6 and 24 h were more aggressive than non-isolated ones. The water in which a group of crabs had been held had an inhibitory effect (not abolished by commercial aquarium filters) on aggressiveness of pairs of crabs grater than plain water or water in which single crabs had been kept. No home-group effect was observed. The effect of group water was enhanced by increasing the concentration of crabs in the duration of confinement, and it is probably due to the accumulation of catabolites in the medium. Crabs kept in groups had a greater excretion than isolated crabs and the nitrogen oupput decreased with an increase in the number of crabs or whith a decrease in the size of the tank (with a constant volume of water and number of crabs).     

Divorce in common murres (Uria aalge): relationship to parental quality

Behavioral precursors of 12 divorces were examined in 30 color-banded pairs of common murres (Uria aalge) over six breeding seasons. Common murres are long-lived seabirds that typically return each year to the same mate and nest site in dense colonies. At least one parent is present continuously from egg lay to chick fledging. Murres, therefore, have considerable opportunities to compare their mates parental behavior with that of several familiar neighbors. Previous reproductive success was lower for divorcing birds than for reuniting pairs. As predicted by the better option hypothesis, there were clear choosers (seven females and five males) that initiated divorce by moving to a new birds site or by courting a new partner at their current site. Choosers improved their reproductive success after the divorce, whereas their previous partners, the victims did not. Yearly divorce rates (average 8.2% per year) were significantly correlated with yearly mortality rates. Divorces appear to be opportunistic: pairs divorced after varying numbers of reproductive failures with the immediate precursor usually being the disappearance (death) of a murre from a successful neighboring site. In contrast to the delays experienced by victims, choosers formed new pairs quickly and laid their eggs no later than reuniting pairs. Prior to the divorce, victims fed their chicks less often than choosers, and some engaged in other behaviors that compromised egg or chick survival. These observations suggest that deficiencies in parental behavior were precursors to the divorce. This report is one of the first cases where reproductive failure of divorcing pairs has been linked to deficits in the parental behavior of the subsequent divorce victim.Communicated by C. Brown     

The establishment of foraging flocks in house sparrows: risk of predation and daily temperature

Summary The foraging decisions of animals often reflect a trade-off between the risk of predation and efficient foraging. One way an animal may reduce the risk of predation, and hence exploit a resource patch in relative safety, is by foraging in a group. Solitary pioneer sparrows often recruit others to a food source by making chirrup calls in order to establish foraging flocks. This study describes the decisions of house sparrows that arrive at food resources of different risks of predation. Four feeding sites at different distances from a perching site and from an observer were presented to sparrows. When the feeder was adjacent to the perching site and far from the observer, the pioneers chirruped less frequently and were more likely to forage alone than when the feeder was in the other three positions. There were differences in the scanning behaviour of sparrows at these sites, suggesting that they were responding to different risks of predation. Furthermore, the chirrup rates of pioneer sparrows in this study and a previous study were found to be negatively correlated with maximum daily temperature. This is consistent with the hypothesis that energy requirements may affect the flock establishment decisions of sparrows, and that the benefits of foraging in flocks may be greater at lower temperatures.     

Comparisons of the feeding activity and diets of male and female cunners Tautogolabrus adspersus (Pisces: Labridae)

The diel distribution of feeding activity during the spawning season was compared for territorial male and female cunners Tautogolabrus adspersus in Conception Bay, Newfoundland, Canada between June and August, 1978. Territorial males feed significantly less often than females and concentrate their feeding activities in the morning, whereas females feed as frequently in the afternoon as in the morning. Based on the contents of alimentary tracts, territorial males, non-territorial males, and females have different diets. Since these three groups share the same habitat, the observed dietary differences probably reflect differences in their foraging behaviour. The relevance of these findings to the concept of time minimizer and energy maximizer is discussed.     

No evidence for acoustic mate-guarding in duetting buff-breasted wrens (Thryothorus leucotis)

There are few empirical tests of the acoustic mate-guarding hypothesis for the function of duetting in birds. This hypothesis states that when females are fertile, males initiate many songs or answer most of their mates solo songs to form duets and repel rival males seeking extra-pair copulations. We tested the hypothesis by comparing song initiation and answer rates of males and females in socially monogamous buff-breasted wrens (Throthorus leucotis) during pre-fertile and fertile periods. During pre-fertile periods, males often sang for short periods before being answered by their mates, yet first duets were formed earlier relative to dawn and more duets were given during the dawn chorus on pre-fertile than fertile mornings. Males initiated more songs during pre-fertile than fertile periods, whereas there was no difference between stages in female song initiation rates. The proportion of songs answered by individuals of both sexes did not differ between breeding stages. Other mate-guarding behaviours, such as frequent copulation and maintaining close proximity to mates when fertile, did not appear to be important in this species, as no copulations were observed and there was no difference in the time pairs spent in close proximity when females were fertile or not. Parentage analysis revealed that only 3% of 31 broods had young that were likely the result of extra-pair paternity. These findings do not support the acoustic mate-guarding hypothesis, and suggest that the low rate of extra-pair paternity in buff-breasted wrens was maintained without the use of acoustic or traditional paternity guards.Communicated by R. Gibson     

The conflict between feeding and territorial defence in the great tit

Summary A field experiment investigated whether feeding and territorial defence competed for time in the activity budget of territorial male great tits during the spring. Feeding tables were placed on the territories of five males. Five other territorial males also fed at these tables, while ten additional males had no access to any feeding tables. A standardised intrusion using playback of territorial song and a stuffed mount of a male great tit was conducted on each of the twenty territories, and the response of the resident male measured. All of the males with access to a feeding table responded more vigorously to the intrusion than the males who received no extra food. These results are not attributable to the fact that males defended their territories more vigorously because of the addition of feeding tables, since males that gained extra food outside the boundaries of their own territories also defended more vigorously than birds with no access to a feeding table. The results support the conclusion that the provisioned males defended more vigorously because they could afford to take more time out from feeding.     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Female-enhanced male competition determines the first mate and principal sire in the spider Linyphia litigiosa (Linyphiidae)

Summary Two to five days before sexual maturation, female sierra dome spiders (Linyphia litigiosa: Linyphiidae) undergo a transformation in their behavior toward males that visit their webs. During this latter part of their penultimate instar, females change from consistently positioning themselves far away from males to actively maintaining close proximity, reactions I call avoidant and associative behavior, respectively. Consistent associative behavior ceases after the female's first mating and thus is limited to soon-to-mature penultimate females. When a mate-seeking male fords an associative female, he attempts to guard her until she matures; this is often a multi-day affair. In contrast, males guard immature avoidant and mature mated females for only a single day. This dichotomy in male guarding times can be understood by the fact that associative behavior signals that the female will soon develop peak reproductive value. Upon completion of their final molt, 98% of females immediately mate with the current guarding male. Secondary suitors are not as likely to achieve mating. Moreover, first mates father 1.8 times more offspring, on average, than secondary mates. Whenever they meet on any female's web, males fight until one of the contestants withdraws. Fights typically are intensive, occasionally deadly, and often result in usurpation of the web by the newly arriving male. Larger males win more fights, but other qualities (e.g., vigor and persistence) appear to be important when contestants differ by less than 10–20% in body weight. Prolonged (i.e., multi-day) guarding of associative females enhances the intrasexual selection process by ensuring that every male that arrives at the web finds it already guarded. Therefore every male that finds the web becomes a participant in a series of male-male conflicts and web usurpations which span the period between the resident female's commencement of associative behavior and her sexual maturation. Since unforced male departures from the webs of associative females are rare, victors are retained on the web until they themselves lose a fight. This facilitates a steady increase in the fighting ability of sequential guards throughout the associative period, up until female maturation and mating. On my study site, first mates represented the final winners in a combative sorting process based on a minimum average of 2 fights; they were heavier and larger than secondary mates and randomly sampled males. The combination of (1) associative behavior by nearly mature females, (2) high mating propensity of newly mature females, and (3) first male sperm priority, constitutes a system whereby females enhance male-male competition and boost the expected fighting prowess of the principal sire of their progeny. Since males appear to make no material contribution toward progeny, the female's behavior probably functions to improve the genetic constitution of the offspring. In addition, the timing of associative behavior may limit prolonged guarding by food-stealing males to a period (1) encompassing the female's pre-molt fast and (2) before the heavy yolking of eggs, thereby ameliorating the nutritional costs of intrasexual selection.     

Conditional lekking in ruff (Philomachus pugnax)

Summary In our study of ruff, a lekking shorebird, we found that the lek ratio — the proportion of the total population of males occurring on leks was low, averaging 12% over the breeding season (Fig. 1D). Off-lek males spent approximately the same proportion of time as lek males in displaying to females (Fig. 4). We defined three spacing tacties that male ruffs use to position themselves to court females: Following — directly pursuing females, and two types of lekking behavior, Intercepting — waiting for females at resource-rich sites, and True Lekking — waiting for females at places without any resources other than the males themselves. Males switched among these tactics, causing the lek ratio to vary over the season (Fig. 1 D). Lek ratio increased when the number of females present in the study area plummeted at the end of May, and was positively correlated throughout the season with the copulation rate of the preceding day, suggesting that males were tracking the behavior, as well as the number, of females available (Table 1, Fig. 1). Early in the season, males off leks spent most of their time feeding (Fig. 4), and lek ratio was positively correlated with temperature (Table 1), suggesting that some males may have been unable to lek during cold weather. Males on leks mated at significantly higher rates than Followers (Table 4). On average, males at our interception lek were less site faithful and less peristent than males at true leks, and the interception lek itself disappeared after females stopped coming to use its adjacent resource (Table 2, Fig. 5). The most successful individuals in our population were the True Lekking males, rather than the Interceptors.In addition to the conditinal lekking tactics described here, ruff display a dimorphism in behaviorat leks. Independent males defend small courts on leks, while Satellite males share courts and mutually display with independents Both independents and satellites may use all three conditional tactics. We propose that satellites evolved as specialized. Followers, adept at tracking the movements of females among leks.     

Feeding deterrence properties of apo-fucoxanthinoids from marine diatoms. I. Chemical structures of apo-fucoxanthinoids produced by Phaeodactylum tricornutum

Methanol extracts of freshly harvested cells of the marine diatom Phaeodactylum tricornutum were found to produce a feeding deterrent effect in the copepod Tigriopus californicus. Bioassay guided fractionation of the methanol extracts led to the isolation of four compounds possessing feeding deterrent activity. The compounds were identified as apo-10-fucoxanthinal (1), apo-12-fucoxanthinal (2), apo-12-fucoxanthinal (3), and apo-13-fucoxanthinone (4) by detailed spectroscopic analysis and comparison with authentic compounds produced semi-synthetically from fucoxanthin. Compounds 1 to 4 exhibited feeding deterrent responses in T. californicus at concentrations of less than 20 ppm.     

Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Extra-pair copulations in a lek: the secondary mating system of monogamous razorbills

Summary I studied the mating strategies of razorbills Alca torda, a monogamous colonial seabird, during 1987–1989 in Wales, U.K. The outstanding feature of the study population's mating system was the performance of most copulations in mating arenas outside the colony. Females visited the arenas and sometimes accepted extra-pair copulations (EPCs), and males aggressively competed for EPCs. During the fertilizable period of most females, the arenas resembled leks because they contained no resources except mates, and males contributed only sperm to extra-pair females. Typical of leks, the operational sex ratio was strongly male-biased, and male mating success was highly skewed. To examine the potential for sexual selection to operate in this system, I correlated male EPC success with variables that could be associated with male-male competition and/or female choice. The frequencies of three behaviors (interference of copulation attempts, fights won against other males, and EPC attempts), accounted for 62% of the variance in EPC success and were strongly intercorrelated, with interference explaining most of the variance in a multivariate test. EPC success was not correlated with body size, age, paired status or either of two estimates of the time spent in the arena. The hypothesis that male EPC success was determined by active female choice was tested by examining the relative success (EPCs per attempt) of males. Relative success was not correlated with any of the three behaviors associated with absolute success, or with age, body size or attendance, suggesting that male-male competition, mainly in the form of copulation interference, is the principal correlate of EPC success. These findings in a monogamous species illustrate that EPC is a secondary mating system of razorbills in which sexual selection operates. The existence of this system outside the colony raises the possibility that lekking for EPCs may also occur within the nesting territories of other monogamous, colonial species but is hidden by competition for nests and breeding partners.     

Unattractiveness of mated females to males in the parasitoid wasp Spalangia endius

Despite common stereotypes, males are not always indiscriminate and eager when it comes to mating. In the parasitoid wasp Spalangia endius, the initial response of males to females was almost always one of apparent excitement; however, this was followed by a clear preference for virgin females over mated females in both no-choice and choice situations. The no-choice data were collected from videotapes of male-female pairs of all possible combinations of mated and virgin individuals. Neither female nor male mating status had a significant effect on likelihood of, or time until, contact or male courtship fanning. However, a males first retreat was sooner when the female was mated than when she was virgin; mated males exhibited their first retreat sooner than did virgin males; and mated females were less likely to be mounted than were virgin females. In addition to the videotapes, male-choice experiments were performed. When given a choice of a virgin and a mated female, both virgin and mated males were more likely to mount and copulate with the virgin. The difference in response to virgin versus mated females seemed to be less in virgin males than in mated males, perhaps due to virgin males greater eagerness to mate: when a virgin male and a mated male were presented with a dead virgin female, the virgin male was usually the first to respond to the female. That males preferentially retreated from and avoided mounting mated females appears to be adaptive given that mated females rarely copulated.Communicated by N. Wedell     

An advantage to mate choice in the seaweed fly,Coelopa frigida

Summary All natural populations studied of the seaweed fly, Coelopa frigida, are polymorphic for the inversion system on chromosome I. It has previously been shown that heterokaryotypes have superior egg-to-adult survival than either homokaryotype, and that there is non-random mating with respect to inversion genotypes. We have inquired whether the production of fitter progeny in larger numbers is the possible reason for the existence of non-random mating behaviour. We measured the fertility, fecundity and progeny survival of adults that have either been allowed to exercise mate preference, or been given a single, randomly chosen, mate. The fertility (% of females producing progeny) and progeny survival (% of larvae surviving to the late 3rd instar stage) are both significantly greater in choice than nochoice females. There are also significant differences in fertility and progeny survival depending on whether the parents were of similar or dissimilar inversion genotypes. An effect of relative adult size on fertility was also observed. There appear to be no effects of mate choice on fecundity. Indirect evidence is presented that adults are choosing their mates not only on the basis of genotypes but on other criteria as well. It is suggested that the superior reproductive performance of animals allowed a choice of mates may account for the evolutionary origin and continued existence of mate discrimination. Furthermore, since over 25% of the genome of Coelopa is located within inversions, the observed pattern of negative assortative mating may maintain a substantial fraction of the genes in a polymorphic state.     

Social and ecological influences on activity budgets of vervet monkeys,and their implications for group living

Summary Activity patterns were documented over a 20-month period in six groups of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya. Group size varied both among groups and within groups through time. The home ranges of two groups were located in Acacia xanthophloea habitat whereas those of four adjacent groups were located in A. tortilis habitat. Repeated measures analysis of variance, three-way analysis of variance, and meta-analysis of group size effects were carried out on feeding, moving, resting, scanning, allo-grooming, and other behavior. Time spent feeding varied seasonally, peaking during periods when A. tortilis seeds were eaten. However, time spent feeding did not vary within or among groups, or between habitats, males and females, or dominants and subordinates. Males spend more time scanning and less time allo-grooming than females, and high-ranking individuals of both sexes spent more time scanning than low-ranking individuals. Among females, subordinates spent more time moving, whereas among males, dominants spent more time moving. Groups living in A. xanthophloea habitat spent more time moving and allo-grooming than groups in A. tortilis habitat but this may have been confounded by the fact that these two groups were on average larger than the others. Significant differences in activity budgets among groups demonstrate that activity patterns observed in one group are not always readily generalized to other groups even when they come from the same population. Group size analyses examined potential costs and benefits of group living. Increased intragroup competition, measured by time spent moving and feeding, was only weakly evident in larger groups. Individuals in larger groups allo-groomed more than individuals in smaller groups. Self-grooming also increased in larger groups. Individuals in larger groups spent less time scanning than individuals in smaller groups, but there was more scanning per group in larger groups than in smaller groups. Correspondence to: L.A. Isbell