Old endemics and new invaders: alternative strategies of passerines for living in the Australian environment

Life history parameters of ”old endemics” and ”new invaders” among the Australian passerines were compared, controlling for phylogeny by using current, phylogenetic research methods. Old endemics were found to have significantly smaller clutches and demonstrated significantly more cooperative breeding than the new invaders, whose fledging periods are significantly longer than those of the old endemics. We argue that breeding under conditions of a small annual food increment during the breeding season, as is probably the situtation in Australia, is possible only when clutch size is small, or while breeding cooperatively, or by extending the fledging period. Our data show that the old endemics use the first two options, while the new invaders employ the third. Received: 10 February 1999 / Received in revised form: 10 November 1999 / Accepted: 20 November 1999     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000     

Parental care of replacement clutches in common terns (Sterna hirundo)

In long-lived seabirds with low annual reproductive output, the renesting decision after breeding failure is critical, and the parents have to weigh benefits of replacement clutches against possible future reproductive costs. In this study, we investigated factors influencing renesting decisions in common terns (Sterna hirundo) and compared aspects of breeding biology and body mass between two breeding attempts by the same pairs in each of 4 years of heavy losses due to predation. Renesting birds were characterized by early laying dates and by a high age. Among early breeders, high egg mass reduced the probability of renesting. A long relaying interval coincided with low mass of replacement eggs in one year, and short intervals with high egg mass in another. Further, egg mass decreased and relaying intervals increased the later the predation events occurred. Evidence of high levels of parental care of replacement clutches came from body mass data: female mass increase prior to egg laying was higher in the second attempt than in the first, whereas male mass was lower during the second courtship period than during the first. Male mass also affected relaying intervals and mass of replacement eggs. We conclude that common terns expend high levels of parental care of replacement clutches. Intrinsic factors related to individual quality (age, body condition) seemed most important for renesting decisions and for the degree of parental care, but foraging conditions seemed to have modifying effects. Received: 13 August 1999 / Received in revised form: 5 February 2000 / Accepted: 13 March 2000     

Length of feeding day and body weight of great tits in a single- and a two-predator environment

The risk of predation may influence the acquisition of energy and the feeding activity of animals. Feeding activity and body reserves of wintering great tits Parus major in response to the priority to food access were studied in two areas differing in incidence of predators. The one-predator area contained sparrowhawks Accipiter nisus only, whereas the two-predator area contained both sparrowhawks and pygmy owls Glaucidium passerinum, whose hunting periods overlap at dawn and dusk. In the two-predator area dominant great tits arrived at feeders significantly later in the morning, and left earlier in the evening, than their subordinate flock-mates. Hence, feeding day length of dominants was found to be significantly shorter. The reverse was true for the one-predator area. In addition, dominants carried significantly greater reserves than subordinates in the area inhabited by two predators. Factors constraining subcutaneous energy reserves were also studied in removal experiments. After the removal of dominant individuals, subordinate great tits did not reduce their body reserves in the two predator area. In contrast, subordinate great tits significantly reduced evening body reserves in the single-predator area. I concluded that the presence of the two predators increases unpredictability in feeding conditions for great tits. Dominant individuals responded to this by shortening their feeding day and increasing body reserves at dusk. Received: 8 December 1999 / Received in revised form: 15 March 2000 / Accepted: 31 March 2000     

Testes mass and reproductive mode of minnows

Most comparative analyses of relative testes mass find that testes are larger in species in which more sperm competition is predicted (multiple males mate with individual females). I tested for differences in adjusted testes mass (for body mass) by spawning mode and by sexual size dimorphism in a comparative analysis of 37 minnow species. No significant differences were found for testes mass by spawning mode or sexual size dimorphism. These results imply a lack of response to selection on testes size from sperm competition in minnow species. Possible explanations for the lack of the expected relationship between testes mass and mating systems in minnows are presented. Received: 8 November 1999 / Received in revised form: 27 January 2000 / Accepted: 13 February 2000     

Effects of paternal care on reproductive success in the polygynous spotless starling Sturnus unicolor

For males of socially polygynous avian species like the spotless starling, there may exist a trade-off between investing in paternal care and controlling several nests. To determine how the intensity of paternal care affects reproductive success per brood sired or expressed as the total number of young raised in all nests controlled by the same male, it is necessary to manipulate paternal care. Testosterone (T) has been shown to depress the tendency for males to care for their young, and induces them to acquire more mates. The effects of paternal care on reproductive success were studied by treating certain male starlings with exogenous T and others with the antiandrogen cyproterone acetate (CA), and comparing the parental behavior of T- and CA-males throughout the breeding season with that of controls. CA-males fed their chicks more during the first week after hatching than T-males, with controls feeding at intermediate rates, both on a per nest basis and as total effort for all nests controlled by the same male. Paternal feeding rates during the first week of chick life had a significant positive effect on the number of fledged young. The hormone treatment significantly affected the number of chicks raised per nest, CA-males having a higher breeding success per nest than T-males, and controls showing intermediate levels of success. There was no significant effect of treatment on total reproductive success attained by males throughout the season. In the polygonous spotless starling, the intensity of paternal care of young affects reproductive success per nest positively but not on a seasonal basis. Received: 6 February 1999 / Received in revised form: 30 June 1999 / Accepted: 11 July 1999     

Weight comparisons and litter size manipulation in Columbian ground squirrels (Spermophilus columbianus) show evidence of costs of reproduction

I studied reproductive costs in the female Columbian ground squirrel (Spermophilus columbianus) using individually marked animals. I compared weight changes during the active season and over winter, and mortality for females that did and did not wean young. Females raising young were heavier at emergence in that spring than unsuccessful ones. Females that did not raise young gained more weight during summer, were heavier than successful females at the time of entry into hibernation, and were heavier emerging from hibernation the following spring. Over-winter mortality was higher for females that reared young compared to reproductively unsuccessful females. A food supplementation experiment showed that energy-rich food can accelerate individuals’ weight gain. Interactions between litter size, birth weight, weight at emergence from the natal burrow, survival of young to yearling age, and maternal fitness were also studied. Litter sizes were experimentally manipulated to evaluate how females cope with costs of rearing one additional young. Birth weight of juveniles was positively correlated with survival to emergence from the natal burrow and with survival to yearling age. Partial litter loss was higher in experimentally enlarged litters than in either experimentally decreased or control litters. Total litter loss, survival of adult females or the probability of weaning young the following year were not affected by the litter size manipulation. Females appear to adjust the size of their litter before birth, and to some extent during lactation, to their ability to wean young. Received: 20 January 2000 / Received in revised form: 12 March 2000 / Accepted: 18 March 2000     

Testosterone and the allocation of reproductive effort in male house finches (Carpodacus mexicanus)

Testosterone has been proposed to serve as the mediator that controls the relative effort that an individual male bird will devote to mating effort versus parental effort. Here, we demonstrate a testosterone-influenced trade-off between parental and mating efforts in male house finches. Male house finches with experimentally elevated testosterone fed nestlings at a significantly lower rate, but sang at a higher rate than males without manipulated testosterone levels. Females mated to testosterone-implanted males fed nestlings at a significantly higher rate than females mated to males without testosterone implants, resulting in similar feeding rates for both treated and untreated pairs. The effects of testosterone on male house finches, however, were not as dramatic as the effects of testosterone observed in some other socially monogamous species of birds. Because extra-pair copulations are uncommon in house finches and males provide substantial amounts of parental care, these more modest effects may be due to differences in how the allocation of reproductive effort affects the costs and benefits of different reproductive behaviors. Received: 6 June 2000 / Accepted: 17 July 2000     

Evolution of male mating behavior: male spadefoot toads preferentially associate with conspecific males

The evolution of male breeding aggregations is difficult to explain because males may reduce their reproductive success by associating with their closest competitors. We examined aggregative behavior by male New Mexico spadefoot toads, Spea multiplicata, which form breeding choruses in rain-filled pools. We specifically asked whether males are attracted to conspecific calls and, if so, whether they preferentially associate with those male calls that are also attractive to females. Field observations revealed that males showed significant clustering with conspecifics within breeding ponds, whereas laboratory phonotaxis experiments revealed that males preferentially associated with conspecific male calls. Moreover, when males were presented with conspecific calls that differed in call rate, smaller males associated with the stimulus preferred by females (average call rate). Thus, males appear to evaluate the attractiveness of competitors using the same trait employed by females to assess potential mates, and males adjust their positions relative to competitors depending on their size. We discuss these results in the light of several current hypotheses on the adaptive significance of male breeding aggregations. Received: 20 December 1999 / Accepted: 18 March 2000     

An analysis of reciprocal exchange of helping behavior in the red-cockaded woodpecker (Picoides borealis)

We examine the frequency of reciprocal exchanges of helping behavior in three red-cockaded woodpecker populations to determine if such exchanges might constitute a significant fitness benefit of helping. Specifically, we determine how often helpers, once they become breeders, are assisted by young they previously helped raise. The estimated frequencies of reciprocal exchange of helping based on survival and status transition probabilities are low (2%). Observed frequencies of reciprocity are significantly higher than estimated frequencies in two of the three populations studied, suggesting that male fledglings more often remain on their natal territory as helpers if another helper is already present. High rates of retention of young males as helpers on high-quality territories or preferential helping of kin, as well as preferential helping of former care-givers, might explain this result. When the analysis is restricted to helpers unrelated to the young they help raise, which controls for preferential helping of kin and largely eliminates effects of territory quality, empirical estimates of the frequency of reciprocity do not differ from estimates calculated from population demography. We conclude that young males do not preferentially help former care-givers and that reciprocal exchange of help is not an important factor in the evolution of helping behavior in the red-cockaded woodpecker. In this species, all known benefits of helping behavior are indirect (i.e., derived through kin selection ) rather than direct. We discuss the occurrence of unrelated helpers in this context. Received: 10 August 1999 / Received in revised form: 13 February 2000 / Accepted: 19 February 2000     

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.     

Cleaner-client interactions on a Caribbean reef: influence of correlates of parasitism

We investigated the influence of known correlates of parasitism, namely fish density, body size and social behaviour, on three highly variable aspects of the interactions between cleaning gobies (Elacatinus spp.) and their clients, on a Barbadian coral reef. We specifically considered (1) variability in client visit rate to cleaning stations, (2) cleaning goby preference for specific clients and (3) variation in the time spent cleaning by cleaners. Using phylogenetically independent contrasts, we found that client species that were abundant on the reef visited cleaning stations more often than rarer client species. This could be due to the positive relationship between ectoparasite load and client density, or alternatively may simply reflect the frequency of contact between cleaners and clients. Cleaning gobies spent more time cleaning large-bodied clients, which usually have higher ectoparasite loads, although cleaning goby preference for clients was influenced by none of the correlates of client parasitism. Overall, factors assumed to correlate with ectoparasite load had a limited influence on the variability observed in the interactions between cleaning gobies and their clients. Received: 27 October 1999 / Received in revised form: 11 January 2000 / Accepted: 24 January 2000     

Experimental evidence of a testosterone-induced shift from paternal to mating behaviour in a facultatively polygynous songbird

Previous studies have suggested that testosterone (T) profiles of male birds reflect a trade-off between mate attraction behaviours (requiring high T levels) and parental care activities (requiring low T levels). In this study, we experimentally elevated T levels of monogamous males in the facultatively polygynous European starling (Sturnus vulgaris), and compared mate attraction and paternal behaviour of T-treated males with those of controls (C-males). T-males significantly reduced their participation in incubation and fed nestlings significantly less often than C-males. Females paired to T-treated males did not compensate for their mate’s lower paternal effort. The observed reduction in a male’s investment in incubating the eggs was accompanied by an increased investment in typical female-attracting behaviours: T-males spent a significantly higher proportion of their time singing to attract additional females. They also occupied more additional nestboxes than C-males, although the differences just failed to be significant, and carried significantly more green nesting materials into an additional nestbox (a behaviour previously shown to serve a courtship function). T-males also behaved significantly more aggressively than C-males. During the nestling period, the frequency of mate-attracting behaviours by T-treated and control males no longer differed significantly. Despite the reduced paternal effort by T-males and the lack of compensation behaviour by females, hatching and breeding success did not differ significantly between T- and C-pairs. Received: 7 February 2000 / Revised: 10 August 2000 / Accepted: 3 September 2000     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

The formation and maintenance of crayfish hierarchies: behavioral and self-structuring properties

Behavioral changes in fighting and the development of dominance relationships were analyzed in groups of juvenile crayfish (Astacus astacus) using quantitative behavioral techniques. When individuals were placed into an aquarium, the number of agonistic challenges, their mean duration, and maximum intensity reached were high initially but then decreased steadily as the hierarchy developed. In all groups, linear hierarchies emerged which became increasingly stable over time. Winning influenced subsequent fighting behavior on two distinct time scales. In the short term, recent winners became progressively less likely to retreat. Second, individuals occupying dominant positions for days became increasingly likely to escalate to higher intensities early in the encounter. Both effects biased the outcome of future interactions such that winning enhanced further success and losing decreased an individual’s subsequent chances for dominance. Received: 11 November 1999 / Revised: 1 March 2000 / Accepted: 3 May 2000     

The modulation of worker behavior by the vibration signal during house hunting in swarms of the honeybee, Apis mellifera

During house hunting, honeybee, Apis melli- fera, workers perform the vibration signal, which may function in a modulatory manner to influence several aspects of nestsite selection and colony movement. We examined the role of the vibration signal in the house-hunting process of seven honeybee swarms. The signal was performed by a small proportion of the older bees, and 20% of the vibrating bees also performed waggle dances for nestsites. Compared to non-vibrating controls, vibrating bees exhibited increased rates of locomotion, were more likely to move into the interiors of the swarms, and were more likely to fly from the clusters and perform waggle dances. Recipients responded to the signal with increased locomotion and were more likely than non- vibrated controls to fly from the swarms. Because vibration signals were intermixed with waggle dances by some vibrators, and because they stimulated flight in recipients, the signals may have enhanced nestsite scouting and recruitment early in the house-hunting process. All swarms exhibited increased vibration activity within 0.5–1 h of departure. During these final periods, numerous vibrating bees wove repeatedly in and out of the clusters while signaling and motion on the swarms increased until it culminated in mass flight. The peaks of vibration activity observed at the end of the house-hunting process may therefore have activated the entire swarm for liftoff once a new nestsite had been selected. Thus, the vibration signal may help to integrate the behavior of numerous groups of workers during nestsite selection and colony relocation. Received: 17 January 2000 / Received in revised form: 5 April 2000 / Accepted: 3 May 2000     

The influence of potential reproductive rate and variation in mate quality on male and female choosiness in the sand goby, Pomatoschistus minutus

Recent models of choosiness in mate choice have identified two particularly important factors: the potential reproductive rate (PRR) of the choosing sex relative to that of the chosen sex, and the variation in quality of potential mates. This experimental study tested how these factors affected choosiness in male and female sand gobies, Pomatoschistus minutus. We manipulated relative PRR by means of water temperature, and mate quality by means of body length. The choosing male or female was offered a choice between two mates with either a small or a large difference in body length representing a small or a large variation in mate quality. Choosiness was measured as (1) preference for the larger mate, and (2) as whether or not spawning occurred with the smaller mate, while the larger mate was visible but screened off. We found that females preferred large males, and that their level of choosiness was affected by variation in male quality, but not by their own relative PRR. Males, on the other hand, seemed unselective in all treatments and were in general more likely than females to spawn with their provided partner. This suggests that in the sand goby, variation in male mate quality has a greater influence than relative PRR on facultative changes in female choosiness. However, a general difference in PRR between males and females may be one important factor explaining the observed sex difference in choosiness. Received: 17 April 2000 / Revised: 24 June 2000 / Accepted: 17 July 2000     

Carotenoids in the egg yolks of collared flycatchers (<Emphasis Type="Italic">Ficedula albicollis</Emphasis>) in relation to parental quality,environmental factors and laying order

Birds may influence the fitness of their offspring by transmission of different amounts of carotenoids to their eggs. Carotenoids play crucial roles in antioxidant protection and immune defence mechanisms, but they may be available to females in limiting amounts. Therefore, their allocation to the eggs may be influenced by the female’s condition, age and environmental circumstances. Furthermore, the quality of the male parent, which affects the reproductive value of the offspring, may also influence this investment. In this correlational study, we investigated proximate and ultimate factors that may lead to variation in yolk lutein, zeaxanthin and β-carotene concentrations among and within clutches of a wild passerine, the collared flycatcher (Ficedula albicollis). We found that carotenoid concentration was positively associated with caterpillar supply at the time of egg formation, which suggests a proximate constraint of carotenoid availability on yolk composition. Neither female condition, body size, age, nor male plumage ornamentation, age and body size correlated with carotenoid deposition. Yolk β-carotene concentration was found to be positively linked to yolk testosterone concentration. We suggest that females allocated more β-carotene to their eggs to mitigate the potentially detrimental effects of elevated steroid concentration. We found that concentration of β-carotene increased with laying order. The possible function of this pattern may be to enhance the resistance to oxidative stress and pathogens of the disadvantaged last-hatching nestling, suggesting that collared flycatchers pursue a compensatory, “brood survival” strategy.