Relationship between body size, provisioning rate, longevity and reproductive success in females of the solitary bee Osmia cornuta

Body size has often been related to reproductive success in bees and wasps. The objective of this 3-year study was to analyze the relationship between nesting female body size, provisioning rate and longevity and their effect on several traits related to parental investment and reproductive success in the solitary bee Osmia cornuta. Body size was not correlated to longevity, and it was only correlated to provisioning rate in the third year (with poor weather conditions during nesting). Variation in fecundity, offspring size and offspring mortality was not well explained by nesting female body size in any of the 3 years. However, in the third year, small females biased their investment toward males, the sex requiring smaller pollen–nectar provisions. Large females were more successful usurpers of other females' nests, but fecundity of usurpers was no higher than fecundity of nonusurpers. Large females were more likely to establish at the release site, probably in relation to size-dependent vigor at emergence. A review of the literature on parental investment in solitary aculeate Hymenoptera showed a stronger relationship between body size and reproductive success in wasps than in bees. In O. cornuta, fecundity was strongly related to longevity and provisioning rate in all 3 years. Offspring size was associated with provisioning rate in 1 year, when females with higher provisioning rates tended to produce larger sons and daughters. Both longevity and provisioning rate appeared to be strongly conditioned by stochastic events.     

Queen longevity,queen adoption,and posthumous indirect fitness in the facultatively polygynous ant Myrmica tahoensis

In many polygynous ant species, established colonies adopt new queens secondarily. Conflicts over queen adoption might arise between queens and workers of established colonies and the newly mated females seeking adoption into nests. Colony members are predicted to base adoption decisions on their relatednesses to other participants, on competition between queens for colony resources, and on the effects that adopted queens have on colony survivorship and productivity. To provide a better understanding of queen-adoption dynamics in a facultatively polygynous ant, colonies of Myrmica tahoensis were observed in the field for 4 consecutive years and analyzed genetically using highly polymorphic microsatellite DNA markers. The extreme rarity of newly founded colonies suggests that most newly mated queens that succeed do so by entering established nests. Queens are closely related on average (rˉ = 0.58), although a sizable minority of queen pairs (29%) are not close relatives. An experiment involving transfers of queens among nests showed that queens are often accepted by workers to which they are completely unrelated. Average queen numbers estimated from nest excavations (harmonic mean = 1.4) are broadly similar to effective queen numbers inferred from the genetic relatedness of colony members, suggesting that reproductive skew is low in this species. Queens appear to have reproductive lifespans of only 1 or 2 years. As a result, queens transmit a substantial fraction of their genes posthumously (through the reproduction of related nestmates), in comparison to direct and indirect reproduction while they are alive. Thus queens and other colony members should often accept new queens when doing so will increase colony survivorship, in some cases even when the adopted queens are not close relatives. Received: 20 February 1996/Accepted after revision: 25 May 1996     

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

Shoal composition, body size and foraging in sticklebacks

Individuals which deviate from the majority in groups are likely to be most vulnerable to predation. This oddity effect, by definition, is frequency dependent, eventually fading at equal frequencies of the phenotypes in a group. It has been hypothesized that the increased predation risk of odd individuals may play an important role in the formation of phenotypically uniform shoals of fish. However, recent work has indicated that individuals may experience, or value, their predation hazard differently depending on their own size in relation to that of other group members: single large fish, but not small ones, appear concerned about their oddity in a shoal. Here I show that the apparent wariness of large fish is also expressed in a frequency-dependent manner, closely conforming to what is predicted if the oddity effect is responsible for their behavior. Using foraging activity of individuals as a means to evaluate their predation risk, I demonstrate with shoals comprising 12 threespine sticklebacks (Gasterosteus aculeatus) that large fish forage least actively when in a shoal consisting of 2 large and 10 small fish. An increase in the number of large fish to 4 among 8 small individuals clearly results in an increase in their foraging activity. However, having reached an equal frequency with small fish in a shoal, large fish do not seem to change their foraging activity much even when their number in a shoal increases further. In contrast, foraging activity of small sticklebacks remains fairly constant throughout the entire range of tested shoal compositions, providing further evidence that small and large fish respond to their oddity differently. Received: 12 February 1998 / Accepted after revision: 7 May 1998     

Queen fertility, egg marking and colony size in the ant Camponotus floridanus

In ant societies, workers do not usually reproduce but gain indirect fitness benefits from raising related offspring produced by the queen. One of the preconditions of this worker self-restraint is sufficient fertility of the queen. The queen is, therefore, expected to signal her fertility. In Camponotus floridanus, workers can recognize the presence of a highly fertile queen via her eggs, which are marked with the queen's specific hydrocarbon profile. If information on fertility is encoded in the hydrocarbon profile of eggs, we expect workers to be able to differentiate between eggs from highly and weakly fertile queens. We found that workers discriminate between these eggs solely on the basis of their hydrocarbon profiles which differ both qualitatively and quantitatively. This pattern is further supported by the similarity of the egg profiles of workers and weakly fertile queens and the similar treatment of both kinds of eggs. Profiles of queen eggs correspond to the cuticular hydrocarbon profiles of the respective queens. Changes in the cuticular profiles are associated with the size of the colony the queen originates from and her current egg-laying rate. However, partial correlation analysis indicates that only colony size predicts the cuticular profile. Colony size is a buffered indicator of queen fertility as it is a consequence of queen productivity within a certain period of time, whereas daily egg-laying rate varies due to cyclical oviposition. We conclude that surface hydrocarbons of eggs and the cuticular profiles of queens both signal queen fertility, suggesting a major role of fertility signals in the regulation of reproduction in social insects.     

Song amplitude and body size in birds

Bird song is a sexually selected multidimensional signal. A fundamental question regarding the evolution of sexually selected signals is what information they convey and how their honesty is maintained. Song amplitude is a performance-related signal trait that varies considerably between individuals, but this signal dimension has been neglected in past studies. I found that median song amplitude in male nightingales (Luscinia megarhynchos) and zebra finches (Taeniopygia guttata) did not vary significantly with body size or residual body mass. In contrast, I found a significant negative correlation between body size (and also residual mass) and the maximum song amplitude during interactive singing in nightingales. However, the function of these more subtle differences in song amplitude remains to be investigated. By and large, the results of this study suggest that mean song amplitude is unlikely to indicate a bird’s body size or current condition (measured as residual mass).     

Effects of mating history on ejaculate size, fecundity, longevity, and copulation duration in the ant-tended lycaenid butterfly, Jalmenus evagoras

The mating system of the Australian lycaenid butterfly, Jalmenus evagoras, is highly unusual compared to most other Lepidoptera. Characteristics of this system, which has been termed an ’explosive mating strategy,’ include the formation of an intensely competitive mating aggregation of males, a highly male biased operational sex ratio, a lack of discrimination and mate choice by both sexes, a high variance in male mating success, and female monogamy. In this study, we tested the hypothesis that multiple mating by males imposes physiological costs resulting in smaller spermatophores, and that this results in a fitness cost to females. We found that male J. evagoras transferred only 2.2% of their eclosion weight during their first mating, consistent with the hypothesis that males of monandrous species produce a relatively small investment. The wet weight of the ejaculate declined by an average of 27% at the second mating and the dry weight by 29%, and an intermating interval of 5–9 days was needed for the ejaculate to return to the size at the first mating, regardless of male size or age. Wet ejaculate mass increased proportionally with male size, though dry mass was proportionally larger in smaller males. Ejaculate mass tended to increase with male age at both first and second matings. Female characteristics, in general, did not affect ejaculate mass, although the wet weight of the ejaculate was positively associated with female weight at the second mating. Copulation duration increased from 2.4 h to approximately 3 h at the second mating, and to over 4 h at the third and fourth matings. Fecundity was positively correlated with female size but not with mating history, copulation duration, or any other characteristics measured for either males or females. Female longevity declined significantly as the number of times the male partner had previously mated increased. We conclude that despite the small male investment in ejaculate, the costs of multiple mating may nonetheless be significant, as indicated by the reduction in ejaculate mass, an increase in copulation duration, and reduction in female lifespan with increasing mating number. Received: 22 January 1999 / Received in revised form: 28 July 1999 / Accepted: 18 September 1999     

Pre- and post-mating sexual selection both favor large males in a rainbowfish

Sexual selection can act through female choice and male–male competition. Although both processes can act simultaneously, they are typically studied independently. Here, we adopt a more integrated approach to studying sexual selection by incorporating measures of both processes using the western rainbowfish Melanotaenia australis, a freshwater fish endemic to northwestern Australia. We assessed male–male competition and female choice separately while measuring the performance of individual males under both processes and used paternity analyses to estimate male reproductive success. We then related the performance of males during each of these stages to their phenotype, which was described using linear measures of size and color pattern traits, and spectrographic measures of the reflectance of color patches. We found that female choice favored relatively large males and that these preferences were consistent within individual females and repeatable between different females. Larger males were also more dominant in the competition trials and sired the majority of offspring produced when females spawned. There was little evidence to suggest that sexual selection acted on male color patterns either via female choice or male contest competition or during subsequent post-mating episodes of sexual selection. We conclude, therefore, that male–male competition and female choice act concordantly to favor relatively large males and that these patterns of mating success are reflected during post-mating episodes of sexual selection.     

Waste management in the leaf-cutting ant Acromyrmex echinatior: the role of worker size, age and plasticity

Division of labour is the hallmark of the success of many social animals. It may be especially important with regard to waste management because waste often contains pathogens or hazardous toxins and worker specialisation can reduce the number of group members exposed to it. Here we examine waste management in a fungus-farming, leaf-cutting ant, Acromyrmex echinatior, in which waste management is necessary to protect their vulnerable fungal crop. By marking ants with task-specific paint colours, we found clear division of labour between workers that engage in waste management and those that forage, at least during the fine timescale of the 3-day marking period. This division of labour was influenced by both age and size, with waste management workers tending to be smaller and younger than foragers. The role of preventing contaminated ants from entering the colony was fulfilled mainly by medium-sized workers. When the level of waste was experimentally increased, most of the ants that responded to remove the waste were workers previously engaged in tasks inside the nest rather than external waste workers or foragers. These responding workers tended to be young and medium-sized. Surprisingly, the responding ants were subsequently able to revert back to working within the fungus garden, but the probability of them doing so depended on their age and the length of time they were exposed to waste. The results demonstrate the importance of division of labour with regard to waste management in A. echinatior and show that this is adaptable to changing needs.     

Initiation and resolution of jumping spider contests: roles for size, proximity, and early detection of rivals

Animals are commonly expected to assess each other during contests in order to economically identify relative status. Escalated or long contests are expected to arise mainly when rivals have difficulty discriminating small differences. Results of the present study of male-male contests in Plexippus paykulli, a jumping spider (Salticidae) with acute vision, are not in accord with this widely held view. Despite the typical finding that size-advantaged rivals are more likely to win contests and that this tendency increases with size disparity, contest dynamics suggest that these tendencies are achieved in the absence of direct size assessment. In contests between different-sized spiders, maximum escalation and overall duration were predicted by the absolute size of the size-disadvantaged spider (usually the loser) rather than the size difference between the rivals. This result suggests that spiders base decisions of persistence on their own size, such that size-disadvantaged rivals usually reach their limits first, and then retreat. This interpretation is further supported by findings that maximum escalation and total duration were both positively related to size in contests between size-matched spiders. Spiders were more likely to win if they oriented and displayed first, and longer, more escalated, contests ensued if the size-disadvantaged spider was the first to orient and display. Proximity of rivals at contest outset also influenced contest dynamics, but not outcome.     

Changes in body size, abundance, and energy allocation in rockfish assemblages of the northeast Pacific.

Fish body size, a key driver of many aspects of fish population biology and ecology, is affected by fisheries that deplete the largest individuals. Rockfish (genus Sebastes) are a diverse group that has been heavily fished on the U.S. West Coast in recent decades. We examined trawl survey data from 1980 to 2001 to determine spatial and temporal trends in body size and density of 16 shelf rockfish species, including six that are considered overfished. Mean individual mass and maximum observed mass declined in the majority of species in one or more zoogeographic regions between central California and Washington. Density changes were far more variable in time and space, but in all regions, density declines were most often associated with large-bodied rockfish. We next estimated the impact of size and density changes on energy consumption and fecundity in a five-species rockfish assemblage that includes bocaccio (S. paucispinis), a large-bodied, overfished species. Indexes of both consumption and fecundity by the assemblage increased in the southern portion of the study area between 1980 and 2001 but decreased in the northern portion. Allocation of energy and reproductive potential within the assemblage shifted dramatically: relative to bocaccio, total energy consumption and fecundity indexes for the other four species increased by orders of magnitude from 1980 to 2001. These changes in community structure may affect the ability of bocaccio and other large rockfish species to recover from overfishing, especially in light of long-term declines in zooplankton production that may also be affecting rockfish size and production. Addressing these issues may require a regional, multispecies management approach.     

Offspring number in pygmy marmosets, Cebuella pygmaea *, in relation to group size and the number of adult males

In many species of birds and mammals with a co-operative breeding and rearing system, offspring survival is positively related to the number of helpers. In the New World callitrichine primates (marmosets and tamarins), adult males are considered as particularly valuable helpers, and female reproductive success may depend strongly on the males' contribution to infant care. We analysed the number of offspring (infants, juveniles) in groups of wild pygmy marmosets, Cebuella pygmaea (Callitrichinae, Cebidae, Primates), in relation to the number of adult males and to the number of adult and subadult group members. In contrast to other callitrichines with a co-operative system of infant care, no relationship was found between the number of adult males and the number of infants and offspring. However, there was a significant positive relationship between the number of juveniles and the number of adult and subadult group members. The lack of a relationship between infant and adult-male number is interpreted as resulting from the reduced importance of adult males as helpers in pygmy marmosets in comparison to other callitrichines, probably due to the reduced costs of infant care. The relationship between the number of juveniles and the number of adult and subadult group members is in accordance with increased offspring survival in larger groups, as observed in other primates. Received: 1 February 1999 / Received in revised form: 5 June 1999 / Accepted: 29 June 1999     

Intraspecific competition affects population size and resource allocation in an ant dispersing by colony fission

Intraspecific competition is a pervasive phenomenon with important ecological and evolutionary consequences, yet its effect in natural populations remains controversial. Although numerous studies suggest that in many cases populations across all organisms are limited by density-dependent processes, this conclusion often relies on correlative data. Here, using an experimental approach, we examined the effect of intraspecific competition on population regulation of the ant Aphaenogaster senilis. In this species females are philopatric while males disperse by flying over relatively long distances. All colonies were removed from 15 experimental plots, except for one focal colony in each plot, while 15 other plots remained unmanipulated. After the first reproductive season, nest density in the experimental plots returned to a level nonsignificantly different from that in the control plots, which was not expected if the populations were indeed regulated by density-independent phenomena. In both the control plots and the experimental plots colonies remained overdispersed throughout the experiment, suggesting colony mutual exclusion. Nests outside the plots rapidly extended their foraging span, but we did not detect any significant inward migration into the experimental plots. Experimental reduction in density did not significantly affect the focal colonies' biomass, measured just before the first reproductive season. However, the ratio of males to workers-pupae biomasses was smaller in experimental plots, suggesting that colonies there had redirected part of the resources normally allocated to male production to the production instead of new workers. Microsatellite analysis indicated that, after the reproductive season, many colonies in the experimental plots were headed by a young queen that was the mother of the brood but not of the old workers, indicating that reduction in colony density stimulated fission of the remaining colonies. Finally, at the end of the experiment, 14 months after experimental reduction in density, colonies that derived from fission were smaller in the experimental than in the control plots, suggesting that the former had undergone fission at a smaller size than in control plots, which presumably allowed them to colonize the emptied areas. We conclude that colonies adjust resource allocation and colony fission to the degree of intraspecific competition.     

Copula duration and testes size in the yellow dung fly,Scathophaga stercoraria (L.) : the effects of diet,body size,and mating history

Summary Seasonal variation in mean hind tibia length and mean testes length is investigated in the yellow dung fly, Scathophaga stercoraria (L.). There is a cycle in mean hind tibia length and mean testes length over a season. The body size curve peaks later than the testes length curve, showing that there is no fixed relationship between the two variables. The causes of variation in testes size and its influence on copula duration are experimentally examined. Increasing the number of Drosophila eaten per day leads to increased mature testes length. Males with larger testes copulate for longer than males with smaller ones, and smaller males copulate for longer than do larger males. While testes shrivel with successive copulas, copula duration remains constant. The more females a male is prepared to copulate with in a day (up to five), the longer he copulates with each. The shrinkage of the testes of males collected throughout a day suggests that males copulate with an average of 4 females per day. The costs of sperm production are thus shown to have a significant influence on the copula duration.Offprint requests to: P.I. Ward at the second address     

Space use,longevity, and reproductive success in meadow voles

Summary We addressed the question of how reproductive success (RS) was limited in the shortlived but highly fecund meadow vole, Microtus pennsylvanicus. In so doing, we asked how differential space use patterns could affect longevity and hence RS in each sex. The sample comprised all voles achieving sexual competency over the course of a 40-week breeding season in a live-trapped population in Manomet, MA USA. Matrilineal families were determined using a radionuclide labelling technique; paternity was estimated using a maximum likelihood model. Individual RS was defined as the number of offspring successfully recruited into the trappable population per adult. We found that the variance in RS among female meadow voles was greater than the variance among males. In an attempt to explain this pattern, reproductively successful individuals were compared to reproductively unsuccessful individuals with regard to survivorship, maximum body weight achieved, and spatial mobility. The only difference between fathers and reproductively unsuccessful males was that fathers were heavier. In contrast, mothers differed from unsuccessful females in every measurement. Females lived longer than males, and mothers lived longer than either fathers or reproductively unsuccessful females. The observed differences in longevity may have been largely the result of differences in levels of mobility, assuming more mobile voles were more susceptible to predation. Mothers were significantly more site tenacious than were either males or unsuccessful females. These patterns explain the distribution of RS in our population if predation differentially affects male and female meadow voles. The meadow vole is the only non-polyandrous vertebrate reported to date in which the variance in RS among females exceeds the variance in RS among males.     

Group size in wedge-capped capuchin monkeys Cebus olivaceus and the reproductive success of males and females

Summary The effect of variation in group size on age-specific survivorship and fecundity rates were examined in a population of wedge-capped capuchin monkeys Cebus olivaceus during a 10 year study. Life tables were constructed separately for four large (15 individuals) and four small groups (<15 individuals). Female reproductive success, and its relative contribution to population growth, was much higher in large groups, primarily through higher age-specific fecundity. Age-specific survivorship was similar in groups of different sizes. The reproductive success of the single breeding male in a group was much higher in large than small groups. Compared to small groups, breeding males in large groups had a longer breeding tenure, and access to greater numbers of reproductive females with a higher average fecundity. Differences in female reproductive success apparently resulted from variation in access to monopolizable fruit trees. Large groups predictably displaced small groups during intergroup encounters. Group rank depended on the number of males resident in groups. The large number of non-breeding males in large groups results from their longer average residency time. I explain the longer residency of males in large groups by the higher average reproductive success of breeding males in these groups.     

Production of sexuals in a fission-performing ant: dual effects of queen pheromones and colony size

Models based on the kin selection theory predict that in social hymenopterans, queens may favor a lower investment in the production of sexuals than workers. However, in perennial colonies, this conflict may be tuned down by colony-level selection because of the trade off between colony survival and reproductive allocation. In this study, we present a survey of sexual production in colonies of Aphaenogaster senilis, a common species of ant in the Iberian Peninsula. Similar to most species that reproduce by fission, males were found in large excess compared to gynes (172:1). Sexuals were more likely to be found in queenless than in queenright (QR) field colonies. However, we also found a few gynes and numerous males in very large QR colonies. We compared these data with those available in the literature for A. rudis, a congeneric species from North America that has independent colony founding. The sex ratio in this species was only five males for each female, and sexuals were mostly found in QR nests, irrespective of colony size. We confirmed queen inhibition of sexual production in A. senilis in laboratory experiments and provide evidence that this inhibition is mediated by a nonvolatile pheromone. To seek the potential source of such a queen pheromone, we analyzed the secretions of two conspicuous exocrine glands, the Dufour’s and postpharyngeal glands (DG and PPG, respectively) in both queens and workers. Both secretions were composed of hydrocarbons, but that of DG also contained small quantities of tetradecanal and hexadecanal. The hydrocarbon profile of the DG and PPG showed notable caste specificity suggesting a role in caste-related behavior. The PPG secretions also differed between colonies suggesting its role in colony-level recognition. We suggest that in A. senilis, there are two modes of colony fission: First, in very large colonies, gynes are produced, probably because of the dilution of the queen pheromone, and consequently one or more gynes leave the mother colony with workers and brood to found a new nest. This is beneficial at the colony level because it avoids the production of costly sexuals in small colonies. However, because the queen and workers have different optima for sexual production, we hypothesize that queens tend to overproduce the pheromone to delay their production. This in turn may drive workers to leave the mother colony during nest relocation and to produce sexuals once they are away from the queen’s influence, creating a second mode of colony fission.