Use of artificial arenas to predict the social organisation of halictine bees: Data for fourteen species from Chile

I studied the behavioural profiles of fourteen species of Chilean halictine bee using the circle tube apparatus. Interactions were classified as aggressive, avoidance or cooperative. One species, Corynura chloris, is believed to be semisocial and its behaviour was largely aggressive, Ruizantheda mutabilis is known to be communal and its behaviour was predominantly cooperative whereas Penapis toroi is a member of a subfamily from which only solitary behaviour is known and it primarily exhibited avoidance behaviours. As a result, I suggest that the relative frequency of these different behaviours may be useful in establishing the social organisation of species. Based upon comparisons of behavioural profiles, I predict the social organizations of the remaining species and suggest that at least some of the bees in the populations of Ruizantheda proxima, Caenohalictus dolator, Ca. species A and Pseudagapostemon pississi investigated exhibit communal behaviour; those of Co. patagonica and Co. herbsti likely have a reproductive division of labour, probably within semisocial societies, and that the populations of Co. corinogaster, Co. melanocladius, Ca. pygosinuatum, Ca. cuprellus and Lasioglossum aricense studied are probably solitary. Phylogenies suggest that solitary behaviour in Ca. pygosinuatum and Ca. cuprellus is reversed from communal behaviour and solitary behaviour in L. aricense may be a reversal from eusociality. The response “back and follow” is not indicative of dominance or cooperation as it correlated only with the frequency of avoidance interactions among the taxa studied. I discuss the utility of the circle tube apparatus in the identification of populations worthy of more detailed sociobiological investigation. Received 10 August 2005; revised 8 March 2006; accepted 21 March 2006.     

The influence of competition between foragers on clutch size decisions in an insect parasitoid with scramble larval competition

The effect of competition between ovipositing females on theirclutch size decisions is studied in animals that lay their eggsin discrete units of larval food (hosts). In such species theeffect of competition depends on the form of the larval competitionwithin such units. In insect parasitoids, there might eitherbe contest (solitary parasitoids) or scramble competition (gregariousparasitoids) between larvae within a host For gregarious parasitoids,a decreasing clutch size with increasing competition betweenforagers is predicted. This prediction is tested in experimentsusing the parasitoid Aphaertta minuta. Parasitoids were eitherkept alone or in groups of four before the experiment, in whichthey were introduced singly in a patch containing unparasitizedhosts. Animals kept together laid on average clutches of 0.74eggs smaller than females kept alone (average clutch is 5.3),thereby confirming the prediction. Clutch size decreased withencounter number, which might be due to the adjustment of thefemale's estimate of the encounter rate with hosts. Finally,the results are compared with those reported for solitary parasitoids(that have scramble larval competition), for which it is predictedthat the clutch size will increase with increasing levels ofcompetition between females.     

The effect of pollen protein concentration on body size in the sweat bee Lasioglossum zephyrum (Hymenoptera: Apiformes)

Adult bees and wasps provide all the food their offspring require to grow from egg to adult. For a given diet, offspring body size generally increases with an increase in the amount of food consumed as a larva, but the extent to which body size is influenced by the type of food consumed is poorly known. Pollen ranges from 2–60% protein among plant species, and bees are extremely efficient at assimilating nitrogen; therefore, it seems likely that either parent bees adjust the size of larval provisions to compensate for differences in pollen protein concentration or bee offspring attain different body size depending on the pollen type(s) consumed as a larva. We presented the generalist sweat bee Lasioglossum zephyrum with pollen diets that differed in protein content and monitored offspring body size during two experiments. In a protein supplementation experiment, diets ranged from 20–66% protein and consisted of Typha pollen amended with soy protein. On a pollen/soy diet, offspring body size increased 25% with a shift from 20–37% protein, but did not increase further at greater protein concentrations. In a multiple pollen experiment, pollen diets ranged from 20–39% protein and consisted of eight pollens that differed naturally in protein concentration. The largest offspring arose from the most protein-rich pollens, whereas much smaller bees developed on protein-poor pollens. Provision size only predicted offspring size when pollen type, and therefore protein quantity, was considered. Adult foragers did not adjust provision size to compensate for pollen protein. Therefore, offspring body size appears to result from a combination of controlled (provision size) and uncontrolled (pollen quality) factors that arise out of bee foraging decisions.     

蜜蜂繁殖冲突与雌性蜜蜂信息素研究进展

在营造社会性生活的蜜蜂群体里,蜂王释放出蜂王信息素来控制工蜂卵巢发育及改造王台特性,并吸引雄蜂为之交配,使蜂群正常繁衍。本文在国内外相关研究的基础上,对蜜蜂各蜂种、亚种以及特殊蜂群无政府主义蜂群、海角蜜蜂等雌性蜜蜂信息素成分以及含量变化进行综合论述,并对蜂王主要信息素对工蜂和雄蜂生理影响以及今后的研究趋势作扼要介绍。     

The influence of marking upon bee behaviour in circle tube experiments with a methodological comparison among studies

Summary. Circle tubes were used to investigate the influence of marking upon the behavioural profiles of two halictid species; one, Penapis toroi Rozen, putatively solitary, the other, Corynura chloris (Spinola) thought to be semisocial. Previous studies have shown that solitary bees tend to exhibit avoidance behaviour and those with a reproductive division of labour tend to behave aggressively within circle tube arenas. Marked pairs of C. chloris were significantly more cooperative than unmarked pairs and marked P. toroi showed significantly increased aggression and decreased cooperation. These findings suggest that marking bees may reduce behavioural differences among taxa with different types of social organization. Recommendations are made on how to use the circle tube apparatus to detect bee populations that might be worth more detailed studies of social organization and also for standardization in the way results of these studies are presented. Further experiments are suggested to clarify the effects of certain variables upon bee behaviour using this apparatus. The data presented here are the first for any member of the subfamily Rophitinae and the first for any bee thought to be primarily semisocial. A narrower range of behaviours were observed in the Rophitine, Penapis toroi Rozen, in comparison to species from other subfamilies of Halictidae and Corynura chloris Spinola had a behavioural profile similar to that of eusocial species.Received 15 December 2003; revised 12 September 2004; accepted 15 October 2004.     

Sperm competition and brain size evolution in mammals

The ‘expensive tissue hypothesis’ predicts a size trade‐off between the brain and other energetically costly organs. A specific version of this hypothesis, the ‘expensive sexual tissue hypothesis’, argues that selection for larger testes under sperm competition constrains brain size evolution. We show here that there is no general evolutionary trade‐off between brain and testis mass in mammals. The predicted negative relationship between these traits is not found for rodents, ungulates, primates, carnivores, or across combined mammalian orders, and neither does total brain mass vary according to the level of sperm competition as determined by mating system classifications. Although we are able to confirm previous reports of a negative relationship between brain and testis mass in echolocating bats, our results suggest that mating system may be a better predictor of brain size in this group. We conclude that the expensive sexual tissue hypothesis accounts for little or none of the variance in brain size in mammals, and suggest that a broader framework is required to understand the costs of brain size evolution and how these are met.     

The influence of soil temperature on the nesting cycle of the halictid bee Lasioglossum malachurum

The physiology and behavior of ectothermic organisms is strongly influenced by temperature. For ground nesting species like the primitively eusocial halictid bee, Lasioglossum malachurum, soil temperature might influence the life cycle as well as the complexity of the social group since the number of broods that can be fitted into the flight season might increase with increasing temperature. Our study populationof L. malachurum at Wuerzburg exhibits a remarkable variability with respect to the number of broods and the pattern of sexual production. Broods are separated by activity pauses during which the larvae develop. In this study we investigate the influence of soil temperature on the pattern of nesting activity (duration of broods and pauses) and on the number of broods in L. malachurum. We observed a total of 1138 nests in 13 aggregations near Wuerzburg. As expected, soil temperature shortened the duration of the pauses, resulting in an overall shortening of the nesting cycle. This is most probably due to a physiological effect of soil temperature on the development of the larvae. With regard to the nesting strategies, we hypothesized that a shortening of the nesting cycle within the limited flight season should enhance the success of a strategy with more worker broods. In fact, patches with higher soil temperature showed more broods. However, this effect was rather weak, suggesting that other factors might have a stronger impact on the variability in nesting strategy within our study population of L. malachurum. Received 20 December 2005; revised 25 April 2006; accepted 2 May 2006. An erratum to this article is available at .     

Sperm competition and maternal effects differentially influence testis and sperm size in Callosobruchus maculatus

The evolutionary factors affecting testis size are well documented, with sperm competition being of major importance. However, the factors affecting sperm length are not well understood; there are no clear theoretical predictions and the empirical evidence is inconsistent. Recently, maternal effects have been implicated in sperm length variation, a finding that may offer insights into its evolution. We investigated potential proximate and microevolutionary factors influencing testis and sperm size in the bruchid beetle Callosobruchus maculatus using a combined approach of an artificial evolution experiment over 90 generations and an environmental effects study. We found that while polyandry seems to select for larger testes, it had no detectable effect on sperm length. Furthermore, population density, a proximate indicator of sperm competition risk, was not significantly associated with sperm length or testis size variation. However, there were strong maternal effects influencing sperm length.     

Season length,body size,and social polymorphism: size clines but not saw tooth clines in sweat bees

1. Annual insects are predicted to grow larger where the growing season is longer. However, transitions from one to two generations per year can occur when the season becomes sufficiently long, and are predicted to result in a sharp decrease in body size because available development time is halved. The potential for resulting saw‐tooth clines has been investigated only in solitary taxa with free‐living larvae. 2. Size clines were investigated in two socially polymorphic sweat bees (Halictidae): transitions between solitary and social nesting occur along gradients of increasing season length, characterised by the absence or presence of workers and offspring that are individually mass provisioned by adults. How the body size changes with season length was examined, and whether transitions in social phenotype generate saw‐tooth size clines. We measured Lasioglossum calceatum and Halictus rubicundus nest foundresses originating from more than 1000 km of latitude, encompassing the transition between social and solitary nesting. 3. Using satellite‐collected temperature data to estimate season length, it was shown that both species were largest where the season was longest. Body size increased linearly with season length in L. calceatum and non‐linearly in H. rubicundus but the existence of saw‐tooth clines was not supported. 4. The present results suggest that because the amount of food consumed by offspring during development is determined by adults, environmental and social influences on the provisioning strategies of adult bees may be more important factors than available feeding time in determining offspring body size in socially polymorphic sweat bees.     

Testis size,epididymis weight,and sperm competition in rhesus macaques

The intensity of sperm competition is often measured using the gonadosomatic index (testes/body weight). But sperm competition could be mediated more by size of the epididymis than by size of the testicles, and little information is available on the relationship between testicular and epididymal size. We found that both organs were positively correlated in size among male rhesus macaques. Body weight accounted for over 70% of the variance in testicle size and volumetric estimates of testicle size accurately reflected testicle weight. We conclude that methods for ascertaining testicle size are accurate, but the covariation in size between testicles and epididymis will hamper understanding of the physiological mechanisms involved in sperm competition in primates. © 1993 Wiley-Liss, Inc.     

Social group size, potential sperm competition and reproductive investment in a hermaphroditic leech, Helobdella papillornata (Euhirudinea: Glossiphoniidae)

Social group size may affect the potential for sperm competition, and this in turn may favour ontogenetic adjustments in testicular mass according to the likely requirements for sperm and spermatophore production. In a number of comparative analyses of testis mass among vertebrate species that differ in mating system or social organization, increasing potential for sperm competition is associated with larger testis size. Intraspecific phenotypic plasticity should be able to produce the same pattern if social group size is heterogenous and reflects differing degrees of average sperm competition, but this intraspecific effect is less well studied. We tested the effect of social groups on both male and female investment in the simultaneously hermaphroditic leech, Helobdella papillornata. Leeches were placed in groups of one, two, four or eight. Sexual investment at the onset of reproductive maturity was quantified as the total testisac volume for male function and total egg volume for female function. We found that testisac volume (statistically adjusted for body size) showed a significant increase with increasing group size. Total egg volume (also adjusted for body size) was unaffected by group size. Our findings indicate adaptive developmental plasticity in male gonad investment in response to the potential for sperm competition.     

Local community size mediates ecological drift and competition in metacommunities

The outcome of competitive interactions is likely to be influenced by both competitive dominance (i.e. niche-based dynamics) and ecological drift (i.e. neutral dynamics governed by demographic stochasticity). However, spatial models of competition rarely consider the joint operation of these two processes. We develop a model based on the original competition-colonization trade-off model that incorporates niche and neutral processes and several realistic facets of ecological dynamics: it allows local competition (i.e. competition within a patch) to occur within communities of a finite size, it allows competitors to vary in the degree of competitive asymmetry, and it includes the role of local migration (i.e. propagule pressure). The model highlights the role of community size, i.e. the number of competitors in the local community, in mediating the relative importance of stochastic and deterministic forces. In metacommunities where local communities are small, ecological drift is substantial enough that strong competitors become effectively neutral, creating abrupt changes in the outcome of competition not predicted by the standard competition-colonization trade-off. Importantly, the model illustrates that, even when other aspects of species interactions (e.g. migration ability, competitive ability) are unchanged, local community size can alter the dynamics of metacommunity persistence. Our work demonstrates that activities which reduce the size of local communities, such as habitat destruction and degradation, effectively compound the extinction debt.     

The influence of horn and body size on the reproductive behavior of the horned rainbow scarab beetlePhanaeus difformis (Coleoptera: scarabaeidae)

The reproductive behavior of horned rainbow scarab beetles,Phanaeus difformis, was studied to determine the influence of morphological traits on intersexual and intrasexual interactions.Phanaeus difformis is a sexually dimorphic dung beetle in which males possess much larger horns than females, and males can be grouped into major and minor male morph categories based on horn size. Male-female pairs cooperated in nest construction and provisioning. In the laboratory, males of both morphs assisted females and were equally successful at copulating. However, in the field larger individuals had a pairing advantage due to greater success in intrasexual competition. Some males used an alternative mating tactic which involved sneaking copulations with paired females. In most cases the sneak male was smaller than the paired male.     

Social competition, corticosterone and survival in female lizard morphs

We examined the selective consequences of variation in behaviour and endocrine physiology in two female throat-colour morphs of the lizard, Uta stansburiana in the wild. Female morphs differed in home-range distribution patterns and corticosterone levels in relation to the density and frequency of their female neighbours. Levels of plasma corticosterone of yellow-throated females increased with increased density of both morphs. In contrast, orange-throated females had reduced levels of corticosterone in response to increased density of orange females. Additionally, females with lower corticosterone survived poorly, suggesting that social interactions and high local densities of orange females may be potentially costly for orange females. These results are consistent with decreased fitness effects and suppression of immune function previously reported for orange female morphs surrounded by more orange neighbours. These correlations, in conjunction with previous work in this system, indicate that corticosterone is likely to be an important physiological mechanism regulating female fitness in nature.     

Foraging and courtship behaviour in males of the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae): thermal physiology and the roles of body size

Abstract.

• 1 The effects of climate and body size on male behaviour were examined in the solitary bee Anthophora plumipes (Hymenoptera: Anthophoridae), which shows resource-based polygyny at floral food sources in Britain in spring.
• 2 Larger males are able to fly at lower temperatures than smaller males, and can therefore court females under conditions in which smaller males cannot fly. This is predicted from patterns of endothermic ability at low temperatures already demonstrated within this species.
• 3 Video analysis of male competition for opportunities to initiate courtship with tethered females showed that larger males are also competitively superior, and can displace smaller males from favoured flight positions immediately behind females.
• 4 The mating system shown by male A plumipes is strongly dependent on male density. At low densities, males show exclusive territoriality at floral sources. As male density increases, this strategy is abandoned in favour of patrolling with considerable spatial overlap between males, and opportunistic Polygyny.
• 5 Despite high endothermic abilities, male behaviour is highly dependent on weather, and particularly ambient temperature. Males bask in the early morning and maintain high thoracic temperatures. Temperature data from freshly killed bees show that thoracic warming from solar sources effectively doubles the thermogenic power generated by the bee alone at low ambient temperatures.
• 6 Male strategies in A.plumipes are compared to female responses to climate. Having controlled for differences in body size there is no difference in endothermic abilities between the sexes. Males do not, however, fly under conditions in which females of the same size would remain active. These results are discussed in the light of differential dependence of reproductive success on flight activity for the two sexes.

     

Reproductive strategies of primates: The influence of body size and diet on litter size

The frequency of multiple births, life history parameters, body size, and diet characteristics were obtained from the literature for 70 primate species. The general pattern within the primate order is to have single infant litters, yet multiple births regularly occur in a number of species in specific phylogenetic groups. Primates which have large litters tend to be small, have short gestation periods and give birth to small infants, which are weaned quickly, and mature rapidly. Species in which multiple births are common also have short interbirth intervals and in the Callitrichidae have males which exhibit paternal care. In addition, they are commonly insectivorous. Although it is difficult to isolate the effects of diet on litter size, independent of body size, analyses suggest that after the influence of body size is statistically removed, as the proportion of insects in the diet increases, animals have larger litters. We suggest that by adopting a mixed diet of insects and fruit primates may be able to ensure access to a seasonally stable food resource that is not greatly restricted by the presence of toxins. This diet would allow a relatively high metabolism and facilitate large litters.