An experimental analysis of joint nesting in the salamander Hemidaetylium scutatum (Caudata: Plethodontidae): the effects of population density

Three nesting behaviour patterns are documented in the plethodontid salamander Hemidactylium scutatum. A female may lay eggs (1) in a solitary nest and brood them, (2) in a joint nest and brood them as well as eggs of other females, or (3) in a joint nest that is brooded by another female. The hypothesis that population density was positively associated with joint nesting was tested by following two populations for 5 years and by experimentally manipulating the population density of nesting females in artificial habitats for the latter 2 years. The proportion of joint nests did not vary with density, although joint nests tended to contain eggs of more females at the high population density. Joint nests were usually brooded by one female; thus, most females that laid eggs in joint nests did not brood them at high density. The reproductive success, as measured by survival of embryos, of solitary and joint nesters was equivalent. Joint nests were deserted less often, however, which decreased the probability of catastrophic mortality. The number of days of brooding was significantly positively correlated with loss of body mass of females, suggesting a cost to brooding behaviour. Joint nesting with solitary brooding is not explained by aggressive usurpation of nests or by brood parasitism.     

Elaborate Nests in a Weaverbird: A Role for Female Choice?

Studies on sexual selection have focused on behaviour and morphology, but several groups of animals build elaborate structures associated with acquiring a mate. I investigated female choice for nests built by male baya weavers (Ploceus philippinus). Nest choice by females should be strong, as nests are obvious direct benefits provided by males. I used a field experiment supplemented with correlational information to ask whether females appear to base mate choice decisions on male behaviour, nest architecture, and nest location. When the nests of highly visited males were exchanged with those of poorly visited males, female visits remained highest at the original male and location. I found no relationship between female choice and male display or other behaviour. Correlational analyses show that nest location was a better predictor of female choice than was nest architecture. These data suggest that current female choice is driven more by access to safe nesting sites rather than to well‐built nests, possibly because all males are able to build nests of adequate quality. However, nest architecture is unlikely to be irrelevant to females, and its role deserves further investigation.     

Dispersed versus gregarious nesting strategies in the mason bee Chalicodoma siculum

Chalicodoma siculum (Rossi) is a mason bee that usually nests in dispersed sites, constructing spherical mud nests on twigs or hemispherical nests on hard surfaces. Under favourable conditions it may nest in aggregations of several hundred bees. At the nesting aggregation, nest usurpation often occurs, either by the laying of eggs in finished but unsealed cells, by opening a sealed cell and replacing its egg, or by taking over an unfinished cell. Mutual construction and provisioning of a single cell by several females was also often observed. Because participant bees display extreme aggression in prolonged disputes over cell ownership, these interactions can be regarded as competitive rather than cooperative. Moreover, since the total investment in such cells is not different from that in cells located in dispersed sites, the female that actually manages to lay an egg gains by participating in a cell under construction.
We discuss the advantages of nesting in aggregations vs. the risks of being usurped, as selective forces underlying nesting behaviour of bees.     

Male short-tailed field voles (Microtus agrestis) build better insulated nests than females

Nest construction is an extremely widespread behaviour. In small endotherms the nest serves primarily to provide insulation, and thereby retard heat loss of the constructor, or its offspring. In arctic and temperate regions many small mammals build nests to protect themselves from low ambient temperatures. We measured the physical properties of nests built by short-tailed field voles Microtus agrestis that were kept in captivity under cold conditions. The most important factor influencing nest insulation was nest wall thickness; however, nests with thick walls also contained more nesting material. Insulative capacity of the nest did not reach an asymptote up to nests containing 20 g of material. Nest insulation was not correlated with resting metabolic rate, body mass or body composition of the vole that constructed the nest. However, nests built by males had greater insulation than those made by females; males also had significantly lower food intake rates when compared to females with nests. No significant difference was observed in either fat mass or whole animal thermal conductance between males and females. Thermal conductance did increase significantly with increasing body mass, although not with resting metabolic rate. Voles with nests for prolonged periods had lower food intakes than voles without nests. The absolute saving averaged 1.9 g and was independent of body mass. This was a 28% saving on intake for a 22-g vole but only an 18% saving for a 40-g individual. When voles had nests for short periods they used the energy they saved to reduce food intake and increase body mass. Accepted: 2 September 1999     

Solitary and group nesting in the orchid bee <Emphasis Type="Italic">Euglossa hyacinthina</Emphasis> (Hymenoptera,Apidae)

Summary Orchid bees (Euglossini) provide a potentially informative contrast for examining origins of advanced social behaviour in bees because they are the only tribe in the apine clade that do not form large colonies or have queens and workers. We investigated natural nests of Euglossa hyacinthina Dressler, an orchid bee that nests singly or in groups. By comparing the two types of nests, we examined if individuals in a group merely share the nest (are communal) or exhibit a level of social organization where there is reproductive division of labour among the females. Observations are consistent with communal nesting, indicating that all females in group nests are reproductively similar to the solitary nesting females because the provisioning of young, as well as the ovary development and mating status of females sharing nests were not different than that of solitary-nesting females. Also, multiple female nests did not produce a female-biased brood as predicted for nests with reproductive division of labour. We also investigated potential advantages of group nesting vs. individual nesting. We demonstrate that per capita offspring production is lower in nests with more than one female. However, we found that nests with single females were left unattended for longer periods of time during foraging, and that there was a high incidence of natural enemy attack in nests when females were absent. Group and solitary nesting may be advantageous under different conditions.Received 3 December 2002; revised 7 March 2003; accepted 2 April 2003.     

Nesting Behaviour of Abert Squirrels (Sciurus aberti)

Nesting behaviour of Abert squirrels (Sciurus aberti), including site selection and use, was studied in the foothills of the Rocky Mountains in Boulder County, Colorado. Only females were observed building nests, although both males and females maintained nests once they were built. Communal nesting by Abert squirrels was rare, but the majority of observed nest sharings involved unrelated male and female pairs. A total of 14 variables were used to evaluate the nests (n = 49) inhabited by Abert squirrels from May 1988 to Jun. 1991. All nests were located in Ponderosa pine (Pinus ponderosa) trees. The majority of nests were constructed of twigs and located in the upper one-third of the canopy, near the trunk, on the south-east side of the tree. Trees with nests were predominantly located in closed stands. Nest trees, when compared with unused control trees that were equally accessible to squirrels, were significantly different from control trees in five of nine variables. Nest tree crowns intertwined with a larger number of adjacent tree crowns than did control tree crowns. Nest trees were also significantly taller than control trees, but subdominant to adjacent trees within a stand. Seasonal and diurnal patterns of nest use indicate that Abert squirrels do not choose nest locations on the south-east sides of trees to facilitate behavioural thermoregulation. Rather, Abert squirrels select nest site locations to (1) maximize accessibility and (2) maximize structural stability which may provide protection from wind and rain.     

Nesting biology and nest re-use ofEulaema nigrita (Hymenoptera: Apidae,Euglossini)

Summary The nesting behavior of the euglossine beeEulaema nigrita was observed in the laboratory after being transferred from three nests to observation boxes. Nests were re-used by successive generations with more than one female working in each re-use process. Associated females were always of the same generation. All females that shared a nest foraged for construction materials (mud, excrement and resin) and each constructed, provisioned and oviposited in her own cells. The number of cells constructed by each female ranged from one to 23. Females stayed in the nests for periods ranging from 15 to 59 days during the hot and wet season and from five to 78 days during the cool and dry season. The egg-to-adult period was related to climatic conditions, and in all re-use processes it was longer than the time of residence of the females in the nests. The meloid beetleMeloetyphlus attacephalus was the only nest parasite. Due to the attack method employed by this parasite, the presence of more than one female in the nest did not result in improved nest defense.     

The nesting of Osmia rufa (L.) (Hymenoptera, Megachilidae) in the Crimea: Structure and composition of nests

The structure and composition of nests of Osmia rufa have been investigated. The weight of the septa between the cells increases regularly from the first to the last cell, whereas the length of cells and weight of the pollen loaf decrease. The difference of weight between cells of males and those of females is sharper. The number of cells and weight of cocoons with adults decrease in the series of nests built by the same female during the nesting season, and the sex ratio shifts to males. Abundant food resources facilitate an increase in the average weight of the offspring, the number of cells, and shift of the sex ratio toward females. The mechanism of the sex ratio regulation in O. rufa nests includes three consecutive elements. The key element is the presence of a marker of the switching to making low-weight pollen loafs. This marker is the moment of the “overloading” of the last cell in a row of cells intended for future females. Daughters of large females are smaller, and those of small females, larger than their mothers. The probable cause of this discrepancy is the allometric relation between the female’s weight and its working capacity.     

Normal and atypical nesting behaviour of Copris lunaris (L.): comparison with related species (Coleoptera, Scarabaeidae)

Abstract. 1. The lifecycle, mating and larval behaviour of Clunaris are described. Adults appeared in the autumn and nested in the following spring. The female beetle remained in the nest with the brood and could nest again the following year.
2. Nesting was initiated when virgin females were mated in the spring. Brood balls were formed by techniques sirnilar to those used by Scarabaeini. The female beetle left the nest soon after the first imagos broke out of the brood balls.
3. Nesting behaviour was readily modified by external conditions. Many parts of the sequence could be repeated or omitted. The female beetle left the nest if the brood was removed, but she remained for longer than usual if younger brood was substituted near the end of the normal nesting period.
4. Certain experimental conditions released behaviour patterns typical of other species. These were formation of superficial nests or of two-chambered nests, oviposition before completing the brood ball, and coating of the brood balls with soil (all found in other Coprina), as well as ball rolling and ball burial (found in Scarabaeini). The results are discussed in relation to the evolution of Copris nesting behaviour.     

Spatial Nest‐Settlement Decisions in Digger Wasps: Conspecifics Matter more than Heterospecifics and Previous Experience

Research into the driving forces behind spatial arrangement of wasp nests has considered abiotic environmental factors, but seldom investigated attraction or repulsion towards conspecifics or heterospecifics. Solitary female digger wasps (Hymenoptera) often nest in dense aggregations, making these insects good models to study this topic. Here, we analysed the nesting patterns in an area shared by three species of the genus Bembix, in a novel study to discover whether female wasps are attracted to or repulsed by conspecific nests, heterospecific nests or their own previously established nests when choosing nest‐digging locations. Early in the season, each species showed a clumping pattern of nests, but later in the season, a random distribution of nests was more common, suggesting an early conspecific attraction. Such behaviour was confirmed by the fact that females started building their nests more frequently where other females of their species were simultaneously digging. The distances between subsequent nests dug by individual females were shorter than those obtained by random simulations. However, this pattern seemed to depend on the tendency to dig close to conspecifics rather than remain in the vicinity of previous nests, suggesting that females' experience matters to future decisions only on a large scale. Nesting patches within nest aggregations largely overlapped between species, but the nests of each species were generally not closer to heterospecific nests than expected by chance, suggesting that females are neither repulsed by, nor attracted to, congenerics within nest aggregations. A role of the spatial distribution of natural enemies on the observed nesting patterns seemed unlikely. Bembix digger wasp nest aggregations seem thus to be primarily the result of female–female attraction during nest‐settlement decisions, in accordance with the ‘copying’ mechanisms suggested for nesting vertebrates.     

Nesting biology and social structure of <Emphasis Type="Italic">Euglossa (Euglossa) townsendi</Emphasis> Cockerell (Hymenoptera,Apidae, Euglossini)

Summary. The nesting behavior of the euglossine bee Euglossa townsendi was studied on the campus of the University of São Paulo-Ribeirão Preto, SP, Brazil, from January 1994 to December 1999, using artificial nesting substrate and observation boxes. Twenty-one nests were founded during the study period, with the highest frequencies of nesting occurring during the hot, wet season. Each nest was founded by a single female and, after the completion of the last cell, she spent most of her time in the nest. The males left the nest immediately after emergence and did not return. Some females left the nest within a few days of eclosing, while others stayed in their natal nests and began to reactivate them. Reactivations were performed by a single female, by one or more females in the presence of the mother, by more than one female in the absence of the mother, and by more than one female in the presence of females that participated in the prior reactivation. According to behavior, the females were classified as forager/egg-laying and egg-laying females. The oviposition by egg-laying females was always preceded by oophagy. All of the forager/egg-laying and egg-laying females that were dissected had been inseminated. The behavior displayed by egg-laying females is characteristic of brood parasitism and fits the parental parasitism hypothesis developed as an alternative pathway by which insect sociality could have arisen. The behaviors displayed by E. townsendi, together with those reported for Euglossa cordata, show that both species have bionomic traits that resemble the conditions suggested as precursors of the origin of eusociality.Received 12 June 2003; revised 6 April 2004; accepted 22 April 2004.     

Nesting patterns of European hedgehogs (Erinaceus europaeus) under northern conditions

Little information exists on the nesting of hedgehogs (Erinaceus europaeus) in the harsh northern environment, although this aspect has specific importance for the species’ survival in this region. We studied the nest types and nesting site selection of radio tagged hedgehogs in urban areas in Eastern Finland. Altogether, four different nest types were discovered and one of them (the pre-hibernation nest) is described here for the first time. Hedgehogs preferred residential areas for nesting during the mating and post-mating seasons while in the pre-hibernation season and hibernation period nesting was more common in forest areas. Nest constructions were closely related to materials found in the nest vicinity. During the active period, males used on average 24 nests and changed them 30 times, whereas females used 17 nests and changed them 26 times. The number of female nests and frequency of nest change increased from post-mating to pre-hibernation seasons, while in males both decreased. In general, female and male nests were situated within their established long-term home range, but in the pre-hibernation season over 30 % of male nests were situated outside of it. In post-mating and pre-hibernation seasons, females avoided other individuals of the same sex; this was demonstrated by the lack of nests in other female core areas. Hedgehogs hibernate for around 223 days, which is the longest reported hibernation period for this species. The results of the present study emphasise the importance of forest and diverse residential areas including forest patches for the hedgehog’s nesting habitat.     

An ecological analysis of mating biology of Xylocopa virginica in southern Ontario

1. Xylocopa virginica virginica Linnaeus is a wide‐ranging species with plastic nesting behaviour that appears to represent an intermediary between solitary and social nesting species. Over 3 years, a natural population was studied with the objective of quantifying the relationship among population dynamics, climate, female nest provisioning behaviour, and male mating strategy. 2. Males in the population congregated around female‐occupied nesting sites before the beginning of nest provisioning by females; both resident and satellite male mating strategies were observed. Overall, the present results are consistent with female defence polygyny. 3. Male mating strategies were consistent across the three breeding seasons of our study, in spite of annual variation in population size, sex ratio, and weather. Male mating behaviour was also consistent with that seen in other populations with longer breeding seasons. 4. Adult non‐breeding females that never leave nests are observed in nests throughout the breeding season and we hypothesise that males continue to defend territories after breeding females have mated because of a small probability they can mate with one of these non‐breeding females. 5. These results are important to our understanding of the relationship between mating systems and the evolution of sociality, contributing data on the role of ecological factors to male mating behaviour. Collection of such data for a variety of species that differ in sociality is necessary for the comparative analysis that is required to fully elucidate coevolution of mating systems and sociality.     

Few, small, and male: multiple effects of reduced nest space on the offspring of the solitary wasp, Euodynerus (Pareuodynerus) posticus (Hymenoptera: Vespidae)

The size of the preexisting wood cavities used as nests by aculeate Hymenoptera is expected to have consequences on fitness parameters such as offspring number and size. We evaluated the consequences of using small and large (three-times more voluminous) trap-nests by the solitary wasp, Euodynerus (Pareuodynerus) posticus (Herrich-Schaeffer). Following life-history and sex allocation theories, a number of non-mutually exclusive hypotheses were formulated: i.e. small nests either produce smaller or fewer offspring and/or more males, the cheaper sex. Wasps built about 28% more, but shorter brood cells in large nests, although their volume was still much higher in large nests. Adult males had smaller body size in small nests, but female size did not differ between large and small nests, possibly as an adaptive response against the future higher foraging costs of size-reduced females. Sex-ratio was often biased towards males in small nests. Mortality did not differ between large and small nests. We conclude that E.?(P.) posticus females would benefit from using larger nests, but that the sex-ratio would be probably overall unbalanced if females would not use also smaller, male-oriented tunnels.     

Anti-social and social behaviour of adolescent yellow-rumped caciques (Icterinae: Cacicus cela)

In an individually marked population in southeastern Peru, males and females of the polygynous, colonial yellow-rumped cacique delayed successful breeding until they were 3 years old. Adolescent (2-year-old) females competed for nest sites, built nests, laid fertile eggs, incubated, and sometimes fed nestlings, but always abandoned nests before they fledged young. Females that completed nests during their adolescent years were also more successful in establishing nest sites the following year. Adolescent females that failed to establish nest sites and those that lost nests to predators tended to leave the study area altogether. By attempting to nest early, adolescent females may facilitate future nesting attempts by forming coalitions with other females and assessing nest site quality in an area. Adolescent males, on the other hand, avoided aggressive interactions with other males, but were very aggressive towards adult females and fledglings. Adolescent males attacked females in nests, chased females to and from the colony, pecked and mounted recent fledglings, and, rarely, pulled nestlings out of nests. This harassment resulted in the death of nestlings and fledglings and was energetically costly to females, which often fought adolescent males while defending their young. Adolescent males preferentially harassed nestling-feeding females, avoided egg-laying females and were never observed attempting to mate with females. Adult males rarely interfered during any of these interactions. Such anti-social behaviour may function as a way of practising adult behaviour, as a means of eliminating future competitors, or may be an epiphenomenon of rising hormone levels and therefore of no adaptive significance.     

Behavioural notes on the Neotropical parasocial spider wasp Ageniella (Lissagenia) flavipennis (Banks) (Hymenoptera: Pompilidae), with host association

1. Ageniella is a species‐rich group of spider wasps restricted to the New World. Knowledge regarding the behaviour of this genus is based mainly on the Nearctic species, which have been reported to nest solitarily in the soil. This study reports for the first time the nesting behaviour, with host association of Ageniella (Lissagenia) flavipennis Banks. 2. Behavioural aspects on the nesting of A. flavipennis were observed from a study of six nests found in an Atlantic Forest conservation area in São Paulo State, Brazil. Host specimens were collected from a nest, as well as while being carried by an A. flavipennis individual. 3. The present study reports the A. flavipennis females cohabiting or nesting solitarily in mud nests, indicating that this spider wasp shows some lower level of parasociality. In addition, the spider Enoploctenus cyclothorax (Bertkau) was reported for the first time as host. As has been observed for other Ageniellini, females of A. flavipennis amputate the host's legs and transport the spider to the nest, flying or walking forward. 4. Communal behaviour has been reported for species of different genera of Pompilidae, such as Macromeris Lepeletier, Paragenia Bingham and Auplopus Spinola. Recent phylogenetic analyses suggest that the Lissagenia species are more closely related to the other Ageniellini genera than to the other Ageniella species. The present information on nesting and prey could contribute towards a more conclusive phylogenetic position of Lissagenia.     

Ecological and Behavioral Aspects of Nest Building in Wild Bonobos (Pan paniscus)

Data on nest building behavior were collected from free ranging bonobos (Pan paniscus). Considering the physical features of both nests and trees used for nest construction, bonobos appeared to show various preferences: (1) Trees with small leaves were chosen more frequently than expected; (2) The majority of nests were constructed in trees of the middle forest layer (15 to 30 m) and at a mean height of 16 m; (3) nests built during the day were located higher than night nests and differed in the general mode of construction (number of trees involved). Comparing nests made by mature males and females it became apparent that females (4) built nests higher, (5) constructed day nests more frequently and (6) used them for a longer time than males. Comparison of data collected at different study sites indicates site-specific differences concerning selection of trees and location of nesting sites. Behavioral observations made clear that nests were not only used for rest but also for other activities like feeding, social grooming and play. Moreover, the study provided the first evidence that bonobos build nests also in order to avoid potential or imminent conflicts with other party members. Field observations indicate that these nests symbolize a taboo zone and show another facet of the social abilities of bonobos for the solution of intra group conflicts. It is suggested that this behavior derives from the spatial intolerance shown by females during the process of weaning.     

A description of nesting behaviors,including factors impacting nest site selection,in black‐and‐white ruffed lemurs (Varecia variegata)

Nest site selection is at once fundamental to reproduction and a poorly understood component of many organisms’ reproductive investment. This study investigates the nesting behaviors of black‐and‐white ruffed lemurs, Varecia variegata, a litter‐bearing primate from the southeastern rainforests of Madagascar. Using a combination of behavioral, geospatial, and demographic data, I test the hypotheses that environmental and social cues influence nest site selection and that these decisions ultimately impact maternal reproductive success. Gestating females built multiple large nests throughout their territories. Of these, females used only a fraction of the originally constructed nests, as well as several parking locations as infants aged. Nest construction was best predicted by environmental cues, including the size of the nesting tree and density of feeding trees within a 75 m radius of the nest, whereas nest use depended largely on the size and average distance to feeding trees within that same area. Microhabitat characteristics were unrelated to whether females built or used nests. Although unrelated to nest site selection, social cues, specifically the average distance to conspecifics’ nest and park sites, were related to maternal reproductive success; mothers whose litters were parked in closer proximity to others’ nests experienced higher infant survival than those whose nests were more isolated. This is likely because nesting proximity facilitated communal crèche use by neighboring females. Together, these results suggest a complex pattern of nesting behaviors that involves females strategically building nests in areas with high potential resource abundance, using nests in areas according to their realized productivity, and communally rearing infants within a network of nests distributed throughout the larger communal territory.     

Seasonal nest characteristics of a facultatively social orchid bee,Euglossa viridissima,in the Yucatan Peninsula,Mexico

Euglossines have long been regarded as largely solitary, though some species are known to exhibit social behavior. We studied the nesting behavior of Euglossa viridissima over an annual cycle, comparing sociality and offspring production across the rainy (RS) and dry seasons (DS) in the Yucatan Peninsula of Mexico. Nests were built in both the RS and the DS, but with greater nest initiation and brood provisioning in the RS, presumably as a consequence of more floral resources at this time of year. Across the year, numerical sex ratios were female biased (0.7 as females/total); sex ratios varied across individual nests from 0.3 to 1.00, though without a clear relationship to sociality. Egg-to-adult development was quicker in females than males and, within a sex, quicker when ambient temperatures were higher. Multi-female (social) nests were only founded at the end of the RS and the beginning of the DS, coincided with the presence of Hymenopteran and Dipteran parasites in nests headed by solitary females. Reduced floral resources and a higher risk of parasitism, possibly coupled with higher female density or reduced nesting sites, may be factors favoring the formation of multi-female associations in this euglossine. Better nourishment of foundress females in the RS may improve lifespan and permit overlapping generations which, coupled with the kin structure of their nests, may favor social nesting in E. viridissima.     

Colony Development and Serial Polygyny in the Primitively Eusocial Wasp Belonogaster juncea juncea (Vespidae: Polistinae)

In Cameroon, the social ropalidiine wasp Belonogaster juncea juncea mostly selects human constructions for nesting, but nests installed in the shelter of large boulders were also noted. Nests were founded throughout the year by one female (25.5%) or by associations of two to eight females. Among the 49 colonies studied from foundation, 24 produced at least one adult, and only 5 produced males among the eight that succeeded in producing sexual individuals. After the first dominant female disappeared (died or abandoned the nest, from 77 to 196 days after foundation) the remaining females fought among themselves, leading to a drastic decrease in the total number of brood. They completely abandoned 22 of 24 surviving nests, over a mean colony cycle lasting about 7 months. Hence the colony and nesting cycles were congruent, illustrating that determinate nesting corresponds to the majority of the cases in this species. In the other two nests, some females remained on the nest and began a new colonial cycle illustrating semi-independence between colony and nesting cycles. Up to four successive cycles were completed in this way in the same nest, thus illustrating the phenomenon of serial polygyny (intermediate between determinate and indeterminate nesting cycles). Serial polygyny in independent founding polistine wasps, previously documented only once, has implications for studies attempting to identify factors involved in regulating the colony and nesting cycles in tropical social wasps.