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.     

Mixed species nesting associations in Darwin's tree finches: nesting pattern predicts predation outcome

Predation is the main cause of passerine nesting failure. Traditionally, large intraspecific group size is thought to accrue individuals with fitness benefits from increased predator vigilance and hence lower predation risk. To date, few studies have investigated interspecific group size in relation to predation risk. In the present study, we examined predation outcome in Darwin's small tree finch, Camarhynchus parvulus , in nests with many or few interspecific neighbours. We tested the predictions: (1) nests in mixed associations have lower predation than do more solitary nests; (2) mixed species nesting associations covary with nest site vegetation characteristics; (3) older (i.e. presumably experienced) males more commonly nest in mixed associations than younger males; (4) older males select more concealed nesting sites; and (5) controlling male age, females prefer to pair with males in mixed associations than at solitary nests. Almost half of all nests occurred in mixed associations (46%) compared to solitary nests (54%), and the overall distribution of nests was decidedly nonrandom, displaying a bimodal pattern. Nest site vegetation characteristics of the focal species were inconsistently associated with nesting pattern, but older males did select more concealed nesting sites. Controlling differences in surrounding vegetation characteristics, mixed nesting associations experienced markedly lower predation than solitary nests, and females showed a preference for males in mixed associations, as demonstrated by higher male pairing success.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 313–324.     

Colony characteristics influence the risk of nest predation of a polydomous ant by a monotreme

Although predation of individual social insect workers has little effect on colony fitness, nest predation may be a significant selective agent because it can result in substantial loss of reproductive success. Surprisingly, the consequences of predation on social insect nests are poorly understood. In the present study, we investigate the factors that correlate with the probability of predation by echidnas, Tachyglossus aculeatus , on nests and colonies of the facultatively polydomous meat ant, Iridomyrmex purpureus. In particular, we investigate whether colony fragmentation provides a mechanism for reducing the costs of echidna predation. Over 2 years, 138 of the 140 colonies on our study site were depredated. Nest predation was most common in woodlands but with no obvious seasonal patterns. The probability of nest predation was positively correlated with the size of the nest, and negatively correlated with the density of surrounding nests. Although polydomous colonies are at a similar risk of predation by echidnas, the proportion of depredated nests is negatively correlated with the number of nests; thus, the probability that one or more nests avoid predation is increased with increasing nest numbers. Surprisingly, we found no influence of the level of echidna predation on colony growth, measured by either changes in the number of nests or the number of nest entrance holes.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 1–8.     

The chemical strategies used by Polistes nimphus social wasp usurpers (Hymenoptera Vespidae)

Polistes foundresses can behave as facultative social parasites when, instead of founding their own nest, they usurp colonies of the same or a different species and temporary use the host workforce to raise their own brood. Conspecific usurpation appears to be common among Polistes wasps, but nothing is known about the mechanisms that these facultative social parasites use to have themselves accepted within usurped colonies. Using behavioural tests, we studied the chemical strategies employed by females of Polistes nimphus when they behave as facultative social parasites in colonies of the same or of a different species. We hypothesized that usurpers would mark host nests with their own odours and/or acquire host nest odours in order to camouflage their real identity from host workers. Our results indicated that P. nimphus usurpers used different chemical strategies depending on host nest species: they acquired conspecific host odours but marked heterospecific host combs with their own odours.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 505–512.     

Testing the function of the domed nests of Saltmarsh Sharp-tailed Sparrows

ABSTRACT.   Saltmarsh Sharp-tailed Sparrows ( Ammodramus caudacutus ) build ground nests, often with a closely-woven dome, in marshes that frequently flood during high tides. To test the hypothesis that domed nests help reduce the loss of eggs and chicks due to flooding or predation, we examined the characteristics and fate of 102 nests at sites along the coast of Connecticut. To test whether nest structure was tailored to suit microhabitat conditions, we also measured vegetation characteristics around nests. Finally, we conducted artificial nest-flooding experiments to determine whether removal of domes reduced egg retention during flooding. We found no significant effects of nest structure on breeding success or failure, and few significant correlations between nest structure and microhabitat. The height of nests above the ground, however, increased with vegetation height, supporting the hypothesis that nest construction is influenced by flooding, but not supporting the hypothesis that predation risk is important. Dome removal experiments showed that domes have a highly significant effect on the retention of eggs during flooding, suggesting that domes help eggs survive the regular tidal flooding of marshes.     

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.     

Appearance and vulnerability of artificial duck nests to avian predators

Use of artificial nests in studies of nest success in birds has become increasingly popular, but this practice has been criticised because the appearance and success of artificial nests may differ from natural nests. Typically, artificial nests are presented with few visible eggs throughout the entire exposure period while natural nests during their life have different appearances, possibly producing stage-dependent predation rates. In ducks, nests may appear with openly visible eggs, eggs covered with nest material and covered by an incubating female. To test how nest appearance may affect nest survival, I simulated such duck nest appearances, and used direct observations to record responses of marsh harriers Circus aeruginosus , the dominant duck nest predator at the Lake Engure, Latvia, during the breeding seasons 2000–2002. I recorded 274 harrier flights over all types of artificial nests, 74 nest discoveries and 54 predation events. Logistic regression analyses showed that nest appearance was a significant predictor of the fate of the nest: nests with openly visible eggs were discovered more often than nests covered with nest material. Nests with dummy females had discovery rates similar to the average of all artificial nest types suggesting that they best reflect the predation risk of artificial nests and could be used in future studies. After discovering nests covered with dummy females, harriers never attacked but tried to scare them off to get access to the eggs. In 52% of cases of nest discovery, harriers gave up before touching the dummy. This shows that duck passive nest defence may prevent clutch predation.     

Brood parasitism is associated with increased bacterial contamination of host eggs: bacterial loads of host and parasitic eggs

Factors related to bacterial environment of nests are of primary interest for understanding the causes of embryo infection and the evolution of antimicrobial defensive traits in birds. Nest visitors such as parasites could act as vectors for bacteria and/or affect the hygienic conditions of nests and hence influence the nest bacterial environment. In the present study, we explored some predictions of this hypothetical scenario in the great spotted cuckoo (Clamator glandarius)–magpie (Pica pica) system of brood parasitism. Great spotted cuckoos visit the nests of their magpie hosts and frequently damage some of the host eggs when laying eggs or on subsequent visits. Therefore, it represents a good system for testing the effect of nest visitors on the bacterial environment of nests. In accordance with this hypothesis, we found that the bacterial load of magpie eggshells was greater in parasitized nests, which may suggest that brood parasitism increases the probability of bacterial infection of magpie eggs. Moreover, comparisons of bacterial loads of cuckoo and magpie eggs revealed that: (1) cuckoo eggshells harboured lower bacterial densities than those of their magpie hosts in the same nests and (2) the prevalence of bacteria inside unhatched eggs was higher for magpies than for great spotted cuckoos. These interspecific differences were predicted because brood parasitic eggs (but not host eggs) always experience the bacterial environments of parasitized nests. Therefore, the results obtained in the present study suggest that parasitic eggs are better adapted to environments with a high risk of bacterial contamination than those of their magpie hosts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 836–848.     

Sunny side up: lethally high, not low, nest temperatures may prevent oviparous reptiles from reproducing at high elevations

Oviparous (egg-laying) lizards and snakes generally inhabit warmer climates than do related viviparous (live-bearing) taxa. This pattern is widely attributed to the failure of oviparous reproduction in cold climates, but the thermal regimes of potential nest-sites above and below the elevational cut-off for oviparous reproduction have never been quantified. We studied oviparous ( Bassiana duperreyi ) and viviparous ( Eulamprus heatwolei ) scincid lizards at such a site in the Brindabella Range of south-eastern Australia. Miniature data-loggers monitored temperatures of nest-sites and lizards in midsummer, partway through the incubation period of eggs in natural nests. Our results contradict the simplistic notion that mean nest temperatures determine this elevational limit for oviparity. Instead, potential nest-sites with average temperatures suitable for embryogenesis in Bassiana are available well above the threshold elevation. However, thermal minima decrease consistently with elevation and thus the maximum temperature needed for any given mean incubation temperature increases rapidly with elevation. Potential nest-sites above the elevational threshold can only attain mean temperatures high enough to sustain embryogenesis by having lethally high thermal maxima. Such nest-sites are available close to the soil surface, but cannot support development. In contrast, behavioural thermoregulation allows viviparous lizards to maintain high mean body temperatures concurrently with relatively low maximum temperatures, regardless of elevation. Paradoxically, oviparous reptiles may be restricted to low elevations not because nests that provide appropriate mean incubation temperatures are unavailable further up the mountain, but because eggs laid in such shallow nests would overheat.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 325–334.     

Multiple maternity in black-headed gull Larus ridibundus clutches as revealed by protein fingerprinting

Social monogamy with biparental care is the norm in gulls Laridae , but egg colour variation suggests that some nests may contain mixed clutches laid by more than one female. Here we use protein fingerprinting of egg albumen to assess the occurrence of mixed maternity clutches in three colonies of black-headed gulls. Among 160 analysed clutches with >1 egg, 34% contained eggs from more than one female, and 15% of the eggs in clutches >1 came from other females than the major female (laying most eggs in nest). Among clutches with 2–3 eggs 28% were mixed, and among clutches with 4 or more eggs 89% contained eggs from two or more females. There were significantly fewer eggs from the major female in mixed nests (mean=2.06±0.63 SD) than in non-mixed nests (mean=2.82±0.43 SD). In nests without evidence of female conflict, hatching success of minority eggs was similar to that of eggs from the major female (12.5 and 8.4%, respectively). In 21% of mixed maternity nests, one or more minority eggs was buried or punctured, and 25% of eggs from major females were also found evicted, suggesting conflict between females and rejection of eggs. Intra-specific nest parasitism seems the most likely cause of mixed clutches, but there are also other possible causes.     

Differences in morphology and colour pattern of shiny cowbird (Molothrus bonariensis) eggs found in nests of two hosts

Genetic differentiation among shiny cowbird (Molothrus bonariensis) females that use different hosts indicates that in this brood parasite, host use is not random at an individual level. We tested whether there exist differences in morphology and coloration between eggs of shiny cowbirds laid in the nests of two different hosts, the chalk‐browed mockingbird (Mimus saturninus) and the house wren (Troglodytes aedon). We took morphometric measures of shiny cowbird eggs found in nests of mockingbirds and wrens and analysed their coloration using digital photography and reflectance spectrometry. We found that shiny cowbird eggs found in mockingbird nests were wider and more asymmetric than those found in wren nests. In addition, cowbird eggs coming from mockingbird nests were brighter and had higher relative red reflectance than those coming from wren nests. Our results show that shiny cowbird eggs laid in nests of two different hosts vary in shape and background colour, but not in spotting pattern. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 838–845.     

Egg Coloration and Recognition of Conspecific Brood Parasitism in Eastern Bluebirds

Individual eastern bluebird (Sialia sialis) females produce clutches of eggs with unique coloration and older females and females in better body condition lay more pigmented blue‐green eggs. Conspecific brood parasitism in this species is not uncommon and bluebirds occasionally reject what appear to be normal eggs by moving them to the periphery of the nest. I used UV‐visual reflectance spectrometry to objectively measure coloration of eggs and nest material. To estimate the conspicuousness of the trait, I calculated the contrast between eggs and background nest material. I found high achromatic and chromatic contrast between the coloration of eggs and of the nests, suggesting that bluebird eggs are highly conspicuous. To test the hypothesis that expression of blue‐green coloration eggs facilitates recognition of eggs laid by conspecific brood parasites, I cross‐fostered individual eggs into host nests during egg laying and monitored the fate of those eggs. I found no support, however, for the hypothesis that egg coloration facilitates discrimination of parasitic eggs from host eggs.     

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.     

Population consequences of huge nesting aggregations ofRopalidia plebeiana (Hymenoptera: Vespidae)

Nest survival rates and reproductive rates of females of the Australian paper wasp, Ropalidia plebeiana, in nest aggregations under a concrete bridge were studied. The annual colony cycle commenced in August with the reutilization of old nests and by the founding of new nests, mainly by associations of foundresses. Distribution patterns of the number of foundresses per nest was nearly random on new nests while contagious on old nests. About one-third of the increase in number of nest was achieved by dividing large, old nests and two-thirds by founding new nests. Nest survival rates for old nests (August to April) and new nests (November to April) were 89.6% and 88.8% respectively, far higher rates than those of other polistine wasps so far reported. The numbers of nests and reproductive females increased during a colony cycle by 2.30 and 10.98 times, respectively. Thus, huge aggregations of nests are probably beneficial for this species by enabling the realization of high reproductive rates. We could find no density-dependent effect in nest growth rate or production of female reproductives. Ratio of cells parasitized by ichneumonid wasps was low. Nests in the central part of an aggregation were prasitized at significantly lower rates than nests on the periphery or scattered nests outside a dense aggregation, suggesting a selfish herd effect.     

Social behaviour ofRopalidia fasciata (Hymenoptera: Vespidae) females on satellite nests and on a nest with multiple combs

The social behaviour ofRopalidia fasciata females on 3 satellite nests and on a nest with multiple, independent combs was observed. Many females often visited satellite nests, and even dominant foundresses, who usually spent more of their time on the nest, became active when satellite nests were being constructed. Three foundresses laid eggs on a satellite nest and eggs laid by 2 of these (who performed foraging) survived. On another satellite nest, only the dominant foundress laid an egg which survived. On a nest consisting of 2 independent combs, females frequently shifted between the 2 combs even after the emergence of many adults. Constructions of satellite nests and of multiple independent combs are considered to be adaptive for this species living in Okinawa, where most of nests are destroyed by frequent typhoons or abandoned after predation by ants.     

Egg morphology fails to identify nests parasitized by conspecifics in common pochard: a test based on protein fingerprinting and including female relatedness

Conspecific brood parasites lay eggs in nests of other females of the same species. A variety of methods have been developed and used to detect conspecific brood parasitism (CBP). Traditional methods may be inaccurate in detecting CBP and in revealing its true frequency. On the other hand more accurate molecular methods are expensive and time consuming. Eadie developed a method for revealing CBP based on differences in egg morphology. That method is based on Euclidean distances calculated for pairs of eggs within a clutch using standardized egg measurements (length, width and weight). We tested the applicability of this method in the common pochard Aythya ferina using nests that were identified as parasitized (39 nests) or non‐parasitized (16 nests) based on protein fingerprinting of eggs. We also analyzed whether we can distinguish between parasitic and host eggs in the nest. We found that variation in MED can be explained by parasitism but there was a huge overlap in MED between parasitized and non‐parasitized nests. MED also increased with clutch size. Using discriminant function analysis (DFA) we found that only 76.4% of nests were correctly assigned as parasitized or non‐parasitized and only 68.3% of eggs as parasitic or host eggs. Moreover we found that MED in parasitized nests increased with relatedness of the females that laid eggs in the nest. This finding was supported by positive correlation between MED and estimated relatedness in female–female pairs. Although variation in egg morphology is associated with CBP, it does not provide a reliable clue for distinguishing parasitized nests from non‐parasitized nests in common pochard.     

Decoding dumping ducks

Conspecific brood parasitism, where females of the same species lay eggs in each other's nests, is common in waterfowl, and is usually considered costly to host females, which are stuck looking after eggs and chicks that are not their own. However, since female waterfowl often exhibit an unusual propensity to nest near where they were born, there has been some uncertainty over whether, in ducks and geese, laying in nests of conspecifics really is parasitism. Do parasitic and host females tend to be related? And is parasitism actually a form of cooperation in disguise? In a population in Hudson Bay, Andersson & Waldeck (this issue) found that ‘parasitic’ eggs in nests of the common eider, Somateria mollissima sedentaria, are more closely related to host eggs than expected by chance. In fact, host and ‘donor’ eggs are more closely related than are females breeding at neighbouring nests. The Hudson Bay population of common eiders is unusual, because unlike in more benign climates, females do not tend to breed near their natal nest. Spatial proximity alone cannot account for the high relatedness between host eggs and ‘dumped’ or donor eggs. Instead, the high relatedness values are probably the result of active recognition, where females favour kin, either when dumping or accepting eggs. These new data, along with evidence indicating that the donor lays the first egg in the nest nearly half the time, suggest that what appears to be parasitism in common eiders may be a form of kin‐based cooperation.     

A nondestructive method for estimating the age of Clapper Rail eggs

ABSTRACT.   Estimates of nest survival rate are often based on the assumption that the rate is constant during the nest cycle. Because violating this assumption can generate biased estimates, determining the age of a nest is important. For precocial marsh birds, including Clapper Rails ( Rallus longirostris ), current estimates of nest survival do not take this source of bias into account because a standardized, nondestructive method for aging eggs in not available. We developed a nondestructive flotation technique using known-age nests that can be used to estimate the projected hatch date of Clapper Rail eggs within 3 d of the actual hatch date. This precision is limited in part due to variance in the length of incubation between nests and the common occurrence of asynchronous hatching. However, for investigators assessing the fate of Clapper Rail nests, our technique provides an effective and nondestructive method for projecting the hatch date of Clapper Rail eggs, and will facilitate unbiased estimates of nest survival rates.     

Incubation regimes of cold-climate reptiles: the thermal consequences of nest-site choice, viviparity and maternal basking

Cold-climate reptiles show three kinds of adaptation to provide warmer incubation regimes for their developing embryos: maternal selection of hot nest sites; prolonged uterine retention of eggs; and increased maternal basking during pregnancy. These traits may evolve sequentially as an oviparous lineage invades colder climates. To compare the thermal consequences of these adaptations, I measured microhabitat temperatures of potential nest sites and actual nests of oviparous scincid lizards ( Bassiana duperreyi ), and body temperatures of pregnant and non-pregnant viviparous scincid lizards ( Eulamprus heatwolei ). These comparisons were made at a site where both species occur, but close to the upper elevational limit for oviparous reptiles in south-eastern Australia. Viviparity and maternal basking effort had less effect on mean incubation temperature than did maternal nest-site selection. Eggs retained in utero experienced bimodal rather than unimodal diel thermal distributions, but similar mean incubation temperatures. Often the published literature emphasizes the ability of heliothermic (basking) reptiles to maintain high body temperatures despite unfavourable ambient weather conditions; this putative ability is central to many hypotheses on selective forces for the evolution of viviparity. In cold climates, however, opportunities for maternal thermoregulation to elevate mean body temperatures (and thus, incubation temperatures) above ambient levels may be severely limited. Hence, at least over the broad elevational range in which oviparous and viviparous species live in sympatry, maternal selection of 'hot' nests may be as effective as is viviparity in providing favourable incubation regimes.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 145–155.     

The influence of oxygen stress on female choice for male nest structure in the common goby

This study examines the role of dissolved oxygen in modifying female mate choice in the common goby, Pomatoschistus microps. Experimental manipulations of nests showed that under normal (saturated) oxygen conditions, females preferred to spawn in nests with the most elaborate construction, that is, those with the most sand on top and the smallest entrance. Such manipulated nests proved less vulnerable to detection by an egg predator, the shore crab, Carcinus maenas, but these small entrances may reduce oxygenation of eggs. Indeed, in a low oxygen environment females with a choice between nests of high build and nests of low build did not significantly prefer either. This may have been due to weakened preference or perhaps detection of a mismatch between the male's parental quality and the increased ventilation needs of eggs when nest entrance sizes are small and ambient oxygen levels are low. When nests were not manipulated, those males that built small entrances fanned their eggs more often, such that hatching success in the absence of a predator was not related to the initial size of the nest entrance area. Thus, under normal oxygen conditions males may initially build nests with the smallest entrance they are capable of ventilating successfully, and females choosing such males gain from nest camouflage. Under low oxygen, the risk that manipulated males may be unable to compensate fully may outweigh such benefits, and females may use other criteria that signal willingness to provide parental care. Copyright 1999 The Association for the Study of Animal Behaviour.