How rapidly can maternal behavior affecting primary sex ratio evolve in a reptile with environmental sex determination?

Theoretical models identify maternal behavior as critical for the maintenance and evolution of sex ratios in organisms with environmental sex determination (ESD). Maternal choice of nest site is generally thought to respond more rapidly to sex ratio selection than environmental sensitivity of offspring sex (threshold temperatures) in reptiles with temperature-dependent sex determination (TSD, a form of ESD). However, knowledge of the evolutionary potential for either of these traits in a field setting is limited. I developed a simulation model using local climate data and observed levels of phenotypic variation for nest-site choice and threshold temperatures in painted turtles (Chrysemys picta) with TSD. Both nest-site choice and threshold temperatures, and hence sex ratios, evolved slowly to simulated climate change scenarios. In contrast to expectations from previous models, nest-site choice evolved more slowly than threshold temperatures because of large climatic effects on nest temperatures and indirect selection on maternally expressed traits. A variant of the model, assuming inheritance of nest-site choice through natal imprinting, demonstrated that natal imprinting inhibited adaptive responses in female nest-site choice to climate change. These results predict that females have relatively low potential to adaptively adjust sex ratios through nest-site choice.     

Making good choices with variable information: a stochastic model for nest-site selection by honeybees

A density-dependent Markov process model is constructed for information transfer among scouts during nest-site selection by honeybees (Apis mellifera). The effects of site quality, competition between sites and delays in site discovery are investigated. The model predicts that bees choose the better of two sites more reliably when both sites are of low quality than when both sites are of high quality and that delay in finding a second site has most effect on the final choice when both sites are of high quality. The model suggests that stochastic effects in honeybee nest-site selection confer no advantage on the swarm.     

Are nest sites actively chosen? Testing a common assumption for three non-resource limited birds

Many widely-accepted ecological concepts are simplified assumptions about complex situations that remain largely untested. One example is the assumption that nest-building species choose nest sites actively when they are not resource limited. This assumption has seen little direct empirical testing: most studies on nest-site selection simply assume that sites are chosen actively (and seek explanations for such behaviour) without considering that sites may be selected randomly. We used 15 years of data from a nestbox scheme in the UK to test the assumption of active nest-site choice in three cavity-nesting bird species that differ in breeding and migratory strategy: blue tit (Cyanistes caeruleus), great tit (Parus major) and pied flycatcher (Ficedula hypoleuca). Nest-site selection was non-random (implying active nest-site choice) for blue and great tits, but not for pied flycatchers. We also considered the relative importance of year-specific and site-specific factors in determining occupation of nest sites. Site-specific factors were more important than year-specific factors for the tit species, while the reverse was true for pied flycatchers. Our results show that nest-site selection, in birds at least, is not always the result of active choice, such that choice should not be assumed automatically in studies of nesting behaviour. We use this example to highlight the need to test key ecological assumptions empirically, and the importance of doing so across taxa rather than for single “model” species.     

Egg-laying behaviour and nest-site selection of domestic hens kept in small floor-pens

The development of egg-laying behaviour and nest-site choice is described in seven flocks of White Leghorn hens housed in small floor-pens. Special interest was directed towards social dominance and the effect of the presence of a cock on the hen's nest-site choice. In the weeks before the hens came into lay, they frequently examined the nests. Nest-entries only occurred some days before, or in the hours preceding, the first oviposition. Some hens were very conservative in their nest-site choice within clutches as well as between clutches. A number of hens only persevered in their nest-site choice within clutches, while a third category did not show any consistency in their choices. Individual hens were consistent in using either the floor or a nest-box as a nesting site.No relationship was found between social status and nest-site choice, nor between social status and floor-laying. Some evidence was provided that the presence of a cock could reduce the number of floor-eggs in a flock provided with roll-away boxes.It is postulated that the nest-examinations in the weeks before laying may serve for the selection of a suitable nest-site.     

Population sex ratios under differing local climates in a reptile with environmental sex determination

Populations that experience different local climates, such as those along a latitudinal gradient, must match life history traits to local environmental conditions. In species with temperature-dependent sex determination, such as many reptiles, population sex ratio is strongly influenced by local climate, yet local climate differs substantially among populations in geographically-widespread species. We studied the painted turtle at three sites across the species’ geographic range to gain a mechanistic understanding of how sex ratios are produced under different local climates. We combined data on maternal nest-site choice, nest incubation temperature, and the resultant offspring sex ratio of populations across a climatic gradient, to demonstrate how geographic variation in behavior and physiology translates into sex ratios among populations of a widely-distributed species. We found that populations across the species’ geographic range match incubation conditions with local climatic conditions through population-specific adjustment of maternal nest-site choice. Incubation temperatures during the thermosensitive period were cooler and clutches were more male-biased in the south, with populations farther north having warmer incubation temperatures and more female-biased sex ratios, yet adult sex ratios were not strongly biased in any population. Most components of maternal nest-site choice varied latitudinally among populations, suggesting that the species may have a considerable repertoire for responding to climate change through adjustment of nest-site choice.     

Retirement investment theory explains patterns in songbird nest-site choice

When opposing evolutionary selection pressures act on a behavioural trait, the result is often stabilizing selection for an intermediate optimal phenotype, with deviations from the predicted optimum attributed to tracking a moving target, development of behavioural syndromes or shifts in riskiness over an individual''s lifetime. We investigated nest-site choice by female golden-winged warblers, and the selection pressures acting on that choice by two fitness components, nest success and fledgling survival. We observed strong and consistent opposing selection pressures on nest-site choice for maximizing these two fitness components, and an abrupt, within-season switch in the fitness component birds prioritize via nest-site choice, dependent on the time remaining for additional nesting attempts. We found that females consistently deviated from the predicted optimal behaviour when choosing nest sites because they can make multiple attempts at one fitness component, nest success, but only one attempt at the subsequent component, fledgling survival. Our results demonstrate a unique natural strategy for balancing opposing selection pressures to maximize total fitness. This time-dependent switch from high to low risk tolerance in nest-site choice maximizes songbird fitness in the same way a well-timed switch in human investor risk tolerance can maximize one''s nest egg at retirement. Our results also provide strong evidence for the adaptive nature of songbird nest-site choice, which we suggest has been elusive primarily due to a lack of consideration for fledgling survival.     

The individual nest box as a super-stimulus for domestic hens

Following reports of inconsistent nest choice by hens in groups, choice of nest boxes by solitary hens was investigated. Thirty-two hens of a White Leghorn strain were each isolated with six nest boxes while they laid six eggs. From 16 hens, eggs were collected daily. Only one of these birds laid all her eggs in one nest box, and others used up to four boxes. The other 16 were allowed to accumulate eggs, and these were significantly more consistent in nest choice. Five used only one box. However, the other 11 of this group showed incomplete consistency.An explanation for inconsistent nest choice was suggested by a second experiment. The importance of enclosure in nest-site selection was tested in hens of White Leghorn and Rhode Island Red strains. Individuals about to lay their first egg were isolated with a choice of two types of nest box, selected from five types of different enclosure. Exposed nests, of similar dimensions to natural nests, were rarely used by either strain. By contrast, more enclosed nests were strongly selected. These results indicate that enclosure is an important stimulus in nest-site selection, and that artificial nest boxes, which are more enclosed than natural sites, act as super-normal stimuli (or “super-stimuli”). Choices between super-stimuli are likely to be equivocal, which accounts for inconsistency in the use of nest boxes and other features of the nesting behaviour of domestic hens.     

Role of histamine as a putative inhibitory transmitter in the honeybee antennal lobe

Background

Nests are built in various animal taxa including fish. In systems with exclusive male parental care, the choice of a nest site may be an important component of male fitness. The nest site may influence male attractiveness as a mate, and male, embryo, and juvenile survival probabilities. Reproductively active three-spined stickleback males establish and defend a territory in which they build a nest. Territories can differ remarkably in qualities that influence male and female reproductive success like predation risk or abiotic factors such as dissolved oxygen concentration or lighting conditions. The latter may be important because in sticklebacks the extended visual capability into the ultraviolet (UV) wave range plays a role in female mate choice. Males are thus expected to be choosy about the habitat in which they will build their nest.

Results

We tested nest-site choice in male three-spined sticklebacks with respect to different UV lighting conditions. Reproductively active males were given the simultaneous choice to build their nest either in an UV-rich (UV+) or an UV-lacking (UV-) environment. Males exhibited no significant nest-site preferences with respect to UV+ or UV-. However, larger males and also heavier ones completed their nests earlier.

Conclusion

We found that UV radiation as well as differences in luminance had no influence on nest-site choice in three-spined sticklebacks. Males that built in the UV-rich environment were not different in any trait (body traits and UV reflection traits) from males that built in the UV-poor environment. There was a significant effect of standard length and body mass on the time elapsed until nest completion in the UV experiment. The larger and heavier a male, the faster he completed his nest. In the brightness control experiment there was a significant effect only of body mass on the duration of nest completion. Whether nest building preferences with respect to UV lighting conditions are context dependent needs to be tested for instance by nest-site choice experiment under increased predation risk.     

Nest-site selection in a fish species with paternal care

Hydrobiologia - Fish that perform paternal care may increase their fitness by choosing nest sites that enhance survival and development of embryos. We studied nest-site choice with respect to...     

Nest-site philopatry and selection for environmental sex determination

The reason for the frequent occurrence of environmental sex determination (ESD) in reptiles is still not well understood, although much effort has been devoted to solving the issue. Stimulated by the occurrence of nest-site philopatry in some species, this paper examines a diploid model of the influence of nest-site philopatry on the evolution of ESD. Analysis shows that nest-site philopatry can lead to ESD because the fitnesses of sons and daughters are not influenced in the same way by nest-site quality. Daughters inherit the nest site and thus benefit more than sons from a high-quality nest site. Conversely, the fitness of daughters at low-quality nest sites is lower compared to the fitness of sons. Therefore, genes causing ESD can spread by causing the production of more sons at low-quality nest sites and more daughters at high-quality nest sites. Suggestions are made to test empirically whether nest-site philopatry led to the evolution of ESD. This revised version was published online in July 2006 with corrections to the Cover Date.     

Nest-site partitioning in a strandveld shrubland bird community

Nest-site selection may vary adaptively among co-existing species as a result of competitive interactions among the species or in response to density-dependent nest predation. We examined nest-site characteristics and degree of partitioning among 14 co-existing bird species breeding in dwarf strandveld shrubland at Koeberg Nature Reserve, South Africa. Habitat characteristics of nest sites differed significantly among species, indicating strong nest-site partitioning. The principal variables distinguishing nest-site characteristics among species (substrate height, cover at 1m, cover at 2m) were features that distinguish tall-shrub species which are strongly associated with patches of Dune Thicket from shorter-shrub species which are strongly associated with more open Sand Plain Fynbos. Two groups of species which differ primarily in their selection of habitat patch type within the patchy nesting landscape were identified. One group nested predominantly in low shrubs associated with open Sand Plain Fynbos. A second group nested at a variety of heights within tall-shrub species associated with Dune Thicket. Species within each of these groups differed from each other in their relative nest height (ratio of nest height to substrate height), distance of nest from foliage edge, nest concealment and choice of particular substrate plant species. On average, stepwise discriminant function analysis classified 52.6% of the nest sites correctly as belonging to one of the 14 species, considerably better than the expected random classification of 11.8%. Individual species' classifications were 2–12 times better than a random classification. We suggest that high levels of nest predation may have selected for strong nest-site partitioning in this community.     

Effect of methodological and ecological approaches on heterogeneity of nest-site selection of a long-lived vulture

The application of scientific-based conservation measures requires that sampling methodologies in studies modelling similar ecological aspects produce comparable results making easier their interpretation. We aimed to show how the choice of different methodological and ecological approaches can affect conclusions in nest-site selection studies along different Palearctic meta-populations of an indicator species. First, a multivariate analysis of the variables affecting nest-site selection in a breeding colony of cinereous vulture (Aegypius monachus) in central Spain was performed. Then, a meta-analysis was applied to establish how methodological and habitat-type factors determine differences and similarities in the results obtained by previous studies that have modelled the forest breeding habitat of the species. Our results revealed patterns in nesting-habitat modelling by the cinereous vulture throughout its whole range: steep and south-facing slopes, great cover of large trees and distance to human activities were generally selected. The ratio and situation of the studied plots (nests/random), the use of plots vs. polygons as sampling units and the number of years of data set determined the variability explained by the model. Moreover, a greater size of the breeding colony implied that ecological and geomorphological variables at landscape level were more influential. Additionally, human activities affected in greater proportion to colonies situated in Mediterranean forests. For the first time, a meta-analysis regarding the factors determining nest-site selection heterogeneity for a single species at broad scale was achieved. It is essential to homogenize and coordinate experimental design in modelling the selection of species' ecological requirements in order to avoid that differences in results among studies would be due to methodological heterogeneity. This would optimize best conservation and management practices for habitats and species in a global context.     

Traditional versus non-traditional nest-site choice: alternative decision strategies for nest-site selection

In order to understand habitat selection, it is important to consider the way individual animals assess the suitability of a future reproductive site. One way of investigating mechanisms (such as those involved in nest site selection) is to examine breeding success and habitat characteristics in terms of animals returning to a place where they have already reproduced and using the same location over successive years or searching for new alternatives. This approach seems especially suitable for testing recent hypotheses suggesting that nest site selection is an integrative process that includes the use of social information (e.g. past breeding success of conspecifics). Determining the factors that elicit conservative or innovative behaviour regarding nest-site selection could be important for improving our understanding of habitat selection decisions in animals. More than half of the nests of the long-distance migratory lesser grey shrike Lanius minor, are built in the same or neighbouring trees. We found no evidence that habitat characteristics influence nest-site tradition. On the contrary, social information in terms of the presence of conspecifics and past reproductive success in terms of complete nest failures due to nest predation (but not detailed information such as variation in fledgling number) influenced nest-site tradition. Hence, social information and past reproductive success may play a role in nest-site choice in this species. Our results further demonstrate that previous experience with a nest site does not appear to be beneficial.     

Testing the importance of individual nest-site selection for a social and group-living vulture

Nest-site selection by species is expected to be adaptive and lead to improved breeding productivity, but in some settings, there exist mismatches between preferred nesting habitat and breeding productivity. We tested the expectation that nest-site selection is adaptive in a sample of 63 nests of a long-lived social species that breeds and forages in groups: the critically endangered white-backed vulture (Gyps africanus). By studying breeding groups in the same area, we controlled for landscape-level effects on habitat selection and investigated how fine-scale nest-site characteristics affect breeding productivity. We developed models to assess how nine characteristics of nest sites selected by breeding vultures compared with 70 random trees and tested associations between these characteristics and breeding productivity. White-backed vultures selected nest sites in taller trees (>7 m), but neither tree height nor any other nest-site characteristics had a clear effect on breeding productivity. Vultures selected nest trees closer to each other than random trees, and the associations between nest density, nearest neighbour distance and breeding success were all positive. These positive associations and the absence of an observable effect between nest-site characteristics and breeding productivity suggest that for this semi-colonial breeder, the social imperative of proximity to conspecifics (i.e., nesting near other vultures and group foraging) may be more important than individual nest-site selection.     

Do Fleas,and/or Old Nest Material,Influence Nest-site preference in Hole-nesting Passerines?

This study examined nest-site choice in a migratory population of pied flycatchers (Ficedula hypoleuca) and sympatric populations of three resident tit species (Parus major, P. caeruleus and P. palustris) in central Sweden. All four species are secondary-cavity nesters which naturally breed in pre-formed tree cavities but readily use artificial nest boxes. We asked whether flycatchers and tits discriminate between nest boxes that: 1. Are ‘empty’; 2. Contain old nests without ectoparasites (fleas Ceratophyllus sp.); or 3. Contain old nests with ectoparasites. We found that pied flycatchers preferred nest boxes containing old nests, regardless of whether these nests held parasites. In contrast, tits did not discriminate between the three types of boxes. Tits may pay a cost for their lack of choosiness: after the breeding season, tit nests contained more fleas than flycatcher nests. Nevertheless, parasites did not affect the choice of a nest site in any of the species studied. We suggest that the migratory flycatchers are in a hurry to start breeding upon arrival and use the presence of an old nest as a shortcut cue to assess nest-site quality. Also, they may save valuable time by copying the choice of previous breeders. Non-migratory tits may have more time to inspect nest sites, but do not seem to use the same cues in nest-site selection as the pied flycatcher.     

Does sex-ratio selection influence nest-site choice in a reptile with temperature-dependent sex determination?

Evolutionary theory predicts that dioecious species should produce a balanced primary sex ratio maintained by frequency-dependent selection. Organisms with environmental sex determination, however, are vulnerable to maladaptive sex ratios, because environmental conditions vary spatio-temporally. For reptiles with temperature-dependent sex determination, nest-site choice is a behavioural maternal effect that could respond to sex-ratio selection, as mothers could adjust offspring sex ratios by choosing nest sites that will have particular thermal properties. This theoretical prediction has generated decades of empirical research, yet convincing evidence that sex-ratio selection is influencing nesting behaviours remains absent. Here, we provide the first experimental evidence from nature that sex-ratio selection, rather than only viability selection, is probably an important component of nest-site choice in a reptile with temperature-dependent sex determination. We compare painted turtle (Chrysemys picta) neonates from maternally selected nest sites with those from randomly selected nest sites, observing no substantive difference in hatching success or survival, but finding a profound difference in offspring sex ratio in the direction expected based on historical records. Additionally, we leverage long-term data to reconstruct our sex ratio results had the experiment been repeated in multiple years. As predicted by theory, our results suggest that sex-ratio selection has shaped nesting behaviour in ways likely to enhance maternal fitness.     

Selection of nest-site habitat by interior least terns in relation to sandbar construction

Federally endangered interior least terns (Sternula antillarum) nest on bare or sparsely vegetated sandbars on midcontinent river systems. Loss of nesting habitat has been implicated as a cause of population declines, and managing these habitats is a major initiative in population recovery. One such initiative involves construction of mid-channel sandbars on the Missouri River, where natural sandbar habitat has declined in quantity and quality since the late 1990s. We evaluated nest-site habitat selection by least terns on constructed and natural sandbars by comparing vegetation, substrate, and debris variables at nest sites (n = 798) and random points (n = 1,113) in bare or sparsely vegetated habitats. Our logistic regression models revealed that a broader suite of habitat features was important in nest-site selection on constructed than on natural sandbars. Odds ratios for habitat variables indicated that avoidance of habitat features was the dominant nest-site selection process on both sandbar types, with nesting terns being attracted to nest-site habitat features (gravel and debris) and avoiding vegetation only on constructed sandbars, and avoiding silt and leaf litter on both sandbar types. Despite the seemingly uniform nature of these habitats, our results suggest that a complex suite of habitat features influences nest-site choice by least terns. However, nest-site selection in this social, colonially nesting species may be influenced by other factors, including spatial arrangement of bare sand habitat, proximity to other least terns, and prior habitat occupancy by piping plovers (Charadrius melodus). We found that nest-site selection was sensitive to subtle variation in habitat features, suggesting that rigor in maintaining habitat condition will be necessary in managing sandbars for the benefit of least terns. Further, management strategies that reduce habitat features that are avoided by least terns may be the most beneficial to nesting least terns. © 2011 The Wildlife Society.     

Deciding on a new home: how do honeybees agree?

A swarm of honeybees (Apis mellifera) is capable of selecting one nest-site when faced with a choice of several. We adapt classical mathematical models of disease, information and competing beliefs to such decision-making processes. We show that the collective decision may be arrived at without the necessity for any bee to make any comparison between sites.     

Morph-dependent resource acquisition and fitness in a polymorphic bird

Understanding genetic colour polymorphism has proved a major challenge, both in terms of the underlying genetic mechanisms and the evolutionarily forces maintaining such genetic variation. In this context, genetic differences in aggression or competitive-related traits may covary with the expression of alternative phenotypes, and affect the evolutionary stability and maintenance of colour polymorphisms. Genetic red and black head-colour morphs of the Gouldian finch (Erythrura gouldiae) co-occur in temporally and geographically stable frequencies in sympatric populations. Gouldian finches are obligate cavity-nesters with highly specific preferences for nest-site morphometry that directly affect reproductive success. Because intra- and interspecific competition for high quality nest-sites is prevalent, and fitness is directly related to nest-site quality, we investigated the relative access (and consequences for reproductive success) of alternative morphs to this critical limiting resource in the wild. Red males defended higher quality nest-sites, and overcame greater levels of nest-site competition against conspecifics and superior heterospecific competitors than black males. Red-headed males also produced more fledglings (especially with red-headed females) than black-headed males, independent of nest-site quality. Finally, the independent (positive) effect of nest-site quality on reproductive success was confirmed. Such competitive asymmetries are important to relative selection among coexisting morphs, and are likely to contribute to the maintenance of alternative sympatric colour-morphs in wild populations.     

Selection through predation, snowfall and microclimate on nest-site preferences in the Water Pipit Anthus spinoletta

Nest-site characteristics can have a strong impact on reproductive success in birds. Nest sites should simultaneously protect from predators, offer shelter and provide a favourable microclimate. We studied the relationship between three agents of natural selection (predators [i.e. Adders and birds/mammals], snowfall and microclimate), nest-site characteristics and reproductive success to determine whether these influenced preference for specific nest-site characteristics in the Water Pipit Anthus spinoletta . Pooled over all nests, the relative importance as agents of natural selection decreased from mammalian/avian predation (15% of all nests) through Adder predation (12%) to snowfall (7%), but there were clear differences in space and time. Predation by Adders selected for nest sites surrounded by few medium-sized shrubs. Selection by mammalian and avian predators favoured no specific nest-site characteristics. Protection from snowfall was best in nests surrounded by relatively few medium-sized shrubs. Microclimate had a strong influence on nestling survival and duration of nestling period. In nests on ENE-facing slopes, where maximum temperatures were reached in the morning, nestling survival was higher than on WSW-facing slopes, where temperature maxima occurred in the afternoon. Our results indicate that weak, but significant, directional selection is acting on preference for certain nest-site characteristics through effects on survival and development of nestlings. As predation and snowfall are unpredictable, the evolution of an optimal nest placement strategy is unlikely on a small scale. On a larger scale, however, choice of one breeding area over another may be favoured because of predictable differences between locations in terms of survival and nestling development.