A diagnosis model of parental care: How parents optimize their provisioning strategy in brood reduction?

Altricial birds often display biased preferences in providing parental care for their dependent offspring, especially during food shortages. During this process, such inflexible rules may result in provisioning errors. To demonstrate how parents optimize their provisioning strategies, we proposed a "diagnosis model" of parental care to posit that parents will undergo a diagnosis procedure to test whether selecting against some particular offspring based on phenotype is an optimal strategy. We tested this model in an asynchronous hatching bird, the Azure-winged Magpie Cyanopica cyanus, based on 10 years of data about demography and parental provisioning behaviors. Given their higher daily survival rates, core offspring (those hatched on the first day) merits an investment priority compared with their marginal brood mates (those hatched on later days). However, a marginal offspring also merited a priority if it displayed greater weight gain than the expected value at the early post-hatching days. Parents could detect such a marginal offspring via a diagnosis strategy, in which they provisioned the brood at the diagnosis stage by delivering food to every nestling that begged, then biased food toward high-value nestlings at the subsequent decision stage by making a negative response to the begging of low-value nestlings. In this provisioning strategy, the growth performance of a nestling became a more reliable indicator of its investment value than its hatching order or competitive ability. Our findings provide evidence for this "diagnosis model of parental care" wherein parents use a diagnosis method to optimize their provisioning strategy in brood reduction.

Turning tables: food availability shapes dynamic aggressive behaviour among asynchronously hatching siblings in red kites Milvus milvus.

Aggression represents the backbone of dominance acquisition in several animal societies, where the decision to interact is dictated by its relative cost. Among siblings, such costs are weighted in the light of inclusive fitness, but how this translates to aggression patterns in response to changing external and internal conditions remains unclear. Using a null-model-based approach, we investigate how day-to-day changes in food provisioning affect aggression networks and food allocation in growing red kite (Milvus milvus) nestlings, whose dominance rank is largely dictated by age. We show that older siblings, irrespective of age, change from targeting only close-aged peers (close-competitor pattern) when food provisioning is low, to uniformly attacking all other peers (downward heuristic pattern) as food conditions improve. While food allocation was generally skewed towards the older siblings, the youngest sibling in the nest increased its probability of accessing food as more was provisioned and as downward heuristic patterns became more prominent, suggesting that different aggression patterns allow for catch-up growth after periods of low food. Our results indicate that dynamic aggression patterns within the nest modulate environmental effects on juvenile development by influencing the process of dominance acquisition and potentially impacting the fledging body condition, with far-reaching fitness consequences.

Food Allocation under Asynchronous Hatching Conditions of Great Tits (Parus major).

The brood reduction hypothesis, which explains asynchronous hatching in birds, as an adaptation that enables selective survival of older nestlings when availability of food is unpredictable. This study was conducted in order to determine whether the brood reduction hypothesis can explain asynchronous hatching in passerines. Infrared cameras were installed inside nest boxes where great tits (Parus major) were attempting to reproduce in order to determine whether the parents practiced selective feeding of older nestlings. According to the results of the study, no significant difference was observed between the hatching order and the average number of feedings per nestling. In addition, when examining the distribution of food according to hatching order over time, every 30 min, beginning at 9 a.m., selective distribution of food to older nestlings was not observed. In conclusion, use of the brood reduction hypothesis, which supports selective provision of beneficial feeding of older and larger nestlings, to explain the asynchronous hatching of passerines is problematic, thus conduct of future studies focusing on other hypotheses in order to explain the asynchronous hatching of this passerine bird will be necessary.

Sibling Cannibalism in Group-Living Larvae of the Solitary Wasp, Isodontia harmandi (Hymenoptera: Sphecidae): Effects of Prey Availability and Size Discrepancy.

Sibling cannibalism has rarely been documented in the resource-based context of sibling rivalry. The solitary wasp Isodontia harmandi (Pérez) (Hymenoptera: Sphecidae) has an exceptional style of nursery 'communal brood cells', in which multiple larvae gregariously develop while relying on maternal prey provisioning. In this species there are several distinct features; e.g., cannibalism-driven brood reduction, single-sex broods, and mass provisioning. Rearing of the wasp larvae in artificial nest chambers provides the novel opportunity to study larval interactions under controlled conditions. To determine the factors that influence the occurrence of cannibalism and its timing, we conducted a rearing experiment using sib and non-sib pairs under two prey-availability treatments, and recorded the occurrence of cannibalism and the processes of larval growth and prey consumption over a 96-hr period. Cannibalism occurred more frequently in the low-prey treatment than in the high-prey treatment, and in female pairs than in male pairs. Cannibalism tended to occur earlier when the amount of prey remaining decreased rapidly. However, cannibalism did not occur only when all prey had been completely consumed, implying that it was not triggered directly by prey exhaustion. Cannibals were always larger than their victims, whose growth rates had often slowed before they were eaten. These results strongly imply that wasp larvae commit sibling cannibalism in the context of resource-based sibling rivalry, ensuring the double benefit of nutritional gains and reduced resource competition. We also investigated the possibility that wasp larvae monitor the size difference and/or body condition of brood-mates as an indicator of prey availability.

Niche construction through a Goldilocks principle maximizes fitness for a nest-sharing brood parasite.

Generalist brood parasites that share nests with host nestlings can optimize resource acquisition from host parents by balancing the benefits that host nest-mates provide, including attracting increased provisions to the nest, against the costs of competing with the same host young over foster parental resources. However, it is unclear how parasitic chicks cope when faced with more nest-mates than are optimal for their survival upon hatching. We suggest that, in the obligate brood parasitic brown-headed cowbird (Molothrus ater), chicks use a niche construction strategy and reduce larger, more competitive host broods to maximize the parasites' survival to fledging. We experimentally altered brood sizes to test for Goldilocks principle patterns (i.e. a 'just right' intermediate brood size) of cowbird survival in nests of prothonotary warbler (Protonotaria citrea) hosts. We found that intermediate brood sizes of two host nestlings maximized cowbird fledging success relative to 0 or 4 host nest-mates at hatching. Specifically, cowbird nestlings lowered host brood sizes towards this optimum when placed in broods with more host nestlings. The results suggest that cowbirds reduce, but do not eliminate, host broods as a niche construction mechanism to improve their own probability of survival.

Functional explanation of extreme hatching asynchrony: Male Manipulation Hypothesis.

Hatching asynchrony in birds is considered an adaptation to facilitate brood reduction because under conditions of food scarcity, the smallest nestling usually dies soon after hatching, thereby minimizing parental effort. However, in species with extreme hatching asynchrony, the last hatchlings paradoxically experience a very low probability of survival and death can take so long that it can hardly be considered an adaptation. Here, we propose and experimentally tested a new adaptive hypothesis explaining the brood reduction paradox, namely the "Male Manipulation Hypothesis". Our hypothesis suggests that by inducing asynchronous hatching, females increase the feeding requirements of the brood, which will induce males to increase provisioning effort. In addition, females may extend the period of male manipulation by feeding the smallest nestling just enough to sustain life. Our study showed that male common blackbirds ( Turdus merula) increased their effort (i.e., number of food items per hour) in experimental asynchronous broods compared to synchronous broods, while females reduced their contribution, as predicted by the hypothesis.

Avian sibling cannibalism: Hoopoe mothers regularly use their last hatched nestlings to feed older siblings.

Sibling cannibalism is relatively common in nature, but its evolution in birds and certain other vertebrates with extended parental care had been discarded. Here, however, we demonstrate its regular occurrence in two European populations of the Eurasian hoopoe (Upupa epops) and explore possible adaptive and non-adaptive explanations. Results showed that sibling cannibalism was more frequently detected in Spain (51.7%) than in Austria (5.9%). In these two populations, the hoopoes laid similar clutch sizes, resulting in similar fledging production, but hatching failures were more frequent in the northern population. Consequently, having more nestlings condemned to die in the southern population may explain the higher incidence of sibling cannibalism. In accordance with this interpretation, hatching span and failure, but not breeding date, explained the probability of sibling cannibalism in the Spanish hoopoes, while all three variables predicted brood reduction intensity. Furthermore, experimental food supply reduced the probability of sibling cannibalism, but not the intensity of brood reduction. Finally, females allocated fewer resources to the smallest nestlings when they were going to starve, but not necessarily when they were going to be used as food for their siblings. These results suggest that hoopoes produce extra eggs that, in the case of reduced hatching failure and food scarcity, produce nestlings that are used to feed older siblings. These findings provide the first evidence that sibling cannibalism occurs regularly in a bird species, thus expanding our evolutionary understanding of clutch size, hatching asynchrony, parent-offspring conflict, infanticide, and sibling cannibalism in the animal kingdom.

The Influence of Biological Factors on Haematological Values in Wild Marsh Harrier (Circus aeruginosus) Nestlings.

Marsh harrier (Circus aeruginosus) is a species with obligatory cainism, in which hatching asynchrony creates a pronounced size hierarchy in nestlings. The size-related competitive advantage of older nestlings means that they tend to dominate the younger ones, and brood reduction occurs in most nests. The aim of the study was to reference values and carry out a haematological examination in order to evaluate the physiological status and health of nestlings with respect to hatching order, brood size, and nest initiation date. To do so, we examined 19 nests with a total of 58 nestlings from a free-living population of this species located in fishpond complexes in the agricultural landscape of eastern Poland. Repeated blood samples (118 in all) were collected from nestlings. The following parameters were measured using fresh full blood: red blood cell count (RBC), haemoglobin level (Hb), white blood cell count (WBC), and plasma glucose level (Glu). The data were analysed using generalized linear mixed models and linear mixed models ((G)LMM). The study revealed that hatching order, but not brood size or nest initiation date, affected the physical condition of marsh harrier nestlings. Hb levels and RBC counts gradually decreased, whereas Glu levels and WBC counts increased from the first- to the last-hatched nestlings. This result points to the generally poorer condition of the youngest nestlings compared with their older siblings. The poor physiological condition of the youngest nestlings may consequently increase the likelihood of their perishing, and hence, of brood reduction.

Pipefish embryo oxygenation, survival, and development: egg size, male size, and temperature effects.

In animals with uniparental care, the quality of care provided by one sex can deeply impact the reproductive success of both sexes. Studying variation in parental care quality within a species and which factors may affect it can, therefore, shed important light on patterns of mate choice and other reproductive decisions observed in nature. Using Syngnathus typhle, a pipefish species with extensive uniparental male care, with embryos developing inside a brood pouch during a lengthy pregnancy, we assessed how egg size (which correlates positively with female size), male size, and water temperature affect brooding traits that relate to male care quality, all measured on day 18, approximately 1/3, of the brooding period. We found that larger males brooded eggs at lower densities, and their embryos were heavier than those of small males independent of initial egg size. However, large males had lower embryo survival relative to small males. We found no effect of egg size or of paternal size on within-pouch oxygen levels, but oxygen levels were significantly higher in the bottom than the middle section of the pouch. Males that brooded at higher temperatures had lower pouch oxygen levels presumably because of higher embryo developmental rates, as more developed embryos consume more oxygen. Together, our results suggest that small and large males follow distinct paternal strategies: large males positively affect embryo size whereas small males favor embryo survival. As females prefer large mates, offspring size at independence may be more important to female fitness than offspring survival during development.

Age-Specific Offspring Mortality Economically Tracks Food Abundance in a Piscivorous Seabird.

Earlier offspring mortality before independence saves resources for kin, which should be more beneficial when food is short. Using 24 years of data on age-specific common tern (Sterna hirundo) chick mortality, best described by the Gompertz function, and estimates of energy consumption per age of mortality, we investigated how energy wasted on nonfledged chicks depends on brood size, hatching order, and annual abundance of herring (Clupea harengus), the main food source. We found mortality directly after hatching (Gompertz baseline mortality) to be high and to increase with decreasing herring abundance. Mortality declined with age at a rate relatively insensitive to herring abundance. The sensitivity of baseline mortality to herring abundance reduced energy wasted on nonfledged chicks when herring was in short supply. Among chicks that did not fledge, last-hatched chicks were less costly than earlier-hatched chicks because of their earlier mortality. However, per hatchling produced, the least energy was wasted on chicks without siblings because their baseline mortality was most sensitive to herring abundance. We suggest that earlier mortality of offspring when food is short facilitates economic adjustment of posthatching parental investment to food abundance but that such economic brood reduction may be constrained by sibling competition.

Early mortality saves energy: estimating the energetic cost of excess offspring in a seabird.

Offspring are often produced in excess as insurance against stochastic events or unpredictable resources. This strategy may result in high early-life mortality, yet age-specific mortality before offspring independence and its associated costs have rarely been quantified. In this study, we modelled age-specific survival from hatching to fledging using 24 years of data on hatching order (HO), growth and age of mortality of more than 15 000 common tern (Sterna hirundo) chicks. We found that mortality peaked directly after hatching, after which it declined rapidly. Mortality hazard was best described with the Gompertz function, and was higher with later HO, mainly due to differences in baseline mortality hazard, rather than age-dependent mortality. Based on allometric mass-metabolism relationships and detailed growth curves of starving chicks, we estimated that the average metabolizable energy intake of non-fledged chicks was only 8.7% of the metabolizable energy intake of successful chicks during the nestling phase. Although 54% of hatchlings did not fledge, our estimates suggest them to have consumed only 9.3% of the total energy consumption of all hatched chicks in the population before fledging. We suggest that rapid mortality of excess offspring is part of an adaptive brood reduction strategy to the benefit of the parents.

Maternal effects as drivers of sibling competition in a parent-offspring conflict context? An experimental test.

Maternal effects occur when the mother's phenotype influences her offspring's phenotype. In birds, differential allocation in egg yolk components can allow mothers to compensate for the competitive disadvantage of junior chicks. We hypothesize that the parent-older chick conflict peaks at intermediate conditions: parents benefit from the younger chick(s) survival, but its death benefits the older chick in terms of growth and survival. We thus expect maternal compensation to follow a bell-shaped pattern in relation to environmental conditions. We studied a black-legged kittiwake (Rissa tridactyla) population where previous results revealed increased allocation of yolk testosterone in younger as compared to older chicks in intermediate conditions, in line with our theoretical framework. We therefore predicted a maternally induced increase in aggressiveness, growth, and survival for younger chicks born in intermediate environmental conditions. Controlling for parental effects and chick sex, we manipulated food availability before egg laying to create a situation with intermediate (Unfed group) and good (Fed group) environmental conditions. Within each feeding treatment, we further created experimental broods where the natural hatching order was reversed to maximize our chances to observe an effect of feeding treatment on the younger chicks' aggressiveness. As predicted, we found that chick aggressiveness was higher in younger chicks born from the Unfed group (i.e., in intermediate environmental conditions), but only when they were put in a senior position, in reversed broods. Predictions on growth and survival were not confirmed. Mothers thus seem to favor the competitiveness of their younger chick in intermediate conditions via egg yolk components, but our study also suggests that hatching asynchrony need to be small for maternal compensation to be efficient. We emphasize the need for further studies investigating other chick behaviors (e.g., begging) and focusing on the relative role of different yolk components in shaping parent-offspring conflict over sibling competition.

Variación en el tamaño de los huevos en garzas (Aves: Ardeidae) que anidan en la ciénaga de Birama, Cuba / Egg size variation in egrets and herons (Aves: Ardeidae) nesting in Birama’s swamp, Cuba

Intraclutch egg size variation in birds depends on many ecological factors and on the evolutive history of each species. In wading birds, a trend to smaller eggs with laying order has been described, but comparative reports are scarce. In this study, egg size variation patterns were described for nine Egrets and Heron species nesting in Birama’ Swamp, Cuba. The patterns were described using external dimensions of 3 142 eggs from 1 875 nests of Butorides virescens, Bubulcus ibis, Ardea alba, Nycticorax nycticorax, Nyctanassa violacea and four Egretta species, taken in the field between 1998 and 2006. Results showed that eggs were 4.9-10% of adult weight and had volume variation coefficients between 6-9%. There were no general and consistent interspecies relationship between clutch size and egg sizes. Average volumes tend to get smaller with laying order, but it is not statistically detectable in Butorides and Bubulcus. Last egg was between 0.2% and 15% smaller than the first, showing an inverse relationship with it. Intraclutch asymmetry is light in E. thula and fluctuating around null in Bubulcus. Size only predicted laying or hatching order for the last egg, in nests with more than two eggs, with 72.4% of confidence.

Las variaciones intranidada de la talla de los huevos de las aves responden a numerosos factores ecológicos y de la historia evolutiva de las especies. En las zancudas aunque se ha descrito una disminución de la talla con el orden de puesta, los trabajos comparativos entre especies son escasos. En este trabajo se describen los patrones de variación del tamaño de los huevos de nueve especies de garzas (Aves: Ardeidae) nidificantes en la ciénaga de Birama, Cuba. Los patrones se describen a partir de las dimensiones de 3 142 huevos, pertenecientes a 1 875 nidos de Butorides virescens, Bubulcus ibis, Ardea alba, Nycticorax nycticorax, Nyctanassa violacea y cuatro especies de Egretta, tomados en la ciénaga de Birama, Cuba, entre 1998 y 2006. Los huevos representaron entre el 4.9-10% del peso adulto y tuvieron un coeficiente de variación del volumen entre el 6-9%. No existió una relación general consistente entre especies entre el tamaño de puesta y las dimensiones de los huevos. El volumen promedio del huevo tiende a disminuir con el orden de puesta, pero no es estadísticamente detectable en Butorides y Bubulcus. El último huevo fue entre un 0.2 y un 15% menor que el primero, disminuyendo su volumen relativo con la talla del primer huevo. La asimetría intranidada es menos marcada en E. thula y prácticamente fluctuante alrededor de cero en Bubulcus. En general, la talla del huevo solo puede predecir el orden de puesta para el último huevo en nidadas mayores de dos con un 72.4% de confianza.

Comparing food limitation among three stages of nesting: supplementation experiments with the burrowing owl.

Food availability is an important limiting factor for avian reproduction. In altricial birds, food limitation is assumed to be more severe during the nestling stage than during laying or incubation, but this has yet to be adequately tested. Using food-supplementation experiments over a 5-year period, we determined the degree and timing of food limitation for burrowing owls (Athene cunicularia) breeding in Canada. Burrowing owls are an endangered species and food limitation during the nestling stage could influence reproductive performance of this species at the northern extent of their range. Supplemented pairs fledged on average 47% more owlets than unfed pairs, except during a year when natural food was not limiting (i.e., a prey irruption year). The difference in fledgling production resulted from high nestling mortality in unfed broods, with 96% of all nestling deaths being attributed to food shortage. Supplemental feeding during the nestling period also increased fledgling structural size. Pairs fed from the start of laying produced the same number of hatchlings as pairs that received no supplemental food before hatch. Furthermore, pairs supplemented from egg laying to fledging and pairs supplemented during the nestling period alone had the same patterns of nestling survival, equal numbers of fledglings, and similar fledgling mass and structural size. Our results provide empirical support for the hypothesis that the nestling period is the most food-limited phase of the breeding cycle. The experimental design we introduce here could be used with other altricial species to examine how the timing of food limitation differs among birds with a variety of life-history strategies. For burrowing owls, and other species with similar life histories, long-term, large-scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation. Our results provide empirical support for the hypothesis that the nestling period is the most food-limited phase of the breeding cycle. For burrowing owls, and other species with similar life histories, long-term, large-scale, and appropriately timed habitat management increasing prey abundance or availability is critical for conservation.

Brood size and its importance for nestling growth in the Biscutate Swift (Streptoprocne biscutata, Aves: Apodidae) / Tamanho da ninhada e sua importância no crescimento de filhotes de Streptoprocne biscutata (Aves: Apodidae)

Many Apodidae, including Streptoprocne biscutata (Sclater, 1866), drop eggs from their nests during incubation. This is interpreted as nest site competition or accident. We provide evidence that egg ejection is deliberate and that this behaviour controls the brood size. Brood sizes were manipulated and nestling growth was measured to test the hypothesis that pairs can regulate brood size during incubation based on current ability to rear nestlings. Natural (control) broods with one, two and three nestlings, and manipulated (experimental) broods reduced to one and increased to two and three young were monitored. Growth rates were measured based on weight, and wing, tail and tarsus lengths of natural and manipulated broods. We compared the slopes of each measure's regression lines of the nestlings of each brood size by t-test. Nestling growth of control nests was similar and relatively little associated with brood size. In broods reduced to one nestling, weight, wing and tail had greater growth rates, and in broods increased to three nestlings growth rates were lower. Weight was most, and tarsus length least influenced by brood size. In general, nestling growth of manipulated nests was inversely proportional to brood size. The results suggest that pairs with larger clutches are in better physical conditions than others. Thus, in experimental broods, pairs are over or under-loaded because feeding activities increase or decrease and these changes affect the growth rate of the nestlings. The present study suggests that egg ejection can control brood size. This behaviour is probably stimulated by physical changes in the adult birds during incubation.

Muitos Apodidae, incluindo Streptoprocne biscutata (Sclater, 1866), derrubam ovos do ninho durante a incubação. Este comportamento tem sido interpretado como competição ou acidente. Este estudo apresenta evidências de que a expulsão de ovos é deliberada e que esse comportamento controla o tamanho da ninhada. Ninhos com diferentes tamanhos de prole foram manipulados para testar a hipótese de que os casais podem regular o tamanho da ninhada durante a incubação com base em suas capacidades de alimentar os filhotes. Ninhadas naturais (controle) com um, dois e três filhotes e ninhadas manipuladas (experimentais) reduzidas para um e aumentadas para dois e três filhotes foram monitoradas. Foram medidas as taxas de crescimento de massa, asa, cauda e tarso de proles naturais e manipuladas. As inclinações das retas de regressão do desenvolvimento de cada medida ao longo da idade dos filhotes de cada tamanho de ninhada foram comparadas pelo t-teste. O crescimento dos filhotes de ninhos controle foi semelhante e esteve relativamente pouco associado ao tamanho da ninhada. Nas ninhadas reduzidas para um filhote, a massa, asa e cauda apresentaram as maiores taxas de crescimento e nas ninhadas aumentadas para três filhotes ocorreram as menores taxas. A massa foi a variável mais influenciada pelos experimentos e o tarso a menos. De forma geral, o crescimento dos filhotes de ninhos manipulados foi inversamente proporcional ao tamanho da ninhada. Os resultados sugerem que os casais com mais filhotes estão em melhores condições físicas do que os outros. Por isso, nas ninhadas experimentais, a taxa de crescimento dos filhotes muda em virtude dos casais se encontrarem sub ou sobrecarregados na tarefa de alimentação. Sugere-se que o comportamento de expulsão de ovos possa ser uma forma de regular o tamanho da ninhada, o qual provavelmente é desencadeado por mudanças nas condições físicas das aves adultas durante o período de incubação.

Hatching asynchrony and brood reduction in Tengmalm's owl Aegolius funereus: the role of temporal and spatial variation in food abundance.

Hatching asynchrony is the consequence of birds initiating incubation before clutch completion. It has been suggested that variation in hatching asynchrony in owls is extensive, and therefore they should be excellent objects to study the effects of spatio-temporal variation in food abundance on this phenomenon. We examined how abundance and predictability of food affected hatching asynchrony in Tengmalm's owl Aegolius funereus (Linnaeus), which mainly feeds on voles which fluctuate in 3- to 4-year cycles in northern Europe. Hatching span averaged 6-7 days (range 0-13 days) and increased with clutch size. Food supply did not directly influence levels of hatching asynchrony but it influenced indirectly via marked among-year changes in clutch size. During the decrease phase of the vole cycle the proportion of hatchlings producing fledglings decreased with asynchrony, suggesting that chick mortality was most common among asynchronous broods when food became scarce. This finding is consistent with Lack's brood reduction hypothesis, i.e. that if food becomes scarce during the nestling period the youngest nestlings would die first without endangering the survival of the whole brood.

Brood reduction facilitates female but not offspring survival in the great tit.

The long-term fitness consequences of brood reduction were examined in two (urban and rural) great tit populations in south-eastern Estonia during 1987-1994. The brood reduction hypothesis in its initial, Lackian sense was not supported since partial brood loss was accompanied by a decrease in fledgling weight and recruitment rate. Female survival was significantly improved in broods with high nestling mortality in the rural population. My results suggest that female great tits might be able to reallocate resources for self-maintenance if food appears to be short for the successful raising of the brood. However, parents are not capable of efficiently reallocating resources between nestlings.

Experiments on clutch size and nest size in passerine birds.

Results of experiments on three passerine species suggest that brood size may be constrained by nest size, since the breeding success of pairs provided with large nestcups was greater than that of those provided with small artificial nestcups. These results may have important implications, e.g. to the design of experiments involving manipulation of clutch and brood size. A small nestcup is requisite for successful hatching during the incubation period, but a large one for successful rearing during the nestling period. In nature this difference may select for types of nesting materials that are elastic, such as mosses and lichens. However, experiments showed that such materials rapidly absorb rainwater but only slowly dry out. In addition, because large nests dry out more slowly than small nests, selection will favour small nests among those open-nesting species that have exposed nests. A further possible nest size constraint on open-nesters is nest predation. However, no difference in the predation rate was found in experiments with small and large artificial nests.