Comparative analyses of the influence of developmental mode on phenotypic diversification rates in shorebirds

Phenotypic diversity is not evenly distributed across lineages. Here, we describe and apply a maximum-likelihood phylogenetic comparative method to test for different rates of phenotypic evolution between groups of the avian order Charadriiformes (shorebirds, gulls and alcids) to test the influence of a binary trait (offspring demand; semi-precocial or precocial) on rates of evolution of parental care, mating systems and secondary sexual traits. In semi-precocial species, chicks are reliant on the parents for feeding, but in precocial species the chicks feed themselves. Thus, where the parents are emancipated from feeding the young, we predict that there is an increased potential for brood desertion, and consequently for the divergence of mating systems. In addition, secondary sexual traits are predicted to evolve faster in groups with less demanding young. We found that precocial development not only allows rapid divergence of parental care and mating behaviours, but also promotes the rapid diversification of secondary sexual characters, most notably sexual size dimorphism (SSD) in body mass. Thus, less demanding offspring appear to facilitate rapid evolution of breeding systems and some sexually selected traits.     

The function of habitat change during brood-rearing in the precocial Kentish plover <Emphasis Type="Italic">Charadrius alexandrinus</Emphasis>

Food availability is often variable during the breeding season. Parents with nonmobile, altricial young have no choice but to accept changes in local food availability, whereas in precocial animals, the parents may lead their young away from poor sites to areas that have rich resources and/or are safe from predators. We investigated the latter hypotheses in the Kentish plover Charadrius alexandrinus, a precocial shorebird that raises its young in two habitats: on lakeshore and in saltmarsh. Parents move with their broods from saltmarsh to lakeshore, especially late in the breeding season, and we hypothesized that lakeshores provide more food than the saltmarsh. Consistent with our hypotheses, plover chicks grew faster on the shore, and the difference in growth rates between the two habitats was amplified later in the breeding season. In addition, brood survival was higher on lakeshore than in saltmarsh and decreased with hatching date. Taken together, our results suggest that Kentish plover parents increase their reproductive success by switching brood-rearing habitats strategically.     

Energetics of growth in semi-precocial shorebird chicks in a warm environment: the African black oystercatcher, Haematopus moquini

We studied prefledging growth, energy expenditure and time budgets of African Black Oystercatcher, Haematopus moquini, chicks on Robben Island, Western Cape, South Africa. The aim of the study was to investigate the effect of parental feeding on the growth and energetics of semi-precocial shorebird chicks. Chicks reached mean fledging mass, 463 g, in 40 days. The growth rate coefficient of African Black Oystercatcher chicks was 2% below the predicted value for a shorebird species of their body mass, but it was smaller than that of other precocial and semi-precocial shorebirds to date. Resting metabolic rate (RMR, measured through respirometry), daily metabolisable energy (DME), defined as daily energy expenditure (DEE, measured with doubly labelled water) plus energy deposited into tissue (E(tis)), and total metabolisable energy (TME) of African Black Oystercatcher chicks were similar to those expected for a species of their body size. DEE was not influenced by weather (ambient temperature, operative temperature and wind speed), therefore, variations in DEE may be explained by body mass alone. The relative RMR of the African Black Oystercatcher was greater, their TME was approximately the same, their average daily metabolisable energy (ADME) was less, and they spent less time foraging (short periods of parental feeding) and more time inactive than three precocial species in the Western Cape. Therefore, the semi-precocial mode of development of African Black Oystercatcher chicks reduced energy costs from thermoregulation and activity, and they were able to grow relatively faster than precocial, self-feeding shorebird species in similar climatic conditions. The growth rate coefficient of African Black Oystercatcher chicks was smaller than that of Eurasian Oystercatcher, Haematopus ostralegus, chicks, which may be a consequence of differences in body size and latitudinal effects.     

The energetic implications of precocial development for three shorebird species breeding in a warm environment

We investigated the effect of body size, parental behaviour and timing of breeding on the chick growth and energetics of three precocial shorebird (Charadrii) species in a warm sub-tropical environment: the Kittlitz's Plover Charadrius pecuarius , Blacksmith Lapwing Vanellus armatus , and Crowned Lapwing V. coronatus . The three species exhibited slower growth, longer pre-fledging periods and lower energy expenditure during the pre-fledging period than Arctic and temperate zone relatives of similar size. It was found that relative growth rates of charadriid and scolopacid species increased by 1.77% per degree latitude. Blacksmith and Crowned Lapwings are of similar size (i.e. about 160 g), about four times the mass of Kittlitz's Plovers. Taking body mass into account, the resting metabolic rate and daily energy expenditure of the three study species were similar. The relative amounts of energy needed for chicks to grow from a hatchling to a fledgling were low for the three species studied. The relative amount of energy needed to complete growth in charadriid and scolopacid species increased by 2.54% per degree latitude. Because Crowned Lapwings exhibited faster growth than Blacksmith Lapwings but similar daily energy expenditure we suggest that Blacksmith Lapwings compensate for greater energy expenditure in colder weather by growing more slowly. We suggest that small growth rate coefficient, low resting metabolic rate and low daily energy expenditure are adaptations to lower food availability and milder ambient temperatures in sub-tropical breeding habitats compared to temperate or Arctic regions.     

Oxygen consumption of brown noddy (Anous stolidus) embryos in a quasiequilibrium state at lowered ambient temperatures

1. The oxygen consumption (MO2) of the semi-precocial Brown Noddy embryos at different stages of development was measured at 36 degrees C and again after 5-hr exposure to lowered ambient temperatures (30 and 32 degrees C). 2. The MO2 measured in a quasiequilibrium state was equal to the value predicted by a temperature coefficient of 2. 3. In contrast to precocial chickens, the semi-precocial Noddy had no apparent metabolic response to cooling before hatching.     

PATTERNS OF GROWTH IN BIRDS

Parameters used to characterize the course of growth are described, and calculated growth parameters are presented for 105 species of birds of many taxonomic groups from a wide range of geographical localities. Growth parameters are found to exhibit as much as 20% variation within a species with respect to geographic locality and time of the nesting season. There is also considerable local variation, irrespective of season and locality, which is related to nutrition and perhaps to an inherited variability. The application of curve-fitting as a method of analysing intraspecific variation is discussed briefly, and the importance of comparative growth studies is emphasized. Growth patterns are correlated with other parameters of the life-history to evaluate the extent of diversity in the course of growth. Low rates of growth and prolonged growth periods occur primarily in species large for their families and in oceanic species. In most others, high rates of growth are maintained for longer periods of time. The shape of the growth curve is not related to the mode of development (i.e. whether precocial or altricial). Overall relative, or weight-specific growth rates, as measured by the constants of fitted growth equations, are most highly correlated with the adult body size of the species, changing as the -0–278 power of adult body weight. Smaller variations in the rate of growth appear to be correlated with differences in nesting success; open-nesting passerines grow faster than hole-nesting species of a similar size. Growth rate is further correlated with brood size. Oceanic species with single egg clutches and tropical land-birds with small clutches have low growth rates. The asymptote of the growth curve of the young (in relation to the adult weight) is related to the foraging behaviour of the adults. Aerial feeders generally have high asymptotes while those of ground feeding species are usually below adult weight. These differences are related to the need in the former for well-developed flight at the time of fledging. The diversity of growth patterns is related to evolutionary trends which are the result of (1) selective forces acting at stages of the life-history cycle other than development, (2) factors which affect the survival of offspring during the growth period, and (3) adjustments made to balance the energy budget of the family group. The last trend is discussed in detail in relation to the correlations found in the analysis. Two hypotheses are presented. Firstly, in species which cannot gather enough food to support even one young at a normal growth rate, the pace of development is reduced to decrease the energetic requirements of the young. Secondly, in species with small clutches, where adjustments to feeding capacities are not readily made by changing brood size, growth rate may be adjusted to accomplish this. The lack of critical energetic data to test these hypotheses is emphasized as a major deficiency in our understanding of the breeding biology of birds.     

Plasticity of Noddy Parents and Offspring to Sea-Surface Temperature Anomalies

Behavioral and/or developmental plasticity is crucial for resisting the impacts of environmental stressors. We investigated the plasticity of adult foraging behavior and chick development in an offshore foraging seabird, the black noddy (Anous minutus), during two breeding seasons. The first season had anomalously high sea-surface temperatures and ‘low’ prey availability, while the second was a season of below average sea-surface temperatures and ‘normal’ food availability. During the second season, supplementary feeding of chicks was used to manipulate offspring nutritional status in order to mimic conditions of high prey availability. When sea-surface temperatures were hotter than average, provisioning rates were significantly and negatively impacted at the day-to-day scale. Adults fed chicks during this low-food season smaller meals but at the same rate as chicks in the unfed treatment the following season. Supplementary feeding of chicks during the second season also resulted in delivery of smaller meals by adults, but did not influence feeding rate. Chick begging and parental responses to cessation of food supplementation suggested smaller meals fed to artificially supplemented chicks resulted from a decrease in chick demands associated with satiation, rather than adult behavioral responses to chick condition. During periods of low prey abundance, chicks maintained structural growth while sacrificing body condition and were unable to take advantage of periods of high prey abundance by increasing growth rates. These results suggest that this species expresses limited plasticity in provisioning behavior and offspring development. Consequently, responses to future changes in sea-surface temperature and other environmental variation may be limited.     

Siberian crane chick calls reflect their thermal state

Chicks can convey information about their needs with calls. But it is still unknown if there are any universal need indicators in chick vocalizations. Previous studies have shown that in some species vocal activity and/or temporal-frequency variables of calls are related to the chick state, whereas other studies did not confirm it. Here, we tested experimentally whether vocal activity and temporal-frequency variables of calls change with cooling. We studied 10 human-raised Siberian crane (Grus leucogeranus) chicks at 3–15 days of age. We found that the cooled chicks produced calls higher in fundamental frequency and power variables, longer in duration and at a higher calling rate than in the control chicks. However, we did not find significant changes in level of entropy and occurrence of non-linear phenomena in chick calls recorded during the experimental cooling. We suggest that the level of vocal activity is a universal indicator of need for warmth in precocial and semi-precocial birds (e.g. cranes), but not in altricial ones. We also assume that coding of needs via temporal-frequency variables of calls is typical in species whose adults could not confuse their chicks with other chicks. Siberian cranes stay on separate territories during their breeding season, so parents do not need to check individuality of their offspring in the home area. In this case, all call characteristics are available for other purposes and serve to communicate chicks’ vital needs.     

Female investment in offspring size and number shifts seasonally in a lizard with single-egg clutches

The timing of reproduction strongly influences reproductive success in many organisms. For species with extended reproductive seasons, the quality of the environment may change throughout the season in ways that impact offspring survival, and, accordingly, aspects of reproductive strategies may shift to maximize fitness. Life-history theory predicts that if offspring environments deteriorate through the season, females should shift from producing more, smaller offspring early in the season to fewer, higher quality offspring later in the season. We leverage multiple iterations of anole breeding colonies, which control for temperature, moisture, and food availability, to identify seasonal changes in reproduction. These breeding colonies varied only by the capture date of the adult animals from the field. We show that seasonal cohorts exhibit variation in key reproductive traits such as inter-clutch interval, egg size and hatchling size consistent with seasonal shifts in reproductive effort. Overall, reproductive effort was highest early in the season due to a relatively high rate of egg production. Later season cohorts produced fewer, but larger offspring. We infer that these results indicate a strategy for differential allocation of resources through the season. Females maximize offspring quantity when environments are favorable, and maximize offspring quality when environments are poor for those offspring. Our study also highlights that subtle differences in methodology (such as capture date of study animals) may influence the interpretation of results. Researchers interested in reproduction must be conscious of how their organism’s reproductive patterns may shift through the season when designing experiments or comparing results across studies.     

Breeding phenology,variation in reproductive effort and offspring size in a tropical population of the woodlouse Porcellionides pruinosus

Summary Most species of woodlice in temperate habitats have discrete breeding seasons. It is hypothesised that breeding synchronises with favourable environmental conditions to maximise offspring growth and survivorship (Willows 1984). We measured the breeding phenology of a species introduced to a tropical environment, primarily to consider the assumption that life histories in the tropics will differ fundamentally from those in temperate habitats. In addition to breeding phenology we considered variation in reproductive effort between individual females and the division of this effort between the size and number of young.A continuous breeding phenology was observed in a synanthropic population of Porcellionides pruinosus within the tropics. Reproductive effort varied between months, showed a weak relationship with female size and was independent of female fecundity. Female sizefecundity relationships varied between samples and when the proportion of reproductive females was high size-fecundity slopes were steeper than at other times. Mean offspring size varied between months and there was a wide range in offspring size within broods. Offspring size was not related to female body mass, reproductive effort or fecundity; consequently brood mass increased linearly with an increase in fecundity. Increased reproductive effort goes into more rather than larger offspring.We propose that the continuous breeding in this population was the result of the constant presence of an environmental cue to reproduction evolved in temperate habitats. Continuous breeding is not necessarily equivocal to high individual reproductive success even though overall population growth may be rapid. However, variation in reproductive effort suggests that individuals respond to current environmental conditions on short time scales.     

A comparison of life‐history and parental care in temperate and tropical wrens

Comparing closely related species that live in different environments is a powerful way to understand selective pressures that influence life‐history evolution. We examined a suite of life‐history traits and parental care in neotropical buff‐breasted wrens Cantorchilus leucotis and north‐temperate Carolina wrens Thryothorus ludovicianus (Family Troglodytidae), to test hypotheses about life‐history evolution. As expected, buff‐breasted wrens exhibited smaller clutch sizes and higher annual adult survival than Carolina wrens. We found minimal support for the nest predation hypothesis, as nest survival and age‐corrected provisioning rates to whole broods were similar between species, and number of breeding attempts and breeding season length were greater in temperate wrens. Critical predictions of the food limitation hypothesis were not supported; in particular age‐corrected provisioning rates per nestling were higher in the tropical than temperate species. The adult survival and offspring quality hypothesis garnered the most support, as buff‐breasted wrens exhibited greater age‐corrected provisioning rates per nestling, a longer nestling period, longer re‐nesting intervals following nest success, and lower annual fecundity than Carolina wrens. Despite similarly prolonged breeding seasons, reproductive strategies differ between species with buff‐breasted wrens investing considerably in single broods to optimize first‐year survival and Carolina wrens investing in multiple small broods to optimize annual fecundity.     

Geographical variation in reproductive investment across avian assemblages in Europe: effects of environmental drivers differ between altricial and precocial species

Reproductive traits provide information about the ways by which available resources are allocated during breeding. We tested for environmental drivers of large scale geographical patterns in assemblage mean clutch size, number of broods and overall reproductive investment per breeding season in European birds. We combined data about geographical distribution with published information about reproductive traits, and calculated mean trait values for avian assemblages occurring in 50 × 50 km grid cells. In total, we employed data from 499 species and 2059 assemblages. As the time available for breeding and the amount of food limit the reproductive effort, we related the geographical variation in reproductive traits to the length of breeding season, normalized difference vegetation index (NDVI) as a surrogate of resource availability, and its seasonality. Geographical patterns in traits may differ between reproductive modes, thus we performed the analyses separately for altricial Passerines (n = 203) and precocial non‐passerine species (n = 164) and controlled for the effect of taxonomy. Large clutches dominated in areas with high NDVI and, in precocial birds, also in areas with high annual seasonality and a long breeding season. High number of broods and high overall reproductive investment dominated in areas with a long breeding season, and high number of broods was found also in areas with low annual seasonality, but only in precocial species. High overall reproductive investment dominated in highly productive areas and also in areas with low annual seasonality in both groups. The increase in reproductive investment is caused mostly by an increase in the number of broods related to the length of season and partly by increase in clutch size related to NDVI. We found a negative correlation between clutch size and the number of broods in Passerines, which might suggest a trade‐off between these traits. Processes behind trait patterns differ between altricial and precocial species.     

Context‐specific influence of water temperature on brook trout growth rates in the field

1. Modelling the effects of climate change on freshwater fishes requires robust field‐based estimates accounting for interactions among multiple factors. 2. We used data from an 8‐year individual‐based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season‐specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third‐order stream in western Massachusetts, U.S.A. 3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context‐dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density. 4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow‐temperature interaction) differed among seasons. 5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures. 6. Our study provided robust, integrative field‐based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold‐water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season‐specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales.     

ADAPTATION, CONSTRAINT, AND COMPROMISE IN AVIAN POSTNATAL DEVELOPMENT

1. This paper discusses factors that influence the evolution of growth rate and determine its variation among species of birds. Growth rate is related to evolutionary fitness through the use of time, energy, and nutrients. In addition, balances between factors favouring rapid growth and those favouring slow growth may be investigated directly by experiment and by comparative observation. 2. David Lack (1968) proposed that the growth rate of the young is the optimum balance between selection for rapid growth to reduce the vulnerable period of development and selection for slow growth to reduce the energy requirements of the young. 3. To test Lack's hypothesis, the growth rates of birds, estimated by fitting sigmoid equations to curves relating weight to age, were surveyed widely from the literature. Among all species examined, growth rate was inversely related to adult weight. Among birds of similar size, most variation in growth rate was related to the degree of maturity of the neonate. Altricial chicks, which depend upon their parents for food and warmth, grow more rapidly than precocial chicks, which are self-sufficient shortly after hatching. Lack's hypothesis, which predicts a direct relationship between growth rate and mortality rate, was not supported. 4. I propose that the key to understanding variation in growth rate among birds lies in the balance between rate of cell proliferation or cell growth, on one hand, and acquisition of mature function, on the other. This idea is consistent with principles of cellular and developmental biology. It is supported by comparisons of (a) the neonates of different species, (b) the individual over the course of the developmental period, and (c) tissues whose use is acquired at different stages of development, wherein more mature individuals or tissues grow more slowly than those with less developed function. 5. Species of birds that are classified as semi-precocial develop precocially but grow rapidly. Although these seemingly violate the general rule relating growth rate to precocity, a closer inspection of their development reveals that they too support the rule. In the Common Tern, the legs, which are the key organ in precocial development, grow at the expected slow rate. The body as a whole grows rapidly because the growth increment of the legs is small and their growth is completed quickly. 6. Growth rates of precocial birds do not decrease abruptly at hatching. This points more to gradual tissue differentiation than to the pattern of procurement and allocation of energy as the primary control for growth rate. 7. Precocious development is favoured when the chicks are capable of self-feeding or when food supplies are distant from the next site and travelling time between one and the other is long. Precocity of the neonates frees both parents to feed at a distant food source. 8. Some species having diets with low levels of protein or other nutrients may grow slowly in order to match nutrient requirements to their availability in the diet. This pattern is indicated especially among the Procellariiformes, which feed an oily diet to their young, and also among tropical fruit-eating birds. 9. Some tropical, pelagically-feeding sea-birds that rear only one offspring at a time may not be able to procure food sufficient to support rapid chick growth. Alternative explanations for slow growth among these species include difficulty in obtaining essential nutrients and more precocious development of activity than in related species having more rapid growth.     

Nest temperature fluctuations in a cavity nester,the southern ground-hornbill

Southern ground-hornbills Bucorvus leadbeateri inhabit savanna and bushveld regions of South Africa. They nest in the austral summer, which coincides with the wet season and hottest daytime temperatures in the region. They are secondary cavity nesters and typically nest in large cavities in trees, cliffs and earth banks, but readily use artificial nest boxes. Southern ground-hornbills are listed as Endangered in South Africa, with reintroductions into suitable areas highlighted as a viable conservation intervention for the species. Nest microclimate, and the possible implications this may have for the breeding biology of southern ground-hornbills, have never been investigated. We used temperature dataloggers to record nest cavity temperature and ambient temperature for one artificial and 11 natural southern ground-hornbill tree cavity nests combined, spanning two breeding seasons. Mean hourly nest temperature, as well as mean minimum and mean maximum nest temperature, differed significantly between southern ground-hornbill nests in both breeding seasons. Mean nest temperature also differed significantly from mean ambient temperature for both seasons. Natural nest cavities provided a buffer against the ambient temperature fluctuations. The artificial nest provided little insulation against temperature extremes, being warmer and cooler than the maximum and minimum local ambient temperatures, respectively. Nest cavity temperature was not found to have an influence on the breeding success of the southern ground-hornbill groups investigated in this study. These results have potentially important implications for southern ground-hornbill conservation and artificial nest design, as they suggest that the birds can tolerate greater nest cavity temperature extremes than previously thought.     

Time constraints in temperate‐breeding species: influence of growing season length on reproductive strategies

Organisms that reproduce in temperate regions have limited time to produce offspring successfully, and this constraint is expected to be more pronounced in areas with short growing seasons. Information concerning how reproductive ecology of endotherms might be influenced by growing season length (GSL) is rare, and species that breed over a broad geographic range provide an opportunity to study the effects of time constraints on reproductive strategies. We analyzed data from a temperate‐breeding bird, the lesser scaup Aythya affinis; hereafter scaup, collected at eight sites across a broad gradient of GSL to evaluate three hypotheses related to reproductive compensation in response to varying time constraints. Clutch initiation date in scaup was unaffected by GSL and was unrelated to latitude; spring thaw dates had a marginal impact on timing of breeding. Clutch size declined during the nesting season, as is reported frequently in bird species, but was also unaffected by GSL. Scaup do not appear to compensate for shorter growing seasons by more rapidly reducing clutch size. This study demonstrates that this species is remarkably consistent in terms of timing of breeding and clutch size, regardless of growing season characteristics. Such inflexibility could make this species particularly sensitive to environmental changes that affect resource availabilities.     

Year class splitting in the woodlouse Philoscia muscorum explained through studies of growth and survivorship

Year class splitting has been observed in a population of the terrestrial isopod Philoscia muscorum (Scopoli) in a fixed dune grassland. In this population, individuals born in one season differentiate into two forms, one of which grows to maturity in one year, the other in two.
Laboratory investigations of growth, fecundity and survivorship of individuals showed fundamental positive relationships between rate of growth and the ambient temperature/day light regime and between parental size and the number of offspring produced. Size also affected survivorship, with adults surviving better than juveniles. A link between survivorship and the temperature/day light regime was only obvious in small juveniles.
No direct physiological cause for the year class split could be derived from the laboratory studies, but a trade-off between the need for delayed or early breeding could result in the patterns of growth and reproduction observed in the field population. Delayed breeding allows continued growth of adults and hence larger numbers of offspring. Early breeding allows more growth of juveniles which greatly improves their chances of winter survival.     

Comparative growth rates and oxygen consumption in young Galliformes

The high correlation between growth rate and adult body weight has been much more thoroughly documented for altricial birds than for precocial species. This paper gathers data from the literature for precocial Galliformes and also reports new growth data on six galliform species for analysis. The onset of homeothermic ability is investigated in Galliformes over a range of body size. The results confirm that (1) large species' chicks grow at a slower rate than those of smaller species, and (2) larger species' chicks can thermoregulate earlier than smaller species' chicks under cold stress situations. Published embryonic body weights are also analysed to determine when growth rate differences appear in the development of precocial species. No interspecific differences appeared in the relative growth rates of embryos, and therefore species body size does not appear to influence growth rate before hatching.     

Communal Nesting in Asocial Abert's Squirrels: the Role of Social Thermoregulation and Breeding Strategy

The social thermoregulation hypothesis states that endothermic species will communally nest to reduce energy expenditures on thermoregulation. The hypothesis predicts that the frequency of communal nesting should increase with decreasing ambient temperature. The potential costs of communal nesting (e.g., increased predation risk, resource competition, cuckoldry, parasite/disease transmission, or infanticide) could decrease the frequency of communal nesting especially for asocial breeding females with dependent offspring. We examined the effects of ambient temperature and seasonal reproductive activities on the probability of communal nesting in Abert's squirrels (Sciurus aberti) in the Pinaleño Mountains, Arizona. Most squirrels nested consistently with the same partner in mixed‐sex pairs. The proportion of individuals engaging in communal nesting increased with decreasing ambient temperature as predicted by the social thermoregulation hypothesis. The onset of the breeding season greatly reduced the proportion of individuals communally nesting. The negative relationship between ambient temperature and communal nesting supports the use of communal nesting in Abert's squirrels as a mechanism to reduce thermoregulatory costs during cold conditions. The abrupt drop in the frequency of communal nesting during the breeding season is likely due to female abandonment of this behavior. By avoiding communally nesting during the breeding season, females may prevent males from mating with them outside of mating chases, reduce resource competition, and protect offspring from infanticide, diseases, and parasites. Males may gain additional fitness benefits from nesting with females because familiarity with females increases dominance rank in mating activities.     

Time‐limited environments affect the evolution of egg–body size allometry

Initial offspring size is a fundamental component of absolute growth rate, where large offspring will reach a given adult body size faster than smaller offspring. Yet, our knowledge regarding the coevolution between offspring and adult size is limited. In time‐constrained environments, organisms need to reproduce at a high rate and reach a reproductive size quickly. To rapidly attain a large adult body size, we hypothesize that, in seasonal habitats, large species are bound to having a large initial size, and consequently, the evolution of egg size will be tightly matched to that of body size, compared to less time‐limited systems. We tested this hypothesis in killifishes, and found a significantly steeper allometric relationship between egg and body sizes in annual, compared to nonannual species. We also found higher rates of evolution of egg and body size in annual compared to nonannual species. Our results suggest that time‐constrained environments impose strong selection on rapidly reaching a species‐specific body size, and reproduce at a high rate, which in turn imposes constraints on the evolution of egg sizes. In combination, these distinct selection pressures result in different relationships between egg and body size among species in time‐constrained versus permanent habitats.