Effects of incubation temperature on behavior of hatchling pine snakes: implications for reptilian distribution

Summary Incubation temperatures in vertebrates affect incubation periods, and in some reptiles incubation temperature determines sex ratios and some limited behavior. Here I present evidence that incubation temperature in pine snakes (Pituophis melanoleucus) affects not only incubation periods and posthatching behavior in the laboratory, but also the behavior of hatchlings required for successful emergence and survival. These behavioral differences have evolutionary implications for selection of hatchlings from particular temperature nests. With increasing temperature, incubation periods decreased in the laboratory. In addition, incubation temperature affected hatching and emergence times as well as movement speed and foraging ability. Hatchlings from medium temperature conditions emerged from nests in the field in less time than hatchlings incubated at high or low temperatures, and hatchlings from low temperatures moved slower and were less able to capture and eat mice in the laboratory than hatchlings incubated at medium or high temperatures. Taken together, these laboratory and field experiments suggest that hatchlings from low temperature nests, compared to those from higher temperature nests, would be less able to emerge, find food, and locate hibernation sites prior to the onset of cold temperatures in the New Jersey Pine Barrens. These results suggest that incubation temperature affects a whole range of behavior and that distributional ranges of reptiles and other poikilotherms could be affected by summer temperatures (via incubation period and subsequent behavior) as well as ambient winter temperatures.     

Locomotion performance of green turtle hatchlings from the Heron Island Rookery,Great Barrier Reef

Locomotion performance plays a vital role in determining hatchling green turtle Chelonia mydas survival in the first few hours after emerging from their nests as hatchlings crawl and swim the gauntlet of predators before reaching the relative safety of the open ocean. Previous laboratory based constant incubation experiments found incubation temperature to influence the size and swimming performance of hatchling green turtles. Here we examine the morphology and crawling and swimming performance of hatchling green turtles as they emerge from nests on Heron Island rookery in the southern Great Barrier Reef to test the hypothesis that nest temperature in the field can influence these attributes. We found inter-nest differences in hatchling mass and dimensions, and that hatchling mass was not correlated with nest temperature. However, hatchlings from warmer nests had smaller carapace dimensions than hatchlings from cool nests suggesting that more yolk was converted to hatchling tissue during embryonic development in cool nests. There was considerable intra- and inter-nest variation in both crawling and swimming performance of hatchlings. Hatchlings from cool nests tended to be faster crawlers than hatchlings from warm nests, but the thrust produced during swimming was not correlated with nest temperature. During the 4 h swimming trial, hatchlings swimming effort decreased significantly during the first 3 h but swimming effort remained relatively constant for the last 1 h. Individual hatchling crawling and swimming performances were not correlated with each other. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of incubation temperature on hatchling quality in the olive ridley turtle, Lepidochelys olivacea, from Alas Purwo National Park, East Java, Indonesia: implications for hatchery management

Nest protection through egg relocation from natural nests into protected hatcheries is a common practice used at rookeries around the world to increase hatchling recruitment into sea turtle populations. However, rarely have the impacts of this practice on hatchling recruitment and quality been assessed. This study investigated the influences of the thermal nest environment of olive ridley turtles Lepidochelys olivacea on emergence success and quality of hatchlings of hatchery nests in Alas Purwo National Park, East Java, Indonesia (2009 and 2010 nesting seasons). Nest temperatures above 34?°C for at least 3 consecutive days during incubation in the hatchery resulted in decreases in emergence success and locomotor performance of hatchlings. The use of the hatchery is recommended due to extremely high predation rate of nests left on the beach; however, altering hatchery management practice by spacing nests one meter apart and providing shade should improve hatchery outcomes now and into the future.     

Incubation temperature has long-term effects on behaviour of young Pine snakes (Pituophis melanoleucus)

Summary Eggs of pine snakes (Pituophis melanoleucus) were incubated at constant temperatures of 21°, 23°, 26°, 28°, 30° and 32° C to determine behavioural differences as a function of incubation temperature. For all behavioural and physiological measures hatchlings from medium incubation temperatures (26°, 28°) performed tests better than those hatchlings from eggs incubated at low temperatures (21°, 23°). For some behavioural tests hatchlings from eggs incubated at high temperature (30°, 32°) performed less well than medium temperature hatchlings. These differences were not due to ambient temperatures or age of testing since these were held constant. Some of the behavioural differences persisted for 24 weeks.     

Influence of emergence success on the annual reproductive output of leatherback turtles

Reproductive output of leatherback turtles (Dermochelys coriacea) is affected by the stochastic nature of emergence success. Average emergence success of nests at Playa Grande, Costa Rica was 0.38 ± 0.27. Incubation temperature affected development of leatherback turtle eggs and emergence of hatchlings from the nest. We found that high temperatures reduced hatching success and emergence rate and increased embryonic mortality both early and late during incubation at Playa Grande. There was a temporal effect on emergence success that resulted in more hatchlings being produced at the beginning of the season, because of higher emergence success, than toward the end. Likewise, production of hatchlings varied from year to year. The average annual reproductive output was 252 ± 141 hatchlings per female. The 2005–2006 nesting season had the highest emergence success and produced the greatest number of hatchlings per female compared to the 2004–2005 (+120%) and 2006–2007 (+41%) seasons. However, average clutch size (62 ± 10) and clutch frequency (9.45 ± 1.63), were not different among years. Turtles that had nested a high number of years exhibited greater clutch frequency and arrived earlier to nest than turtles that had nested in fewer numbers of years. Nesting when environmental conditions favor high developmental success and emergence rate may constitute an advantageous reproductive strategy.     

Patterns in the emergence of green (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) and loggerhead (<Emphasis Type="Italic">Caretta caretta</Emphasis>) turtle hatchlings from their nests

The emergence patterns of both green (Chelonia mydas) and loggerhead (Caretta caretta) turtle hatchlings were observed in great detail over three seasons at Alagadi beach, northern Cyprus. In total, 38 green turtle and 50 loggerhead turtle nests were monitored, accounting for the emergence of 2,807 and 2,259 hatchlings, respectively. We quantified these emergences into 397 green turtle and 302 loggerhead turtle emergence groups. Overall, 85.0% of green turtle and 79.5% of loggerhead turtle groups emerged at night; these accounted for 85.5 and 90.8% of hatchlings, respectively. The remaining emergences were dispersed throughout the day for green turtle nests but confined to the morning in loggerhead turtle nests. Hatchling emergence from individual nests occurred over periods of between 1 and 7 nights, with most hatchlings typically emerging on the first night. Group sizes of green turtles emerging during the day were significantly smaller than those emerging at night. Hatchlings of both species that emerged from nests during the day had longer emergence durations than those that emerged from nests at night only.Communicated by R.J. Thompson, St. Johns     

Intraspecific nest parasitism in the swallow Hirundo rustica: the importance of neighbors

Summary Intraspecific nest parasitism in the swallow Hirundo rustica involves several parasite and anti-parasite tactics. Neighboring swallows breed asynchronously, perhaps because neighboring nests with overlapping egg-laying periods have higher frequency of intraspecific nest parasitims than neighboring nests with no such overlap. Since nest guarding prevents nest parasitism, the effect of nest guarding was removed by putting up old swallow nests. Such experimental nests were parasitized by swallows more often when the nests contained eggs, than when empty, and when the nests were far away from, rather than close to, an active neighboring nest. Close experimental nests with eggs were, however, parasitized less frequently if asynchronous (non-overlapping egg-laying periods), rather than synchronous, to active neighboring nests. Aggressive anti-parasite behavior of nest owners was determined from responses towards female intruders. Responses were more often aggressive before and especially during egg laying than during incubation. Nest owners also behaved aggressively more often when the intruder approached close to as compared to at some distance from their nest. Focal pairs behaved aggressively more often towards closely approaching intruders during the incubation period than towards distantly approaching intruders during the egg-laying period. Swallows reduce the frequency of intraspecific nest parasitism by nesting asynchronously with and close to a neighboring nest. Aggression by neighbors may reduce the success of potential parasitic swallows.     

Nest concealment and parental behaviour interact in affecting nest survival in the blackcap (Sylvia atricapilla): an experimental evaluation of the parental compensation hypothesis

Nest concealment varies strongly within populations of many species. Although some studies have revealed the beneficial effects of concealment in mitigating predation pressure on nests, other studies were unable to find similar effects. One potential reason for the mixed results is that parental behaviour may compensate for the effects of nest cover, and specifically designed experimental studies are needed to reveal this compensation. I studied the effects of concealment on the probability of nest survival in the blackcap (Sylvia atricapilla), by experimentally manipulating the degree of nest-foliage cover. There was a significant effect of the treatment depending on nest type and the phase of nesting. Whereas there was no effect of concealment on nest survival in natural nests, there was a positive effect in real nests baited with plasticine clutches (i.e. without parental activity). Parents probably behaviourally compensated for poor concealment in natural nests (nest guarding, defence). In line with this, there was no effect of concealment on nest survival during incubation, whereas there was probably a positive effect in the nestling phase. Parents spent more time on the nest during incubation (80%) than during the care of nestlings (40%) and, consequently, had more opportunities to compensate for poor cover. In general, we cannot use single measures of behaviours or states (nest concealment) as an indication of predation risk because of the capacity for compensation in other behaviours.Communicated by C. Brown     

Experimental analysis of worker division of labor in bumblebee nest thermoregulation (<Emphasis Type="Italic">Bombus huntii</Emphasis>, Hymenoptera: Apidae)

Bumblebee colonies experience daily and seasonal fluctuations in ambient temperature, but proper brood development requires a stable nest temperature. This study examined how adaptive colony responses to changing ambient temperature are achieved through the in-nest workers’ behavioral plasticity. We studied three Bombus huntii colonies in the laboratory. In the first experiment, we manipulated ambient temperature and recorded brood cell incubation and wing fanning by individually marked, known-age bees. The colonies maintained their nests closer to appropriate brood development temperatures (28 to 32°C) when exposed to a range of ambient temperatures from 10.3 to 38.6°C. Incubation activity was greater in cooler treatment conditions, whereas in the highest temperature treatment, some bees fanned and others moved off the brood. As the ambient temperature dropped, workers increased the duration of their incubating bouts, but, except at the highest temperature, the number of workers that incubated did not differ significantly among treatments. A subset of the bees incubated significantly more than their nest mates, some of which never incubated. Worker body size, but not age, was a good predictor of incubation rates, and smaller bees incubated at higher rates. In the second experiment, we removed the most actively incubating workers. Immediately after removals, the total colony incubation effort was lower than pre-removal levels, but incubation effort rebounded toward pre-removal levels after 24 h. The increased thermoregulatory demand after removals was met primarily by bees increasing their rates of incubation rather than by bees switching from a different task to incubation. We conclude that some B. huntii workers specialize on nest thermoregulation, and that changes in work rates are more important than task switching in meeting thermal challenges.     

Seasonal fluctuations in sand temperature: effects on the incubation period and mortality of loggerhead sea turtle (<Emphasis Type="Italic">Caretta</Emphasis><Emphasis Type="Italic">caretta</Emphasis>) pre-emergent hatchlings in Minabe,Japan

Incubation period, hatching success, and emergence percentage in loggerhead (Caretta caretta) nests were quantified during the 1993 and 1995 nesting seasons and following incubation seasons in Minabe, Wakayama, Japan. Sand and nest temperatures were also monitored. Over the seasons, daily mean sand temperature at nest depth fluctuated between 18.0°C and 33.3°C, with a steep increase in the second week of July and a peak in late August. Temperatures inside the nest chambers were a few degrees above those of the surrounding sand at the end of incubation. The incubation period ranged from 46 to 82 days. A significant negative correlation was found between mean sand temperature and incubation period. The relationship conformed to the day-degree concept. There was no significant seasonal trend in hatching success, but many pre-emergent hatchlings were found dead in most of the clutches during the warmest part of the season. Emergence percentage was correlated with mean sand temperature calculated for 4 days before emergence, suggesting that mortality may be due to heat. This heat-related mortality is considered to be a common phenomenon at our study site, because the peak in emergences coincides with the peak in high temperatures. These temperature effects on hatchling mortality must be taken into account in estimates of hatchling sex ratios. Because sand temperatures already exceed the optimal thermal range for incubation, this population is vulnerable to even small temperature increases resulting from global warming.     

Emergence pattern of loggerhead turtle (<Emphasis Type="Italic">Caretta caretta</Emphasis>) hatchlings from Kyparissia Bay,Greece

The present study describes the emergence pattern of loggerhead sea turtle hatchlings (Caretta caretta) from a nesting beach in Kyparissia Bay (Greece). We try to establish the role played by hatchling biometry, nest relocation and distance from nest to the sea on this emergence pattern. We surveyed a total of 32 nests, and found long emergence periods (mean = 6.7 nights). The majority of emergences occurred at night, mainly between 0030 and 0100 hours, and in small groups. Most of the hatchlings emerged from the nests the first night. We found no clear trend when we studied the effect of hatchling biometry between successive emergence days. We also found that relocation of the nests did not significantly affect the emergence pattern. However, we noted that in the relocated nests, hatchlings emerged in smaller groups. Emergence periods were inversely related to distance from the sea. In short, factors such as climate conditions, relocation and nest distance to the sea appear to have some effect on the emergence pattern. Therefore, they should be taken into account in both biological studies and management plans for sea turtle nesting beaches. Our results suggest leaving an extended period between the first emergence of hatchlings and the excavation of nests by researchers in future studies in the area.     

Why do most tropical animals reproduce seasonally? Testing hypotheses on an Australian snake

Most species reproduce seasonally, even in the tropics where activity occurs year-round. Squamate reptiles provide ideal model organisms to clarify the ultimate (adaptive) reasons for the restriction of reproduction to specific times of year. Females of almost all temperate-zone reptile species produce their eggs or offspring in the warmest time of the year, thereby synchronizing embryogenesis with high ambient temperatures. However, although tropical reptiles are freed from this thermal constraint, most do not reproduce year-round. Seasonal reproduction in tropical reptiles might be driven by biotic factors (e.g., peak periods of predation on eggs or hatchlings, or food for hatchlings) or abiotic factors (e.g., seasonal availability of suitably moist incubation conditions). Keelback snakes (Tropidonophis mairii, Colubridae) in tropical Australia reproduce from April to November, but with a major peak in May-June. Our field studies falsify hypotheses that invoke biotic factors as explanations for this pattern: the timing of nesting does not minimize predation on eggs, nor maximize food availability or survival rates for hatchlings. Instead, our data implicate abiotic factors: female keelbacks nest most intensely soon after the cessation of monsoonal rains when soils are moist enough to sustain optimal embryogenesis (wetter nests produce larger hatchlings, that are more likely to survive) but are unlikely to become waterlogged (which is lethal to eggs). Thus, abiotic factors may favor seasonal reproduction in tropical as well as temperate-zone animals.     

How Kentish plovers,<Emphasis Type="Italic"> Charadrius alexandrinus</Emphasis>, cope with heat stress during incubation

Biparental incubation is frequent among shorebirds and is expected when the survival prospects of offspring increase relative to uniparental incubation. To understand why this occurs, it is important to identify the factors that constrain uniparental incubation. It is assumed that birds choose nesting sites that provide an appropriate microclimate for incubation. Many shorebirds nest in sites with no or little cover, where ambient temperatures at ground level might be >50°C during very hot days. Shorebirds nest in exposed sites because predation risk on incubating adults is higher in covered sites. In hot environments, incubating shorebirds might experience heat stress in exposed sites, and this may compromise nesting success if adults are unable to attend their nests continuously, limiting the possibilities of uniparental incubation and thus the expression of a sexual conflict over incubation. The operative temperatures of Kentish plovers (Charadrius alexandrinus) were recorded in exposed and covered sites, and the thermal behaviour and incubating tactics of pair members were studied in a hot environment. During the hottest part of the day, there was a difference of 10–15°C in the operative temperatures of plovers between covered and exposed sites. Plovers in covered sites did not exhibit any thermoregulatory behaviour indicative of thermal stress, probably because the thermal range encountered by them in such places during most of the daytime was close to the thermo-neutral zone. The frequency with which plovers in exposed sites exhibited thermoregulatory behaviour was related to ambient temperature. Under very hot conditions, incubating birds were probably unable to maintain homeostasis for long periods and pair members resorted to shortening incubation bouts. Female Kentish plovers mainly incubate in the daytime and males during the night. However, the probability of diurnal incubation by males increased with ambient temperature in exposed nests, but not in covered ones. In fact, the frequency of participation in diurnal incubation by males was greater in exposed than in covered sites, suggesting that the participation of males in diurnal incubation may be related to the inability of females to stay at the nest during long periods when the ambient temperature is high. Even after resorting to shortened incubation bouts, the plovers may be unable to attend their nests continuously during heat waves, and the nests may be deserted. The propensity of plovers to desert their nests was affected by proximity to water, with nests located close to water being deserted less frequently. It seems likely that susceptibility to thermal stress changed in relation to proximity to water because in sites close to water it was possible to belly-soak, which would allow a more continuous nest attendance. Therefore, despite the adoption of behavioural solutions to face heavy heat loads, nesting success was vulnerable to these solutions because heat stress during extended periods may constrain parental nest attendance, and this may limit the opportunities for sexual conflicts over incubation.Communicated by J. Graves     

Variation in nesting patterns affecting nest temperatures in two populations of painted turtles (<Emphasis Type="Italic">Chrysemys picta</Emphasis>) with temperature-dependent sex determination

Mechanisms maintaining sex ratios in populations with temperature-dependent sex determination (TSD) remain elusive. Although geographic variation in embryonic sex determination (i.e., pivotal temperature) has been widely investigated in reptiles exhibiting TSD, no previous studies have directly addressed geographic variation in maternal behavior affecting nest thermal conditions. I evaluated patterns of nest-site selection and its effects on thermal and hydric nest conditions for a population of painted turtles ( Chrysemys picta bellii) exhibiting TSD in New Mexico. These results are compared to data collected from a well-studied, conspecific population experiencing relatively cooler climatic conditions in Illinois. Since canopy vegetation cover reduces nest temperatures in Illinois, I expected females in New Mexico to nest under high amounts of canopy vegetation cover. However, females from New Mexico placed nests under significantly less canopy vegetation cover, but closer to standing water, than did females from Illinois. Experimental nests in New Mexico demonstrated that increased canopy vegetation cover and soil moisture reduced nest temperatures. By nesting close to standing water rather than under canopy vegetation cover, females in New Mexico nested in habitats more closely associated with maximizing moisture around nests rather than reducing nest temperatures through shading. Mean July nest temperatures were similar for both populations. Since nest hydric conditions affect hatching success and hatchling size in C. picta, nesting patterns in New Mexico may primarily reflect selection for microhabitats affecting offspring survivorship or size.     

Reproductive defence priorities of male willow ptarmigan (Lagopus lagopus): enhancing mate survival or extending paternity options?

Summary Cock willow ptarmigan (Lagopus lagopus) closely guard their mates from predators and conspecific males, and vigorously defend their nests and young. In view of potential costs and benefits of behavioural guarding descisions, I designed a test to examine if, when and how males altered defence priorities. Cock willow ptarmigan were very attentive to their mates throughout the breeding season, unaccompanied hens comprising less than 3% of the sightings of mated females. From the second week of incubation until shortly after hatch, males attempted to guard both mates and offspring. Males guarded mates preferentially over nests until the third week of incubation; they then changed their priority to defence of offspring (nest and/or chicks). The overall defence response initially was strong, but decreased after the first week of incubation. Strength of response increased again in late incubation, and declined as chicks began to fly. Consistency of defence decisions was highest during prenesting and egg laying and lowest during late incubation. Since males defended their mates for a longer period than required for protection of paternity, hypotheses predicting enhanced mate survival and extended paternity options through renesting were examined. Removal of males did not result in reduced survival or increased weight loss of widows during incubation. In light of heavy nest depredation and displacement pressures by conspecifics, mate guarding throughout incubation enhanced male fitness by ensuring paternity in renesting attempts.     

Nesting characteristics of the olive ridley turtle (<Emphasis Type="Italic">Lepidochelys olivacea</Emphasis>) in Cabo Pulmo,southern Baja California

Baja California represents the northernmost reproductive area for Lepidochelys olivacea. This region is characterized by its high temperature and low humidity. Thus, nest site selection plays an important role in reproductive success. To evaluate the factors affecting site selection for oviposition Las Barracas Beach, located in the south of the peninsula, was visited, from August to November 2000, in weekly periods. A density of 2.55 nests ha^–1 was estimated from a total of 55 nests counted. A hatching success of 73.7% was observed, with 23.7% mortality and 2.5% of eggs without apparent development (eggs counted=4,031). On the day of oviposition, the relative surface humidity (between 0.75% and 1.75%) and temperature (32°C) preferred by the females for nest construction was determined at the nest site, as well as the mean distance between the nests and the high tide line for that day (17.65 m). Hatching success was affected by humidity and distance. In addition, hatching success was higher between 10 and 30 m above the high tide line measured on the day of oviposition (surface humidity ca. 1%). A total of 874 hatchlings were measured, with an average of 42.1 mm straight carapace length. Larger sizes were found in October and smaller ones in November. Generally speaking, the sizes found here were larger than those obtained in other areas. This is probably due to two factors: (1) the nests in Las Barracas were kept in situ, whereas in other areas nests are sometimes incubated artificially and (2) the nesting population of southern Baja California is different from the eastern Pacific population.Communicated by P.W. Sammarco, Chauvin     

Behaviorally mediated spatial and temporal refuges from a cleptoparasite,Argochrysis armilla (Hymenoptera: Chrysididae), attacking a ground-nesting wasp,Ammophila dysmica (Hymenoptera: Sphecidae)

Summary It has been suggested that parasite pressure favors the evolution of sociality within the Hymenoptera. I analyzed the impact of a chrysidid nest parasite, Argochrysis armilla, on its host, the solitary ground-nesting wasp, Ammophila dysmica, to assess the role of parasitism in favoring two steps towards sociality: aggregated nesting and delayed nest provisioning. The foraging strategy of Argochrysis armilla involves discovering host nests during excavation, learning the locations of discovered nest, and returning to oviposit in nests during nest provisioning; I therefore assessed the influence of host behavior on (1) parasite discovery of nests and (2) parasite oviposition in nests. Significantly fewer parasites discovered host nests that were excavated during the early morning and late afternoon, due to partial asynchrony of host and parasite activity. Nests excavated in areas of low nest density were also less likely to be discovered; use of low density nest sites increased during periods of high parasite activity. Due to a rapidly decelerating rate of parasite recruitment to nests under excavation, the duration of nest digging had only a limited influence on nest discovery by parasites. The probability of parasite oviposition in a host nest was determined by the number of parasites discovering the nest during excavation and by the time between nest excavation and provisioning; delaying nest provisioning reduced the risk of parasite oviposition. Delayed provisionings primarily appeared to be a result of the stochastic process of hunting and prey encounter. The number of provisions placed in a nest (one vs two) had no effect on the probability of nest parasitism. Spatial patterns of parasitism were directly density dependent in 1984 and density independent in 1986. In this system parasite pressure acts against the formation of nesting aggregations and in favor of delayed nest provisioning. The dependence of these results on species-specific aspects of the parasite's foraging strategy and the host's defensive strategy suggests, however, that different parasite species may generate qualitatively different selection pressures, potentially contributing to the diversity of nesting behavior in the Hymenoptera.     

Climate and predation dominate juvenile and adult recruitment in a turtle with temperature-dependent sex determination

Conditions experienced early in life can influence phenotypes in ecologically important ways, as exemplified by organisms with environmental sex determination. For organisms with temperature-dependent sex determination (TSD), variation in nest temperatures induces phenotypic variation that could impact population growth rates. In environments that vary over space and time, how does this variation influence key demographic parameters (cohort sex ratio and hatchling recruitment) in early life stages of populations exhibiting TSD? We leverage a 17-year data set on a population of painted turtles, Chrysemys picta, to investigate how spatial variation in nest vegetation cover and temporal variation in climate influence early life-history demography. We found that spatial variation in nest cover strongly influenced nest temperature and sex ratio, but was not correlated with clutch size, nest predation, total nest failure, or hatching success. Temporal variation in climate influenced percentage of total nest failure and cohort sex ratio, but not depredation rate, mean clutch size, or mean hatching success. Total hatchling recruitment in a year was influenced primarily by temporal variation in climate-independent factors, number of nests constructed, and depredation rate. Recruitment of female hatchlings was determined by stochastic variation in nest depredation and annual climate and also by the total nest production. Overall population demography depends more strongly on annual variation in climate and predation than it does on the intricacies of nest-specific biology. Finally, we demonstrate that recruitment of female hatchlings translates into recruitment of breeding females into the population, thus linking climate (and other) effects on early life stages to adult demographics.     

High social density increases foraging and scouting rates and induces polydomy in Temnothorax ants

Many organisms live in crowded groups where social density affects behavior and fitness. Social insects inhabit nests that contain many individuals where physical interactions facilitate information flow and organize collective behaviors such as foraging, colony defense, and nest emigration. Changes in nest space and intranidal crowding can alter social interactions and affect worker behavior. Here, I examined the effects of social density on foraging, scouting, and polydomy behavior in ant colonies—using the species Temnothorax rugatulus. First, I analyzed field colonies and determined that nest area scaled isometrically with colony mass—this indicates that nest area changes proportionally with colony size and suggests that ants actively control intranidal density. Second, laboratory experiments showed that colonies maintained under crowded conditions had greater foraging and scouting activities compared to the same colonies maintained at a lower density. Moreover, crowded colonies were significantly more likely to become polydomous. Polydomous colonies divided evenly based on mass between two nests but distributed fewer, heavier workers and brood to the new nests. Polydomous colonies also showed different foraging and scouting rates compared to the same colonies under monodomous conditions. Combined, the results indicate that social density is an important colony phenotype that affects individual and collective behavior in ants. I discuss the function of social density in affecting communication and the organization of labor in social insects and hypothesize that the collective management of social density is a group level adaptation in social insects.     

Nest-maintenance effort and health status in chinstrap penguins, Pygoscelis antarctica: the functional significance of stone-provisioning behaviour

Stone provisioning is a nest-maintenance activity accomplished by pygoscelid penguins after reliefs during the incubation/brooding period. The functional significance of this behaviour has been mainly explained as a parental strategy preventing nest flooding under detrimental weather conditions. In addition, and in the light of recent studies, this behaviour could also fit into the sexual selection process. In this study, we tested the first idea, that is, whether stone provisioning is a nest-maintenance behaviour to increase egg/nestling survival by lowering the risk of nest flooding, and can thus be considered a form of parental care. Additionally, we investigated if the effort invested by parents in nest maintenance is constrained by physiologically limiting resources. The effort of stone collection and the perceived risk of nest flooding were experimentally manipulated during the incubation and early brooding phases in a chinstrap penguin, Pygoscelis antarctica, colony. Three groups of nests were established. After weighing, control nests were left unmanipulated. In a second group of nests (reduced group), only one-half of the initial weight of stones was returned to the nests. In a third group of nests (snow-added group), we both reduced nest weight by a half and added snow outside the nest bowl over 6 consecutive days. Ten days after manipulation, the difference in nest weight between initial and final conditions was significantly related to treatment: penguins increased stone provisioning in the reduced group (44% of half-reduced nests), but drastically more in the reduced and snow-added group (123% of half-reduced nests), while the weight of control nests was unchanged compared to premanipulation conditions. The intensity of stone provisioning was affected by nest date, peaking about hatching time and shortly after, and declining with advancing chick age. These results suggest that stone provisioning is a mechanism that has evolved to prevent egg or chick mortality by nest flooding. The haematocrit, but not leukocyte numbers as expressed by the buffy coat layer, varied with the experimental conditions. Penguins investing more time in nest maintenance had a lower haematocrit, suggesting a physiological trade-off probably mediated by competition between the time devoted to nest maintenance versus foraging activities. The amount of stones collected and the haematocrit were positively related to the number of neighbour nests, so those individuals surrounded by more nests seemed to obtain benefits in the availability of nest material and energy savings. This study indicates that stone-provisioning behaviour is a nest-maintenance activity evolved to improve thermal nest characteristics potentially increasing offspring survival, and competing in time and energy with other reproductive activities. Stone provisioning in penguins should therefore be regarded as a form of parental care and an important part of individual reproductive effort in species breeding in harsh environments. Furthermore, nest size and nest-maintenance effort should be considered reproductive traits indicative of parental quality and thus could also be involved in the post-mating sexual selection process.