Journal of Experimental Zoology Part A: Ecological and Integrative Physiology

Developing in suboptimal temperatures can have widespread negative effects in organisms, but early exposure to thermal challenges can also trigger increased investment in thermoregulatory capacity. Within birds, the effects of cold exposure during incubation may differ between precocial and altricial species due to differences in the ontogeny of thermoregulation. To date, most of the evidence showing that embryonic cold exposure shapes future thermoregulatory capacity comes from a few precocial species in laboratory settings. Less is known about sensitivity to low-temperature incubation in altricial species which only become fully endothermic after hatching. In this study, we tested the effects of cold temperatures during embryonic development by temporarily cooling the nest boxes of free-living tree swallows (Tachycineta bicolor) during late incubation and subsequently exposing nestlings to an acute cold challenge. Cold exposure during early development made nestlings better at maintaining their body temperatures during the acute challenge. Developmentally cold-exposed birds also had higher thyroxine (T4) levels when sampled at lower ambient temperatures, whereas control birds showed the opposite relationship. Among nestlings that failed to maintain their body temperature during the acute cold challenge, developmentally cold-exposed nestlings secreted more corticosterone during the acute cold challenge than control nestlings. Despite these changes, cold-exposed birds did not differ from controls in cold-induced metabolic rates, body mass, or bacteria killing ability. These results are, to our knowledge, the first to show that developmental cold exposure increases the capacity to maintain body temperature during future cold challenges in a songbird. These findings suggest that experiencing even a mild decline in ambient temperatures during embryonic development could prime birds to cope more effectively with harsh or variable environmental conditions later in life.

Heat-induced sterility is widespread across ectotherm species, yet its underlying mechanisms remain poorly understood. In mated females, thermal stress can disrupt female reproductive physiology while also affecting sperm stored within the reproductive tract, making the mechanisms of fertility loss difficult to distinguish. Here, we investigated the extent and recovery dynamics of female heat-induced fertility loss across seven Drosophila species. We tracked fertility after acute heat exposure, tested whether remating restored fertility, quantified stored sperm in two closely related species, and examined whether reproductive thermal sensitivity covaried with adult heat tolerance. We found that heat exposure reduced egg-hatching success in all seven species, but recovery differed among species. Fertility recovered within several days in most species, whereas D. simulans showed persistent declines and D. melanogaster showed limited recovery. Remating restored or maintained fertility in both species. Stored sperm declined through time in both D. simulans and D. mauritiana, but D. mauritiana retained more sperm at later sampling points, consistent with their contrasting patterns of fertility decline and maintenance. Across species, greater adult heat tolerance was associated with smaller immediate reductions in fertility, but not with cumulative reproductive loss across subsequent days. Together, these findings show that similar initial fertility losses can lead to markedly different recovery trajectories and highlight the importance of comparative, multi-species approaches that integrate female physiology, sperm-storage dynamics, mating behavior, and whole-organism thermal tolerance when assessing reproductive resilience to climate warming.

Traditional veterinary pathology investigations on wild animals generally conclude a primary cause of death based on a combination of observed clinical signs, gross and microscopic lesions, microbial culture and PCR. However, these methods may fail to capture the broader microbial community of the host. We employed metatranscriptomic sequencing to characterise the potential pathobiome of necropsied marine turtles, comparing these findings with corresponding veterinary pathology reports. Tissue samples were collected from 76 individuals spanning three species of marine turtles -- the green turtle (Chelonia mydas), the hawksbill turtle (Eretmochelys imbricata), and the loggerhead turtle (Caretta caretta) -- across the coastline of New South Wales, Australia. Reported causes of mortality ranged from infectious disease to trauma. We identified a diverse range of parasites within the genera Neospirochis, Hapalotrema, Paraorgymatobothrium, Eimeria and Callitetrarhynchus, and bacteria including the genera Aeromonas, Clostridium, and Vibrio. Novel viruses of the families Astroviridae, Caliciviridae, Hepaciviridae, Hepeviridae, Nyamivirdae, Orthomyxoviridae, Picobirnaviridae and Rhabdoviridae were also identified. In cases where infectious disease was noted in the pathology report, we frequently observed high pathogen read counts consistent with the listed primary diagnoses. However, we also detected a range of incidental infections involving eukaryotic parasites, bacteria, and viruses. The diversity of infectious agents detected in marine turtles suggests that the observed disease outcomes are unlikely to be driven by a single causative agent, but rather by complex interactions between host health, the pathobiome, and environmental stressors. We discuss the potential implications for turtle health and conservation.

Parasites decrease host fitness, more so during physiologically challenging life stages such as migration. However, parasites are unevenly distributed among individuals, and hosts with different physiological condition may face different costs of migration with infections. We tested whether the relationship between infection by haemosporidian parasites and oxidative balance during autumn migration in blackcaps depends on individual age and sex, two major sources of variation in prevalence and physiological condition. We captured blackcaps at stopover in central Spain and measured infection status and physiological parameters. Blackcaps physiologically more prepared to depart from stopover had lower antioxidant capacity of plasma, probably because of physiological adjustments prior to the upcoming flight stage. Plasmodium parasites occurred only in coinfections, which were more frequent among young blackcaps. Regardless of individual sex and age, coinfected birds had lower antioxidant capacity of plasma, and blackcaps coinfected with Plasmodium had more oxidative damage to lipids in their red blood cells. Our results support the idea that haemosporidian infections impair the oxidative balance of blackcaps during autumn migration, which could impair individual performance and ultimately fitness, with increased population impacts during the first year of life when a larger proportion of individuals migrate coinfected with harmful Plasmodium parasites.

Reliable morphological criteria for sexing birds without the need for invasive procedures are scientifically and ethically valuable. Smith et al (2005) reported that iris colour assigns the genetic sex of Starlings Sturnus vulgaris with 98% accuracy, and bill base colour with 100% accuracy when birds are in breeding condition, but these figures have not been tested independently. We applied their criteria to 31 captive Starlings in breeding condition, using three observers blind to genetic sex, two of them new to Starling sexing. Bill base colour correctly predicted genetic sex for all 29 birds scored, with complete observer agreement. Mean iris score was correct for 27 of 31 (87.1%), fewer than the 94 of 96 reported by Smith et al (2005) for birds in breeding condition (Fisher exact test, P = 0.031), although with so few errors in either study this comparison is imprecise. Observer agreement was high, so these errors reflect iris colouration conflicting with genetic sex rather than inconsistent scoring; bill base colour was correct for all four misassigned birds. Body mass and tarsus length classified 77% and 58% of birds correctly, their published thresholds missing in opposite directions. We conclude that bill base colour should be the primary criterion for sexing Starlings in breeding condition; the accuracy of iris colour should not be assumed to be 98%.

Despite being a topic of early experimental studies, the effects of immersion-emersion cycles and temperature on the heart rate of intertidal Mytilus mussels have not been fully characterized. Using a recently released intertidal chamber paired with our own software package, we conducted a multiday experiment and continuously monitored the heart rate of the blue mussel Mytilus edulis. Under normal conditions, we detected a subtle spike in heart rate upon emersion before the eventual decline, as well as an overshoot upon reimmersion. Additionally, heart rate was sensitive to temperature changes during immersion but less responsive during emersion. These patterns were markedly altered following an unintended prolonged emersion event, which led to the loss of the emersion spike, an extended reimmersion overshoot, and heightened thermal sensitivity on top of an overall elevation in heart rate. However, the heart rate gradually returned to the pre-event baseline over the following days, demonstrating the significant resilience of the mussels. Our study highlights the potential of high-resolution physiological data in assessing the effects of environmental stressors on coastal ecosystems.

Marine heatwaves cause widespread coral bleaching and mortality, yet thermal responses often differ between adult and juvenile corals, with the latter frequently being more tolerant. Whether the observed enhanced tolerance of juveniles reflects intrinsic physiological differences across life stages or simply reduced environmental stress due to shaded microhabitats occupied by juveniles, or a combination of both factors, remains unclear. To test whether colony size determines thermal responses and if a threshold size exists for tolerance, we exposed 89 colonies of the reef-building coral Acropora aff. digitifera to a 6-week, 32.5 C simulated marine heatwave under controlled light conditions, with cumulative stress reaching 20 C-weeks. Contrary to expectations, ~2 C-weeks less heat stress was required to trigger bleaching and mortality in coral juveniles compared to adults. Mortality divergence across colony sizes was greatest at ~10 C-weeks, when the smallest corals exceeded 80% mortality, whereas the largest colonies showed <10% mortality. A breakpoint in survival odds was apparent at 13.4 cm diameter, corresponding to the size at onset of sexual maturity in the native wild population for this species. These results indicate an ontogenetic shift in physiological heat tolerance, with larger reproductively mature colonies showing enhanced tolerance. The apparent survival advantage of juveniles in the reef frequently reported during bleaching events could be more likely due to micro-environmental buffering than intrinsic physiological tolerance. Our findings suggest that juvenile corals may be more vulnerable to climate change than previously thought, particularly as declining habitat complexity could reduce microhabitat refugia on future reefs.

Climate change is threatening coral reef ecosystems around the world, causing significant declines in reef organisms, including sponges. The health of sponges is intimately linked to their symbiosis with microorganisms, yet little is known about their integrated response to ocean warming. Here, we present a comprehensive view of the effect of elevated sea surface temperatures (SST) on the model sponge Ianthella basta. I. basta individuals were exposed to temperatures up to 32 {degrees}C, 2-3 {degrees}C above the mean annual maxima. Host gene expression profiles showed an initial response to heat stress after 2 days, and a decreased expression of genes involved in maintaining tissue integrity, pumping capacity, vitamin transport, and stable symbioses after 4 to 8 days. Congruently, heat stressed samples started to show tissue regression and a reduced abundance of the dominant symbionts after 4 days. A significant loss of the symbionts occurred after 8 days, together with a reduction in microbially driven nitrogen cycling and amino-acid transport and an introduction of opportunistic microbes, which could lead to nutrient imbalance and further destabilisation of the holobiont. Together, this work provides important insights into the avenues through which future SST can impact host-microbe interactions in coral reef sponges.

Immune defenses may vary across environments due to differences in life history or abiotic and biotic characteristics, but few studies have compared the strength of immune responses across broad geographical gradients. Here we investigated how bacteria killing ability (BKA), a measure of constitutive innate immune function, varies among populations of tree swallows (Tachycineta bicolor) inhabiting different environments. We hypothesized that populations living in unpredictable environments, which also have shorter breeding seasons and thus fewer opportunities to renest in the event of reproductive failure, would invest in reproduction at the expense of immunity. To test this hypothesis, we measured BKA and monitored reproductive activity in four populations located across most of the breeding range of tree swallows, in Tennessee, New York, Wyoming and Alaska. Consistent with our predictions, BKA was lowest in Alaska and Wyoming and highest in Tennessee. In most populations BKA was also lower during incubation than during nestling provisioning, and lower in males than in females. These results suggest that the high reproductive investment seen in populations experiencing more unpredictable breeding conditions may come at a cost to immune function. While reduced immune investment could increase susceptibility to parasites, this potential cost is likely offset by lower parasite pressure in these environments, limiting the need for maintaining strong constitutive innate defenses like BKA.

Fledglings of burrow-nesting seabirds are vulnerable to disorientation by artificial light at night (ALAN) and grounding in coastal human settlements, where most individuals presumably perish if not rescued. Accurate estimates of the effects of stranding on seabird populations remain elusive due to several levels of uncertainty, such as the number of stranded seabirds estimated to remain undiscovered by rescuers. One species subjected to stranding is the Atlantic puffin (Fratercula arctica). We hypothesized that the detection probability of puffin fledglings by the Puffin Patrol rescue program, operating in Newfoundland, Canada, is affected by light level, concealment level, and search effort. We deployed realistic puffin decoys in locations of varying light and concealment levels throughout two sites that are searched nightly by rescuers and measured the amount of time and search effort required until each decoy was found. Approximately 78% more search effort was required to find a decoy in darkness (32.0 min) compared with illuminated conditions (18.0 min), and up to 100% more search effort was required to find a decoy placed in concealed conditions (35.5 min) than one in the open (17.5 min). After two hours of searching, on average over 10% of decoys were still not found, although each additional minute of search effort increased detection rate of an individual decoy by 0.6%. In combination with recent advances in our knowledge of seabird post-stranding behaviour, our study offers a first step in the development of correction factors which can be adapted to site- and species-specific requirements of other rescue programs around the world that rely on volunteers to search for stranded birds. Application of such correction factors will hopefully result in better estimation of mortality rates and their resultant effects on populations.

In Baltic grey seals (Halichoerus grypus), interannual variation in sea-ice availability can alter the proportion of females breeding on land, generating substantial variation in local breeding density among years. We investigated preweaning survival, post-weaning residence, and weaning body condition at a Baltic grey seal land-breeding colony over two years with contrasting colony densities. Drone imagery provided repeated counts of pups in distinct developmental stages and estimates of moulting pup mass as an index of weaning body condition. A Bayesian state-space population model estimated substantially greater pup production in the high- (282.5 [95% CrI = 252.8 to 320.9] pups) than the low-density (188.3 [95% CrI =168.9 to 210.8] pups) year, but similar preweaning survival (0.66 [95% CrI = 0.53 to 0.81] and 0.67 [95% CrI = 0.56 to 0.79], respectively). In contrast, post-weaning residence differed substantially between the high- (11.8 [95% CrI = 9.2 to 15.7] days) and low-density (30.3 [95% CrI = 23.1 to 39.1] days) years. Greater exposure to other pups during early life was associated with lower weaning body mass ({beta} = -0.45 [-0.89 to -0.01], P[{beta} < 0] = 0.98). Together, these results suggest that the consequences of higher breeding density may emerge through poorer weaning condition and altered post-weaning residence even when effects on preweaning survival are not detectable.

Vertebrates cope with rapid temperature fluctuations by plastically adjusting their behaviour and physiology. This phenotypic plasticity is partly mediated by plastic changes in glucocorticoid levels; however, whether individual differences in endocrine plasticity are associated with fitness remains unknown. Here, we examined the relationship between individual variation in glucocorticoid plasticity in response to air temperature changes and reproductive success in a wild great tit population (Parus major). Over nine years, we collected repeated measurements of baseline and stress-induced corticosterone during nestling rearing. We found consistent reaction norm patterns in hormone response to temperature variation -- as shown in a previous study conducted over a shorter period. This provides robust evidence of individual differences in average hormone levels (intercepts) and the strength of the plastic response (slopes), indicating potential for evolution. However, neither intercepts nor slopes were associated with reproductive success (fledgling number or brood mass), suggesting that individual variation in endocrine plasticity was not under natural selection, at least within the environmental conditions experienced. This study provides a first empirical test of the evolutionary implications of endocrine plasticity in the wild and highlights the need to assess its fitness relevance under more extreme conditions, such as those expected under future climate scenarios.

Metabolic rate governs the pace of life through the allocation and expenditure of energy, yet metabolic rate data are unavailable for most species. External morphological traits, such as gill slits, have been used to infer respiratory physiology and metabolism in sharks, yet, whether that pattern is generalizable across rays is unknown. In rays (Batoidea), spiracle length seems adaptive to oxygen uptake capacity, environmental oxygen availability, and activity level. Thus, we evaluate the allometry of external respiratory morphology (spiracles and gill slits) in rays using 550 museum specimens representing 88 species. We test whether spiracle length and gill slit height grow isometrically with body size (total length or disc width) within and across species. Overall, we find spiracles and gill slits are morphologically diverse, varying widely in shape within and between families. Spiracle length and gill slit height scale isometrically and are a consistent proportion of body size, despite such high morphological variability. While that proportion is consistent through ontogeny, each species proportion differs (e.g. gill slit height ranges from 0.93% of total length in Spotted Legskate to 2.93% in Pacific Electric Ray). Of the three swimming styles, pectoral oscillating species (Mobulid and Eagle Rays) may have steeper scaling of spiracle length with gill slit height - consistent with greater respiratory capacity in species with more active lifestyles. Our findings are consistent with literature hypothesizing external respiratory morphology (spiracles and gill slits) is tightly related to metabolic rate and activity level, and demonstrate the value of natural history collections in ecological research. Importantly, the isometry of both respiratory traits means that a single measure of any individual, irrespective of size, will be representative of the species thus providing a basis for expanding our understanding of metabolic morphology, enabling future work on life history, extinction risk, and ecological function.

1. Organisms rely on environmental cues, such as moonlight illumination and weather, to modulate nocturnal behaviors that reduce predation risk and conserve energy during demanding periods. Urbanization can alter the reliability of cues, potentially impairing organisms' ability to respond appropriately with consequences in increasingly urbanized landscapes. 2. We leveraged the Motus Wildlife Tracking System to identify roost initiation, roost departure and nocturnal restlessness of dark-eyed juncos (Junco hyemalis) overwintering along an urban to rural gradient to determine if activity patterns are affected by weather, moonlight illumination, and urbanization. 3. Nocturnal behavior of overwintering juncos varied with environmental factors, including moonlight illumination and temperature. However, the strength of these responses differed among sites, suggesting that natural cues of moonlight illumination were masked in urban areas, where individuals were more responsive to temperature. 4. Our results suggest that urbanization weakens birds' responsiveness to natural light cues, likely because artificial lighting reduces the visibility of moonlight; representing a key mechanism by which human-altered landscapes can disrupt nocturnal ecology in birds.

Early environmental conditions can have long-lasting effects on survival and reproductive fitness. Using a 23 year-long monitoring data set of captive mouse lemur's life history traits, we tested a relationship between maternal allocation to offspring (N = 1365) and their adulthood survival. Maternal characteristics such as age or body condition did not affect allocation to offspring whatever the litter size (1 to 3). Although birth mass depended on size and composition of the litters, mouse lemur survival was highly correlated with body mass acquired after weaning in both sexes, through potential sibling competition. Moreover, female's reproductive success correlated with this body mass and was consistently associated with increased longevity. These findings suggest the presence of a silver spoon effect under constant captive conditions. However, cumulative effects of genetic and adulthood social conditions strongly interact to affect adult survival. Deaths related to intra- or inter-sexes aggressive social interactions may outweigh effect of early environment and therefore minimize the silver-spoon effect on captive mouse lemur's survival. However, having a high body mass after weaning appeared to be a determinant factor in individual survival for mouse lemurs.

1. Temperature shapes life histories through its effects on energy allocation among growth, reproduction and maintenance. The resulting temperature dependence of performance is commonly characterised using thermal performance curves (TPCs), which are often treated as static descriptions that apply uniformly throughout life. Yet the response to thermal variability may change across ontogeny. 2. We recorded growth, reproduction and survival throughout the lifetimes of individual Daphnia magna under four thermal regimes: Constant Low (15C), Constant Rearing (20C), Constant High (25C) and Variable (15C to 25C). The mean temperature under Variable closely matched that under Constant Rearing. Using an allometric energy-allocation model, we derived age specific and lifetime performance measures and quantified their uncertainty with a stratified individual-level bootstrap. Our principal quantity was the Jensen contrast, defined as performance under Variable minus that under Constant Rearing. 3. Age specific point estimates of the Jensen contrast changed sign across ontogeny. A positive aggregate contrast in full trajectory growth concealed a negative mid-life contrast; reproductive point estimates were positive during early and mid life but negative later. Bootstrap support was strong for the positive early life and negative mid-life growth contrasts, and for the positive early life reproductive contrast. Under a second-order approximation, this pattern of estimated signs is consistent with ontogenetic changes in effective thermal curvature. 4. The positive early life reproductive response did not carry through to the lifetime outcome: expected lifetime reproduction was lower under Variable than under Constant Rearing, with strong bootstrap support for the negative lifetime contrast. Because lifetime reproduction is survival weighted, lower survival under Variable reduced later reproductive contributions; across the estimated trajectories, the accumulated negative late life contribution out-weighed the earlier positive contributions. 5. These results show why forecasts based on mean temperature, a single life stage or an aggregate TPC may misrepresent lifetime performance under variable thermal regimes. Accounting for both ontogeny and temperature variability may therefore improve forecasts of life-history performance in increasingly variable thermal environments.

Urbanisation imposes novel environmental conditions that can shape phenotypic traits including plumage, yet, its effects on characteristics beyond colouration remain poorly understood. Feather microstructure plays an important role in thermoregulation, protection from solar radiation and waterproofing, and its expression is sensitive to environmental conditions. Here, we quantified feather barbule density, a key component of feather microstructure, in great tits (Parus major) from five urban-rural paired populations across Europe to assess its association with urbanisation, temperature and elevation. We found no significant effects of the urban environment or temperature. However, feather microstructure significantly varied between the studied areas and was positively associated with elevation and ultraviolet B (UV-B) radiation, suggesting a potential adaptive response to variation in solar radiation.

Sea turtle conservation relies largely on the identification of ecologically significant marine areas. Green sea turtles (Chelonia mydas) are resident in Aotearoa New Zealand waters, yet much of their spatial ecology remains undefined. To address this, satellite telemetry was used to map juvenile and sub-adult green sea turtle spatial movements, residency patterns, and habitat associations. Our findings provide the first evidence of individual fine-scale habitat use and long-term residency in New Zealand waters. Data will contribute to regional and global efforts to identify ecologically significant sea turtle habitat, including Important Marine Turtle Areas (IMTAs) and Blue Corridors for Turtles (BC4T), and help advance conservation efforts.

Fluctuating asymmetry (FA) is widely used as a proxy for developmental instability, although reduced rather than increased FA, has occasionally been reported under particularly challenging conditions, possibly because environmental filtering removes less developmentally stable individuals. Here, we investigated temporal variation in wing shape asymmetry in males of Drosophila sturtevanti collected in 2022, 2024, and 2026 at the same sampling site, an urban Atlantic Forest fragment. Sampling in 2022 and 2026 were completed during single field expeditions in early April, whereas in 2024 repeated collections were required until mid-May because of markedly low capture success. We therefore examined whether this difference was associated with unusual environmental conditions and whether FA was higher in 2024, as expected under greater developmental perturbation. Daily temperature and relative humidity were examined from early March onward, encompassing approximately one month before adult emergence. The 2024 sampling period was warmer and drier than the corresponding periods in 2022 and 2026 and coincided with the final phase of the strong 2023-2024 El Ni&ntildeo event. Contrary to our prediction, FA was significantly lower in 2024. The magnitude of directional asymmetry remained similar among years, although the spatial pattern of the DA vector differed between 2024 and 2026. The coincidence of reduced capture success and reduced FA is consistent with the hypothesis that adverse conditions may impose demographic filtering, such that individuals expressing greater developmental instability are less likely to contribute to the sampled adult population.

Social relationships can buffer individuals against environmental challenges, reducing the physiological costs of disturbance. Assessing how social and environmental conditions jointly shape physiological stress is critical for predicting individual responses to disturbance, particularly in managed populations. We evaluated whether social and environmental stressors act additively or whether social context modifies physiological responses to environmental challenges. We combined behavioural data and faecal glucocorticoid metabolite samples from a small, intensively monitored population of African elephants (Loxodonta africana) in a fenced reserve. Using a Bayesian framework, we tested how social factors interacted with variation in the biophysical environment, including natural environmental and anthropogenic disturbances, to predict glucocorticoid concentrations. Natural environmental variables, including weather, water availability, and season, did not predict glucocorticoid concentrations. In contrast, helicopter use, a common management intervention, was associated with higher glucocorticoid concentrations. Social effects were context dependent. Following disturbance, low-ranking individuals showed higher glucocorticoid concentrations than high-ranking individuals, whereas this pattern reversed under low-disturbance conditions. Individuals with more social connections had lower glucocorticoid concentrations, but this effect was evident only under non-disturbance conditions. Measures of betweenness centrality did not predict glucocorticoid levels. These results demonstrate that social factors mediate physiological responses to anthropogenic disturbance and that management actions can impose uneven physiological costs across individuals. Quantifying interactions between social factors and the biophysical environment may improve conservation and management strategies for social species. Integrating physiological responses to interactions between social factors and the biophysical environment can guide conservation strategies and improve management interventions in social species.