Extensive regional variation in the phenology of insects and their response to temperature across North America.

Climate change models often assume similar responses to temperatures across the range of a species, but local adaptation or phenotypic plasticity can lead plants and animals to respond differently to temperature in different parts of their range. To date, there have been few tests of this assumption at the scale of continents, so it is unclear if this is a large-scale problem. Here, we examined the assumption that insect taxa show similar responses to temperature at 96 sites in grassy habitats across North America. We sampled insects with Malaise traps during 2019-2021 (N = 1041 samples) and examined the biomass of insects in relation to temperature and time of season. Our samples mostly contained Diptera (33%), Lepidoptera (19%), Hymenoptera (18%), and Coleoptera (10%). We found strong regional differences in the phenology of insects and their response to temperature, even within the same taxonomic group, habitat type, and time of season. For example, the biomass of nematoceran flies increased across the season in the central part of the continent, but it only showed a small increase in the Northeast and a seasonal decline in the Southeast and West. At a smaller scale, insect biomass at different traps operating on the same days was correlated up to ~75 km apart. Large-scale geographic and phenological variation in insect biomass and abundance has not been studied well, and it is a major source of controversy in previous analyses of insect declines that have aggregated studies from different locations and time periods. Our study illustrates that large-scale predictions about changes in insect populations, and their causes, will need to incorporate regional and taxonomic differences in the response to temperature.

Pre-fledging quality and recruitment in an aerial insectivore reflect dynamics of insects, wetlands and climate.

Wetland systems, including shallow palustrine ponds, are hotspots for emergent aquatic insects but are globally threatened by land-use practices and climate change. Loss of insects is hypothesized as a key driver of population declines in aerial insectivores, but studies of climate-driven fluctuations in pond abundance during wet-dry periods and aerial insects on nestling quality and apparent recruitment are lacking. Using tree swallow (Tachycineta bicolor) data spanning 14-28 years we evaluated: (1) whether nestling quality based on pre-fledging (~ 12 days old) body mass changed over the time series; (2) how annual estimates of aerial insect biomass and variability, temperature, and pond abundance influenced nestling mass; and (3) whether the annual number of recruits produced was related to the annual mean mass of nestlings, aerial insects, and pond abundance in their year of hatching. Average nestling body mass varied annually but no long-term temporal trends were detected. Nestlings were heavier when raised during periods of stable insect biomass, warmer temperatures, and higher pond abundance. Pond abundance consistently had strong effects on nestling mass and inter-annual apparent recruitment, suggesting that this metric provides a complementary index of either higher prey abundance or higher-quality aquatic prey. Overall, pre-fledging quality and annual recruitment of nestling tree swallows reflects dynamic interannual changes in climate, pond availability, and aerial insect food supply. Our findings further suggest the abundance of ponds in this semi-arid prairie landscape is likely a strong predictor of regional population stability in tree swallows and possibly other ecologically similar species.

The relative contribution of individual quality and changing climate as drivers of lifetime reproductive success in a short-lived avian species.

Animal populations are influenced strongly by fluctuations in weather conditions, but long-term fitness costs are rarely explored, especially in short-lived avian species. We evaluated the relative contributions of individual characteristics and environmental conditions to lifetime reproductive success (LRS) of female tree swallows (Tachycineta bicolor) from two populations breeding in contrasting environments and geographies, Saskatchewan and British Columbia, Canada. Female swallows achieved higher LRS by breeding early in the season and producing more fledglings. Other measures of female quality had virtually no influence on LRS. Genetic factors did not predict LRS, as there was no correlation between life-history components for sister pairs nor between mothers and their daughters. Instead, climate variability-indexed by spring pond density (i.e., abundance of wetland basins holding water) during years when females bred-had strong positive effects on female LRS in more arid Saskatchewan but only weak positive effects of moisture conditions were detected in wetter British Columbia. Overall, several life history trait correlates of LRS were similar between populations, but local environmental factors experienced by individuals while breeding produced large differences in LRS. Consequently, variable and extreme environmental conditions associated with changing climate are predicted to influence individual fitness of distinct populations within a species' range.

A range-wide domino effect and resetting of the annual cycle in a migratory songbird.

Latitudinal differences in timing of breeding are well documented but how such differences carry over to influence timing of events in the annual cycle of migratory birds is not well understood. We examined geographical variation in timing of events throughout the year using light-level geolocator tracking data from 133 migratory tree swallows ( Tachycineta bicolor) originating from 12 North American breeding populations. A swallow's breeding latitude influenced timing of breeding, which then carried over to affect breeding ground departure. This resulted in subsequent effects on the arrival and departure schedules at autumn stopover locations and timing of arrival at non-breeding locations. This 'domino effect' between timing events was no longer apparent by the time individuals departed for spring migration. Our range-wide analysis demonstrates the lasting impact breeding latitude can have on migration schedules but also highlights how such timing relationships can reset when individuals reside at non-breeding sites for extended periods of time.

Assessing costs of carrying geolocators using feather corticosterone in two species of aerial insectivore.

Despite benefits of using light-sensitive geolocators to track animal movements and describe patterns of migratory connectivity, concerns have been raised about negative effects of these devices, particularly in small species of aerial insectivore. Geolocators may act as handicaps that increase energetic expenditure, which could explain reported effects of geolocators on survival. We tested this 'Energetic Expenditure Hypothesis' in 12 populations of tree swallows (Tachycineta bicolor) and barn swallows (Hirundo rustica) from North America and Europe, using measurements of corticosterone from feathers (CORTf) grown after deployment of geolocators as a measure of physiology relevant to energetics. Contrary to predictions, neither among- (both species) nor within-individual (tree swallows only) levels of CORTf differed with respect to instrumentation. Thus, to the extent that CORTf reflects energetic expenditure, geolocators apparently were not a strong handicap for birds that returned post-deployment. While this physiological evidence suggests that information about migration obtained from returning geolocator-equipped swallows is unbiased with regard to levels of stress, we cannot discount the possibility that corticosterone played a role in reported effects of geolocators on survival in birds, and suggest that future studies relate corticosterone to antecedent factors, such as reproductive history, and to downstream fitness costs.

No selection on immunological markers in response to a highly virulent pathogen in an Arctic breeding bird.

In natural populations, epidemics provide opportunities to look for intense natural selection on genes coding for life history and immune or other physiological traits. If the populations being considered are of management or conservation concern, then identifying the traits under selection (or 'markers') might provide insights into possible intervention strategies during epidemics. We assessed potential for selection on multiple immune and life history traits of Arctic breeding common eiders (Somateria mollissima) during annual avian cholera outbreaks (summers of 2006, 2007 & 2008). We measured prelaying body condition, immune traits, and subsequent reproductive investment (i.e., clutch size) and survival of female common eiders and whether they were infected with Pasteurella multocida, the causative agent of avian cholera. We found no clear and consistent evidence of directional selection on immune traits; however, infected birds had higher levels of haptoglobin than uninfected birds. Also, females that laid larger clutches had slightly lower immune responses during the prelaying period reflecting possible downregulation of the immune system to support higher costs of reproduction. This supports a recent study indicating that birds investing in larger clutches were more likely to die from avian cholera and points to a possible management option to maximize female survival during outbreaks.

Sex-specific effects of increased incubation demand on innate immunity in black guillemots.

Life-history theory predicts that there should be negative fitness consequences, in terms of future reproduction and survival, for parents with increased reproductive effort. We examined whether increased incubation demand affected innate immunity and body condition by performing a clutch-size manipulation experiment in black guillemots (Cepphus grylle). We found that plasma from males incubating experimentally enlarged clutches exhibited significantly reduced lysis titers compared with plasma from males incubating control clutches, while this was not observed in females. The increased incubation demand also impacted agglutination titers differently in males and females, although the effect of treatment was not significant in either sex. Among all birds, lysis titers increased and haptoglobin concentrations decreased from mid- to late incubation. Natural antibody-mediated agglutination titers and body condition were highly repeatable within the incubation bout and between years. This suggests that agglutination titers may serve as a reliable and resilient index of the immunological character of individuals in future studies. Overall, this study demonstrates that increased incubation demand impacts indices of innate immunity differently in males and females. The potential for different components of the immune system to be impacted sex-specifically should be considered in future studies linking immune function and life-history trade-offs.

Sex-specific effects of handling time on an index of immune activity in zebra finches.

Recently, there has been considerable interest in the role of the immune system in shaping life-history evolution, sexual selection strategies, and indexes of individual quality. The most frequently used assay of immune function, particularly in avian field studies, is the phytohemagglunitin (PHA) skin test. PHA is injected subcutaneously into the wing web, and the magnitude of the resultant swelling has traditionally been interpreted as an index of an individual's cell-mediated immunocompetence. The test follows one of two protocols: the traditional two-wing injection protocol, with one wing web injected with PHA and the other with phosphate-buffered saline (PBS), or the simplified one-wing protocol that omits the PBS injection. In this technical comment, we alert researchers to the importance of considering handling time when performing the PHA test. We show that zebra finches (Taeniopygia guttata) subjected to the two-wing protocol had a lower wing-web swelling than individuals injected in one wing. In males, handling time explained over 50% of the variation in an individual's skin swelling response; females were relatively unaffected by handling time. We suggest that caution should be exercised when comparing the magnitude of wing-web swelling across studies in which the alternate protocol was followed. In addition, the recording of handling time, and its inclusion in subsequent statistical analyses, may aid in the detection of subtle differences across treatments.