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Artificial light at night (ALAN) is a globally widespread and expanding form of anthropogenic change that impacts arthropod biodiversity. ALAN alters interspecific interactions between arthropods, including predation and parasitism. Despite their ecological importance as prey and hosts, the impact of ALAN on larval arthropod stages, such as caterpillars, is poorly understood. We examined the hypothesis that ALAN increases top-down pressure on caterpillars from arthropod predators and parasitoids. We experimentally illuminated study plots with moderate levels (10–15 lux) of LED lighting at light-naive Hubbard Brook Experimental Forest, New Hampshire. We measured and compared between experimental and control plots: (i) predation on clay caterpillars, and (ii) abundance of arthropod predators and parasitoids. We found that predation rates on clay caterpillars and abundance of arthropod predators and parasitoids were significantly higher on ALAN treatment plots relative to control plots. These results suggest that moderate levels of ALAN increase top-down pressure on caterpillars. We did not test mechanisms, but sampling data indicates that increased abundance of predators near lights may play a role. This study highlights the importance of examining the effects of ALAN on both adult and larval life stages and suggests potential consequences of ALAN on arthropod populations and communities. [ABSTRACT FROM AUTHOR]

Sexual selection studies widely estimate several metrics, but values may be inaccurate because standard field methods for studying wild populations produce limited data (e.g., incomplete sampling, inability to observe copulations directly). We compared four selection metrics (Bateman gradient, opportunity for sexual selection, opportunity for selection, and s max ′ ) estimated with simulated complete and simulated limited data for 15 socially monogamous songbird species with extrapair paternity (4%–54% extrapair offspring). Inferring copulation success from offspring parentage creates nonindependence between these variables and systematically underestimates copulation success. We found that this introduces substantial bias for the Bateman gradient, opportunity for sexual selection, and s max ′ . Notably, 47.5% of detected Bateman gradients were significantly positive for females, suggesting selection on females to copulate with multiple males, although the true Bateman gradient was zero. Bias generally increased with the extent of other sources of data limitations tested (nest predation, male infertility, and unsampled floater males). Incomplete offspring sampling introduced bias for all of the metrics except the Bateman gradient, while incomplete sampling of extrapair sires did not introduce additional bias when sires were a random subset of breeding males. Overall, our findings demonstrate how biases due to field data limitations can strongly impact the study of sexual selection. [ABSTRACT FROM AUTHOR]

The Indian subcontinent is the primary wintering ground and stopover site for migratory shorebirds to refuel along the Central Asian and South Asian Flyways. Despite the conservation importance of this region for migratory shorebirds, we lack information on the distribution and abundance of over-summering shorebirds— migrants that remain on their wintering grounds during the breeding season—to evaluate the impacts of anthropogenic change on this poorly understood life history strategy. We conducted weekly surveys of over-summering shorebirds at mudflats, mangroves, and sand beaches at the Kadalundi- Vallikkunnu Community Reserve on the southwest coast of India, 2005–2018. We examined long-term patterns of oversummering shorebird abundance, species composition, and habitat use and estimated proportional changes in oversummering abundance of each shorebird species documented at this globally important site. Over the 14 yr study, we documented 7 over-summering species, including Lesser Sand Plover (Charadrius mongolus), Whimbrel (Numenius phaeopus), Greater Sand Plover (Charadrius leschenaultia), Kentish Plover (Charadrius alexandrinus), Common Sandpiper (Actitis hypoleucos), Ruddy Turnstone (Arenaria interpres), and Pacific Golden-Plover (Pluvialis fulva). Mean abundance and species richness were higher at mudflats than at mangroves and sand beaches, with fewer individuals and species observed over-summering after 2010. Mudflats also had the highest species diversity and evenness compared to mangroves and sand beaches. Management plans that aim to restore vulnerable mudflats and mangroves and to reduce anthropogenic threats such as sand mining and waste dumping are needed to prevent the loss of important oversummering, foraging habitat for migratory shorebirds in southern India and along the Central Asian and South Asian Flyways. [ABSTRACT FROM AUTHOR]

In cooperatively breeding animals, genetic relatedness among group members often determines the extent of reproductive sharing, cooperation and competition within a group. Studies of species for which cooperative behaviour is not entirely based on kinship are key for understanding the benefits favouring the evolution and maintenance of cooperative breeding among nonrelatives. In the cooperatively breeding chestnut-crested yuhina, Yuhina everetti , a songbird endemic to Borneo, we tested whether unrelated helpers are more likely to gain parentage than are related helpers consistent with the hypothesis that inbreeding risk constrains reproduction by related helpers. We also examined whether related or unrelated helpers provision broods more because of differences in their potential indirect or direct fitness benefits of helping. Kin structure of breeding groups (breeding pair and up to eight helpers of both sexes, median = 2 helpers, 96% of 57 pairs had helpers) based on genetic analysis was mixed; 48% of 76 breeder/helper dyads were first-order (26%) or second-order (22%) relatives of one or both members of the breeding pair, and 52% were nonrelatives. Only unrelated male and female helpers gained parentage, and helpers did not differ in their provisioning rate according to their relatedness to the broods. We documented quasi-parasitism or co-breeding by female helpers in 14% of 29 broods and extrapair paternity by male helpers in 21% of 47 broods. This rate of extrapair paternity is relatively high among the few tropical species examined but fit with predictions for mixed-kin groups where inbreeding is avoided. These findings support the emerging pattern for cooperative breeding in birds with mixed-kin groups, wherein unrelated helpers are more likely to gain parentage than are related helpers and helping effort is not necessarily predicted by kinship. • Chestnut-crested yuhina breeding groups contained a mix of kin and nonkin helpers. • Only male and female helpers unrelated to the breeding pair gained parentage. • Female helpers laid eggs in 14% of 29 broods sired by the male breeder. • Male helpers sired extrapair young in 21% of 47 broods laid by the female breeder. • Provisioning rate was not associated with helper–brood genetic relatedness. [ABSTRACT FROM AUTHOR]

Thermal variation poses a problem for nesting birds and can result in reduced offspring growth rates and survival. To increase the thermal stability of the nest, females can adjust nest characteristics and nest attendance in response to changes in environmental conditions. However, it is unclear how and to what extent females modify parental behaviors during various stages of offspring development. We tested the hypothesis that females adjust nest characteristics and brooding patterns in response to thermal variation during the nest-building and nestling stages, respectively. We examined elevational variation in nest location, nest construction, and brooding patterns in the migratory Black-throated Blue Warbler (Setophaga caerulescens) across a 2°C gradient at the Hubbard Brook Experimental Forest, New Hampshire, USA. Density of woody stems at nest sites and nest wall thickness increased from low to high elevation, corresponding to decreasing temperatures, but we found no relationship between weather during nest building and nest characteristics. However, weather during the nestling stage was associated with female brooding patterns: at lower temperatures and with higher rainfall, females spent more time off the nest, which was associated with lower nestling mass near fledging. These results suggest that thermal cues during nest building may be unreliable as predictors of future conditions for developing nestlings and also that females might favor their own self-maintenance and compromise nestling growth under adverse thermal conditions. [ABSTRACT FROM AUTHOR]

Molecular studies have revealed that social groups composed mainly of nonrelatives may be widespread in group-living vertebrates, but the benefits favoring such sociality are not well understood. In the Old World, birds often form conspecific foraging groups that are maintained year-round and offspring usually disperse to other social groups. We tested the hypothesis that nonbreeding group members are largely unrelated and gain direct fitness benefits through breeding opportunities (males) and brood parasitism (females) in the tropical gray-throated babbler, Stachyris nigriceps, in Malaysian Borneo. Babblers foraged in social groups containing one or more breeding pairs (median = 8 group members of equal sex ratio), but group members rarely assisted with breeding (9% of 67 breeding pairs had a third helper; exhibiting facultative cooperative breeding). Although 20% of 266 group member dyads were firstorder relatives of one or both members of the breeding pairs, 80% were unrelated. Male group members gained direct fitness benefits through extrapair and extra-group paternity (25% of 73 offspring), which was independent of their relatedness to the breeding pair and increased with decreasing group size. In contrast, females did not gain direct fitness benefits through brood parasitism. The low levels of relatedness and helping in social groups suggest that most group members do not gain indirect fitness benefits by helping to raise unrelated offspring. These findings highlight the importance of examining benefits of sociality for unrelated individuals that largely do not help and broaden the direct fitness benefits of group foraging beyond assumed survival benefits. [ABSTRACT FROM AUTHOR]

Many studies of sexual selection assume that individuals have equal mating opportunities and that differences in mating success result from variation in sexual traits. However, the inability of sexual traits to explain variation in male mating success suggests that other factors moderate the strength of sexual selection. Extrapair paternity is common in vertebrates and can contribute to variation in mating success and thus serves as a model for understanding the operation of sexual selection. We developed a spatially explicit, multifactor model of all possible female-male pairings to test the hypothesis that ecological (food availability) and social (breeding density, breeding distance, and the social mate's nest stage) factors influence an individual's opportunity for extrapair paternity in a socially monogamous bird, the black-throated blue warbler, Setophaga caerulescens. A male's probability of siring extrapair young decreased with increasing distance to females, breeding density, and food availability. Males on food-poor territories were more likely to sire extrapair young, and these offspring were produced farther from the male's territory relative to males on food-abundant territories. Moreover, males sired extrapair young mostly during their socialmates' incubation stage, especially males on food-abundant territories. This study demonstrates how ecological and social conditions constrain the spatial and temporal opportunities for extrapair paternity that affect variation inmating success and the strength of sexual selection in socially monogamous species. [ABSTRACT FROM AUTHOR]

Single-nucleotide polymorphisms ( SNPs) are preferred over microsatellite markers in many evolutionary studies, but have only recently been applied to studies of parentage. Evaluations of SNPs and microsatellites for assigning parentage have mostly focused on special cases that require a relatively large number of heterozygous loci, such as species with low genetic diversity or with complex social structures. We developed 120 SNP markers from a transcriptome assembled using RNA-sequencing of a songbird with the most common avian mating system-social monogamy. We compared the effectiveness of 97 novel SNPs and six previously described microsatellites for assigning paternity in the black-throated blue warbler, Setophaga caerulescens. We show that the full panel of 97 SNPs (mean Ho = 0.19) was as powerful for assigning paternity as the panel of multiallelic microsatellites (mean Ho = 0.86). Paternity assignments using the two marker types were in agreement for 92% of the offspring. Filtering individual samples by a 50% call rate and SNPs by a 75% call rate maximized the number of offspring assigned with 95% confidence using SNPs. We also found that the 40 most heterozygous SNPs (mean Ho = 0.37) had similar power to assign paternity as the full panel of 97 SNPs. These findings demonstrate that a relatively small number of variable SNPs can be effective for parentage analyses in a socially monogamous species. We suggest that the development of SNP markers is advantageous for studies that require high-throughput genotyping or that plan to address a range of ecological and evolutionary questions. [ABSTRACT FROM AUTHOR]

Background: Although the highest diversity of birds occurs in tropical regions, little is known about the genetic mating systems of most tropical species. We describe microsatellite markers isolated in the chestnut-crested yuhina (Staphida everetti), endemic to the island of Borneo, and the grey-throated babbler (Stachyris nigriceps), widely distributed across Southeast Asia. Both species belong to the avian family Timaliidae and are highly social, putatively cooperatively breeding birds in which helpers attend the nests of members of their social group. We obtained DNA from individuals in social groups breeding in Kinabalu Park, Malaysian Borneo. Results: We used a shotgun sequencing approach and 454-technology to identify 36 microsatellite loci in the yuhina and 40 in the babbler. We tested 13 primer pairs in yuhinas and 20 in babblers and characterized eight polymorphic loci in 20 unrelated female yuhinas and 21 unrelated female babblers. Polymorphism at the yuhina loci ranged from 3 to 9 alleles, observed heterozygosities from 0.58 to 1.00, and expected heterozygosities from 0.64 to 0.81. Polymorphism at the babbler loci ranged from 3 to 12 alleles, observed heterozygosities from 0.14 to 0.90 and expected heterozygosities from 0.14 to 0.87. One locus in the yuhina deviated significantly from Hardy-Weinberg equilibrium. We detected nonrandom allele associations between two pairs of microsatellite loci in each species. Conclusions: Microsatellite markers will be used to describe the genetic mating system of these socially complex species and to measure genetic parentage and relatedness within social groups. [ABSTRACT FROM AUTHOR]

An emerging question in animal behaviour is whether and how behavioural plasticity will enable organisms to adjust to human-induced, rapid environmental changes that affect breeding conditions. Adaptive behavioural plasticity in response to changing resource conditions will depend on the sensitivity of the neuroendocrine system to food stimuli and on constraints in the mechanisms mediating the expression of reproductive behaviours. We tested the hypotheses that food availability mediates plasticity in androgen and corticosterone (CORT) responses, and that circulating concentrations of these hormones in turn mediate investment into competing mating and parental behaviours in a migratory songbird. We provided supplemental food to black-throated blue warblers, Setophaga caerulescens , breeding in habitats with natural low and high food abundance. The effects of supplemental feeding were most pronounced in food-poor habitat. During their social mates' fertile stage, fed males sang less near their nest sites, had lower plasma androgen and CORT levels, and had higher residual mass than control males. Fed males benefited indirectly from their mates' access to supplemental food because fed females provided more parental effort than control females. Fed males did not increase their own parental effort. Our results suggest that fed males increased mate-guarding effort over extrapair mate attraction while their social mates were fertile and invested additional resources into territorial defence or self-maintenance during the parental stage. These findings reveal plastic patterns of covariation among endocrine parameters and mating behaviours, but not parental behaviours, in response to food supplementation. These responses enabled individuals to adjust their reproductive effort rapidly to changes in resource conditions. Our findings suggest that adaptive reproductive investment in response to environmental change is unlikely to be constrained by the hormonal mechanisms underlying the mediation of competing mating and parental behaviours. [ABSTRACT FROM AUTHOR]

Many studies investigating edge effects on forest-nesting birds have focused on nest success. Fewer have examined edge effects on other components of fitness. Nestling growth rates have been positively correlated with food availability, which may differ at edges compared to the forest interior. However, previous work has not examined growth as a function of distance to edge and edge type. We investigated whether edge type, distance to edge, hatching date, brood size, brood parasitism, year, and site influenced growth rates of wings, tarsi, and mass of nestling Wood Thrushes (Hylocichla mustelina) using regression tree analyses. Simultaneously, we examined edge effects on daily nest survival. We conducted our study in southwestern Michigan in 2002 and 2003. We located 175 nests and measured nestlings from 61 nests. Contrary to expectations, tarsal and mass growth rates were more rapid near edges than in the forest interior and tarsal growth rates were more rapid near gradual edge types than near abrupt edge types. Wing chord growth rates were more rapid in the forest interior than at edges. Mass and tarsal growth rates were more rapid later in the breeding season, when brood sizes were smaller. We found no effect of edge type or distance to edge on nest survival. The influence of edge on nestling growth rate but not nest survival suggests nestling growth rates may indicate edge-interior and edge type differences in habitat quality, even when high regional fragmentation levels overwhelm potential edge-interior differences in nest survival. [ABSTRACT FROM AUTHOR]

In socially monogamous species, male reproductive success consists of "within‐pair" offspring produced with their socially paired mate(s), and "extra‐pair" offspring produced with additional females throughout the population. Both reproductive pathways offer distinct opportunities for selection in wild populations, as each is composed of separate components of mate attraction, female fecundity, and paternity allocation. Identifying key sources of variance and covariance among these components is a crucial step toward understanding the reproductive strategies that males use to maximize fitness both annually and over their lifetimes. We use 16 years of complete reproductive data from a population of black‐throated blue warblers (Setophaga caerulescens) to partition variance in male annual and lifetime reproductive success, and thereby identify if the opportunity for selection varies over the lifetimes of individual males and what reproductive strategies likely favor maximum lifetime fitness. The majority of variance in male reproduction was attributable to within‐pair success, but the specific effects of individual components of variance differed between total annual and total lifetime reproductive success. Positive overall lifetime covariance between within‐pair and extra‐pair components indicates that males able to maximize within‐pair success, particularly with double‐brooding females, likely achieve higher overall lifetime fitness via both within‐pair and extra‐pair reproductive pathways. [ABSTRACT FROM AUTHOR]

Extrapair paternity should contribute to sexual selection by increasing the number of potential mates available to each individual. Potential copulation partners are, however, limited by their proximity. Spatial constraints may therefore reduce the impact of extrapair paternity on sexual selection. We tested the effect of spatial constraints on sexual selection by simulating extrapair copulations for 15 species of socially monogamous songbirds with varying rates of extrapair paternity. We compared four metrics of sexual selection between simulated populations without spatial constraints and populations where extrapair copulations were restricted to first- and second-order neighbors. Counter to predictions, sexual selection as measured by the Bateman gradient (the association between the number of copulation partners and offspring produced) increased under spatial constraints. In these conditions, repeated extrapair copulations between the same individuals led to more offspring per copulation partner. In contrast, spatial constraints did somewhat reduce sexual selection—as measured by the opportunity for selection, s' max, and the selection gradient on male quality—when the association between simulated male quality scores and copulation success (e.g., female preferences or male–male competition) was strong. Sexual selection remained strong overall in those populations even under spatial constraints. Spatial constraints did not substantially reduce sexual selection when the association between male quality and copulation success was moderate or weak. Thus, spatial constraints on extrapair copulations are insufficient to explain the absence of strong selection on male traits in many species. [ABSTRACT FROM AUTHOR]

Environmental factors can shape reproductive investment strategies and influence the variance in male mating success. Environmental effects on extrapair paternity have traditionally been ascribed to aspects of the social environment, such as breeding density and synchrony. However, social factors are often confounded with habitat quality and are challenging to disentangle. We used both natural variation in habitat quality and a food supplementation experiment to separate the effects of food availability -- one key aspect of habitat quality -- on extrapair paternity (EPP) and reproductive success in the black-throated blue warbler, Setophaga caerulescens. High natural food availability was associated with higher within-pair paternity (WPP) and fledging two broods late in the breeding season, but lower EPP. Food-supplemented males had higher WPP leading to higher reproductive success relative to controls, and when in low-quality habitat, food-supplemented males were more likely to fledge two broods but less likely to gain EPP. Our results demonstrate that food availability affects trade-offs in reproductive activities. When food constraints are reduced, males invest in WPP at the expense of EPP. These findings imply that environmental change could alter how individuals allocate their resources and affect the selective environment that drives variation in male mating success. [ABSTRACT FROM AUTHOR]

Understanding the demographic drivers of range contractions is important for predicting species' responses to climate change; however, few studies have examined the effects of climate change on survival and recruitment across species' ranges. We show that climate change can drive trailing edge range contractions through the effects on apparent survival, and potentially recruitment, in a migratory songbird. We assessed the demographic drivers of trailing edge range contractions using a long‐term demography dataset for the black‐throated blue warbler (Setophaga caerulescens) collected across elevational climate gradients at the trailing edge and core of the breeding range. We used a Bayesian hierarchical model to estimate the effect of climate change on apparent survival and recruitment and to forecast population viability at study plots through 2040. The trailing edge population at the low‐elevation plot became locally extinct by 2017. The local population at the mid‐elevation plot at the trailing edge gradually declined and is predicted to become extirpated by 2040. Population declines were associated with warming temperatures at the mid‐elevation plot, although results were more equivocal at the low‐elevation plot where we had fewer years of data. Population density was stable or increasing at the range core, although warming temperatures are predicted to cause population declines by 2040 at the low‐elevation plot. This result suggests that even populations within the geographic core of the range are vulnerable to climate change. The demographic drivers of local population declines varied between study plots, but warming temperatures were frequently associated with declining rates of population growth and apparent survival. Declining apparent survival in our study system is likely to be associated with increased adult emigration away from poor‐quality habitats. Our results suggest that demographic responses to warming temperatures are complex and dependent on local conditions and geographic range position, but spatial variation in population declines is consistent with the climate‐mediated range shift hypothesis. Local populations of black‐throated blue warblers near the warm‐edge range boundary at low latitudes and low elevations are likely to be the most vulnerable to climate change, potentially leading to local extirpation and range contractions. [ABSTRACT FROM AUTHOR]

Because elevated glucocorticoid levels can impair reproduction, populations or species that engage in particularly valuable reproductive attempts may down-regulate the glucocorticoid stress response during reproduction (the brood value hypothesis). It is not clear, however, whether individuals rapidly modulate glucocorticoid responses based on shifting cues about the likelihood of reproductive success. By manipulating brood size to create broods that differed in potential value, we tested whether female barn swallows ( Hirundo rustica ) rapidly modulated the glucocorticoid stress response to promote investment in high-value broods, and whether nestling phenotype was influenced by treatment. Within-individual changes in female corticosterone, body mass, and measures of oxidative stress were unrelated to brood size treatment. Standard offspring provisioning rate did not differ across treatments; however, in the presence of a model predator, females raising enlarged broods maintained higher offspring feeding rates relative to control broods. Brood size did influence nestling phenotype. Nestlings from enlarged broods had lower body mass and higher baseline corticosterone than those from reduced broods. Finally, in adult females both baseline and stress-induced corticosterone were individually repeatable. Thus, while under moderately challenging environmental conditions brood size manipulations had context-dependent effects on parental investment, and influenced nestling phenotype, maternal glucocorticoid levels were not modulated based on brood value but were individually consistent features of phenotype during breeding. [ABSTRACT FROM AUTHOR]

Numerous studies have correlated the advancement of lay date in birds with warming climate trends, yet the fitness effects associated with this phenological response have been examined in only a small number of species. Most of these species–primarily insectivorous cavity nesters in Europe–exhibit fitness declines associated with increasing asynchrony with prey. Here, we use 25 years of demographic data, collected from 1986 to 2010, to examine the effects of spring temperature on breeding initiation date, double brooding, and annual fecundity in a Nearctic - Neotropical migratory songbird, the black-throated blue warbler (Setophaga caerulescens). Data were collected from birds breeding at the Hubbard Brook Experimental Forest, New Hampshire, USA, where long-term trends toward warmer springs have been recorded. We found that black-throated blue warblers initiated breeding earlier in warmer springs, that early breeders were more likely to attempt a second brood than those starting later in the season, and that double brooding and lay date were linked to higher annual fecundity. Accordingly, we found selection favored earlier breeding in most years. However, in contrast to studies of several other long-distance migratory species in Europe, this selection pressure was not stronger in warmer springs, indicating that these warblers were able to adjust mean lay date appropriately to substantial inter-annual variation in spring temperature. Our results suggest that this North American migratory songbird might not experience the same fecundity declines as songbirds that are unable to adjust their timing of breeding in pace with spring temperatures. [ABSTRACT FROM AUTHOR]

Threats to a species' persistence are likely to change as conservation measures reduce some threats, while natural and anthropogenic changes increase others. Despite a variety of potential underlying mechanisms, extinction threats will be manifested through one of the 3 components of population dynamics: reducing population growth potential, increasing population variability, or lowering the population ceiling. Consequently, effective management can be guided by monitoring programs and population models that examine each of these components. We examined the potential for a coupled monitoring and modeling effort to guide management of species-at-risk while accounting for evolving risks using the case study of the threatened San Clemente sage sparrow (Amphispiza belli clementeae). Originally listed due to a low population ceiling imposed by severe habitat loss, we found that the major threat to San Clemente sage sparrow persistence has shifted to low population growth potential driven by high juvenile mortality. We further found that successful mitigation of high juvenile mortality will shift the primary threat to drought frequency, which is predicted to increase on San Clemente Island as a consequence of global climate change. The latter shift is a consequence of the boom-bust ecology exhibited by San Clemente sage sparrows in response to rainfall-likely a common characteristic of short-lived terrestrial vertebrates in arid environments. Our ability to successfully recover this species hinges on a comprehensive monitoring and modeling program incorporating all 3 components of population dynamics informing changes in management priorities to reflect shifting threats. Our study indicates that the next critical step to recovering sage sparrows is to understand and mitigate the causes of high juvenile mortality. In response to these predictions, the United States Navy has funded a radio-telemetry study to determine the cause(s) of juvenile mortalities. [ABSTRACT FROM AUTHOR]