Immune gene expression and epigenetic potential affect the consumption of risky food by female house sparrows.

When organisms move into new areas, they are likely to encounter novel food resources. Even if they are nutritious, these foods can also be risky, as they might be contaminated by parasites. The behavioural immune system of animals could help them avoid the negative effects of contaminated resources, but our understanding of behavioural immunity is limited, particularly whether and how behavioural immunity interacts with physiological immunity. Here, we asked about the potential for interplay between these two traits, specifically how the propensity of an individual house sparrow (Passer domesticus) to take foraging risks was related to its ability to regulate a key facet of its immune response to bacterial pathogens. Previously, we found that sparrows at expanding geographic range edges were more exploratory and less risk-averse to novel foods; in those same populations, birds tended to over-express Toll-like receptor 4 (TLR4), a pattern-recognition receptor that distinguishes cell-wall components of Gram-negative bacteria, making it the major sensor of potentially lethal gut microbial infections including salmonellosis. When we investigated how birds would respond to a typical diet (i.e., mixed seeds) spiked with domesticated chicken faeces, birds that expressed more TLR4 or had higher epigenetic potential for TLR4 (more CpG dinucleotides in the putative gene promoter) ate more food, spiked or not. Females expressing abundant TLR4 were also willing to take more foraging risks and ate more spiked food. In males, TLR4 expression was not associated with risk-taking. Altogether, our results indicate that behaviour and immunity covary among individual house sparrows, particularly in females where those birds that maintain more immune surveillance also are more disposed to take foraging risks.

A telemetry study shows that an endangered nocturnal avian species roosts in extremely dry habitats to avoid predation.

Describing animal space use is essential for understanding their ecological needs and for planning effective conservation schemes. Notably, certain biomes and life histories are understudied due to methodological challenges in tracking animals in their natural habitats. Specifically, both arid environments and nocturnal species are not sufficiently researched compared to diurnal species and to other biomes. This knowledge gap hinders our ability to properly prioritize habitats for species protection in areas undergoing human-related development. Here, we investigate the movement ecology of the Egyptian Nightjar (Caprimulgus aegyptius) in the arid Dead-sea region of Israel, the Palestinian Authority (the West Bank) and Jordan. This nocturnal insectivore is a cryptic desert-dweller and was considered locally extinct until it was rediscovered in 2016. For this work we tracked twelve individuals using GPS tags to determine how this resource-poor environment affects their home range, (predicting large areas), habitat use, and day-roost ecology. We found that the tracked Egyptian Nightjars had a much larger home range area than other Nightjar species, commuting nightly between foraging grounds and day-roosts. We found, as expected, intensive foraging activity at agricultural fields, where artificial irrigation likely supports higher resource (insect) density. Additionally, we found that individuals showed very high roosting site fidelity, often returning to the same specific site, located in extremely dry and exposed habitats, presumably for predator avoidance. This finding highlights the ecological value of these barren habitats that are often considered "lifeless" and therefore of lower priority for conservation. Consequently, our research demonstrates the importance of describing the space-use of nocturnal animals in arid habitats for conservation efforts.

The effect of local species composition on the distribution of an avian invader.

Estimating the potential distribution of invasive species has been primarily achieved by employing species distribution models (SDM). Recently introduced joint species distribution models (JSDM) that include species interactions are expected to improve model output. Here we compare the predictive ability of SDM and JSDM by modelling the distribution of one of the most prolific avian invaders in the world, the common myna (Acridotheres tristis), in a recent introduction in Israel. Our results indicate that including information on the local species composition did not improve model accuracy, possibly because of the unique characteristics of this species that include broad environmental tolerance and behavior flexibility. However, the JSDM provided insights into co-occurrence patterns of common mynas and their local heterospecifics, suggesting that at this time point, there is no evidence of species exclusion by common mynas. Our findings suggest that the invasion potential of common mynas depends greatly on urbanization and less so on the local species composition and reflect the major role of anthropogenic impact in increasing the distribution of avian invaders.

Food for Sex in Bats Revealed as Producer Males Reproduce with Scrounging Females.

Food sharing is often evolutionarily puzzling, because the provider's benefits are not always clear. Sharing among kin may increase indirect fitness [1], but when non-kin are involved, different mechanisms were suggested to act. Occasionally, "tolerated theft" [2, 3] is observed, merely because defending a resource is not cost effective. Sharing may also be explained as "costly signaling" [4, 5], where individuals signal their high qualities by distributing acquired resources, as has been suggested to occur in certain human cultures [6]. Alternatively, a transferred food item might be compensated for in later interactions [7]. In vampire bats, blood sharing reflects reciprocity between non-kin colony members [8-10], and long-term social bonds affect food sharing in chimpanzees [11]. Food may also be exchanged for other goods or social benefits [12-14]. One reciprocity-based explanation for intersexual food sharing is the food-for-sex hypothesis [15-17]. This hypothesis proposes that males share food with females in exchange for mating opportunities. Studies on human hunter-gatherer societies suggest that males with increased foraging success have higher reproductive success [18, 19]. Male chimpanzees, which in contrast to humans do not maintain pair bonds, were suggested to share food with females to increase their mating opportunities [16] (but see [20]). Bats, which are long-lived social mammals [21, 22], provide an opportunity to study long-term social reciprocity mechanisms. We monitored producer-scrounger interactions of a captive Egyptian fruit bat (Rousettus aegyptiacus) colony for more than a year and genetically determined the paternity of the pups that were born in the colony. We found that females carry the young of males from which they used to scrounge food, supporting the food-for-sex hypothesis in this species.

Age-dependent survival rate of the colonial Little Tern (Sternula albifrons).

Many ground-nesting bird species are suffering from habitat loss and population decline. Data on population ecology and demography in colonies of threatened species are thus essential for designing effective conservation protocols. Here, we used extensive ringing and observation data to estimate directly, for the first time, the survival rate of juvenile and adult Little Tern (Sternula albifrons), as well as testing for a possible effect of age on probability of survival. We estimated adult annual survival rate to be 0.77, and juvenile (first year) survival to be 0.49 with a possible linear decrease in the survival rate of the juveniles that ranged from 0.681 to 0.327. We found no evidence that survival was age-dependent among the early age classes after the first year. We discuss these findings in light of survival estimates for other species, and their implications for the Little Tern conservation.

Taxonomic revision of Israeli snakes belonging to the Platyceps rhodorachis species complex (Reptilia: Squamata: Colubridae).

The Platyceps rhodorachis species complex encompasses a widespread group of morphologically similar colubrid snakes. The number and identities of species from this complex in Israel have recently been debated. Studies from the previous decade concluded that there are two species in Israel and its vicinity (compared with one previously recognized), but their identity remained contested. We estimated the number of species and their taxonomic identity using morphological and molecular data. We found some evidence for clinal variation in many of the characters used to differentiate the species, and a great overlap in traits of putative species. Genetic data revealed very low sequence divergence, with all putative species being paraphyletic. Platyceps rogersi emerged as genetically closer to Platyceps saharicus rather than to its putative conspecific, P. karelini. The phylogenetic and taxonomic results thus indicate that the Israeli populations of the P. rhodorachis complex all belong to a single species, Platyceps saharicus (Schätti McCarthy 2004).

Persistent producer-scrounger relationships in bats.

Social foraging theory suggests that group-living animals gain from persistent social bonds, which lead to increased tolerance in competitive foraging and information sharing. Bats are among the most social mammals, often living in colonies of tens to thousands of individuals for dozens of years, yet little is known about their social foraging dynamics. We observed three captive bat colonies for over a year, quantifying >13,000 social foraging interactions. We found that individuals consistently used one of two foraging strategies, either producing (collecting) food themselves or scrounging it directly from the mouth of other individuals. Individual foraging types were consistent over at least 16 months except during the lactation period when females shifted toward producing. Scroungers intentionally selected whom to interact with when socially foraging, thus generating persistent nonrandom social relationships with two to three specific producers. These persistent producer-scrounger relationships seem to reduce aggression over time. Finally, scrounging was highly correlated with vigilance, and we hypothesize that vigilant-prone individuals turn to scrounging in the wild to mitigate the risk of landing on a potentially unsafe fruit tree. We find the bat colony to be a rich and dynamic social system, which can serve as a model to study the role that social foraging plays in the evolution of mammalian sociality. Our results highlight the importance of considering individual tendencies when exploring social behavior patterns of group-living animals. These tendencies further emphasize the necessity of studying social networks over time.

Phenotypic divergence despite low genetic differentiation in house sparrow populations.

Studying patterns of phenotypic variation among populations can shed light on the drivers of evolutionary processes. The house sparrow (Passer domesticus) is one of the world's most ubiquitous bird species, as well as a successful invader. We investigated phenotypic variation in house sparrow populations across a climatic gradient and in relation to a possible scenario of an invasion. We measured variation in morphological, coloration, and behavioral traits (exploratory behavior and neophobia) and compared it to the neutral genetic variation. We found that sparrows were larger and darker in northern latitudes, in accordance with Bergmann's and Gloger's biogeographic rules. Morphology and behavior mostly differed between the southernmost populations and the other regions, supporting the possibility of an invasion. Genetic differentiation was low and diversity levels were similar across populations, indicating high gene flow. Nevertheless, the southernmost and northern populations differed genetically to some extent. Furthermore, genetic differentiation (F ST) was lower in comparison to phenotypic variation (P ST), indicating that the phenotypic variation is shaped by directional selection or by phenotypic plasticity. This study expands our knowledge on evolutionary mechanisms and biological invasions.

Mother doesn't always know best: Maternal wormlion choice of oviposition habitat does not match larval habitat choice.

Habitat choice is an important decision that influences animals' fitness. Insect larvae are less mobile than the adults. Consequently, the contribution of the maternal choice of habitat to the survival and development of the offspring is considered to be crucial. According to the "preference-performance hypothesis", ovipositing females are expected to choose habitats that will maximize the performance of their offspring. We tested this hypothesis in wormlions (Diptera: Vermileonidae), which are small sand-dwelling insects that dig pit-traps in sandy patches and ambush small arthropods. Larvae prefer relatively deep and obstacle-free sand, and here we tested the habitat preference of the ovipositing female. In contrast to our expectation, ovipositing females showed no clear preference for either a deep sand or obstacle-free habitat, in contrast to the larval choice. This suboptimal female choice led to smaller pits being constructed later by the larvae, which may reduce prey capture success of the larvae. We offer several explanations for this apparently suboptimal female behavior, related either to maximizing maternal rather than offspring fitness, or to constraints on the female's behavior. Female's ovipositing habitat choice may have weaker negative consequences than expected for the offspring, as larvae can partially correct suboptimal maternal choice.

Crowd vocal learning induces vocal dialects in bats: Playback of conspecifics shapes fundamental frequency usage by pups.

Vocal learning, the substrate of human language acquisition, has rarely been described in other mammals. Often, group-specific vocal dialects in wild populations provide the main evidence for vocal learning. While social learning is often the most plausible explanation for these intergroup differences, it is usually impossible to exclude other driving factors, such as genetic or ecological backgrounds. Here, we show the formation of dialects through social vocal learning in fruit bats under controlled conditions. We raised 3 groups of pups in conditions mimicking their natural roosts. Namely, pups could hear their mothers' vocalizations but were also exposed to a manipulation playback. The vocalizations in the 3 playbacks mainly differed in their fundamental frequency. From the age of approximately 6 months and onwards, the pups demonstrated distinct dialects, where each group was biased towards its playback. We demonstrate the emergence of dialects through social learning in a mammalian model in a tightly controlled environment. Unlike in the extensively studied case of songbirds where specific tutors are imitated, we demonstrate that bats do not only learn their vocalizations directly from their mothers, but that they are actually influenced by the sounds of the entire crowd. This process, which we term "crowd vocal learning," might be relevant to many other social animals such as cetaceans and pinnipeds.

The effect of previous experience on trap construction and movement distance in a pit-building predator.

Wormlion larvae are sit-and-wait predators that construct cone-shaped pits in sandy patches to capture prey. Wormlions select microhabitats that feature favorable conditions for pit construction, in a similar way to other trap-building predators, like spiders and antlions. We investigated whether wormlions exhibit an experience-based behavioral plasticity in their pit construction behavior. In a laboratory experiment, pit sizes and relocation distances were compared between larvae that experienced either a period of unfavorable conditions, i.e., surface obstacles, shallow or coarse sand, or a period of favorable conditions, i.e., clear, deep, and fine sand and were able to construct pits undisturbed. We expected that wormlions experiencing improving conditions would build larger pits than those experiencing deteriorating conditions. In addition, we expected that larvae experiencing unfavorable conditions would be less choosy in their new microhabitat and move over shorter distances. We observed a certain effect of recent experience on the trap-building behavior; however, it was not consistent among treatments. Additionally, we detected a correlation between larval body mass, relocation distance, and pit area. These findings might suggest that past experience does not influence wormlion foraging behavior in a simple manner but that different types of experience induce different behavioral responses.

Expression of multiple sexual signals by fathers and sons in the East-Mediterranean barn swallow: are advertising strategies heritable?

The level of expression of sexually selected traits is generally determined by genes, environment and their interaction. In species that use multiple sexual signals which may be costly to produce, investing in the expression of one sexual signal may limit the expression of the other, favoring the evolution of a strategy for resource allocation among signals. As a result, even when the expression of sexual signals is condition dependent, the relative level of expression of each signal may be heritable. We tested this hypothesis in the East-Mediterranean barn swallow (Hirundo rustica transitiva), in which males have been shown to express two uncorrelated sexual signals: red-brown ventral coloration, and long tail streamers. We show that variation in both signals may partially be explained by age, as well as by paternal origin (genetic father-son regressions), but that the strongest similarity between fathers and sons is the relative allocation towards one trait or the other (relative expression index), rather than the expression of the traits themselves. These results suggest that the expression of one signal is not independent of the other, and that genetic strategies for resource allocation among sexual signals may be selected for during the evolution of multiple sexual signals.

Signatures of natural selection in the mitochondrial genomes of Tachycineta swallows and their implications for latitudinal patterns of the 'pace of life'.

Latitudinal variation in avian life histories can be summarized as a slow-fast continuum, termed the 'pace of life', that encompasses patterns in life span, reproduction, and rates of development among tropical and temperate species. Much of the variation in avian pace of life is tied to differences in rates of long-term metabolic energy expenditure. Given the vital role of the mitochondrion in metabolic processes, studies of variation in the mitochondrial genome may offer opportunities to establish mechanistic links between genetic variation and latitudinal 'pace of life' patterns. Using comparative genomic analyses, we examined complete mitochondrial genome sequences obtained from nine, broadly distributed Tachycineta swallow species to test for signatures of natural selection across the mitogenome within a phylogenetic framework. Our results show that although purifying selection is the dominant selective force acting on the mitochondrial genome in Tachycineta, three mitochondrial genes (ND2, ND5, and CYTB) contain regions that exhibit signatures of diversifying selection. Two of these genes (ND2 and ND5) encode interacting subunits of NADH dehydrogenase, and amino residues that were inferred to be targets of positive selection were disproportionately concentrated in these genes. Moreover, the positively selected sites exhibited a phylogenetic pattern that could be indicative of adaptive divergence between "fast" and "slow" lineages. These results suggest that functional variation in cytochrome b and NADH dehydrogenase could mechanistically contribute to latitudinal 'pace of life' patterns in Tachycineta.

Environmental harshness is positively correlated with intraspecific divergence in mammals and birds.

Life on Earth is conspicuously more diverse in the tropics. Although this intriguing geographical pattern has been linked to many biotic and abiotic factors, their relative importance and potential interactions are still poorly understood. The way in which latitudinal changes in ecological conditions influence evolutionary processes is particularly controversial, as there is evidence for both a positive and a negative latitudinal gradient in speciation rates. Here, we identify and address some methodological issues (how patterns are analysed and how latitude is quantified) that could lead to such conflicting results. To address these issues, we assemble a comprehensive data set of the environmental correlates of latitude (including climate, net primary productivity and habitat heterogeneity) and combine it with biological, historical and molecular data to explore global patterns in recent divergence events (subspeciation). Surprisingly, we find that the harsher conditions that typify temperate habitats (lower primary productivity, decreased rainfall and more variable and unpredictable temperatures) are positively correlated with greater subspecies richness in terrestrial mammals and birds. Thus, our findings indicate that intraspecific divergence is greater in regions with lower biodiversity, a pattern that is robust to both sampling variation and latitudinal biases in taxonomic knowledge. We discuss possible causal mechanisms for the link between environmental harshness and subspecies richness (faster rates of evolution, greater likelihood of range discontinuities and more opportunities for divergence) and conclude that this pattern supports recent indications that latitudinal gradients of diversity are maintained by simultaneously higher potentials for both speciation and extinction in temperate than tropical regions.

Multiple sexual signals and behavioral reproductive isolation in a diverging population.

Sexual trait divergence has been shown to play a role in the evolution of reproductive isolation. While variation in multiple sexual signals is common among closely related species, little is known about the role of these different axes of phenotype variation with respect to the evolution of behavioral reproductive isolation. Here we study a unique population of barn swallows (Hirundo rustica transitiva) that can be distinguished phenotypically from its neighboring populations only on the basis of two features of male plumage: exaggerated expression of both long tail streamers and dark ventral coloration. Using phenotype manipulation experiments, we conducted a paternity study to examine whether both traits are sexually selected. Our results show that an exaggerated form of the local male phenotype (with both tail elongation and color darkening) is favored by local females, whereas males whose phenotypes were manipulated to look like males of neighboring subspecies suffered paternity losses from their social mates. These results confirm the multiple signaling role of the unique tail and color combination in our diverging population and suggest a novel possibility according to which multiple sexual signals may also be used to discriminate among males from nearby populations when prezygotic reproductive isolation is adaptive.

Latitudinal clines: an evolutionary view on biological rhythms.

Properties of the circadian and annual timing systems are expected to vary systematically with latitude on the basis of different annual light and temperature patterns at higher latitudes, creating specific selection pressures. We review literature with respect to latitudinal clines in circadian phenotypes as well as in polymorphisms of circadian clock genes and their possible association with annual timing. The use of latitudinal (and altitudinal) clines in identifying selective forces acting on biological rhythms is discussed, and we evaluate how these studies can reveal novel molecular and physiological components of these rhythms.

Species trees for the tree swallows (Genus Tachycineta): an alternative phylogenetic hypothesis to the mitochondrial gene tree.

The New World swallow genus Tachycineta comprises nine species that collectively have a wide geographic distribution and remarkable variation both within- and among-species in ecologically important traits. Existing phylogenetic hypotheses for Tachycineta are based on mitochondrial DNA sequences, thus they provide estimates of a single gene tree. In this study we sequenced multiple individuals from each species at 16 nuclear intron loci. We used gene concatenated approaches (Bayesian and maximum likelihood) as well as coalescent-based species tree inference to reconstruct phylogenetic relationships of the genus. We examined the concordance and conflict between the nuclear and mitochondrial trees and between concatenated and coalescent-based inferences. Our results provide an alternative phylogenetic hypothesis to the existing mitochondrial DNA estimate of phylogeny. This new hypothesis provides a more accurate framework in which to explore trait evolution and examine the evolution of the mitochondrial genome in this group.

Clock gene variation in Tachycineta swallows.

Many animals use photoperiod cues to synchronize reproduction with environmental conditions and thereby improve their reproductive success. The circadian clock, which creates endogenous behavioral and physiological rhythms typically entrained to photoperiod, is well characterized at the molecular level. Recent work provided evidence for an association between Clock poly-Q length polymorphism and latitude and, within a population, an association with the date of laying and the length of the incubation period. Despite relatively high overall breeding synchrony, the timing of clutch initiation has a large impact on the fitness of swallows in the genus Tachycineta. We compared length polymorphism in the Clock poly-Q region among five populations from five different Tachycineta species that breed across a hemisphere-wide latitudinal gradient (Fig. 1). Clock poly-Q variation was not associated with latitude; however, there was an association between Clock poly-Q allele diversity and the degree of clutch size decline within breeding seasons. We did not find evidence for an association between Clock poly-Q variation and date of clutch initiation in for any of the five Tachycineta species, nor did we found a relationship between incubation duration and Clock genotype. Thus, there is no general association between latitude, breeding phenology, and Clock polymorphism in this clade of closely related birds.Figure 1Photos of Tachycineta swallows that were used in this study: A) T. bicolor from Ithaca, New York, B) T. leucorrhoa from Chascomús, Argentina, C) T. albilinea from Hill Bank, Belize, D) T. meyeni from Puerto Varas, Chile, and E) T. thalassina from Mono Lake, California, Photographers: B: Valentina Ferretti; A, C-E: David Winkler.

Phylogeny of the Tachycineta genus of New World swallows: insights from complete mitochondrial genomes.

The Tachycineta genus of swallows is comprised of nine species that range from Alaska to southern Chile. We sequenced the entire mitochondrial genome of each member of Tachycineta and generated a completely resolved phylogenetic hypothesis for the corresponding mitochondrial gene tree. Our analyses confirm the presence of two sub-clades within Tachycineta that are associated with geography: a North American/Caribbean clade and a South/Central American clade. We found considerable variation among regions of the mitochondrial genome in both substitution rates and the level of information that each region supplied for phylogenetic reconstruction. We found no evidence of positive directional selection within mitochondrial coding regions, but we identified numerous sites under purifying selection. This finding suggests that, despite differences in life history traits and distributions, mitochondrial genes in Tachycineta are predominantly under purifying selection for conserved function.

Sexual signaling: climatic carry-over.

A long term study of warblers in the Himalayas reveals a surprising contrast in the effects of warm springs as opposed to warm summers on a signaling trait, emphasizing the need to consider year-round influences of the environment on morphological variation.