High prevalence of haemosporidian parasites in Eurasian jays.

Avian haemosporidians are vector-borne parasites, infecting a great variety of birds. The order Passeriformes has the highest average infection probability; nevertheless, some common species of Passeriformes have been rather poorly studied. We investigated haemosporidians in one such species, the Eurasian jay Garrulus glandarius (Corvidae), from a forest population in Hesse, Central Germany. All individuals were infected with at least one haemosporidian genus (overall prevalence: 100%). The most common infection pattern was a mixed Haemoproteus and Leucocytozoon infection, whereas no Plasmodium infection was detected. Results on lineage diversity indicate a rather pronounced host-specificity of Haemoproteus and Leucocytozoon lineages infecting birds of the family Corvidae.

Brain surface morphology and ecological and macroevolutionary inferences of avian New World suboscines (Aves, Passeriformes, Tyrannides).

The New World suboscines (Passeriformes and Tyrannides) are one of the biggest endemic vertebrate radiations in South America, including the families Furnariidae and Tyrannidae. Avian brain morphology is a reliable proxy to study their evolution. The aim of this work is to elucidate whether the brains of these families reflect the ecological differences (e.g., feeding behavior) and to clarify macroevolutionary aspects of their neuroanatomy. Our hypotheses are as follows: Brain size is similar between both families and with other Passeriformes; brain morphology in Tyrannides is the result of the pressure of ecological factors; and brain disparity is low since they share ecological traits. Skulls of Furnariidae and Tyrannidae were micro-computed tomography-scanned, and three-dimensional models of the endocast were generated. Regression analyses were performed between brain volume and body mass. Linear and surface measurements were used to build phylomorphospaces and to calculate the amount of phylogenetic signal. Tyrannidae showed a larger brain disparity than Furnariidae, although it is not shaped by phylogeny in the Tyrannides. Furnariidae present enlarged Wulsts (eminentiae sagittales) but smaller optic lobes, while in Tyrannidae, it is the opposite. This could indicate that in Tyrannides there is a trade-off between the size of these two visual-related brain structures.

Adaptive expansion of ERVK solo-LTRs is associated with Passeriformes speciation events.

Endogenous retroviruses (ERVs) are ancient retroviral remnants integrated in host genomes, and commonly deleted through unequal homologous recombination, leaving solitary long terminal repeats (solo-LTRs). This study, analysing the genomes of 362 bird species and their reptilian and mammalian outgroups, reveals an unusually higher level of solo-LTRs formation in birds, indicating evolutionary forces might have purged ERVs during evolution. Strikingly in the order Passeriformes, and especially the parvorder Passerida, endogenous retrovirus K (ERVK) solo-LTRs showed bursts of formation and recurrent accumulations coinciding with speciation events over past 22 million years. Moreover, our results indicate that the ongoing expansion of ERVK solo-LTRs in these bird species, marked by high transcriptional activity of ERVK retroviral genes in reproductive organs, caused variation of solo-LTRs between individual zebra finches. We experimentally demonstrated that cis-regulatory activity of recently evolved ERVK solo-LTRs may significantly increase the expression level of ITGA2 in the brain of zebra finches compared to chickens. These findings suggest that ERVK solo-LTRs expansion may introduce novel genomic sequences acting as cis-regulatory elements and contribute to adaptive evolution. Overall, our results underscore that the residual sequences of ancient retroviruses could influence the adaptive diversification of species by regulating host gene expression.

White-crested elaenias (Elaenia albiceps chilensis) breeding across Patagonia exhibit similar spatial and temporal movement patterns throughout the year.

For migratory birds, events happening during any period of their annual cycle can have strong carry-over effects on the subsequent periods. The strength of carry-over effects between non-breeding and breeding grounds can be shaped by the degree of migratory connectivity: whether or not individuals that breed together also migrate and/or spend the non-breeding season together. We assessed the annual cycle of the White-crested Elaenia (Elaenia albiceps chilensis), the longest-distance migrant flycatcher within South America, which breeds in Patagonia and spends the non-breeding season as far north as Amazonia. Using light-level geolocators, we tracked the annual movements of elaenias breeding on southern Patagonia and compared it with movements of elaenias breeding in northern Patagonia (1,365 km north) using Movebank Repository data. We found that elaenias breeding in southern Patagonia successively used two separate non-breeding regions while in their Brazilian non-breeding grounds, as already found for elaenias breeding in the northern Patagonia site. Elaenias breeding in both northern and southern Patagonia also showed high spread in their non-breeding grounds, high non-breeding overlap among individuals from both breeding sites, and similar migration phenology, all of which suggests weak migratory connectivity for this species. Elucidating the annual cycle of this species, with particular emphasis on females and juveniles, still requires further research across a wide expanse of South America. This information will be critical to understanding and possibly predicting this species' response to climate change and rapid land-use changes.

Association between the gut microbiome and carotenoid plumage phenotype in an avian hybrid zone.

Vertebrates host complex microbiomes that impact their physiology. In many taxa, including colourful wood-warblers, gut microbiome similarity decreases with evolutionary distance. This may suggest that as host populations diverge, so do their microbiomes, because of either tight coevolutionary dynamics, or differential environmental influences, or both. Hybridization is common in wood-warblers, but the effects of evolutionary divergence on the microbiome during secondary contact are unclear. Here, we analyse gut microbiomes in two geographically disjunct hybrid zones between blue-winged warblers (Vermivora cyanoptera) and golden-winged warblers (Vermivora chrysoptera). We performed 16S faecal metabarcoding to identify species-specific bacteria and test the hypothesis that host admixture is associated with gut microbiome disruption. Species identity explained a small amount of variation between microbiomes in only one hybrid zone. Co-occurrence of species-specific bacteria was rare for admixed individuals, yet microbiome richness was similar among admixed and parental individuals. Unexpectedly, we found several bacteria that were more abundant among admixed individuals with a broader deposition of carotenoid-based plumage pigments. These bacteria are predicted to encode carotenoid biosynthesis genes, suggesting birds may take advantage of pigments produced by their gut microbiomes. Thus, host admixture may facilitate beneficial symbiotic interactions which contribute to plumage ornaments that function in sexual selection.

Social restructuring during harsh environmental conditions promotes cooperative behaviour in a songbird.

Cooperation may emerge from intrinsic factors such as social structure and extrinsic factors such as environmental conditions. Although these factors might reinforce or counteract each other, their interaction remains unexplored in animal populations. Studies on multilevel societies suggest a link between social structure, environmental conditions and individual investment in cooperative behaviours. These societies exhibit flexible social configurations, with stable groups that overlap and associate hierarchically. Structure can be seasonal, with upper-level units appearing only during specific seasons, and lower-level units persisting year-round. This offers an opportunity to investigate how cooperation relates to social structure and environmental conditions. Here, we study the seasonal multilevel society of superb fairy-wrens (Malurus cyaneus), observing individual responses to experimental playback of conspecific distress calls. Individuals engaged more in helping behaviour and less in aggressive/territorial song during the harsher non-breeding season compared to the breeding season. The increase in cooperation was greater for breeding group members than for members of the same community, the upper social unit, comprised of distinct breeding groups in association. Results suggest that the interaction between social structure and environmental conditions drives the seasonal switch in cooperation, supporting the hypothesis that multilevel societies can emerge to increase cooperation during harsh environmental conditions.

Higher body condition with infection by Haemoproteus parasites in Bananaquits (Coereba flaveola).

Parasite transmission is a heterogenous process in host-parasite interactions. This heterogeneity is particularly apparent in vector-borne parasite transmission where the vector adds an additional level of complexity. Haemosporidian parasites, a widespread protist, cause a malaria-like disease in birds globally, but we still have much to learn about the consequences of infection to hosts' health. In the Caribbean, where malarial parasites are endemic, studying host-parasites interactions may give us important insights about energetic trade-offs involved in malarial parasites infections in birds. In this study, we tested the consequences of Haemoproteus infection on the Bananaquit, a resident species of Puerto Rico. We also tested for potential sources of individual heterogeneity in the consequences of infection such as host age and sex. To quantify the consequences of infection to hosts' health we compared three complementary body condition indices between infected and uninfected individuals. Our results showed that Bananaquits infected by Haemoproteus had higher body condition than uninfected individuals. This result was consistent among the three body condition indices. Still, we found no clear evidence that this effect was mediated by host age or sex. We discuss a set of non-mutually exclusive hypotheses that may explain this pattern including metabolic syndrome, immunological responses leading to host tolerance or resistance to infection, and potential changes in consumption rates. Overall, our results suggest that other mechanisms, may drive the consequences of avian malarial infection.

Risk-sensitive learning is a winning strategy for leading an urban invasion.

In the unpredictable Anthropocene, a particularly pressing open question is how certain species invade urban environments. Sex-biased dispersal and learning arguably influence movement ecology, but their joint influence remains unexplored empirically, and might vary by space and time. We assayed reinforcement learning in wild-caught, temporarily captive core-, middle-, or edge-range great-tailed grackles-a bird species undergoing urban-tracking rapid range expansion, led by dispersing males. We show, across populations, both sexes initially perform similarly when learning stimulus-reward pairings, but, when reward contingencies reverse, male-versus female-grackles finish 'relearning' faster, making fewer choice-option switches. How do male grackles do this? Bayesian cognitive modelling revealed male grackles' choice behaviour is governed more strongly by the 'weight' of relative differences in recent foraging payoffs-i.e., they show more pronounced risk-sensitive learning. Confirming this mechanism, agent-based forward simulations of reinforcement learning-where we simulate 'birds' based on empirical estimates of our grackles' reinforcement learning-replicate our sex-difference behavioural data. Finally, evolutionary modelling revealed natural selection should favour risk-sensitive learning in hypothesised urban-like environments: stable but stochastic settings. Together, these results imply risk-sensitive learning is a winning strategy for urban-invasion leaders, underscoring the potential for life history and cognition to shape invasion success in human-modified environments.

Responses in the breeding parameters of the collared flycatcher to the changing climate.

Global climate change involves various aspects of climate, including precipitation changes and declining surface wind speeds, but studies investigating biological responses have often focused on the impacts of rising temperatures. Additionally, related long-term studies on bird reproduction tend to concentrate on breeding onset, even though other aspects of breeding could also be sensitive to the diverse weather aspects. This study aimed to explore how multiple aspects of breeding (breeding onset, hatching delay, breeding season length, clutch size, fledgling number) were associated with different weather components. We used an almost four-decade-long dataset to investigate the various aspects of breeding parameters of a collared flycatcher (Ficedula albicollis) population in the Carpathian Basin. Analyses revealed some considerable associations, for example, breeding seasons lengthened with the amount of daily precipitation, and clutch size increased with the number of cool days. Parallel and opposing changes in the correlated pairs of breeding and weather parameters were also observed. The phenological mismatch between prey availability and breeding time slightly increased, and fledgling number strongly decreased with increasing mistiming. Our results highlighted the intricate interplay between climate change and the reproductive patterns of migratory birds, emphasizing the need for a holistic approach. The results also underscored the potential threats posed by climate change to bird populations and the importance of adaptive responses to changing environmental conditions.

Taxonomic revision of the Scaled Antbird Drymophila squamata (Aves: Thamnophilidae) reveals a new and critically endangered taxon from northeastern Brazil.

The Scaled Antbird Drymophila squamata is a suboscine passerine endemic to the Atlantic Forest of eastern Brazil. Two subspecies, putatively diagnosed by the presence/absence of white spots on the crown, have traditionally been recognized: the nominate, ranging from Pernambuco to Bahia in northeastern Brazil, and D. squamata stictocorypha, from Minas Gerais to Santa Catarina in southeastern and southern Brazil. Here we combine morphological, acoustic, and genetic data to examine geographic variation in and revise the taxonomy of D. squamata. We show that there are two separately evolving population lineages in D. squamata, one south and the other north of the So Francisco River. The latter is unnamed and is thus described herein. We found that crown variation is not as geographically structured as previously thought, and thus we suggest that D. squamata stictocorypha is not a valid taxon. Finally, we also provide evidence of clinal variation in the species vocalizations and underscore the importance of broad geographic sampling when assessing species limits using vocalizations.

Environmental harshness does not affect the propensity for social learning in great tits, Parus major.

According to the harsh environment hypothesis, natural selection should favour cognitive mechanisms to overcome environmental challenges. Tests of this hypothesis to date have largely focused on asocial learning and memory, thus failing to account for the spread of information via social means. Tests in specialized food-hoarding birds have shown strong support for the effects of environmental harshness on both asocial and social learning. Whether the hypothesis applies to non-specialist foraging species remains largely unexplored. We evaluated the relative importance of social learning across a known harshness gradient by testing generalist great tits, Parus major, from high (harsh)- and low (mild)-elevation populations in two social learning tasks. We showed that individuals use social learning to find food in both colour-associative and spatial foraging tasks and that individuals differed consistently in their use of social learning. However, we did not detect a difference in the use or speed of implementing socially observed information across the elevational gradient. Our results do not support predictions of the harsh environment hypothesis suggesting that context-dependent costs and benefits as well as plasticity in the use of social information may play an important role in the use of social learning across environments. Finally, this study adds to the accumulating evidence that the harsh environment hypothesis appears to have more pronounced effects on specialists compared to generalist species.

Investigations of the tick burden on passeriform, water-associated and predatory birds reveal new tick-host associations and habitat-related factors of tick infestation.

BACKGROUND: Previous studies on the tick infestation of birds in the Carpathian Basin focused on songbirds (Passeriformes). Thus, the primary aim of the present work was to extend the scope of previous studies, i.e. to include aquatic (water-associated) bird species in a similar context, especially considering that these birds are usually long-distance migrants. METHODS: Between March 2021 and August 2023, 11,919 birds representing 126 species were checked for the presence of ticks. From 352 birds belonging to 40 species, 905 ixodid ticks were collected. Tick species were identified morphologically and/or molecularly. RESULTS: Ticks from avian hosts belonged to seven species: Ixodes ricinus (n = 448), I. frontalis (n = 31), I. festai (n = 2), I. arboricola (n = 36), I. lividus (n = 4), Haemaphysalis concinna (n = 382) and Dermacentor reticulatus (n = 2). Nymphs of I. ricinus occurred with a single activity peak around March-May, whereas its larvae typically infested birds in May, June or July. By contrast, H. concinna usually had its activity maximum during the summer (nymphs in June-July, larvae later in July-August). Interestingly, two ornithophilic species, I. frontalis and I. arboricola, were most active around winter months (between October and April). A significantly lower ratio of aquatic birds was found tick-infested than songbirds. Several new tick-host associations were revealed, including I. ricinus from Greylag Goose (Anser anser) and D. reticulatus from Great Egret (Ardea alba) and Sedge Warbler (Acrocephalus schoenobaenus). Ticks were collected for the first time in Europe from two species of predatory birds as well as from Little Bittern (Ixobrychus minutus). Bird species typically inhabiting reedbeds were most frequently infested with H. concinna, and most ticks localized at their throat, as opposed to forest-dwelling avian hosts, on which I. ricinus predominated and ticks were more evenly distributed. CONCLUSIONS: In the evaluated region, aquatic birds appear to be less important in tick dispersal than songbirds. However, newly revealed tick-host associations in this category attest to their hitherto neglected contribution. The results suggest that the habitat type will have significant impact not only on the species composition but also on the feeding location of ticks on birds.

Dorsal and Ventral Plumage Coloration Evolve as Distinct Modules with Different Environmental Correlations.

AbstractMany animals exhibit contrast between their dorsal coloration and their ventral coloration. If selection acts differently on dorsal versus ventral coloration, ancestral covariance between these traits should break down, eventually leading to independent modules of trait evolution. Here, we compare the evolution of feather color across body regions for a clade of Australasian songbirds (Meliphagoidea). We find evidence for three modules of covarying color regions. Among these modules, ventral feathers evolve with high lability, evolving at three times the rate of dorsal plumage and 20 times the rate of flight feathers. While both dorsal plumage and ventral plumage are darker in areas with more precipitation and vegetation, we find that dorsal plumage is twice as similar to colors in satellite photos of background substrates. Overall, differential selection on ventral and dorsal colors likely maintains these as distinct modules over evolutionary timescales-a novel explanation for dorsoventral contrast in pigmentation.

A RETROSPECTIVE EVALUATION OF MORBIDITY AND MORTALITY IN ASSOCIATION WITH INFESTATION BY THE MITE STERNOSTOMA TRACHEACOLUM IN AN AVIARY-HOUSED POPULATION OF GOULDIAN FINCHES (CHLOEBIA GOULDIAE).

The Rhynonyssid mesostigmatic mite, Sternostoma tracheacolum, is a well-documented endoparasitic hematophagous arthropod of the respiratory tracts of multiple avian species, particularly Estrildid finches and canaries. In this retrospective study, 175 medical and 278 pathology records for the Gouldian finch (Chloebia gouldiae) population (N = 377) at the San Diego Zoo between 2013 and 2021 were analyzed to evaluate the effectiveness of ivermectin-based prophylaxis. A multivariable negative binomial regression model was constructed to evaluate the population effects of monthly treatments on morbidity or mortality associated with respiratory mites. While controlling for other factors in the model, the prophylactic treatment did not significantly reduce the monthly rate of mite-associated morbidity or mortality (IRR = 1.017, 95% CI: 0.997-1.036, P = 0.0759); however, low proportions of the population were prophylactically treated over time. Different factors were significant when separately evaluating adjusted associations with respiratory morbidity and mortality. The findings suggest increased rates of respiratory morbidity for each successive year of the study period (IRR = 1.180, 95% CI: 1.046-1.342, P = 0.0090) and increased rates of mite-associated mortality occurring annually between May and October (IRR = 1.697, 95% CI: 1.034-2.855, P = 0.0404) compared to the wet winter season. Our findings highlight the need to continually evaluate and optimize treatment regimens in zoological collections. Further investigations into this host-parasite relationship and potential treatments and preventive therapies are warranted.

Auditory risk recognition is socially transmitted across territory borders in wild birds.

Prey species commonly assess predation risk based on acoustic signals, such as predator vocalizations or heterospecific alarm calls. The resulting risk-sensitive decision-making affects not only the behavior and life-history of individual prey, but also has far-reaching ecological consequences for population, community, and ecosystem dynamics. Although auditory risk recognition is ubiquitous in animals, it remains unclear how individuals gain the ability to recognize specific sounds as cues of a threat. Here, it has been shown that free-living birds (Wood Warblers Phylloscopus sibilatrix) can learn to recognize unfamiliar, complex sounds (samples of punk rock songs) as cues of a threat from conspecifics holding adjacent territories during the spring breeding season. In a playback experiment, Wood Warblers initially ignored the unfamiliar sounds, but after repeatedly hearing that these sounds trigger alarm calling reaction of neighbors, most individuals showed an anti-predator response to them. Moreover, once learned soon after nestlings hatching, the anti-predator response of parents toward previously unfamiliar sounds was then retained over the entire nestlings rearing period. These results demonstrate that social learning via the association of unfamiliar sounds with known alarm signals enables the spread of anti-predator behavior across territory borders and provides a mechanism explaining the widespread abilities of animals to assess predation risk based on acoustic cues.

Genotype-environment associations reveal genes potentially linked to avian malaria infection in populations of an endemic island bird.

Patterns of pathogen prevalence are, at least partially, the result of coevolutionary host-pathogen interactions. Thus, exploring the distribution of host genetic variation in relation to infection by a pathogen within and across populations can provide important insights into mechanisms of host defence and adaptation. Here, we use a landscape genomics approach (Bayenv) in conjunction with genome-wide data (ddRADseq) to test for associations between avian malaria (Plasmodium) prevalence and host genetic variation across 13 populations of the island endemic Berthelot's pipit (Anthus berthelotii). Considerable and consistent spatial heterogeneity in malaria prevalence was observed among populations over a period of 15 years. The prevalence of malaria infection was also strongly positively correlated with pox (Avipoxvirus) prevalence. Multiple host loci showed significant associations with malaria prevalence after controlling for genome-wide neutral genetic structure. These sites were located near to or within genes linked to metabolism, stress response, transcriptional regulation, complement activity and the inflammatory response, many previously implicated in vertebrate responses to malarial infection. Our findings identify diverse genes - not just limited to the immune system - that may be involved in host protection against malaria and suggest that spatially variable pathogen pressure may be an important evolutionary driver of genetic divergence among wild animal populations, such as Berthelot's pipit. Furthermore, our data indicate that spatio-temporal variation in multiple different pathogens (e.g. malaria and pox in this case) may have to be studied together to develop a more holistic understanding of host pathogen-mediated evolution.

Odours of caterpillar-infested trees increase testosterone concentrations in male great tits.

Trees release Herbivore-Induced Plant Volatiles (HIPVs) into the air in response to damage inflicted by insects. It is known that songbirds use those compounds to locate their prey, but more recently the idea emerged that songbirds could also use those odours as cues in their reproductive decisions, as early spring HIPVs may contain information about the seasonal timing and abundance of insects. We exposed pre-breeding great tits (Parus major) to the odours of caterpillar-infested trees under controlled conditions, and monitored reproduction (timing of egg laying, number of eggs, egg size) and two of its main hormonal drivers (testosterone and 17ß-estradiol in males and females, respectively). We found that females exposed to HIPVs did not advance their laying dates, nor laid larger clutches, or larger eggs compared to control females. 17ß-estradiol concentrations in females were also similar between experimental and control birds. However, males exposed to HIPVs had higher testosterone concentrations during the egg-laying period. Our study supports the hypothesis that insectivorous songbirds are able to detect minute amounts of plant odours. The sole manipulation of plant scents was not sufficient to lure females into a higher reproductive investment, but males increased their reproductive effort in response to a novel source of information for seasonal breeding birds.

Nightly reductions in body temperature and effect of transmitter attachment method in free-living welcome swallows (Hirundoneoxena).

Small birds and mammals face similar energetic challenges, yet use of torpor to conserve energy while resting is considered less common among birds, especially within the most specious order Passeriformes. We conducted the first study to record the natural thermoregulatory physiology of any species from the family Hirundinidae, which we predicted would use torpor because of their specialised foraging by aerial pursuit of flying insects, that are less active during cold or windy weather. We used temperature telemetry on wild-living welcome swallows (Hirundo neoxena, 13 to 17 g) and found that skin temperature declined during nightly resting by an average by 5 °C, from daytime minima of 41.0 ± 0.8 °C to nightly minima of 36.3 ± 0.8 °C, and by a maximum of 8 °C to a minimum recorded skin temperature of 32.0 °C. The extent of reduction in skin temperature was greater on cold nights and following windy daytime (foraging) periods. Further, we found that transmitters glued directly to the skin between feather tracts (i.e., an apterium) provided a less variable and probably also more accurate reflection of body temperature than transmitters applied over closely trimmed feathers. A moderate decrease in skin temperature, equivalent to shallow torpor, would provide energy savings during rest. Yet, deeper torpor was not observed, despite a period of extreme rainfall that presumedly decreased foraging success. Further studies are needed to understand the resting thermoregulatory energetics of swallows under different environmental conditions. We advocate the importance of measuring thermal biology in wild-living birds to increase our knowledge of the physiology and ecological importance of torpor among passerine birds.