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The species complex comprising the rufous-naped lark Corypha africana , Sharpe's lark Corypha sharpii , the red-winged lark Corypha hypermetra , the Somali long-billed lark Corypha somalica and Ash's lark Corypha ashi encompasses 31 recognised taxa across sub-Saharan Africa, many of which are extremely poorly known and some not observed for decades. Only 17 taxa have been studied molecularly and none comprehensively for morphology, vocalisations or other behaviours. Here, we undertake comprehensive integrative taxonomic analyses based on plumage and morphometrics (for 97% of the taxa), mitochondrial and nuclear loci (77%), ≤ 1.3 million genome-wide single nucleotide polymorphisms (68%), song (many described for the first time; 52%) and additional behavioural data (45%). All polytypic species as presently circumscribed are paraphyletic, with eight primary clades separated by ≤ 6.3–6.8 Myr, broadly supported by plumage, morphometrics, song and other behaviours. The most recent divergences concern sympatric taxon pairs usually treated as separate species, whereas the divergence of all clades including C. africana subspecies is as old as sister species pairs in other lark genera. We propose the recognition of nine instead of five species, while C. ashi is synonymised with C. somalica rochei as C. s. ashi. The geographical distributions are incompletely known, and although the nine species are generally para-/allopatric, some might be sympatric. [ABSTRACT FROM AUTHOR]

Background: In contrast with macroorganisms, that show well-documented biogeographical patterns in distribution associated with local adaptation of physiology, behavior and life history, strong biogeographical patterns have not been found for microorganisms, raising questions about what determines their biogeography. Thus far, large-scale biogeographical studies have focused on free-living microbes, paying little attention to host-associated microbes, which play essential roles in physiology, behavior and life history of their hosts. Investigating cloacal gut microbiota of closely-related, ecologically similar free-living songbird species (Alaudidae, larks) inhabiting desert, temperate and tropical regions, we explored influences of geographical location and host species on α-diversity, co-occurrence of amplicon sequence variants (ASVs) and genera, differentially abundant and dominant bacterial taxa, and community composition. We also investigated how geographical distance explained differences in gut microbial community composition among larks. Results: Geographic location did not explain variation in richness and Shannon diversity of cloacal microbiota in larks. Out of 3798 ASVs and 799 bacterial genera identified, 17 ASVs (< 0.5%) and 43 genera (5%) were shared by larks from all locations. Desert larks held fewer unique ASVs (25%) than temperate zone (31%) and tropical larks (34%). Five out of 33 detected bacterial phyla dominated lark cloacal gut microbiomes. In tropical larks three bacterial classes were overrepresented. Highlighting the distinctiveness of desert lark microbiota, the relative abundances of 52 ASVs differed among locations, which classified within three dominant and 11 low-abundance phyla. Clear and significant phylogenetic clustering in cloacal microbiota community composition (unweighted UniFrac) showed segregation with geography and host species, where microbiota of desert larks were distinct from those of tropical and temperate regions. Geographic distance was nonlinearly associated with pairwise unweighted UniFrac distances. Conclusions: We conclude that host-associated microbiota are geographically structured in a group of widespread but closely-related bird species, following large-scale macro-ecological patterns and contrasting with previous findings for free-living microbes. Future work should further explore if and to what extent geographic variation in host-associated microbiota can be explained as result of co-evolution between gut microbes and host adaptive traits, and if and how acquisition from the environmental pool of bacteria contributes to explaining host-associated communities. [ABSTRACT FROM AUTHOR]

Background: Gut microbiotas play a pivotal role in host physiology and behaviour, and may affect host life-history traits such as seasonal variation in host phenotypic state. Generally, seasonal gut microbiota variation is attributed to seasonal diet variation. However, seasonal temperature and day length variation may also drive gut microbiota variation. We investigated summer–winter differences in the gut bacterial community (GBC) in 14 homing pigeons living outdoors under a constant diet by collecting cloacal swabs in both seasons during two years. Because temperature effects may be mediated by host metabolism, we determined basal metabolic rate (BMR) and body mass. Immune competence is influenced by day length and has a close relationship with the GBC, and it may thus be a link between day length and gut microbiota. Therefore, we measured seven innate immune indices. We expected the GBC to show summer–winter differences and to correlate with metabolism and immune indices. Results: BMR, body mass, and two immune indices varied seasonally, other host factors did not. The GBC showed differences between seasons and sexes, and correlated with metabolism and immune indices. The most abundant genus (Lachnoclostridium 12, 12%) and associated higher taxa, were more abundant in winter, though not significantly at the phylum level, Firmicutes. Bacteroidetes were more abundant in summer. The Firmicutes:Bacteroidetes ratio tended to be higher in winter. The KEGG ortholog functions for fatty acid biosynthesis and linoleic acid metabolism (PICRUSt2) had increased abundances in winter. Conclusions: The GBC of homing pigeons varied seasonally, even under a constant diet. The correlations between immune indices and the GBC did not involve consistently specific immune indices and included only one of the two immune indices that showed seasonal differences, suggesting that immune competence may be an unlikely link between day length and the GBC. The correlations between the GBC and metabolism indices, the higher Firmicutes:Bacteroidetes ratio in winter, and the resemblance of the summer–winter differences in the GBC with the general temperature effects on the GBC in the literature, suggest that temperature partly drove the summer–winter differences in the GBC in homing pigeons. [ABSTRACT FROM AUTHOR]

The innate immune system is essential for survival, yet many immune traits are highly variable between and within individuals. In recent years, attention has shifted to the role of environmental factors in modulating this variation. A key environmental factor is food availability, which plays a major role in shaping life histories, and may affect resource allocation to immune function through its effect on nutritional state.We developed a technique to permanently increase foraging costs in seed‐eating birds, and leveraged this technique to study the effects of food availability on the innate immune system over a 3‐year period in 230 zebra finches housed in outdoor aviaries. The immune components we studied were haptoglobin, ovotransferrin, nitric oxide, natural antibodies through agglutination, complement‐mediated lysis, and killing capacity of Escherichia coli and Candida albicans, covering a broad spectrum of the innate immune system. We explored the effects of food availability in conjunction with other potentially important variables: season, age, sex and manipulated natal brood size.Increased foraging costs affected multiple components of the immune system, albeit in a variable way. Nitric oxide and agglutination levels were lower under harsh foraging conditions, while Escherichia coli killing capacity was increased. Agglutination levels also varied seasonally, but only at low foraging costs. C. albicans killing capacity was lower in winter, and even more so for animals in harsh foraging conditions that were raised in large broods. Effects of food availability on ovotransferrin were also seasonal, and only apparent in males. Haptoglobin levels were independent of foraging costs and season.Males had higher levels of immune function than females for three of the measured immune traits. Innate immune function was independent of age and manipulated natal brood size.Our finding that food availability affects innate immune function suggests that fitness effects of food availability may at least partially be mediated by effects on the immune system. However, food availability effects on innate immunity varied in direction between traits, illustrating the complexity of the immune system and precluding conclusions on the level of disease resistance. [ABSTRACT FROM AUTHOR]

Background: In a diverse microbial world immune function of animals is essential. Diverse microbial environments may contribute to extensive variation in immunological phenotypes of vertebrates, among and within species and individuals. As maternal effects benefit offspring development and survival, whether females use cues about their microbial environment to prime offspring immune function is unclear. To provide microbial environmental context to maternal effects, we asked if the bacterial diversity of the living environment of female zebra finches Taeniopygia guttata shapes maternal effects on egg immune function. We manipulated environmental bacterial diversity of birds and tested if females increased immunological investment in eggs in an environment with high bacterial diversity (untreated soil) versus low (gamma-sterilized soil). We quantified lysozyme and ovotransferrin in egg albumen and IgY in egg yolk and in female blood, and we used 16S rRNA gene sequencing to profile maternal cloacal and eggshell microbiotas. Results: We found a maternal effect on egg IgY concentration that reflected environmental microbial diversity: females who experienced high diversity deposited more IgY in their eggs, but only if maternal plasma IgY levels were relatively high. We found no effects on lysozyme and ovotransferrin concentrations in albumen. Moreover, we uncovered that variation in egg immune traits could be significantly attributed to differences among females: for IgY concentration in yolk repeatability R = 0.80; for lysozyme concentration in albumen R = 0.27. Furthermore, a partial least squares path model (PLS-PM) linking immune parameters of females and eggs, which included maternal and eggshell microbiota structures and female body condition, recapitulated the treatment-dependent yolk IgY response. The PLS-PM additionally suggested that the microbiota and physical condition of females contributed to shaping maternal effects on egg immune function, and that (non-specific) innate egg immunity was prioritized in the environment with low bacterial diversity. Conclusions: The microbial environment of birds can shape maternal effects on egg immune function. Since immunological priming of eggs benefits offspring, we highlight that non-genetic maternal effects on yolk IgY levels based on cues from the parental microbial environment may prove important for offspring to thrive in the microbial environment that they are expected to face. [ABSTRACT FROM AUTHOR]

Seasonal variation in immune function can be attributed to life history trade-offs, and to variation in environmental conditions. However, because phenological stages and environmental conditions co-vary in temperate and arctic zones, their separate contributions have not been determined. We compared immune function and body mass of incubating (female only), chick-feeding (female and male), and non-breeding (female and male) red-capped larks Calandrella cinerea breeding year-round in three tropical equatorial (Kenya) environments with distinct climates. We measured four immune indices: haptoglobin, nitric oxide, agglutination, and lysis. To confirm that variation in immune function between breeding (i.e., incubating or chick-feeding) and non-breeding was not confounded by environmental conditions, we tested if rainfall, average minimum temperature (Tmin), and average maximum temperature (Tmax) differed during sampling times among the three breeding statuses per location. Tmin and Tmax differed between chick-feeding and non-breeding, suggesting that birds utilized environmental conditions differently in different locations for reproduction. Immune indices did not differ between incubating, chick-feeding and non-breeding birds in all three locations. There were two exceptions: nitric oxide was higher during incubation in cool and wet South Kinangop, and it was higher during chick-feeding in the cool and dry North Kinangop compared to non-breeding birds in these locations. For nitric oxide, agglutination, and lysis, we found among-location differences within breeding stage. In equatorial tropical birds, variation in immune function seems to be better explained by among-location climate-induced environmental conditions than by breeding status. Our findings raise questions about how within-location environmental variation relates to and affects immune function. [ABSTRACT FROM AUTHOR]

Life history theory suggests a trade-off between costly activities such as breeding and migration and somatic self-maintenance. However, how the short-term cost of parental effort is expressed in species with a slow pace-of-life is not well understood. Also, investigating carry-over effects of migration is most meaningful when comparing migratory strategies within the same population, but this has rarely been done. We explore this hypothesis in a long-lived, pelagic seabird, the Cory's Shearwater, Calonectris borealis, where males display partial migration. By manipulating reproductive effort and taking advantage of the natural variation in migratory strategy, we investigate whether early reproductive failure and migratory strategy had implications on the physical condition of males on return to the colony the following year. We experimentally induced breeding failure from mid-incubation, tracked the over-winter movements of these males and of males that invested in parental effort, and assessed innate immunity, stress, and residual body mass the following year. Early breeding failure resulted in earlier return to the colony among all males, associated with greater probability of reproductive success. Residents had a lower tail feather fault bar intensity, an indicator of stress during the non-breeding period, compared to migrants. Reproductive effort and migratory strategy had no impact on physiological condition otherwise. Our results provide evidence that in species with a slow-pace of life, such as the Cory's Shearwater, somatic maintenance is prioritised, with the costs of reproduction and migration paid in delayed arrival date. [ABSTRACT FROM AUTHOR]

Geographic variation in aridity determines environmental productivity patterns, including large-scale variability in pathogens, vectors and associated diseases. If disease risk decreases with increasing aridity and is matched by immune defense, we predict a decrease in innate immune function along a gradient of increasing aridity from the cool-wet forest to the hot-dry Sahel, from south to north in Nigeria. We sampled blood and measured five innate immune indices from 286 Common Bulbuls Pycnonotus barbatus between 6 and 13°N. We sampled in the dry season; we resampled the first location (Jos) also as the last sample location to test temporal change in immune function. Immune indices did not decrease with aridity. One immune index, nitric oxide concentration showed a weak quadratic pattern. In Jos, ovotransferrin concentration, haemagglutination and haemolysis titres increased 12 weeks into the dry season, contrary to expectations that immune indices should decrease with increased dryness. In this tropical system, innate immune function does not decrease with increasing aridity but temporal factors within a location may influence immune function more strongly than spatial variation in aridity, suggesting that immune variation does not follow a simple environmental productivity pattern. Consequently, caution should probably be exercised in predicting effects of climate variability on immune function or disease risk. [ABSTRACT FROM AUTHOR]

Predation is one of the most relevant selective forces in nature. However, the physiological mechanisms behind anti‐predator strategies have been overlooked, despite their importance to understand predator–prey interactions. In this context, the immune system could be especially revealing due to its relationship with other critical functions and its ability to enhance prey's probabilities of survival to a predator's attack. Developing organisms (e.g. nestlings) are excellent models to study this topic because they suffer a high predation pressure while undergoing the majority of their development, which maximizes potential trade‐offs between immunity and other biological functions. Using common blackbirds Turdus merula as model species, we experimentally investigated whether an elevated nest predation risk during the nestling period affects nestlings' immunity and its possible interactions with developmental conditions (i.e. body condition and growth). Experimental nestlings modified some components of their immunity, but only when considering body condition and growth rate, indicating a multifaceted immunological response to predation risk and an important mediator role of nestlings' developmental conditions. Predation risk induced a suppression of IgY but an increase in lymphocytes in nestlings with poor body condition. In addition, experimental but not control nestlings showed a negative correlation between growth and heterophils, demonstrating that nest predation risk can affect the interaction between growth and immunity. This study highlights the importance of immunity in anti‐predator response in nestlings and shows the relevance of including physiological components to the study of predation risk. [ABSTRACT FROM AUTHOR]

Home‐range studies have received considerable attention from ecologists but are greatly skewed towards the north temperate areas. Tropical areas offer an ideal setting to tease apart hypotheses about weather, food availability and social interactions as important factors influencing home‐range. In this study, we investigated home‐range and movement patterns of the tropical Red‐capped Lark Callandrella cineria, a year‐round breeding bird with a dynamic social structure. We tracked 56 individuals using radiotransmitters and colour‐ring readings over a 23‐month period. Our objective was to understand year‐round variation in home‐range size in the context of the highly aseasonal and unpredictable variation in weather and resources typical of many equatorial habitats, in addition to the birds' changing social structure and year‐round breeding. The mean composite monthly home‐range of Red‐capped Larks was 58.0 ha, and the mean individual home‐range size was 19.9 ha, but this varied considerably between individuals. The total number of nests found per month (breeding intensity) best predicted home‐range size of non‐breeding birds, and of breeding and non‐breeding birds combined. We show for the first time that breeding intensity decreases the home‐range size of non‐breeding individuals. Our study also underlines the relevance of conducting more studies in aseasonal tropical areas in order to disentangle effects of weather, food availability and breeding that vary in parallel, peaking simultaneously in most seasonal areas. [ABSTRACT FROM AUTHOR]

Diet alteration may lead to nutrient limitations even in the absence of food limitation, and this may affect physiological functions, including immunity. Nutrient limitations may also affect the maintenance of body mass and key life‐history events that may affect immune function. Yet, variation in immune function is largely attributed to energetic trade‐offs rather than specific nutrient constraints.To test the effect of diet on life‐history traits, we tested how diet composition affects innate immune function, body mass and moult separately and in combination with each other, and then used path analyses to generate hypotheses about the mechanistic connections between immunity and body mass under different diet compositions.We performed a balanced parallel and crossover design experiment with omnivorous common bulbuls Pycnonotus barbatus in out‐door aviaries in Nigeria. We fed 40 wild‐caught bulbuls ad libitum on fruits or invertebrates for 24 weeks, switching half of each group between treatments after 12 weeks. We assessed innate immune indices (haptoglobin, nitric oxide and ovotransferrin concentrations, and haemagglutination and haemolysis titres), body mass and primary moult, fortnightly. We simplified immune indices into three principal components (PCs), but we explored mechanistic connections between diet, body mass and each immune index separately.Fruit‐fed bulbuls had higher body mass, earlier moult and showed higher values for two of the three immune PCs compared to invertebrate‐fed bulbuls. These effects were reversed when we switched bulbuls between treatments after 12 weeks. Exploring the correlations between immune function, body mass and moult, showed that an increase in immune function was associated with a decrease in body mass and delayed moult in invertebrate‐fed bulbuls, while fruit‐fed bulbuls maintained body mass despite variation in immune function. Path analyses indicated that diet composition was most likely to affect body mass and immune indices directly and independently from each other. Only haptoglobin concentration was indirectly linked to diet composition via body mass.We demonstrated a causal effect of diet composition on innate immune function, body mass and moult: bulbuls were in a better condition when fed on fruits than invertebrates, confirming that innate immunity is nutrient specific. Our results are unique because they show a reversible effect of diet composition on wild adult birds whose immune systems are presumably fully developed and adapted to wild conditions—demonstrating a short‐term consequence of diet alteration on life‐history traits. [ABSTRACT FROM AUTHOR]

In some tropical birds, breeding seasonality is weak at the population level, even where there are predictable seasonal peaks in environmental conditions. It therefore remains unclear whether individuals are adapted to breeding at specific times of the year or flexible to variable environmental conditions. We tested whether the relative year‐round breeding activity of the Common Bulbul Pycnonotus barbatus arises due to within‐individual variability in breeding dates. We collected data from 827 birds via mist‐netting over 2 years with corresponding local weather data. We used a combination of climate envelope and generalized linear mixed models to explore how the timing of breeding is influenced by time of year, individual variation, rainfall and temperature in a West African savannah where seasonal precipitation determines annual variation in environmental conditions. We also pooled 65 breeding records from 19 individuals recorded between 2006 and 2017 based on brood patch occurrence and behavioural observation to compare within‐individual and population variability in breeding dates. We show that the breeding dates of individuals may be as variable as for the population as a whole. However, we observed a seasonal peak in juvenile occurrence that varies significantly between years. Models suggest no relationship between nesting and moult, and within‐year variation in rainfall and temperature, and birds were unlikely to breed during moult but may do so afterwards. Moult was very seasonal, correlating strongly with day length. We suggest that because environmental conditions permit year‐round breeding, and because reproductive output is subject to high predation risk, there is probably a weak selection for individuals to match breeding with variable peak conditions in the environment. Instead, moult, which always occurs annually and successfully, is probably under strong selection to match variable peak conditions in the environment so that long‐term survival ensures future reproduction. [ABSTRACT FROM AUTHOR]

Seasonal variation in innate immunity is often attributed to either temporal environmental variation or to life‐history trade‐offs that arise from specific annual cycle stages but decoupling them is difficult in natural populations.Here, we effectively decouple seasonal environmental variation from annual cycle stage effects by exploiting cross‐seasonal breeding and moult in the tropical Common Bulbul Pycnonotus barbatus. We test how annual cycle stage interacts with a key seasonal environmental variable, rainfall, to determine immunity at population and individual level. If immune challenge varies with precipitation, we might expect immune function to be higher in the wet season due to increased environmental productivity. If breeding or moult imposes resource constraints on birds, depending on or independent of precipitation, we might expect lower immune indices during breeding or moult.We sampled blood from 818 birds in four annual cycle stage categories: breeding, moult, simultaneous breeding and moulting, or neither. We quantified indices of innate immunity (haptoglobin, nitric oxide (NOx) and ovotransferrin concentrations, and haemagglutination and haemolysis titres) over two annual cycles of wet and dry seasons.Environment (but not annual cycle stage or interactions between both) explained variation in all immune indices, except NOx. NOx concentration differed between annual cycle stages but not between seasons. However, within the wet season, haptoglobin, NOx, ovotransferrin and haemolysis differed significantly between breeding and non‐breeding females. Aside from some recorded inconsistencies, population level results were largely similar to results within individuals that were measured repeatedly. Unexpectedly, most immune indices were higher in the dry season and during breeding.Higher immune indices may be explained if fewer or poorer quality resources force birds to increase social contact, thereby exposing individuals to novel antigens and increased infection risk, independently of environmental productivity. Breeding birds may also show higher immunity if less immune‐competent and/or infected females omit breeding. We conclude that seasonal environmental variation impacts immunity more directly in natural animal populations than via resource trade‐offs. In addition, immune indices were more often variable within than among individuals, but some indices are characteristic of individuals, and so may offer selective advantages if heritable. Tropical environments are considered highly immune‐challenging for animals because of their relative higher organic productivity. However, this study from a seasonally arid tropical environment where annual cycle stages are decoupled from seasonal environmental variation, provides evidence that variation in innate immune function does not follow differences in annual cycle stages or simple environmental productivity pattern but may arise from interactions between individuals and their environment, and this may apply to variation in disease risk. [ABSTRACT FROM AUTHOR]

Bergmann’s rule, defined as the tendency for endotherms to be larger in colder environments, is a biophysical generalization of body size variation that is frequently tested along latitudinal gradients, even though latitude is only a proxy for temperature variation. We test whether variation in temperature and aridity determine avian body size conformity to Bergmann’s rule independent of latitude differences, using the ubiquitous Common Bulbul Pycnonotus barbatus, along a West African environmental gradient. We trapped 538 birds in 22 locations between latitudes 6 and 13°N in Nigeria, and estimated average body surface area to mass ratio per location. We then modelled body surface to mass ratio using general linear models, with latitude, altitude and one of 19 bioclimatic variables extracted from http://www.worldclim.org/bioclim as predictors. We sequentially dropped latitude and altitude from each model to obtain the R2 of the resultant models. Finally, we compared the R2 of univariate models, where bioclimatic variables predicted body surface area to mass ratio significantly (14 out of 19), to multivariate models including latitude, altitude and a bioclimatic variable, using the Wilcoxon matched pairs test. We found that multivariate models did not perform better than univariate models with only bioclimatic variables. Six temperature and eight precipitation variables significantly predicted variation in body surface area to mass ratio between locations; in fact, 50% (seven out of 14) of these better explained variation in body surface area to mass ratio than the multivariate models. Birds showed a larger body surface area relative to body mass ratio in hotter environments independent of latitude or altitude, which conforms to Bergmann’s rule. Yet, a combination of morphometric analyses and controlled temperature-exposure experiments is required to prove the proposed relationship between relative body surface area and thermoregulation in endotherms. [ABSTRACT FROM AUTHOR]

Most altricial birds remove their nestlings’ feces from the nest, but the evolutionary forces driving this behavior are poorly understood. A possible adaptive explanation for this could be that birds avoid the attraction of nest predators to their nests due to the visual or olfactory cues produced by feces (nest predation hypothesis). This hypothesis has received contrasting support indicating that additional experimental studies are needed, particularly with respect to the visual component of fecal sacs. To test this hypothesis, we conducted an experiment manipulating the presence of fecal sacs on inactive Woodlark (Lullula arborea) nests. This ground nester has highly cryptic nests that are mainly depredated by visually oriented nest predators (i.e., corvids) in our study population, making it an excellent system to test for the nest predation hypothesis. Our results showed that the presence of fecal sacs in the nest does not seem to be an important factor explaining nest predation. Interestingly, the effect of nest concealment, the most important factor explaining nest predation in Woodlark nests, depended on whether the nest was depredated the previous year or not, supporting the importance of using different nesting sites between years. Our findings indicate that this important nest sanitation behavior is not likely motivated by nest predation and highlight the need to explore alternative selective pressures in this context. [ABSTRACT FROM AUTHOR]

Nest survival is critical to breeding in birds and plays an important role in life‐history evolution and population dynamics. Studies evaluating the proximate factors involved in explaining nest survival and the resulting temporal patterns are biased in favor of temperate regions. Yet, such studies are especially pertinent to the tropics, where nest predation rates are typically high and environmental conditions often allow for year‐round breeding. To tease apart the effects of calendar month and year, population‐level breeding activity and environmental conditions, we studied nest survival over a 64‐month period in equatorial, year‐round breeding red‐capped larks Calandrella cinerea in Kenya. We show that daily nest survival rates varied with time, but not in a predictable seasonal fashion among months or consistently among years. We found negative influences of flying invertebrate biomass and rain on nest survival and higher survival of nests when nests were more abundant, which suggests that nest predation resulted from incidental predation. Although an increase in nest predation is often attributed to an increase in nest predators, we suggest that in our study, it may be caused by altered predator activity resulting from increased activity of the primary prey, invertebrates, rather than activity of the red‐capped larks. Our results emphasize the need to conduct more studies in Afro‐tropical regions because proximate mechanisms explaining nest predation can be different in the unpredictable and highly variable environments of the tropics compared with the relatively predictable seasonal changes found in temperate regions. Such studies will aid in better understanding of the environmental influences on life‐history variation and population dynamics in birds. [ABSTRACT FROM AUTHOR]

Predation risk is thought to modify the physiology of prey mainly through the stress response. However, little is known about its potential effects on the immunity of animals, particularly in young individuals, despite the importance of overcoming wounding and pathogen aggression following a predator attack.We investigated the effect of four progressive levels of nest predation risk on several components of the immune system in common blackbird (Turdus merula) nestlings by presenting them with four different calls during 1 h: non-predator calls, predator calls, parental alarm calls and conspecific distress calls to induce a null, moderate, high and extreme level of risk, respectively. Nest predation risk induced an increase in ovotransferrin, immunoglobulin and the number of lymphocytes and eosinophils. Thus, the perception of a potential predator per se could stimulate the mobilization of a nestling's immune function and enable the organism to rapidly respond to the immune stimuli imposed by a predator attack. Interestingly, only high and extreme levels of risk caused immunological changes, suggesting that different immunological parameters are modulated according to the perceived level of threat. We also found a mediator role of parasites (i.e. Leucocytozoon) and the current health status of the individual, as only nestlings not parasitized or in good body condition were able to modify their immune system. This study highlights a previously unknown link between predation risk and immunity, emphasizing the complex relationship among different selective pressures (predation, parasitism) in developing organisms and accentuating the importance of studying predation from a physiological point of view. [ABSTRACT FROM AUTHOR]

Background: Variation in growth and immune function within and among populations is often associated with specific environmental conditions. We compared growth and immune function in nestlings of year-round breeding equatorial Red-capped Lark Calandrella cinerea from South Kinangop, North Kinangop and Kedong (Kenya), three locations that are geographically close but climatically distinct. In addition, we studied growth and immune function of lark nestlings as a function of year-round variation in breeding intensity and rain within one location. We monitored mass, wing, and tarsus at hatching (day 1) and at 4, 7, and 10 days post-hatch, and we quantified four indices of immune function (haptoglobin, agglutination, lysis and nitric oxide) using blood samples collected on day 10. Results: Nestling body mass and size at hatching, which presumably reflect the resources that females allocated to their eggs, were lowest in the most arid location, Kedong. Contrary to our predictions, nestlings in Kedong grew faster than nestlings in the two other cooler and wetter locations of South and North Kinangop. During periods of peak reproduction within Kedong, nestlings were heavier at hatching, but they did not grow faster over the first 10 days post-hatch. In contrast, rainfall, which did not relate to timing of breeding, had no effect on hatching mass, but more rain did coincide with faster growth post-hatch. Finally, we found no significant differences in nestling immune function, neither among locations nor with the year-round variation within Kedong. Conclusion: Based on these results, we hypothesize that female body condition determines nestling mass and size at hatching, but other independent environmental conditions subsequently shape nestling growth. Overall, our results suggest that environmental conditions related to food availability for nestlings are relatively unimportant to the timing of breeding in equatorial regions, while these same conditions do have consequences for nestling size and growth. [ABSTRACT FROM AUTHOR]

Timing of reproduction in birds is important for reproductive success and is known to depend on environmental cues such as day length and food availability. However, in equatorial regions, where day length is nearly constant, other factors such as rainfall and temperature are thought to determine timing of reproduction. Rainfall can vary at small spatial and temporal scales, providing a highly fluctuating and unpredictable environmental cue. In this study we investigated the extent to which spatio-temporal variation in environmental conditions can explain the timing of breeding of Red-capped Lark, Calandrella cinerea, a species that is capable of reproducing during every month of the year in our equatorial east African study locations. For 39 months in three climatically-distinct locations, we monitored nesting activities, sampled ground and flying invertebrates, and quantified rainfall, maximum (Tmax) and minimum (Tmin) temperatures. Among locations we found that lower rainfall and higher temperatures did not coincide with lower invertebrate biomasses and decreased nesting activities, as predicted. Within locations, we found that rainfall, Tmax, and Tmin varied unpredictably among months and years. The only consistent annually recurring observations in all locations were that January and February had low rainfall, high Tmax, and low Tmin. Ground and flying invertebrate biomasses varied unpredictably among months and years, but invertebrates were captured in all months in all locations. Red-capped Larks bred in all calendar months overall but not in every month in every year in every location. Using model selection, we found no clear support for any relationship between the environmental variables and breeding in any of the three locations. Contrary to popular understanding, this study suggests that rainfall and invertebrate biomass as proxy for food do not influence breeding in equatorial Larks. Instead, we propose that factors such as nest predation, female protein reserves, and competition are more important in environments where weather and food meet minimum requirements for breeding during most of the year. [ABSTRACT FROM AUTHOR]

Birds should store body reserves if starvation risk is anticipated; this is known as an 'interrupted foraging response'. If foraging remains unrestricted, however, body mass should remain low to limit the predation risk that gaining and carrying body reserves entails. In temperate environments mass gain in female birds during breeding is often attributed to egg formation and mass loss after incubation to flight adaptation or the effect of reproductive workload, rather than as a result of an adaptive interrupted foraging response to the limited foraging time or unpredictable foraging conditions that breeding demands. In tropical environments, foraging conditions vary more within the breeding season than in temperate environments, and so studies in tropical environments are more suited to decouple the potentially confounded effects of increase in body reserves versus egg formation on the body mass of breeding birds. In this study, we test whether breeding results in an interrupted foraging response in a tropical savannah system using body mass data collected over a 15-year period from female common bulbuls Pycnonotus barbatus. This species breeds both in the wet and dry season, despite fewer resources being available in the dry season. Breeding stage predicted female body mass: body mass peaked abruptly during incubation, but was not closely associated with the egg-laying stage, and declined during brood rearing. Breeding females were heavier in the dry season than in the wet season. In the dry season, heavier birds were more likely to incubate eggs or brood chicks. These observations suggest that increased body reserves are required to buffer the consequence of limited foraging time or impoverished foraging conditions, which may be most pronounced during incubation and in the dry season, respectively. Such mass increases are consistent with an interrupted foraging response, which may apply to temperate zone birds experiencing foraging restrictions during breeding. [ABSTRACT FROM AUTHOR]

The family Alaudidae, larks, comprises 93–100 species (depending on taxonomy) that are widely distributed across Africa and Eurasia, with single species extending their ranges to North and northernmost South America and Australia. A decade-old molecular phylogeny, comprising ∼80% of the species, revealed multiple cases of parallel evolution and large variation in rates of morphological evolution, which had misled taxonomists into creating many non-monophyletic genera. Here, we reconstruct the phylogeny of the larks, using a dataset covering one mitochondrial and 16 nuclear loci and comprising all except one of the currently recognised species as well as several recently proposed new species (in total 133 taxa; not all loci available for all species). We provide additional support using genome-wide markers to infer a genus-level phylogeny based on near-complete generic sampling (in total 51 samples of 44 taxa across 40 species). Our results confirm the previous findings of rampant morphological convergence and divergence, and reveal new cases of paraphyletic genera. We propose a new subfamily classification, and also that the genus Mirafra is divided into four genera to produce a more balanced generic classification of the Alaudidae. Our study supports recently proposed species splits as well as some recent lumps, while also questioning some of the latter. This comprehensive phylogeny will form an important basis for future studies, such as comparative studies of lark natural history, ecology, evolution and conservation. [ABSTRACT FROM AUTHOR]

Many animal species have populations in which some individuals migrate and others remain on the breeding grounds. This phenomenon is called partial migration. Despite substantial theoretical work, empirical data on causes and consequences of partial migration remain scarce, mainly because of difficulties associated with tracking individuals over large spatial scales. We used stable hydrogen isotopes in claw material to determine whether skylarks Alauda arvensis from a single breeding population in the Netherlands had migrated or remained resident in the previous winter and investigated whether there were causes or consequences of either strategy. Age and sex had no influence on the propensity to migrate, but larger individuals were more likely to be residents. The wintering strategy was not fixed within individuals. Up to 45% of individuals measured in multiple years switched strategies. Reproductive parameters were not related to the wintering strategy, but individuals that wintered locally experi-enced lower future return rates, and this was directly correlated with two independent measures of immune function. Our results suggest that partial migration in skylarks is based neither on genetic dimorphism nor on an age-and sex-dependent condition. Instead, the wintering strategy is related to structural size and immune function. These new insights on causes and consequences of partial migration advance our understanding of the ecology, evolution, and coexistence of different life-history strategies. [ABSTRACT FROM AUTHOR]

Microbial invasion of egg contents is a cause of embryonic death. To counter infection risks, the embryo is protected physically by the eggshell and chemically by antimicrobial proteins. If microbial pressure drives embryo mortality, then females may have evolved, through natural selection, to adapt their immune investment into eggs. Although frequently hypothesized, this match between immune allocation and microorganisms has not been explored yet. To examine if correlations between microbes on eggs and immunity in eggs exist, we collected eggs from red-capped larks (Calandrella cinerea) and simultaneously examined their bacterial communities and antimicrobial components—pH, lysozyme and ovotransferrin—during natural incubation. Using molecular techniques, we find that bacterial communities are highly dynamic: bacterial abundance increases from the onset to late incubation, Shannon’s α-diversity index increases during early incubation stages, and β-diversity analysis shows that communities from 1 day-old clutches are phylogenetically more similar to each other than the older ones. Regarding the antimicrobials, we notice a decrease of pH and lysozyme concentration, while ovotransferrin concentration increases during incubation. Interestingly, we show that two eggs of the same clutch share equivalent immune protection, independent of clutch age. Lastly, our results provide limited evidence of significant correlation between antimicrobial compounds and bacterial communities. Our study examined simultaneously, for the first time in a wild bird, the dynamics of bacterial communities present on eggshells and of albumen-associated antimicrobial components during incubation and investigated their relationship. However, the link between microorganisms and immunity of eggs remains to be elucidated further. Identifying invading microbes and their roles in embryo mortality, as well as understanding the role of the eggshell microbiome, might be key to better understand avian strategies of immune maternal investment. [ABSTRACT FROM AUTHOR]

Introduction All bird eggs are exposed to microbes in the environment, which if transmitted to the developing embryo, could cause hatching failure. However, the risk of trans-shell infection varies with environmental conditions and is higher for eggs laid in wetter environments. This might relate to generally higher microbial abundances and diversity in more humid environments, including on the surface of eggshells, as well as the need for moisture to facilitate microbial penetration of the eggshell. To protect against microbial infection, the albumen of avian eggs contains antimicrobial proteins, including lysozyme and ovotransferrin. We tested whether lysozyme and ovotransferrin activities varied in eggs of larks (Alaudidae) living along an arid-mesic gradient of environmental aridity, which we used as a proxy for risk of trans-shell infection. Results Contrary to expectations, lysozyme activity was highest in eggs from hotter, more arid locations, where we predicted the risk of trans-shell infection would be lower. Ovotransferrin concentrations did not vary with climatic factors. Temperature was a much better predictor of antimicrobial protein activity than precipitation, a result inconsistent with studies stressing the importance of moisture for trans-shell infection. Conclusions Our study raises interesting questions about the links between temperature and lysozyme activity in eggs, but we find no support for the hypothesis that antimicrobial protein deposition is higher in eggs laid in wetter environments. [ABSTRACT FROM AUTHOR]

Microorganisms are closely associated with eggs and may play a determinant role in embryo survival. Yet, the majority of studies focusing on this association relied on culture-based methodology, eventually leading to a skewed assessment of microbial communities. By targeting the 16 S r RNA gene and internal transcribed spacer ( ITS) region, we, respectively, described bacterial and fungal communities on eggshells of the homing pigeon Columba livia. We explored their structure, abundance, and composition. Firstly, we showed that sampling technique affected the outcome of the results. While broadly used, the egg swabbing procedure led to a lower DNA extraction efficiency and provided different profiles of bacterial communities than those based on crushed eggshell pieces. Secondly, we observed shifts in bacterial and fungal communities during incubation. At late incubation, bacterial communities showed a reduction in diversity, while their abundance increased, possibly due to the competitive advantage of some species. When compared to their bacterial counterparts, fungal communities also decreased in diversity at late incubation. In that case, however, the decline was associated with a diminution of their overall abundance. Conclusively, our results showed that although incubation might inhibit microbial growth when compared to unincubated eggs, we observed the selective growth of specific bacterial species during incubation. Moreover, we showed that fungi are a substantial component of the microbial communities associated with eggshells and require further investigations in avian ecology. Identifying the functional roles of these microorganisms is likely to provide news insights into the evolutionary strategies that control embryo survival. [ABSTRACT FROM AUTHOR]

Trade-offs between immune function and other physiological and behavioural processes are central in ecoimmunology, but one important problem is how to distinguish a reallocation of resources away from the immune system from a reallocation or redistribution within the immune system. While variation in baseline values of individual immune parameters is well established, studies in wild animals on multiple parameters during an immune response are lacking. It also remains to be tested whether and how immune responses correlate with baseline values that vary, for example, over the course of an annual cycle. We studied immunological responses to an endotoxin challenge in skylarks (Alauda arvensis), a partial migrant bird breeding in temperate zones. We compared birds injected with the endotoxin LPS with un-injected controls, characterizing immunological responses with leukocyte profiles, titres of lytic enzymes and natural antibodies, and concentrations of haptoglobin and heat shock proteins. We did this in five annual-cycle stages to test whether the response varied throughout the year. The endotoxin challenge affected six of 10 measured parameters. Lysis titres and proportions of heterophils increased; haptoglobin concentrations and proportions of lymphocytes, basophils and eosinophils decreased. The variable effects on different immune components demonstrate the complexity of an Immune response. We found no evidence that the response differed between annual-cycle stages. The response was independent of baseline measures taken directly upon capture in the field, indicating that birds were facing no immunological ceiling when mounting an immune response. Values of five parameters collected under field conditions were significantly related to values taken under standardized laboratory conditions. We conclude that multiple parts of the immune system are modulated during an immunological response and that responses are not re-organized throughout the annual cycle. [ABSTRACT FROM AUTHOR]

Intercellular and covalently bound lipids within the stratum corneum (SC), the outermost layer of the epidermis, are the primary barrier to cutaneous water loss (CWL) in birds. We compared CWL and intercellular SC lipid composition in 20 species of birds from desert and mesic environments. Furthermore, we compared covalently bound lipids with CWL and intercellular lipids in the lark family (Alaudidae). We found that CWL increases in birds from more mesic environments, and this increase was related to changes in intercellular SC lipid composition. The most consistent pattern that emerged was a decrease in the relative amount of cerebrosides as CWL increased, a pattern that is counterintuitive based on studies of mammals with Gaucher disease. Although covalently bound lipids in larks did not correlate with CWL, we found that covalently bound cerebrosides correlated positively with intercellular cerebrosides and intercellular cholesterol ester, and intercellular cerebrosides correlated positively with covalently bound free fatty acids. Our results led us to propose a new model for the organization of lipids in the avian SC, in which the sugar moieties of cerebrosides lie outside of intercellular lipid layers, where they may interdigitate with adjacent intercellular cerebrosides or with covalently bound cerebrosides. [ABSTRACT FROM AUTHOR]

Most birds rely on flight for survival. Yet as an energetically taxing and physiologically integrative process, flight has many repercussions. Studying pigeons (Columba h'via) and employing physiological and immunological indices that are relevant to ecologists working with wild birds, we determined what, if any, acute immune-like responses result from bouts of intense, non-migratory flight. We compared the effects of flight with the effects of a simulated bacterial infection. We also investigated indices in terms of their post-flight changes within individuals and their relationship with flight speed among individuals. Compared to un-flown controls, flown birds exhibited significant elevations in numbers of heterophils relative to numbers of lymphocytes and significant reductions in numbers of eosinophils and monocytes. Furthermore, within-individual changes in concentrations of an acute phase protein were greater in flown birds than in controls. However, none of the flight-affected indices showed any evidence of being related to flight speed. While some of the effects of flight were comparable to the effects of the simulated bacterial infection, other effects were observed only after one of these two physiological challenges. Our study suggests that flight by pigeons yields immune-like responses, and these responses have the potential to complicate the conclusions drawn by ecologists regarding immune function in free-living birds. Still, a better understanding of the repercussions of flight can help clarify the ties between the physiology of exercise and the disease ecology of migration and will ultimately assist in the broader goal of accounting for immunological variation within and among species. [ABSTRACT FROM AUTHOR]

Immune defense may vary as a result of trade-offs with other life-history traits or in parallel with variation in antigen levels in the environment. We studied lark species (Alaudidae) in the Arabian Desert and temperate Netherlands to test opposing predictions from these two hypotheses. Based on their slower pace of life, the trade-off hypothesis predicts relatively stronger immune defenses in desert larks compared with temperate larks. However, as predicted by the antigen exposure hypothesis, reduced microbial abundances in deserts should result in desert-living larks having relatively weaker immune defenses. We quantified host-independent and host-dependent microbial abundances of culturable microbes in ambient air and from the surfaces of birds. We measured components of immunity by quantifying concentrations of the acute-phase protein haptoglobin, natural antibody-mediated agglutination titers, complement- mediated lysis titers, and the microbicidal ability of whole blood. Desert-living larks were exposed to significantly lower concentrations of airborne microbes than temperate larks, and densities of some bird-associated microbes were also lower in desert species. Haptoglobin concentrations and lysis titers were also significantly lower in desert-living larks, but other immune indexes did not differ. Thus, contrary to the trade-off hypothesis, we found little evidence that a slow pace of life predicted increased immunological investment. In contrast, and in support of the antigen exposure hypothesis, associations between microbial exposure and some immune indexes were apparent. Measures of antigen exposure, including assessment of host-independent and host-dependent microbial assemblages, can provide novel insights into the mechanisms underlying immunological variation. [ABSTRACT FROM AUTHOR]

A central hypothesis of ecological immunology is that immune defences are traded off against competing physiological and behavioural processes. During energetically demanding periods, birds are predicted to switch from expensive inflammatory responses to less costly immune responses. Acute phase responses (APRs) are a particularly costly form of immune defence, and, hence, seasonal modulations in APRs are expected. Yet, hypotheses about APR modulation remain untested in freeliving organisms throughout a complete annual cycle. We studied seasonal modulations in the APRs and in the energy budgets of skylarks Alauda arvensis, a partial migrant bird from temperate zones that experiences substantial ecological changes during its annual cycle. We characterized throughout the annual cycle changes in their energy budgets by measuring basal metabolic rate (BMR) and body mass. We quantified APRs by measuring the effects of a lipopolysaccharide injection on metabolic rate, body mass, body temperature, and concentrations of glucose and ketone. Body mass and BMR were lowest during breeding, highest during winter and intermediate during spring migration, moult and autumn migration. Despite this variation in energy budgets, the magnitude of the APR, as measured by all variables, was similar in all annual cycle stages. Thus, while we find evidence that some annual cycle stages are relatively more energetically constrained, we find no support for the hypothesis that during these annual cycle stages birds compromise an immune defence that is itself energetically costly. We suggest that the ability to mount an APR may be so essential to survival in every annual cycle stage that skylarks do not trade off this costly form of defence with other annual cycle demands. [ABSTRACT FROM AUTHOR]

Changes in agricultural management have been identified as the most probable cause for the decline of Skylark (Alauda arvensis) populations in Europe. However, parasitic infections have not been considered as a possible factor influencing this process. Four hundred and thirty-four Skylarks from the Southern Italy and the Netherlands were screened for haemosporidian parasites (Haemosporida) using the microscopy and polymerase chain reaction (PCR)-based methods. The overall prevalence of infection was 19.5%; it was 41.8% in Italian birds and 8.3% in Dutch birds. The prevalence of Plasmodium spp. was 34.1% and 6.5% in Skylarks from Italy and Netherlands, respectively. Approximately 15% of all recorded haemosporidian infections were simultaneous infections both in Italian and Dutch populations. Six different mitochondrial cytochrome b (cyt b) lineages of Plasmodium spp. and three lineages of Haemoproteus tartakovskyi were found. The lineage SGS1 of Plasmodium relictum was the most prevalent at both study sites; it was recorded in 24.7% of birds in Italy and 5.5% in the Netherlands. The lineages SYAT05 of Plasmodium vaughani and GRW11 of P. relictum were also identified with a prevalence of <2% at both study sites. Two Plasmodium spp. lineages (SW2 and DELURB4) and three H tartakovskyi lineages have been found only in Skylarks from Italy. Mitochondrial cyt b lineages SYAT05 are suggested for molecular identification of P. vaughani, a cosmopolitan malaria parasite of birds. This study reports the greatest overall prevalence of malaria infection in Skylarks during the last 100 years and shows that both Plasmodium and Haemoproteus spp. haemosporidian infections are expanding in Skylarks so it might contribute to a decrease of these bird populations in Europe. [ABSTRACT FROM AUTHOR]

The extent to which organisms can protect themselves from disease depends on both the immune defenses they maintain and the pathogens they face. At the same time, immune systems are shaped by the antigens they encounter, both over ecological and evolutionary time. Ecological immunologists often recognize these interactions, yet ecological immunology currently lacks major efforts to characterize the environmental, host-independent, antigenic pressures to which all animals are exposed. Failure to quantify relevant diseases and pathogens in studies of ecological immunology leads to contradictory hypotheses. In contrast, including measures of environmental and host-derived commensals, pathogens, and other immune-relevant organisms will strengthen the field of ecological immunology. In this article, we examine how pathogens and other organisms shape immune defenses and highlight why such information is essential for a better understanding of the causes of variation in immune defenses. We introduce the concept of “operative protection” for understanding the role of immunologically relevant organisms in shaping immune defense profiles, and demonstrate how the evolutionary implications of immune function are best understood in the context of the pressures that diseases and pathogens bring to bear on their hosts. We illustrate common mistakes in characterizing these immune-selective pressures, and provide suggestions for the use of molecular and other methods for measuring immune-relevant organisms. [ABSTRACT FROM AUTHOR]

The immune system is a complex collection of interrelated and overlapping solutions to the problem of disease. To deal with this complexity, researchers have devised multiple ways to measure immune function and to analyze the resulting data. In this way both organisms and researchers employ many tactics to solve a complex problem. One challenge facing ecological immunologists is the question of how these many dimensions of immune function can be synthesized to facilitate meaningful interpretations and conclusions. We tackle this challenge by employing and comparing several statistical methods, which we used to test assumptions about how multiple aspects of immune function are related at different organizational levels. We analyzed three distinct datasets that characterized 1) species, 2) subspecies, and 3) among- and within-individual level differences in the relationships among multiple immune indices. Specifically, we used common principal components analysis (CPCA) and two simpler approaches, pair-wise correlations and correlation circles. We also provide a simple example of how these techniques could be used to analyze data from multiple studies. Our findings lead to several general conclusions. First, relationships among indices of immune function may be consistent among some organizational groups (e.g. months over the annual cycle) but not others (e.g. species); therefore any assumption of consistency requires testing before further analyses. Second, simple statistical techniques used in conjunction with more complex multivariate methods give a clearer and more robust picture of immune function than using complex statistics alone. Moreover, these simpler approaches have potential for analyzing comparable data from multiple studies, especially as the field of ecological immunology moves towards greater methodological standardization. [ABSTRACT FROM AUTHOR]

One route to gain insight into the causes and consequences of ecological differentiation is to understand the underlying physiological mechanisms. We explored the relationships between immunological and oxidative status and investigated how birds cope physiologically with the effects of immune-derived oxidative damage. We successively implemented two experimental manipulations to alter physiological status in a model bird species: the homing pigeon (Columba livia). The first manipulation, an immune supplementation, was achieved by oral administration of lysozyme, a naturally occurring and non-specific antimicrobial enzyme. The second manipulation, an immune challenge, took the form of an injection with lipopolysaccharide, a bacterial endotoxin. Between groups of lysozyme-treated and control birds, we compared lipopolysaccharide-induced changes in reactive oxygen metabolites, total antioxidant capacity, haptoglobin, oxygen consumption, body mass and cloacal temperature. Lysozyme supplementation intensified the lipopolysaccharide-induced inflammatory response and generated short-term oxidative and metabolic costs. We identified significant interactions between immune supplementation and immune challenge in terms of reactive oxygen metabolites, haptoglobin and oxygen consumption. Our study provides alternative interpretations of differences in oxidative and immunological indices and demonstrates that these indices can also fluctuate and interact across very short time scales, reflecting something akin to current 'health status' or 'physiological condition'. These ephemeral effects highlight the need to broadly consider current physiological condition when drawing conclusions that relate physiology to ecology and evolution. [ABSTRACT FROM AUTHOR]

Migration has fascinated researchers for years and many active areas of study exist. However, the question of how migratory species stay healthy within the context of their annual cycle remains relatively unexplored. This article addresses this question using Red Knots (Calidris canutus) as a model migrant species. We review recent research on immune function in Red Knots and integrate this work with the broader eco-immunological literature to introduce a conceptual model. This model synthesizes earlier ideas about resource allocation and the costs of immunity with recent increases in our knowledge about the vertebrate immune system and then puts these concepts into the context of defense against real pathogens in environments where a myriad of factors change in time and space. We also suggest avenues for further research, which will help to test the model and better link measures of immune function to pressure from pathogens and to optimal defense against disease. [ABSTRACT FROM AUTHOR]

The article discusses a study on the difference between the immune functions of shorebirds that were still recovering from protein lost during migratory flight and fueling birds that were depositing fat during their short stopover in Delaware Bay. It is explained that the constitutive immune function of birds is their most important defense during stopovers. The total leukocyte concentrations have been measured to gather information about other aspects of immune function such as infection.

In response to unbalanced energy budgets, animals must allocate resources among competing physiological systems to maximize fitness. Constraints can be imposed on energy availability or energy expenditure, and adjustments can be made via changes in metabolism or trade-offs with competing demands such as body-mass maintenance and immune function. This study investigates changes in constitutive immune function and the acute-phase response in shorebirds (red knots) faced with limited access time to food. We separated birds into two experimental groups receiving either 6 h or 22 h of food access and measured constitutive immune function. After 3 wk, we induced an acute-phase response, and after 1 wk of recovery, we switched the groups to the opposite food treatment and measured constitutive immune function again. We found little effect of food treatment on constitutive immune function, which suggests that even under resource limitation, a baseline level of immune function is maintained. However, birds enduring limited access to food suppressed aspects of the acute-phase response (decreased feeding and mass to maintain energy intake, and they downregulated thermoregulatory adjustments to food treatment to maintain body temperature during simulated infection. Thus, under resource-limited conditions, birds save energy on the most costly aspects of immune defense. [ABSTRACT FROM AUTHOR]

The winter immunoenhancement hypothesis associates long nights and increased exposure to melatonin with enhanced immune function in winter when resource availability is low and the chances of becoming ill are high. Thus, increased exposure to melatonin in the winter could be adaptive for species facing difficult winter conditions. This idea has found some support in studies of resident mammals. In birds, the link between day length and melatonin over the annual cycle is weaker, and contributions of melatonin to seasonal timing are unclear. Furthermore, many species, especially migrants, do not experience the most difficult conditions of their annual cycle in winter. In this study, we tested whether the winter immunoenhancement hypothesis holds in an avian species, the red knot Calidris canutus. We found that melatonin duration and amplitude varied significantly over the annual cycle with the highest values occurring in winter. However, peaks did not correspond to the winter solstice or with annual variation in immune function. Our findings do not support the winter immunoenhancement hypothesis in knots and question whether the idea that immune function should be bolstered in winter can be generalized to systems where winter is not the most difficult time of the year. [ABSTRACT FROM AUTHOR]

Throughout the annual cycle, demands on competing physiological systems change, and animals must allocate resources to maximize fitness. Immune function is one such system and is important for survival. Yet detailed empirical data tracking immune function over the entire annual cycle are lacking for most wild animals. We measured constitutive immune indices once a month for a year on captive red knots (Calidris canutus). We also examined temperature as an environmental contributor to immune variation by manipulating ambient temperature to vary energy expenditure. To identify relationships among immune indices, we performed principal- component analysis. We found significant repeatability in immune indices over the annual cycle and covariation of immune indices within and among individuals. This covariation suggests immune strategies as individual traits among individuals and the use of different immune strategies during different annual-cycle stages within individuals. Over the annual cycle, both higher-cost phagocyte- based immunity and lower-cost lymphocyte-based immunity were high during mass change, but there was a clear shift toward lower-cost lymphocyte-based immunity during peak molt. Experimental manipulation of temperature had little effect on annual variation in immune function. This suggests that other environmental factors, such as food availability and disease, should also be examined in the future. [ABSTRACT FROM AUTHOR]

Experiments on captive animals, in which conditions can be controlled, are useful for examining complex biological phenomena such as immune function. Such experiments have increased our understanding of immune responses in the context of trade-offs and pathogen pressure. However, few studies have examined how captivity itself affects immune function. We used microbial killing, leukocyte concentrations and complement-natural antibody assays to examine non-induced (constitutive) immunity in captive and free-living red knots Calidris canutus. Univariate and multivariate analyses indicated that captive and free-living birds differed in their immune strategies. Captive birds showed reduced S. aureus killing, C. albicans killing, heterophils, and eosinophils. In a principal component analysis, the affected variables fell onto a single axis, that reflected phagocytosis and inflammation based immunity. We discuss possible reasons for this result in an immune cost and protective benefit framework. [ABSTRACT FROM AUTHOR]

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We measured resting and peak metabolic rates (RMR and PMR, respectively) during development of chicks of seven species of shorebirds: least sandpiper (Calidris minutilla; adult mass 20—22 g), dunlin (Calidris alpina; 56—62 g), lesser yellowlegs (Tringa flavipes 88—92 g), short-billed dowitcher (Limnodromus griseus; 85—112 g), lesser golden plover (Pluvialis dominicana; 150—156 g), Hudsonian godwit (Limosa haemastica; 205—274 g), and whimbrel (Numenius phaeopus; 380 g). We tested two opposing hypotheses: the growth rate-maturity hypothesis, which posits that growth rate in chicks is inversely related to functional maturity of tissues, and the fast growth rate-high metabolism hypothesis, which suggests that rapid growth is possible only with a concomitant increase in either RMR or PMR. We have found no evidence that chicks of shorebirds with fast growth rates have lower RMRs or lower PMRs, as would be predicted by the growth rate-maturity hypothesis, but our data suggested that faster-growing chest muscles resulted in increased thermogenic capacity, consistent with the fast growth-high metabolism hypothesis. The development of homeothermy in smaller species is a consequence primarily of greater metabolic intensities of heat-generating tissues. The maximum temperature gradient between a chick's body and environment that can be maintained in the absence of a net radiative load increased rapidly with body mass during development and was highest in least sandpipers and lowest among godwits. Chicks of smaller species could maintain a greater temperature gradient at a particular body mass because of their higher mass-specific maximum metabolic rates. [ABSTRACT FROM AUTHOR]

In wild birds, relatively little is known about intra- or interspecific variation in immunological capabilities, and even less is known about the effects of stress on immune function. Immunological assays adaptable to field settings and suitable for a wide variety of taxa will prove most useful for addressing these issues. We describe a novel application of an in vitro technique that measures the intrinsic bacteria-killing abilities of blood. We assessed the capacities of whole blood and plasma from free-living individuals of five tropical bird species to kill a nonpathogenic strain of E. coli before and after the birds experienced an acute stress. Killing invasive bacteria is a fundamental immune function, and the bacteria-killing assay measures constitutive, innate immunity integrated across circulating cell and protein components. Killing ability varied significantly across species, with common ground doves exhibiting the lowest levels and blue-crowned motmots exhibiting the highest levels. Across species, plasma killed bacteria as effectively as whole blood, and higher concentrations of plasma killed significantly better. One hour of acute stress reduced killing ability by up to 40%. This assay is expected to be useful in evolutionary and ecological studies dealing with physiological and immunological differences in birds. [ABSTRACT FROM AUTHOR]

We call into question the idea that birds have not evolved unique physiological adaptations to desert environments. The rate at which desert larks metabolize energy is lower than in mesic species within the same family, and this lower rate of living translates into a lower overall energy requirement in the wild. We argue that selection has reduced oxygen consumption at the tissue level under basal conditions for birds living in deserts. We document that total evaporative water loss--the sum of cutaneous water loss (CWL) and respiratory water loss--is reduced in desert birds, and present evidence that changes in CWL are responsible for this pattern. The diminution in CWL is attributable to changes in the lipid structure of the stratum corneum of the skin, the physical barrier to diffusion of water vapor. Finally, we show linkages between physiology and life-history attributes of larks along an aridity gradient; birds from deserts have not only a reduced rate of metabolism but also a small clutch size and slow nestling development. Hence, attributes of physiology are correlated with traits that directly affect reproductive success. Our hope is that we will prompt students to question the notion that birds do not possess physiological adaptations to the desert environment, and raise the specter of doubt about "preadaptation" in birds living in deserts. [ABSTRACT FROM AUTHOR]

We compared physiological, demographic, and ecological variables of larks to gain insights into life history variation along an aridity gradient, incorporating phylogenetic relationships in analyses when appropriate. Quantifying field metabolic rate (FMR) and water influx rate (WIR) of parents feeding nestlings as measures of parental effort, we found that parental FMR and WIR of larks decreased with increasing aridity. Water and energy requirements of 6-8 d old nestlings were reduced in desert species. Nestling growth rate, clutch size, and number of clutches decreased with increasing aridity, and nest predation rates increased with increasing aridity. We combined FMR and WIR of parents and chicks, energy and water accumulated during growth, and brood size to establish energy and water budgets of parent-brood units. Parent-offspring energy budgets equaled 261 kJ/d for Hoopoe Lark, 164 kJ/d for Dunn's Lark, 388 kJ/d for Skylark, and 347 kJ/d for Woodlark, a 27% reduction in the desert species when taking into account mass differences. Family unit WIRs were 46.6 g/d for Hoopoe Lark, 37.3 g/d for Dunn's Lark, 101.6 g/d for Skylark, and 83.2 g/d for Woodlark. Parent-brood units of arid-zone species used 28-51% less water per gram mass than species from mesic areas. These results support the hypothesis that decreasing food and water availability favor lower energy and water requirements of parents and young, reduced growth rates, and smaller clutch sizes with increasing aridity. The decrease in parental effort with increasing aridity might reflect a lower fitness value of a single brood for arid-zone species than for larks from mesic habitats, suggesting that the probability of adult survival is higher in arid than in mesic areas. [ABSTRACT FROM AUTHOR]

We explored the relationship between lipids of the stratum corneum (SC), the barrier to water-vapor diffusion of the skin, and cutaneous water loss (CWL) of species of free-living larks along a temperature-moisture gradient. Our results showed that free fatty acids, cholesterol, and ceramides were the major constituents of SC in larks from different environments including the Netherlands, a mesic environment; Iran, a semiarid region; and several areas in Saudi Arabia, a hot dry desert. We found that CWL was reduced among larks inhabiting deserts, but our data did not support the hypothesis that birds from desert environments have larger quantities of lipids per unit dry mass of the SC than larks from more mesic environments. Instead, larks in arid environments had a higher proportion of ceramides, especially the more polar fractions 4/6, and a smaller proportion of free fatty acids in their SC, an adjustment that apparently reduced their CWL. Subtle changes in the ratios of lipid classes can apparently alter the movement of water vapor through the skin. We hypothesize that desert birds have higher proportions of ceramides in their SC and lower proportions of free fatty acids because this combination allows the lipid lamellae to exist in a more highly ordered crystalline phase and consequently creates a tighter barrier to water-vapor diffusion. [ABSTRACT FROM AUTHOR]

When vertebrates invaded land during the Carboniferous period, they were exposed not only to new ecological opportunities but also to a desiccating environment. To maintain cellular water homeostasis, natural selection modified the integument of pioneering terrestrial animals, enabling them to reduce water loss through the skin. In desert environments, where ambient temperatures (T[sub a]) can reach 50°C, relative humidities are low and drinking water is scarce, integumentary modifications that reduce cutaneous water loss (CWL) could be fundamental to survival. Previous research has shown that hoopoe larks (Alaemon alaudipes) from the Arabian desert reduced CWL when acclimated to 35°C compared with individuals at 15°C, but skylarks (Alauda arvensis) and woodlarks (Lullula arborea), from The Netherlands, and Dunn's larks (Eremalauda dunni), also from the Arabian desert, did not. Here, we test the idea that hoopoe larks acclimated to 35°C would alter the lipid composition of their stratum corneum (SC), resulting in a decrease in CWL, but that skylarks, woodlarks and Dunn's larks would not. Specifically, we hypothesized that hoopoe larks, acclimated to 35°C, would increase the proportions of polar ceramide content and decrease the proportions of free fatty acids in their SC compared with individuals acclimated to 15°C. Results showed that hoopoe larks at 35°C had lower CWL and higher proportions of total ceramides but lower proportions of free fatty acids and sterols in their SC. We demonstrate that adjustments in ratios of lipid classes in the SC are associated with changes in CWL in hoopoe larks. [ABSTRACT FROM AUTHOR]

We investigated interindividual variation and intra-individual phenotypic flexibility in basal metabolic rate (BMR), total evaporative water loss (TEWL), body temperature (T[sub b]), the minimum dry heat transfer coefficient (h), and organ and muscle size of five species of larks geographically distributed along an aridity gradient. We exposed all species to constant environments of 15°C or 35°C, and examined to what extent interspecific differences in physiology can be attributed to acclimation. We tested the hypothesis that birds from deserts display larger intra-individual phenotypic flexibility and smaller interindividual variation than species from mesic areas. Larks from arid areas had lower BMR, TEWL, and h, but did not have internal organ sizes different from birds from mesic habitats. BMR of 15°C-acclimated birds was 18.0%, 29.1%, 12.2%, 25.3%, and 4.7% higher than of 35°C-acclimated Hoopoe Larks, Dunn's Larks, Spike-heeled Larks, Skylarks, and Woodlarks, respectively. TEWL of 15°C-acclimated Hoopoe Larks exceeded values for 35°C-acclimated individuals by 23% but did not differ between 15°C- and 35°C-acclimated individuals in the other species. The dry heat transfer coefficient was increased in 15°C-acclimated individuals of Skylarks and Dunn's Larks, but not in the other species. Body temperature was on average 0.4°C ± 0.15°C (mean ± 1 SEM) lower in 15°C-acclimated individuals of all species. Increased food intake in 15°C-acclimated birds stimulated enlargement of intestine (26.9-38.6%), kidneys (9.824.4%), liver (16.5-27.2%), and stomach (22.0-31.6%). The pectoral muscle increased in 15°C-acclimated Spike-heeled Larks and Skylarks, remained unchanged in Hoopoe Larks, and decreased in 15°C-acclimated Woodlarks and Dunn's Larks. We conclude that the degree of intra-individual flexibility varied between physiological traits and among species, but that acclimation does not account for interspecific differences in BMR, TEWL, and h in larks. We found no general support for the hypothesis that species from desert environments display larger intra-individual phenotypic flexibility than those from mesic areas. The coefficient of variation of larks acclimated to their natural environment was smaller in species from arid areas than in species from mesic areas for mass-corrected BMR and surface-specific h, but not for mass-corrected TEWL. The high repeatabilities of BMR, TEWL, and h in several species indicated a within-individual consistency on which natural selection could operate. [ABSTRACT FROM AUTHOR]

Birds from deserts generally have lower total evaporative water loss (TEWL), the sum of cutaneous (CWL) and respiratory water loss (RWL), than species from mesic areas. We investigated the role of CWL and RWL as a function of air temperature (T[suba]) in hoopoe larks (Alaemon alaudipes) and Dunn's larks (Eremalauda dunni) from the Arabian Desert and skylarks (Alauda arvensis) and woodlarks (Lullula arborea) from temperate mesic grasslands. The proportional contribution of CWL to TEWL in all larks at moderate T[suba] ranged from 50% to 70%. At high T[suba] (40°-45°C), larks enhanced CWL by only 45%-78% and relied on an increase in RWL by 676%-2,733% for evaporative cooling. Surface-specific CWL at 25°C was 29% lower in the arid-zone species than in the mesic larks. When acclimated to constant T[suba], 15°C-acclimated hoopoe larks increased CWL by 22% compared with 35°C-acclimated birds, but the other species did not change CWL. This study is consistent with the hypothesis that larks from deserts have a reduced CWL at moderate and low T[suba] but provided no support for the hypothesis that at high T[suba] larks from arid regions rely more on CWL than larks from mesic environments, interspecific differences in CWL cannot be attributed to acclimation to environmental temperature and are possibly the result of genetic differences due to natural selection or of phenotypically plastic responses to divergent environments during ontogeny. [ABSTRACT FROM AUTHOR]