Antipredator vigilance in birds: modelling the 'edge' effect

Many animals spend a large proportion of their time either foraging for food or watching out for predators (antipredator vigilance). There have been many theoretical and empirical studies investigating the trade-off between these two activities, especially in birds. Previous models of antipredator vigilance assume that all birds within the group spend the same amount of time feeding. However, many empirical studies have shown that individuals on the edge of flocks are more vigilant. Here we describe a vigilance model which investigates the effect of position on the birds' strategies by dividing the feeding area into an inner and outer region. The model examines how various parameters such as food availability and predation risk affect the strategies of individual birds according to whether they are in the inner or outer region. Our model predicts that birds in the outer group are always more vigilant than those in the inner region. Birds in the centre receive a higher payoff in each of the wide range of scenarios that we have considered, and so our model also indicates why dominant birds would choose to feed in the centre of the group; a prediction in accord with several empirical studies.     

A comparative study of the chromosomes of birds

Karyotype analysis and morphometric measurement of the chromosomes of eleven species of Indian birds are described. The unequivocal identification of W chromosome in the females of five species of the present investigation further strengthens the generalisation that, at least in Carinatae, the sex chromosome constitutions are of ZZ and ZW types in males and females respectively. — The chromosomes of different species of birds so far worked out in each order have been compared using quantitative methods and tentative conclusions have been drawn regarding chromosomal affinities between species of different taxonomic categories.     

Sensory basis of vigilance behavior in birds: synthesis and future prospects

Birds gather visual information through scanning behavior to make decisions relevant for survival (e.g., detecting predators and finding food). The goal of this study was (a) to review some visual properties involved in scanning behavior (retinal specialization for visual resolution and motion detection, visual acuity, and size of the blind area), and (b) hypothesize how the inter-specific variability in these properties may lead to different scanning strategies. The avian visual system has a high degree of heterogeneity in visual performance across the visual field, with some sectors providing higher levels of visual resolution and motion detection (e.g., retinal specializations) than others (e.g., peripheral retina and blind area). Thus, information quality will vary in different parts of the visual field, which contradicts some theoretical assumptions on information gathering. Birds need to move their eyes and heads to align the retinal specializations to different sectors of visual space. The rates of eye and head movements can then be used as proxies for scanning strategies. I propose specific predictions as to how each of the visual properties studied can affect scanning strategies in the context of predator detection in different habitat types and with different levels of predation risk. Establishing the degree of association between sensory specializations and scanning strategies can enhance our understanding of the evolution of anti-predator behavior.     

Covariation of sexual dichromatism and plumage colours in lekking and non-lekking birds: A comparative analysis

The extreme polygyny expressed by male lekking birds leads to the expectation that sexual dimorphism should be greater in lekkers than related non-lekkers. However, evidence for this association is weak, and many lekkers are actually monomorphic in size and plumage. To better understand the kinds of plumages associated with lekking, I characterized plumage variation for combinations of sexual dichromatism and colourfulness-and-conspicuousness (COCO) among lekking and related non-lekking birds. Compared in this way, the plumages of lekkers and non-lekkers differ dramatically for both sexes. Correlations between sexual dichromatism and COCO for phylogenetically independent contrasts are significant for male lekkers (positive) and female non-lekkers (negative), but not for female lekkers or male non-lekkers. Moreover, the total number of character–state combinations, and multivariate measures of variability, are greater in non-lekkers than lekkers.The characteristic plumages of lekkers (duller monochromatic, brighter dichromatic and intermediate between these extremes) comprise just a subset of those observed among non-lekkers, and exclude extremely dull dichromatic and extremely bright monochromatic plumages. I suggest that predation, and foraging behaviours compatible with lekking, may restrict plumage variation among lekkers. Thus ecological rather than overt sexual characteristics may explain monomorphism in birds under intense mate competition, as well as the paradox of strong female mate preferences on leks, where males appear to contribute only sperm to female reproductive efforts.     

Resting breathing frequency in aquatic birds: a comparative analysis with terrestrial species

Several studies have indicated that in birds breathing frequency ( f , breaths min⁻¹) scales to the −1/3 of body weight ( W , kg); this is different from the −1/4 of mammals. We wondered if this discrepancy was due to the peculiar scaling pattern of aquatic birds, as is the case of aquatic mammals. In fact, we had noted previously that the allometric scaling of f differs considerably between aquatic and terrestrial mammals, respectively, W ^−0.42 and W ^−0.25. Measurements of f were obtained in 48 aquatic birds of 22 species and in 35 terrestrial birds of 27 species, during resting conditions on land. Additional data from 11 aquatic and 14 terrestrial species, different from the ones measured, were obtained from the literature. The allometric curve of all species combined (terrestrial and aquatic, n =74) was f =13.3 W ^−0.36, similar to what is reported in previous studies. However, the allometric curve of the aquatic species ( n =33, f =14.5 W ^−0.56) differed greatly ( P <0.001) from that of the terrestrial species ( n =41, f =13.4 W ^−0.26). On average, f of aquatic birds of the 3–5 kg range was 63%, and that of birds of larger size was 57%, of the values of terrestrial birds of similar W . We conclude that, as in mammals, also in terrestrial birds f scales to the −1/4 exponent of W . The similarity of the scaling patterns of f between aquatic birds and mammals suggests a common breathing adaptation to life in the aquatic environment irrespective of phylogenetic relations.     

A species assemblage approach to comparative phylogeography of birds in southern Australia

We present a novel approach to investigating the divergence history of biomes and their component species using single-locus data prior to investing in multilocus data. We use coalescent-based hierarchical approximate Bayesian computation (HABC) methods (MsBayes) to estimate the number and timing of discrete divergences across a putative barrier and to assign species to their appropriate period of co-divergence. We then apply a coalescent-based full Bayesian model of divergence (IMa) to suites of species shown to have simultaneously diverged. The full Bayesian model results in reduced credibility intervals around divergence times and allows other parameters associated with divergence to be summarized across species assemblages. We apply this approach to 10 bird species that are wholly or patchily discontinuous in semi-arid habitats between Australia's southwest (SW) and southeast (SE) mesic zones. There was substantial support for up to three discrete periods of divergence. HABC indicates that two species wholly restricted to more mesic habitats diverged earliest, between 594,382 and 3,417,699 years ago, three species from semi-arid habitats diverged between 0 and 1,508,049 years ago, and four diverged more recently, between 0 and 396,843 years ago. Eight species were assigned to three periods of co-divergence with confidence. For full Bayesian analyses, we accounted for uncertainty in the two remaining species by analyzing all possible suites of species. Estimates of divergence times from full Bayesian divergence models ranged between 429,105 and 2,006,355; 67,172 and 663,837; and 24,607 and 171,085 for the earliest, middle, and most recent periods of co-divergence, respectively. This single-locus approach uses the power of multitaxa coalescent analyses as an efficient means of generating a foundation for further, targeted research using multilocus and genomic tools applied to an understudied biome.     

The evolution of stereopsis and the Wulst in caprimulgiform birds: a comparative analysis

Owls possess stereopsis (i.e., the ability to perceive depth from retinal disparity cues), but its distribution amongst other birds has remained largely unexplored. Here, we present data on species variation in brain and telencephalon size and features of the Wulst, the neuroanatomical substrate that subserves stereopsis, in a putative sister-group to owls, the order Caprimulgiformes. The caprimulgiforms we examined included nightjars (Caprimulgidae), owlet-nightjars (Aegothelidae), potoos (Nyctibiidae), frogmouths (Podargidae) and the Oilbird (Steatornithidae). The owlet-nightjars and frogmouths shared almost identical relative brain, telencephalic and Wulst volumes as well as overall brain morphology and Wulst morphology with owls. Specifically, the owls, frogmouths and owlet-nightjars possess relatively large brains and telencephalic and Wulst volumes, had a characteristic brain shape and displayed prominent laminae in the Wulst. In contrast, potoos and nightjars both had relatively small brains and telencephala, and Wulst volumes that are typical for similarly sized birds from other orders. The Oilbird had a large brain, telencephalon and Wulst, although these measures were not quite as large as those of the owls. This gradation of owl-like versus nightjar-like brains within caprimulgiforms has significant implications for understanding the evolution of stereopsis and the Wulst both within the order and birds in general.     

Abundance-range size relationships of breeding and wintering birds in Britain: a comparative analysis

Both breeding and wintering assemblages of birds in Britain exhibit positive interspecific relationships between population size and geographic range size, such that the average density of species is greater if they are more widely distributed Species in common to both assemblages, that is resident species, had greater population sizes, geographic range sizes, and densities in winter In contrast, whilst winter migrants had higher abundances than summer migrants, the range sizes of the former were disproportionately larger still, resulting in a lower density for species that only winter in Britain than for those that only breed Such differences aside, the overall form of the abundance-range size relationship is remarkably similar between the two assemblages and their constituent subsets of species     

Why migrate during the day: a comparative analysis of North American birds

Migration can take place primarily during the day or at night, or during both depending on the species. Why the timing of migration varies among species has been the object of much research but the underlying ecological processes are still unclear. Proximally, migration timing may be influenced by the timing of other activities or may be more prevalent in species that migrate over long distances. Adaptive scenarios emphasize the reduction in flight costs at night especially in smaller species and the advantages of travelling in groups during the day to locate staging sites more efficiently. I used phylogenetic independent contrasts to examine these hypotheses in all North American nesting birds. I uncovered 24 evolutionary transitions in migration timing, most of which involved a switch from nocturnal to diurnal migration. Few of these transitions involved a concomitant change in the timing of foraging habits or migration distance. However, species in diurnal clades were larger, travelled in larger flocks and were generally more sociable than their nocturnal counterparts. The results support the hypotheses that a reduction in flight costs and the ability to pool information from companions are associated with migration timing in North American bird species.     

Life-history variation within a taxon: a comparative analysis of birds and mammals

Most studies on life-history evolution discuss the necessity of distinguishing between extrinsic and intrinsic sources of variability in life-history traits. I use log/log plots of yearly neonate production in daugters (b) versus adult mortality (Ma) for 75 bird species and 88 mammal species to compare graphically life-history "fields" arranged by these selective forces along a "slow-fast continuum". Under the assumptions of steady-state and linear relationship between adult mortality and reproductive effort, as well as between juvenile survival and relative neonate weight, it is possible to place additional axes in the two-dimentional plot, and to predict covariations among demographic and individual growth traits. The functional regression analysis shows, that the assumptions are completely fulfilled, at least for birds, but mammals show nonlinear relationship between adult mortality and reproductive effort. This can be explained by peculiarities of metabolism and parental care in small mammals with high reproductive output. Hence, for birds the axis of relative neonate weight approximately coincides in direction with the juvenile survivorship axis, but this is not a case for mammals. In both taxa, the relative neonate weight is an invariant in relation to fecundity and adult mortality (but not in relation to adult body weight). This important feature, together with other intrinsic (energetic and phylogenetic) constraints, explains well-documented close covariations among traits, even when the effect of body size is factored out. It is argued that life-history and body size variations in birds and mammals mainly depend on a pattern of temporal resource deficiency, although this impact cannot be separated from that of extrinsic juvenile mortality.     

Insights from comparative analyses of aging in birds and mammals

Many laboratory models used in aging research are inappropriate for understanding senescence in mammals, including humans, because of fundamental differences in life history, maintenance in artificial environments, and selection for early aging and high reproductive rate. Comparative studies of senescence in birds and mammals reveal a broad range in rates of aging among a variety of taxa with similar physiology and patterns of development. These comparisons suggest that senescence is a shared property of all vertebrates with determinate growth, that the rate of senescence has been modified by evolution in response to the potential life span allowed by extrinsic mortality factors, and that most variation among species in the rate of senescence is independent of commonly ascribed causes of aging, such as oxidative damage. Individuals of potentially long‐lived species, particularly birds, appear to maintain high condition to near the end of life. Because most individuals in natural populations of such species die of aging‐related causes, these populations likely harbor little genetic variation for mechanisms that could extend life further, or these mechanisms are very costly. This, and the apparent evolutionary conservatism in the rate of increase in mortality with age, suggests that variation in the rate of senescence reflects fundamental changes in organism structure, likely associated with the rate of development, rather than physiological or biochemical processes influenced by a few genes. Understanding these evolved differences between long‐lived and short‐lived organisms would seem to be an essential foundation for designing therapeutic interventions with respect to human aging and longevity.     

Neural sites and pathways regulating food intake in birds: a comparative analysis to mammalian systems

The paper reviews hypotheses explaining the regulation of food intake in mammals that have addressed specific anatomical structures in the brain. An hypothesis, poikilostasis, is introduced to describe multiple, homeostatic states whereby the regulation of metabolism and feeding occur in birds. Examples are given for both wild and domestic avian species, illustrating dynamic shifts in homeostasis responsible for the changes in body weights that are seen during the course of an annual cycle or by a particular strain of bird. The following neural structures are reviewed as each has been shown to affect food intake in birds or in mammals: ventromedial hypothalamic nucleus (n.), lateral hypothalamic area, paraventricular hypothalamic n., n. tractus solitarius and area postrema, amygdala, parabrachial n., arcuate n. and bed n. of the stria terminalis. Two neural pathways are described which have been proposed to regulate feeding. The trigeminal sensorimotor pathway is the most complete neural pathway characterized for this behavior and encompasses the mechanics of pecking, grasping and mandibulating food particles from the tip of the bill to the back of the buccal cavity. A second pathway, the visceral forebrain system (VFS), affects feeding by regulating metabolism and the balance of the autonomic nervous system. Wild, migratory birds are shown to exhibit marked changes in body weight which are hypothesized to occur due to shifts in balance between the sympathetic and parasympathetic nervous systems. Domestic avian species, selected for a rapid growth rate, are shown to display a dominance of the parasympathetic nervous system. The VFS is the neural system proposed to effect poikilostasis by altering the steady state of the autonomic nervous system in aves and perhaps is applicable to other classes of vertebrates as well.     

A comparative study of aerodynamic function and flexural stiffness of outer tail feathers in birds

To transmit aerodynamic forces to the body, tail feathers should be stiff to resist lift forces with minimum deformation. Because aerodynamic theory predicts that such feathers do not produce lift forces beyond the point of the maximum continuum width of the tail, species with deeply forked tails should not require stiff outer rectrices distal to that point. I tested this prediction by comparing the relative thickness of the outer rectrix rachis between species with deeply forked tails to those with triangular or shallowly forked tails. Eleven pairs of closely related species belonging to families Fregatidae, Phalacrocoracidae, Accipitridae, Sternidae, Caprimulgidae, Trochilidae, Coraciidae, Tyrannidae, Cotingidae, and Hirundinidae were compared. All but one of the phylogenetically independent comparisons showed that the species with triangular or shallowly forked tails have higher relative rachis thickness than their deeply forked relatives. In addition, nine out of eleven of the species with deeply forked tails showed a proportionately greater increase in relative rachis thickness from distal to proximal parts of the feather. In contrast, triangular and shallowly forked tails showed an approximately linear relation between relative rachis thickness and relative rachis length. These results considered together are consistent with the idea that the distal part of outer rectrix rachis in species with deeply forked tails has not been selected to resist lift forces and may be adaptively reduced to attenuate the costs of a hypertrophied ornament.     

A comparative methodical study of the faecal steroid analysis on birds: looking for a valid method of testosterone determination

In a comparative study, a relatively simple and high sensitivity method was developed for analysis of testosterone-equivalent(s) in the faeces of different bird species. To determine the recovery of extractions and purifications, tritium-labelled testosterone was added to the wet samples. Then the samples were treated with sodium dodecil sulphate (SDS), an emulsificator to "open-up" the complex, lipid-coated particles of faecal samples. This emulsification resulted in the decrease of the quantity of interfering substances after diethyl-ether extraction and the linearity of the measured testosterone equivalents from aliquots in the range of 2 and 10 mg of faeces. In the RIA, we applied a group specific polyclonal testosterone antibody which cross-reacted with reduced metabolites and at a certain level with sulphate conjugates as well. The use of Helix enzymes did not modified significantly the results of the analysis relating to a low level of conjugated androgens in the faecal extracts. The biological validity of the method was tested on domestic cockerels, where between the plasma and faecal testosterone values a four hours phase shift was observed, with a correlation of 0.6355. This method is suitable for "non invasive", behavioural-ethological studies.     

Variation in mating system among birds: ecological basis revealed by hierarchical comparative analysis of mate desertion

Since most bird species are socially monogamous, variation among species in social mating systems is determined largely by variation in the frequency of mate desertion. Mate desertion is expected to occur when the benefits, in terms of additional reproductive opportunities, outweigh the costs, in terms of reduced reproductive success from the present brood. However, despite much research, the relative importance of costs and benefits in explaining mating system variation is not well understood. Here, we investigate this problem using a comparative method. We analyse changes in the frequency of mate desertion at different phylogenetic levels. Differences between orders and families in the frequency of desertion are negatively associated with changes in the potential costs of desertion, but are not associated with changes in the potential benefits of desertion. Conversely, differences among genera and species in the frequency of desertion are positively associated with increases in the potential benefits of desertion, but not with changes in the potential costs of desertion. Hence, we suggest that mate desertion in birds originates through a combination of evolutionary predisposition and ecological facilitation. In particular, ancient changes in life-history strategy determine the costs of desertion and predispose certain lineages to polygamy, while contemporary changes in the distribution of resources determine the benefits of desertion and thereby the likelihood that polygamy will be viable within these lineages. Thus, monogamy can arise via two very different evolutionary pathways. Groups such as albatrosses (Procellariidae) are constrained to social monogamy by the high cost to desertion, irrespective of the potential benefits. However, in groups such as the accentors (Prunellidae), which are predisposed to desertion, monogamy occurs only when the benefits of desertion are very limited. These conclusions emphasise the additional power which a hierarchical approach contributes to the modern comparative method.