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In species that produce broods of multiple offspring, parents need to partition resources among simultaneously growing neonates that often differ in growth requirements. In birds, multiple ovarian follicles develop inside the female at the same time, resulting in a trade-off of resources among them and potentially limiting maternal ability for sex-specific allocation. We compared resource acquisition among oocytes in relation to their future sex and ovulation order in two populations of house finches with contrasting sex-biased maternal strategies. In a native Arizona population, where mothers do not bias offspring sex in relation to ovulation order, the male and female oocytes did not show sex-specific trade-offs of resources during growth and there was no evidence for spatial or temporal segregation of male and female oocytes in the ovary. In contrast, in a recently established Montana population where mothers strongly bias offspring sex in relation to ovulation order, we found evidence for both intra-sexual trade-offs among male and female oocytes and sex-specific clustering of oocytes in the ovary. We discuss the importance of sex-specific resource competition among offspring for the evolution of sex-ratio adjustment and sex-specific maternal resource allocation. [ABSTRACT FROM AUTHOR]

Comparative analyses of avian population fluctuations have shown large interspecific differences in population variability that have been difficult to relate to variation in general ecological characteristics. Here we show that interspecific variation in demographic stochasticity, caused by random variation among individuals in their fitness contributions, can be predicted from a knowledge of the species' position along a "slow-fast" gradient of life-history variation, ranging from high reproductive species with short life expectancy at one end to species that often produce a single offspring but survive well at the other end of the continuum. The demographic stochasticity decreased with adult survival rate, age at maturity, and generation time or the position of the species toward the slow end of the slow-fast life-history gradient. This relationship between life-history characteristics and demographic stochasticity was related to interspecific differences in the variation among females in recruitment as well as to differences in the individual variation in survival. Because reproductive decisions in birds are often subject to strong natural selection, our results provide strong evidence for adaptive modifications of reproductive investment through life-history evolution of the influence of stochastic variation on avian population dynamics. [ABSTRACT FROM AUTHOR]

Maternal phenotype and maternal environment can profoundly affect the phenotype and fitness of offspring. Yet the causes of variation in such maternal effects are rarely known. Embryos in avian eggs cannot develop without being incubated and this creates an opportunity for maternal control of duration and onset of offspring development. However, females might adjust the start of incubation (e.g., coincident with the first egg or delayed until after egg-laying) in response to environmental conditions that they experience at the time of breeding. We studied two populations of the house finch (Carpodacus mexicanus) that breed at the climatic extremes of the species' geographical range (Montana and Alabama) and found that in both populations, the timing of incubation onset was closely associated with the bias in the sequence in which male and female eggs were laid within a clutch. When females started incubation with the first egg, they produced sons and daughters in highly biased sequence, when females delayed the onset of incubation until after the egg-laying, the sequence of sons and daughters was not biased. Because in both populations, onset of incubation was associated with the ambient temperature, these results emphasize that maternal effects on offspring can be influenced by ecological conditions experienced by parental generation. [ABSTRACT FROM AUTHOR]

Presents a study which examined the evolution of sexual size dimorphism in the house finch. Materials and methods; Results; Discussion.

Discusses avian studies that demonstrated the importance of an organismal perspective on the expression of sexual displays and preferences. Investigation on the forces behind the evolution of sexual displays; Solution for the problem presented by context-dependence of the benefits obtained from mate choice; Implications for the coevolution of references and preferred traits of males and females.

Examines the relationship between song and plumage elaborations in cardueline finches. Relation of both song complexity and carotenoid-based plumage traits to overall individual condition; Different signaling strategies in variable environments; Effects of potential covariates on song attributes of cardueline finches.

Presents a study that examined the divergence in sexual dimorphism of adults between established Montana and Alabama populations of the house finch Carpodacus mexicanus. Attributes to population differences in growth of males and females; Comparisons of the ontogeny of males and females in diverged populations that differ in sexual dimorphism of adults.

Examines the pattern of covariation in avian life history strategies along an elevational gradient. Patterns in life history strategies; Suggestions on the negative relationship between offspring number duration and duration of parental care; Effects of developmental cost variations and maintenance of secondary sexual traits.

Examines the ontogeny of sexual size dimorphism in an isolated population of the house finch Carpodacus mexicanus. Identification of sex; Growth patterns in males and females; Sexual dimorphism for bill traits in relation to age.

Presents information on a study which examined the evolution of sexual size dimorphism in a passerine bird called the house finch. Discussion on sexual dimorphism; Methodology; Results and discussion.

Presents information on a study which analyzed the evolution of sexual morphological variance-covariance matrices for seven house finch population. Methodology of the study; Results and discussion on the study.

Presents a study which determined the role of current net selection in shaping and maintaining contemporary sexual dimorphism in an established population of the house finch, Carpodacus mexicanus in Montana. Methodology; Results; Discussion.

Investigates the sensitivity of mandibular development of shrews to stress. Increase in asymmetry for shrews born under stressful conditions; Association of asymmetry with lower physiological conditions; Sensitivity of males to stressful conditions; Relation between differential sensitivity to developmental morphological integration.

Summary Tarsal spurs play an important role in intrasexual competition for females among male wild turkeys (Meleagris gallopavo ). Thus variation in spur development may have important fitness consequences. Fitness correlates of spur development were studied in a free-living population of wild turkeys and it was found that heavier males and males with longer beards had longer spurs. Males that had longer spurs spent more time on display areas during the breeding season and less time moving among these areas compared to males with shorter spurs, independently of their body mass. Otherwise ideally symmetrical spurs showed fluctuating asymmetry between left and right tarsi, the degree of absolute asymmetry decreased with spur length in adults, but not in subadults, and males that survived at least one winter had more symmetrical spurs compared to males that did not. We conclude that if the ability to produce symmetrical spurs has a genetic basis, then spur length and spur asymmetry could reliably indicate individual quality and that these traits are under directional selection for increased size and symmetry in wild turkeys. [ABSTRACT FROM AUTHOR]

Examines ecological and behavioral factors that affect variation in seasonal ranges of eastern wild turkeys. Habitat distribution; Age; Sex; Body mass.

Focuses on sexual dichromatism in birds with emphasis on the importance of nest predation and nest location for females as opposed to males. Cause of sexual dichromatism in birds; Significance of more concealed nest sites.

Studies nest site fidelity of female Wild Turkeys. Background on the study; Site fidelity, age, and nest predation; Reproductive performance and site fidelity; Spring dispersal; Previous nesting habitat.

Studies the influence of habitat characteristics on the evolution of song attributes in Phylloscopus and Hippolais warblers. Factors constraining the amount of environmental variation in song properties; Effects of body mass and habitat structure on variation in song structure; Correlation of habitat structure with temporal characteristics of songs.

Secondary sexual traits are thought to be costly to produce and maintain, and the allocation of energy to sexual traits could result in reduced investment in traits associated with growth and basic maintenance. It has been suggested that the trade-off between sexual traits and growth and maintenance traits should manifest itself in the correlation between development of sexual traits and resistance to environmental variability. Interspecifically, this hypothesis predicts that species with greater sexual ornamentation and dimorphism should be able to tolerate a narrower width of environmental condition compared torelated species with less developed sexual traits. We tested this prediction by examining whether sexual dimorphism in cardueline fincheswas negatively associated with the width of elevational range occupied during breeding. We assumed that the range in elevations where breeding can occur represents a measure of tolerance of environmental variability. We found a positive rather than a negative relationship between breeding range and sexual dimorphism. Finches that were capableof breeding over a large range of elevations were also more dimorphic in plumage. Possible explanations for the observed relationship between sexual dichromatism and ecological breadth could include interspecific differences in food availability and foraging ability, as wellas variation in energy required for baseline metabolism. [ABSTRACT FROM AUTHOR]

Elevation has long been considered a major influence on the evolution of life-history traits. Most elevation-induced variation in life history traits has been attributed to changes in climate, duration of breeding season, predation, and food limitation. I use a phylogenetic approach to show that life histories are closely associated with breeding elevation in extant cardueline finches. Finches at high elevations had smaller clutches, fewer broods, and longer incubation periods. Neither food limitation nor nest predation appear to readily account for this strong elevational variation in cardueline life histories. However, juvenile survival may be greater at higher elevations as a result of prolonged parental care and shorter natal dispersal and can potentially compensate for reduced fecundity in high-elevation finches. [ABSTRACT FROM AUTHOR]

We report the first case of mycoplasmosis in the western range of the House Finch (Carpodacus mexicanus). This disease originated in the eastern United States and has been previously documented only in eastern introduced House Finch populations where it reached epizootic proportions causing extensive and widespread mortality. Documentation of this disease in western Montana suggests that previously disjunct eastern and western populations of House Finches are now mixing in the northern part of their range. More importantly, as native House Finches are highly susceptible to this novel pathogen, western populations may now be at risk of high mortality, similar to that experienced by non-native eastern populations. Close monitoring of this disease in the western part of the House Finch range will provide important insight into the dynamics of the emerging disease and evolution of resistance to the pathogen. Received 10 May 2002, accepted 5 February 2003. [ABSTRACT FROM AUTHOR]

Most species of birds can lay only one egg per day until a dutch is complete, and the order in which eggs are laid often has strong and sex-specific effects on offspring growth and survival In two recently established populations of the house finch (Carpodacus mexicanus) in Montana and Alabama, breeding females simultaneously adjusted the sex and growth of offspring in relation to their position in the laying order, thereby reducing the mortality of sons and daughters by 10 to 20% in both environments. We show experimentally that the reduction in mortality is produced by persistent and sex-specific maternal effects on the growth and morphology of offspring. These strong parental effects may have facilitated the rapid adaptive divergence among populations of house finches. [ABSTRACT FROM AUTHOR]

All organisms depend on input of exogenous compounds that cannot be internally produced. Gain and loss of such dependencies structure ecological communities and drive species' evolution, yet the evolution of mechanisms that accommodate these variable dependencies remain elusive. Here, we show that historical cycles of gains and losses of external dependencies in avian carotenoid-producing networks are linked to their evolutionary diversification. This occurs because internalization of metabolic controls—produced when gains in redundancy of dietary inputs coincide with increased branching of their derived products—enables rapid and sustainable exploration of an existing network by shielding it from environmental fluctuations in inputs. Correspondingly, loss of internal controls constrains evolution to the rate of the gains and losses of dietary precursors. Because internalization of a network's controls necessarily bridges diet-specific enzymatic modules within a network, it structurally links local adaptation and continuous evolution even for traits fully dependent on contingent external inputs. The mechanisms that accommodate variable external dependencies in evolution are not clear. Here, the authors show that switches between external and internal metabolic controls of carotenoid-producing networks in birds are linked to shifts in evolutionary rates, with internalization of control resulting in bursts of evolutionary diversification. [ABSTRACT FROM AUTHOR]

1. Ecological convergence in morphology among taxa of distinct evolutionary histories is a common illustration of the efficacy of natural selection. Ecological convergence is often enabled by functional redundancy of complex morphological structures, such that modification of existing morphologies in response to similar functional requirements can lead to the development and evolution of morphological diversity. Thus, studies of the mechanisms that enable the development of similar adaptations in taxa with distinct morphologies provide important insights into both the evolution of past adaptations and patterns of future evolutionary divergence. 2. Here, we examine mechanisms that have enabled ecological convergence in foraging morphology among four geographically isolated and morphologically distinct populations of shrews: south-eastern Arizona and north-central New Mexico populations of the montane shrews ( Sorex monticolus) and northern California and north-central Montana populations of the vagrant shrew ( S. vagrans). 3. We show that despite overlap in diet, populations had distinct skeletal and muscular morphologies of the mandible. This association between ecological convergence and morphological uniqueness among populations was enabled by versatility of foraging morphologies that generated similar functional outputs. 4. In addition, we found that populations exhibited unique skeletal and muscular correlations with diet suggesting that distinct muscular and morphological components of the complex foraging apparatus can be used for a particular resource. This result corroborates a previous finding that extensive modularity in mandibular development allows diverse morphologies to generate equivalent functions and utilize similar resources across taxa. 5. Synthesis. We conclude that the observed functional and ecological convergences resulted from population-specific musculoskeletal interactions, and suggest that the differences in skeletal and muscular morphologies observed among these populations reflect evolved differences in plasticity of the skeletal and muscular components of the mandible. [ABSTRACT FROM AUTHOR]

Diversity in organismal forms among taxa is thought to reflect distinct selection pressures across environments. The central assumption underlying this expectation is that taxa experiencing similar selection have similar response to that selection. However, because selection acts on trait function, taxa similarity in selection response depends crucially on the relationship between function and morphology. Further, when a trait consists of multiple parts, changes in function in response to selection can result from modification of different parts, and adaptation to the same environment might result in functional but not morphological similarity. Here, we address the extent to which functional and morphological diversity in masticatory apparatus of soricid shrews reflects a shared ecological characteristic of their diet type. We examine the factors limiting morphological variation across shrew species by assessing the relative contribution of trait function (biomechanics of the jaw), ecology, and phylogeny to species similarity in mandibular traits. We found that species that shared diet type were functionally but not morphologically similar. The presence of multiple semi-independently varying traits enabled functional equivalence of composite foraging morphologies and resulted in variable response to selection exerted by similar diet. We show that functional equivalence of multiple morphologies enabled persistence of differences in habitat use (e.g., habitat moisture and coverage) among species that specialize on the same diet. We discuss the importance of developmental and functional integration among traits for evolutionary diversification of morphological structures that generate equivalent functions. [ABSTRACT FROM AUTHOR]

The evolutionarily persistent and widespread use of carotenoid pigments in animal coloration contrasts with their biochemical instability. Consequently, evolution of carotenoid-based displays should include mechanisms to accommodate or limit pigment degradation. In birds, this could involve two strategies: (i) evolution of a moult immediately prior to the mating season, enabling the use of particularly fast-degrading carotenoids and (ii) evolution of the ability to stabilize dietary carotenoids through metabolic modification or association with feather keratins. Here, we examine evolutionary lability and transitions between the two strategies across 126 species of birds. We report that species that express mostly unmodified, fast-degrading, carotenoids have pre-breeding moults, and a particularly short time between carotenoid deposition and the subsequent breeding season. Species that expressed mostly slow-degrading carotenoids in their plumage accomplished this through increased metabolic modification of dietary carotenoids, and the selective expression of these slow-degrading compounds. In these species, the timing of moult was not associated with carotenoid composition of plumage displays. Using repeated samples from individuals of one species, we found that metabolic modification of dietary carotenoids significantly slowed their degradation between moult and breeding season. Thus, the most complex and colourful ornamentation is likely the most biochemically stable in birds, and depends less on ecological factors, such as moult timing and migration tendency. We suggest that coevolution of metabolic modification, selective expression and biochemical stability of plumage carotenoids enables the use of unstable pigments in long-term evolutionary trends in plumage coloration. [ABSTRACT FROM AUTHOR]

Study of morphogenesis and its regulation requires analytical tools that enable simultaneous assessment of processes operating at cellular level, such as synthesis of transcription factors (TF), with their effects at the tissue scale. Most current studies conduct histological, cellular and immunochemical (IHC) analyses in separate steps, introducing inevitable biases in finding and alignment of areas of interest at vastly distinct scales of organization, as well as image distortion associated with image repositioning or file modifications. These problems are particularly severe for longitudinal analyses of growing structures that change size and shape. Here we introduce a python-based application for automated and complete whole-slide measurement of expression of multiple TFs and associated cellular morphology. The plugin collects data at customizable scale from the cell-level to the entire structure, records each data point with positional information, accounts for ontogenetic transformation of structures and variation in slide positioning with scalable grid, and includes a customizable file manager that outputs collected data in association with full details of image classification (e.g., ontogenetic stage, population, IHC assay). We demonstrate the utility and accuracy of this application by automated measurement of morphology and associated expression of eight TFs for more than six million cells recorded with full positional information in beak tissues across 12 developmental stages and 25 study populations of a wild passerine bird. Our script is freely available as an open-source Fiji plugin and can be applied to IHC slides from any imaging platforms and transcriptional factors., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2024 The Authors. Published by Elsevier B.V.)

Robustness against environmental fluctuations within an adaptive state should preclude exploration of new adaptive states when the environment changes. Here, we study transitions between adaptive associations of feather structure and carotenoid uptake to understand how robustness and evolvability can be reconciled. We show that feather modifications induced by unfamiliar carotenoids during a range expansion are repeatedly converted into precise coadaptations of feather development and carotenoid accommodation as populations persist in a region. We find that this conversion is underlain by a uniform and coordinated increase in the sensitivity of feather development to local carotenoid uptake, indicative of cooption and modification of the homeostatic mechanism that buffers feather growth in the evolution of new adaptations. Stress-buffering mechanisms are well placed to alternate between robustness and evolvability and we suggest that this is particularly evident in adaptations that require close integration between widely fluctuating external inputs and intricate internal structures.

Evolution of adaptive phenotypic flexibility requires a system that can dynamically restore and update a functional phenotype in response to environmental change. The architecture of such a system evolves under the conflicting demands of versatility and robustness, and resolution of these demands should be particularly evident in organisms that require external inputs for reiterative trait production within a generation, such as in metabolic networks that underlie yearly acquisition of diet-dependent coloration in birds. Here, we show that a key structural feature of carotenoid networks-redundancy of biochemical pathways-enables these networks to translate variable environmental inputs into consistent phenotypic outcomes. We closely followed life-long changes in structure and utilization of metabolic networks in a large cohort of free-living birds and found that greater individual experience with dietary change between molts leads to wider occupancy of the metabolic network and progressive accumulation of redundant pathways in a functionally active network. This generated a regime of emergent buffering whereby greater dietary experience was mechanistically linked to greater robustness of resulting traits and an increasing ability to retain and implement previous adaptive solutions. Thus, experience-related buffering links evolvability and robustness in carotenoid-metabolizing networks and we argue that this mechanistic principle facilitates the evolution of phenotypic flexibility., (© 2018 The Author(s). Evolution © 2018 The Society for the Study of Evolution.)

Evolution of adaptation requires both generation of novel phenotypic variation and retention of a locally beneficial subset of this variation. Such retention can be facilitated by genetic assimilation, the accumulation of genetic and molecular mechanisms that stabilize induced phenotypes and assume progressively greater control over their reliable production. A particularly strong inference into genetic assimilation as an evolutionary process requires a system where it is possible to directly evaluate the extent to which an induced phenotype is progressively incorporated into preexisting developmental pathways. Evolution of diet-dependent pigmentation in birds-where external carotenoids are coopted into internal metabolism to a variable degree before being integrated with a feather's developmental processes-provides such an opportunity. Here we combine a metabolic network view of carotenoid evolution with detailed empirical study of feather modifications to show that the effect of physical properties of carotenoids on feather structure depends on their metabolic modification, their environmental recurrence, and biochemical redundancy, as predicted by the genetic assimilation hypothesis. Metabolized carotenoids caused less stochastic variation in feather structure and were more closely integrated with feather growth than were dietary carotenoids of the same molecular weight. These patterns were driven by the recurrence of organism-carotenoid associations: commonly used dietary carotenoids and biochemically redundant derived carotenoids caused less stochastic variation in feather structure than did rarely used or biochemically unique compounds. We discuss implications of genetic assimilation processes for the evolutionary diversification of diet-dependent animal coloration.

The network of the interactions among genes, proteins, and metabolites delineates a range of potential phenotypic diversifications in a lineage, and realized phenotypic changes are the result of differences in the dynamics of the expression of the elements and interactions in this deterministic network. Regulatory mechanisms, such as hormones, mediate the relationship between the structural and dynamic properties of networks by determining how and when the elements are expressed and form a functional unit or state. Changes in regulatory mechanisms lead to variable expression of functional states of a network within and among generations. Functional properties of network elements, and the magnitude and direction of evolutionary change they determine, depend on their location within a network. Here, we examine the relationship between network structure and the dynamic mechanisms that regulate flux through a metabolic network. We review the mechanisms that control metabolic flux in enzymatic reactions and examine structural properties of the network locations that are targets of flux control. We aim to establish a predictive framework to test the contributions of structural and dynamic properties of deterministic networks to evolutionary diversifications., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Covariation among organismal traits is nearly universal, occurring both within and among species (static and evolutionary allometry, respectively). If conserved developmental processes produce similarity in static and evolutionary allometry, then when species differ in development, it should be expressed in discordance between allometries. Here, we investigate whether rapidly evolving developmental processes result in discordant static and evolutionary allometries attributable to trade-offs in resource acquisition, allocation, or growth across 30 species of aquatic beetles. The highly divergent sperm phenotypes of these beetles might be an important contributor to allometric evolution of testis and accessory gland mass through altered requirements for the production of sperm and seminal fluids. We documented extensive discordance between static and evolutionary allometries, indicating that allometric relationships are flexibly modified over short time periods but subject to constraint over longer time spans. Among species, sperm phenotype did not influence relative investment in accessory glands but was weakly associated with investment in testes. Furthermore, except when sperm were long and simple, sperm phenotype was not associated with species-specific modification of the allometry of testis/accessory gland mass and body size. Our results demonstrate the utility of allometric discordance to infer species differences in the provisioning and growth of concurrently developing traits.

The complexity of organismal organization channels and accommodates novel genomic and developmental modifications. Here, I extend this perspective to suggest that emergent processes that dominate homeostasis-co-option, re-use, and recombination of accumulated elements-can create configurations and dependencies among these elements that strongly reduce the number of evolutionary steps needed for the evolution of precise novel adaptations. Evolutionary retention and environmental matching of such configurations are further facilitated when they include elements of homeostasis that are responsive to particular environmental cues. I apply this perspective to the study of evolution of sex-biased egg-laying in birds, a phenomenon that combines precision, complexity, context-dependency, and reversibility. I show that homeostatic hitchhiking can overcome the main difficulty in the evolution of this adaptation-the perceived necessity of de novo co-evolution of oogenesis, sex-determination, and order of ovulation in each environmental context-something that would require unrealistic expectations of evolutionary rates and population sizes and is not a desirable outcome for a process that needs to retain substantial environmental sensitivity. First, I explain the rationale behind the homeostatic-hitchhiking hypothesis and outline its predictions specifically for studies of sex-bias in order of egg-laying. Second, I show that a combination of self-regulatory and emergent processes and ubiquitous re-use of conserved growth factors make oogenesis particularly amendable to homeostatic hitchhiking. Third, I review empirical evidence for this mechanism in the rapid evolution of adaptive sex-biased order of egg-laying that accompanied colonization of North America by the house finch (Carpodacus mexicanus).

In complex organisms, neutral evolution of genomic architecture, associated compensatory interactions in protein networks and emergent developmental processes can delineate the directions of evolutionary change, including the opportunity for natural selection. These effects are reflected in the evolution of developmental programmes that link genomic architecture with a corresponding functioning phenotype. Two recent findings call for closer examination of the rules by which these links are constructed. First is the realization that high dimensionality of genotypes and emergent properties of autonomous developmental processes (such as capacity for self-organization) result in the vast areas of fitness neutrality at both the phenotypic and genetic levels. Second is the ubiquity of context- and taxa-specific regulation of deeply conserved gene networks, such that exceptional phenotypic diversification coexists with remarkably conserved generative processes. Establishing the causal reciprocal links between ongoing neutral expansion of genomic architecture, emergent features of organisms' functionality, and often precisely adaptive phenotypic diversification therefore becomes an important goal of evolutionary biology and is the latest reincarnation of the search for a framework that links development, functioning and evolution of phenotypes. Here I examine, in the light of recent empirical advances, two evolutionary concepts that are central to this framework-natural selection and inheritance-the general rules by which they become associated with emergent developmental and homeostatic processes and the role that they play in descent with modification.

The social environment is a critical determinant of fitness and, in many taxa, is shaped by an individual's behavioral discrimination among social contexts, suggesting that animals can actively influence the selection they experience. In competition to attract females, males may modify sexual selection by choosing social environments in which they are more attractive relative to rivals. Across the population, such behaviors should influence sexual selection patterns by altering the relationship between male mating success and sexual ornament elaboration. Here we use network analysis to examine patterns of male social behavior in relation to plumage ornamentation and mating success in a free-living population of house finches. During the nonbreeding season, less elaborate males changed associations with distinct social groups more frequently, compared to more elaborate males that showed greater fidelity to a single social group. By the onset of pair formation, socially labile males effectively increased their attractiveness relative to other males in the same flocks. Consequently, males that frequently moved between social groups had greater pairing success than less social individuals with equivalent sexual ornamentation. We discuss these results in relation to conditional mating tactics and the role of social behavior in evolutionary change by sexual selection.

The link between adaptation and evolutionary change remains the most central and least understood evolutionary problem. Rapid evolution and diversification of avian beaks is a textbook example of such a link, yet the mechanisms that enable beak's precise adaptation and extensive adaptability are poorly understood. Often observed rapid evolutionary change in beaks is particularly puzzling in light of the neo-Darwinian model that necessitates coordinated changes in developmentally distinct precursors and correspondence between functional and genetic modularity, which should preclude evolutionary diversification. I show that during first 19 generations after colonization of a novel environment, house finches (Carpodacus mexicanus) express an array of distinct, but adaptively equivalent beak morphologies-a result of compensatory developmental interactions between beak length and width in accommodating microevolutionary change in beak depth. Directional selection was largely confined to the elimination of extremes formed by these developmental interactions, while long-term stabilizing selection along a single axis-beak depth-was mirrored in the structure of beak's additive genetic covariance. These results emphasize three principal points. First, additive genetic covariance structure may represent a historical record of the most recurrent developmental and functional interactions. Second, adaptive equivalence of beak configurations shields genetic and developmental variation in individual components from depletion by natural selection. Third, compensatory developmental interactions among beak components can generate rapid reorganization of beak morphology under novel conditions and thus greatly facilitate both the evolution of precise adaptation and extensive diversification, thereby linking adaptation and adaptability in this classic example of Darwinian evolution.

When faced with changing environments, organisms rapidly mount physiological and behavioural responses, accommodating new environmental inputs in their functioning. The ubiquity of this process contrasts with our ignorance of its evolutionary significance: whereas within-generation accommodation of novel external inputs has clear fitness consequences, current evolutionary theory cannot easily link functional importance and inheritance of novel accommodations. One hundred and twelve years ago, J. M. Baldwin, H. F. Osborn and C. L. Morgan proposed a process (later termed the Baldwin effect) by which non-heritable developmental accommodation of novel inputs, which makes an organism fit in its current environment, can become internalized in a lineage and affect the course of evolution. The defining features of this process are initial overproduction of random (with respect to fitness) developmental variation, followed by within-generation accommodation of a subset of this variation by developmental or functional systems ('organic selection'), ensuring the organism's fit and survival. Subsequent natural selection sorts among resultant developmental variants, which, if recurrent and consistently favoured, can be inherited when existing genetic variance includes developmental components of individual modifications or when the ability to accommodate novel inputs is itself heritable. Here, I show that this process is consistent with the origin of novel adaptations during colonization of North America by the house finch. The induction of developmental variation by novel environments of this species's expanding range was followed by homeostatic channelling, phenotypic accommodation and directional cross-generational transfer of a subset of induced developmental outcomes favoured by natural selection. These results emphasize three principal points. First, contemporary novel adaptations result mostly from reorganization of existing structures that shape newly expressed variation, giving natural selection an appearance of a creative force. Second, evolutionary innovations and maintenance of adaptations are different processes. Third, both the Baldwin and parental effects are probably a transient state in an evolutionary cycle connecting initial phenotypic retention of adaptive changes and their eventual genetic determination and, thus, the origin of adaptation and evolutionary change.

As is the case with any metaphor, parental effects mean different things to different biologists--from developmental induction of novel phenotypic variation to an evolved adaptation, and from epigenetic transference of essential developmental resources to a stage of inheritance and ecological succession. Such a diversity of perspectives illustrates the composite nature of parental effects that, depending on the stage of their expression and whether they are considered a pattern or a process, combine the elements of developmental induction, homeostasis, natural selection, epigenetic inheritance and historical persistence. Here, we suggest that by emphasizing the complexity of causes and influences in developmental systems and by making explicit the links between development, natural selection and inheritance, the study of parental effects enables deeper understanding of developmental dynamics of life cycles and provides a unique opportunity to explicitly integrate development and evolution. We highlight these perspectives by placing parental effects in a wider evolutionary framework and suggest that far from being only an evolved static outcome of natural selection, a distinct channel of transmission between parents and offspring, or a statistical abstraction, parental effects on development enable evolution by natural selection by reliably transferring developmental resources needed to reconstruct, maintain and modify genetically inherited components of the phenotype. The view of parental effects as an essential and dynamic part of an evolutionary continuum unifies mechanisms behind the origination, modification and historical persistence of organismal form and function, and thus brings us closer to a more realistic understanding of life's complexity and diversity.

Differences in relative fitness of male and female offspring across ecological and social environments should favour the evolution of sex-determining mechanisms that enable adjustment of brood sex ratio to the context of breeding. Despite the expectation that genetic sex determination should not produce consistent bias in primary sex ratios, extensive and adaptive modifications of offspring sex ratio in relation to social and physiological conditions during reproduction are often documented. Such discordance emphasizes the need for empirical investigation of the proximate mechanisms for modifying primary sex ratios, and suggests epigenetic effects on sex-determining mechanisms as the most likely candidates. Birds, in particular, are thought to have an unusually direct opportunity to modify offspring sex ratio because avian females are heterogametic and because the sex-determining division in avian meiosis occurs prior to ovulation and fertilization. However, despite evidence of strong epigenetic effects on sex determination in pre-ovulatory avian oocytes, the mechanisms behind such effects remain elusive. Our review of molecular and cytological mechanisms of avian meiosis uncovers a multitude of potential targets for selection on biased segregation of sex chromosomes, which may reflect the diversity of mechanisms and levels on which such selection operates in birds. Our findings indicate that pronounced differences between sex chromosomes in size, shape, size of protein bodies, alignment at the meiotic plate, microtubule attachment and epigenetic markings should commonly produce biased segregation of sex chromosomes as the default state, with secondary evolution of compensatory mechanisms necessary to maintain unbiased meiosis. We suggest that it is the epigenetic effects that modify such compensatory mechanisms that enable context-dependent and precise adjustment of primary sex ratio in birds. Furthermore, we highlight the features of avian meiosis that can be influenced by maternal hormones in response to environmental stimuli and may account for the precise and adaptive patterns of offspring sex ratio adjustment observed in some species.

A tremendous diversity of avian color displays has stimulated numerous studies of natural and sexual selection. Yet, the developmental mechanisms that produce such diversification, and thus the proximate targets of selection pressures, are rarely addressed and poorly understood. In particular, because feathers are colored during growth, the dynamics of feather growth play a deterministic role in the variation in ornamentation. No study to date, however, has addressed the contribution of feather growth to the expression of carotenoid-based ornamentation. Here, we examine the developmental basis of variation in ornamental feather shapes in male house finches (Carpodacus mexicanus)-a species in which carotenoid displays are under strong natural and sexual selection. First, we use geometric morphometrics to partition the observed shape variation in fully grown feathers among populations, ages, degrees of elaboration, ornamental body parts, and individuals. Second, we use a biologically informed mathematical model of feather growth to predict variation in shape of ornamental feathers due to simulated growth rate, angle of helical growth of feather barbs, initial number of barb ridges, rate of addition of new barbs, barb diameter, and ramus-expansion angle. We find close concordance between among-individual variation in feather shape and hue of entire ornament, and show that this concordance can be attributed to a shared mechanism-growth rate of feather barbs. Predicted differences in feather shape due to rate of addition of barbs and helical angle of feather growth explained observed variation in ornamental area both among individuals and between populations, whereas differences in helical angle of growth and the number of barbs in the feather follicle explained differences in feather shape between ornamental parts and among males of different ages. The findings of a close association of feather growth dynamics and overall ornamentation identify the proximate targets of selection for elaboration of sexual displays. Moreover, the close association of feather growth and pigmentation not only can reinforce condition-dependence in color displays, but can also enable phenotypic and genetic accommodation of novel pigments into plumage displays providing a mechanism for the observed concordance of within-population developmental processes and between-population diversification of color displays.

Evolutionary diversifications are commonly attributed to the continued modifications of a conserved genetic toolkit of developmental pathways, such that complexity and convergence in organismal forms are assumed to be due to similarity in genetic mechanisms or environmental conditions. This approach, however, confounds the causes of organismal development with the causes of organismal differences and, as such, has only limited utility for addressing the cause of evolutionary change. Molecular mechanisms that are closely involved in both developmental response to environmental signals and major evolutionary innovations and diversifications are uniquely suited to bridge this gap by connecting explicitly the causes of within-generation variation with the causes of divergence of taxa. Developmental pathways of bone formation and a common role for bone morphogenetic proteins (BMPs) in both epigenetic bone remodeling and the evolution of major adaptive diversifications provide such opportunity. We show that variation in timing of ossification can result in similar phenotypic patterns through epigenetically induced changes in gene expression and propose that both genetic accommodation of environmentally induced developmental pathways and flexibility in development across environments evolve through heterochronic shifts in bone maturation relative to exposure to unpredictable environments. We suggest that such heterochronic shifts in ossification can not only buffer development under fluctuating environments while maintaining epigenetic sensitivity critical for normal skeletal formation, but also enable epigenetically induced gene expression to generate specialized morphological adaptations. We review studies of environmental sensitivity of BMP pathways and their regulation of formation, remodeling, and repair of cartilage and bone to examine the hypothesis that BMP-mediated skeletal adaptations are facilitated by evolved reactivity of BMPs to external signals. Surprisingly, no empirical study to date has identified the molecular mechanism behind developmental plasticity in skeletal traits. We outline a conceptual framework for future studies that focus on mediation of phenotypic plasticity in skeletal development by the patterns of BMP expression.

Examination of historical persistence of integration patterns provides an important insight into understanding the origin and evolution of complex traits. Specifically, the distinct effects of developmental and functional integration on the evolution of complex traits are often overlooked. Because patterns of functional integration are commonly shaped by selection exerted by the external environment, whereas patterns of developmental integration can be determined by relatively environment-independent selection for developmental homeostasis, examination of historical persistence of morphological integration patterns among species should reveal the relative importance of current selection in the evolution of complex traits. We compared historical persistence of integration patterns produced by current developmental versus ecological requirements by examining the evolution of complex mandibular structures in nine species of soricid shrews. We found that, irrespective of phylogenetic relatedness of species, patterns of developmental and functional integration were highly concordant, suggesting that strong selection for developmental homeostasis favors concordant channeling of both internal and external variation. Overall, our results suggest that divergence in mandible shape among species closely follows variation in functional demands and ecological requirements regardless of phylogenetic relatedness among species.