Tactile bill-tip organs in seabirds suggest conservation of a deep avian symplesiomorphy.

Birds' bills are their main tactile interface with the outside world. Tactile bill-tip organs associated with specialized foraging techniques are present in several bird groups, yet remain understudied in most clades. One example is Austrodyptornithes, the major seabird clade uniting Procellariiformes (albatrosses and petrels) and Sphenisciformes (penguins). Here, we describe the mechanoreceptor arrangement and neurovascular anatomy in the premaxillae of Austrodyptornithes. Using a wide phylogenetic sample of extant birds (361 species), we show that albatrosses and penguins exhibit complex tactile bill-tip anatomies, comparable to birds with known bill-tip organs, despite not being known to use tactile foraging. Petrels (Procellariidae, Hydrobatidae and Oceanitidae) lack these morphologies, indicating an evolutionary transition in bill-tip mechanosensitivity within Procellariiformes. The bill-tip organ in Austrodyptornithes may be functionally related to nocturnal foraging and prey detection under water, or courtship displays involving tactile stimulation of the bill. Alternatively, these organs may be vestigial as is likely the case in most palaeognaths (e.g. ostriches and emu). Ancestral state reconstructions fail to reject the hypothesis that the last common ancestor of Austrodyptornithes had a bill-tip organ; thus, tactile foraging may be ancestral for this major extant clade, perhaps retained from a deeper point in crown bird evolutionary history.

A rapid-response soft end effector inspired by the hummingbird beak.

Biology is a wellspring of inspiration in engineering design. This paper delves into the application of elastic instabilities-commonly used in biological systems to facilitate swift movement-as a power-amplification mechanism for soft robots. Specifically, inspired by the nonlinear mechanics of the hummingbird beak-and shedding further light on it-we design, build and test a novel, rapid-response, soft end effector. The hummingbird beak embodies the capacity for swift movement, achieving closure in less than [Formula: see text]. Previous work demonstrated that rapid movement is achieved through snap-through deformations, induced by muscular actuation of the beak's root. Using nonlinear finite element simulations coupled with continuation algorithms, we unveil a representative portion of the equilibrium manifold of the beak-inspired structure. The exploration involves the application of a sequence of rotations as exerted by the hummingbird muscles. Specific emphasis is placed on pinpointing and tailoring the position along the manifold of the saddle-node bifurcation at which the onset of elastic instability triggers dynamic snap-through. We show the critical importance of the intermediate rotation input in the sequence, as it results in the accumulation of elastic energy that is then explosively released as kinetic energy upon snap-through. Informed by our numerical studies, we conduct experimental testing on a prototype end effector fabricated using a compliant material (thermoplastic polyurethane). The experimental results support the trends observed in the numerical simulations and demonstrate the effectiveness of the bio-inspired design. Specifically, we measure the energy transferred by the soft end effector to a pendulum, varying the input levels in the sequence of prescribed rotations. Additionally, we demonstrate a potential robotic application in scenarios demanding explosive action. From a mechanics perspective, our work sheds light on how pre-stress fields can enable swift movement in soft robotic systems with the potential to facilitate high input-to-output energy efficiency.

Geometric morphometric analysis of beak shape of Columbimorphae (Columbas, Van, Mardin and Dönek).

Selective breeding over centuries has led to the emergence of numerous pigeon breeds from the single species Columba livia, showcasing unparalleled phenotypic diversity. In Eastern Turkey, Van pigeons originate, while Tumbler pigeon varieties thrive countrywide, reflecting local traditions. The avian beak, vital for survival, offers insights into domestication and traits influenced by natural selection. Geometric morphometrics, a shape analysis method, allows for a comprehensive examination of beak shapes among domestic pigeons, providing a nuanced understanding of their complexity. Understanding beak diversity in pigeons, especially those found in Turkey, enhances our knowledge of avian evolution and adaptation processes. The study utilized 48 skulls from 4 pigeon breeds, including wild rock pigeons and domestic pigeons of Tumbler, Mardin and Van breeds, all free from pathological lesions and adults. Geometric morphometric analyses of beak shape were conducted using dorsal and lateral photographs, with landmarks placed using tpsDig software. MorphoJ software facilitated procrustes analysis and principal component analysis (PCA) to assess morphological variability and differentiate pigeon breeds based on shape patterns, with significance set at p < 0.05. PCA revealed significant shape variations among pigeon breeds, with 47 principal components identified for lateral beak views and 36 principal components for dorsal views. Canonical variates analysis further distinguished morphological patterns among breeds, indicating distinct shape variations in both dorsal and lateral views, elucidating the unique characteristics of each breed's beak morphology. Our results demonstrate statistically significant differences in pigeon beak shape, particularly in the lateral view, confirming the importance of these variations (p < 0.05) and suggesting that such differences occur less than 5% of the time under the null hypothesis.

Bill shape variation in selected species in birds of prey.

At the top of many ecosystems, raptors, also known as birds of prey, hold major influence. They shape their surroundings through their powerful hunting skills and complex interactions with their environment. This study investigates the beak morphology of four prominent raptor species, Golden eagle (Aquila chrysaetos), Common buzzard (Buteo buteo), Peregrine falcon (Falco peregrinus) and Common kestrel (Falco tinnunculus), found in Türkiye. By employing geometric morphometric methods, we investigate shape variations in the beaks of these species to unravel the adaptive significance of their cranial structures. This analysis reveals distinct beak morphologies among the studied raptors, reflecting adaptations to their feeding habits, hunting techniques and ecological niches. The results from Principal component analysis and Canonical variate analysis demonstrate significant differences in beak morphology between the Falconiformes and Accipitriformes clades, as well as among all three groups. The overall mean beak shapes of Golden Eagles are quite similar to Common Buzzards, with both species having longer beaks. In contrast, Falcons exhibit a distinctly different beak morphology, characterized by wider and shorter beaks. Changes in beak shape can lead to changes depending on the skull. It is thought that skull shape variations among predator families may have an impact on beak shape. These findings highlight the importance of integrating morphometric analyses with ecological insights to enhance our understanding of the evolutionary processes shaping raptor beak morphology.

Birds optimize fruit size consumed near their geographic range limits.

Animals can adjust their diet to maximize energy or nutritional intake. For example, birds often target fruits that match their beak size because those fruits can be consumed more efficiently. We hypothesized that pressure to optimize diet-measured as matching between fruit and beak size-increases under stressful environments, such as those that determine species' range edges. Using fruit-consumption and trait information for 97 frugivorous bird and 831 plant species across six continents, we demonstrate that birds feed more frequently on closely size-matched fruits near their geographic range limits. This pattern was particularly strong for highly frugivorous birds, whereas opportunistic frugivores showed no such tendency. These findings highlight how frugivore interactions might respond to stressful conditions and reveal that trait matching may not predict resource use consistently.

Genetic parameter estimation and molecular foundation of chicken beak shape.

The bird beak is mainly functioned as feeding and attacking, and its shape has extremely important significance for survival and reproduction. In chickens, since beak shape could lead to some disadvantages including pecking and waste of feed, it is important to understand the inheritance of chicken beak shape. In the present study, we firstly established 4 indicators to describe the chicken beak shapes, including upper beak length (UL), lower beak length (LL), distance between upper and lower beak tips (DB) and upper beak curvature (BC). And then, we measured the 4 beak shape indicators as well as some production traits including body weight (BW), shank length (SL), egg weight (EW), eggshell strength (ES) of a layer breed, Rhode Island Red (RIR), in order to estimate genetic parameters of chicken beak shape. The heritabilities of UL and LL were 0.41 and 0.37, and the heritabilities of DB and BC were 0.22 and 0.21, indicating that beak shape was a highly or mediumly heritable. There were significant positive genetic and phenotypic correlations among UL, LL, and DB. And UL was positively correlated with body weight (BW18) and shank length (SL18) at 18 weeks of age in genetics, and DB was positively correlated with BC in terms of genetics and phenotype. We also found that layers of chicken cages played a role on beak shape, which could be attributed to the difference of lightness in different cage layers. By a genome-wide association study (GWAS) for the chicken UL, we identified 9 significant candidate genes associated with UL in RIR. For the variants with low minor allele frequencies (MAF <0.01) and outside of high linkage disequilibrium (LD) regions, we also conducted rare variant association studies (RVA) and GWAS to find the association between genotype and phenotype. We also analyzed transcriptomic data from multiple tissues of chicken embryos and revealed that all of the 9 genes were highly expressed in beak of chicken embryos, indicating their potential function for beak development. Our results provided the genetic foundation of chicken beak shape, which could help chicken breeding on beak related traits.

Complex patterns of morphological diversity across multiple populations of an urban bird species.

Urbanization presents a natural evolutionary experiment because selection pressures in cities can be strongly mismatched with those found in species' historic habitats. However, some species have managed to adapt and even thrive in these novel conditions. When a species persists across multiple cities, a fundamental question arises: do we see similar traits evolve in similar novel environments? By testing if and how similar phenotypes emerge across multiple urban populations, we can begin to assess the predictability of population response to anthropogenic change. Here, we examine variation within and across multiple populations of a songbird, the dark-eyed junco (Junco hyemalis). We measured morphological variations in juncos across urban and nonurban populations in Southern California. We investigated whether the variations we observed were due to differences in environmental conditions across cities. Bill shape differed across urban populations; Los Angeles and Santa Barbara juncos had shorter, deeper bills than nonurban juncos, but San Diego juncos did not. On the other hand, wing length decreased with the built environment, regardless of the population. Southern Californian urban juncos exhibit both similarities and differences in morphological traits. Studying multiple urban populations can help us determine the predictability of phenotypic evolutionary responses to novel environments.

Correlated evolution of beak and braincase morphology is present only in select bird clades.

Complex morphological structures, such as skulls or limbs, are often composed of multiple morphological components (e.g., bones, sets of bones) that may evolve in a covaried manner with one another. Previous research has reached differing conclusions on the number of semi-independent units, or modules, that exist in the evolution of structures and on the strength of the covariation, or integration, between these hypothesized modules. We focus on the avian skull as an example of a complex morphological structure for which highly variable conclusions have been reached in the numerous studies analyzing support for a range of simple to complex modularity hypotheses. We hypothesized that past discrepancies may stem from both the differing densities of data used to analyze support for modularity hypotheses and the differing taxonomic levels of study. To test these hypotheses, we applied a comparative method to 3D geometric morphometric data collected from the skulls of a diverse order of birds (the Charadriiformes) to test support for 11 distinct hypotheses of modular skull evolution. Across all Charadriiformes, our analyses suggested that charadriiform skull evolution has been characterized by the semi-independent, but still correlated, evolution of the beak from the rest of the skull. When we adjusted the density of our morphometric data, this result held, but the strength of the signal varied substantially. Additionally, when we analyzed subgroups within the order in isolation, we found support for distinct hypotheses between subgroups. Taken together, these results suggest that differences in the methodology of past work (i.e., statistical method and data density) as well as clade-specific dynamics may be the reasons past studies have reached varying conclusions.

Geometric morphometrics casts light on phylogenetic relevance of cephalopod beak morphological.

The feeding organ of cephalopod species, the beak, can be used to reveal important ecological information. In this study, geometric morphometric approaches were employed to investigate the phylogenetic relevance and classification effect of beak lateral profile shape. The two-dimensional beak morphologies of 1164 pairs of 24 species from 13 genera and five families were constructed, and their evolutionary relationships and taxonomic status were confirmed using geometric morphometrics and molecular biology approaches. We also assessed the phylogenetic signals of beak shape. The analysis results show shape variation in the beak mainly in the rostrum, hood, and lateral wall. The overall shape parameters (all PCs) of the upper and lower beak are more useful for species identification. The shapes of the upper and lower beak show a strong phylogenetic signal, and the phenogram based on the beak shape basically reflected the families' taxonomic positions. We also hypothesized that the shape variation in the beaks of cephalopods may be ascribed to genetic and environmental differences. In summary, beaks are a reliable material for the classification of cephalopod species. Geometric morphometric approaches are a powerful tool to reveal the identification, phylogenetic relevance and phenotypic diversity of beak shape in cephalopods.

Community-wide genome sequencing reveals 30 years of Darwin's finch evolution.

A fundamental goal in evolutionary biology is to understand the genetic architecture of adaptive traits. Using whole-genome data of 3955 of Darwin's finches on the Galápagos Island of Daphne Major, we identified six loci of large effect that explain 45% of the variation in the highly heritable beak size of Geospiza fortis, a key ecological trait. The major locus is a supergene comprising four genes. Abrupt changes in allele frequencies at the loci accompanied a strong change in beak size caused by natural selection during a drought. A gradual change in Geospiza scandens occurred across 30 years as a result of introgressive hybridization with G. fortis. This study shows how a few loci with large effect on a fitness-related trait contribute to the genetic potential for rapid adaptive radiation.

New reptile shows dinosaurs and pterosaurs evolved among diverse precursors.

Dinosaurs and pterosaurs have remarkable diversity and disparity through most of the Mesozoic Era1-3. Soon after their origins, these reptiles diversified into a number of long-lived lineages, evolved unprecedented ecologies (for example, flying, large herbivorous forms) and spread across Pangaea4,5. Recent discoveries of dinosaur and pterosaur precursors6-10 demonstrated that these animals were also speciose and widespread, but those precursors have few if any well-preserved skulls, hands and associated skeletons11,12. Here we present a well-preserved partial skeleton (Upper Triassic, Brazil) of the new lagerpetid Venetoraptor gassenae gen. et sp. nov. that offers a more comprehensive look into the skull and ecology of one of these precursors. Its skull has a sharp, raptorial-like beak, preceding that of dinosaurs by around 80 million years, and a large hand with long, trenchant claws that firmly establishes the loss of obligatory quadrupedalism in these precursor lineages. Combining anatomical information of the new species with other dinosaur and pterosaur precursors shows that morphological disparity of precursors resembles that of Triassic pterosaurs and exceeds that of Triassic dinosaurs. Thus, the 'success' of pterosaurs and dinosaurs was a result of differential survival among a broader pool of ecomorphological variation. Our results show that the morphological diversity of ornithodirans started to flourish among early-diverging lineages and not only after the origins of dinosaurs and pterosaurs.

Beak shape and nest material use in birds.

The evolution of behaviour can both influence, and be influenced by, morphology. Recent advances in methods and data availability have facilitated broad-scale investigations of physical form and behavioural function in many contexts, but the relationship between animal morphology and object manipulation-particularly objects used in construction-remains largely unknown. Here, we employ a new global database of nest materials used by 5924 species of birds together with phylogenetically informed random forest models to evaluate the link between beak shape and these nest-building materials. We find that beak morphology, together with species diet and access to materials, can predict nest-material use above chance and with high accuracy (68-97%). Much of this relationship, however, is driven by phylogenetic signal and sampling biases. We therefore conclude that while variation in nest material use is linked with that of beak shape across bird species, these correlations are modulated by the ecological context and evolutionary history of these species. This article is part of the theme issue 'The evolutionary ecology of nests: a cross-taxon approach'.

Grey parrot (Psittacus erithacus) beak papillae and nerves identified using novel 2-D and 3-D imaging modalities.

OBJECTIVE: The avian beak is a complex organ containing bone, neurovascular tissue, and keratinized covering (rhamphotheca). Nerve-rich papillae extend through bone into rhamphotheca providing sensory input from the beak tip. Beak trimming is a common procedure in avian species and is used for corrective, cosmetic, and behavioral modification purposes. Yet, practitioners are not well versed in complete beak anatomy, and therefore, beak trimming often disregards neurovasculature, injuring the patient and hampering recovery. Here, using comprehensive anatomical description, we aim to provide recommendations on how to safely perform beak trimming without damaging underlying sensory papillae. ANIMALS: Here, we evaluated beaks of 2 deceased grey parrots (Psittacus erithacus). PROCEDURES: In one, we used a novel stain and microcomputed tomography to visualize papillae in the upper and lower beaks. In a second, we hand isolated the upper and lower beak dermal papillae and used high-resolution photography plus traditional paraffin histology. RESULTS: Papillae and their nerves were easily identified in these 2- and 3-dimensional approaches. This allowed us to determine the approximate lengths of papillae within the upper and lower beak. CLINICAL RELEVANCE: Based on these findings, the authors recommend lateral radiographs of the bird's head and beak to identify the location of the underlying bone relative to the overlying rhamphotheca before performing beak trims. Specifically in grey parrots, the authors recommend the upper and lower beak should not be trimmed closer than 8 to 10 mm from the underlying bone. Further work is needed to support these recommendations and provide guidelines for other species.

Biomechanics illuminates form-function relationships in bird bills.

The field of comparative biomechanics examines how form, mechanical properties and environmental interactions shape the function of biological structures. Biomechanics has advanced by leaps and bounds as rapid technological progress opens up new research horizons. In this Review, I describe how our understanding of the avian bill, a morphologically diverse multifunctional appendage, has been transformed by employing a biomechanical perspective. Across functions from feeding to excavating hollows in trees and as a vocal apparatus, the study of the bill spans both solid and fluid biomechanics, rendering it useful to understand general principles across disciplines. The different shapes of the bill across bird species result in functional and mechanical trade-offs, thus representing a microcosm of many broader form-function questions. Using examples from diverse studies, I discuss how research into bird bills has been shaped over recent decades, and its influence on our understanding of avian ecology and evolution. Next, I examine how bill material properties and geometry influence performance in dietary and non-dietary contexts, simultaneously imposing trade-offs on other functions. Following an examination of the interactions of bills with fluids and their role as part of the vocal apparatus, I end with a discussion of the sensory biomechanics of the bill, focusing specifically on the bill-tip mechanosensory organ. With these case studies, I highlight how this burgeoning and consequential field represents a roadmap for our understanding of the function and evolution of biological structures.

Intrinsically disordered BMP4 morphogen and the beak of the finch: Co-option of an ancient axial patterning system.

Darwin's finches, with the primary diversity in the shape and size of their beaks, represent an excellent model system to study speciation and adaptive evolution. It is generally held that evolution depends on the natural selection of heritable phenotypic variations originating from the genetic mutations. However, it is now increasingly evident that epigenetic transgenerational inheritance of phenotypic variation can also guide evolutionary change. Several studies have shown that the bone morphogenetic protein BMP4 is a major driver of beak morphology. A recent study explored variability of the morphological, genetic, and epigenetic differences in the adjacent "urban" and "rural" populations of two species of Darwin's finches on the Galápagos Islands and revealed significant changes in methylation patterns in several genes including those involved in the BMP/TGFß pathway in the sperm DNA compared to erythrocyte DNA. These observations indicated that epigenetic changes caused by environmental fluctuations can be passed on to the offspring. Nonetheless, the mechanism by which dysregulated expression of BMP4 impacts beak morphology remains poorly understood. Here, we show that BMP4 is an intrinsically disordered protein and present a causal a link between epigenetic changes, BMP4 dysregulation and the evolution of the beak of the finch by natural selection.

Comparative morphology and soft tissue histology of the remote-touch bill-tip organ in three ibis species of differing foraging ecology.

Ibises (order: Pelecaniformes, family: Threskiornithidae) are probe-foraging birds that use 'remote-touch' to locate prey items hidden in opaque substrates. This sensory capability allows them to locate their prey using high-frequency vibrations in the substrate in the absence of other sensory cues. Remote-touch is facilitated by a specialised bill-tip organ, comprising high densities of mechanoreceptors (Herbst corpuscles) embedded in numerous foramina in the beak bones. Each foramen and its associated Herbst corpuscles make up a sensory unit, called a 'sensory pit'. These sensory pits are densely clustered in the distal portion of the beak. Previous research has indicated that interspecific differences in the extent of sensory pitting in the beak bones correlate with aquatic habitat use of ibises, and have been suggested to reflect different levels of remote-touch sensitivity. Our study investigates the interspecific differences in the bone and soft tissue histology of the bill-tip organs of three species of southern African ibises from different habitats (mainly terrestrial to mainly aquatic). We analysed the external pitting pattern on the bones, as well as internal structure of the beak using micro-CT scans and soft tissue histological sections of each species' bill-tip organs. The beaks of all three species contain remote-touch bill-tip organs and are described here in detail. Clear interspecific differences are evident between the species' bill-tip organs, both in terms of bone morphology and soft tissue histology. Glossy Ibises, which forage exclusively in wetter substrates, have a greater extent of pitting but lower numbers of Herbst corpuscles in each pit, while species foraging in drier substrates (Hadeda and Sacred Ibises) have more robust beaks, fewer pits and higher densities of Herbst corpuscles. Our data, together with previously published histological descriptions of the bill-tip organs of other remote-touch foraging bird species, indicate that species foraging in drier habitats have more sensitive bill-tip organs (based on their anatomy). The vibrations produced by prey (e.g., burrowing invertebrates) travel poorly in dry substrates compared with wetter ones (i.e., dry soil vs. mud or water), and thus we hypothesise that a more sensitive bill-tip organ may be required to successfully locate prey in dry substrates. Furthermore, our results indicate that the differences in bill-tip organ anatomy between the species reflect complex trade-offs between morphological constraints of beak shape and remote-touch sensitivity requirements, both of which are likely related to each species' foraging behaviour and substrate usage. Our study suggests that structures in the bone of the bill-tip organ could provide valuable osteological correlates for the associated soft tissues, and consequently may provide information on the sensory ecology and habitat usage of the birds in the absence of soft tissues.

[Small scale evolution]. / L'évolution à petite échelle.

Small-scale evolution or microevolution concerns evolution at the intra-specific level or between closely related species. At the intra-specific level, it allows the analysis of the evolutionary forces at work: mutation, genetic drift, migration and selection. Moreover, because of the short evolutionary time, it is easier to identify the genetic basis of observed phenotypic differences. Most studies focus on current populations but more and more analyses are performed on ancient DNA. This provides important information for tracing the history of populations and also allows the reconstruction of phenotypes of individuals that disappeared several thousand years ago. In this short review, I present studies showing how pre-zygotic or post-zygotic barriers involved in species formation are set up using the example of the geographical barrier due to the formation of the Isthmus of Panama and that of the heterochromatin divergence in Drosophilidae. I also describe the different approaches that have been used to identify the genetic basis of well known phenotypic variations: candidate gene approach (about melanism in felines), QTL mapping (variation in the number of lateral bone plates in sticklebacks), association study (pigmentation in the Asian ladybird). Finally, I illustrate the key impact of natural selection with the iconic example of the evolution of the beak of Galapagos finches, and the role of certain developmental genes in its morphological diversification.

Title: L'évolution à petite échelle. Abstract: L'évolution à petite échelle ou microévolution concerne l'évolution au niveau intra-spécifique ou entre espèces proches. Au niveau intra-spécifique, elle permet d'analyser les forces évolutives en action : mutation, dérive génétique, migration et sélection. De plus, en raison de ce temps évolutif court, il est plus facile d'identifier les bases génétiques des différences phénotypiques observées. La plupart des études porte sur des populations actuelles mais de plus en plus de travaux analysent l'ADN ancien. Ces derniers apportent non seulement des informations importantes pour retracer l'histoire des populations mais permettent également de reconstituer les phénotypes d'individus disparus depuis plusieurs milliers d'années. Dans cette courte revue, je présente des travaux montrant comment se mettent en place des barrières pré-zygotiques ou post-zygotiques impliquées dans la formation d'espèces, avec l'exemple de la barrière géographique due à la formation de l'isthme de Panama et celui de la divergence de l'hétérochromatine chez les drosophilidés. Par ailleurs, à propos de cas bien établis, je décris les différentes approches qui ont été utilisées pour identifier les bases génétiques de variations phénotypiques : approche gène-candidat pour ce qui concerne le mélanisme chez les félins, cartographie QTL (Quantitative trait loci) pour la variation du nombre de plaques osseuses latérales chez les épinoches, étude d'association pour la pigmentation chez la coccinelle asiatique. Enfin, j'illustre le rôle de la sélection naturelle avec l'exemple iconique de l'évolution du bec des pinsons des Galapagos et l'implication de certains gènes du développement dans sa diversification morphologique.

Change in beak overhangs of cliff swallows over 40 years: Partly a response to parasites?

Some birds exhibit a maxillary overhang, in which the tip of the upper beak projects beyond the lower mandible and may curve downward. The overhang is thought to help control ectoparasites on the feathers. Little is known about the extent to which the maxillary overhang varies spatially or temporally within populations of the same species. The colonial cliff swallow (Petrochelidon pyrrhonota) has relatively recently shifted to almost exclusive use of artificial structures such as bridges and highway culverts for nesting and consequently has been exposed to higher levels of parasitism than on its ancestral cliff nesting sites. We examined whether increased ectoparasitism may have favored recent changes in the extent of the maxillary overhang. Using a specimen collection of cliff swallows from western Nebraska, USA, spanning 40 years and field data on live birds, we found that the extent of the maxillary overhang increased across years in a nonlinear way, peaking in the late 2000's, and varied inversely with cliff swallow colony size for unknown reasons. The number of fleas on nestling cliff swallows declined in general over this period. Those birds with perceptible overhangs had fewer swallow bugs on the outside of their nest, but they did not have higher nesting success than birds with no overhangs. The intraspecific variation in the maxillary overhang in cliff swallows was partly consistent with it having a functional role in combatting ectoparasites. The temporal increase in the extent of the overhang may be a response by cliff swallows to their relatively recent increased exposure to parasitism. Our results demonstrate that this avian morphological trait can change rapidly over time.

Analyzing Disparity and Rates of Morphological Evolution with Model-Based Phylogenetic Comparative Methods.

Understanding variation in rates of evolution and morphological disparity is a goal of macroevolutionary research. In a phylogenetic comparative methods framework, we present three explicit models for linking the rate of evolution of a trait to the state of another evolving trait. This allows testing hypotheses about causal influences on rates of phenotypic evolution with phylogenetic comparative data. We develop a statistical framework for fitting the models with generalized least-squares regression and use this to discuss issues and limitations in the study of rates of evolution more generally. We show that the power to detect effects on rates of evolution is low in that even strong causal effects are unlikely to explain more than a few percent of observed variance in disparity. We illustrate the models and issues by testing if rates of beak-shape evolution in birds are influenced by brain size, as may be predicted from a Baldwin effect in which presumptively more behaviorally flexible large-brained species generate more novel selection on themselves leading to higher rates of evolution. From an analysis of morphometric data for 645 species, we find evidence that both macro- and microevolution of the beak are faster in birds with larger brains, but with the caveat that there are no consistent effects of relative brain size.[Baldwin effect; beak shape; behavioral drive; bird; brain size; disparity; phylogenetic comparative method; rate of evolution.].

Independent evolution of intermediate bill widths in a seabird clade.

Interspecific introgression can occur between species that evolve rapidly within an adaptive radiation. Pachyptila petrels differ in bill size and are characterised by incomplete reproductive isolation, leading to interspecific gene flow. Salvin's prion (Pachyptila salvini), whose bill width is intermediate between broad-billed (P. vittata) and Antarctic (P. desolata) prions, evolved through homoploid hybrid speciation. MacGillivray's prion (P. macgillivrayi), known from a single population on St Paul (Indian Ocean), has a bill width intermediate between salvini and vittata and could also be the product of interspecies introgression or hybrid speciation. Recently, another prion population phenotypically similar to macgillivrayi was discovered on Gough (Atlantic Ocean), where it breeds 3 months later than vittata. The similarity in bill width between the medium-billed birds on Gough and macgillivrayi suggest that they could be closely related. In this study, we used genetic and morphological data to infer the phylogenetic position and evolutionary history of P. macgillivrayi and the Gough medium-billed prion relative other Pachyptila taxa, to determine whether species with medium bill widths evolved through common ancestry or convergence. We found that Gough medium-billed prions belong to the same evolutionary lineage as macgillivrayi, representing a new population of MacGillivray's prion that originated through a colonisation event from St Paul. We show that macgillivrayi's medium bill width evolved through divergence (genetic drift) and independently from that of salvini, which evolved through hybridisation (gene flow). This represents the independent convergence towards a similarly medium-billed phenotype. The newly discovered MacGillivray's prion population on Gough is of utmost conservation relevance, as the relict macgillivrayi population in the Indian Ocean is very small.