The importance of migratory drop-off for island colonization in birds.

Seasonal migration is an underappreciated driver of animal diversification. Changes in migratory behaviour may favour the establishment of sedentary founder populations and promote speciation if there is sufficient reproductive isolation between sedentary and migratory populations. From a systematic literature review, we here quantify the role of migratory drop-off-the loss of migratory behaviour-in promoting speciation in birds on islands. We identify at least 157 independent colonization events likely initiated by migratory species that led to speciation, including 44 cases among recently extinct species. By comparing, for all islands, the proportion of island endemic species that derived from migratory drop-off with the proportion of migratory species among potential colonizers, we showed that seasonal migration has a larger effect on island endemic richness than direct dispersal. We also found that the role of migration in island colonization increases with the geographic isolation of islands. Furthermore, the success of speciation events depends in part on species biogeographic and ecological factors, here positively associated with greater range size and larger flock sizes. These results highlight the importance of shifts in migratory behaviour in the speciation process and calls for greater consideration of migratory drop-off in the biogeographic distribution of birds.

A global analysis of avian island diversity-area relationships in the Anthropocene.

Research on island species-area relationships (ISAR) has expanded to incorporate functional (IFDAR) and phylogenetic (IPDAR) diversity. However, relative to the ISAR, we know little about IFDARs and IPDARs, and lack synthetic global analyses of variation in form of these three categories of island diversity-area relationship (IDAR). Here, we undertake the first comparative evaluation of IDARs at the global scale using 51 avian archipelagic data sets representing true and habitat islands. Using null models, we explore how richness-corrected functional and phylogenetic diversity scale with island area. We also provide the largest global assessment of the impacts of species introductions and extinctions on the IDAR. Results show that increasing richness with area is the primary driver of the (non-richness corrected) IPDAR and IFDAR for many data sets. However, for several archipelagos, richness-corrected functional and phylogenetic diversity changes linearly with island area, suggesting that the dominant community assembly processes shift along the island area gradient. We also find that archipelagos with the steepest ISARs exhibit the biggest differences in slope between IDARs, indicating increased functional and phylogenetic redundancy on larger islands in these archipelagos. In several cases introduced species seem to have 're-calibrated' the IDARs such that they resemble the historic period prior to recent extinctions.

Brain size predicts bees' tolerance to urban environments.

The rapid conversion of natural habitats to anthropogenic landscapes is threatening insect pollinators worldwide, raising concern regarding the negative consequences on their fundamental role as plant pollinators. However, not all pollinators are negatively affected by habitat conversion, as certain species find appropriate resources in anthropogenic landscapes to persist and proliferate. The reason why some species tolerate anthropogenic environments while most find them inhospitable remains poorly understood. The cognitive buffer hypothesis, widely supported in vertebrates but untested in insects, offers a potential explanation. This theory suggests that species with larger brains have enhanced behavioural plasticity, enabling them to confront and adapt to novel challenges. To investigate this hypothesis in insects, we measured brain size for 89 bee species, and evaluated their association with the degree of habitat occupancy. Our analyses revealed that bee species mainly found in urban habitats had larger brains relative to their body size than those that tend to occur in forested or agricultural habitats. Additionally, urban bees exhibited larger body sizes and, consequently, larger absolute brain sizes. Our results provide the first empirical support for the cognitive buffer hypothesis in invertebrates, suggesting that a large brain in bees could confer behavioural advantages to tolerate urban environments.

AVONET: morphological, ecological and geographical data for all birds.

Functional traits offer a rich quantitative framework for developing and testing theories in evolutionary biology, ecology and ecosystem science. However, the potential of functional traits to drive theoretical advances and refine models of global change can only be fully realised when species-level information is complete. Here we present the AVONET dataset containing comprehensive functional trait data for all birds, including six ecological variables, 11 continuous morphological traits, and information on range size and location. Raw morphological measurements are presented from 90,020 individuals of 11,009 extant bird species sampled from 181 countries. These data are also summarised as species averages in three taxonomic formats, allowing integration with a global phylogeny, geographical range maps, IUCN Red List data and the eBird citizen science database. The AVONET dataset provides the most detailed picture of continuous trait variation for any major radiation of organisms, offering a global template for testing hypotheses and exploring the evolutionary origins, structure and functioning of biodiversity.

Niche shifts after island colonization spurred adaptive diversification and speciation in a cosmopolitan bird clade.

Islands have long been recognized as key contributors to biodiversity because they facilitate geographic isolation and ecological divergence from mainland ancestors. However, island colonization has traditionally been considered an evolutionary dead-end process, and its consequences for continental biodiversity remain understudied. Here, we use the evolutionary radiation of Columbiformes (i.e. pigeons and doves) to examine if ecological niche shifts on islands shaped biological diversification and community composition on continents. We show that the colonization of islands by continental, terrestrial-foraging lineages led to the exploitation of a new ecological niche (i.e. arboreal foraging). This transition towards arboreal foraging was associated with evolutionary adaptation towards a new morphological optimum. In addition, arboreal-foraging lineages of islands experienced an increase in speciation rates, which was associated with successful range expansions to other islands as well as back colonization of continents. Our results provide empirical evidence that diversification on continents can only be fully understood when studying the diversification processes that took place on islands, challenging the view of islands as mere sinks of evolutionary diversity.

Feeding specialization and longer generation time are associated with relatively larger brains in bees.

Despite their miniature brains, insects exhibit substantial variation in brain size. Although the functional significance of this variation is increasingly recognized, research on whether differences in insect brain sizes are mainly the result of constraints or selective pressures has hardly been performed. Here, we address this gap by combining prospective and retrospective phylogenetic-based analyses of brain size for a major insect group, bees (superfamily Apoidea). Using a brain dataset of 93 species from North America and Europe, we found that body size was the single best predictor of brain size in bees. However, the analyses also revealed that substantial variation in brain size remained even when adjusting for body size. We consequently asked whether such variation in relative brain size might be explained by adaptive hypotheses. We found that ecologically specialized species with single generations have larger brains-relative to their body size-than generalist or multi-generation species, but we did not find an effect of sociality on relative brain size. Phylogenetic reconstruction further supported the existence of different adaptive optima for relative brain size in lineages differing in feeding specialization and reproductive strategy. Our findings shed new light on the evolution of the insect brain, highlighting the importance of ecological pressures over social factors and suggesting that these pressures are different from those previously found to influence brain evolution in other taxa.

Relative Brain Size and Its Relation with the Associative Pallium in Birds.

Despite growing interest in the evolution of enlarged brains, the biological significance of brain size variation remains controversial. Much of the controversy is over the extent to which brain structures have evolved independently of each other (mosaic evolution) or in a coordinated way (concerted evolution). If larger brains have evolved by the increase of different brain regions in different species, it follows that comparisons of the whole brain might be biologically meaningless. Such an argument has been used to criticize comparative attempts to explain the existing variation in whole-brain size among species. Here, we show that pallium areas associated with domain-general cognition represent a large fraction of the entire brain, are disproportionally larger in large-brained birds and accurately predict variation in the whole brain when allometric effects are appropriately accounted for. While this does not question the importance of mosaic evolution, it suggests that examining specialized, small areas of the brain is not very helpful for understanding why some birds have evolved such large brains. Instead, the size of the whole brain reflects consistent variation in associative pallium areas and hence is functionally meaningful for comparative analyses.

Undiscovered bird extinctions obscure the true magnitude of human-driven extinction waves.

Birds are among the best-studied animal groups, but their prehistoric diversity is poorly known due to low fossilization potential. Hence, while many human-driven bird extinctions (i.e., extinctions caused directly by human activities such as hunting, as well as indirectly through human-associated impacts such as land use change, fire, and the introduction of invasive species) have been recorded, the true number is likely much larger. Here, by combining recorded extinctions with model estimates based on the completeness of the fossil record, we suggest that at least ~1300-1500 bird species (~12% of the total) have gone extinct since the Late Pleistocene, with 55% of these extinctions undiscovered (not yet discovered or left no trace). We estimate that the Pacific accounts for 61% of total bird extinctions. Bird extinction rate varied through time with an intense episode ~1300 CE, which likely represents the largest human-driven vertebrate extinction wave ever, and a rate 80 (60-95) times the background extinction rate. Thus, humans have already driven more than one in nine bird species to extinction, with likely severe, and potentially irreversible, ecological and evolutionary consequences.

Evolutionary transitions in diet influence the exceptional diversification of a lizard adaptive radiation.

BACKGROUND: Diet is a key component of a species ecological niche and plays critical roles in guiding the trajectories of evolutionary change. Previous studies suggest that dietary evolution can influence the rates and patterns of species diversification, with omnivorous (animal and plant, 'generalist') diets slowing down diversification compared to more restricted ('specialist') herbivorous and carnivorous diets. This hypothesis, here termed the "dietary macroevolutionary sink" hypothesis (DMS), predicts that transitions to omnivorous diets occur at higher rates than into any specialist diet, and omnivores are expected to have the lowest diversification rates, causing an evolutionary sink into a single type of diet. However, evidence for the DMS hypothesis remains conflicting. Here, we present the first test of the DMS hypothesis in a lineage of ectothermic tetrapods-the prolific Liolaemidae lizard radiation from South America. RESULTS: Ancestral reconstructions suggest that the stem ancestor was probably insectivorous. The best supported trait model is a diet-dependent speciation rate, with independent extinction rates. Herbivory has the highest net diversification rate, omnivory ranks second, and insectivory has the lowest. The extinction rate is the same for all three diet types and is much lower than the speciation rates. The highest transition rate was from omnivory to insectivory, and the lowest transition rates were between insectivory and herbivory. CONCLUSIONS: Our findings challenge the core prediction of the DMS hypothesis that generalist diets represent an 'evolutionary sink'. Interestingly, liolaemid lizards have rapidly and successfully proliferated across some of the world's coldest climates (at high elevations and latitudes), where species have evolved mixed arthropod-plant (omnivore) or predominantly herbivore diets. This longstanding observation is consistent with the higher net diversification rates found in both herbivory and omnivory. Collectively, just like the evolution of viviparity has been regarded as a 'key adaptation' during the liolaemid radiation across cold climates, our findings suggest that transitions from insectivory to herbivory (bridged by omnivory) are likely to have played a role as an additional key adaptation underlying the exceptional diversification of these reptiles across extreme climates.

Niche expansion and adaptive divergence in the global radiation of crows and ravens.

The processes that allow some lineages to diversify rapidly at a global scale remain poorly understood. Although earlier studies emphasized the importance of dispersal, global expansions expose populations to novel environments and may also require adaptation and diversification across new niches. In this study, we investigated the contributions of these processes to the global radiation of crows and ravens (genus Corvus). Combining a new phylogeny with comprehensive phenotypic and climatic data, we show that Corvus experienced a massive expansion of the climatic niche that was coupled with a substantial increase in the rates of species and phenotypic diversification. The initiation of these processes coincided with the evolution of traits that promoted dispersal and niche expansion. Our findings suggest that rapid global radiations may be better understood as processes in which high dispersal abilities synergise with traits that, like cognition, facilitate persistence in new environments.

Neuron numbers link innovativeness with both absolute and relative brain size in birds.

A longstanding issue in biology is whether the intelligence of animals can be predicted by absolute or relative brain size. However, progress has been hampered by an insufficient understanding of how neuron numbers shape internal brain organization and cognitive performance. On the basis of estimations of neuron numbers for 111 bird species, we show here that the number of neurons in the pallial telencephalon is positively associated with a major expression of intelligence: innovation propensity. The number of pallial neurons, in turn, is greater in brains that are larger in both absolute and relative terms and positively covaries with longer post-hatching development periods. Thus, our analyses show that neuron numbers link cognitive performance to both absolute and relative brain size through developmental adjustments. These findings help unify neuro-anatomical measures at multiple levels, reconciling contradictory views over the biological significance of brain expansion. The results also highlight the value of a life history perspective to advance our understanding of the evolutionary bases of the connections between brain and cognition.

Madagascar's extraordinary biodiversity: Threats and opportunities.

Madagascar's unique biota is heavily affected by human activity and is under intense threat. Here, we review the current state of knowledge on the conservation status of Madagascar's terrestrial and freshwater biodiversity by presenting data and analyses on documented and predicted species-level conservation statuses, the most prevalent and relevant threats, ex situ collections and programs, and the coverage and comprehensiveness of protected areas. The existing terrestrial protected area network in Madagascar covers 10.4% of its land area and includes at least part of the range of the majority of described native species of vertebrates with known distributions (97.1% of freshwater fishes, amphibians, reptiles, birds, and mammals combined) and plants (67.7%). The overall figures are higher for threatened species (97.7% of threatened vertebrates and 79.6% of threatened plants occurring within at least one protected area). International Union for Conservation of Nature (IUCN) Red List assessments and Bayesian neural network analyses for plants identify overexploitation of biological resources and unsustainable agriculture as the most prominent threats to biodiversity. We highlight five opportunities for action at multiple levels to ensure that conservation and ecological restoration objectives, programs, and activities take account of complex underlying and interacting factors and produce tangible benefits for the biodiversity and people of Madagascar.

Loss of functional diversity through anthropogenic extinctions of island birds is not offset by biotic invasions.

Human impacts reshape ecological communities through the extinction and introduction of species. The combined impact of these factors depends on whether non-native species fill the functional roles of extinct species, thus buffering the loss of functional diversity. This question has been difficult to address, because comprehensive information about past extinctions and their traits is generally lacking. We combine detailed information about extinct, extant, and established alien birds to quantify historical changes in functional diversity across nine oceanic archipelagos. We found that alien species often equal or exceed the number of anthropogenic extinctions yet apparently perform a narrower set of functional roles as current island assemblages have undergone a substantial and ubiquitous net loss in functional diversity and increased functional similarity among assemblages. Our results reveal that the introduction of alien species has not prevented anthropogenic extinctions from reducing and homogenizing the functional diversity of native bird assemblages on oceanic archipelagos.

Coexistence of predators in time: Effects of season and prey availability on species activity within a Mediterranean carnivore guild.

The degree of coexistence among predators can determine the structure of ecological communities. Niche partitioning is a common strategy applied by species to enhance their coexistence. Diet, habitat, or time use can be responsible for segregation among carnivore species, the latter factor being the least studied in Mediterranean ecosystems. Terrestrial medium-sized carnivores (i.e., mesocarnivores) carry out important functions in ecosystems, and identifying their interactions is essential for their conservation.In this study, we explore the activity of a terrestrial mesocarnivore guild in order to determine seasonal differences in daily activity patterns of competitors and prey. We also investigate how the abundance of a common mesocarnivore prey in the region, small mammals, influences the activity of predators.During a year, camera trap devices (n = 18) were installed in Montseny Natural Park (Catalan Pre-Coastal Range, North-East Iberian Peninsula), a region that hosts five mesocarnivore species. Camera trapping detections were used to estimate their daily activity patterns and corresponding overlaps. We also surveyed small mammal plots (n = 5) in order to calculate prey abundance and test its effect on the relative activity of each carnivore species.Despite all target mesocarnivores are mainly nocturnal, the activity overlap among them varies according to species particularities and season. Red fox (Vulpes vulpes) appears as a generalist species in terms of time use, whereas stone marten (Martes foina) and genet (Genetta genetta) show the most similar activity patterns and both of them seem to be positively influenced by small mammal abundance. Overall, the diversity found in the way mesocarnivore species use time could facilitate their coexistence.Despite activity pattern similarities among carnivore species should not be directly translated to negative interactions, they can have a strong influence in habitat and resource-limited ecosystems. Therefore, activity overlaps should be taken into account when discussing wildlife management actions.

Behavioural plasticity is associated with reduced extinction risk in birds.

Behavioural plasticity is believed to reduce species vulnerability to extinction, yet global evidence supporting this hypothesis is lacking. We address this gap by quantifying the extent to which birds are observed behaving in novel ways to obtain food in the wild; based on a unique dataset of >3,800 novel behaviours, we show that species with a higher propensity to innovate are at a lower risk of global extinction and are more likely to have increasing or stable populations than less innovative birds. These results mainly reflect a higher tolerance of innovative species to habitat destruction, the main threat for birds.

Larger brains spur species diversification in birds.

Evidence is accumulating that species traits can spur their evolutionary diversification by influencing niche shifts, range expansions, and extinction risk. Previous work has shown that larger brains (relative to body size) facilitate niche shifts and range expansions by enhancing behavioral plasticity but whether larger brains also promote evolutionary diversification is currently backed by insufficient evidence. We addressed this gap by combining a brain size dataset for >1900 avian species worldwide with estimates of diversification rates based on two conceptually different phylogenetic-based approaches. We found consistent evidence that lineages with larger brains (relative to body size) have diversified faster than lineages with relatively smaller brains. The best supported trait-dependent model suggests that brain size primarily affects diversification rates by increasing speciation rather than decreasing extinction rates. In addition, we found that the effect of relatively brain size on species-level diversification rate is additive to the effect of other intrinsic and extrinsic factors. Altogether, our results highlight the importance of brain size as an important factor in evolution and reinforce the view that intrinsic features of species have the potential to influence the pace of evolution.

Are Urban Vertebrates City Specialists, Artificial Habitat Exploiters, or Environmental Generalists?

Although urbanization is a major threat to biodiversity, some species are able to thrive in cities. This might be because they have specific adaptations to urban conditions, because they are able to cope with artificial habitats in general or because they are generalists that can live in a wide range of conditions. We use the latest version of the IUCN database to distinguish these possibilities in 25,985 species of the four classes of terrestrial vertebrates with the help of phylogenetically controlled methods. We first compare species occurrence in cities with that of the five other artificial habitats recognized by the IUCN and use principal components analyses to ask which of these most resembles cities. We then test whether urban species have a wider habitat breadth than species occurring in other, non-urban, artificial habitats, as well as species that occur only in natural habitats. Our results suggest that the proportion of terrestrial vertebrates that occur in urban environments is small and that, among the species that do occur in cities, the great majority also occur in other artificial habitats. Our data also show that the presence of terrestrial vertebrates in urban habitats is skewed in favor of habitat generalists. In birds and mammals, species occurrence in urban areas is most similar to that of rural gardens, while in reptiles and amphibians, urban areas most resemble pasture and arable land. Our study suggests that cities are likely not unique, as is often thought, and may resemble other types of artificial environments, which urban exploiters can adapt to because of their wide habitat breadth.

Predictable evolution towards larger brains in birds colonizing oceanic islands.

Theory and evidence suggest that some selective pressures are more common on islands than in adjacent mainland habitats, leading evolution to follow predictable trends. The existence of predictable evolutionary trends has nonetheless been difficult to demonstrate, mainly because of the challenge of separating in situ evolution from sorting processes derived from colonization events. Here we use brain size measurements of >1900 avian species to reveal the existence of one such trend: increased brain size in island dwellers. Based on sister-taxa comparisons and phylogenetic ancestral trait estimations, we show that species living on islands have relatively larger brains than their mainland relatives and that these differences mainly reflect in situ evolution rather than varying colonization success. Our findings reinforce the view that in some instances evolution may be predictable, and yield insight into why some animals evolve larger brains despite substantial energetic and developmental costs.

The life-history basis of behavioural innovations.

The evolutionary origin of innovativeness remains puzzling because innovating means responding to novel or unusual problems and hence is unlikely to be selected by itself. A plausible alternative is considering innovativeness as a co-opted product of traits that have evolved for other functions yet together predispose individuals to solve problems by adopting novel behaviours. However, this raises the question of why these adaptations should evolve together in an animal. Here, we develop the argument that the adaptations enabling animals to innovate evolve together because they are jointly part of a life-history strategy for coping with environmental changes. In support of this claim, we present comparative evidence showing that in birds, (i) innovative propensity is linked to life histories that prioritize future over current reproduction, (ii) the link is in part explained by differences in brain size, and (iii) innovative propensity and life-history traits may evolve together in generalist species that frequently expose themselves to novel or unusual conditions. Combined with previous evidence, these findings suggest that innovativeness is not a specialized adaptation but more likely part of a broader general adaptive system to cope with changes in the environment.

Environmental variation and the evolution of large brains in birds.

Environmental variability has long been postulated as a major selective force in the evolution of large brains. However, assembling evidence for this hypothesis has proved difficult. Here, by combining brain size information for over 1,200 bird species with remote-sensing analyses to estimate temporal variation in ecosystem productivity, we show that larger brains (relative to body size) are more likely to occur in species exposed to larger environmental variation throughout their geographic range. Our reconstructions of evolutionary trajectories are consistent with the hypothesis that larger brains (relative to body size) evolved when the species invaded more seasonal regions. However, the alternative-that the species already possessed larger brains when they invaded more seasonal regions-cannot be completely ruled out. Regardless of the exact mechanism, our findings provide strong empirical support for the association between large brains and environmental variability.