Aim  As urban landscapes expand, shifts in biodiversity are occurring. This is leading biogeographers and ecologists to consider human‐dominated landscapes in their current work. One question that arises is: what characterizes those species that are widespread in the most highly urban environments compared with those restricted to less urbanized areas in the city? Here, we aim to identify the traits that enable species to become urban exploiters, i.e. to dominate highly urbanized surroundings. Identifying these traits may help us better predict and possibly mitigate the biotic homogenization occurring in these areas. Location  Israel in general, with special focus on the city of Jerusalem. Methods  Combining literature and field‐based data for birds in Israel we compared phenotypic, behavioural and life‐history traits between urban exploiters and urban adapters. The latter occur in urban landscapes, but are characteristic of the less urbanized parts of the city. We then examined the trends along a finer field‐sampled gradient of increasing urbanization from sub‐natural to downtown areas within the city of Jerusalem. Results  Urban exploiters and adapters differed primarily in social structure and migratory status: exploiters were significantly more social and sedentary than urban adapters. Clear trends were also seen for dietary preferences along a gradient of increasing urbanization in Jerusalem, such that, with increasing urbanization, the proportion of granivorous species increased whereas the proportion of species feeding on invertebrates declined. In contrast, neither relative brain size nor behavioural flexibility, as measured by feeding innovations, differed significantly among urban exploiters and adapters in Israel or along the urbanization gradient in Jerusalem specifically. Main conclusions  The results of our study suggest that being successful in more vs. less urbanized environments in the city is not necessarily a factor of brain size nor of how flexible and behaviourally innovative the species is; rather, it depends on a combination of traits, including diet, degree of sociality, sedentariness and preferred nesting sites.

Multivariate analyses of brain composition in mammals, amphibians and fish have revealed the evolution of ‘cerebrotypes’ that reflect specific niches and/or clades. Here, we present the first demonstration of similar cerebrotypes in birds. Using principal component analysis and hierarchical clustering methods to analyze a data set of 67 species, we demonstrate that five main cerebrotypes can be recognized. One type is dominated by galliforms and pigeons, among other species, that all share relatively large brainstems, but can be further differentiated by the proportional size of the cerebellum and telencephalic regions. The second cerebrotype contains a range of species that all share relatively large cerebellar and small nidopallial volumes. A third type is composed of two species, the tawny frogmouth (Podargus strigoides) and an owl, both of which share extremely large Wulst volumes. Parrots and passerines, the principal members of the fourth group, possess much larger nidopallial, mesopallial and striatopallidal proportions than the other groups. The fifth cerebrotype contains species such as raptors and waterfowl that are not found at the extremes for any of the brain regions and could therefore be classified as ‘generalist’ brains. Overall, the clustering of species does not directly reflect the phylogenetic relationships among species, but there is a tendency for species within an order to clump together. There may also be a weak relationship between cerebrotype and developmental differences, but two of the main clusters contained species with both altricial and precocial developmental patterns. As a whole, the groupings do agree with behavioral and ecological similarities among species. Most notably, species that share similarities in locomotor behavior, mode of prey capture or cognitive ability are clustered together. The relationship between cerebrotype and behavior/ecology in birds suggests that future comparative studies of brain-behavior relationships will benefit from adopting a multivariate approach.

The possible relationships between relative brain size and developmental mode and between relative brain size and five measures of the length of the development period were tested across over 1400 species of birds. Using both conventional statistics and phylogenetically based comparative methods, significant differences in relative brain size were detected among modes of development. Across all species, there were significant relationships between relative brain size and each of the following developmental traits: incubation period, age of fledging, duration of postfledging parental care, and total period of parental care. In contrast, the age of first flight was not significantly correlated with relative brain size. The relationships between these five developmental traits and relative brain size varied among developmental modes and orders such that significant relationships were present within some modes and orders but not in others. Thus, developmental differences play a significant role in the evolution of brain-size differences, but the role depends upon the taxonomic level being investigated. This is likely due to the differential lengths of periods of neural and behavioural development in young birds. Our conclusions support the contention of previous studies that developmental differences have played a key role in avian brain evolution.

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.

Endocranial volumes of vertebrate skulls and brain masses are often used interchangeably in comparative analyses of brain size. We test whether endocranial volume can be used as a reliable estimate of brain size in birds by comparing endocranial volumes with brain masses across 82 species using absolute values and with respect to body size. The results of paired tests across all 82 species and within two orders, Passeriformes and Psittaciformes, did not yield a significant difference between the two measures. These results were supported by correlational analyses that showed a significant positive relationship between endocranial volume and brain mass. Unpaired tests within short-tailed shearwaters (Puffinus tenuirostris) and paired tests within budgerigars (Melopsittacus undulatus) also yielded no significant differences between endocranial volume and brain mass. Thus, a combination of interspecific and intraspecific comparisons indicates that endocranial volume does provide a reliable estimate of brain size. Although this may enable more rapid collection of avian brain size data, endocranial volume should be used with caution because it cannot account for seasonal and age-related variation and cannot be used to measure differences in brain structure.