Selection on individuals of introduced species starts before the actual introduction.

Biological invasion is a global problem with large negative impacts on ecosystems and human societies. When a species is introduced, individuals will first have to pass through the invasion stages of uptake and transport, before actual introduction in a non-native range. Selection is predicted to act during these earliest stages of biological invasion, potentially influencing the invasiveness and/or impact of introduced populations. Despite this potential impact of pre-introduction selection, empirical tests are virtually lacking. To test the hypothesis of pre-introduction selection, we followed the fate of individuals during capture, initial acclimation, and captivity in two bird species with several invasive populations originating from the international trade in wild-caught pets (the weavers Ploceus melanocephalus and Euplectes afer). We confirm that pre-introduction selection acts on a wide range of physiological, morphological, behavioral, and demographic traits (incl. sex, age, size of body/brain/bill, bill shape, body mass, corticosterone levels, and escape behavior); these are all traits which likely affect invasion success. Our study thus comprehensively demonstrates the existence of hitherto ignored selection acting before the actual introduction into non-native ranges. This could ultimately change the composition and functioning of introduced populations, and therefore warrants greater attention. More knowledge on pre-introduction selection also might provide novel targets for the management of invasive species, if pre-introduction filters can be adjusted to change the quality and/or quantity of individuals passing through such that invasion probability and/or impacts are reduced.

Experimental evidence that matching habitat choice drives local adaptation in a wild population.

Matching habitat choice is a unique, flexible form of habitat choice based on self-assessment of local performance. This mechanism is thought to play an important role in adaptation and population persistence in variable environments. Nevertheless, the operation of matching habitat choice in natural populations remains to be unequivocally demonstrated. We investigated the association between body colour and substrate use by ground-perching grasshoppers (Sphingonotus azurescens) in an urban mosaic of dark and pale pavements, and then performed a colour manipulation experiment to test for matching habitat choice based on camouflage through background matching. Naturally, dark and pale grasshoppers occurred mostly on pavements that provided matching backgrounds. Colour-manipulated individuals recapitulated this pattern, such that black-painted and white-painted grasshoppers recaptured after the treatment aggregated together on the dark asphalt and pale pavement, respectively. Our study demonstrates that grasshoppers adjust their movement patterns to choose the substrate that confers an apparent improvement in camouflage given their individual-specific colour. More generally, our study provides unique experimental evidence of matching habitat choice as a driver of phenotype-environment correlations in natural populations and, furthermore, suggests that performance-based habitat choice might act as a mechanism of adaptation to changing environments, including human-modified (urban) landscapes.

Biased movement drives local cryptic coloration on distinct urban pavements.

Explanations of how organisms might adapt to urban environments have mostly focused on divergent natural selection and adaptive plasticity. However, differential habitat choice has been suggested as an alternative. Here, we test for habitat choice in enhancing crypsis in ground-perching grasshoppers colonizing an urbanized environment, composed of a mosaic of four distinctly coloured substrates (asphalt roads and adjacent pavements). Additionally, we determine its relative importance compared to present-day natural selection and phenotypic plasticity. We found that grasshoppers are very mobile, but nevertheless approximately match the colour of their local substrate. By manipulating grasshopper colour, we confirm that grasshoppers increase the usage of those urban substrates that resemble their own colours. This selective movement actively improves crypsis. Colour divergence between grasshoppers on different substrates is not or hardly owing to present-day natural selection, because observed mortality rates are too low to counteract random substrate use. Additional experiments also show negligible contributions from plasticity in colour. Our results confirm that matching habitat choice can be an important driver of adaptation to urban environments. In general, studies should more fully incorporate that individuals are not only selective targets (i.e. selected on by the environment), but also selective agents (i.e. selecting their own environments).

Selection on a behaviour-related gene during the first stages of the biological invasion pathway.

Human-induced biological invasions are common worldwide and often have negative impacts on wildlife and human societies. Several studies have shown evidence for selection on invaders after introduction to the new range. However, selective processes already acting prior to introduction have been largely neglected. Here, we tested whether such early selection acts on known behaviour-related gene variants in the yellow-crowned bishop (Euplectes afer), a pet-traded African songbird. We tested for nonrandom allele frequency changes after trapping, acclimation and survival in captivity. We also compared the native source population with two independent invasive populations. Allele frequencies of two SNPs in the dopamine receptor D4 (DRD4) gene-known to be linked to behavioural activity in response to novelty in this species-significantly changed over all early invasion stages. They also differed between the African native population and the two invading European populations. The two-locus genotype associated with reduced activity declined consistently, but strongest at the trapping stage. Overall genetic diversity did not substantially decrease, and there is little evidence for new alleles in the introduced populations, indicating that selection at the DRD4 gene predominantly worked on the standing genetic variation already present in the native population. Our study demonstrates selection on a behaviour-related gene during the first stages of a biological invasion. Thus, pre-establishment stages of a biological invasion do not only determine the number of propagules that are introduced (their quantity), but also their phenotypic and genetic characteristics (their quality).

Seed dispersal by macaws shapes the landscape of an Amazonian ecosystem.

Seed dispersal is one of the most studied plant-animal mutualisms. It has been proposed that the dispersal of many large-seeded plants from Neotropical forests was primarily conducted by extinct megafauna, and currently by livestock. Parrots can transport large fruits using their beaks, but have been overlooked as seed dispersers. We demonstrate that three macaws (Ara ararauna, A. glaucogularis and A. severus) are the main dispersers of the large-seeded motacú palm Attalea princeps, which is the biomass-dominant tree in the Bolivian Amazonian savannas. Macaws dispersed fruits at high rates (75-100% of fruits) to distant (up to 1200 m) perching trees, where they consumed the pulp and discarded entire seeds, contributing to forest regeneration and connectivity between distant forests islands. The spatial distribution of immature palms was positively associated to the proximity to macaws' perching trees and negatively to the proximity to cattle paths. The disperser role of livestock, presumably a substitute for extinct megafauna, had little effect due to soil compaction, trampling and herbivory. Our results underscore the importance of macaws as legitimate, primary dispersers of large-seeded plants at long distances and, specifically, their key role in shaping the landscape structure and functioning of this Amazonian biome.