The ecological impacts of commensal species: black rats, Rattus rattus, at the urban–bushland interface

Exotic species have had devastating impacts worldwide and are a major threat to native wildlife. Human commensal species (hereafter commensals) are a special class of exotic species that live largely off the resources associated with human activity. The encroachment of commensals from an urban area into surrounding bushland has been frequently overlooked as an important component of urban impacts, even though human-commensals are common to many urban regions globally. In this review, we present theoretical and empirical evidence for the processes and outcomes occurring when exotic commensal species encroach into native bushland. Specifically we ask when, how and why exotic commensal species encroach into bushland, what determines whether they establish, and what are the ecological consequences. We focus on the black rat, Rattus rattus, arguably the archetypal commensal species with a cosmopolitan distribution and the greatest potential for ecological damage of all the commensal rodents. We expect that the processes that we outline apply to other commensal species more broadly. We argue that commensals are in fact natives of the urban milieu and only become alien when they encroach into peri-urban bushland. We propose that the mechanisms of this encroachment will be different from those of other, non-commensal exotic species because urban areas act as dispersal hubs to overcome many of the barriers of invasion that other exotic species face. We suggest that resource supplementation by urban areas creates a great potential for promoting encroachment, invasion as well as impact. However, biotic and abiotic barriers to invasion are still relevant for commensals, highlighting the need to maintain the integrity of ecosystems and wildlife populations in urban edges so as to prevent commensal incursion. We examine how commensal black rats affect wildlife via three fundamental mechanisms, namely, predation, disease transfer and competition for resources, and also consider their possible positive impacts acting as functional replacements for lost natives. We conclude the review with an outline of research priorities and future directions that are essential for progressing our understanding of the ecology of commensal species.