Interference competition between introduced black rats and endemic Galápagos rice rats.

Replicated field experiments were used to quantify and to describe the mechanism of competition between the introduced black rat Rattus rattus and the endemic Santiago rice rat Nesoryzomys swarthi on Santiago Island, Galápagos Islands, Ecuador. The removal of R. rattus significantly slowed the rate of seasonal population decline in N. swarthi. This effect was particularly evident for female, relative to male, N. swarthi and appeared to be driven solely by enhanced immigration; no other fitness or space use parameters were affected. The candidate hypotheses to explain the mechanism of competition were exploitation competition, interference by resource defense, and interference by aggressive encounter. To distinguish between hypotheses, we conducted a replicated resource supplementation experiment with patchy food, scattered food, and no food (control) treatments. The opportunistic R. rattus responded to the extra resources with increased adult immigration and juvenile recruitment, resulting in a significant abundance boost of sevenfold on patchy grids and fourfold on scattered grids. Females increased in body mass, and the breeding season was lengthened. In contrast, there was no change in the abundance of N. swarthi and no obvious benefit to reproduction. Instead, the costs of interference apparently outweighed the benefits of extra food: female N. swarthi increased in mass with supplementary food, but female (relative to male) immigration and residency were repressed on all supplemented areas. This response supported the hypothesis of interference by aggressive encounter, and we were able to rule out the alternative hypotheses. Although periodic population crashes of R. rattus on the arid north coast of Santiago may ameliorate its competitive impact, climate change may tip the balance. Control or eradication of R. rattus should improve future survival prospects for N. swarthi, but wildlife managers must be prepared for the potential eruption of the introduced house mouse Mus musculus, because this species experienced a release from interference competition and immigrated to removal areas.

Space invaders? A search for patterns underlying the coexistence of alien black rats and Galápagos rice rats.

The introduction and spread of the black rat Rattus rattus is believed to have caused the worst decline of any vertebrate taxon in Galápagos. However, the "extinct" Santiago rice rat Nesoryzomys swarthi has recently been rediscovered in sympatry with R. rattus providing the first exception to this general pattern of displacement. We carried out an exploratory investigation of this novel system with the aim of identifying patterns that may facilitate the apparent coexistence of the two species. We carried out an extensive survey of Santiago Island to map the current distribution of the endemic rice rat and to explore broad scale distribution-habitat associations. We then used live-trapping, radio-tracking, and spool-and-line tracking to quantify abundance-habitat correlations and to test for evidence of interspecific spatial segregation, alteration of N. swarthi activity patterns (spatial and temporal), and microhabitat partitioning. We found that N. swarthi has disappeared from part of its historical range and appears to be restricted to a 14 km stretch of the north-central coast, characterised by high density of the cactus Opuntia galapageia. In contrast, the generalist R. rattus was found at all survey sites. We found no evidence of spatial segregation, and home range size, temporal activity and density of N. swarthi did not vary with local density of R. rattus. However, pre-dawn and post-dusk N. swarthi activity levels increased with R. rattus density perhaps reflecting an increase in foraging effort necessary to compensate for the costs of interspecific exploitation or interference competition. The distribution, microhabitat selection, and abundance-habitat relations of N. swarthi suggest that the endemic cactus O. galapageia may facilitate interspecific coexistence. Further research should include a comparison of inter-seasonal resource preference and foraging activity of the two species coupled with replicated field experiments to confirm and quantify competition and to elucidate the mechanism of competitive coexistence.