An experimental examination of colonization as a generator of biotic nestedness

Species assemblages of systems of islands and other fragmented habitats frequently show a "nested-subset" structure in which the biotas of sites with low species richness are non-random subsets of the biotas of richer sites. Much literature suggests that extinction is more likely to produce strongly nested patterns than colonization, although very few experiments have been conducted on the generation of nested-subset patterns. Here, we describe an experiment on nestedness of benthic invertebrates occupying rocks in the littoral zone of a lake in western Victoria, Australia. Data collected from previous work indicated that the invertebrate biotas of rocks were nested. We used initially defaunated, different-sized habitat patches (half house-bricks, full house-bricks, double house-bricks) and followed the time-course of occupation to assess whether nested-subset patterns would emerge, and whether extinction or colonization best accounted for the patterns. We predicted that nestedness would increase through time up to the end of the experiment. This was found to be the case with colonization dominating the establishment of a strongly nested system; extinction appeared to be of relatively little importance over the duration of the experiment. These results suggest that at least in some circumstances, differential colonization may be influential in producing nested-subset patterns but experiments conducted over longer times may be needed to more completely understand the respective roles of extinction and colonization in generating nested-subset patterns in this system.

Nestedness in fragmented landscapes: birds of the box-ironbark forests of south-eastern Australia

Nestedness in biota as a function of species richness – biota of depauperate assemblages being non-random subsets of richer biotas – has been widely documented in recent years (see Wright et al. 1998, Oecologia 113: 1–20). Ordering sites by richness maximizes nestedness indices; however, ordering by other criteria such as area or isolation may be more ecologically interpretable. We surveyed birds in true fragments (35 in all), and in "reference areas" in large extant forest blocks (30 locations), of the same range of areas (10, 20, 40, 80 ha). The avifauna was divided into "bush birds"– species dependent on forest and woodland, and "open country" species. We looked at nestedness in four data sets: "bush birds" in fragments and reference areas, and "all birds" in fragments and in reference areas. All data sets were significantly nested. Ordering by area in all cases was not significantly less nested than ordering by richness. Ordering by area in fragments was significantly greater than in reference areas, but the differences in standardized nestedness indices were small (<15%). We identified those birds that had distributions among fragments that conformed strongly with area, those that were more randomly distributed and some species that were more likely to occupy the smallest fragments. Among the latter was a hyperaggressive, invasive, colonial native species (noisy miner Manorina melanocephala). A suite of small, insectivorous birds were more likely to strongly conform with expected distributions in relation to area, which was consistent with observations of their vulnerability to the effects of the noisy miner in smaller fragments.