Determinants of regional and local patterns in the floras of braided riverbeds in New Zealand

Aim To determine the composition and biogeographical origins of the native and naturalized flora of braided shingle riverbeds in New Zealand and whether the proportion of naturalized species is greater than in similar habitats elsewhere in the world. To test whether regional species pools, landscape matrix configuration, and local riverbed environment are all equally important in determining community structure in these systems. Location The braided reaches of four catchments on each of the eastern and western sides of North Island and South Island, New Zealand. Methods Plotless records of all native and naturalized seed plant and fern species on disturbed mobile surfaces were made over the length of four rivers. Altitude, climate variables, riverbed attributes, and surrounding land-cover from maps were recorded for each site. For all species, the taxonomy, life form, origin and history, and presence within the province through which the rivers flowed, was obtained from published floras. Direct and indirect ordination and variance partitioning were employed to examine how native and naturalized species composition varies among and within rivers, and the degree to which this variation reflects climate, characteristic of riverbeds, and the surrounding land-cover. Regression was used to determine how much introduction date and native geographical range influence the frequency of naturalized species. Results The total riverbed flora of 289 species comprises 40% native species and 60% naturalized species, both dominated by Asteraceae and Poaceae. The relative contributions of other plant families differ, and the two groups comprise different life-form spectra. Native species occur across fewer rivers (mean 1.6 rivers) than naturalized species (mean 2.1 rivers). Species common in at least one river system tend to be widespread, occurring in at least three rivers. The rivers differ in their floras, with distinctions between North Island and South Island, and eastern and western rivers. The South Island rivers have more native species and a higher proportion of their regional native species pools than North Island rivers, whereas they have a lower proportion of their naturalized species pools. Introduction date and native geographical range are correlated with frequency of individual naturalized species. Geographical position and climate, riverbed variables of substrate size and seepage presence, and the type of adjacent land-cover, differ between rivers and are significantly related to species composition. Land-cover variables alone account for 46% of the explained variation in species composition, environmental variables alone 32% and components of the environment and land-cover that co-vary, 22%. Main conclusions New Zealand shingle riverbeds are among the riparian communities in the world most invaded by naturalized species but comparisons are difficult because studies of similar riverbeds are rare. New Zealand riverbeds are dominated by short-lived terrestrial species from Eurasia. Native species are most frequent in South Island rivers draining extensive high mountains, and their abundances are determined to a greater extent by riverbed processes than are those of the naturalized species, which are more abundant when the surrounding landscape is modified. The distribution of the naturalized flora is not yet at equilibrium with the environment. Each river has a distinctive flora determined by ambient environmental factors, aspects of the riverbed environment itself, adjacent land-cover reflecting the presence of native vegetation, the intensity of human modification or use, and invasion history.