Turf wars: experimental tests for alternative stable states in a two-phase coastal ecosystem.

Alternative stable states have long been thought to exist in natural communities, but direct evidence for their presence and for the environmental switches that cause them has been scarce. Using a combination of greenhouse and field experiments, we investigated the environmental drivers associated with two distinctive herbaceous communities in coastal ecosystems in New Zealand. In a mosaic unrelated to micro-topography, a community dominated largely by native turf species (notably Leptinella dioica, Samolus repens, and Selliera radicans) alternates with vegetation comprising exotic (i.e., nonnative) pasture species (notably Agrostis stolonifera, Holcus lanatus, Lolium perenne, and Trifolium repens). The species of these two communities differ in functional characters related to leaf longevity and growth rate, and occupy soils of differing nitrogen levels. Both spatial and environmental factors influenced the species composition locally. Reciprocal transplants of soil, with and without associated vegetation, showed that a native turf community developed when sward or soil from either community was bounded by turf, and a pasture community developed when sward or soil from either community was surrounded by pasture. In artificial mixed communities in the greenhouse, turf was able to invade the pasture community where the vegetation was clipped to simulate grazing, and also where Trifolium was removed and/or salt spray was applied. The pasture community invaded the turf where Trifolium was present or nitrogen was added. These results were supported by trends in experimentally manipulated field plots, where the amount of turf cover increased when nitrogen was kept low and when salt spray was applied, whereas pasture cover increased in the absence of salt spray. Thus, persistence of the native turf community is dependent on grazing, both directly and via its effect on keeping nitrogen levels low by excluding the exotic, nitrogen-fixing Trifolium, and by exposing the vegetation to salt spray. If any of these factors change, there could be a state change to pasture dominance that might be resistant to reversion to turf. Managing such coastal herbaceous communities therefore requires an understanding of the environmental and species characteristics that maintain alternative states.

Is New Zealand vegetation really 'problematic'? Dansereau's puzzles revisited.

Over four decades ago, Pierre Dansereau, the noted North American ecologist, proposed six features of New Zealand vegetation as being problematic or unusual in a global context. We examine his propositions in the light of current ecological knowledge to determine whether or not these can still be considered unusual characteristics of New Zealand vegetation. (1) 'Climatic change is still progressing' resulting in disequilibrium between species' distributions and the present climate. New data and methods of analysis now available have removed the impression that Dansereau gained of imprecise zonation, unclear vegetation/climate relations and missing vegetation types. Communities cited as having regeneration failure can now be seen as even-aged stands that developed after major disturbance, although there are other, also non-climatic, explanations. However, the cause of the Westland 'Nothofagus gap' has become more, rather than less, controversial. (2) 'Continuity of community composition defies classification' and 'Very few New Zealand associations have faithful species' are correct observations, but perhaps equally true of vegetation elsewhere. Dansereau's assertion of low species richness in New Zealand is not supported by the comparative data available. (3) 'Lack of intolerant [i.e. mid-seral] trees …' is not evident with newer information. The order of species in succession, seen as unclear by Dansereau, has been determined by a range of approaches, largely confirming each other. (4) 'Discrepancies of form and function …' in divaricate shrubs and widespread heteroblasty are still controversial, with many more explanations. Several abiotic explanations have failed to stand up to investigation. Explanations in terms of herbivory have been well supported, although the extinction of the large avian herbivores makes certainty impossible. (5) 'Incidence of hybridization …' remains problematic. We do not know whether the incidence is unusually high, as Dansereau alleged, but the limited comparative data available suggest not. (6) The 'overwhelming … competing power of exotics' is strongly context dependent. They are prominent in many non-forest habitats. It seems that they are drivers of the vegetation change in some habitats, yet passengers after disturbance in others. Invasions can be slow, and may still be very incomplete in some ecosystem types. Whether exotics will eventually take over in most communities, or whether the native species will 'laugh them to scorn' as Cockayne suggested, only time will tell. In conclusion, some aspects of New Zealand's vegetation seem less unusual with increased knowledge, but others remain 'problems'.

Does niche overlap control relative abundance in French lacustrine fish communities? A new method incorporating functional traits.

1. The mechanisms that structure biological communities hold the key to understanding ecosystem functioning and the maintenance of biodiversity. Patterns of species abundances have been proposed as a means of differentiation between niche-based and neutral processes, but abundance information alone cannot provide unequivocal discrimination. 2. We combined species niche information and species' relative abundances to test the effects of two opposing structuring mechanisms (environmental filtering and niche complementarity) on species' relative abundances in French lacustrine fish communities. The test involved a novel method comparing the abundance-weighted niche overlap within communities against that expected when relative abundances were randomized among species within the community. 3. Observed overlap was consistently significantly lower than expected at random for two (swimming ability and trophic status) of four primary niche axes across lakes of differing physical environments. Thus, for these niche axes, pairs of abundant species tended to have relatively low niche overlap, while rare species tended to have relatively high niche overlap with abundant species. 4. This suggests that niche complementarity may have acted to enhance ecosystem function and that it is important for species coexistence in these fish communities. The method used may be easily applied to any sort of biological community and thus may have considerable potential for determining the generality of niche complementarity effects on community structure.

Is there really insufficient support for Tilman's R* concept? A comment on Miller et al.

Miller et al. (2005), in the American Naturalist (165:439-448), critically reviewed the applicability of Tilman's resource-ratio hypothesis. One of their conclusions was that only eight experimental papers support the R* concept, while five do not. We are familiar with some of the latter studies, and we question this conclusion. Our evaluation shows that 12 of the 13 articles investigated by Miller et al. support R* prediction, while one article does not fit the experimental conditions for a proper test. Moreover, the microbial and aquatic literature contains many more competition experiments consistent with the R* prediction. We therefore conclude that there is strong experimental support for the R* concept, at least from studies with bacteria, phytoplankton, and zooplankton.

Is the abundance of species determined by their functional traits? A new method with a test using plant communities.

The relation between functional traits and abundance of species has the potential to provide evidence on the mechanisms that structure local ecological communities. The niche-limitation/limiting-similarity hypothesis, derived from MacArthur and Levins' original concept, predicts that species that are similar to others in terms of functional traits will suffer greater competition and hence be less abundant. On the other hand, the environment-filtering/habitat-optimum hypothesis predicts that groups of species with functional traits that are close to the optimum for that environment, and are therefore similar to other species, will be more abundant. We propose a new niche-assembly model for predicting the relative abundance of species in communities from their functional traits, which can detect the patterns that would be expected from either of these hypotheses. The model was fitted to eight plant communities sampled in the Lake Ohau district of New Zealand. For seven of the sites, the patterns could not be distinguished from that expected under a null model. However, in one site there was highly significant departure from the null model in the direction expected from the niche-limitation hypothesis. The site was probably the most productive of those examined. It is possible that competition for light rather than belowground resources, or faster recovery from disturbance, allowed greater predictability. Surprisingly, the predictability was seen when just the presences of a species' neighbours in trait space were taken into account, but not when the potential effects of those neighbours were weighted by their abundance. For three of the four model types, the effects of species on each other were consistently negative: a significant trend. These results contradict the various neutral models of ecological communities.

Niche overlap estimates based on quantitative functional traits: a new family of non-parametric indices.

The concept of niche overlap appears in studies of the mechanisms of the maintenance of species diversity, in searches for assembly rules, and in estimation of within-community species redundancy. For plant traits measured on a continuous scale, existing indices are inadequate because they split the scale into a number of categories thus losing information. An index is easy to construct if we assume a normal distribution for each trait within a species, but this assumption is rarely true. We extend and apply an index, NO(K), which is based on kernel density functions, and can therefore work with distributions of any shape without prior assumptions. For cases where the ecologist wishes to downweight traits that are inter-correlated, we offer a variant that does this: NO(Kw). From either of these indices, an index of the mean niche overlap in a community can be calculated: NO(K,community) and NO(Kw,community). For all these indices, the variance can be calculated and formulae for this are given. To give examples of the new indices in use, we apply them to a coastal fish dataset and a sand-dune plant dataset. The former exhibits considerable non-normality, emphasising the need for kernel-based indices. Accordingly, there was a considerable difference in index values, with those for an index based on a normal distribution being significantly higher than those from an index which, being based on kernel fitting, is not biased by an assumption for the distribution. The NO(K) values were ecologically consistent for the fish species concerned, varying from 0.02 to 0.53. The sand-dune plant data also showed a wide range of overlap values. Interestingly, the least overlap was between two graminoids, which would have been placed in the same functional group in the coarse classification often used in functional-type/ecosystem-function work.

Functional regularity: a neglected aspect of functional diversity.

Functional diversity has been identified as a key to understanding ecosystem and community functioning. However, due to the lack of a sound definition its nature and measurement are still poorly understood. In the same way that species diversity can be split into species richness and species evenness, so functional diversity can be split into functional richness (i.e. the amount of functional trait/character/attribute space filled) and functional evenness (i.e. the evenness of abundance distribution in functional trait space). We propose a functional regularity index (FRO) as a measure of functional evenness for situations where species are represented only by a single functional trait value (e.g. mean, median or mode), and species abundances are known. This new index is based on the Bulla O index of species evenness. When dealing with functional types or categorical functional traits, the Bulla O or any other accepted species evenness index may be used directly to measure functional evenness. The advantage of FRO is that it supplies a measure of functional evenness for continuous trait data. The FRO index presented in this paper fulfils all the a priori criteria required. We demonstrate with two example datasets that a range of FRO values may be obtained for both plant and animal communities. Moreover, FRO was strongly related to ecosystem function as seen in photosynthetic biomass in plant communities, and was able to differentiate sampling stations in a lagoon based on the functional traits of fish. Thus, the FRO index is potentially a highly useful tool for measuring functional diversity in a variety of ecological situations.

Can we tell how a community was constructed? A comparison of five evenness indices for their ability to identify theoretical models of community construction.

Evenness indices provide a simple measure of community structure. It might therefore be possible to use them to identify the process by which a community has been assembled. To test whether this is practicable, we constructed simulated community samples using five stochastic models of community construction, proposed by Tokeshi. We then examined the ability of five evenness indices to identify the model under which the community samples were produced. Each model produced samples with a range of evenness values, but mean evenness differed between the models. The dominance decay (DD) model produced community samples with the greatest evenness, followed by MacArthur fraction (MF). Evenness was lowest under the dominance pre-emption (DP) model. The differences between models were quite consistent across indices and consistent between 5-species and 15-species communities. Samples produced under the DD and MF models varied least in evenness between 5-species and 15-species samples, and those produced under the random assortment (RA) model varied most, irrespective of the evenness index used. Evenness varied considerably between replicate samples, as expected with stochastic processes. In 5-species communities, the greatest robustness in evenness across replicates was seen using indices O or E'. In 15-species communities, O and E(Q) were the most robust. The index best able to identify the model which had generated the sample differed between models and with species richness. If the model of interest is not known in advance, the best index for identifying the generating model is E(var) for communities of more than 10 species for RF, RA and DP models and O for other communities. The number of samples required in a data set before it could be effectively identified was, for example, more than 30 for 5-species samples produced under the random fraction (RF) model, and 3 for 15-species RF samples. In contrast, a 5-species DD data set could be effectively identified when it contained 15 or more samples. We conclude that evenness can be used to identify the process by which the community has been constructed, out of the five models considered here. The best evenness index for doing this varies with the species richness and the evenness of the community but we can suggest the use of O without knowing the model because its the more stable one against species richness and the more robust and unbiased against simulated samples and it encompass a good discriminating power between the models in most of the cases studied.

The myth of constant predator: prey ratios.

Apparent constancy in the ratio of predator species to prey species has been offered as evidence that ecological communities are structured by interspecific interactions. If significantly different from random expectation, this effect would be one of the few sound pieces of evidence for community structure. The evidence was re-evaluated by using the data from previous studies to form species pools, and forming simulated 'communities' by drawing species at random from these pools (with replacement). Using a correlation coefficient (number of predator species versus number of prey species), and also the statistic used by the original workers (where different), the observed predator:prey correlation was compared to that for the random communities. In five studies, the observed predator:prey ratio was not significantly different from random expectation. In the only two studies where there was significant departure from the null model, it was with more variation in the ratio than expected on a random basis. It is concluded that there is as yet no evidence for near-constant predator:prey ratios.

A null model of guild proportionality, applied to stratification of a New Zealand temperate rain forest.

Much ecological theory assumes that the number of species that can coexist (by 'species packing') is limited, because competitive exclusion occurs when any pair of species within a guild is too similar - 'species saturation' or 'niche limitation'. If such niche limitation occurs, the proportion of species in each guild should be relatively constant - 'guild proportionality'. This concept is applied to the guilds represented by strata in a forest. A method is produced, and used to examine a New Zealand temperate rain forest. Most strata showed no deviation from a null model of no niche limitation, i.e. no tendency to guild proportionality. The proportion of lianes was more variable than in the null model, tending to be inversely related to the proportion of epiphytes, Canopy tree proportion was significantly more constant than in the null model, but this could be interpreted as a limit caused by the size of a canopy tree individual.

A comparison of realised niche relations of species in New Zealand and Britain.

Many plant species prominent in the native vegetation of the dry shingle banks at Dungeness (Britain) are also prominent as exotics in the dry Upper Clutha catchment (New Zealand). To examine the realised niche relations of these species, vegetation was sampled in the two areas. Inverse classification and ordination were used to determine the relative beta niches of the species in the two areas. There was little agreement; it seems that the exotic species in the Upper Clutha were pre-adapted to different niches from those in their native range.