Suzanne M. Prober

Conservation of the Grassy White Box Woodlands: Relative Contributions of Size and Disturbance to Floristic Composition and Diversity of Remnants

Before European settlement, grassy white box woodlands were the dominant vegetation in the east of the wheat-sheep belt of south-eastern Australia. Tree clearing, cultivation and pasture improvement have led to fragmentation of this once relatively continuous ecosystem, leaving a series of remnants which themselves have been modified by livestock grazing. Little-modified remnants are extremely rare. We examined and compared the effects of fragmentation and disturbance on the understorey flora of woodland remnants, through a survey of remnants of varying size, grazing history and tree clearing. In accordance with fragmentation theory, species richness generally increased with remnant size, and, for little-grazed remnants, smaller remnants were more vulnerable to weed invasion. Similarly, tree clearing and grazing encouraged weed invasion and reduced native species richness. Evidence for increased total species richness at intermediate grazing levels, as predicted by the intermediate disturbance hypothesis, was equivocal. Remnant quality was more severely affected by grazing than by remnant size. All little-grazed remnants had lower exotic species abundance and similar or higher native species richness than grazed remnants, despite their extremely small sizes (< 6 ha). Further, small, littlegrazed remnants maintained the general character of the pre-European woodland understorey, while grazing caused changes to the dominant species. Although generally small, the little-grazed remnants are the best representatives of the pre-European woodland understorey, and should be central to any conservation plan for the woodlands. Selected larger remnants are needed to complement these, however, to increase the total area of woodland conserved, and, because most little-grazed remnants are cleared, to represent the ecosystem in its original structural form. For the maintenance of native plant diversity and composition in little-grazed remnants, it is critical that livestock grazing continues to be excluded. For grazed remnants, maintenance of a site in its current state would allow continuation of past management, while restoration to a pre-European condition would require management directed towards weed removal, and could take advantage of the difference noted in the predominant life-cycle of native (perennial) versus exotic (annual or biennial) species.

Integrative modelling reveals mechanisms linking productivity and plant species richness

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

Identifying ecological barriers to restoration in temperate grassy woodlands: soil changes associated with different degradation states

Temperate grassy woodlands were once the dominant vegetation across many agricultural regions of south-eastern Australia, but most of these are now highly degraded and fragmented. Adequate conservation of these woodlands is dependent on successful ecological restoration; however, ecological barriers often limit ecosystem recovery once degrading processes are removed. To help identify these barriers, we used a state and transition framework to compare topsoils of little-disturbed (reference) and variously degraded remnants of grassy Eucalyptus albens Benth. and E. melliodora Cunn. ex Schauer woodlands. Topsoils of degraded remnants showed a repeated pattern, with the most compacted, most acidic and most depleted topsoils occurring in remnants dominated by Aristida ramosa R.Br. or Austrodanthonia H.P.Linder and Austrostipa scabra (Lindl.) S.W.L.Jacobs & J.Everett; the least compacted and most nutrient rich topsoils in remnants dominated by annual exotics; and generally intermediate topsoils in remnants dominated by Bothriochloa macra (Steud.) S.T.Blake or Austrostipa bigeniculata (Hughes) S.W.L.Jacobs & J.Everett. Surprisingly, topsoils beneath trees in reference sites (supporting Poa sieberiana Spreng.) were similar to topsoils supporting annual exotics for most soil properties. Chemical properties of topsoils from open areas of reference sites [supporting Themeda australis (R.Br.) Stapf] were usually intermediate and similar to Bothriochloa macra and Austrostipa bigeniculata topsoils. The most striking exception to these trends was for soil nitrate, which was extremely low in all reference topsoils and showed a high correlation with annual exotic abundance. We discuss the potential for positive feedbacks between soil nitrogen cycling and understorey composition and the need for intervention to assist possible nitrate-dependent transitions between annual and perennial understorey states. Dominant grasses, trees and annual weed abundance may be useful indicators of soil conditions and could inform selection of target sites, species and techniques for restoration projects.

Regional Contingencies in the Relationship between Aboveground Biomass and Litter in the World's Grasslands

Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex.

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4–8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Abundance of introduced species at home predicts abundance away in herbaceous communities

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Australian Aboriginal Peoples’ Seasonal Knowledge: a Potential Basis for Shared Understanding in Environmental Management

Natural resource scientists and managers increasingly recognize traditional ecological knowledge (TEK) for its potential contribution to contemporary natural resource management (NRM) and, through this, to more resilient social-ecological systems. In practice, however, inadequate cross-cultural means to organize and communicate TEK can limit its effective inclusion in management decisions. Indigenous seasonal knowledge involving temporal knowledge of biota, landscapes, weather, seasonal cycles, and their links with culture and land uses is one type of TEK relevant to this issue. We reviewed the literature on Australian Aboriginal seasonal knowledge to characterize contemporary and potential applications to NRM. This knowledge was often documented through cross-cultural collaboration in the form of ecological calendars. Our analysis revealed a variety of basic and applied environmental information in Aboriginal seasonal descriptions and calendars that can contribute directly to NRM. Documented applications have been limited to date, but include fire management, inclusion as general material in NRM plans, and interpretative information about environments. Emerging applications include water management and climate change monitoring. Importantly, seasonal knowledge can also contribute indirectly to NRM outcomes by providing an organizing framework for the recovery, retention, and cross- cultural communication of TEK and linking to its broader cultural and cosmological contexts. We conclude that by facilitating the combination of experiential with experimental knowledge and fostering complementarity of different knowledge systems, Aboriginal seasonal knowledge can increasingly contribute to more resilient social-ecological outcomes in NRM. Nevertheless, the seasonal framework should augment, rather than override, other approaches to cross-cultural NRM such as those with spatial and/or social-ecological emphasis.

Species abundance at home predicts abundance away in grasslands

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Plasticity of functional traits varies clinally along a rainfall gradient in Eucalyptus tricarpa

Widespread species often occur across a range of climatic conditions, through a combination of local genetic adaptations and phenotypic plasticity. Species with greater phenotypic plasticity are likely to be better positioned to cope with rapid anthropogenic climate changes, while those displaying strong local adaptations might benefit from translocations to assist the movement of adaptive genes as the climate changes. Eucalyptus tricarpa occurs across a climatic gradient in south-eastern Australia, a region of increasing aridity, and we hypothesized that this species would display local adaptation to climate. We measured morphological and physiological traits reflecting climate responses in nine provenances from sites of 460 to 1040 mm annual rainfall, in their natural habitat and in common gardens near each end of the gradient. Local adaptation was evident in functional traits and differential growth rates in the common gardens. Some traits displayed complex combinations of plasticity and genetic divergence among provenances, including clinal variation in plasticity itself. Provenances from drier locations were more plastic in leaf thickness, whereas leaf size was more plastic in provenances from higher rainfall locations. Leaf density and stomatal physiology (as indicated by δ(13)C and δ(18)O) were highly and uniformly plastic. In addition to variation in mean trait values, genetic variation in trait plasticity may play a role in climate adaptation.

Genome-wide scans detect adaptation to aridity in a widespread forest tree species

Patterns of adaptive variation within plant species are best studied through common garden experiments, but these are costly and time‐consuming, especially for trees that have long generation times. We explored whether genome‐wide scanning technology combined with outlier marker detection could be used to detect adaptation to climate and provide an alternative to common garden experiments. As a case study, we sampled nine provenances of the widespread forest tree species, Eucalyptus tricarpa, across an aridity gradient in southeastern Australia. Using a Bayesian analysis, we identified a suite of 94 putatively adaptive (outlying) sequence‐tagged markers across the genome. Population‐level allele frequencies of these outlier markers were strongly correlated with temperature and moisture availability at the site of origin, and with population differences in functional traits measured in two common gardens. Using the output from a canonical analysis of principal coordinates, we devised a metric that provides a holistic measure of genomic adaptation to aridity that could be used to guide assisted migration or genetic augmentation.