Jodi N. Price

Advances in restoration ecology: rising to the challenges of the coming decades

Simultaneous environmental changes challenge biodiversity persistence and human wellbeing. The science and practice of restoration ecology, in collaboration with other disciplines, can contribute to overcoming these challenges. This endeavor requires a solid conceptual foundation based in empirical research which confronts, tests and influences theoretical developments. We review conceptual developments in restoration ecology over the last 30 years. We frame our review in the context of changing restoration goals which reflect increased societal awareness of the scale of environmental degradation and the recognition that inter-disciplinary approaches are needed to tackle environmental problems. Restoration ecology now encompasses facilitative interactions and network dynamics, trophic cascades, and above- and below ground linkages. It operates in a non-equilibrium, alternative states framework, at the landscape scale, and in response to changing environmental, economic and social conditions. Progress has been marked by conceptual advances in the fields of trait-environment relationships, community assembly, and understanding the links between biodiversity and ecosystem functioning. Conceptual and practical advances have been enhanced by applying evolving technologies, including treatments to increase seed germination and overcome recruitment bottlenecks, high throughput DNA sequencing to elucidate soil community structure and function, and advances in satellite technology and GPS tracking to monitor habitat use. The synthesis of these technologies with systematic reviews of context dependencies in restoration success, model based analyses and consideration of complex socio-ecological systems will allow generalizations to inform evidence based interventions. Ongoing challenges include setting realistic, socially acceptable goals for restoration under changing environmental conditions, and prioritizing actions in an increasingly space-competitive world. Ethical questions also surround the use of genetically modified material, translocations, taxon substitutions, and de-extinction, in restoration ecology. Addressing these issues, as the Ecological Society of America looks to its next century, will require current and future generations of researchers and practitioners, including economists, engineers, philosophers, landscape architects, social scientists and restoration ecologists, to work together with communities and governments to rise to the environmental challenges of the coming decades.

Functional species pool framework to test for biotic effects on community assembly

Functional trait differences among species are increasingly used to infer the effects of biotic and abiotic processes on species coexistence. Commonly, the trait diversity observed within communities is compared to patterns simulated in randomly generated communities based on sampling within a region. The resulting patterns of trait convergence and divergence are assumed to reveal abiotic and biotic processes, respectively. However, biotic processes such as competition can produce both trait divergence and convergence, through either excluding similar species (niche differences, divergence) or excluding dissimilar species (weaker competitor exclusion, convergence). Hence, separating biotic and abiotic processes that can produce identical patterns of trait diversity, or even patterns that neutralize each other, is not feasible with previous methods. We propose an operational framework in which the functional trait dissimilarity within communities (FDcomm) is compared to the corresponding trait dissimilarity expected from the species pool (i.e., functional species pool diversity, FDpool). FDpool includes the set of potential species for a site delimited by the operating environmental and dispersal limitation filters. By applying these filters, the resulting pattern of trait diversity is consistent with biotic processes, i.e., trait divergence (FDcomm > FDpool) indicates niche differentiation, while trait convergence (FDcomm < FDpool) indicates weaker competitor exclusion. To illustrate this framework, with its potential application and constraints, we analyzed both simulated and field data. The functional species pool framework more consistently detected the simulated trait diversity patterns than previous approaches. In the field, using data from plant communities of typical Northern European habitats in Estonia, we found that both niche-based and weaker competitor exclusion influenced community assembly, depending on the traits and community considered. In both simulated and field data, we demonstrated that only by estimating the species pool of a site is it possible to differentiate the patterns of trait dissimilarity produced by operating biotic processes. The framework, which can be applied with both functional and phylogenetic diversity, enables a reinterpretation of community assembly processes. Solving the challenge of defining an appropriate reference species pool for a site can provide a better understanding of community assembly.

Small-scale grassland assembly patterns differ above and below the soil surface

The existence of deterministic assembly rules for plant communities remains an important and unresolved topic in ecology. Most studies examining community assembly have sampled aboveground species diversity and composition. However, plants also coexist belowground, and many coexistence theories invoke belowground competition as an explanation for aboveground patterns. We used next-generation sequencing that enables the identification of roots and rhizomes from mixed-species samples to measure coexisting species at small scales in temperate grasslands. We used comparable data from above (conventional methods) and below (molecular techniques) the soil surface (0.1 x 0.1 x 0.1 m volume). To detect evidence for nonrandom patterns in the direction of biotic or abiotic assembly processes, we used three assembly rules tests (richness variance, guild proportionality, and species co-occurrence indices) as well as pairwise association tests. We found support for biotic assembly rules aboveground, with lower variance in species richness than expected and more negative species associations. Belowground plant communities were structured more by abiotic processes, with greater variability in richness and guild proportionality than expected. Belowground assembly is largely driven by abiotic processes, with little evidence for competition-driven assembly, and this has implications for plant coexistence theories that are based on competition for soil resources.

Drivers of change in the Social-Ecological Systems of the Gwydir Wetlands and Macquarie Marshes in northern New South Wales, Australia

The Murray–Darling Basin is a Social-Ecological System (SES) of major importance to Australia and includes extensive wetland areas in the north-western parts of New South Wales. The Gwydir Wetlands and the Macquarie Marshes are the particular focus of this paper. These two wetland SES have undergone five successive adaptive cycles (phases) since they were first visited by Europeans in the early 19th century and the ecological, economic and social drivers initiating each transformation to a new cycle are described and analysed. The arrival of the European settlers with their domestic livestock rapidly displaced the Indigenous SES and the wetlands were extensively grazed; during wet periods the livestock were moved out of the wetlands and moved back in as the water receded. More recent land-use changes resulted from the building of major dams to enable storage of water for use in irrigated agriculture. A consequence of dam construction and water use has been a reduction in the frequency and extent of flooding, which has allowed many parts of the wetlands to be continually grazed. Furthermore, as machinery capable of cultivating the very heavy textured soils became available, dryland cropping became a major enterprise in areas of the floodplain where the likelihood of flooding was reduced. With the reduction in flooding, these wetland sites have been seriously degraded. The final phase has seen the invasion by an exotic weed, lippia [Phyla canescens (Kunth) Greene], which is a perennial that grows mat-like between other species of plants and spreads to produce a virtually mono-specific stand. The domestic livestock carrying capacity of the land becomes more or less zero and the conservation value of the wetlands is also dramatically decreased. Therefore, we suggest that lippia should be classed as an ecosystem engineer that has caused the latest transformation of these wetland SES and suggest research directions to investigate how they can be managed to revert to a state in which lippia is no longer dominant.

Species-Specific Effects of Woody Litter on Seedling Emergence and Growth of Herbaceous Plants

The effect of litter on seedling establishment can influence species richness in plant communities. The effect of litter depends on amount, and also on litter type, but relatively little is known about the species-specific effects of litter. We conducted a factorial greenhouse experiment to examine the effect of litter type, using two woody species that commonly co-occur in boreonemoral forest—evergreen spruce (Picea abies), deciduous hazel (Corylus avellana), and a mixture of the two species—and litter amount—shallow (4 mm), deep (12 mm) and leachate—on seedling emergence and biomass of three understorey species. The effect of litter amount on seedling emergence was highly dependent on litter type; while spruce needle litter had a significant negative effect that increased with depth, seedling emergence in the presence of hazel broadleaf litter did not differ from control pots containing no litter. Mixed litter of both species also had a negative effect on seedling emergence that was intermediate compared to the single-species treatments. Spruce litter had a marginally positive (shallow) or neutral effect (deep) on seedling biomass, while hazel and mixed litter treatments had significant positive effects on biomass that increased with depth. We found non-additive effects of litter mixtures on seedling biomass indicating that high quality hazel litter can reduce the negative effects of spruce. Hazel litter does not inhibit seedling emergence; it increases seedling growth, and creates better conditions for seedling growth in mixtures by reducing the suppressive effect of spruce litter, having a positive effect on understorey species richness.

Seasonality and facilitation drive tree establishment in a semi-arid floodplain savanna

A popular hypothesis for tree and grass coexistence in savannas is that tree seedlings are limited by competition from grasses. However, competition may be important in favourable climatic conditions when abiotic stress is low, whereas facilitation may be more important under stressful conditions. Seasonal and inter-annual fluctuations in abiotic conditions may alter the outcome of tree–grass interactions in savanna systems and contribute to coexistence. We investigated interactions between coolibah (Eucalyptus coolabah) tree seedlings and perennial C4 grasses in semi-arid savannas in eastern Australia in contrasting seasonal conditions. In glasshouse and field experiments, we measured survival and growth of tree seedlings with different densities of C4 grasses across seasons. In warm glasshouse conditions, where water was not limiting, competition from grasses reduced tree seedling growth but did not affect tree survival. In the field, all tree seedlings died in hot dry summer conditions irrespective of grass or shade cover, whereas in winter, facilitation from grasses significantly increased tree seedling survival by ameliorating heat stress and protecting seedlings from herbivory. We demonstrated that interactions between tree seedlings and perennial grasses vary seasonally, and timing of tree germination may determine the importance of facilitation or competition in structuring savanna vegetation because of fluctuations in abiotic stress. Our finding that trees can grow and survive in a dense C4 grass sward contrasts with the common perception that grass competition limits woody plant recruitment in savannas.

New insights into vegetation patterns and processes

A basic question in vegetation science is – what ecological and evolutionary processes create the patterns that we observe (Fig. 1)? Considerable descriptive information about vegetation structure is available for many regions of the world. Major forces linked to particular vegetation patterns are also known: evolution (availability of regional species pools), dispersal (how organisms reach a location), abiotic (climatic, soil conditions) and biotic (competition, facilitation, parasitism, mutualism) factors. However, the question of how these mechanisms interact to shape vegetation patterns remains largely open. This question becomes increasingly relevant as novel ecosystems emerge, native species change their ranges and alien species arrive, establish and become dominant, having profound impacts on a range of ecosystem functions and ecosystem services. This is the context in which the 56th Annual Symposium of the International Association for Vegetation Science (IAVS), ‘Vegetation patterns and their underlying processes’, was held at the University of Tartu, Estonia, on 26–30 June 2013 (Fig. 2). The meeting attracted over 450 participants from 41 countries.

Multi-decadal increases in shrub abundance in non-riverine red gum (Eucalyptus camaldulensis) woodlands occur during a period of complex land-use history

The history of land-use was examined in Eucalyptus camaldulensis Dehnh. woodland in the Victoria Valley of the Grampians National Park, south-eastern Australia, to help interpret changes in vegetation there during the last 50 years. We used aerial photography and dendrochronological data to quantify the amount of, and the rate of change in, the woody-vegetation cover between 1948 and 1997, and historical data to document land-use changes during this time. Aerial photographs indicated that in 1948, 56% of study area had 50% woody-plant cover. The native shrub Leptospermum scoparium J.R. Forst & G. Forst (Myrtaceae) was predominantly responsible for the increases in cover. Demographic analyses indicated that recruitment has been ongoing rather than episodic; large numbers of shrubs, however, have recruited since 1994. We hypothesise that the vegetation changes observed are likely a response to changes in land-use that have occurred since European occupation. Increased woody-plant cover followed the removal of sheep grazing in the long-term absence of fire. It is very likely that the long history of stock grazing, coupled with selective logging and associated soil disturbance, initiated a change in understorey vegetation by reducing competitive native tussock grasses and fuel loads to carry fires and this reduction was initially responsible for the encroachment of shrubs into the woodland. Recruitment has been ongoing in the absence of any recent land-use changes (most utilisation ceasing after the declaration of National Park status in 1984) and hence, this transformation from species-rich herbaceous woodland to shrubby woodland is expected to continue in the future.

Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands—those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)—had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.Analysis of 65 grasslands worldwide from the Nutrient Network experiment reveals that plant communities with higher α- and β-diversity have higher levels of ecosystem multifunctionality, and that this effect is amplified across scales.

Periodic rest from grazing provided no control of an invasive perennial forb

This study examined if one grazing strategy (namely seasonal rest) was effective in the control of the invasive forbPhylacanescens(Kunth)Greene(hereafterlippia).Weexaminedifrestfromgrazingcouldincreasethecompetitiveness of native palatable species by allowing time to recover from defoliation, thereby altering competitive interactions between native species and lippia. In a field trial, we manipulated cattle grazing to determine its effects on the biomass of lippia and nativespecies.Wecomparedrestfromgrazingatdifferenttimesoftheyearwithyear-longgrazing(lowintensitycontinuous and high intensity short duration) and no grazing (permanently excluding large grazing animals). Experimental plots were stratified into different hydrological areas (approximately annual flooding and flooded less than once every 5 years) to include flood dynamics in the management scenarios. We detected no negative impacts of seasonal rest on lippia, but some positiveeffectsonnativespecies.Wefoundthatcompleteexclusionfromgrazinginareasthatalreadyhavesubstantiallippia invasion (and no flooding) may actually favour the expansion of lippia (at certain times). This study does not suggest that grazing management cannot be used as a tool for lippia control - simply that seasonal resting had no effect over a 3-year period.Thisislikelyduetothedynamicsofaboom-and-bustlandscapeinwhichiftheabioticconditionsarenotsuitablefor growth, then native species will not grow whether grazed or rested.