The assessment of a global marine ecosystem model on the basis of emergent properties and ecosystem function: a case study with ERSEM

Ecosystem models are often assessed using quantitative metrics of absolute ecosystem state, but these model-data comparisons are disproportionately vulnerable to discrepancies in the location of important circulation features. An alternative method is to demonstrate the models capacity to represent ecosystem function; the emergence of a coherent natural relationship in a simulation indicates that the model may have an appropriate representation of the ecosystem functions that lead to the emergent relationship. Furthermore, as emergent properties are large-scale properties of the system, model validation with emergent properties is possible even when there is very little or no appropriate data for the region under study, or when the hydrodynamic component of the model differs significantly from that observed in nature at the same location and time. A selection of published meta-analyses are used to establish the validity of a complex marine ecosystem model and to demonstrate the power of validation with emergent properties. These relationships include the phytoplankton community structure, the ratio of carbon to chlorophyll in phytoplankton and particulate organic matter, the ratio of particulate organic carbon to particulate organic nitrogen and the stoichiometric balance of the ecosystem. These metrics can also inform aspects of the marine ecosystem model not available from traditional quantitative and qualitative methods. For instance, these emergent properties can be used to validate the design decisions of the model, such as the range of phytoplankton functional types and their behaviour, the stoichiometric flexibility with regards to each nutrient, and the choice of fixed or variable carbon to nitrogen ratios.

An unusually large phytoplankton spring bloom drives rapid changes in benthic diversity and ecosystem function

In 2012, the Western English Channel experienced an unusually large and long-lived phytoplankton spring bloom. When compared with data from the past 20 years, average phytoplankton biomass at Station L4 (part of the Western Channel Observatory) was approximately 3× greater and lasted 50% longer than any previous year. Regular (mostly weekly) box core samples were collected from this site before, during and after the bloom to determine its impact on macrofaunal abundance, diversity, biomass, community structure and function. The spring bloom of 2012 was shown to support a large and rapid response in the majority of benthic taxa and functional groups. However, key differences in the precise nature of this response, as well as in its timing, was observed between different macrofauna feeding groups. Deposit feeders responded almost instantly at the start of the bloom, primarily thorough an increase in abundance. Suspension feeders and opportunistic/predatory/carnivorous taxa responded slightly more slowly and primarily with an increase in biomass. At the end of the bloom a rapid decline in macrobenthic abundance, diversity and biomass closely followed the decline in phytoplankton biomass. With suspension feeders showing evidence of this decline a few weeks before deposit feeders, it was concluded that this collapse in benthic communities was driven primarily by food availability and competition. However, it is possible that environmental hypoxia and the presence of toxic benthic cyanobacteria could also have contributed to this decline. This study shows evidence for strong benthic–pelagic coupling at L4; a shallow (50 m), coastal, fine-sand habitat. It also demonstrates that in such habitats, it is not just planktonic organisms that demonstrate clear community phenology. Different functional groups within the benthic assemblage will respond to the spring bloom in specific manner, with implications for key ecosystem functions and processes, such as secondary production and bioturbation. Only by taking integrated benthic and pelagic observations over such fine temporal scales (weekly) was the current study able to identify the intimate structure of the benthic response. Similar studies from other habitats and under different bloom conditions are urgently needed to fully appreciate the strength of benthic–pelagic coupling in shallow coastal environments.

Detecting the effects of diversity on measures of ecosystem function: experimental design, null models and empirical observations

Here we present a novel experimental approach to examine the relationship between diversity and ecosystem Function. We develop four null predictive models, with which to differentiate between the 'sampling effect' - the chance inclusion of a highly productive species, and 'species complementarity' - the complementary use of resources by species that differ in their niche or resource use. We investigate the effects of manipulating species and functional richness on ecosystem function in marine benthic system and using empirical data from our own experiments we illustrate the application of these models.

Effects of Phytophthora cinnamomi on heathland fauna and ecosystem function

Progression of disease caused by the plant pathogen Phytophthora cinnamomi was correlated to rainfall events and resulted in a loss of plant species diversity in heathland vegetation at Anglesea, Victoria. Lower captures of small mammals were recorded in diseased areas. Management of disease using the chemical phosphite was also evaluated.

Friend or foe : exotic flora and ecosystem function

Exotic flora, particularly weeds, are renowned for out-competing and displacing native flora, consequently affecting native fauna and pollinator relationships. Nonetheless, it stands to reason that weeds must provide some compensatory ecological value. This study assessed whether weeds are friend or foe to ecosystem function by considering the quality and quantity of pollen offered by widespread weeds in Australian ecosystems. Using the Honeybee Apis mellifera as a case study, and information derived from highly experienced commercial apiarists, we determined that 32 exotic plants are important pollen sources. Most species offered high to very high quality pollen. Pollen quality varied temporally, spatially and infraspecifically. Fifteen species were considered more beneficial to A. mellifera than others; only seven species were considered less beneficial. Thus, exotic flora contribute pollen resources that are valuable to maintain ecosystem function, particularly at times when flowering native species are few.

Functional replacement across species pools of vertebrate scavengers separated at a continental scale maintains an ecosystem function

Summary: The composition of species pools can vary in space and time. While many studies are focused on understanding which factors influence the make-up of species pools, the question to which degree biogeographic variation in species composition propagates to biogeographic variation in ecological function is rarely examined. If different local species assemblages operate in ways that maintain specific ecological processes across continents, they can be regarded as functionally equivalent. Alternatively, variation in species assemblages might result in the loss of ecological function if different species fulfil different functions, and thereby fail to maintain the ecological process. Here, we test whether ecological function is affected by differences in the composition of species pools across a continental scale, comparing a tropical with a temperate pool. The model systems are assemblages of vertebrates foraging on ocean beaches, and the ecological function of interest is the consumption of wave-cast carrion, a pivotal process in sandy shore ecosystems. We placed fish carcasses (n = 179) at the beach-dune interface, monitored by motion-triggered cameras to record scavengers and quantify the detection and removal of carrion. Scavenging function was measured on sandy beaches in two distinct biogeographic regions of Australia: tropical north Queensland and temperate Victoria. The composition of scavenging assemblages on sandy beaches varied significantly across the study domain. Raptors dominated in the tropics, while invasive red foxes were prominent in temperate assemblages. Notwithstanding the significant biogeographic change in species composition, ecological function - as indexed by carcass detection and removal - was maintained, suggesting strong functional replacement at the continental scale. Species pools of vertebrate scavengers that are assembled from taxonomically distinct groups (birds vs. mammals) and located in distinct climatic regions (temperate vs. tropical) can maintain an ecological process via replacement of species with comparable functional traits.

Strongly interactive carnivore species: maintaining and restoring ecosystem function

By emphasizing Australian carnivores as exemplars of flesh-eaters in other parts of the world, this book will be an important reference for researchers, wildlife managers and students worldwide.

Are shrubs really a sign of declining ecosystem function? Disentangling the myths and truths of woody encroachment in Australia

Since European settlement, there has been a dramatic increase in the density, cover and distribution of woody plants in former grassland and open woodland. There is a widespread belief that shrub encroachment is synonymous with declines in ecosystem functions, and often it is associated with landscape degradation or desertification. Indeed, this decline in ecosystem functioning is considered to be driven largely by the presence of the shrubs themselves. This prevailing paradigm has been the basis for an extensive program of shrub removal, based on the view that it is necessary to reinstate the original open woodland or grassland structure from which shrublands are thought to have been derived. We review existing scientific evidence, particularly focussed on eastern Australia, to question the notion that shrub encroachment leads to declines in ecosystem functions. We then summarise this scientific evidence into two conceptual models aimed at optimising landscape management to maximise the services provided by shrub-encroached areas. The first model seeks to reconcile the apparent conflicts between the patch- and landscape-level effects of shrubs. The second model identifies the ecosystem services derived from different stages of shrub encroachment. We also examined six ecosystem services provided by shrublands (biodiversity, soil C, hydrology, nutrient provision, grass growth and soil fertility) by using published and unpublished data. We demonstrated the following: (1) shrub effects on ecosystems are strongly scale-, species- and environment-dependent and, therefore, no standardised management should be applied to every case; (2) overgrazing dampens the generally positive effect of shrubs, leading to the misleading relationship between encroachment and degradation; (3) woody encroachment per se does not hinder any of the functions or services described above, rather it enhances many of them; (4) no single shrub-encroachment state (including grasslands without shrubs) will maximise all services; rather, the provision of ecosystem goods and services by shrublands requires a mixture of different states; and (5) there has been little rigorous assessment of the long-term effectiveness of removal and no evidence that this improves land condition in most cases. Our review provides the basis for an improved, scientifically based understanding and management of shrublands, so as to balance the competing goals of providing functional habitats, maintaining soil processes and sustaining pastoral livelihoods.

Shifts in microbial communities do not explain the response of grassland ecosystem function to plant functional composition and rainfall change

Ecosystem functioning in grasslands is regulated by a range of biotic and abiotic factors, and the role of microbial communities in regulating ecosystem function has been the subject of much recent scrutiny. However, there are still knowledge gaps regarding the impacts of rainfall and vegetation change upon microbial communities and the implications of these changes for ecosystem functioning. We investigated this issue using data from an experimental mesotrophic grassland study in south-east England, which had been subjected to four years of rainfall and plant functional composition manipulations. Soil respiration, nitrogen and phosphorus stocks were measured, and the abundance and community structure of soil microbes were characterised using quantitative PCR and multiplex-TRFLP analysis, respectively. Bacterial community structure was strongly related to the plant functional composition treatments, but not the rainfall treatment. However, there was a strong effect of both rainfall change and plant functional group upon bacterial abundance. There was also a weak interactive effect of the two treatments upon fungal community structure, although fungal abundance was not affected by either treatment. Next, we used a statistical approach to assess whether treatment effects on ecosystem function were regulated by the microbial community. Our results revealed that ecosystem function was influenced by the experimental treatments, but was not related to associated changes to the microbial community. Overall, these results indicate that changes in fungal and bacterial community structure and abundance play a relatively minor role in determining grassland ecosystem function responses to precipitation and plant functional composition change, and that direct effects on soil physical and chemical properties and upon plant and microbial physiology may play a more important role.

Biodiversity and Ecosystem Function in the Gulf of Maine: Pattern and Role of Zooplankton and Pelagic Nekton

This paper forms part of a broader overview of biodiversity of marine life in the Gulf of Maine area (GoMA), facilitated by the GoMA Census of Marine Life program. It synthesizes current data on species diversity of zooplankton and pelagic nekton, including compilation of observed species and descriptions of seasonal, regional and cross-shelf diversity patterns. Zooplankton diversity in the GoMA is characterized by spatial differences in community composition among the neritic environment, the coastal shelf, and deep offshore waters. Copepod diversity increased with depth on the Scotian Shelf. On the coastal shelf of the western Gulf of Maine, the number of higher-level taxonomic groups declined with distance from shore, reflecting more nearshore meroplankton. Copepod diversity increased in late summer, and interdecadal diversity shifts were observed, including a period of higher diversity in the 1990s. Changes in species diversity were greatest on interannual scales, intermediate on seasonal scales, and smallest across regions, in contrast to abundance patterns, suggesting that zooplankton diversity may be a more sensitive indicator of ecosystem response to interannual climate variation than zooplankton abundance. Local factors such as bathymetry, proximity of the coast, and advection probably drive zooplankton and pelagic nekton diversity patterns in the GoMA, while ocean-basin-scale diversity patterns probably contribute to the increase in diversity at the Scotian Shelf break, a zone of mixing between the cold-temperate community of the shelf and the warm-water community offshore. Pressing research needs include establishment of a comprehensive system for observing change in zooplankton and pelagic nekton diversity, enhanced observations of "underknown'' but important functional components of the ecosystem, population and metapopulation studies, and development of analytical modeling tools to enhance understanding of diversity patterns and drivers. Ultimately, sustained observations and modeling analysis of biodiversity must be effectively communicated to managers and incorporated into ecosystem approaches for management of GoMA living marine resources.

Impact of elevated levels of CO2 on animal mediated ecosystem function: The modification of sediment nutrient fluxes by burrowing urchins

A mesocosm experiment was conducted to quantify the relationships between the presence and body size of two burrowing heart urchins (Brissopsis lyrifera and Echinocardium cordatum) and rates of sediment nutrient flux. Furthermore, the impact of seawater acidification on these relationships was determined during this 40-day exposure experiment. Using carbon dioxide (CO2) gas, seawater was acidified to pHNBS 7.6, 7.2 or 6.8. Control treatments were maintained in natural seawater (pH = 8.0). Under normocapnic conditions, burrowing urchins were seen to reduce the sediment uptake of nitrite or nitrate whilst enhancing the release of silicate and phosphate. In acidified (hypercapnic) treatments, the biological control of biogeochemical cycles by urchins was significantly affected, probably through the combined impacts of high CO2 on nitrifying bacteria, benthic algae and urchin behaviour. This study highlights the importance of considering biological interactions when predicting the consequences of seawater acidification on ecosystem function.

Microbes as engines of ecosystem function : When does community structure enhance predictions of ecosystem processes?

Peer reviewed

Tree species diversity and ecosystem function: Can tropical multi-species plantations generate greater productivity?

Results from the humid tropics of Australia demonstrate that diverse plantations can achieve greater productivity than monocultures. We found that increases in both the observed species number and the effective species richness were significantly related to increased levels of productivity as measured by stand basal area or mean individual tree basal area. Four of five plantation species were more productive in mixtures with other species than in monocultures, offering on average, a 55% increase in mean tree basal area. A general linear model suggests that species richness had a significant effect on mean individual tree basal area when environmental variables were included in the model. As monoculture plantations are currently the preferred reforestation method throughout the tropics these results suggest that significant productivity and ecological gains could be made if multi-species plantations are more broadly pursued. (c) 2006 Elsevier B.V. All rights reserved.