Ross M. Thompson

META‐ANALYSIS: TROPHIC LEVEL, HABITAT, AND PRODUCTIVITY SHAPE THE FOOD WEB EFFECTS OF RESOURCE SUBSIDIES

Studies of the effects of cross-habitat resource subsidies have been a feature of food web ecology over the past decade. To date, most studies have focused on demonstrating the magnitude of a subsidy or documenting its effect in the recipient habitat. Ecologists have yet to develop a satisfactory framework for predicting the magnitude of these effects. We used 115 data sets from 32 studies to compare consumer responses to resource subsidies across recipient habitat type, trophic level, and functional group. Changes in consumer density or biomass in response to subsidies were inconsistent across habitats, trophic, and functional groups. Responses in stream cobble bar and coastline habitats were larger than in other habitats. Contrary to expectation, the magnitude of consumer response was not affected by recipient habitat productivity or the ratio of productivity between donor and recipient habitats. However, consumer response was significantly related to the ratio of subsidy resources to equivalent resources in the recipient habitat. Broad contrasts in productivity are modified by subsidy type, vector, and the physical and biotic characteristics of both donor and recipient habitats. For this reason, the ratio of subsidy to equivalent resources is a more useful tool for predicting the possible effect of a subsidy than coarser contrasts of in situ productivity. The commonness of subsidy effects suggests that many ecosystems need to be studied as open systems.

Food webs: Reconciling the structure and function of biodiversity

The global biodiversity crisis concerns not only unprecedented loss of species within communities, but also related consequences for ecosystem function. Community ecology focuses on patterns of species richness and community composition, whereas ecosystem ecology focuses on fluxes of energy and materials. Food webs provide a quantitative framework to combine these approaches and unify the study of biodiversity and ecosystem function. We summarise the progression of food-web ecology and the challenges in using the food-web approach. We identify five areas of research where these advances can continue, and be applied to global challenges. Finally, we describe what data are needed in the next generation of food-web studies to reconcile the structure and function of biodiversity.

TROPHIC LEVELS AND TROPHIC TANGLES: THE PREVALENCE OF OMNIVORY IN REAL FOOD WEBS

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.

Mesocosm Experiments as a Tool for Ecological Climate-Change Research

Abstract Predicting the ecological causes and consequences of global climate change requires a variety of approaches, including the use of experiments, models, and surveys. Among experiments, mesocosms have become increasingly popular because they provide an important bridge between smaller, more tightly controlled, microcosm experiments (which can suffer from limited realism) and the greater biological complexity of natural systems (in which mechanistic relationships often cannot be identified). A new evaluation of the contribution of the mesocosm approach, its potential for future research, as well as its limitations, is timely. As part of this review, we constructed a new database of over 250 post-1990 studies that have explored different components of climate change across a range of organisational levels, scales, and habitats. Issues related to realism, reproducibility and control are assessed in marine, freshwater, and terrestrial systems. Some general patterns emerged, particularly at the ecosystem level, such as consistent and predictable effects on whole-system respiration rates. There are, however, also many seemingly idiosyncratic, contingent responses, especially at the community level, both within and among habitat types. These similarities and differences in both the drivers and responses highlight the need for caution before making generalisations. Finally, we assess future directions and prospects for new methodological advances and the need for greater international coordination and interdisciplinarity.

Stepping in Elton’s footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.

Evolutionary refugia and ecological refuges: key concepts for conserving Australian arid zone freshwater biodiversity under climate change.

Refugia have been suggested as priority sites for conservation under climate change because of their ability to facilitate survival of biota under adverse conditions. Here, we review the likely role of refugial habitats in conserving freshwater biota in arid Australian aquatic systems where the major long-term climatic influence has been aridification. We introduce a conceptual model that characterizes evolutionary refugia and ecological refuges based on our review of the attributes of aquatic habitats and freshwater taxa (fishes and aquatic invertebrates) in arid Australia. We also identify methods of recognizing likely future refugia and approaches to assessing the vulnerability of arid-adapted freshwater biota to a warming and drying climate. Evolutionary refugia in arid areas are characterized as permanent, groundwater-dependent habitats (subterranean aquifers and springs) supporting vicariant relicts and short-range endemics. Ecological refuges can vary across space and time, depending on the dispersal abilities of aquatic taxa and the geographical proximity and hydrological connectivity of aquatic habitats. The most important are the perennial waterbodies (both groundwater and surface water fed) that support obligate aquatic organisms. These species will persist where suitable habitats are available and dispersal pathways are maintained. For very mobile species (invertebrates with an aerial dispersal phase) evolutionary refugia may also act as ecological refuges. Evolutionary refugia are likely future refugia because their water source (groundwater) is decoupled from local precipitation. However, their biota is extremely vulnerable to changes in local conditions because population extinction risks cannot be abated by the dispersal of individuals from other sites. Conservation planning must incorporate a high level of protection for aquifers that support refugial sites. Ecological refuges are vulnerable to changes in regional climate because they have little thermal or hydrological buffering. Accordingly, conservation planning must focus on maintaining meta-population processes, especially through dynamic connectivity between aquatic habitats at a landscape scale.

A New Dolphin Species, the Burrunan Dolphin Tursiops australis sp. nov., Endemic to Southern Australian Coastal Waters

Small coastal dolphins endemic to south-eastern Australia have variously been assigned to described species Tursiops truncatus, T. aduncus or T. maugeanus; however the specific affinities of these animals is controversial and have recently been questioned. Historically ‘the southern Australian Tursiops’ was identified as unique and was formally named Tursiops maugeanus but was later synonymised with T. truncatus. Morphologically, these coastal dolphins share some characters with both aforementioned recognised Tursiops species, but they also possess unique characters not found in either. Recent mtDNA and microsatellite genetic evidence indicates deep evolutionary divergence between this dolphin and the two currently recognised Tursiops species. However, in accordance with the recommendations of the Workshop on Cetacean Systematics, and the Unified Species Concept the use of molecular evidence alone is inadequate for describing new species. Here we describe the macro-morphological, colouration and cranial characters of these animals, assess the available and new genetic data, and conclude that multiple lines of evidence clearly indicate a new species of dolphin. We demonstrate that the syntype material of T. maugeanus comprises two different species, one of which is the historical ‘southern form of Tursiops’ most similar to T. truncatus, and the other is representative of the new species and requires formal classification. These dolphins are here described as Tursiops australis sp. nov., with the common name of ‘Burrunan Dolphin’ following Australian aboriginal narrative. The recognition of T. australis sp. nov. is particularly significant given the endemism of this new species to a small geographic region of southern and south-eastern Australia, where only two small resident populations in close proximity to a major urban and agricultural centre are known, giving them a high conservation value and making them susceptible to numerous anthropogenic threats.

What does biodiversity actually do? A review for managers and policy makers

Conservation managers and policy makers must often justify the need for protection of biodiversity. However, results of scientific studies testing for a positive value of biodiversity in terms of community stability and ecosystem function have been complex and inconsistent. We review recent information on the consequences of loss of biodiversity for natural systems. The relationships described vary with scale of interest – for instance, biodiversity at local scales typically has strong effects on ecosystem function, although the opposite relationship is often found at regional scales. These inconsistencies lead to some concern as to whether these relationships can be used to justify biodiversity protection. This is particularly relevant to policy, where holistic protection of biodiversity has most often been mooted and justified. For managers, who most often work to protect single species, communities or ecosystem functions, biodiversity research has failed to address questions of critical concern such as consequences of the loss of rare species or the identification of functional keystone species. For the general public, we believe that the confusion and debate surrounding biodiversity and ecosystem function relationships is in danger of eroding the positive value society places on biodiversity. We further warn that using those relationships in policy documents as justifications for biodiversity protection is fraught with difficulties. Finally, we contend that biodiversity research has largely not addressed issues of concern to conservation managers, and list a set of priorities for relevant research on the consequences of biodiversity loss.

The Effect of Seasonal Variation on the Community Structure and Food-Web Attributes of Two Streams: Implications for Food-Web Science

Benthic communities from two grassland streams in New Zealand were contrasted to quantify seasonal variation in food-web and community attributes. Connectivity food webs were used to describe trophic links and energy flows Community attributes webs were used to describe trophic links and energy flows. Community attributes ere used to describe the distribution of species and biomass across taxa and functional feeding groups. The food webs produced are amongst the largest published reflecting detailed taxonomy across fish, algae and macro-invertebrates. All food-web attributes - species richness, number of links, connectance, mean chain length. average number of links down and prey:predator ratio - showed significant variation across seasons. Community structure showed less seasonality, and low variation Variation in food supply was important in structuring the community, e.g. primary production was correlated with the vertical dimensions (mean chain length) of the food webs and seemed to determine the proportion of species and biomass that were composed of browsers. Organic matter availability (as seston or standing crop) seemed to be important in dictating the proportion of species and biomass that were composed of collector-gatherers and filter feeders. Cluster analysis reveals that the two streams differed in fundamental ways. Dempsters Stream was characterised by a structurally complex, species-rich community, with a high proportion of predators and a predominance of browsers. This appears to be based on high algal productivity. Sutton Stream was characterised by a simpler food web with a greater contribution of collector-gatherers and filter feeders, reflecting higher availability and standing crop of organic matter. When clustered by food-web attributes, species richness was important in dictating which communities clustered together, identifying the species-rich spring and particularly summer food webs of Dempsters Stream as distinct from the other communities. On the other hand, when clustered according to community attributes, the spring and summer Dempsters communities grouped with the autumn and summer Sutton communities, whereas the autumn Dempsters community was most distinctive, due to a high species richness of Plecoptera and a high predator biomass. A comparison of seasonal and whole year summary food webs showed that web size. the number of links, and the average number of links down were over-estimated in the summary webs. Other attributes of the summary food webs, such as connectance. mean chain lengths and prey:predator ratios approximated the average of those from the seasonal food webs, the summary webs clustered separately from all the other food webs due to their higher species richness. Thus a food web gathered in a single season does not accurately reflect the food-web structure during any other season, or of the whole year.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.