Loreto Rossi

Connecting the green and brown worlds : allometric and stoichiometric predictability of above- and below-ground networks

Abstract We examine the potential of trait-based parameters of taxa for linking above- and below-ground ecological networks (hereafter ‘green’ and ‘brown’ worlds) to understand and predict community dynamics. This synthesis considers carbon, nitrogen and phosphorus-related traits, the abundance of component species and their size distribution across trophic levels under different forms of management. We have analysed existing and novel databases on plants, microbes and invertebrates that combine physico-chemical and biological information from (agro)ecosystems spanning the globe. We found (1) evidence that traits from above- and below-ground systems may be integrated in the same model and (2) a much greater than expected stoichiometric plasticity of plants and microbes which has implications for the entire food-web mass–abundance scaling. Nitrogen and phosphorus are primary basal resources (hence, drivers) and more retranslocation of P than of N from leaves will lead to higher N:P in the litter and soil organic matter. Thus, under nutrient-rich conditions, higher foliar concentrations of N and P are reflected by lower N:P in the brown litter, suggesting less P retranslocated than N. This apparent stoichiometric dichotomy between green and brown could result in shifts in threshold elemental ratios critical for ecosystem functioning. It has important implications for a general food-web model, given that resource C:N:P ratios are generally assumed to reflect environmental C:N:P ratios. We also provide the first evidence for large-scale allometric changes according to the stoichiometry of agroecosystems. Finally, we discuss insights that can be gained from integrating carbon and nitrogen isotope data into trait-based approaches, and address the origin of changes in Δ 13 C and Δ 15 N fractionation values in relation to consumer–resource body-mass ratios.

Cascading effects of predatory fish exclusion on the detritus-based food web of a lake littoral zone (Lake Vico, central Italy)

Abstract. An exclosure experiment was carried out in the reed-dominated littoral zone of a volcanic lake (Lake Vico, central Italy) to test whether the impact of predatory fish on benthic invertebrates cascades on fungal colonisation and breakdown of leaf detritus. The abundance, biomass, and Shannon diversity index of the invertebrate assemblage colonising Phragmites australis leaf packs placed inside: (1) full-exclosure cages, (2) cages allowing access only to small-sized fish predators, and (3) cageless controls, were monitored over a 45-day period together with the mass loss and associated fungal biomass of leaf packs. The species composition of the fungal assemblage was further assessed at the end of the manipulation. In general, invertebrate predators did not show any significant response to fish exclusion, either on a trophic guild or on a single taxon level. In contrast, the exclusion of large predatory fish induced a diverse spectrum of changes in the abundance and population size-structure of dominant detritivore taxa, ultimately increasing the biomass and Shannon diversity index of the whole detritivorous guild. These changes corresponded with significant variations in leaf detritus decay rates as well as in the biomass and assemblage structure of associated fungal colonisers. Our experimental findings provide evidence that in Lake Vico effects of fish predators on invertebrate detritivores influence the fungal conditioning and breakdown of the detrital substrate. We conclude that in lacustrine littoral zones predator-driven constraints may structure lower trophic levels of detritus-based food webs and affect the decomposition of leaf detritus originated from the riparian vegetation.

Sediment structural properties mediating dominant feeding types patterns in soft-bottom macrobenthos of the Northern Adriatic Sea

In sediments sampled in Northern Adriatic Sea invertebrate benthic macrofauna was examined in comparison with both substratum bulk parameters (average grain size, sorting, total organic matter content) and attributes of grain size classes (specific organic matter content of dimensional fractions). The aims were: 1) to determine whether variations of community overall parameters (total spatial density, number of taxa) and in numerically dominant feeding types (deposit-feeders and suspension-feeders) patterns were related to changes in sediment general properties; 2) to evaluate whether the observed patterns of abundance for deposit- and filter-feeders had more valid correlates in specific properties of substratum dimensional fractions.Number of taxa and animal spatial density resulted conventionally related to substratum bulk parameters while feeding types patterns in comparison with sediment total organic matter content exibited a reciprocal negative effect; only when both feeding types and sediment structure were resolved to their specific attributes in terms of prevalence of tubicolous taxa among deposit-feeders and organic matter richness of single dimensional fractions, the analysis emphasized patterns unequally affected by specific size fractions abundance and organic matter content: deposit feeders demonstrated to be related to the presence in the substrate of intermediate grain size classes necessary for tube-building. Those intermediate fractions proved to be more adequate descriptors than average grain size or other overall sediment parameters per se of relations actually established between feeding types and the complex nature of bottom sediments.Our results emphasize that in the Northern Adriatic Sea, sediment organic matter content may represent a factor of minor importance in comparison with other substrate attributes, for which it may be necessary a detailed analysis of sediment structure. Thus, we concluded that the complexity of soft-bottom communities may defy any simple paradigm relating macrobenthic patterns to any single sediment bulk attribute, and we propose a shift in focus towards an higher resolution of both functional groups in macrobenthic associations (as already suggested in other investigations) and of substratum structural description.

Interactions between detritivores and microfungi during the leaf detritus decomposition in a volcanic lake (Lake Vico – central Italy)

The role of biota in the mass loss of Phragmites australis (Cav.) Trin. ex Steud was studied in the littoral belt of a central Italy volcanic lake. The research focussed on the feeding interactions between detritivores and decomposing fungi as drivers of the leaf litter decomposition. The litterbag technique was used to assess the leaf mass loss, the number of colonizing fungi and the patterns of leaf colonization by detritivores during 40 days of submersion in 16 sampling sites. Cores of bottom sediment were collected to estimate the organic content and ergosterol concentration as measure of fungal mass. The rate of leaf mass loss showed significant variability among the sampling sites and was non-linearly related to the quantity of organic depositions onto the lake bottom, peaking at about 40% of the dry matter. The rate was also positively correlated with the density of detritivore mass relative to the leaf unit mass, which increased with time. On the 20th day of litterbag immersion, when 40% of the initial leaf litter remained, we observed the best accordance between the two measures as well as the lowest difference in the detritivore mass density among sampling sites. In the absence of animals, the decomposition rate was positively related to the number of fungi on the decaying litter. The feeding activity of detritivores changed both the species richness and composition of the fungal community on the litter. The substrate reduction due to intense animal feeding appeared to limit the ability of fungi to regrow after grazing. As a result, an inverse relationship between the number of fungi and the decomposition rate was observed.

δ15N variation in Ulva lactuca as a proxy for anthropogenic nitrogen inputs in coastal areas of Gulf of Gaeta (Mediterranean Sea)

We tested the capacity of Ulva lactuca to mark N sources across large marine areas by measuring variation in its δ(15)N at several sites in the Gulf of Gaeta. Comparisons were made with the macroalga Cystoseira amentacea. Variation of δ(15)N values was assessed also in the coastal waters off the Circeo Natural Park, where U. lactuca and C. amentacea were harvested, as these waters are barely influenced by human activities and were used as reference site. A small fragment from each frond was preserved before deployment in order to characterize the initial isotopic values. After 48 h of submersion, U. lactuca was more responsive than C. amentacea to environmental variation and δ(15)N enrichment in the Gulf of Gaeta was observed. The spatial distribution of δ(15)N enrichment indicated that different macro-areas in the Gulf were affected by N inputs from different origins. Comparison of the δ(15)N values of fragments taken from the same transplanted frond avoided bias arising from natural isotopic variability.

Competition between two aquatic detritivorous isopods – a laboratory study

The role of adult faeces in juvenile nutrition of two isopod species, Proasellus coxalis s.l. and Asellus aquaticus (L.), with similar trophic strategies and different reproductive output, has been studied in laboratory. Our aim was to consider the possible competitive mechanisms occurring at the beginning of the species coexistence using allopatric populations in single and mixed species experiments. Two series of competition experiments were performed. In the first, adult specimens were used for breeding and feeding trials. Both population dynamics and the percentage of ovigerous females and juveniles were evaluated during 10 months. Adult densities and juvenile percentage of A.aquaticus were lower in the presence of P. coxalis s.l. than when alone. At the end of the breeding experiments the dietary preferences of adults on a set of fungally conditioned leaf discs were not different among treatments. In the second series of experiments, the influence of coexistence on the feeding rates of young asellids and the relative importance of faeces and decaying plant material in their diet were investigated. Individual consumption by wild juveniles in multiple-choice laboratory experiments was measured by radioisotopes (322P). Juveniles of P. coxalis s.l. showed the highest ingestion rates. In co-occurrence, contrary to A.aquaticus, they were able to further increase feeding on parental faeces. The role of parental faeces in the diet of the two species juveniles and the competitive dominance of P. coxalis s.l. are discussed.

Predators and resources influence phosphorus transfer along an invertebrate food web through changes in prey behaviour.

Predators play a fundamental role in prey trophic behaviour, with indirect consequences for species coexistence and ecosystem functioning. Resource quality and availability also influence prey trophic behaviour, with potential effects on predator-prey dynamics. Although many studies have addressed these topics, little attention has been paid to the combined effects of predators and resources on prey species coexistence and nutrient transfer along food chains, especially in detritus-based systems. To determine the influence of predators and resource quality on the movement and P uptake of detritivores, we carried out a field experiment on the River Kelvin (Scotland) using 32P to test the hypothesis of reduced prey vagility among resource patches as a strategy to avoid predation. Thirty leaf sacks containing alder leaves and two detritivore prey populations (Asellus aquaticus and Lymnaea peregra) were placed in cages, half of them with two predator species (Dendrocoelum lacteum and Erpobdella octoculata) and the other half without predators. Five alder leaf bags, each individually inoculated with a different fungus strain to simulate a patchy habitat, were placed inside each leaf sack. One bag in each sack was labelled with 32P, in order to assess the proportion of detritivores using it as food and thus their movement among the five resource patches. Three replicates for each labelled fungus and each predation treatment (i.e. with and without predators) were left on the riverbed for 7 days. The presence of predators had negligible effects on the number of detritivores in the leaf bags, but it did reduce the proportion of 32P-labelled detritivores and their P uptake. The most strongly affected species was A. aquaticus, whose vagility, trophic overlap with L. peregra and P uptake were all reduced. The results confirm the importance of bottom-up and top-down forces acting simultaneously to regulate nutrient transfer along food chains in patchy habitats.

Effect of multiple disturbances on food web vulnerability to biodiversity loss in detritus-based systems

Global biodiversity is affected by human pressure and climate change, and the present rate of biodiversity loss is probably higher than ever before. Community composition is also changing, and interspecific interactions are under severe pressure. The extinction of one species within a food web can result in further secondary extinctions, due to bottom-up effects that can be even more intense and less predictable than the direct effects of disturbance, undermining our capacity for ecosystem management and conservation. Here we investigated a metric for assessing the structural stability of food webs in the face of species loss, referred to as “Resistance”, based on two fundamental web properties: (1) the proportion of key species in the web, a “key” species being one whose deletion leads to at least one secondary extinction, and (2) the mean number of secondary extinctions observed per key species deletion. We compared web Resistance with web Robustness (Dunne et al. 2002) based on 12 detritus-based riverine food webs under four species extinction scenarios on various temporal and spatial scales. We investigated the effect of multiple disturbances (extreme flood and river basin urbanization) on community vulnerability to biodiversity loss, assessing the behavior of Robustness and Resistance under the applied species extinction scenarios and testing their dependence on web topology. We estimated the contribution of the rarest and the most dominant species, and that of the most and least connected species, to web Resistance. Urbanization negatively affected community vulnerability to biodiversity loss. Only food web Resistance showed a significant flood effect and interaction between flood and urbanization. The most connected species contributed the most to food web resistance, whereas the rarest and the most abundant species had a similar, intermediate structural importance. Both food web Resistance and the role of selected key species varied across web description scales. Food web Resistance values were coherent across species extinction scenarios, demonstrating the suitability of the proposed approach for quantifying community vulnerability to species loss and the importance of considering food webs in monitoring and impact assessment programs. The approach is thus seen to be a promising research pathway supporting ecosystem management.

Seasonal abundance and trophic position of the Atlantic blue crab Callinectes sapidus Rathbun 1896 in a Mediterranean coastal habitat

The blue crab Callinectes sapidus, native to the western coasts of the Atlantic Ocean, has been introduced in Mediterranean waters where it is currently considered an invasive species. Here, we verified the occurrence of an established population of C. sapidus in the Torre Colimena basin, a Mediterranean habitat located in SE Italy, and provided a first assessment of its functional role in a Mediterranean coastal ecosystem. Crab traps were used to estimate the species abundance at a seasonal frequency; in addition, its trophic position was estimated in summer using nitrogen stable isotopes. Estimations were performed using both a generally adopted value of 3.4 ‰ for the nitrogen trophic level fractionation factor ∆15N, and species-specific estimations obtained from the literature. C. sapidus was sampled throughout the year, reaching peak abundances in summer. This result, taken together with the observed patterns of variation in the average body size and sex ratio of captured specimens, provided evidence of a fully established population in the Torre Colimena basin. The estimated trophic position of the crab was consistent with the literature information on western Atlantic populations, and resulted, irrespective of the fractionation factor used, significantly higher than those characterizing an autochthonous brachyuran (Pachygrapsus marmoratus) and a fish predator (Sparus aurata). The present study provided novel information on the occurrence and potential functional impact of this non-indigenous species in Mediterranean coastal habitats, highlighting current knowledge lacunae and identifying future research lines on marine alien species.

A COADAPTED TROPHIC NICHE IN TWO SPECIES OF CRUSTACEA (ISOPODA): ASELLUS AQUATICUS (L.) AND PROASELLUS COXALIS DOLFF

In present-day ecologic literature, the hypothesis that competition is an important factor in coevolutionary processes of antagonist species is widely shared (MacArthur, 1970; Colwell and Fuentes, 1975; Roughgarden, 1979; Connell, 1980). In spite of the many theoretical papers devoted to this subject, it is still to be assessed how and to what extent the notion can be applied to real communities and what circumstances actually promote the coevolutionary partitioning of resources among coexisting and ecologically similar species. It is uncommon to find direct evidence of coevolution in present species, but we can observe allopatric and sympatric populations and assume that the former condition precedes the latter in time. If the studied areas are basically similar it is possible that differences between conspecific populations are a response to competition rather than adaptation to the environment. This is perhaps the best of non-experimental methods, especially in benthic communities of lotic environments: in practice, it is extremely difficult in natural waterways to control experiments involving manipulation of the number of species in order to obtain direct evidence of coevolution (Connell, 1980). On the other hand, very little is known about coevolutionary trophic partitioning between benthic detritivorous animals. The importance of microfungi present on plant detritus in the diet of this class of animals has been ascertained: detritivorous animals select detritus according to the type of microfungal colonization (Birlocher and Kendrick, 1973; Cummins and KlIug, 1979; Rossi et al., 1981). Moreover, in populations of two species of isopods co-occurring in a biotope in southern Italy, we showed a strong trophic dependence of the animals on the fungi and noted a high interspecific differentiation in the capacity to exploit the various fungus strains offered as food (Rossi and Fano, 1979). Our data did not permit speculation as to the evolutionary nature of the phenomenon, so in the present research attention was given to the trophic niches of allopatric and sympatric populations of the isopods Asellus aquaticus and Proasellus coxalis. Our investigations particularly concerned the potential of trophic exploitation of many microfungus strains present on the detritus of waterways. We intended to find an answer to two questions. 1) Do populations of two species of isopods co-occurring in the same biotope and two allopatrically occurring populations result in a different potential trophic separation? 2) Do levels of trophic separation of the two species depend on the frequencies of the microfungal resources in nature?