Similar breakdown rates and benthic macroinvertebrate assemblages on native and Eucalyptus globulus leaf litter in Californian streams

1. Eucalyptus globulus, a tree species planted worldwide in many riparian zones, has been reported to affect benthic macroinvertebrates negatively. Although there is no consensus about the effects of Eucalyptus on aquatic macrobenthos, its removal is sometimes proposed as a means of ecological restoration. 2. We combined the sampling of macroinvertebrates with measurement of the colonisation of leaf packs in mesh bags, to examine the effects of riparian Eucalyptus and its litter on benthic macroinvertebrates in three small streams in California, U.S.A. Each stream included one reach bordered by Eucalyptus (E‐site) and a second bordered by native vegetation (N‐site). 3. The macrobenthos was sampled and two sets of litter bags were deployed at each site: one set with Eucalyptus litter (Euc‐bags) and one with mixed native tree litter (Nat‐bags) containing Quercus, Umbellularia, Acer and Alnus. Bags were exposed for 28, 56 and 90 days and this experiment was repeated in the autumn, winter and spring to account for effects of changing stream flow and insect phenology. 4. Litter input (average dry mass: 950 g m^?2 year^?1 in E‐sites versus 669 g m^?2 year^?1 in N‐sites) was similar, although in‐stream litter composition differed between E‐ and N‐sites. Litter broke down at similar rates in Euc‐bags and Nat‐bags (0.0193 day^?1 versus 0.0134 day^?1), perhaps reflecting the refractory nature of some of the leaves of the native trees (Quercus agrifolia). 5. Summary metrics for macroinvertebrates (taxon richness, Shannon diversity, pollution tolerance index) did not differ significantly between the E and N sites, or between Euc‐bags and Nat‐bags. No effect of exposure time or site was detected by ordination of the taxa sampled. However, distinct seasonal ordination clusters were observed in winter, spring and autumn, and one of the three streams formed a separate cluster. 6. The presence of Eucalyptus was less important in explaining the taxonomic composition of the macrobenthos than either ‘season’ or ‘stream’. Similarly, these same two factors (but not litter species) also helped explain the variation in leaf breakdown. We conclude that patches of riparian Eucalyptus and its litter have little effect on stream macrobenthos in this region.     

Leaf diversity influences in-stream litter decomposition through effects on shredders

1. The functioning of many aquatic ecosystems is controlled by surrounding terrestrial ecosystems. In a view of growing interest in linking biodiversity to ecosystem‐level processes, we examined whether and how leaf diversity influences litter decomposition and consumers in streams. 2. We tested experimentally the hypothesis that the effects of leaf diversity on decomposition are determined by the responses of leaf consumers to resource–habitat heterogeneity. Leaves from three common riparian trees, beech (Fagus sylvatica), hazel (Corylus avellana) and ash (Fraxinus excelsior), were exposed alone and in all possible mixtures of two and three species in a stream. We analysed individual leaf species for decomposition rate, microbial respiration and mycelial biomass, and we determined the species composition, abundance and biomass of shredders in leaf bags. 3. We found that the decomposition of the fastest decomposing leaves (hazel and ash) was substantially stimulated (up to twofold higher than single species leaf packs) in mixtures containing beech leaves, which are refractory. In contrast, the decomposition of beech leaves was not affected by leaf mixing. Such species‐specific behaviour of leaves in species mixtures has been overlooked in previous studies that examined the overall decomposition of litter mixtures. 4. The effects of leaf diversity on decomposition varied with the abundance and biomass of shredders but not with microbial parameters. Beech leaves alone were less attractive to shredders than leaf packs made of hazel, ash or any mixture of species. Moreover, the presence of beech leaves in mixtures led to higher shredder abundance and biomass than we had expected from data from single species exposed alone. Lastly, we found that early instars of the caddisfly Potamophylax (the dominant shredder in terms of biomass) almost exclusively used the toughest material (i.e. beech leaves) to construct their cases. 5. Leaf pack heterogeneity may have altered shredder‐mediated decomposition. Shredders colonising diverse leaf packs benefited from the stable substratum provided by beech leaves, whereas ash and hazel leaves were primarily used as food. Thus, our findings provide strong evidence for an intimate linkage between the diversity of riparian vegetation and aquatic communities.     

Breakdown of four leaf litter species and associated fauna in a Basque Country forested stream

Leaves of 4 species (Quercus robur, Castanea sativa, Corylus avellana and Pinus pinaster) were incubated in a forested headwater stream using bags (1 mm and 5 mm net size) and trays (5 mm net size). The type of treatment influenced litter breakdown rates. Differences in loss rates were noted between Pinus and the deciduous species, and between Quercus and Corylus. Numbers of invertebrates per sample and per gram leaf AFDW generally increased with exposure time. The taxa colonizing the four leaf species were largely identical but colonization was greatly dependent on type of treatment. Taxa richness (Hmax) increased with time. Differences between Hmax and specific diversity (H) were high, due to the importance of Chironomids. Collectors and scrapers, among other functional groups, exhibited the highest densities. Differences in loss rates between fine and coarse treatments were related to differences in shredder biomass, corroborating the importance of this functional group in litter processing.     

Trophic cascading effects of predatory fish on leaf litter processing in a Japanese stream

A manipulative field experiment to test for trophic cascading effects of predatory fish on detritus processing by benthic invertebrates was performed in stream channels running through a wetland forest in northern Japan. To control for fish effects on benthic invertebrates, two simple treatments (fish-present and fish-absent) were established for 4 weeks, with two common predatory fish, rainbow trout (Oncorhynchus mykiss) and freshwater sculpin (Cottus nozawae), being introduced into and excluded from stream cages. At the end of experiment, the biomass of the dominant detritivore, an amphipod (Jesogammarus jezoensis), was significantly less in the fish-present treatment (0.56 g m^–2 in dry mass on average) than that in the fish-absent treatment (1.32 g m^–2), there being no significant treatment effect evident for the second-dominant detritivore, coleopteran larvae (Optioservus kubotai). The loss of oak leaves (Quercus crispla) from litter bags in the fish-present treatment (0.31 g week^–1 in dry mass on average) was significantly less than in the fish-absent treatment (0.54 g week^–1). Predator-induced lower biomass and likely lowered foraging activities of the J. jezoensis were responsible for the suppression of litter processing efficiency. In contrast, the standing crop of fine particulate organic matter did not differ significantly between the treatments. The experimental results revealed that the predatory fish had an indirect but significant effect on leaf litter processing in the stream.     

Disparate effects of plant genotypic diversity on foliage and litter arthropod communities

Intraspecific diversity can influence the structure of associated communities, though whether litter-based and foliage-based arthropod communities respond to intraspecific diversity in similar ways remains unclear. In this study, we compared the effects of host-plant genotype and genotypic diversity of the perennial plant, Solidago altissima, on the arthropod community associated with living plant tissue (foliage-based community) and microarthropods associated with leaf litter (litter-based community). We found that variation among host-plant genotypes had strong effects on the diversity and composition of foliage-based arthropods, but only weak effects on litter-based microarthropods. Furthermore, host-plant genotypic diversity was positively related to the abundance and diversity of foliage-based arthropods, and within the herbivore and predator trophic levels. In contrast, there were minimal effects of plant genotypic diversity on litter-based microarthropods in any trophic level. Our study illustrates that incorporating communities associated with living foliage and senesced litter into studies of community genetics can lead to very different conclusions about the importance of intraspecific diversity than when only foliage-based community responses are considered in isolation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The importance of bacterivorous protists in the decomposition of stream leaf litter

1. Allochthonous organic matter, in the form of senesced leaves, is a major source of carbon supporting detrital food webs. While studies have documented the role of bacteria and fungi in the decomposition of leaf litter, little information is available regarding the role of protists in the decomposition process. 2. We tested the hypothesis that the presence of stream‐dwelling bacterivorous protists leads to an increased rate of leaf decomposition through grazing pressure on bacteria. We isolated live protists from decomposing leaves collected in a stream in Northern Virginia, U.S.A. (Goose Creek) and established laboratory cultures of common bacterivorous protists. 3. Recently senesced leaves from the field were used in laboratory microcosm experiments to determine if the rate of litter decomposition differed between four treatments: bacteria only, bacteria + flagellates, bacteria + flagellates + ciliates, autoclaved stream water (control). We determined the dry weight of leaf remaining, bacterial abundance, flagellate abundance and ciliate abundance for each replicate on days 0, 7, 14, 30, 60 and 120. 4. The rate of leaf decomposition was significantly higher in treatments with protists than without and bacterial abundance declined in protist treatments compared with bacteria only treatment. Weight loss in the presence of flagellates was three to four times higher when protists were present compared with treatments with bacteria alone. These results provide experimental evidence that protists could play a significant role in the detrital processes of streams.     

Species richness at the guild level: effects of species pool and local environmental conditions on stream macroinvertebrate communities

1.?A fundamental question in ecology is which factors determine species richness. Here, we studied the relative importance of regional species pool and local environmental characteristics in determining local species richness (LSR). Typically, this question has been studied using whole communities or a certain taxonomic group, although including species with widely varying biological traits in the same analysis may hinder the detection of ecologically meaningful patterns. 2.?We studied the question above for whole stream macroinvertebrate community and within functional feeding guilds. We defined the local scale as a riffle site and the regional scale (i.e. representing the regional species pool) as a stream. Such intermediate-sized regional scale is rarely studied in this context. 3.?We sampled altogether 100 sites, ten riffles (local scale) in each of ten streams (regional scale). We used the local-regional richness regression plots to study the overall effect of regional species pool on LSR. Variation partitioning was used to determine the relative importance of regional species pool and local environmental conditions for species richness. 4.?The local-regional richness relationship was mainly linear, suggesting strong species pool effects. Only one guild showed some signs of curvilinearity. However, variation partitioning showed that local environmental characteristics accounted for a larger fraction of variance in LSR than regional species pool. Also, the relative importance of the fractions differed between the whole community and guilds, as well as among guilds. 5.?This study indicates that the importance of the local and regional processes may vary depending on feeding guild and trophic level. We conclude that both the size of the regional species pool and local habitat characteristics are important in determining LSR of stream macroinvertebrates. Our results are in agreement with recent large-scale studies conducted in highly different study systems and complement the previous findings by showing that the interplay of regional and local factors is also important at intermediate regional scales.     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

Variable effects of air-drying on leaching losses from tree leaf litter

Leaching of soluble substances may be an important first step in leaf litter decomposition in small streams, but recent research has suggested that large leaching losses (up to 30% of initial mass in 48 h) may be an artifact created by using air-dried leaves in decomposition experiments. In laboratory experiments, we compared 3 d leaching losses from freshly fallen and air-dried senescent leaves of 27 tree species from different regions across Canada. Air-dried leaves from all species leached measurable amounts of original mass (3.6–32.8% dry mass), but leaching losses from fresh leaves (0–35%) were detectable in all but two species. Air-drying increased leaching losses in many species, but in others it reduced leaching losses or had no measurable effect. Results for leaves of the same species collected in different regions or in different years were generally similar, but species within the same genus often behaved very differently. Neither moisture content (fresh or air-dried), leaf thickness, nor cuticle thickness proved of any value as predictors of leaching losses or the effect of air-drying. The propensity of autumn-fallen leaves to leach, whether fresh or air-dried, appears to be a property of the individual tree species.     

The effects of bankside management on chalk stream invertebrate communities

SUMMARY 1. Communities of aquatic macroinvertebrates and the terrestrial adult phases of aquatic insects were investigated from short stretches of English chalk streams with two different bankside vegetation types: simply structured grazed grass (grazed) and structurally complex herbaceous vegetation with scattered trees (ungrazed). Macroinvertebrates were sampled in spring, summer, autumn and winter 1996–97 from three aquatic habitats: mid-channel gravel, patches of the aquatic macrophyte Ranunculus and marginal emergent macrophytes. The terrestrial adult phases of aquatic insects were sampled in spring, summer and autumn from bankside vegetation.
2. Total macroinvertebrate abundance did not differ between stretches with different bankside vegetation. Taxon richness of mid-channel gravel was, however, significantly higher in ungrazed compared with grazed stretches and Shannon diversity ( H ') of mid-channel gravel and marginal vegetation was significantly higher in ungrazed compared with grazed stretches. Total abundance, taxon richness and Shannon diversity ( H ') of the terrestrial adult phases of aquatic insect were significantly higher from the bankside vegetation of ungrazed compared with grazed stretches.
3. Ordination of communities of aquatic macroinvertebrates and terrestrial adults demonstrated that individual families of both groups were generally more abundant in ungrazed stretches. Many more families were significantly associated with ungrazed stretches than with grazed stretches.
4. This investigation has shown that high structural diversity of bankside vegetation along lowland chalk streams is accompanied at the reach scale by increased diversity of both aquatic macroinvertebrates and the terrestrial adult phases of aquatic insects. The conservation potential of such streams may thus be lowered by management practices that result in the removal or simplification of bankside vegetation along extensive stream stretches.     

Dynamics of yeast populations recovered from decaying leaves in a nonpolluted stream: a 2-year study on the effects of leaf litter type and decomposition time

Here we report on the results of a survey of the yeast populations occurring on submerged leaves (alder, eucalyptus and oak) in a natural mountain stream, during different phases of their decomposition and through two consecutive years. Leaf litter mass loss, total yeast counts, Shannon-Weiner index (H'), yeast community structure and physiologic abilities were analyzed to evaluate the dynamics of yeast communities during decay. Seventy-two yeast taxa were recorded, and in all litter types, species of basidiomycetous affinity predominated over ascomycetous ones. Discriminant analysis of presence/absence data (yeast species) showed significant differences both among substrate types (P<0.0026) and with decomposition time (P<0.0001). Carbon and nitrogen source utilization by yeast strains also varied with the substrate (P<0.0001) and decomposition time (P<0.0001). Further conclusions were that: (1) all litter types have in common ubiquitous yeast species, such as Cryptococcus albidus, Debaryomyces hansenii and Rhodotorula glutinis, among the common 20 yeast species; (2) only a few species were dominant, and most species were rare, being recorded once or twice throughout decomposition; and (3) the order of yeast appearance, and their substrate assimilation patterns, strongly suggest a succession phenomenon. Finally, explanations for the distribution patterns and variations in yeast communities are discussed.     

Effects of eucalyptus afforestation on leaf litter dynamics and macroinvertebrate community structure of streams in Central Portugal

To test the hypothesis whether afforestation with Eucalyptus globulus affects litter dynamics in streams and the structure of macroinvertebrate aquatic communities, we compared streams flowing through eucalyptus and deciduous forests, paying attention to: (i) litterfall dynamics, (ii) accumulation of organic matter, (iii) processing rates of two dominant leaf species: eucalyptus and chestnut, and (iv) macroinvertebrate community structure. The amount of allochthonous inputs was similar in both vegetation types, but the seasonality of litter inputs differed between eucalyptus and natural deciduous forests. Eucalyptus forest streams accumulated more organic matter than deciduous forest streams. Decomposition of both eucalyptus and chestnut leaf litter was higher in streams flowing through deciduous forests. The eucalyptus forest soils were highly hydrophobic resulting in strong seasonal fluctuations in discharge. In autumn the communities of benthic macroinvertebrates of the two stream types were significantly different. Deciduous forest streams contained higher numbers of invertebrates and more taxa than eucalyptus forest streams. Mixed forest streams (streams flowing through eucalyptus forests but bordered by deciduous vegetation) were intermediate between the two other vegetation types in all studied characteristics (accumulation of benthic organic matter, density and diversity of aquatic invertebrates). These results suggest that monocultures of eucalyptus affect low order stream communities. However, the impact may be attenuated if riparian corridors of original vegetation are kept in plantation forestry.     

Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

Consumption of leaf detritus by a stream shredder: Influence of tree species and nutrient status

Four species of riparian vegetation (alder, birch, willow and poplar) were fertilized with nitrogen, phosphorus, nitrogen + phosphorus, or no fertilizer (control). The resulting leaf detritus (leached but not microbially colonized) was offered to a stream shredder, Hydatophylax variabilis (Trichoptera: Limnephilidae). In one experiment, shredder consumption of leaf detritus from different nutrient treatments (within tree species) was compared, and in a second experiment, consumption of different tree species (within nutrient treatments) was compared. Larvae preferred leaf detritus from nitrogen + phosphorus treatments (except in poplar where nitrogen treatment was preferred). Alder was preferred over other tree species for all treatments. Chemical and physical analyses of leaf litter showed differences between tree species and nutrient treatments in nutrient content, tannins and leaf toughness. Leaf consumption by larvae was positively associated with nitrogen content and negatively associated with condensed tannin content. Species composition and nutrient status of riparian vegetation may strongly influence detrital food webs in streams.     

Evidence of parasite-mediated disruptive selection on genetic diversity in a wild fish population

Identifying the processes maintaining genetic variability in wild populations is a major concern in conservation and evolutionary biology. Parasite-mediated selection may strongly affect genetic variability in wild populations. The inbreeding depression theory predicts that directional selection imposed by parasites should act against the most inbred hosts, thus favouring genetic diversity in wild populations. We have tested this prediction by evaluating the strength and shape of the relationship between the load of a harmful fin-feeder ectoparasite ( Tracheliastes polycolpus ) and the genome-wide genetic diversity (i.e. heterozygosity measured at a set of 15 microsatellites) of its fish host, the rostrum dace ( Leuciscus leuciscus ). Contrary to expectation, we found a nonlinear relationship between host genetic diversity and ectoparasite load, with hosts that were either homozygous or heterozygous harbouring significantly fewer parasites than hosts with an intermediate level of heterozygosity. This relationship suggests that parasites could increase the variance of global heterozygosity in this host population through disruptive selection on genetic diversity. Moreover, when genetic diversity was measured at each locus separately, we found two very strong positive associations between host genetic diversity and the ectoparasite load. This latter result has three main implications: (i) genome-wide effect cannot alone explain the nonlinear relationship between global heterozygosity and ectoparasite load, (ii) negative non-additive allelic interactions (i.e. underdominance) may be a mechanism for resisting ectoparasite infection, and (iii) ectoparasites may favour homozygosity at some loci in this host population.     

The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities

The prevalence of non-indigenous species (NIS) in southern California embayments was assessed from 123 Van Veen grab samples collected in nine bays and harbors during the summer of 1998. NIS occurred in all but two samples. They accounted for only 4.3% of the 633 taxa but contributed 27.5% of the abundance. There was no significant difference in the proportion of NIS abundance among ports harboring large vessels, small boat marinas, and areas where boats were not moored. Three species accounted for 92% of the NIS abundance: a spionid polychaete worm Pseudopolydora paucibranchiata, a mytilid bivalve Musculista senhousia, and a semelid bivalve Theora ubrica. The NIS did not appear to have a negative impact at the overall community level since NIS abundance was positively correlated with the abundance and richness of other species. This may be due to biogenic structures built by P. paucibranchiata and M. senhousia that enhance the abundances of other macrofauna.     

Nutrient dependent effects of consumer identity and diversity on freshwater ecosystem function

1. Over the past decade, ecologists have tried to determine how changes in species composition and diversity affect ecosystem structure and function. Until recently, the majority of these studies have been conducted in terrestrial ecosystems and have not taken into account environmental variability. The purpose of this research was to determine how species identity and diversity in the freshwater zooplankton affected biomass of algae and zooplankton at two levels of nutrient enrichment.
2. Several species of cladocerans were grown alone and together in microcosms at both ambient and raised phosphorus concentrations to determine if the effects of consumer identity and diversity were nutrient dependent.
3. Total zooplankton biomass was greater, while algal biomass was lower, in mixed culture than in monoculture. The effects of zooplankton diversity on algal biomass, however, were only observed at raised phosphorus concentrations, suggesting that diversity effects were nutrient dependent. Specifically, diversity effects appeared to be related with biological mechanisms such as complementarity in resource use and/or facilitation.
4. More diverse communities of zooplankton appear to be better able to control algae than single species of zooplankton at high nutrient concentrations; therefore, zooplankton diversity may provide a buffer against eutrophication in freshwater ecosystems.