Ecosystem Linkages between Southern Appalachian Headwater Streams and Their Banks: Leaf Litter Breakdown and Invertebrate Assemblages

We examined red maple (Acer rubrum L.) leaf litter breakdown in streams and riparian zones at two sites in the southern Appalachian Mountains to understand how differences in abiotic and biotic factors influence leaf breakdown rates. Litterbags were placed in three riparian habitats differing in litter layer moisture: stream > bank > upland. Invertebrates colonizing litterbags at one site were also examined to determine how variations in community and functional structure affect breakdown rates. Leaves broke down fastest in streams and slowest in upland habitats, whereas bank habitats were intermediate and characterized by high variability. Faster leaf breakdown rates in streams appeared to be a function of greater moisture availability, a more stable thermal regime, and a higher biomass of leaf-shredding invertebrates, especially the stonefly Tallaperla. In addition, patterns of leaf breakdown and invertebrate community structure provided evidence for a stronger than expected ecological connection between the stream and the bank. Overall, detritus processing within this narrow riparian ecosystem varied considerably depending on the availability of moisture. Results from this study show that stream channel–floodplain interactions in riparian ecosystems of steep forested mountains are analogous to ones in larger downstream or low-gradient systems. Riparian zones throughout a river network display a remarkable heterogeneity in their ability to process organic matter, which is ultimately driven by changes in hydrological conditions. Received 6 March 2001; accepted 3 July 2001.     

Identifying the Relative Importance of Leaf versus Shredder Species Loss on Litter Decomposition in Streams

Understanding how species loss influences ecosystem function is a contemporary issue in ecology. However, most research has focused on species loss at one trophic‐level. We explored the relationship between functional diversity (FD) and species richness separately for trees and aquatic leaf‐shredding detritivores. For trees, we collected information on species‐specific leaf tissue chemistry and species co‐occurrences in the mid‐Atlantic region (USA). For shredders, we used a published trait database with information on communities from 38 streams in the same region. We used a clustering algorithm to estimate FD for each community and for randomly assembled communities. If FD was high, we concluded that species loss was important to change in function; if low, species were functionally redundant and insensitive to species loss. We found tree FD to be significantly different than expected, but shredders exhibited FD levels similar to patterns based on random assembly. Furthermore, there were more leaf species exclusively associated with very high or very low levels of functional diversity compared to shredders. This approach revealed greater implications for leaf than shredder species loss for litter breakdown. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Aquatic Hyphomycete Communities Associated with Decomposing Alder Leaf Litter in Reference Headwater Streams of the Basque Country (northern Spain)

The community of aquatic hyphomycetes associated with decomposing alder leaf litter was studied during autumn-winter in nine headwater reference streams of the Basque Country (northern Spain). In order to study the spatial variability in composition and community structure, three streams from each of three different river basins were compared. The colonization dynamics and community changes throughout the decomposition process were also followed in three of the rivers (one per basin). The taxonomic richness and community structure of these fungi varied among rivers, including similar streams of a given watershed. However, neither species diversity nor total abundance was statistically related to environmental variables. Only the conidial production of two of the species, Flagellospora curvula and Lunulospora curvula appeared to be enhanced by nitrate availability in the water. The taxonomic richness and the reproductive activity (sporulation rate) were positively related to the leaf litter decomposition rate. The changes in conidial production along the process were similar for all the streams and helped explain leaf litter quality dynamics.     

Leaf Litter as a Source of Dissolved Organic Carbon in Streams

Dissolved organic carbon (DOC) is an abundant form of organic matter in stream ecosystems. Most research has focused on the watershed as the source of DOC in streams, but DOC also comes from leaching of organic matter stored in the stream channel. We used a whole-ecosystem experimental approach to assess the significance of leaching of organic matter in the channel as a source of DOC in a headwater stream. Inputs of leaf litter were excluded from a forested Appalachian headwater stream for 3 years. Stream-water concentration, export, and instream generation of DOC were reduced in the litter-excluded stream as compared with a nearby untreated reference stream. The proportion of high molecular weight (HMW) DOC (more than 10,000 daltons) in stream water was not altered by litter exclusion. Mean DOC concentration in stream water was directly related to benthic leaf-litter standing stock. Instream generation of DOC from leaf litter stored in the stream channel contributes approximately 30% of daily DOC exports in this forested headwater stream. This source of DOC is greatest during autumn and winter and least during spring and summer. It is higher during increasing discharge than during base flow. We conclude that elimination of litter inputs from a forested headwater stream has altered the biogeochemistry of DOC in this ecosystem. Received 2 September 1997; accepted 27 January 1998.     

Spatial Variability in Litterfall,Litter Standing Crop and Litter Quality in a Tropical Rain Forest Region

Understanding the spatial variability in plant litter processes is essential for accurate comprehension of biogeochemical cycles and ecosystem function. We assessed spatial patterns in litter processes from local to regional scales, at sites throughout the wet tropical rain forests of northern Australia. We aimed to determine the controls (e.g., climate, soil, plant community composition) on annual litter standing crop, annual litterfall rate and in situ leaf litter decomposability. The level of spatial variance in these components, and leaf litter N, P, Ca, lignin, α‐cellulose and total phenolics, was determined from within the scale of subregion, to site (1 km transects) to local/plot (~30 m²). Overall, standing crop was modeled with litterfall and its chemical composition, in situ decomposability, soil Na, and topography (r² = 0.69, 36 plots). Litterfall was most closely aligned with plant species richness and stem density (negative correlation); leaf decomposability with leaf‐P and lignin, soil Na, and dry season moisture (r² = 0.89, 40 plots). The predominant scale of variability in litterfall rates was local (plot), while litter standing crop and α‐cellulose variability was more evenly distributed across spatial scales. Litter decomposability, N, P and phenolics were more aligned with subregional differences. Leaf litter C, lignin and Ca varied most at the site level, suggesting more local controls. We show that variability in litter quality and decomposability are more easily accounted for spatially than litterfall rates, which vary widely over short distances possibly in response to idiosyncratic patterns of disturbance.     

Responses of Aquatic Fungal Communities on Leaf Litter to Temperature‐Change Events

Increases of extreme weather events are predicted to occur with ongoing climate change, but impacts to freshwaters have rarely been examined. We assessed the effects of temperature on leaf‐litter associated fungi by exposing leaves colonized in a stream to 18 °C (control), 25 °C, or 18 °C after freezing. Treatments altered fungal dominance on leaves; Lunulospora curvula sporulation was stimulated by increased temperature and stopped by the freeze‐thaw treatment. Fungal biomass and diversity decreased at 18 °C after freezing, but not at 25 °C. Leaf decomposition was retarded by the freeze‐thaw treatment (k = –0.024 day^–1) and stimulated at 25 °C (k = –0.069 day^–1). Results suggest that occasional freezing may constrain fungal diversity and their ecological functions, while warming appears to accelerate plant‐litter decomposition in streams. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effect of Drought Frequency and Other Reach Characteristics on Invertebrate Communities and Litter Breakdown in the Intermittent Mediterranean River Pula (Sardinia,Italy)

Benthic invertebrate communities were sampled in spring and fall, and breakdown of reed litter was measured in winter, spring, summer and fall, at 23 sites in an intermittent Mediterranean river basin (Pula, Sardinia, Italy), and their variations were related to drought frequency and other reach characteristics (altitude, order, stream channel width and depth, vegetation cover, sediment type, organic content, and water quality). 78% of the sites were affected by drought events, mainly in summer and fall. Benthic taxonomic richness was higher and most taxa were more widely distributed in spring than in fall. Canonical correspondence analysis showed that measured environmental variables accounted for 72% of the variance in invertebrate community structure. Reed decay rates varied among seasons, and during the dry season among stream orders as well. Drought events slowed down breakdown and explained 44% of spatial variability in decay rates (Stepwise multiple regression analysis), an additional 8% being explained by water chemistry. Results suggest that drought events affect ecosystem functioning (litter breakdown) more than structure (invertebrate communities). (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of Zn,Fe and Mn on Leaf Litter Breakdown by Aquatic Fungi: a Microcosm Study

We investigated the effects of heavy metals on leaf litter decomposition in streams. Leaves were immersed (10 days) at a reference (R) and a metal‐impacted (I) site and exposed in microcosms with increased Zn, Mn or Fe content, and to stream water from site R or I. Fungal biomass was higher in microcosms with leaves colonized at I and water from R. Fungal sporulation was higher in microcosms with leaves and water from R. Concentrations of 4.9, 9.6 and 5 ppm of Zn, Mn and Fe decrease fungal sporulation. The number of fungal species (spore counts and DGGE fingerprints) was lower in leaves colonized at site I. Cluster analyses of DGGE showed that Fe was the metal that most altered the structure of fungal community. Our results suggest that metal pollution affect leaf‐associated fungi depending on metal identity and concentration, and effects appear to be less pronounced in metal‐adapted communities. (© 2010 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Regional Intra‐Subject Variability in Abdominal Adiposity Limits Usefulness of Computed Tomography

Objective: Computed tomography (CT) and magnetic resonance imaging, the most accurate methods of abdominal fat measurement, have been applied using a number of protocols, ranging from single‐slice area determination to multiple‐slice volume calculation. The aim of this study was to assess the validity of single‐slice CT for abdominal fat area measurement by estimating the intra‐subject variability in abdominal fat areas and comparing the ranking of subjects across four contiguous abdominal levels. Research Methods and Procedures: Nineteen premenopausal women (age, 35.3 ± 1.4 years; mean ± SE) were studied. CT was used to measure intra‐abdominal fat (IAF) area, percentage of total intra‐abdominal area (%IAF), subcutaneous abdominal fat (SAF) area, and IAF/SAF at four adjacent cross‐sectional lumbar levels (L2–L4). Intra‐subject variability (percentage) was defined as SD/mean × 100. Total body fat was measured by DXA, which was further analyzed for central abdominal fat. Results: Mean body mass index was 24.9 ± 1.0 kg/m². The average (range) intra‐subject variability was 28% (8% to 61%) for IAF, 46% (19% to 124%) for %IAF, 26% (14% to 38%) for SAF area, and 19% (7% to 71%) for IAF/SAF. The pattern of this variability was not uniform between subjects, because their ranking by IAF area was markedly different at each CT level. Discussion: We demonstrated significant intra‐subject variability in CT‐measured adipose tissue areas across four predetermined sites. This resulted in a difference in the ordering of subjects by IAF at each of the four imaging sites, suggesting that the usefulness of single‐slice CT in the assessment of abdominal adiposity in premenopausal women may be limited, particularly when performed for the purpose of making comparisons between subjects based on abdominal fat area.     

A Comparative Study of Leaf Breakdown of Three Native Tree Species in a Slowly‐Flowing Headwater Stream in the Colombian Andes

The dynamics of leaf breakdown in a headwater Colombian stream were evaluated for the native tree species Myrsine guianensis, Cupania latifolia and Nectandra lineatifolia using coarse and fine mesh litter bags. Ten bags of each species (five of each mesh size) were retrieved from the stream at 1, 8, 15, 30, 60 and 120 days. k values ranged from 0.0008 to 0.0058 day^–1 and density of macroinvertebrates from 35 to 55 individuals per leaf bag, peaking at day 8. Myrsine guianensis degraded more rapidly than the other species for both coarse and fine mesh bags. This species and Nectandra lineatifolia presented differences in k values between coarse and fine mesh bags, suggesting that macroinvertebrates influenced the decay rate. Despite the low densities of macroinvertebrates found, shredders represented 12.7% of individuals and 50 to 68% of the invertebrate biomass in bags, indicating that this functional feeding group was an important component of fauna associated with litter breakdown in this first order tropical stream. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Chemical and Morphological Inter‐ and Intrapopulation Variability in Natural Populations of Gentiana pneumonanthe L.

Inter‐ and intrapopulation variability in six natural populations of the rare species Gentiana pneumonanthe was examined based on morphological and chemical data. Population size and linear morphometric parameters differed significantly among populations, but without a clear connection to habitat conditions, i. e. water supply and light availability. Leaf shape varied from ovate to lanceolate in all populations, and one population was distinctive in having the largest number of leaves of transitional shape. HPLC analyses of six secondary metabolites were performed separately for belowground parts, and aboveground vegetative and reproductive parts of individual plants (6 populations ×7 individuals ×3 plant parts, n=126) in order to examine differences at the population and individual levels. Three secoiridoids (swertiamarin (SWM), sweroside (SWZ), and gentiopicrin (GP)), one xanthone (mangiferin (MGF)), and two flavones (isoorientin (IO) and isovitexin (IV)) were detected and quantified in the analyzed samples: sweroside dominated in the aboveground reproductive part, mangiferin in the aboveground vegetative part, and gentiopicrin in the belowground part. At the population level, differences in contents of the analyzed chemicals among populations were significant only for a few metabolites. At the individual level, a pronounced organ‐dependent distribution of secondary metabolites was revealed. The results of this study contribute to a better understanding of natural variability within populations of the rare and threatened G. pneumonanthe, and provide data on the contents and within‐plant distribution of secondary metabolites, which are important as pharmacologically active compounds and may be useful for further biotechnological procedures regarding this species.     

Effects of Fungal Inocula and Habitat Conditions on Alder and Eucalyptus Leaf Litter Decomposition in Streams of Northern Spain

We investigated how fungal decomposer (aquatic hyphomycetes) communities colonizing alder and eucalyptus leaf litter respond to changes in habitat characteristics (transplantation experiment). We examined the breakdown of leaf materials and the associated fungal communities at two contrasting sites, a headwater stream (H) and a midreach (M). Agroforestry increased from headwater to midreach. One month after the start of experiments at both sites, some leaf samples from the midreach site were transplanted to the headwater site (M–H treatment). Although both sites showed similar dissolved inorganic nutrient concentrations, eucalyptus leaves initially incubated at the midreach site (M, M–H) increased their breakdown rate compared to those incubated along the experiment at the headwater site (H). Alder breakdown rate was not enhanced, suggesting that their consumption was not limited by nutrient availability. Sporulation rates clearly differed between leaf types (alder > eucalyptus) and streams (H > M), but no transplantation effect was detected. When comparing conidial assemblages after transplantation, an inoculum effect (persistence of early colonizing species) was clear in both leaf species. Substrate preference and shifts in the relative importance of some fungal species along the process were also observed. Overall, our results support the determining role of the initial conditioning phase on the whole litter breakdown process, highlighting the importance of intrinsic leaf characteristics and those of the incubation habitat.     

Fungal and Bacterial Colonization of Submerged Leaf Litter in a Mediterranean Stream

Microbial colonization dynamics of fungi and bacteria were analyzed in an intermittent Mediterranean forested stream using two different leaf substrata (Platanus acerifolia and Populus nigra). Results showed that fungal and bacterial biomass accumulation was stimulated on both leaves due to a flooding episode that increased dissolved inorganic nitrogen (DIN) and dissolved oxygen (DO) availability in the stream water. Leaf mass loss coincided with the parallel increase in microbial biomass and extracellular enzymatic activities after the flood event. Differences in litter quality favoured bacterial biomass accumulation and β‐glucosidase and cellobiohydrolase enzymatic activities in the soft Populus species. Microbial heterotrophs colonization of submerged leaf litter and organic matter use in Mediterranean‐type streams are modulated by environmental conditions, especially the hydrological variability. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Comparison of Leaf Litter Decomposition Rates in Restored and Mature Riparian Forests on the Sacramento River, California

One of the largest riparian restoration projects in the United States is currently taking place in California on the Sacramento River. Nearly 2,000 ha of land adjacent to the river channel have been revegetated with native riparian species in an effort to reestablish riparian forests. The objective of this study was to compare leaf litter decomposition rates in restored riparian forests to those in mature, naturally established riparian forests, in order to monitor the development of this ecosystem function in restored forests. Leaf litter decomposition rates were measured over 1 year in six restored riparian forests (4, 7, and 9 years old) and two mature remnant riparian forests (>50 years old), in order to test two hypotheses: (1) decomposition rates of restored and mature forests are significantly different and (2) decomposition rates in the chronosequence of restored forests are moving along a trajectory, approaching the decomposition rates characteristic of mature forests as they age. Statistical analyses revealed no significant differences in annual decay rates among the four different forest ages and no trajectory among leaf litter decomposition rates in restored forests. These results suggest that a functionally equivalent process of leaf litter decomposition occurs in both restored and naturally established forests and show promise for the efficiency of nutrient cycling processes in these restored forests.     

Regional patterns and controls of leaf decomposition in Australian tropical rainforests

Future climates have the potential to alter decomposition rates in tropical forest with implications for carbon emissions, nutrient cycling and retention of standing litter. However, our ability to predict impacts, particularly for seasonally wet forests in the old world, is limited by a paucity of data, a limited understanding of the relative importance of different aspects of climate and the extent to which decomposition rates are constrained by factors other than climate (e.g. soil, vegetation composition). We used the litterbag method to determine leaf litter decay rates at 18 sites distributed throughout the Australian wet tropics bioregion over a 14‐month period. Specifically, we investigated regional controls on litter decay including climate, soil and litter chemical quality. We used both in situ litter collected from litterfall on site and a standardized control leaf litter substrate. The control litter removed the effect of litter chemical quality and the in situ study quantified decomposition specific to the site. Decomposition was generally slower than for other tropical rainforests globally except in our wet and nutrient‐richer sites. This is most likely attributable to the higher latitude, often highly seasonal rainfall and very poor soils in our system. Decomposition rates were best explained by a combination of climate, soil and litter quality. For in situ litter (native to the site) this included: average leaf wetness in the dry season (LWDS; i.e. moisture condensation) and the initial P content of the leaves, or LWDS and initial C. For control litter (no litter quality effect) this included: rainfall seasonality (% dry season days with 0‐mm rainfall), soil P and mean annual temperature. These results suggest that the impact of climate change on decomposition rates within Australian tropical rainforests will be critically dependent on the trajectory of dry season moisture inputs over the coming decades.     

Leaf Litter Dynamics and Its Use by Invertebrates in a High‐Altitude Tropical Andean Stream

This study evaluates litter dynamics and its relation to macroinvertebrate communities (assemblages and feeding habits) at Piburja stream (3300 m, Ecuador). Annual litterfall (216 g AFDM/m²) was not related to rainfall, but differed significantly between months. Transport of Coarse Particulate Organic Matter (CPOM) did not differ between seasons, but retention was higher during the dry season. Thus, CPOM standing stock was higher in the dry (125.28 g AFDM/m²) compared to the wet season (12.27 g AFDM/m²). Macroinvertebrate richness and diversity were positively related to Coarse Benthic Organic Matter (CBOM) especially during the wet season. Gut content analysis revealed that, regardless of season, CPOM and Fine Particulate Organic Matter (FPOM) were the most important food items in the diet of most invertebrate taxa, including those that are not considered shredders or gathering collectors in the literature. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Leaf‐Litter Mixtures Affect Breakdown and Macroinvertebrate Colonization Rates in a Stream Ecosystem

Previous work in terrestrial and aquatic ecosystems has suggested that the relationship between breakdown rates of leaf litter and plant species richness may change unpredictability due to non‐additive effects mediated by the presence of key‐species. By using single‐ and mixed‐species leaf bags (7 possible combinations of three litter species differing in toughness; common alder [Alnus glutinosa ], sweet chestnut [Castanea sativa ], and Spanish oak [Quercus ilex ilex ]), I tested whether leaf species diversity, measured as richness and composition, affects breakdown dynamics and macroinvertebrate colonization (abundance, richness and composition) during 90 days incubation in a stream. Decomposition rates were additive, i.e., observed decomposition rates were not different from expected ones. However, decomposition rates of individual leaf species were affected by the mixture, i.e., there were species‐specific responses to mixing litter. The invertebrate communities colonizing the mixtures were not richer and more diverse in mixtures than in single‐species leaf bags. On the opposite, mixing leaf species had a negative, non‐additive effect on rates of shredder and taxa colonization and on macroinvertebrate diversity. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Catchment land-use, macroinvertebrates and detritus processing in headwater streams: taxonomic richness versus function

1. We used a litter bag technique to assess the effect of catchment land-use (forest, wetland, agriculture, urban) on the processing of red maple ( Acer rubrum L.) litter in 17 streams in Maine, U.S.A. Litter processing by fungi was predicted to increase with nutrient concentrations along a gradient of land use, from relatively unmodified to highly modified. Litter processing by litter-shredding macroinvertebrates was predicted to decline along this gradient because of a decline in their taxonomic richness and biomass.
2. Land use was associated with the anticipated gradient in nutrient and macroinvertebrate attributes, and a significant relationship was found between land use and nitrate concentration. There was, however, no significant relationship between land use and soluble reactive phosphorus (SRP) concentration. Similarly, shredder taxonomic richness was significantly related to land use type, whereas shredder biomass showed no significant relationship to land use.
3. Attributes of the shredder assemblage structure and nutrient concentrations were both strong determinants of litter processing. Increasing biomass and taxonomic richness of shredders was significantly related to increasing rates of litter mass loss. Increasing concentrations of nitrate and SRP were significantly related to increasing rates of litter softening below threshold concentrations (approximately 0.20 mg NO₃-N L^–1 and 5 μg SRP L^–1).
4. The potentially additive effects of nitrate and SRP concentrations or shredder richness and biomass on litter processing rates were confounded by the lack of significant correlation between these pairs of variables. Consequently, rates of litter processing (as rates of softening or mass loss) did not vary systematically among different land use regimes.     

Among-habitat and Temporal Variability of Selected Macroinvertebrate Based Metrics in a Mediterranean Shallow Lake (NW Spain)

According to the European Water Framework Directive, waterbodies have to be classified on the basis of their ecological status using biological quality elements, such as macroinvertebrates. This needs to take into consideration the influence of natural variation (both spatial and temporal) of reference biological communities as this may obscure the effects caused by anthropogenic disturbance. We studied the influence of among-habitat and temporal (seasonal and interannual) changes on the macroinvertebrate communities of an Iberian shallow lake and the variability of 21 measures potentially useful for bioindication purposes. Two series of data were examined: (a) macroinvertebrate samples taken on four occasions over an annual cycle were used to assess the effects of seasonality and among-habitat variability; (b) macroinvertebrate samples collected in three consecutive summers were used to assess interannual variability. Coefficients of variation, expressed as percentage, were used to quantify the effect of among-habitat and temporal variability on the selected metrics. According to our results, % Insecta, Shannon–Wiener diversity index and the qualitative taxonomic metrics (measures based on number of taxa) were robust in terms of temporal (seasonal and interannual) and among-habitat variability. Abundance ratio and some metrics based on functional feeding groups were highly variable. Therefore, qualitative taxonomic metrics may be promising tools in biomonitoring programs of Mediterranean shallow lakes due to their comparatively low variability.