Abstract Leaf breakdown is an important process in forested headwater streams. A common method used to quantify the role of macroinvertebrate and microbial communities in leaf litter breakdown involves using paired mesh bags that either allow or exclude macroinvertebrate access to leaves. We examined common assumptions of the paired litterbag method to test (1) whether mesh size alters microbial respiration and (2) whether the effects of abrasive flows (e.g., from water and sediment) differ between coarse‐ and fine‐mesh litterbags. We measured rates of microbial respiration on Acer rubrum and Rhododendron maximum leaves incubated in coarse‐ and fine‐mesh litterbags. We also measured rates of abrasion using aerated concrete blocks in pairs of coarse‐ and fine‐mesh bags in ten streams across a gradient of discharge. We found that rates of microbial respiration on Acer rubrum leaves conditioned in fine‐mesh bags were 65% greater than the rates of respiration in paired coarse‐mesh bags, but respiration rates on Rhododendron maximum were similar in coarse‐ and fine‐mesh bags. Abrasion was, on average, 56% greater in coarse‐mesh than paired fine‐mesh bags, and these effects were greater in streams with higher discharge. These results suggest that more caution is required when attributing the difference in leaf breakdown between coarse‐ and fine‐mesh bags to macroinvertebrates. Because the effect of mesh size on microbial respiration of Acer leaves and abrasion are opposite in direction, the effect that dominates and creates bias likely depends on both environmental context and experimental design.
more »
« less
Decomposing decomposition: isolating direct effects of temperature from other drivers of detrital processing
Understanding the observed temperature dependence of decomposition (i.e., its apparent activation energy) requires separation of direct effects of temperature on consumer metabolism (i.e., the inherent activation energy) from those driven by indirect seasonal patterns in phenology and biomass, and by longer‐term, climate‐driven shifts in acclimation, adaptation, and community assembly. Such parsing is important because studies that relate temperature to decomposition usually involve multi‐season data and/or spatial proxies for long‐term shifts, and so incorporate these indirect factors. The various effects of such factors can obscure the inherent temperature dependence of detrital processing. Separating the inherent temperature dependence of decomposition from other drivers is important for accurate prediction of the contribution of detritus‐sourced greenhouse gases to climate warming and requires novel approaches to data collection and analysis. Here, we present breakdown rates of red maple litter incubated in coarse‐ and fine‐mesh litterbags (the latter excluding macroinvertebrates) for serial approximately one‐month increments over one year in nine streams along a natural temperature gradient (mean annual: 12.8°–16.4°C) from north Georgia to central Alabama, USA. We analyzed these data using distance‐based redundancy analysis and generalized additive mixed models to parse the dependence of decomposition rates on temperature, seasonality, and shredding macroinvertebrate biomass. Microbial decomposition in fine‐mesh bags was significantly influenced by both temperature and seasonality. Accounting for seasonality corrected the temperature dependence of decomposition rate from 0.25 to 0.08 eV. Shredder assemblage structure in coarse‐mesh bags was related to temperature across both sites and seasons, shifting from “cold” stonefly‐dominated communities to “warm” communities dominated by snails or crayfish. Shredder biomass was not a significant predictor of either coarse‐mesh or macroinvertebrate‐mediated (i.e., coarse‐ minus fine‐mesh) breakdown rates, which were also jointly influenced by temperature and seasonality. Unlike fine‐mesh bags, however, temperature dependence of litter breakdown did not differ between models with and without seasonality for either coarse‐mesh (0.36 eV) or macroinvertebrate‐mediated (0.13 eV) rates. We conclude that indirect (non‐thermal) seasonal and site‐level effects play a variable and potentially strong role in shaping the apparent temperature dependence of detrital breakdown. Such effects should be incorporated into studies designed to estimate inherent temperature dependence of slow ecological processes.
more » « less- Award ID(s):
- 1655956
- NSF-PAR ID:
- 10443844
- Publisher / Repository:
- Wiley Blackwell (John Wiley & Sons)
- Date Published:
- Journal Name:
- Ecology
- Volume:
- 102
- Issue:
- 10
- ISSN:
- 0012-9658
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
-
more » « less
Abstract Decomposition of coarse detritus (e.g., dead organic matter larger than ~1 mm such as leaf litter or animal carcasses) in freshwater ecosystems is well described in terms of mass loss, particularly as rates that compress mass loss into one number (e.g., a first‐order decay coefficient, or breakdown rate, “
k ”); less described are temporal changes in the elemental composition of these materials during decomposition, with important implications for elemental cycling from microbes to ecosystems. This stands in contrast with work in the terrestrial realm, where a focus on detrital elemental cycling has provided a sharper mechanistic understanding of decomposition, especially with specific processes such as immobilization and mineralization. Notably, freshwater ecologists often measure carbon (C), nitrogen (N), and phosphorus (P), and their stoichiometric ratios in decomposing coarse materials, including carcasses, wood, leaf litter, and more, but these measurements remain piecemeal. These detrital nutrients are measurements of the entire detrital–microbial complex and are integrative of numerous processes, especially nutrient immobilization and mineralization, and associated microbial growth and death. Thus, data relevant to an elemental, mechanistically focused decomposition ecology are available in freshwaters, but have not been fully applied to that purpose. We synthesized published detrital nutrient and stoichiometry measurements at a global scale, yielding 4038 observations comprising 810 decomposition time series (i.e., measurements within a defined cohort of decomposing material through time) to build a basis for understanding the temporality of elemental content in freshwater detritus. Specifically, the dataset focuses on temporally and ontogenetically (mass loss) explicit measurements of N, P, and stoichiometry (C:N, C:P, N:P). We also collected ancillary data, including detrital characteristics (e.g., species, lignin content), water physiochemistry, geographic location, incubation system type, and methodological variables (e.g., bag mesh size). These measurements are important to unlocking mechanistic insights into detrital ontogeny (the temporal trajectory of decomposing materials) that can provide a deeper understanding of heterotroph‐driven C and nutrient cycling in freshwaters. Moreover, these data can help to bridge aquatic and terrestrial decomposition ecology, across plant or animal origin. By focusing on temporal trajectories of elements, this dataset facilitates cross‐ecosystem comparisons of fundamental decomposition controls on elemental fluxes. It provides a strong starting point (e.g., via modeling efforts) for comparing processes such as immobilization and mineralization that are understudied in freshwaters. Time series from decomposing leaf litter, particularly in streams, are common in the dataset, but we also synthesized ontogenies of animal‐based detritus, which tend to decompose rapidly compared with plant‐based detritus that contains high concentrations of structural compounds such as lignin and cellulose. Although animal‐based data were rare, comprising only three time series, their inclusion in this dataset underscores the opportunities to develop an understanding of decomposition that encompasses all detrital types, from carrion to leaf litter. There are no copyright or proprietary restrictions on the dataset; please cite this data paper when reusing these materials. -
more » « less
Abstract Models estimating decomposition rates of dead wood across space and time are mainly based on studies carried out in temperate zones where microbes are dominant drivers of decomposition. However, most dead wood biomass is found in tropical ecosystems, where termites are also important wood consumers. Given the dependence of microbial decomposition on moisture with termite decomposition thought to be more resilient to dry conditions, the relative importance of these decomposition agents is expected to shift along gradients in precipitation that affect wood moisture.
Here, we investigated the relative roles of microbes and termites in wood decomposition across precipitation gradients in space, time and with a simulated drought experiment in tropical Australia. We deployed mesh bags with non‐native pine wood blocks, allowing termite access to half the bags. Bags were collected every 6 months (end of wet and dry seasons) over a 4‐year period across five sites along a rainfall gradient (ranging from savanna to wet sclerophyll to rainforest) and within a simulated drought experiment at the wettest site. We expected microbial decomposition to proceed faster in wet conditions with greater relative influence of termites in dry conditions.
Consistent with expectations, microbial‐mediated wood decomposition was slowest in dry savanna sites, dry seasons and simulated drought conditions. Wood blocks discovered by termites decomposed 16–36% faster than blocks undiscovered by termites regardless of precipitation levels. Concurrently, termites were 10 times more likely to discover wood in dry savanna compared with wet rainforest sites, compensating for slow microbial decomposition in savannas. For wood discovered by termites, seasonality and drought did not significantly affect decomposition rates.
Taken together, we found that spatial and seasonal variation in precipitation is important in shaping wood decomposition rates as driven by termites and microbes, although these different gradients do not equally impact decomposition agents. As we better understand how climate change will affect precipitation regimes across the tropics, our results can improve predictions of how wood decomposition agents will shift with potential for altering carbon fluxes.
Read the free
Plain Language Summary for this article on the Journal blog. -
more » « less
Abstract Climate change is increasing the frequency, severity, and extent of wildfires and drought in many parts of the world, with numerous repercussions for the physical, chemical, and biological characteristics of streams. However, information on how these perturbations affect top predators and their impacts on lower trophic levels in streams is limited.
The top aquatic predator in southern California streams is native
Oncorhynchus mykiss , the endangered southern California steelhead trout (trout). To examine relationships among the distribution of trout, environmental factors, and stream invertebrate resources and assemblages, we sampled pools in 25 stream reaches that differed in the presence (nine reaches) or absence (16 reaches) of trout over 12 years, including eight reaches where trout were extirpated during the study period by drought or post‐fire flood disturbances.Trout were present in deep pools with high water and habitat quality. Invertebrate communities in trout pools were dominated by a variety of medium‐sized collector–gatherer and shredder invertebrate taxa with non‐seasonal life cycles, whereas tadpoles and large, predatory invertebrates (Odonata, Coleoptera, Hemiptera [OCH]), often with atmospheric breather traits, were more abundant in troutless than trout pools.
Structural equation modelling of the algal‐based food web indicated a trophic cascade from trout to predatory invertebrates to collector–gatherer taxa and weaker direct negative trout effects on grazers; however, both grazers and collector–gatherers also were positively related to macroalgal biomass. Structural equation modelling also suggested that bottom‐up interactions and abiotic factors drove the detritus‐based food web, with shredder abundance being positively related to leaf litter (coarse particulate organic matter) levels, which, in turn, were positively related to canopy cover and negatively related to flow. These results emphasise the context dependency of trout effects on prey communities and of the relative importance of top‐down versus bottom‐up interactions on food webs, contingent on environmental conditions (flow, light, nutrients, disturbances) and the abundances and traits of component taxa.
Invertebrate assemblage structure changed from a trout to a troutless configuration within a year or two after trout were lost owing to post‐fire scouring flows or drought. Increases in OCH abundance after trout were lost were much more variable after drought than after fire. The reappearance of trout in one stream resulted in quick, severe reductions in OCH abundance.
These results indicate that climate‐change induced disturbances can result in the extirpation of a top predator, with cascading repercussions for stream communities and food webs. This study also emphasises the importance of preserving or restoring refuge habitats, such as deep, shaded, perennial, cool stream pools with high habitat and water quality, to prevent the extirpation of sensitive species and preserve native biodiversity during a time of climate change.
-
more » « less
Abstract Increases in nitrogen (N) and phosphorus (P) availability are changing animal communities, partly by altering stoichiometric imbalances between consumers and their food. Testing relationships between resource stoichiometry and consumer assemblage structure requires ecosystem‐level manipulations that have been lacking to date.
We analysed patterns of macroinvertebrate community composition in five detritus‐based headwater streams subject to experimental whole‐stream N and P additions that spanned a steep gradient in dissolved N:P ratio (2:1, 8:1, 16:1, 32:1, 128:1) over 2 years, following a 1‐year pre‐treatment period.
We predicted that shifts in leaf litter stoichiometry would drive overall patterns of community composition via greater responses of shredders to enrichment than other taxa, as shredders dominate primary consumer biomass and experience larger consumer–resource elemental imbalances than other functional groups in stream ecosystems. Specifically, we expected litter C:P to be a significant predictor of shredder biomass given the greater relative imbalances between shredder and litter C:P than C:N. Finally, we tested whether shredder responses to enrichment were related to other taxon‐level traits, including body size and stoichiometry, larval life span and growth rate.
Whole‐community composition shifted similarly across the five streams after enrichment, largely driven by increased shredder and predator biomass. These shifts were limited to the autumn/winter seasons and related to decreased leaf litter C:P, highlighting important links between the quality of seasonal litter subsidies and community phenology.
Among 10 taxa that drove structural shifts, two declined while other taxa from the same functional/taxonomic groups responded positively, suggesting that specific life‐history traits may determine sensitivity to enrichment.
Increases in total shredder biomass, and in biomass of several common shredders, were associated with lower litter C:P. Body C:P did not predict shredder response to enrichment. However, weak negative relationships between shredder response and body size, and larval life span, suggest that small‐bodied and short‐lived taxa may be more responsive to shifting resource stoichiometry.
Moderate anthropogenic increases in N and P availability affect resource stoichiometry and can alter animal communities, influencing additional food web and ecosystem properties. We provide support for ecological stoichiometry as a framework for predicting such outcomes based on changes in the elemental composition of resource pools.
A
plain language summary is available for this article.
