Does Facilitation of Faunal Recruitment Benefit Ecosystem Restoration? An Experimental Study of Invertebrate Assemblages in Wetland Mesocosms

We used wetland mesocosms (1) to experimentally assess whether inoculating a restored wetland site with vegetation/sediment plugs from a natural wetland would alter the development of invertebrate communities relative to unaided controls and (2) to determine if stocking of a poor invertebrate colonizer could further modify community development beyond that due to simple inoculation. After filling mesocosms with soil from a drained and cultivated former wetland and restoring comparable hydrology, mesocosms were randomly assigned to one of three treatments: control (a reference for unaided community development), inoculated (received three vegetation/sediment cores from a natural wetland), and stocked + inoculated (received three cores and were stocked with a poorly dispersing invertebrate group—gastropods). All mesocosms were placed 100 m from a natural wetland and allowed to colonize for 82 days. Facilitation of invertebrate colonization led to communities in inoculated and stocked + inoculated treatments that contrasted strongly with those in the unaided control treatment. Control mesocosms had the highest taxa richness but the lowest diversity due to high densities and dominance of Tanytarsini (Diptera: Chironomidae). Community structure in inoculated and stocked + inoculated mesocosms was more similar to that of a nearby natural wetland, with abundance more evenly distributed among taxa, leading to diversity that was higher than in the control treatment. Inoculated and stocked + inoculated communities were dominated by non‐aerial invertebrates, whereas control mesocosms were dominated by aerial invertebrates. These results suggest that facilitation of invertebrate recruitment does indeed alter invertebrate community development and that facilitation may lead to a more natural community structure in less time under conditions simulating wetland restoration.     

An invasive frog, Eleutherodactylus coqui, increases new leaf production and leaf litter decomposition rates through nutrient cycling in Hawaii

A frog endemic to Puerto Rico, Eleutherodactylus coqui, invaded Hawaii in the late 1980s, where it can reach densities of 50,000 individuals ha⁻¹. Effects of this introduced insectivore on invertebrate communities and ecosystem processes, such as nutrient cycling, are largely unknown. In two study sites on the Island of Hawaii, we studied the top-down effects of E. coqui on aerial, herbivorous, and leaf litter invertebrates; herbivory, plant growth, and leaf litter decomposition rates; and leaf litter and throughfall chemistry over 6 months. We found that E. coqui reduced all invertebrate communities at one of the two study sites. Across sites, E. coqui lowered herbivory rates, increased NH₄⁺ and P concentrations in throughfall, increased Mg, N, P, and K in decomposing leaf litter, increased new leaf production of Psidium cattleianum, and increased leaf litter decomposition rates of Metrosideros polymorpha. In summary, E. coqui effects on invertebrates differed by site, but E. coqui effects on ecosystem processes were similar across sites. Path analyses suggest that E. coqui increased the number of new P. cattleianum leaves and leaf litter decomposition rates of M. polymorpha by making nutrients more available to plants and microbes rather than through changes in the invertebrate community. Results suggest that E. coqui in Hawaii has the potential to reduce endemic invertebrates and increase nutrient cycling rates, which may confer a competitive advantage to invasive plants in an ecosystem where native species have evolved in nutrient-poor conditions.     

The interactive effects of livestock exclusion and mammalian pest control on the restoration of invertebrate communities in small forest remnants

Abstract

In many agricultural landscapes, significant biodiversity gains can be made by improving the ecological condition of degraded remnants of semi‐natural habitat. Recent emphasis has been on the level of management intervention required to initiate vegetation recovery in small forest remnants, but no comparable emphasis has been placed on benefits for invertebrate communities. In the Waikato region, New Zealand, we tested the effects of livestock exclusion, mammalian pest control, and their interaction, on leaf‐litter invertebrate communities in 30 forest remnants, using a space‐for‐time substitution approach. A total of 87 376 invertebrates were extracted from 964 leaf‐litter samples. Invertebrate density was an order of magnitude lower in remnants than in nearby large forest reserves. For key taxa, such as Diplopoda, Isopoda, Coleoptera and Mollusca, 10‐ to 100‐fold lower densities were recorded in remnants with no pest control, particularly where livestock were not excluded. By contrast, other taxa such as Thysanoptera and For‐micidae (Hymenoptera) had up to 100‐fold greater densities in remnants with recent stock exclusion and pest control. These changes led to a significant livestock exclusion x pest control interaction effect on the degree of invertebrate community dissimilarity between forest remnants and forest reserves. Using structural equation modelling, we found that treatment effects were largely mediated by a cascading series of indirect causal paths involving altered soil chemistry, vegetation composition, and litter mass relative to large forest reserves, although the livestock exclusion × pest control interaction was inadvertently confounded with differing slopes and areas of remnants in different treatments. Livestock exclusion and mammalian pest control have significant, but contrasting, effects on invertebrates in the first 10–20 years following livestock exclusion from forest remnants, with mammalian pest control having limited benefit for the leaf‐litter invertebrate fauna without livestock exclusion.     

Animal-habitat relationships in the Knysna Forest,South Africa: discrimination between forest types by birds and invertebrates

Summary Effects of forest plant species composition and physiognomy on bird and invertebrate communities were investigated in three discrete, relatively undisturbed forest types along a dry-wet soil moisture gradient. Using discriminant function analysis, a 100% floristic and a 78% vegetation structural discrimination were obtained between the three forest types. However, the bird communities of these different forest types were very similar in species composition, and had much lower densities than those normally encountered in other, superficially similar forests. Although an 81% discrimination between forest types was attained through analysis of ground surface invertebrates, measures of litter and aerial invertebrate abundance were also of limited use as discriminators. Historical and biogeographic factors, as well as the low nutritional levels in the soil and vegetation may be the causes of low bird and invertebrate density and diversity. It is concluded that floristics and vegetation structure have, at best, a minor influence on bird community structure, and possibly also on invertebrate community structure in the Knysna Forest.     

Effects of organic matter and season on leaf litter colonisation and breakdown in cave streams

1. Low organic matter availability is thought to be a primary factor influencing evolutionary and ecological processes in cave ecosystems. We examined links among organic matter abundance, macroinvertebrate community structure and breakdown rates of red maple (Acer rubrum) and corn litter (Zea mays) in coarse‐ (10 × 8 mm) and fine‐mesh (500‐μm) litter bags over two seasonal periods in four cave streams in the south‐eastern U.S.A. 2. Organic matter abundance differed among cave streams, averaging from near zero to 850 g ash‐free dry mass m^?2. Each cave system harboured a different macroinvertebrate community. However, trophic structure was similar among caves, with low shredder biomass (2–17% of total biomass). 3. Corn litter breakdown rates (mean k = 0.005 day^?1) were faster than red maple (mean k = 0.003 day^?1). Breakdown rates in coarse‐mesh bags (k = 0.001–0.012 day^?1) were up to three times faster than in fine‐mesh bags (k = 0.001–0.004 day^?1). Neither invertebrate biomass in litter bags nor breakdown rates were correlated with the ambient abundance of organic matter. Litter breakdown rates showed no significant temporal variation. Epigean (surface‐adapted) invertebrates dominated biomass in litter bags, suggesting that their effects on cave ecosystem processes may be greater than hypogean (cave‐adapted) taxa, the traditional focus of cave studies. 4. The functional diversity of our cave communities and litter breakdown rates are comparable to those found in previous litter breakdown studies in cave streams, suggesting that the factors that control organic matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas.     

Invasive litter, not an invasive insectivore, determines invertebrate communities in Hawaiian forests

In Hawaii, invasive plants have the ability to alter litter-based food chains because they often have litter traits that differ from native species. Additionally, abundant invasive predators, especially those representing new trophic levels, can reduce prey. The relative importance of these two processes on the litter invertebrate community in Hawaii is important, because they could affect the large number of endemic and endangered invertebrates. We determined the relative importance of litter resources, represented by leaf litter of two trees, an invasive nitrogen-fixer, Falcataria moluccana, and a native tree, Metrosideros polymorpha, and predation of an invasive terrestrial frog, Eleutherodactylus coqui, on leaf litter invertebrate abundance and composition. Principle component analysis revealed that F. moluccana litter creates an invertebrate community that greatly differs from that found in M. polymorpha litter. We found that F. moluccana increased the abundance of non-native fragmenters (Amphipoda and Isopoda) by 400% and non-native predaceous ants (Hymenoptera: Formicidae) by 200%. E. coqui had less effect on the litter invertebrate community; it reduced microbivores by 40% in F. moluccana and non-native ants by 30% across litter types. E. coqui stomach contents were similar in abundance and composition in both litter treatments, despite dramatic differences in the invertebrate community. Additionally, our results suggest that invertebrate community differences between litter types did not cascade to influence E. coqui growth or survivorship. In conclusion, it appears that an invasive nitrogen-fixing tree species has a greater influence on litter invertebrate community abundance and composition than the invasive predator, E. coqui.     

Role of arthropod communities in bioenergy crop litter decomposition†

The extensive land use conversion expected to occur to meet demands for bioenergy feedstock production will likely have widespread impacts on agroecosystem biodiversity and ecosystem services, including carbon sequestration. Although arthropod detritivores are known to contribute to litter decomposition and thus energy flow and nutrient cycling in many plant communities, their importance in bioenergy feedstock communities has not yet been assessed. We undertook an experimental study quantifying rates of litter mass loss and nutrient cycling in the presence and absence of these organisms in three bioenergy feedstock crops—miscanthus (Miscanthus x giganteus), switchgrass (Panicum virgatum), and a planted prairie community. Overall arthropod abundance and litter decomposition rates were similar in all three communities. Despite effective reduction of arthropods in experimental plots via insecticide application, litter decomposition rates, inorganic nitrogen leaching, and carbon–nitrogen ratios did not differ significantly between control (with arthropods) and treatment (without arthropods) plots in any of the three community types. Our findings suggest that changes in arthropod faunal composition associated with widespread adoption of bioenergy feedstock crops may not be associated with profoundly altered arthropod‐mediated litter decomposition and nutrient release.     

Size distribution of aquatic invertebrates in two prairie wetlands, with and without fish, with implications for community production

1. We compared the size distribution of aquatic invertebrates in two prairie wetlands, one supporting a population of fathead minnows and the other fishless. Both wetlands were sampled in three depth zones on three dates, allowing assessment of temporal and spatial variation.
2. We determined biomass of aquatic invertebrates in 17 log₂ size classes, and used these data to develop normalized size spectra. We also coupled size distributions with an allometric model to estimate relative production at the community level.
3. The composition of the invertebrate communities differed greatly between sites, and invertebrate biomass was higher in nearly all size classes in the fishless wetland. Intercepts of normalized size spectra were significantly different between wetlands, but slopes generally were not, indicating differences in standing-stock biomass but similar size structures between the two invertebrate communities. Higher standing-stock biomass in the fishless wetland resulted in higher relative production per unit area, but similar size distributions resulted in similar mass-specific production (P/B) between wetlands.
4. Our results indicate that invertebrate communities in prairie wetlands may have relatively consistent size structures in spite of large differences in community composition and standing-stock biomass. We hypothesize that the observed differences are because of predation by the minnow population and/or differences in the macrophyte communities between the two sites. However, the relative importance of macrophytes and fish predation in structuring invertebrate communities in prairie wetlands is poorly known.     

How do patch quality and spatial context affect invertebrate communities in a natural moss microlandscape?

Globally, moss associated invertebrates remain poorly studied and it is largely unknown to what extent their diversity is driven by local environmental conditions or the landscape context. Here, we investigated small scale drivers of invertebrate communities in a moss landscape in a temperate forest in Western Europe. By comparing replicate quadrats of 5 different moss species in a continuous moss landscape, we found that mosses differed in invertebrate density and community composition. Although, in general, richness was similar among moss species, some invertebrate taxa were significantly linked to certain moss species. Only moss biomass and not relative moisture content could explain differences in invertebrate densities among moss species. Second, we focused on invertebrate communities associated with the locally common moss species Kindbergia praelonga in isolated moss patches on dead tree trunks to look at effects of patch size, quality, heterogeneity and connectivity on invertebrate communities. Invertebrate richness was higher in patches under closed canopies than under more open canopies, presumably due to the higher input of leaf litter and/or lower evaporation. In addition, increased numbers of other moss species in the same patch seemed to promote invertebrate richness in K. praelonga, possibly due to mass effects. Since invertebrate richness was unaffected by patch size and isolation, dispersal was probably not limiting in this system with patches separated by tens of meters, or stochastic extinctions may be uncommon. Overall, we conclude that invertebrate composition in moss patches may not only depend on local patch conditions, in a particular moss species, but also on the presence of other moss species in the direct vicinity.     

Coqui frog invasions change invertebrate communities in Hawaii

The Puerto Rican coqui frog (Eleutherodactylus coqui) invaded Hawaii in the late 1980s. Because the coqui reaches high densities and consumes large quantities of invertebrates, it was hypothesized to change invertebrate communities where it invades. Previous research found that coquis can change invertebrate communities, but these studies used highly manipulative, small-scale experiments. The objective of this research was to determine whether coquis create community-level changes in invertebrate communities at the landscape scale. We collected leaf litter, flying, and foliage invertebrates on both sides of 15 coqui invasion fronts across the island of Hawaii. Multivariate analyses show that coquis are associated with changes in leaf-litter communities, primarily reductions in Acari, but are not associated with overall changes in flying or foliage communities. Across sites, coquis reduced the total number of leaf-litter invertebrates by 27%, specifically by reducing Acari by 36%. Across sites, coquis increased flying Diptera by 19%. Changes were greater where coqui densities were higher. We suggest that coquis changed leaf-litter communities primarily through direct predation, but that they increased Diptera through the addition of frog carcasses and excrement. Results support previous studies conducted in more controlled settings, but add to our understanding of the invasion by showing that coqui effects on invertebrate communities are measurable at the landscape scale.     

Cave invertebrate assemblages differ between native and exotic leaf litter

Abstract Allochtonous leaf litter is an important source of energy and nutrients for invertebrates in cave ecosystems. A change to the quality or quantity of litter entering caves has the potential to disrupt the structure and function of cave communities. In this study, we adopted an experimental approach to examine rates of leaf litter decomposition and the invertebrate assemblages colonizing native and exotic leaf litter in limestone caves in the Jenolan Caves Karst Conservation Reserve, New South Wales, Australia. We deployed traps containing leaf litter from exotic sycamore (Acer pseudoplatanus) and radiata pine (Pinus radiata) trees and native eucalypts (Eucalyptus spp.) in twilight zones (near the cave entrance) and areas deep within the caves for 3 months. Thirty‐two invertebrate morphospecies were recorded from the litter traps, with greater richness and abundance evident in the samples from the twilight zone compared with areas deep within the cave. Sycamore litter had significantly greater richness and abundance of invertebrates compared with eucalypt and pine litter in samples from the twilight zone, but there was no difference in richness or abundance among litter samples placed deep within the cave. Relative rates of decay of the three litters were sycamore > eucalypt > pine. We discuss the potential for the higher decomposition rates and specific leaf area in sycamores to explain their higher invertebrate diversity and abundance. Our findings have important implications for the management of exotic plants and the contribution of their leaf litter to subterranean ecosystems.     

Arthropod Abundance and Diversity in a Lowland Tropical Forest Floor in Panama: The Role of Habitat Space vs. Nutrient Concentrations

Tropical forest floor characteristics such as depth and nutrient concentrations are highly heterogeneous even over small spatial scales and it is unclear how these differences contribute to patchiness in forest floor arthropod abundance and diversity. In a lowland tropical forest in Panama we experimentally increased litter standing crop by removing litter from five plots (L−) and adding it to five other plots (L+); we had five control plots. After 32 mo of treatments we investigated how arthropod abundance and diversity were related to differences in forest floor physical (mass, depth, water content) and chemical properties (pH, nutrient concentrations). Forest floor mass and total arthropod abundance were greater in L+ plots compared with controls. There were no treatment differences in nutrient concentrations, pH or water content of the organic horizons. Over all plots, the mass of the fermentation horizon (Oe) was greater than the litter horizon (Oi); arthropod diversity and biomass were also greater in the Oe horizon but nutrient concentrations tended to be higher in the Oi horizon. Arthropod abundance was best explained by forest floor mass, while arthropod diversity was best explained by phosphorus, calcium and sodium concentrations in the Oi horizon and by phosphorus concentrations in the Oe horizon. Differences in arthropod community composition between treatments and horizons correlated with phosphorus concentration and dry mass of the forest floor. We conclude that at a local scale, arthropod abundance is related to forest floor mass (habitat space), while arthropod diversity is related to forest floor nutrient concentrations (habitat quality).     

粤东地区速生桉树林与天然林枯枝落叶层无脊椎动物多样性比较

研究比较了粤东地区天然林和速生桉树林枯枝落叶层无脊椎动物的群落结构以及菌食性蓟马的物种多样性.结果表明,螨类、弹尾目、双翅目幼虫、膜翅目、缨翅目和鞘翅目构成了该地区天然林枯枝落叶层无脊椎动物群落的主要成分,它们的个体数占天然林枯枝落叶层无脊椎动物总个体数的96.5%;而螨类、弹尾目、双翅目幼虫和鳞翅目幼虫则是速生桉树林枯枝落叶层无脊椎动物群落的主要成分,它们的个体数占全部个体数的96.3%.天然林的多样性指数H′、均匀性指数J、DG指数均比速生桉树林高,Simpson优势度指数D比桉树林小,菌食性蓟马种类和个体数也显著多于桉树林,说明速生桉树林凋落物无脊椎动物多样性不及天然林丰富,但螨类数量巨大,占桉树林无脊椎动物总数的77.6%,以致两种林分凋落物无脊椎动物的平均密度差异不显著.表明保留桉树人工林的林下植被和凋落物对提高速生桉树林枯枝落叶层无脊椎动物的群落多样性具有重要意义.     

Highly reduced mass loss rates and increased litter layer in radioactively contaminated areas

The effects of radioactive contamination from Chernobyl on decomposition of plant material still remain unknown. We predicted that decomposition rate would be reduced in the most contaminated sites due to an absence or reduced densities of soil invertebrates. If microorganisms were the main agents responsible for decomposition, exclusion of large soil invertebrates should not affect decomposition. In September 2007 we deposited 572 bags with uncontaminated dry leaf litter from four species of trees in the leaf litter layer at 20 forest sites around Chernobyl that varied in background radiation by more than a factor 2,600. Approximately one quarter of these bags were made of a fine mesh that prevented access to litter by soil invertebrates. These bags were retrieved in June 2008, dried and weighed to estimate litter mass loss. Litter mass loss was 40 % lower in the most contaminated sites relative to sites with a normal background radiation level for Ukraine. Similar reductions in litter mass loss were estimated for individual litter bags, litter bags at different sites, and differences between litter bags at pairs of neighboring sites differing in level of radioactive contamination. Litter mass loss was slightly greater in the presence of large soil invertebrates than in their absence. The thickness of the forest floor increased with the level of radiation and decreased with proportional loss of mass from all litter bags. These findings suggest that radioactive contamination has reduced the rate of litter mass loss, increased accumulation of litter, and affected growth conditions for plants.     

Plant diversity does not buffer drought effects on early‐stage litter mass loss rates and microbial properties

Human activities are decreasing biodiversity and changing the climate worldwide. Both global change drivers have been shown to affect ecosystem functioning, but they may also act in concert in a non‐additive way. We studied early‐stage litter mass loss rates and soil microbial properties (basal respiration and microbial biomass) during the summer season in response to plant species richness and summer drought in a large grassland biodiversity experiment, the Jena Experiment, Germany. In line with our expectations, decreasing plant diversity and summer drought decreased litter mass loss rates and soil microbial properties. In contrast to our hypotheses, however, this was only true for mass loss of standard litter (wheat straw) used in all plots, and not for plant community‐specific litter mass loss. We found no interactive effects between global change drivers, that is, drought reduced litter mass loss rates and soil microbial properties irrespective of plant diversity. High mass loss rates of plant community‐specific litter and low responsiveness to drought relative to the standard litter indicate that soil microbial communities were adapted to decomposing community‐specific plant litter material including lower susceptibility to dry conditions during summer months. Moreover, higher microbial enzymatic diversity at high plant diversity may have caused elevated mass loss of standard litter. Our results indicate that plant diversity loss and summer drought independently impede soil processes. However, soil decomposer communities may be highly adapted to decomposing plant community‐specific litter material, even in situations of environmental stress. Results of standard litter mass loss moreover suggest that decomposer communities under diverse plant communities are able to cope with a greater variety of plant inputs possibly making them less responsive to biotic changes.     

More food,less habitat: how necromass and leaf litter decomposition combine to regulate a litter ant community

1. In brown food webs of the forest floor, necromass (e.g. insect carcasses and frass) falling from the canopy feeds both microbes and ants, with the former decomposing the homes of the latter. In a tropical litter ant community, we added necromass to 1 m² plots, testing if it added as a net food (increasing ant colony growth and recruitment) or destroyer of habitat (by decomposing leaf litter). 2. Maximum, but not mean, colony growth rates were higher on +food plots. However, neither average colony size, nor density was higher on +food plots. In contrast, +food plots saw diminished availability of leaf litter and higher microbial decomposition of cellulose, a main component of the organic substrate that comprises litter habitat. 3. Furthermore, necromass acted as a limiting resource to the ant community only when nest sites were supplemented on +food plots in a second experiment. Many of these +food +nest plots were colonised by the weedy species Wasmannia auropunctata. 4. Combined, these results support the more food–less habitat hypothesis and highlight the importance of embedding studies of litter ant ecology within broader decomposer food web dynamics.     

Effects of litter addition on ectomycorrhizal associates of a lodgepole pine (Pinus contorta) stand in Yellowstone National Park

Increasing soil nutrients through litter manipulation, pollution, or fertilization can adversely affect ectomycorrhizal (EM) communities by inhibiting fungal growth. In this study, we used molecular genetic methods to determine the effects of litter addition on the EM community of a Pinus contorta stand in Yellowstone National Park that regenerated after a stand-replacing fire. Two controls were used; in unmodified control plots nothing was added to the soil, and in perlite plots perlite, a chemically neutral substance, was added to maintain soil moisture and temperature at levels similar to those under litter. We found that (i) species richness did not change significantly following perlite addition (2.6 +/- 0.3 species/core in control plots, compared with 2.3 +/- 0.3 species/core in perlite plots) but decreased significantly (P < 0.05) following litter addition (1.8 +/- 0.3 species/core); (ii) EM infection was not affected by the addition of perlite but increased significantly (P < 0.001) in response to litter addition, and the increase occurred only in the upper soil layer, directly adjacent to the added litter; and (iii) Suillus granulatus, Wilcoxina mikolae, and agaricoid DD were the dominant organisms in controls, but the levels of W. mikolae and agaricoid DD decreased significantly in response to both perlite and litter addition. The relative levels of S. granulatus and a fourth fungus, Cortinariaceae species 2, increased significantly (P < 0.01 and P < 0.05, respectively) following litter addition. Thus, litter addition resulted in some negative effects that may be attributable to moisture-temperature relationships rather than to the increased nutrients associated with litter. Some species respond positively to litter addition, indicating that there are differences in their physiologies. Hence, changes in the EM community induced by litter accumulation also may affect ecosystem function.     

Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.     

Role of transported particulate organic matter in the macroinvertebrate colonization of litter bags in streams

1. The exposure of mesh litter bags has been widely used to investigate the role of benthic macroinvertebrates in leaf processing in freshwaters. In this sense, several studies have related litter bag breakdown rates to the presence of colonizing invertebrates. A possible confounding factor in such experiments is that the litter bags trap suspended particulate organic matter that can itself attract invertebrate colonists, irrespective of the intended experimental treatment.
2. We attempted to quantify the accumulation of particulate organic matter (POM) within litter bags and to investigate its possible impact on macroinvertebrate density and richness. In seven headwater forested streams we exposed mesh bags filled either with beech leaves ( Fagus sylvatica ) or with plastic strips of an equal surface area.
3. Principal component analysis (PCA) showed that bag type and stream were the main explanatory variables for invertebrate colonization and POM accumulation within the bags. In contrast, there was little variation among sampling dates (6.4% of the total inertia).
4. The accumulated POM within the bags was substantial (up to 8.83 g ash-free dry mass (AFDM)) but highly variable among sites (mean from 0.32 to 4.58 g AFDM). At each of the seven sites, both richness and abundance of invertebrates were positively correlated with the mass of accumulated POM in bags. Macroinvertebrate colonization (notably taxon richness) was directly linked with the quantity of POM accumulated.
5. Our findings provide evidence of a potential pitfall in linking invertebrates to litter processing in mesh bags, particularly when large amounts of POM, entrained in stream flow, accumulate within the bags. An evaluation of the POM mass trapped in litter bags could account for the erratic patterns sometimes observed in their colonization by invertebrates.     

The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.