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.     

Degradation of native and exotic riparian plant leaf litter in a floodplain pond

1. A litter‐bag experiment was undertaken in a pond on the margins of a large temperate floodplain in south‐western France to assess the potential influence of the replacement of native by exotic riparian species on organic matter degradation. We determined initial litter chemical composition, breakdown rates and the invertebrate assemblages associated with the litter for five pairs of native dominant and exotic invasive species co‐occurring at different stages along a successional gradient. 2. Litter chemical composition, breakdown rates and abundance and diversity of detritivorous invertebrates were similar for the exotic and native species overall. No overall changes in organic matter degradation can thus be predicted from the replacement of dominant natives by exotic invasives. Breakdown rates were primarily driven by the C/N ratio. 3. One invasive species (Buddleja davidii) showed significantly higher breakdown rates than its native counterpart (Populus nigra), resulting in the disappearance of leaf litter 6 months prior to the next litterfall. In some cases, therefore, invasion by exotic species may result in discontinuity of resource supply for decomposers.     

Effect of native and exotic leaf litter on macroinvertebrate communities and decomposition in a western Montana stream

Invasion by exotic trees into riparian areas has the potential to impact terrestrial and aquatic systems. To test the effect of different streamside tree species on the aquatic food web in a stream in Montana, we compared decomposition rates of leaf litter and invertebrate assemblages associated with the leaf litter of the exotic Acer platanoides and the dominant native Populus trichocarpa trees. Macroinvertebrate family richness, evenness, and diversity increased with days of aquatic processing; however, there was no effect of leaf species. Leaves of the A. platanoides were associated with 70% greater density of macroinvertebrates of the family Nemouridae. This family consists primarily of detritivores and had the greatest overall density and frequency of occurrence relative to other macroinvertebrate families. The density of a family of predatory macroinvertebrates (Rhyacophilidae) was also generally (73%) greater in association with A. platanoides than P. trichocarpa leaves. The density of Ephemerellidae and Rhyacophilidae increased over time. In contrast to studies comparing leaves of exotic vs. native trees, we observed no difference in leaf decomposition rates; however, the amount of leaf inputs are likely to differ between native and invaded forests. The results indicate that replacement of native riparian trees with exotics affected the most common family of macroinvertebrates and possibly a common family of predatory macroinvertebrates (Rhyacophilidae), which may affect the detrital food web.     

Convergence in leaf phenology traits of two understory ferns in the northwestern Iberian Peninsula

Aims Plants control leaf phenology to maximize annual photosynthetic product. Although ferns play an important ecological role in many habitats, especially forests, their phenology traits have been poorly studied. Here, we investigate the leaf phenology of two ferns of the forest understorey and analyse the relationship between the timing of leaf emergence and spore dispersal and the effect of between-year climatic variation.Methods We compared the leafing and sporing phenologies of two ferns with very large (>2 m), overwintering leaves: Culcita macrocarpa and Woodwardia radicans. We regularly monitored individuals of six populations in the northwestern Iberian Peninsula during a 3-year study. We studied eight phenology variables: leafing start date, leafing end date, leaf expansion time, number of new leaves per individual, between-individual synchrony, within-individual synchrony, percentage of fertile leaves and spore release date. We also determined leaf mass per area (LMA) and gathered data on air temperature and humidity.Important findings Both C. macrocarpa and W. radicans produce few leaves (~2 leaves individual^-1 year^-1), which expand simultaneously for a very long period (from midwinter to early summer), are retained for more than 1 year (37 and 19 months, respectively) and have relatively high LMAs. Such traits, together with large leaf size, have also been found in seed plants from the forest understorey and represent adaptations to this light-limited environment. Spores of both study ferns are simultaneously released in late winter, with little between-year variation caused by differences in air humidity. This remarkable similarity between species suggests that the convergence in timing of leaf emergence favours the convergence in timing of spore dispersal.     

华南4种乡土阔叶树种枯落叶分解能力

采用分解袋法,研究了樟树(Cinnamomum camphora (L.) Presl)、米老排(Mytilaria laosensis Lec.)、火力楠(Michelia macclurei Dandy)和黎蒴(Castanopsis fissa Rehd. et Wils.) 4种华南乡土阔叶树种枯落叶在阔叶混交人工林和湿地松人工林下的分解能力.4个树种枯落叶的分解速率大小依次为:樟树>米老排>火力楠>黎蒴,它们在混交林中的分解速率均大于松林.4个树种枯落叶的分解动态符合Olson指数模型,分解系数(K)在混交林中均大于人工林,且在两种林地中均表现为樟树＞米老排＞火力楠＞黎蒴.不同时间枯落叶养分动态分析结果显示,4个树种枯落叶的N元素分解均出现先富集,后释放的特点,但仅在松林下出现N元素的净释放;K元素释放方式表现为先释放,后富集,仅在阔叶林下出现K元素净释放;Ca、Mg和B元素均表现为分解前期少量富积,之后开始大量释放的变化趋势,它们在阔叶林和松林下均出现净释放.4个树种在P元素释放方式上差异较大,趋势不明显.     

Reciprocal effects of litter from exotic and congeneric native plant species via soil nutrients

Invasive exotic plant species are often expected to benefit exclusively from legacy effects of their litter inputs on soil processes and nutrient availability. However, there are relatively few experimental tests determining how litter of exotic plants affects their own growth conditions compared to congeneric native plant species. Here, we test how the legacy of litter from three exotic plant species affects their own performance in comparison to their congeneric natives that co-occur in the invaded habitat. We also analyzed litter effects on soil processes. In all three comparisons, soil with litter from exotic plant species had the highest respiration rates. In two out of the three exotic-native species comparisons, soil with litter from exotic plant species had higher inorganic nitrogen concentrations than their native congener, which was likely due to higher initial litter quality of the exotics. When litter from an exotic plant species had a positive effect on itself, it also had a positive effect on its native congener. We conclude that exotic plant species develop a legacy effect in soil from the invaded range through their litter inputs. This litter legacy effect results in altered soil processes that can promote both the exotic plant species and their native congener.     

Effects of leaf litter species on macroinvertebrate community properties and mosquito yield in Neotropical tree hole microcosms

Detritus quality and quantity affect macroinvertebrate productivity and distribution in many freshwater ecosystems. This study experimentally investigated the effects of leaf litter from Ceiba pentandra, Dipteryx panamensis, Ficus yoponensis, and Platypodium elegans on macroinvertebrate species composition, richness, and abundance in artificial water-filled tree holes in a lowland moist forest of Panama. Species composition was similar among treatments, but species richness and longevity differed among litter types and were consistently highest with Platypodium litter. Similar patterns were observed in natural tree holes of the focal tree species. The mosquito Culex mollis was the most abundant species in the field experiment. Average conductivity and dissolved oxygen concentration differed among leaf species, but pH did not. Leaf toughness was positively correlated with mean macroinvertebrate abundance and cumulative species richness. A laboratory experiment measured C. mollis yield and pupation time in tree hole microcosms containing the four litter species. Cumulative mosquito mass and time to pupation differed among leaf litter species, with Platypodium litter supporting the greatest yield. Pupation was slowest on Ceiba litter. Grazing by mosquito larvae facilitated leaf decomposition in all treatments. Results suggest that differences in macroinvertebrate species richness and mosquito yield can be attributed to differences in nutritional quality among litter species. Received: 14 October 1998 / Accepted: 21 February 1999     

泰山4种优势造林树种叶片凋落物分解对凋落物内细菌群落结构的影响

为研究泰山不同造林树种凋落物叶分解对细菌群落的影响。以泰山4种主要优势造林树种刺槐(Robinia pseucdoacacia)、麻栎(Quercus acutissima)、油松(Pinus tabulaeformis)和赤松(Pinus densiflora)为研究对象,采用凋落物分解袋法及Illumina Miseq测序平台对细菌16S rDNA V4—V5区扩增产物进行双端测序,分析了4种树种叶片凋落物分解对细菌群落结构及多样性的影响。结果表明:(1)4种树种叶片分解速率差异显著(P0.05),刺槐分解速率显著高于其他3个树种(P0.05),表现为刺槐赤松油松麻栎。(2)4种叶凋落物分解一年后化学元素含量与初始化学元素相比均存在显著差异。C、木质素含量均显著降低(P0.05);N、P含量显著升高(P0.05)。(3)所有样品一共获得643440条有效序列,分属于35门,92纲,121目,246科,410属,206种。细菌群落NMDSβ-多样性分析显示除油松和赤松间差异较小外,其他树种间差异程度均较大。其中,细菌群落相对丰度在5%以上的优势类群是变形菌门、放线菌门、拟杆菌门、酸杆菌门,且在4种处理之间差异显著(PSymbol|@@0.05)。在纲水平上,α-变形菌纲、β-变形菌纲、不明放线菌纲、鞘脂杆菌纲、γ-变形菌纲、δ-变形菌纲为主要的优势纲,其中不明放线菌纲和鞘脂杆菌纲差异显著(PSymbol|@@0.05)。在种水平上,Bradyrhizobium elkanii和Luteibacter rhizovicinus在4个处理中都为优势种,每个处理也都有自己所特有的优势种。(4)4个处理细菌丰富度(OUT、观测到的物种数和ACE指数)和系统发育多样性(PD指数)之间差异显著(PSymbol|@@0.05),且阔叶树种刺槐和麻栎显著高于针叶树种赤松和油松。(5)叶片凋落物性状和细菌群落NMDS分析表明,细菌群落多样性受到凋落物化学性质的影响,尤其是凋落物初始C/N比和木质素/N比。此外,在细菌群落多样性和叶片凋落物化学性质两个因素中,分解速率与凋落物化学性质相关性更大。研究结果有助于理解细菌群落结构和多样性对森林生态系统叶片凋落物分解的影响。     

Leaf litter age regulates the effect of native and exotic tree species on understory herbaceous vegetation of riparian forests

Decomposing litter is regarded as the most important source of allelochemicals released into soil. In this study, a greenhouse experiment was designed to assess the net effect of differently aged leaf litter from exotic (Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila) and native riverine trees (Populus alba, Populus nigra and Ulmus minor) on the germination and growth of three herb species (Trifolium repens, Dactylis glomerata and Chenopodium album). We also characterized the chemical composition of litter samples at different litter ages (0, 1, 2 and 3 months) based on phenolic compounds, fibers and ergosterol (as a measure of fungal biomass) contents. Overall, litter from both native and exotic species had a negative effect on shoot and root growth of target species, indicating that phytotoxic effects of litter predominate over positive effects. The inhibition effect of the exotic species was similar or even lower than that of the natives, which does not support the Novel Weapons Hypothesis. Among exotic trees, U. pumila showed the highest inhibition effect on the growth of the target species. T. repens was the most sensitive target species. The importance of litter age varied with both target and donor species. In general, D. glomerata was more inhibited by fresh litter, C. album by half-decomposed litter of U. pumila and R. pseudoacacia and by fresh litter of A. altissima and T. repens was more inhibited by fresh litter of A. altissima and P. alba and by highly decomposed litter of U. minor. The concentration of total phenolics and flavonoids decreased while acid detergent fiber, lignin and ergosterol increased with increasing litter age. Hydroxybenzoic and protocatechuic acids and the flavonoid quercetin were detected in all litter species and at most of the litter ages, while gallic, chlorogenic, vanillic, coumaric and rosmarinic acids were species-specific and they were only detected in fresh litter. Ergosterol concentration appeared as the strongest constrictor of inhibitory effects of litter on understory species. The results of this study contribute to the understanding of the net effect of fresh and decomposed litter from exotic and native trees on the growth of understory species.     

森林演替过程中优势树种凋落叶对土壤微生物组成的影响

为了进一步认识森林群落演替机理,研究了黄土高原马栏林区处于不同演替阶段的森林土壤微生物组成,以及优势树种凋落叶对土壤微生物组成的影响.采用凋落叶浸提液处理不同类型的土壤,利用土壤浸提液固体培养基测定微生物数量,以系统聚类和主成分分析等方法进行数据分析.结果表明:微生物总数量及土壤细菌数量按白桦(Betulaplatyphylla)林→油松(Pinus tabulaeformis)林→辽东栎(Quercus liaotungensis)林递增;在先锋森林群落向顶极群落演替过程中,微生物组成的综合性指标Shannon-Wiener指数呈下降趋势;土壤微生物3大类群中,数量较少者对Shannon-Wiener指数的大小变化起着较大作用;依据可培养微生物组成,8种土样可划分为3个类群,辽东栎及油松凋落叶对土壤微生物组成的影响不同,辽东栎凋落叶对土壤微生物学性质的作用更强;马栏林区森林土壤微生物组成与植被类型及演替阶段关系密切;演替高级阶段森林优势树种凋落叶可能通过其对土壤微生物组成的改造优势,逐渐改变土壤的微生物学性质,进而促进植被演替.     

The production of hybrids with high ecological amplitude between exotic Spartina densiflora and native S. maritima in the Iberian Peninsula

Aim Hybridization is a common and potent mechanism of plant evolution that has the potential to be evolutionary significant in its own right, and hybrids are common between invasive and native congeneric species. Our aims were to document the existence and nature of new Spartina hybrids arising between introduced Spartina densiflora and native S. maritima in tidal marshes of the Iberian Peninsula, to examine the actual and potential range of hybrids in the intertidal zone, and to analyse the seed set of hybrids. Location South‐West Iberian Peninsula. Methods Hybrids were characterized using chloroplast and nuclear DNA, and ploidy assessments. The ecological tolerance of the hybrids was studied using vegetation surveys and transplant experiments. Results We found that both parental species have been seed parents to hybrids and that all hybrids had an additive pattern of species‐specific nuclear markers consistent with F₁ hybrids. Hybrid chromosome numbers varied between 2n = ca. 65 and 2n = ca. 94–97, while S. maritima had 2n = ca. 60 and S. densiflora had 2n = ca. 70. Hybrids grew in three discrete locations along the intertidal zone but were capable of growing throughout the ranges of both parental species in transplanted gardens, and in most cases, grew better than the parental species. While the potential exists for the origination of another invasive Spartina hybrid species, thus far hybrid plants are not fertile, limiting their ability to invade and spread. Main conclusions We recommend the eradication of all of the currently quite limited hybrid cordgrass and to fight the S. densiflora invasion in the Iberian Peninsula to prevent the origination of a new invasive allopolyploid Spartina species.     

Influence of tree species composition on soil and soil solution properties in two mixed spruce-beech stands with contrasting history in Southern Germany

The properties of the soil and soil solution of two mixed spruce-beech forests in Southern Germany were investigated in order to identify non-additive effects of tree species compared to the monocultures. At five subplots in each of the two monocultures and at 10 subplots in each of the two mixed stands humus morphology, topsoil acidity as well as nitrate and sulphate concentrations in seepage water below the rooting zone were measured. At the Höglwald site, an 8×8 m sampling grid was also installed. Tree species composition within a 10 m circle surrounding each sampling point was correlated with the soil properties and the soil solution properties using the method of breakpoint estimation as well as a linear and a cubic model. At both sites, thickness and acidity of the forest floor as well as sulphate and nitrate concentrations in seepage water were significantly higher in the spruce monocultures than in the beech monocultures. Non-linear patterns in the correlation between both the thickness and the acidity of the forest floor and tree species composition occurred at both sites. In addition, nitrate concentration in the seepage water showed a non-linear correlation pattern at the Höglwald site. The correlation patterns were site specific and depended on stand history. While the influence of spruce at the Höglwald site was greater than expected from spruce monocultures, the opposite was found at the Schongau site. Interaction effects on nitrate concentration were absent at the Schongau site. Interaction effects on sulphate concentrations were absent at both sites. Current knowledge about the complex processes and patterns in mixed species stands is still limited. At least for certain properties, mixed species stands cannot be treated as a summation of the corresponding monocultures.     

Living in the litter: the influence of tree leaf litter on wetland communities

Empirical research in streams has demonstrated that terrestrial subsidies of tree leaf litter influence multiple community factors including composition, diversity and growth of individuals. However, little research has examined the importance of tree litter species on wetlands, which are ubiquitous across the landscape and serve as important habitats for a unique and diverse community of organisms. Using outdoor mesocosms, we assessed the impact of 12 litter monocultures and three litter mixtures (from both broadleaf and conifer trees) on pond communities containing gray tree frog tadpoles Hyla versicolor, periphyton, phytoplankton and zooplankton. We found that leaf litter species had substantial and differential impacts on all trophic groups in the community including effects on algal abundance, zooplankton density and amphibian growth. In many instances, patterns of responses were specific to individual litter species yet some responses, including both pH values and periphyton biomass, were generalizable to broad taxonomic groups. In addition, while most responses of litter mixtures were additive, we found evidence for antagonistic effects of litter mixing among responses of periphyton and amphibian body mass. Our results highlight the potential impact of human and naturally driven changes in forest composition on wetland communities through associated changes in leaf litter.     

Partitioning the effect of composition and diversity of tree communities on leaf litter decomposition and soil respiration

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance‐weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.     

不同化学性质叶凋落物添加对土壤有机碳矿化及激发效应的影响

凋落物输入可显著影响土壤有机碳(SOC)矿化速率,但添加不同化学性质叶凋落物对土壤有机碳矿化释放CO₂及激发效应的影响及其机理仍不清楚。本研究将亚热带6种树种¹³C标记的叶凋落物添加至天然次生林0～10 cm原位土柱中,比较不同树种叶凋落物添加对土壤总CO₂、外源凋落物和土壤来源CO₂释放速率和累积量以及激发效应的影响,并量化叶凋落物化学性质与土壤CO₂释放累积量、激发效应的相关关系。结果表明: 添加叶凋落物能够显著提高土壤总CO₂和土壤来源CO₂释放量,存在显著正激发效应,激发效应值为68%～128%。不同树种叶凋落物添加对土壤有机碳矿化和激发效应的影响存在显著差异。Pearson相关分析和逐步多元线性回归分析发现,凋落物来源CO₂释放累积量与叶凋落物C、P和纤维素含量呈显著负相关,而土壤来源CO₂释放量与叶凋落物C:N和木质素:N呈显著正相关。综上,不同化学性质的叶凋落物对土壤有机碳矿化和激发效应的影响存在异质性,在亚热带地区森林类型转变过程中营造具有高质量叶凋落物的人工林将有助于减少森林土壤碳损失。     

Effect of a native tree on seedling establishment of two exotic invasive species in a semiarid ecosystem

Theory predicts that in more stressful environments, positive plant-plant interactions should be more important than negative ones. For instance, in arid and semiarid regions, amelioration of soil drought produced by the shade of established plants could facilitate establishment of other species, in spite of light reduction. However, this theory has not been tested widely in the context of plant invasion. In this paper we evaluated the hypothesis that in a semiarid ecosystem of central Chile, the native tree, Lithrea caustica, should facilitate through positive shading effects, the seedling establishment of two widely planted and invasive forestry species, Pinus radiata and Eucalyptus globulus. We assessed the seedling establishment examining two processes: seedling recruitment (including germination) and subsequent seedling survival. We sowed seeds (to assess recruitment) and planted 8 months old seedlings (to assess seedling survival) of each exotic species under Lithrea patches, open sites and under an artificial shade mimicking Lithrea shading. The study was repeated in a north-facing and a south-facing slope in the study area located in a xeric zone within the distribution range of plantations of these species in central Chile. Our results show that in a north-facing slope Lithrea had positive effects on recruitment of both species, which was produced by shading. These effects were counteracted by negative effects on seedling survival but through a different mechanism, which suggests that Lithrea would have no significant effect on the whole seedling establishment process of Pinus radiata nor Eucalyptus globulus in this habitat. In turn, in a south-facing slope Lithrea had no significant effect on recruitment but had a negative effect on seedling survival, which was not produced by shading. This suggests that in this habitat Lithrea has a negative effect on the seedling establishment of these exotic species. Our results suggest that the effect of the native Lithrea caustica on the seedling establishment of these exotic species is dependent upon the life-cycle phase (recruitment or seedling survival) and habitat even within the same semiarid ecosystem. In contrast to the expected positive effects Lithrea is unlikely to facilitate seedling establishment of these exotic species in this area, and in fact in some habitats this effect could be negative. However, our results also suggest that a common mechanism proposed to resist invasion in forest ecosystems such as shading, probably is not sufficient to inhibit invasion in a semiarid region.     

Stream carbon and nitrogen supplements during leaf litter decomposition: contrasting patterns for two foundation species

Leaf litter decomposition plays a major role in nutrient dynamics in forested streams. The chemical composition of litter affects its processing by microorganisms, which obtain nutrients from litter and from the water column. The balance of these fluxes is not well known, because they occur simultaneously and thus are difficult to quantify separately. Here, we examined C and N flow from streamwater and leaf litter to microbial biofilms during decomposition. We used isotopically enriched leaves (¹³C and ¹⁵N) from two riparian foundation tree species: fast-decomposing Populus fremontii and slow-decomposing Populus angustifolia, which differed in their concentration of recalcitrant compounds. We adapted the isotope pool dilution method to estimate gross elemental fluxes into litter microbes. Three key findings emerged: litter type strongly affected biomass and stoichiometry of microbial assemblages growing on litter; the proportion of C and N in microorganisms derived from the streamwater, as opposed to the litter, did not differ between litter types, but increased throughout decomposition; gross immobilization of N from the streamwater was higher for P. fremontii compared to P. angustifolia, probably as a consequence of the higher microbial biomass on P. fremontii. In contrast, gross immobilization of C from the streamwater was higher for P. angustifolia, suggesting that dissolved organic C in streamwater was used as an additional energy source by microbial assemblages growing on slow-decomposing litter. These results indicate that biofilms on decomposing litter have specific element requirements driven by litter characteristics, which might have implications for whole-stream nutrient retention.