Insect biological control accelerates leaf litter decomposition and alters short‐term nutrient dynamics in a Tamarix‐invaded riparian ecosystem

Insect herbivory can strongly influence ecosystem nutrient dynamics, yet the indirect effects of herbivore‐altered litter quality on subsequent decomposition remain poorly understood. The northern tamarisk beetle Diorhabda carinulata was released across several western states as a biological control agent to reduce the extent of the invasive tree Tamarix spp. in highly‐valued riparian ecosystems; however, very little is currently known about the effects of this biocontrol effort on ecosystem nutrient cycling. In this study, we examined alterations to nutrient dynamics resulting from beetle herbivory in a Tamarix‐invaded riparian ecosystem in the Great Basin Desert in northern Nevada, USA, by measuring changes in litter quality and decomposition, as well as changes in litter quantity. Generally, herbivory resulted in improved leaf litter chemical quality, including significantly increased nitrogen (N) and phosphorus (P) concentrations and decreased carbon (C) to nitrogen (C:N), C:P, N:P, and lignin:N ratios. Beetle‐affected litter decomposed 23% faster than control litter, and released 16% more N and 60% more P during six months of decomposition, as compared to control litter. Both litter types showed a net release of N and P during decomposition. In addition, herbivory resulted in significant increases in annual rates of total aboveground litter and leaf litter production of 82% and 71%, respectively, under the Tamarix canopy. Our finding that increased rates of N and P release linked with an increased rate of mass loss during decomposition resulting from herbivore‐induced increases in litter quality provides new support to the nutrient acceleration hypothesis. Moreover, results of this study demonstrate that the introduction of the northern tamarisk beetle as biological control to a Tamarix‐invaded riparian ecosystem has lead to short‐term stimulation of nutrient cycling. Alterations to nutrient dynamics could have implications for future plant community composition, and thus the potential for restoration of Tamarix‐invaded ecosystems.     

Differing influences of natural and artificial disturbances on riparian cottonwoods from prairie to mountain ecoregions in Alberta, Canada

Aim Ecoregions represent biophysical zones where environmental factors enable the development of particular plant communities. Ecoregions are generally large but abrupt transitions occur in areas with rapid physical change. A particularly abrupt transitional sequence occurs in the Rocky Mountain region of south‐western Alberta where fescue prairie, aspen parkland and mountain ecoregions occur within 15 km. To investigate plant adaptation across ecoregions, our study investigated the influences of a natural disturbance (flooding) and an artificial disturbance (cattle grazing) on reproductive and population processes of black cottonwood (Populus balsamifera subsp. trichocarpa, Torr. & Gray), the dominant riparian tree. Location We studied cottonwoods throughout their elevational range along two free‐flowing, first‐order streams, Yarrow and Drywood creeks. Cottonwood was the only prominent tree in the prairie ecoregion, the dominant riparian tree in the parkland and extended upward through the montane ecoregion where it was a pioneer species for the mixed coniferous–deciduous woodland. Cottonwoods did not occur in the higher elevation sub‐alpine ecoregion. Methods Thirty‐six cross‐sectional sampling transects were located across the three ecoregions with cottonwoods, and in ungrazed and grazed areas of each ecoregion. Rectangular 100 m² tree and 2 m² seedling quadrats were positioned along the transects, and substrate and vegetation were assessed. Historic hydrological data were analysed relative to flood recurrences and seasonal flow patterns. Results Overall, the cottonwoods displayed a sawtooth shaped ‘punctuated progressive age structure’ with many young trees, progressively fewer older trees, and about four pulses of increased recruitment over the past century. This was considered to provide a healthy cottonwood population and recruitment pulses were apparently associated with flood events with appropriate peak timing and magnitude and a gradual post‐flood stage recession. However, analyses of tree, sapling and seedling data indicated that flood‐associated seedling recruitment was less important and clonal processes were more important for cottonwood recruitment in the montane ecoregion, the highest ecoregion with cottonwoods. The correlation between flood events and cottonwood recruitment was strongest in the mid‐elevation parkland ecoregion suggesting greater reliance on flood‐associated seedling recruitment. There was little correlation with flooding and limited recruitment in the fescue prairie ecoregion in recent decades and the disturbed age structure probably results from cattle impacts that have prevented recruitment and produced a decrepit cottonwood forest population. Main conclusions These analyses suggested that a healthy cottonwood population displayed a sawtooth shaped ‘punctuated progressive age structure’ and that cottonwood reproduction processes varied across ecoregions with increased clonality in the highest montane ecoregion. Cattle grazing impacts on reproduction were most severe in the lowest prairie ecoregion that is treeless except for the riparian zone. We conclude that appropriate strategies of instream flow regulation, land‐use policies and practices, and conservation and restoration efforts should be refined according to ecoregion to recognize the differences in cottonwood reproductive and population ecology.     

Seedling competition between native cottonwood and exotic saltcedar: implications for restoration

Altered hydrology of southwestern United States rivers has led to a decline in native cottonwood (Populus deltoides). Areas historically dominated by cottonwood have been replaced by invasive saltcedar (Tamarix chinensis). Restoration of historic hydrology through periodic flooding of riparian areas has been a means of restoring native species. However, due to similarity in germination requirements of cottonwoods and saltcedars, flooding may create an unwanted increase in the number of saltcedar seedlings. Therefore, we evaluated competitive aspects of these co-occurring species in an extant riparian habitat in the arid southwestern US. We measured effects of competition between cottonwood and saltcedar seedlings and among cottonwood seedlings during the first growing season following seedling establishment in 360, 0.5 × 0.5-m plots at the Bosque del Apache National Wildlife Refuge, New Mexico. We used five interspecific density treatments and five intraspecific density treatments. Cottonwood seedling biomass and height were twice that of saltcedar seedlings across all density treatments. As density of cottonwood increased, intraspecific competition increased in severity and biomass of cottonwood seedlings decreased. At 4 plants/0.25 m², cottonwood seedlings had the greatest biomass; whereas, survival was highest at 10 plants/0.25 m². Our results support greenhouse studies and suggest that if favorable germination conditions are established for cottonwood in floodplains, saltcedar seedlings that cogerminate could be outcompeted by native cottonwood seedlings.     

Effects of flooding on leaf development,transpiration, and photosynthesis in narrowleaf cottonwood,a willow-like poplar

The narrowleaf cottonwood, Populus angustifolia, occurs in occasionally flooded, low elevation zones along river valleys near the North American Rocky Mountains. This small poplar has narrow leaves and fine branching and thus resembles willows, which are commonly flood-tolerant. We investigated the flood response of narrowleaf cottonwoods and a related native hybrid, jackii cottonwood (P. × jackii = P. balsamifera × P. deltoides), by studying saplings of 24 clones in a greenhouse, with some pots being inundated to provide the flood treatment. Flooding slightly reduced leaf numbers (−10%), and leaf sizes were reduced by about 21% in female P. angustifolia versus a 50% reduction in the female hybrids. Flooding-reduced stomatal conductance and net photosynthetic rate, and reduced transpiration particularly in P. × jackii. The effects on foliar gas exchange declined over a 5-week interval, suggesting compensation. The moderate impact of flooding supports the hypothesis that narrowleaf cottonwoods are flood-tolerant, and we anticipate that these trees could provide traits to increase the flood tolerance of fast-growing hybrid poplars. The results further indicate that female cottonwoods may be more flood-tolerant than males, and females could be more successful in lower, flood-prone sites.     

Leaf nutrient concentration as an indicator of Populus and Tamarix response to flooding

Plants growing in infertile environments are able to produce more biomass per unit of nutrient taken up than plants of fertile habitats, and also to minimize nutrients loss by resorbing them from senescing leaves. The leaf nutrient concentration variability of two co-existing riparian tree genera (Populus and Tamarix) along a flood inundation gradient was examined to infer nutrient limitation and to compare nutrient use strategies in the two genera. To that end, seasonal and spatial variability in leaf nitrogen (N) and phosphorus (P) concentration (i.e., % dry mass of N and P) were analyzed in 720 samples of leaves (2 tree genera × 3 seasons × 12 sites × 10 tree replicates). Both Populus and Tamarix showed strong seasonal variability in leaf N and P concentrations, with values decreasing throughout the growing season. However, while N:P atomic ratio remained seasonally constant in Populus (N:P = 33), Tamarix shifted from N:P = 29 in spring to N:P = 36 and 37 in summer and fall. %N, %P and N:P atomic ratios were also spatially variable, but leaf litter N and P concentration (i.e., nutrient resorption proficiency) and leaf litter N:P generally followed the local flood inundation gradient as shown by linear mixed effects models. In particular, nutrient resorption was usually less proficient (higher terminal nutrient concentrations) at higher flood durations (in gravel bars and natural levees), whereas N:P increased in the drier sites (floodplain terrace). At floodplain level, a P-limitation that is higher than N-limitation seems to characterize the plant nutrient circulation in the riparian ecosystem studied. Tamarix was slightly more proficient in P resorption than Populus. The study shows that leaf nutrient concentration (e.g., N and P) derived from nutrient availability is partly controlled by the flood inundation regime and can be used as an indicator of nutrient limitation in forested floodplains. Subtle differences between tree genera provide an additional, novel explanation for the recent expansion of Tamarix in many arid and semi-arid rivers with altered hydrogeomorphic regimes.     

Leaf litter identity alters the timing of lotic nutrient dynamics

相似文献   

Mechanisms of Riparian Cottonwood Decline Along Regulated Rivers

Decline of riparian forests has been attributed to hydrologic modifications to river flows. However, little is known about the physiological and structural adjustments of riparian forests subject to modified flow regimes, and the potential for forest restoration using historic flow regimes is poorly understood. In this paired river study, we compared hydrology, water relations, and forest structure in cottonwood-dominated floodplains of the regulated Green River to those of the unregulated Yampa River. We measured floodplain groundwater levels, soil water availability, cottonwood xylem pressure (Ψ_xp), and leaf-level stomatal conductance (g_s) to assess current impacts of river regulation on the water status of adult cottonwoods. We also simulated a flood on the former floodplain of the regulated river to evaluate its impact on cottonwood water relations. Canopy and root structure were quantified with estimates of cottonwood leaf area and percent live canopy and root density and biomass, respectively. Regulation of the Green River has lowered the annual peak flow yet raised minimum flows in most years, resulting in a 60% smaller stage change, and lowered soil water availability by as much as 70% compared to predam conditions. Despite differences in water availability, daily and seasonal trends in Ψ_xp and g_s were similar for cottonwoods on the regulated and unregulated rivers. In addition, soil water added with the experimental flood had little effect on cottonwood water relations, contrary to our expectations of alleviated water stress. Green River cottonwoods had 10%–30% lower stand leaf area, 40% lower root density, and 25% lower root biomass compared with those for Yampa River cottonwoods. Our results suggest that water relations at the leaf and stem level are currently similar for Yampa and Green River trees due to structural adjustments of cottonwood forests along the Green River, triggered by river regulation.     

Genetic and environmental influences on leaf phenology and cold hardiness of native and introduced riparian trees

To explore the roles of plasticity and genetic variation in the response to spatial and temporal climate variation, we established a common garden consisting of paired collections of native and introduced riparian trees sampled along a latitudinal gradient. The garden in Fort Collins, Colorado (latitude 40.6°N), included 681 native plains cottonwood (Populus deltoides subsp. monilifera) and introduced saltcedar (Tamarix ramosissima, T. chinensis and hybrids) collected from 15 sites at 29.2–47.6°N in the central United States. In the common garden both species showed latitudinal variation in fall, but not spring, leaf phenology, suggesting that the latitudinal gradient in fall phenology observed in the field results at least in part from inherited variation in the critical photoperiod, while the latitudinal gradient in spring phenology observed in the field is largely a plastic response to the temperature gradient. Populations from higher latitudes exhibited earlier bud set and leaf senescence. Cold hardiness varied latitudinally in both fall and spring for both species. For cottonwood, cold hardiness began earlier and ended later in northern than in southern populations. For saltcedar northern populations were hardier throughout the cold season than southern populations. Although cottonwood was hardier than saltcedar in midwinter, the reverse was true in late fall and early spring. The latitudinal variation in fall phenology and cold hardiness of saltcedar appears to have developed as a result of multiple introductions of genetically distinct populations, hybridization and natural selection in the 150 years since introduction.     

Insect outbreaks alter nutrient dynamics in a southern African savanna: patchy defoliation of Colophospermum mopane savanna by Imbrasia belina larvae

Invertebrate herbivore outbreaks have important impacts on system biogeochemical cycling, but these effects have been poorly documented in African savanna ecosystems. In semi‐arid African savannas, outbreaks of the lepidopteran Imbrasia belina (mopane worm) affect discrete patches of the dominant Colophospermum mopane trees; larvae may completely defoliate trees for up to six weeks during each of the early and late growing seasons. We studied the impact of mopane worm outbreaks on the availability of nitrogen (N), phosphorus (P), and potassium (K) within mopane savanna by quantifying major nutrient pools in defoliated and non‐defoliated savanna patches, including leaves, leaf litter, worm frass, and the soil beneath trees. Within an outbreak area, approximately 44 percent of trees were infested, supporting ~29,000 worms/ha, leading to ~640 kg/ha dry weight frass deposition at 1.4 g of frass/day‐individual (fourth or fifth instar), compared with an average 1645 kg/ha dry weight of leaf on trees most of which should be deposited by litterfall at the end of the growing season. Frass had twofold higher P, 10 percent higher K, but equivalent N content than litter. Taking frass and litter deposition together, the N, P, and K contents added due to the outbreak event at our study site were 0.88, 5.8, and 2.8 times those measured in non‐outbreak patches, a pattern which was reflected in the nutrient contents of soil surfaces beneath defoliated trees. Invertebrate herbivory appears to be an important driver for mopane savanna but has been largely neglected.     

炼锌废渣-修复植物-凋落物体系的生态化学计量学研究

为揭示金属冶炼废渣堆场生态修复多年后,废渣-植物-凋落物系统中养分循环和系统维持机制。该研究以实现生态修复6 a的黔西北铅锌冶炼废渣堆场上土荆芥(Dysphania ambrosioides)、芦竹(Arundo donax)、刺槐(Robinia pseudoacacia)、构树(Broussonetia papyrifera)和柳杉(Cryptomeria fortunei)五种优势修复植物为对象,分析它们的主要营养器官(细根、粗根、茎/干、枝、叶片)、地表凋落物、植被下方表层废渣(0～10 cm)中碳(C)、氮(N)、磷(P)含量及化学计量特征,探讨它们之间的相互关系。结果表明:不同植物、不同营养器官间C、N、P的含量具有显著差异(P<0.05),C平均含量在两种草本植物中为茎>叶片>根>凋落物,在三种乔木中为干>枝>细根>粗根>叶片>凋落物; N和P的分布在草本植物中分别为叶片>凋落物>根>茎和叶片>根>凋落物>茎,在三种乔木中均为叶片>细根>凋落物>粗根>枝>干。五种植物中,柳杉各营养器官及凋落物中C含量均高于其他植物,N、P含量呈相反的规律; 刺槐中N含量最高。C:N和C:P在五种植物营养器官与凋落物中的变化规律跟N、P的分布相反,说明C:N和C:P分别主要受N和P含量影响。相关性分析指出,草本植物的N:P受N和P共同影响,三种乔木的N:P主要由N的分布决定,同时受到枝和叶片中P含量影响。五种植物中,仅豆科类刺槐的叶片N:P大于16,在系统中生长受P限制,其他植物生长均受N限制,说明刺槐更能适应贫瘠的废渣环境,建议在修复贫瘠的废渣堆场时优先选择豆科类植物作为先锋植物,改善基质养分条件。植被下方表层废渣中C、N、P含量基本都低于植物各营养器官及凋落物,不同修复植物下方对应的表层废渣中C、N、P含量间具有显著差异(P<0.05),草本植物修复下的废渣中C、N、P含量低于乔木修复下的含量。废渣-植物-凋落物体系中N、P、N:P之间的相关性分析显示,植物细根和凋落物中N、P含量与废渣中N、P含量及化学计量比关系更密切。     

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

High‐frequency fire alters C : N : P stoichiometry in forest litter

Fire is a major driver of ecosystem change and can disproportionately affect the cycling of different nutrients. Thus, a stoichiometric approach to investigate the relationships between nutrient availability and microbial resource use during decomposition is likely to provide insight into the effects of fire on ecosystem functioning. We conducted a field litter bag experiment to investigate the long‐term impact of repeated fire on the stoichiometry of leaf litter C, N and P pools, and nutrient‐acquiring enzyme activities during decomposition in a wet sclerophyll eucalypt forest in Queensland, Australia. Fire frequency treatments have been maintained since 1972, including burning every 2 years (2yrB), burning every 4 years (4yrB) and no burning (NB). C : N ratios in freshly fallen litter were 29–42% higher and C : P ratios were 6–25% lower for 2yrB than NB during decomposition, with correspondingly lower 2yrB N : P ratios (27–32) than for NB (34–49). Trends in litter soluble and microbial N : P ratios were similar to the overall litter N : P ratios across fire treatments. Consistent with these, the ratio of activities for N‐acquiring to P‐acquiring enzymes in litter was higher for 2yrB than NB, whereas 4yrB was generally intermediate between 2yrB and NB. Decomposition rates of freshly fallen litter were significantly lower for 2yrB (72 ± 2% mass remaining at the end of experiment) than for 4yrB (59 ± 3%) and NB (62 ± 3%), a difference that may be related to effects of N limitation, lower moisture content, and/or litter C quality. Results for older mixed‐age litter were similar to those for freshly fallen litter although treatment differences were less pronounced. Overall, these findings show that frequent fire (2yrB) decoupled N and P cycling, as manifested in litter C : N : P stoichiometry and in microbial biomass N : P ratio and enzymatic activities. Furthermore, these data indicate that fire induced a transient shift to N‐limited ecosystem conditions during the postfire recovery phase.     

Growth of an invertebrate shredder on native (Populus) and non-native (Tamarix, Elaeagnus) leaf litter

1. Large-scale invasions of riparian trees can alter the quantity and quality of allochthonous inputs of leaf litter to streams and thus have the potential to alter stream organic matter dynamics. Non-native saltcedar ( Tamarix sp.) and Russian olive ( Elaeagnus angustifolia ) are now among the most common trees in riparian zones in western North America, yet their impacts on energy flow in streams are virtually unknown.
2. We conducted a laboratory feeding experiment to compare the growth of the aquatic crane fly Tipula (Diptera: Tipulidae) on leaf litter from native cottonwood ( Populus ) and non-native Tamarix and Elaeagnus . Tipula showed positive growth on leaf litter of all three species; however, after 7 weeks, larvae fed Tamarix leaves averaged 1.7 and 2.5 times the mass of those fed Elaeagnus and Populus , respectively. Tipula survival was highest on Populus , intermediate on Tamarix and lowest on Elaeagnus .
3. High Tipula growth on Tamarix probably reflects a combination of leaf chemistry and morphology. Conditioned Tamarix leaf litter had intermediate carbon : nitrogen values (33 : 1) compared to Populus (40 : 1) and Elaeagnus (26 : 1), and it had intermediate proportions of structural carbon (42%) compared to Elaeagnus (57%) and Populus (35%). Tamarix leaves are also relatively small and possibly more easily ingested by Tipula than either Elaeagnus or Populus .
4. Field surveys of streams in the western U.S.A. revealed that Tamarix and Elaeagnus leaf packs were rare compared to native Populus , probably due to the elongate shape and small size of the non-native leaves. Thus we conclude that, in general, the impact of non-native riparian invasion on aquatic shredders will depend not only on leaf decomposition rate and palatability but also on rates of leaf litter input to the stream coupled with streambed retention and subsequent availability to consumers.     

Nitrogen and phosphorus economy of the riparian shrub <Emphasis Type="Italic">Salix gracilistyla</Emphasis> in western Japan

Salix gracilistyla is one of the dominant plants in the riparian vegetation of the upper-middle reaches of rivers in western Japan. This species colonizes mainly sandy habitats, where soil nutrient levels are low, but shows high potential for production. We hypothesized that S.␣gracilistyla uses nutrients conservatively within stands, showing a high resorption efficiency during leaf senescence. To test this hypothesis, we examined seasonal changes in nitrogen (N) and phosphorus (P) concentrations in aboveground organs of S. gracilistyla stands on a fluvial bar in the Ohtagawa River, western Japan. The concentrations in leaves decreased from April to May as leaves expanded. Thereafter, the concentrations showed little fluctuation until September. They declined considerably in autumn, possibly owing to nutrient resorption. We converted the nutrient concentrations in each organ to nutrient amounts per stand area on the basis of the biomass of each organ. The resorption efficiency of N and P in leaves during senescence were estimated to be 44 and 46%, respectively. Annual net increments of N and P in aboveground organs, calculated by adding the amounts in inflorescences and leaf litter to the annual increments in perennial organs, were estimated to be 9.9 g and 0.83 g m⁻² year⁻¹, respectively. The amounts released in leaf litter were 6.7 g N and 0.44 g P m⁻². These values are comparable to or larger than those of other deciduous trees. We conclude that S. gracilistyla stands acquire large amounts of nutrients and release a large proportion in leaf litter.     

Leaf litter quality affects aquatic insect emergence: contrasting patterns from two foundation trees

Reciprocal subsidies between rivers and terrestrial habitats are common where terrestrial leaf litter provides energy to aquatic invertebrates while emerging aquatic insects provide energy to terrestrial predators (e.g., birds, lizards, spiders). We examined how aquatic insect emergence changed seasonally with litter from two foundation riparian trees, whose litter often dominates riparian streams of the southwestern United States: Fremont (Populus fremontii) and narrowleaf (Populus angustifolia) cottonwood. P. fremontii litter is fast-decomposing and lower in defensive phytochemicals (i.e., condensed tannins, lignin) relative to P. angustifolia. We experimentally manipulated leaf litter from these two species by placing them in leaf enclosures with emergence traps attached in order to determine how leaf type influenced insect emergence. Contrary to our initial predictions, we found that packs with slow-decomposing leaves tended to support more emergent insects relative to packs with fast-decomposing leaves. Three findings emerged. Firstly, abundance (number of emerging insects m^?2 day^?1) was 25 % higher on narrowleaf compared to Fremont leaves for the spring but did not differ in the fall, demonstrating that leaf quality from two dominant trees of the same genus yielded different emergence patterns and that these patterns changed seasonally. Secondly, functional feeding groups of emerging insects differed between treatments and seasons. Specifically, in the spring collector-gatherer abundance and biomass were higher on narrowleaf leaves, whereas collector-filterer abundance and biomass were higher on Fremont leaves. Shredder abundance and biomass were higher on narrowleaf leaves in the fall. Thirdly, diversity (Shannon’s H′) was higher on Fremont leaves in the spring, but no differences were found in the fall, showing that fast-decomposing leaves can support a more diverse, complex emergent insect assemblage during certain times of the year. Collectively, these results challenge the notion that leaf quality is a simple function of decomposition, suggesting instead that aquatic insects benefit differentially from different leaf types, such that some use slow-decomposing litter for habitat and its temporal longevity and others utilize fast-decomposing litter with more immediate nutrient release.     

Branch sacrifice: cavitation-associated drought adaptation of riparian cottonwoods

In their native riparian zones (floodplains), Populus deltoides (prairie cottonwood) and P. fremontii (Fremont cottonwood) commonly experience substantial branch die-back. These trees occur in semi-arid areas of North America and unexpectedly given the dry regions, they are exceptionally vulnerable to xylem cavitation, drought-induced air embolism of xylem vessels. We propose that the vulnerability to cavitation and branch die-back are physiologically linked; drought-induced cavitation underlies branch die-back that reduces transpirational demand enabling the remaining shoot to maintain a favorable water balance. This proposal follows field observation along various western North American rivers as precocious branch senescence, the yellowing and death of leaves on particular branches during mid- to late summer, was common for P. deltoides and P. fremontii during hot and dry periods of low stream-flow. Branches displaying precocious senescence were subsequently dead the following year. The proposed association between cavitation, precocious senescence and branch die-back is also supported by experiments involving external pressurization of branches to about 2.5 MPa with a branch collar or through an adjacent cut-branch. The treatments induced xylem cavitation and increased leaf diffusive resistance (stomatal closure) that was followed by leaf senescence and branch death of P. deltoides. P. trichocarpa (black cottonwood) appeared to be less affected by the pressurization treatment and this species as well P. angustifolia (narrowleaf cottonwood) and P. balsamifera (balsam poplar) seldom display the patchy summer branch senescence typical of P. deltoides and P. fremontii. ’Branch sacrifice’ describes this cavitation-associated senescence and branch die-back that may provide a drought adaptation for the prairie and Fremont cottonwoods. Received: 13 May 1999 / Accepted: 4 November 1999     

云南药山自然保护区黄背栎林和巧家五针松林生态化学计量特征

以云南药山自然保护区黄背栎林和巧家五针松林的4块样地为研究对象,旨在揭示这2种森林生态系统营养元素含量状况和土壤养分的供给能力。通过分析4块样地"叶片-凋落物-土壤"C、N、P含量、生态化学计量特征及其相关性,结果表明:(1)不同样地同一组分的C、N含量差异显著,P含量差异不显著,同一样地各组分间C、N含量差异显著,均为叶片凋落物土壤,P含量则为叶片土壤凋落物;(2)不同样地叶片C∶P、N∶P比值和凋落物与土壤C∶N比值差异不显著,其余指标差异均显著。同一样地叶片、凋落物、土壤的C∶N、C∶P、N∶P比值差异显著,均为凋落物叶片土壤;(3)黄背栎林叶片-土壤C含量、C∶P比值和凋落物-叶片N∶P比值呈极显著或显著相关,巧家五针松林凋落物-叶片C、N含量和叶片-土壤P含量、C∶N、N∶P比值呈极显著或显著相关;(4)土壤N元素缺乏是限制植物生长的主要因素,P元素主要源于土壤矿物风化释放,而非生物小循环。     

Palatability, decomposition and insect herbivory: patterns in a successional old-field plant community

We tested the hypothesis that selective feeding by insect herbivores in an old‐field plant community induces a shift of community structure towards less palatable plant species with lower leaf and litter tissue quality and may therefore affect nutrient cycling. Leaf palatability of 20 herbaceous plant species which are common during the early successional stages of an old‐field plant community was assayed using the generalist herbivores Deroceras reticulatum (Mollusca: Agriolomacidae) and Acheta domesticus (Ensifera: Gryllidae). Palatability was positively correlated with nitrogen content, specific leaf area and water content of leaves and negatively correlated with leaf carbon content and leaf C/N‐ratio. Specific decomposition rates were assessed in a litter bag experiment. Decomposition was positively correlated with nitrogen content of litter, specific leaf area and water content of living leaves and negatively correlated with leaf C/N‐ratio. When using phylogentically independent contrasts the correlations between palatability and decomposition versus leaf and litter traits remained significant (except for specific leaf area) and may therefore reflect functional relationships. As palatability and decomposition show similar correlations to leaf and litter traits, the correlation between leaf palatability and litter decomposition rate was also significant, and this held even in a phylogenetically controlled analysis. This correlation highlights the possible effects of invertebrate herbivory on resource dynamics. In a two‐year experiment we reduced the density of above‐ground and below‐ground insect herbivores in an early successional old‐field community in a two‐factorial design by insecticide application. The palatability ranking of plants showed no relationship with the specific change of cover abundance of plants due to the reduction of above‐ or below‐ground herbivory. Thus, changes in the dominance structure as well as potentially associated changes in the resource dynamics are not the result of differences in palatability between plant species. This highlights fundamental differences between the effects of insect herbivory on ecosystems and published results from vertebrate‐grazing systems.     

Nutrient content and seasonal fluctuations in the leaf component of coark-oak (Quercus suber L.) litterfall

Nutrient content and seasonality of the leaf component in cork-oak litterfal were studied over a two year period in two cork-oak forest sites differing in biomass and edaphic condition in the north-eastern Iberian peninsula. Fallen senescent leaves compared to young leaves showed higher non-mobile nutrient concentrations and lower mobile nutrient concentrations, specially P, N, K, and Mg. At both sites, seasonal fluctuations affected both leaf production and leaf mineral content. The maximum leaf fall period correspond to the start of the vegetative growth and to the lowest N and P concentration in the falling leaves. The opposite was true for the winter, when litterfal was minimal and N and P content in falling leaves was at a maximum.The comparison between falling leaves and canopy leaves suggests that the analysis of fallen leaves can be a useful measure of N and P nutrition in cork-oak. We found site dependent differences in nutrient content and nutrient remobilization. In comparison with Q. ilex, although litter production was in the same range, nutrient retranslocation was greater for Q. suber.