Foliar and Litter Nutrients, Nutrient Resorption, and Decomposition in Hawaiian Me t rosideros polymorpha

The native tree Metrosideros polymorpha dominates Hawaiian forests across a very wide range of soil fertility, including both sites where forest production is limited by nitrogen (N) and others where it is limited by phosphorus (P). Five long-term fertilization experiments have further broadened the range of nutrient availabilities experienced by Metrosideros. Adding P to P-limited sites increased foliar P concentrations threefold and litter P concentrations up to 10-fold; lignin concentrations decreased, and the decomposability of leaf litter increased from 32%–35% to 36%–46% mass loss in the first year. Adding N to N-limited sites increased leaf and litter N concentrations by only 15%–20%, with little or no effect on the decomposability of tissue. Received 22 January 1998; accepted 4 May 1998.     

Nutrient Limitation in a Fire‐derived,Nitrogen‐rich Hawaiian Grassland1

Grasslands created by grass invasions into shrublands or woodlands followed by fire are now a dominant feature of many seasonally dry environments. In Hawaii Volcanoes National Park, introduced perennial grasses dominate grasslands created by fire in grass‐invaded woodlands. In a previous study, we found that net primary production in these grasslands is substantially lower than in unburned woodlands. Yet, our estimates of annual net nitrogen (N) mineralization showed higher rates in these savannas than in the unburned woodlands, rates that appear to greatly exceed annual N demand by the vegetation. We therefore hypothesized that N should not be limited to the plants growing in these sites. We tested this hypothesis with a 2‐yr fertilization experiment. At peak biomass, we found a 30 percent increase in live biomass in plots with N added and no increase in production with only phosphorus (P) added. N and P together were synergistic, suggesting that co‐limitation or P limitation becomes important when N is more available. Plants responded to added N by increasing individual leaf area and shoot length by over 50 percent. Tissue N was higher with added N; hence, biomass N was substantially higher. Tissue P concentrations declined with N addition but were elevated by P addition despite lack of a growth response to P alone. Overall, N limitation exists despite high annual rates of net N mineralization, and co‐limitation of production by P may occur when N is abundant. Here, asynchrony between plant nutrient demand and N availability may contribute to N limitation.     

Nitrogen Fixation in Bryophytes, Lichens, and Decaying Wood along a Soil-age Gradient in Hawaiian Montane Rain Forest

We determined rates of acetylene reduction and estimated total nitrogen fixation associated with bryophytes, lichens, and decaying wood in Hawaiian montane rain forest sites with underlying substrate ranging in age from 300 to 4.1 million years. Potential N fixation ranged from ca 0.2 kg/ha annually in the 300‐year‐old site to ca 1 kg/ha annually in the 150,000‐year‐old site. Rates of acetylene reduction were surprisingly uniform along the soil‐age gradient, except for high rates in symbiotic/associative fixers at the 150,000‐year‐old site and in heterotrophic fixers at the 2100‐year‐old site. Low fixation at the youngest site, where plant production is known to be N‐limited, suggests that demand for N alone does not govern N fixation. Total N fixation was highest in sites with low N:P ratios in leaves and stem wood, perhaps because epiphytic bryophytes and lichens depend on canopy leachate for mineral nutrients and because heterotrophic fixation is partly controlled by nutrient supply in the decomposing substrate; however, differences in substrate cover, rather than in fixation rates, had the largest effect on the total N input from fixation at these sites.     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

Changes in Production and Nutrient Cycling across a Wetness Gradient within a Floodplain Forest

Floodplain forest ecosystems are highly valuable to society because of their potential for water quality improvement and vegetation productivity, among many other functions. Previous studies have indicated that hydrology influences productivity but that the relationship between hydroperiod and productivity is a complex one. Consequently, we compared multiple indexes of productivity, nutrient circulation, and hydroperiod among three communities on the Flint River floodplain, Georgia, that differed in terms of inundation frequency. We hypothesized that (a) the wettest community would have the lowest total net primary production (NPP) values because of saturated soil conditions; (b) as wetness increases, nutrient circulation in litterfall would decrease because of the hypothesized lower productivity in the wetter community; and (c) as wetness increases, internal translocation would become more efficient. The study site was partitioned into three wetness types—somewhat poorly drained (SPD), intermediate (I) and poorly drained (PD). We found that belowground biomass was greatest on the SPD, litterfall was similar for all three sites, and that woody biomass current annual increment (CAI) was greatest in the PD community. However, when the three variables were totaled for each site, the PD had the greatest NPP, thus disproving hypothesis (a). For hypothesis (b), we observed that P content in litterfall, although not significant, followed the predicted trend; nitrogen (N) content displayed the opposite pattern (PD > I > SPD). As wetness increased, internal translocation became more efficient for phosphorus (support for hypothesis [c]), but the SPD community was more efficient at retranslocating N (contradiction of hypothesis [c]). Received 19 June 2000; accepted 19 October 2000.     

Ecosystem Consequences of Exotic Earthworm Invasion of North Temperate Forests

The invasion of north temperate forests by exotic species of earthworms is an important issue that has been overlooked in the study and management of these forests. We initiated research to address the hypothesis that earthworm invasion will have large consequences for nutrient retention and uptake in these ecosystems. In this special feature of Ecosystems, we present five papers describing results from our experiment. In this paper, we (a) introduce our experimental approach and conceptual model of how earthworms influence forest ecosystem processes, (b) describe the characteristics of the study areas and earthworm communities at our two study locations, and (c) provide a brief overview and synthesis of the main findings. The most dramatic effect of earthworm invasion was the loss of the forest floor at an undisturbed forest site, which altered the location and nature of nutrient cycling activity in the soil profile. Invasion changed soil total carbon (C) and phosphorus (P) pools, carbon–nitrogen (C:N) ratios, the loss and distribution of different soil P fractions, and the distribution and function of roots and microbes. Response to invasion varied with site characteristics and earthworm species. Our results suggest that exotic earthworm invasion is a significant factor that will influence the structure and function of northern temperate forest ecosystems over the next few decades. Regional evaluations of these forests will need to consider the presence or absence of earthworms along with other important ecosystem drivers, such as pollution, climate, and underlying soil characteristics.     

Invasion by a Perennial Herb Increases Decomposition Rate and Alters Nutrient Availability in Warm Temperate Lowland Forest Remnants

We determined the impact of the invasive herb, Tradescantia fluminensis Vell., on litter decomposition and nutrient availability in a remnant of New Zealand lowland podocarp–broadleaf forest. Using litter bags, we found that litter beneath mats of Tradescantia decomposed at almost twice the rate of litter placed outside the mat. Values of k (decomposition quotient) were 9.44±0.42 yrs for litter placed beneath Tradescantia and 5.42±0.42 yrs for litter placed in native, non-Tradescantia plots. The impact of Tradescantia on decomposition was evident through the smaller forest floor mass in Tradescantia plots (2.65±1.05 t ha⁻¹) compared with non-Tradescantia plots (5.05±1.05 t ha⁻¹), despite similar quantities of annual leaf litterfall into Tradescantia plots (6.85±0.85 t ha⁻¹ yr⁻¹) and non-Tradescantia plots (7.45±1.05 t ha⁻¹ yr⁻¹). Moreover, there was increased plant nitrate available, as captured on resin bags, in Tradescantia plots (25.77 ± 8.32 cmol(−)/kg resin) compared with non-Tradescantia plots (9.55±3.72 cmol(−)/kg resin). Finally, the annual nutrient uptake by Tradescantia represented a large proportion of nutrients in litterfall (41% N, 61% P, 23% Ca, 46% Mg and 83% K), exceeded the nutrient content of the forest floor (except Ca), but was a small proportion of the topsoil nutrient pools. Taken together, our results show that Tradescantia increases litter decomposition and alters nutrient availability, effects that could influence the long-term viability of the majority of podocarp–broadleaf forest remnants affected with Tradescantia in New Zealand. These impacts are likely mostly due to Tradescantia's vegetation structure (i.e., tall, dense mats) and associated microclimate, compared with native ground covers. This revised version was published online in July 2006 with corrections to the Cover Date.     

The Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

The Effect of Water on Decomposition Dynamics in Mesic to Wet Hawaiian Montane Forests

I used a mesic to wet precipitation gradient on Maui, Hawaii, to test whether variation in rainfall regulates decomposition in tropical wet forest. Decomposition rates of leaves and roots from the dominant tree species, Metrosideros polymorpha, were measured at six sites similar in temperature regime, parent material, ecosystem age, vegetation, and topographical relief, whereas mean annual precipitation (MAP) at these six sites varied from 2200 to over 5000 mm/y. In situ decomposition rates of leaves placed on the soil surface declined by a factor of 6.4 with increased precipitation, whereas the decomposition rate of roots placed below ground declined by a factor of 2.3 across the gradient. Leaves collected from the 2200-mm site and placed at all sites on the gradient decomposed faster on the soil surface than they did below ground, whereas both above- and belowground decomposition rates of the common leaves decreased by a factor of 2.5 with increased precipitation. Of the environmental variables that changed with MAP, soil oxygen availability appeared to be the proximal factor that limited decomposition rates across the gradient, both above and below ground. When plant tissue collected from all sites across the gradient was decomposed at a common site, leaves from the wettest sites decomposed almost three times more slowly than leaves from the mesic sites. In contrast, roots from across the gradient all decomposed at a similar rate in a common site. Of tissue chemistry variables, high lignin concentration was correlated consistently with slow decomposition for roots and leaves. These results suggest that soil oxygen limitation combined with poorly decomposable leaves caused slower rates of decomposition and nutrient release with increased rainfall in these upland forests. Received 14 April 2000; Accepted 11 December 2000.     

Asymbiotic Nitrogen Fixation and Litter Decomposition on a Long Soil-Age Gradient in Hawaiian Montane Rain Forest1

We determined rates of decomposition and asymbiotic nitrogen fixation in the leaf litter of Cheirodendron spp. on the Hawaiian Islands. Leaf litter was collected from four sites on a long soil-age gradient (300 yr to 4.1 M yr) and decomposed at two sites that differed widely in substrate age and nutrient availability. Rates of decomposition were higher in litter decomposed at the older site, where nutrient availability was greater. A substantial amount of nitrogen and phosphorus immobilization occurred in litter decomposed at the older site, with more immobilization occurring in litter with lower initial nitrogen and phosphorus concentrations, suggesting both supply and demand controls on nutrient immobilization. Potential rates of nitrogen fixation were very low in the first 25 d (0–5 nmol acetylene/gdw/h), rose to much higher rates by 70 d (20–45 nmol), and then declined by 140 d. We found no significant difference in rates of potential nitrogen fixation between sites of decomposition, but there was a strong substrate effect, with higher rates in litter with low lignin, low nitrogen, and high phosphorus. Where significant immobilization of nitrogen occurred for decomposing Cheirodendron, nitrogen fixation could have comprised no more than 10 percent of immobilized nitrogen. Overall, rates of nitrogen fixation were dependent on the source of the decomposing substrate but not on the site of decomposition, while short-term decomposition and nutrient immobilization were strongly dependent on the site of decomposition but not as much on the source of the decomposing substrate.     

Effects of Exotic Plant Invasions on Soil Nutrient Cycling Processes

Although it is generally acknowledged that invasions by exotic plant species represent a major threat to biodiversity and ecosystem stability, little attention has been paid to the potential impacts of these invasions on nutrient cycling processes in the soil. The literature on plant–soil interactions strongly suggests that the introduction of a new plant species, such as an invasive exotic, has the potential to change many components of the carbon (C), nitrogen (N), water, and other cycles of an ecosystem. I have reviewed studies that compare pool sizes and flux rates of the major nutrient cycles in invaded and noninvaded systems for invasions of 56 species. The available data suggest that invasive plant species frequently increase biomass and net primary production, increase N availability, alter N fixation rates, and produce litter with higher decomposition rates than co-occurring natives. However, the opposite patterns also occur, and patterns of difference between exotics and native species show no trends in some other components of nutrient cycles (for example, the size of soil pools of C and N). In some cases, a given species has different effects at different sites, suggesting that the composition of the invaded community and/or environmental factors such as soil type may determine the direction and magnitude of ecosystem-level impacts. Exotic plants alter soil nutrient dynamics by differing from native species in biomass and productivity, tissue chemistry, plant morphology, and phenology. Future research is needed to (a) experimentally test the patterns suggested by this data set; (b) examine fluxes and pools for which few data are available, including whole-site budgets; and (c) determine the magnitude of the difference in plant characteristics and in plant dominance within a community that is needed to alter ecosystem processes. Such research should be an integral component of the evaluation of the impacts of invasive species.     

林冠附生物在森林生态系统养分循环中的作用

1　引　言林冠是森林生态系统中重要的组成部分。由于认识上、技术上和其它方面的原因 ,过去一直都把林冠当作“黑箱”来研究。近 2 0年来 ,随着对林冠生物多样性、林冠附生物在生态系统功能过程影响认识和研究技术上的提高 ,对林冠附生物的研究已逐步从个体水平转移到系统水平上。林冠现在被认为是一个适宜于许多动植物种类生存的场所 ,其物种数量比原先所想象的更为丰富。通过对林冠的研究 ,促进了对天然林生态系统中养分循环的了解 ,以及林冠干扰或破坏对自然和人类活动的影响。其中林冠附生物质在森林生态系统养分循环过程中的作用和影…     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

红松人工林枝叶分解速度及养分动态的研究

修枝作为培育无节良材的重要经营措施已被广泛应用于生产,修落的枝叶能为林地提供大量的有机物,分解后提高林地土壤肥力。为了探明红松人工纯林枝叶分解过程中各种营养元素的变化规律,我们于1984年至1986年在草河     

Exotic Earthworm Invasion and Microbial Biomass in Temperate Forest Soils

Invasion of north temperate forest soils by exotic earthworms has the potential to alter microbial biomass and activity over large areas of North America. We measured the distribution and activity of microbial biomass in forest stands invaded by earthworms and in adjacent stands lacking earthworms in sugar maple-dominated forests in two locations in New York State, USA: one with a history of cultivation and thin organic surface soil horizons (forest floors) and the other with no history of cultivation and a thick (3–5 cm) forest floor. Earthworm invasion greatly reduced pools of microbial biomass in the forest floor and increased pools in the mineral soil. Enrichment of the mineral soil was much more marked at the site with thick forest floors. The increase in microbial biomass carbon (C) and nitrogen (N) in the mineral soil at this site was larger than the decrease in the forest floor, resulting in a net increase in total soil profile microbial biomass in the invaded plots. There was an increase in respiration in the mineral soil at both sites, which is consistent with a movement of organic matter and microbial biomass into the mineral soil. However, N-cycle processes (mineralization and nitrification) did not increase along with respiration. It is likely that the earthworm-induced input of C into the mineral soil created a microbial sink for N, preventing an increase in net mineralization and nitrification and conserving N in the soil profile.     

一个农牧结合生态系统营养循环的源,库,流

一个农牧结合生态系统营养循环的源、库、流曾江海,张玉铭（中国科学院石家庄农业现代化研究所０５００２１）Ｓｏｕｒｃｅ,ＰｏｏｌａｎｄＦｌｕｘｏｆＮｕｔｒｉｅｎｔＣｙｃｌｉｎｇｉｎａＣｏｍｂｉｎｅｄＡｇｒｏ－ＡｎｉｍａｌＨｕｓｂａｎｄｒｙＥｃｏｓｙｓｔｅ...