Effects of Nutrient Enrichment on Decomposition and Fungal Colonization of Sweet Chestnut Leaves in an Iberian Stream (Central Portugal)

This study assessed the effect of nutrient enrichment on rates of decomposition, ergosterol concentrations (as a measure of fungal biomass), and rates of fungal sporulation of sweet chestnut (Castanea sativa Miller) leaves in a 3rd order stream (Central Portugal), with medium to high background values of nutrients. Coarse and fine mesh leaf bags were attached to nutrient diffusing substrata containing NaNO₃, KH₂PO₄, both nutrients, or no additions. Leaf breakdown rates were similar in the four treatments and in the two mesh sizes (k=−0.0155 to −0.0219 day⁻¹). Phosphorus content of P or N + P enriched leaves was higher than in the other treatments after 28 days, but there were no differences in N concentrations. Ergosterol concentrations associated with decomposing leaves were similar among treatments. The peak sporulation rates of aquatic hyphomycetes were stimulated by the addition of N + P and N but not by P alone. Results from the experiment provide evidence that leaf breakdown in the study stream, as a model for streams with naturally medium to high level of nutrients, was not nutrient-limited, and that fungal reproductive activity was limited by dissolved N but not by dissolved P in stream water.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Seasonal growth and biomass ofTrapa japonica Flerov in Ojaga-Ike Pond,Chiba, Japan

In order to determine the seasonal growth and biomass ofTrapa japonica Flerov, field observations were carried out at Ojaga-ike Pond, Chiba, Japan, during 1979 and 1980. In spring, the plant showed exponential growth (c. 0.080 g g⁻¹ day⁻¹) and shoot elongation was as rapid as 10 cm day⁻¹. The plant attained its maximum biomass (380.5±35.1 g m⁻²) in late August, and about 50% of this was concentrated in the topmost 30-cm stratum (645.7±33.1 g m⁻³); maximum total stem length exceeded 6m. The plant produced large (500–800 mg per fruit), but small numbers of nut-like fruit (maximum, 5 fruits per rosette). Defoliation occurred almost linearly with time at a rate of 30.6 leaves m⁻² day⁻¹; annual net leaf production was estimated to be about twice as large as the seasonal maximum leaf biomass. While the number of leaves per rosette showed moderate seasonal change, rosette density, rosette area and leaf dry weight changed considerably during the year. From the negative log-log correlation between mean total leaf dry weight per rosette and rosette density, density-dependent rosette growth was assumed. The cause of the wide spread of this species in aquatic habitats is briefly discussed in terms of its seed size and morphology.     

Does leaf quality mediate the stimulation of leaf breakdown by phosphorus in Neotropical streams?

1. Lowland tropical streams have a chemically diverse detrital resource base, where leaf quality could potentially alter the effect of high nutrient concentrations on leaf breakdown. This has important implications given the extent and magnitude of anthropogenic nutrient loading to the environment. 2. Here, we examine if leaf quality (as determined by concentrations of cellulose, lignin and tannins) mediates the effects of high ambient phosphorus (P) concentration on leaf breakdown in streams of lowland Costa Rica. We hypothesised that P would have a stronger effect on microbial and insect processing of high‐ than of low‐quality leaves. 3. We selected three species that represented extremes of quality as measured in leaves of eight common riparian species. Species selected were, from high‐ to low‐quality: Trema integerrima > Castilla elastica > Zygia longifolia. We incubated single‐species leaf packs in five streams that had natural differences in ambient P concentration (10–140 μg soluble reactive phosphorus (SRP) L^?1), because of variable inputs of solute‐rich groundwater and also in a stream that was experimentally enriched with P (approximately 200 μg SRP L^?1). 4. The breakdown rate of all three species varied among the six streams: T. integerrima (k‐values range: 0.0451–0.129 day^?1); C. elastica (k‐values range: 0.0064–0.021 day^?1); and Z. longifolia (k‐values range: 0.002–0.008 day^?1). Both ambient P concentration and flow velocity had significant effects on the breakdown rate of the three species. 5. Results supported our initial hypothesis that litter quality mediates the effect of high ambient P concentration on leaf processing by microbes and insects. The response of microbial respiration, fungal biomass and invertebrate density to high ambient P concentration was greater in Trema (high quality) than in Castilla or Zygia (low quality). Variation in flow velocity, however, confounded our ability to determine the magnitude of stimulation of breakdown rate by P. 6. Cellulose and lignin appeared to be the most important factors in determining the magnitude of P‐stimulation. Surprisingly, leaf secondary compounds did not have an effect. This contradicts predictions made by other researchers, regarding the key role of plant secondary compounds in affecting leaf breakdown in tropical streams.     

Habitat Use of Juvenile Fish in the Lower Danube and the Danube Delta: Implications for Ecotone Connectivity

Functional processes in freshwater ecosystems are highly influenced by acidic conditions. Foodwebs are affected and macroinvertebrate species diversity is decreased. This study aims to investigate leaf decomposition at very low pH in the acidic Banyupahit–Banyuputih river originating from the acidic crater lake Kawah Ijen in Indonesia. Leaf decomposition experiments were carried out for 200 days in the acidic river at pHs of approximately 0.7, 2.3 and 3.0 and in the neutral Kali Sengon river, using leaves from teak, Tectona grandis, and bamboo, Bambusa sp. Two different types of leaf packs were used: fine mesh size packs were used to exclude macroinvertebrates and coarse mesh size packs allowed macroinvertebrate colonization. Clear differences in decomposition rate were observed between the neutral Kali Sengon and the acidic Banyupahit–Banyuputih river with decomposition in the Kali Sengon river proceeding significantly faster for both leaf types. In the Kali Sengon k values (d⁻¹) over 46 days were 0.0202 for fine teak, 0.0236 for coarse teak, 0.0114 for fine bamboo and 0.0151 for coarse bamboo. No significant differences were observed between the three sites in the acidic Banyupahit–Banyuputih river with k values of 0.0034–0.0066 for fine teak, 0.0002–0.0057 for coarse teak, 0.0029–0.0054 for fine bamboo and 0.0000–0.0068 for coarse bamboo. Moreover, no clear adaptation of macroinvertebrates or microbes to low pH conditions could be detected. The coarse mesh leaf packs in the neutral Kali Sengon river revealed that macroinvertebrates are important in the breakdown process. Fine mesh packs revealed that microbial activity is depressed under acidic conditions. Based on this evidence, we conclude that the toxicity at low pH conditions, and probably also the precipitation of metals on the leaf material, seriously affects leaf decomposition.     

Fungi on Leaf Blades of <Emphasis Type="Italic">Phragmites australis</Emphasis> in a Brackish Tidal Marsh: Diversity,Succession, and Leaf Decomposition

Although fungi are known to colonize and decompose plant tissues in various environments, there is scanty information on fungal communities on wetland plants, their relation to microhabitat conditions, and their link to plant litter decomposition. We examined fungal diversity and succession on Phragmites australis leaves both attached to standing shoots and decaying in the litter layer of a brackish tidal marsh. Additionally, we followed changes in fungal biomass (ergosterol), leaf nitrogen dynamics, and litter mass loss on the sediment surface of the marsh. Thirty-five fungal taxa were recorded by direct observation of sporulation structures. Detrended correspondence analysis and cluster analysis revealed distinct communities of fungi sporulating in the three microhabitats examined (middle canopy, top canopy, and litter layer), and indicator species analysis identified a total of seven taxa characteristic of the identified subcommunities. High fungal biomass developed in decaying leaf blades attached to standing shoots, with a maximum ergosterol concentration of 548 ± 83 μg g^–1 ash-free dry mass (AFDM; mean ± SD). When dead leaves were incorporated in the litter layer on the marsh surface, fungi experienced a sharp decline in biomass (to 191 ± 60 μg ergosterol g^–1 AFDM) and in the number of sporulation structures. Following a lag phase, species not previously detected began to sporulate. Leaves placed in litter bags on the sediment surface lost 50% of their initial AFDM within 7 months (k = −0.0035 day^–1) and only 21% of the original AFDM was left after 11 months. Fungal biomass accounted for up to 34 ± 7% of the total N in dead leaf blades on standing shoots, but to only 10 ± 4% in the litter layer. These data suggest that fungi are instrumental in N retention and leaf mass loss during leaf senescence and early aerial decay. However, during decomposition on the marsh surface, the importance of living fungal mass appears to diminish, particularly in N retention, although a significant fraction of total detrital N may remain associated with dead hyphae.     

Leaf breakdown in streams differing in catchment land use

1. The impact of changes in land use on stream ecosystem function is poorly understood. We studied leaf breakdown, a fundamental process of stream ecosystems, in streams that represent a range of catchment land use in the Piedmont physiographic province of the south‐eastern United States. 2. We placed bags of chalk maple (Acer barbatum) leaves in similar‐sized streams in 12 catchments of differing dominant land use: four forested, three agricultural, two suburban and three urban catchments. We measured leaf mass, invertebrate abundance and fungal biomass in leaf bags over time. 3. Leaves decayed significantly faster in agricultural (0.0465 day^?1) and urban (0.0474 day^?1) streams than in suburban (0.0173 day^?1) and forested (0.0100 day^?1) streams. Additionally, breakdown rates in the agricultural and urban streams were among the fastest reported for deciduous leaves in any stream. Nutrient concentrations in agricultural streams were significantly higher than in any other land‐use type. Fungal biomass associated with leaves was significantly lower in urban streams; while shredder abundance in leaf bags was significantly higher in forested and agricultural streams than in suburban and urban streams. Storm runoff was significantly higher in urban and suburban catchments that had higher impervious surface cover than forested or agricultural catchments. 4. We propose that processes accelerating leaf breakdown in agricultural and urban streams were not the same: faster breakdown in agricultural streams was due to increased biological activity as a result of nutrient enrichment, whereas faster breakdown in urban streams was a result of physical fragmentation resulting from higher storm runoff.     

The Estimated Impact of Fungi on Nutrient Dynamics During Decomposition of Phragmites australis Leaf Sheaths and Stems

Decomposition of culms (sheaths and stems) of the emergent macrophyte Phragmites australis (common reed) was followed for 16 months in the litter layer of a brackish tidal marsh along the river Scheldt (the Netherlands). Stems and leaf sheaths were separately analyzed for mass loss, litter-associated fungal biomass (ergosterol), nutrient (N and P), and cell wall polymer concentrations (cellulose and lignin). The role of fungal biomass in litter nutrient dynamics was evaluated by estimating nutrient incorporation within the living fungal mass. After 1 year of standing stem decay, substantial fungal colonization was found. This corresponded to an overall fungal biomass of 49 ± 8.7 mg g⁻¹ dry mass. A vertical pattern of fungal colonization on stems in the canopy is suggested. The litter bag experiment showed that mass loss of stems was negligible during the first 6 months, whereas leaf sheaths lost almost 50% of their initial mass during that time. Exponential breakdown rates were −0.0039 ± 0.0004 and −0.0026 ± 0.0003 day⁻¹ for leaf sheaths and stems, respectively (excluding the initial lag period). In contrast to the stem tissue—which had no fungal colonization—leaf sheaths were heavily colonized by fungi (93 ± 10 mg fungal biomass g⁻¹ dry mass) prior to placement in the litter layer. Once being on the sediment surface, 30% of leaf sheath's associated fungal biomass was lost, but ergosterol concentrations recovered the following months. In the stems, fungal biomass increased steadily after an initial lag period to reach a maximal biomass of about 120 mg fungal biomass g⁻¹ dry mass for both plant parts at the end of the experiment. Fungal colonizers are considered to contain an important fraction of nutrients within the decaying plant matter. Fungal N incorporation was estimated to be 64 ± 13 and 102 ± 15% of total available N pool during decomposition for leaf sheaths and stems, respectively. Fungal P incorporation was estimated to be 37 ± 9 and 52 ± 15% of total available P during decomposition for leaf sheaths and stems, respectively. Furthermore, within the stem tissue, fungi are suggested to be active immobilizers of nutrients from the external environment because fungi were often estimated to contain more than 100% of the original nutrient stock.     

Decomposition of Fire Exposed Eucalyptus Leaves in a Portuguese Lowland Stream

We compared fire exposed with normal abscised eucalyptus leaves incubated in a stream running through eucalyptus plantations in central Portugal, in terms of breakdown rates, microbial activity, diversity and macroinvertebrate abundance. Although leaves exposed to fire had lower nutritional value, mass loss was similar for both leaf types (k = 0.0089–0.0095 d^–1 for fire and k = 0.0084–0.00103 d^–1 for normal leaves). Fungal biomass was similar among treatments, whereas sporulation and microbial respiration were lower in fire exposed leaves. Both leaf types had similar aquatic hyphomycetes communities. Physical fragmentation was important in fire exposed leaves breakdown. Invertebrates colonized leaves in low numbers in both treatments. Alteration of leaf litter quality determined by fires in streams does not seem to determine changes in ecosystem functioning in a short term. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Decomposition of aboveground and belowground organs of wild rice (Zizania latifolia): mass loss and nutrient changes

Decomposition of aboveground and belowground organs of the emergent macrophyte Z.latifolia was investigated using a litterbag technique for a period of 359 days in a freshwater marsh in Japan. Aboveground parts were classified into: leaves, sheaths and stems. Belowground parts were classified into: horizontal rhizomes (new rhizome, hard rhizome, soft rhizome) and vertical rhizome (stembase). The decay rate (k) was 0.0036 day⁻¹, 0.0033 day⁻¹ and 0.0021 day⁻¹ for leaves, sheaths and stems, respectively. For belowground parts, the decay rate varied considerably from 0.0018 day⁻¹ to 0.0079 day⁻¹, according to differences in the initial chemical compositions of rhizomes. After 359 days of decay, new rhizomes lost 94% of their original dry mass, compared with a loss of 48–84% for the other rhizomes. There was a significant positive relationship between litter quality and decay rate for horizontal rhizomes. For the new rhizomes, which had an internal nitrogen content of 24.2 mg N g⁻¹ dry mass, the mass loss was 40% higher than that of soft rhizomes, which had an internal N content of 9.8 mg N g⁻¹ dry mass. Over the period of 359 days, the nitrogen concentration in all rhizome types decreased to levels lower than initial values, but the phosphorus concentration remained constant after an initial leaching loss. Most nitrogen and phosphorus were lost during the first 45 days of decay. Changes in carbon to nitrogen (C:N) and carbon to phosphorus (C:P) ratios basically followed inversed trends of the nitrogen and phosphorus concentrations.     

Effects of litter patch area on macroinvertebrate assemblage structure and leaf breakdown in Central Amazonian streams

Leaf breakdown in streams is affected by several factors, such as leaf characteristics, water chemistry, microbial activity, and abundance of shredders. In turn, shredders may be resource-controlled. We hypothesized that the size of litter patches affects leaf breakdown, because large patches should be stable over time and therefore harbor high densities of shredders. We selected litter patches (area 0.25–10 m²) in 10 pools of three first-order streams (Manaus, Brazil). We installed 10 leaf packs of Mabea speciosa (Euphorbiaceae) in each patch, and sampled one after 1 day and three after 5, 19, and 28 days. The leaf packs were quickly colonized by the shredding caddisflies Triplectides and Phylloicus. The leaf breakdown rate (mean k = 0.026 ± 0.0015 SE) was high and similar to values reported for other tropical and temperate streams, although much higher than values reported for the adjacent Cerrado biome. Assemblage composition varied over time, but was not related to the size of litter patches. Contrary to our hypothesis, litter patch area did not affect breakdown rates (r ² = 0.012, P = 0.766) or abundance of shredders after 5, 19, and 28 days (r ² < 0.243, P > 0.147). We found, however, a positive relationship between the abundance of tropical shredders and leaf breakdown after 19 days (r ² = 0.572, P = 0.011), suggesting that shredders play an important role in leaf breakdown in these headwater streams. Our study indicates that leaf breakdown rates in tropical streams are variable and can be as high as those of temperate streams.     

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.     

Do tadpoles affect leaf decomposition in neotropical streams?

1. Of the relatively few studies that have examined consequences of amphibian declines on stream ecosystems, virtually all have focused on changes in algae (or algal‐based food webs) and little is known about the potential effects of tadpoles on leaf decomposition. We compared leaf litter decomposition dynamics in two neotropical streams: one with an intact community of tadpoles (with frogs) and one where tadpoles were absent (frogless) as a result of a fungal pathogen that had driven amphibians locally extinct. The stream with tadpoles contained a diverse assemblage (23 species) of larval anurans, and we identified five species of glass frog (Centrolenidae) tadpoles that were patchily distributed but commonly associated with leaf detritus and organic sediments in pools. The latter reached total densities of 0–318 tadpoles m^?2. 2. We experimentally excluded tadpoles from single‐species leaf packs incubated over a 40‐day period in streams with and without frogs. We predicted that decomposition rates would be higher in control (allowing access of tadpoles) treatments in the study stream with frogs than in the frogless stream and, in the stream with frogs, in the control than in the tadpole exclusion treatment. 3. In the stream with frogs, Centrolene prosoblepon and Cochranella albomaculata tadpoles were patchily distributed in leaf packs (0.0–33.3 m^?2). In contrast to our predictions, leaf mass loss and temperature‐corrected leaf decomposition rates in control treatments were almost identical in our stream with frogs (41.01% AFDM lost, k_{degree day} = ?0.028 day^?1) and in the frogless stream (41.81% AFDM lost, k_{degree day} = ?0.027 day^?1) and between control and tadpole exclusion treatments within each stream. Similarly, there were no significant differences in leaf pack bacterial biomass, microbial respiration rates or macroinvertebrate abundance between treatments or streams. Invertebrate assemblages on leaf packs were similar between treatments (SIMI = 0.97) and streams (SIMI = 0.95) and were dominated by larval Chironomidae, Simuliidae (Diptera) and larval Anchytarsus spp. (Coleoptera). 4. In contrast to dramatic effects of grazing tadpoles on algal communities observed previously, tadpoles had no major effects on decomposition. While centrolenid tadpoles were common in the stream with frogs, their patchy distribution in both experimental and natural leaf packs suggests that their effects on detrital dynamics and microbes are probably more localised than those of grazing tadpoles on algae.     

Litter decomposition in a Cerrado savannah stream is retarded by leaf toughness, low dissolved nutrients and a low density of shredders

1. To assess whether the reported slow breakdown of litter in tropical Cerrado streams is due to local environmental conditions or to the intrinsic leaf characteristics of local plant species, we compared the breakdown of leaves from Protium brasiliense, a riparian species of Cerrado (Brazilian savannah), in a local and a temperate stream. The experiment was carried out at the time of the highest litter fall in the two locations. An additional summer experiment was conducted in the temperate stream to provide for similar temperature conditions. 2. The breakdown rates (k) of P. brasiliense leaves in the tropical Cerrado stream ranged from 0.0001 to 0.0008 day⁻¹ and are among the slowest reported. They were significantly (F = 20.12, P < 0.05) lower than in the temperate stream (0.0046–0.0055). The maximum ergosterol content in decomposing leaves in the tropical Cerrado stream was 106 μg g⁻¹, (1.9% of leaf mass) measured by day 75, which was lower than in the temperate stream where maximum ergosterol content of 522 μg g⁻¹ (9.5% of leaf mass) was achieved by day 30. The ATP content, as an indicator of total microbial biomass, was up to four times higher in the tropical Cerrado than in the temperate stream (194.0 versus 49.4 nmoles g⁻¹). 3. Unlike in the temperate stream, leaves in the tropical Cerrado were not colonised by shredder invertebrates. However, in none of the experiments did leaves exposed (coarse mesh bags) and unexposed (fine mesh bags) to invertebrates differ in breakdown rates (F = 1.15, P > 0.05), indicating that invertebrates were unable to feed on decomposing P. brasiliense leaves. 4. We conclude that the slow breakdown of P. brasiliense leaves in the tropical Cerrado stream was because of the low nutrient content in the water, particularly nitrate (0.05 mgN L⁻¹), which slows down fungal activity and to the low density of invertebrates capable of using these hard leaves as an energy source.     

Decomposition of dominant submerged macrophytes: implications for nutrient release in Myall Lake, NSW, Australia

Breakdown and nutrient dynamics of submerged macrophytes were studied in Myall Lake, Australia. Mass loss of Myriophyllum sulsagineum was the lowest (64.90%) among the studied macrophytes during the 322 days followed by charophytes (60.79%), whereas Najas marina and Vallisneria gigantea lost 91.15 and 86.02% of their respective initial mass during that time. The overall exponential breakdown rates of Najas marina and Vallisneria gigantea were similar, with k-values of 0.24 and 0.23 day⁻¹, respectively. These rates were significantly higher than the break down rates of charophytes (0.007 day⁻¹) and M. sulsagineum (0.008 day⁻¹). During growth phase, water column depicted lower nutrient concentrations while during decay period, significant increase in water column nutrients resulted. Release of nutrients from decomposing macrophytes and incorporation of these nutrients into sedimentary phase as well as uptake of nutrients by the growing macrophytes, can present a considerable cycling pathway of nutrients in Myall lake system. The results of this study suggest that different submerged macrophytes may differ appreciably in quality and may exhibit different decomposition rates, patterns and nutrient dynamics in aquatic ecosystems in general, and Myall lakes in particular.     

The relation between above- and belowground biomass allocation patterns and competitive ability

Summary In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m⁻² yr⁻¹). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.     

Relative Importance of Bacteria and Fungi in a Tropical Headwater Stream: Leaf Decomposition and Invertebrate Feeding Preference

Bacteria and fungi provide critical links between leaf detritus and higher trophic levels in forested headwater food webs, but these links in tropical streams are not well understood. We compared the roles of bacteria and fungi in the leaf decomposition process and determining feeding preference for two species of freshwater shrimp found in the Luquillo Experimental Forest, Puerto Rico, using experimental microcosms. We first tested the effects of four treatments on decomposition rates for leaves from two common riparian species, Cecropia scheberiana (Moraceae) and Dacryodes excelsa (Burseraceae), in laboratory microcosms. Treatments were designed to alter the microbial community by minimizing the presence of bacteria or fungi. The fastest decay rate was the control treatment for D. excelsa where both bacteria and fungi were present (k = −0.0073 day⁻¹) compared to the next fastest rate of k = −0.0063 day⁻¹ for the bacterial-conditioned D. excelsa leaves. The fastest decay rate for C. scheberiana was also the control treatment (k = −0.0035 day⁻¹), while the next fastest rate was for fungal-conditioned leaves (k = −0.0029 day⁻¹). The nonadditive effect for leaf decomposition rates observed in the control treatments where both fungi and bacteria were present indicate that bacteria and fungi perform different functions in processing leaf litter. Additionally, leaf types differed in microbial colonization patterns. We next tested feeding preference for leaf type and microbe treatment in microcosms using two species of freshwater shrimp: Xiphocaris elongata, a shredder, and Atya lanipes, a scraper/filterer. To estimate feeding preferences of individual shrimp, we measured change in leaf surface area and the amount of particles generated during 5-day trials in 16 different two-choice combinations. X. elongata preferred D. excelsa over C. scheberiana, and leaves with microbial conditioning over leaves without conditioning. There was no clear preference for fungal-conditioned leaves over bacterial-conditioned leaves. This lack of preference for which microbes were responsible for the conditioning demonstrates the importance of both bacterial and fungal resources in these tropical stream food web studies.     

What determines interspecific variation in relative growth rate of <Emphasis Type="Italic">Eucalyptus</Emphasis> seedlings?

The present study examines relative growth rate (RGR) and its determinants in seedlings of nine Eucalyptus species. Species were selected from mesic (1,800 mm a⁻¹ rainfall) through to semi-arid habitats (300 mm a⁻¹), and thus, notionally vary in “stress” tolerance. Seedlings were grown in a glasshouse during early summer and received between 33 mol and 41 mol PAR m⁻² day⁻¹ . The mean RGR varied among species—from a minimum of 66 mg g⁻¹ day⁻¹ in E. hypochlamydea to a maximum of 106 mg g⁻¹ day⁻¹ in E. delegatensis. RGR was positively related to rainfall at the sites of seed collection. Neither specific leaf area (SLA) nor net assimilation rate was related to rainfall or RGR. While the absence of relationships with SLA and net assimilation rate contrasts with other studies and species, we cannot rule out the effects of sample size (n=9 species) and modest ranges in SLA and RGR. The ratio of leaf mass to total mass (LMR) varied from 0.49±0.07 g g⁻¹ in E. socialis to 0.74±0.04 g g⁻¹ in E. delegatensis and was strongly positively related with rainfall (r ²=0.77). Interspecific differences in RGR were strongly related to LMR (positive relationship, r ²=0.50) and the rate of dry matter production per mol of leaf nitrogen (positive relationship, r ²=0.64). Hence, the slow RGR of low-rainfall species was functionally related to a lower growth rate per mol of leaf nitrogen than high-rainfall species. Furthermore, slow RGR of low-rainfall species was related to greater allocation to roots at the expense of leaves. Increasing allocation to roots versus leaves is likely an adaptation to soil and atmospheric water deficits, but one that comes at the expense of a slow RGR.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

Quantitative analysis of carbon balance in the reproductive organs and leaves of <Emphasis Type="Italic">Cinnamomum camphora</Emphasis> (L.) Presl

We investigated seasonal changes in dry mass and CO₂ exchange rate in fruit and leaves of the evergreen tree Cinnamomum camphora with the aim of quantitatively determining the translocation balance between the two organs. The fruit dry mass growth peaked in both August and October: the first increase was due to fruit pulp development and the second to seed development. Fruit respiration also increased with the rapid increase in fruit dry mass. Therefore, the carbohydrates required for fruit development showed two peaks during the reproductive period. Fruit photosynthesis was relatively high in early August, when fruit potentially re-fixed 75% of respired CO₂, indicating that fruit photosynthesis contributed 15–35% of the carbon requirement for fruit respiration. Current-year leaves completed their growth in June when fruit growth began. Current-year leaves translocated carbohydrates at a rate of approximately 10–25 mg dry weight (dw) leaf⁻¹ day⁻¹ into other organs throughout the entire fruit growth period. This rate of translocation from current-year leaves was much higher than the amount of carbohydrate required for reproduction (ca. 3 mg dw fruit⁻¹ day⁻¹). Given the carbon balance between fruit and current-year leaves, carbohydrates for reproduction were produced within the current-year fruit-bearing shoots. C. camphora would be adaptive for steadily supplying enough amount of carbohydrate to the fruits, as there was little competition for carbohydrates between the two organs. As assimilates by leaves are used for processes such as reproduction and the formation of new shoots, photosynthesis by reproductive organs is considered to be important to compensate for reproductive cost.