Leaf litter decomposition and macroinvertebrate communities in headwater streams draining pine and hardwood catchments

Benthic invertebrates, litter decomposition, andlitterbag invertebrates were examined in streamsdraining pine monoculture and undisturbed hardwoodcatchments at the Coweeta Hydrologic Laboratory in thesouthern Appalachian Mountains, USA. Bimonthlybenthic samples were collected from a stream draininga pine catchment at Coweeta during 1992, and comparedto previously collected (1989–1990) benthic data froma stream draining an adjacent hardwood catchment. Litter decomposition and litterbag invertebrates wereexamined by placing litterbags filled with pine ormaple litter in streams draining pine catchments andhardwood catchments during 1992–1993 and 1993–1994. Total benthic invertebrate abundance and biomass inthe pine stream was ca. 57% and 74% that of thehardwood stream, respectively. Shredder biomass wasalso lower in the pine stream but, as a result ofhigher Leuctra spp. abundance, shredderabundance was higher in the pine stream than thehardwood stream. Decomposition rates of both pine andred maple litter were significantly faster in pinestreams than adjacent hardwood streams (p<0.05). Total shredder abundance, biomass, and production weresimilar in maple bags from pine and hardwood streams. However, trichopteran shredder abundance and biomass,and production of some trichopteran taxa such asLepidostoma spp., were significantly higher in maplelitterbags from pine streams than hardwood streams(p<0.05). In contrast, plecopteran shredders(mainly Tallaperla sp.) were more important inmaple litterbags from hardwood streams. Shredderswere well represented in pine litterbags from pinestreams, but low shredder values were obtained frompine litterbags in hardwood streams. Resultssuggest conversion of hardwood forest to pinemonoculture influences taxonomic composition of streaminvertebrates and litter decomposition dynamics. Although the impact of this landscape-leveldisturbance on invertebrate shredder communitiesappeared somewhat subtle, significant differences indecomposition dynamics indicate vital ecosystem-levelprocesses are altered in streams draining pinecatchments.     

Land use change affects macroinvertebrate community size spectrum in streams: the case of Pinus radiata plantations

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An Unexpected Nitrate Decline in New Hampshire Streams

Theories of forest nitrogen (N) cycling suggest that stream N losses should increase in response to chronic elevated N deposition and as forest nutrient requirements decline with age. The latter theory was supported initially by measurements of stream NO₃ ⁻ concentration in old-growth and successional stands on Mount Moosilauke, New Hampshire (Vitousek and Reiners 1975; Bioscience 25:376–381). We resampled 28 of these and related streams to evaluate their response to 23 years of forest aggradation and chronic N deposition. Between 1973–74 and 1996–97, mean NO₃ ⁻ concentration in quarterly samples from Mount Moosilauke decreased by 71% (25 μmol/L), Ca²⁺ decreased by 24% (8 μmol/L), and Mg²⁺ decreased by 22% (5 μmol/L). Nitrate concentrations decreased in every stream in every season, but spatial patterns among streams persisted: Streams draining old-growth stands maintained higher NO₃ ⁻ concentrations than those draining successional stands. The cause of the NO₃ ⁻ decline is not evident. Nitrogen deposition has changed little, and although mechanisms such as insect defoliation and soil frost may contribute to the temporal patterns of nitrate loss, they do not appear to fully explain the NO₃ ⁻ decline across the region. Although the role of climate remains uncertain, interannual climate variation and its effects on biotic N retention may be responsible for the synchronous decrease in NO₃ ⁻ across all streams, overriding expected increases due to chronic N deposition and forest aging. Received 4 December 2001; accepted 30 May 2002.     

Phosphorus retention in streams draining pine and hardwood catchments in the southern Appalachian Mountains

SUMMARY. 1. This study was designed to determine how catchment use affects stream phosphorus retention by comparing retention in streams draining three mixed hardwood catchments and three catchments that were planted in white pine in the 1950s.
2. Catchments of similar area and stream discharge were chosen and phosphorus uptake was measured monthly in each catchment along with temperature, discharge, velocity, coarse particulate organic matter (CPOM), fine particulate organic matter (FPOM), and microbial respiration associated with FPOM.
3. On an annual basis, average phosphorus retention was not different between streams draining pine and hardwood catchments nor were there significant differences between physical (temperature, velocity and discharge) or biological (CPOM, FPOM and respiration) parameters based on catchment type. However, discharge was more variable in streams draining pine catchments.
4. Because phosphorus uptake was correlated with discharge, phosphorus retention was also more variable in streams draining pine catchments. Storms caused a greater increar.e in discharge and loss of phosphorus in pine streams than in mixed hardwood streams, but discharge returned to baseline more quickly in pine streams.
5. We suggest that discharge regimes and phosphorus dynamics of streams draining pine catchments are less resistant to change but more resilient than streams draining mixed hardwood forests.     

Beaver invasion alters terrestrial subsidies to subantarctic stream food webs

North American beavers (Castor canadensis) were introduced to Tierra del Fuego Island in 1946 for their fur, and have since spread across the archipelago and onto the South American mainland. We assessed the impact of invasive beavers on streams of these forested watersheds by quantifying the trophic basis of production (TBP) and consumptive organic matter flows of benthic macroinvertebrate assemblages. TBP was determined in two streams: clear- and black-water. Stable isotopes were used across four streams to further elucidate food web structure and dominant pathways. TBP and stable isotopes showed that terrestrially derived organic matter (amorphous detritus, leaves, and wood) supported a majority of secondary production in the benthic food webs at all sites (forested reaches, beaver ponds, and sections downstream of ponds with foraged riparian zones). The magnitude of these flows was enhanced in beaver-modified sites compared with forested habitats (4.0–5.3× increase g AFDM m⁻² year⁻¹ in pond habitats, 1.1–2.1× increase in downstream habitats). Diatoms were the only autochthonous resource identified in macroinvertebrate guts, but their contribution to secondary production was small. Consumptive flows mirrored trends in TBP (i.e., dominance of terrestrial sources and greater magnitude in beaver ponds). Collector–gatherer consumption of amorphous detrital material dominated food web flows in all habitats, but was higher in beaver ponds relative to other habitats. Food web structure was simplified in beaver ponds; only two of the five possible functional groups contributed >1% of total organic matter flow in ponds (collector–gatherers and predators). Consumptive flows to predators increased in ponds, and stable isotopes of nitrogen and carbon (δ¹⁵N and δ¹³C) corroborated a relatively greater importance of predators (greater trophic distance), as well as less diversity of basal resources (less variation in δ¹³C) in ponds. Our findings indicate that invasive beaver’s engineering activities resulted in greater flows of terrestrial organic matter subsidies to in-stream food webs, which had a relatively greater change in the clear-water than in the black-water stream. Owing to the fact that these streams were naturally dependent on allochthonous resources for a majority of production and material flows, changes wrought by beavers to streams in forested environments are probably less than in watersheds with inherently greater dependence on autochthonous production such as the adjacent steppe biome.     

Effects of nutrient enrichment on C and N stable isotope ratios of invertebrates,fish and their food resources in boreal streams

Carbon and nitrogen stable isotopes are frequently used to study energy sources and food web structure in ecosystems, and more recently, to study the effects of anthropogenic stress on aquatic ecosystems. We investigated the effect of nutrient enrichment on δ¹³C and δ¹⁵N in fine (FPOM), coarse (CPOM) particulate organic matter, periphyton, invertebrates and fish in nine boreal streams in south-central Sweden. In addition, we analysed the diet of benthic consumers using stable isotope data. Increases in δ¹⁵N of periphyton (R ² = 0.88), CPOM (0.78), invertebrates (0.92) and fish (0.89) were related to nutrient enrichment. In contrast, δ¹³C signatures did not change along the nutrient gradient. Our results show that δ¹⁵N has potential as a sensitive indicator of nutrient enrichment in boreal streams. Carbon and nitrogen isotopes failed to elucidate putative diets of selected aquatic consumers. Indeed, comparison of low- and high-impact sites showed that δ¹³C of many consumers were found outside the ranges of basal resource δ¹³C. Moreover, ranges of basal resource δ¹³C and δ¹⁵N overlapped at both low and high sites, making discrimination between the importance of allochthonous and autochthonous production difficult. Our findings show that a fractionation rate of 3.4‰ is not always be appropriate to assess trophic interactions, suggesting that more studies are needed on fractionation rates along gradients of impairment. Handling editor: M. Power     

The effects of land use on environmental features and functional organization of macroinvertebrate communities in Patagonian low order streams

Benthic invertebrates, water quality variables, chlorophyll a and particulate organic matter (POM) were studied in 18 sites of mountain streams in Patagonia (Argentina) subjected to six different land uses: native forest, pine plantation, pasture, harvest forest, urban and reference urban. Three streams of each land use were studied in March 2006. Macroinvertebrates were sampled in three riffles and three pools (n = 108) and biomass of detrital fractions of POM were quantified. Overall benthic POM biomass was higher at native and harvest forest than pastures, whereas fine fraction (FPOM) was higher in harvest forest than in pastures. Regarding to autotrophic subsidies bryophytes were the only that changed consistently among uses. These differences in energy resources were correlated with changes in community attributes. Shredder richness was clearly higher at native and harvest forest than exotic pine plantations, collector gatherers density was consistently high at harvest sites, and total density was significantly higher at urban and harvest forest. Multidimensional scaling ordination based on macroinvertebrate density data showed a clear separation of forested (either native or exotic species) from riparian modified areas (pasture, urban and harvest sites). Environmental variables having explanation power on macroinvertebrate assemblages were mostly related with: detritus availability (wood and leaves biomass) and impairment (total phosphorous and % sand). These results confirm that macroinvertebrate assemblage structure in Patagonian low order streams can be altered by land use practices. Among guild structure measures, those indicators based on benthic community functional attributes, shredders richness and collectors density, resulted good candidates to assess land use impacts. On account of riparian corridor management may be critical to the distribution of benthic taxa, the maintenance of good conditions of vegetation adjacent to rivers will enhance water quality and the environment for highly endemic headwater communities of Patagonian streams.     

Modelling Inter- and Intra-specific Competition in Loblolly Pine (Pinus taeda L.) Plantations on Cutover, Site-prepared Lands

The presence of volunteer hardwood species in loblolly pineplantations demands studies on both intra- and inter-specificcompetition in order to build growth and yield models and toguide vegetation management for these stands. This paper, basedon analyses of data collected from a thinning study, reportsan investigation of responses of loblolly pine and hardwoodspecies towards intra- and inter-specific competition. Underhigh levels of overall competition, hardwood species were morecompetitive, both intra- and inter-specifically. Intra-specificcompetition was more effective in reducing hardwood basal areagrowth than inter-specific competition under high levels ofoverall competition. However, under low levels of overall competition,intra- and inter-specific competition were quantitatively similartoward loblolly pine basal area growth. The basal area growthof hardwoods was significantly related to levels of inter-specificcompetition, but not with intra-specific effects under low levelsof overall competition. More than half of the variability inloblolly pine basal area growth under unthinned control andlight thin treatments could be accounted for by competitioneffects. However, only one third of the variability could beexplained for loblolly pine under heavy thin treatment. Forhardwood species, the percentage of growth variation accountedfor by competition was about 45%, and did not change among thethinning treatments. Different resource demands between thetwo categories of tree species and a certain amount of thinningshock were suggested by the modelling results.Copyright 1994,1999 Academic Press Neighbourhood approach, competition index, hardwood, no-interaction model, basal area, reciprocal yield law     

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH₄NO₃) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m^-2 y^-1), high N (15 g N m^-2 y^-1), and low N + sulfur (5 g N m^-2 y^-1 plus 7.4 g S m^-2 y^-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of ¹⁵N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996. Received 2 April 1999; Accepted 29 October 1999.     

Land Use Influences Niche Size and the Assimilation of Resources by Benthic Macroinvertebrates in Tropical Headwater Streams

It is well recognized that assemblage structure of stream macroinvertebrates changes with alterations in catchment or local land use. Our objective was to understand how the trophic ecology of benthic macroinvertebrate assemblages responds to land use changes in tropical streams. We used the isotope methodology to assess how energy flow and trophic relations among macroinvertebrates were affected in environments affected by different land uses (natural cover, pasture, sugar cane plantation). Macroinvertebrates were sampled and categorized into functional feeding groups, and available trophic resources were sampled and evaluated for the isotopic composition of ¹³C and ¹⁵N along streams located in the Cerrado (neotropical savanna). Streams altered by pasture or sugar cane had wider and more overlapped trophic niches, which corresponded to more generalist feeding habits. In contrast, trophic groups in streams with native vegetation had narrower trophic niches with smaller overlaps, suggesting greater specialization. Pasture sites had greater ranges of resources exploited, indicating higher trophic diversity than sites with natural cover and sugar cane plantation. We conclude that agricultural land uses appears to alter the food base and shift macroinvertebrate assemblages towards more generalist feeding behaviors and greater overlap of the trophic niches.     

Understory development in canopy gaps of pine and pine-hardwood forests of the upper Coastal Plain of Virginia

Canopygaps are important in establishing a pool of natural regeneration in manytemperate forest ecosystems. Information on the role of gaps in loblolly pine(Pinus taeda L.) and pine-hardwood foreststands in the southeastern Coastal Plain of the United States is lacking.Accordingly, 12 small canopy gaps in mature pine and pine-hardwood standsin Petersburg National Battlefield, Virginia, were studied. Loblolly pineregeneration was significantly more abundant in canopy gaps as compared to theadjacent forest in both forest cover types. In four stands dominated by loblollypine, there were 750 saplings/ha in the gaps compared to only 125saplings/ha in the adjacent forest. Pine saplings dominated the regenerationspectrum in the gaps in the pine stands, while red maple (Acerrubrum L.) was more important in the adjacent forest. In fourpine-hardwood stands, regeneration in both the gaps and adjacent forestwas dominated by sweetgum (Liquidambar styracifluaL.) with importance values of 27% and 28%, respectively.There were no loblolly pine seedlings in the adjacent forest, but an average 313per ha in the gaps of the pine-hardwood stands. Within thegaps in both cover types, loblolly pine saplings were lower in stature thancompeting hardwood stems, leading to the conclusion that the gaps may form atemporary pool of pine regeneration. Without further stand disturbance, theprocess of gap closure may reduce the pine component to a secondary status, orperhaps eliminate it altogether.     

Natural 15N abundance in two nitrogen saturated forest ecosystems

Natural ¹⁵N abundance values were measured in needles, twigs, wood, soil, bulk precipitation, throughfall and soil water in a Douglas fir (Pseudotsuga menziesii (Mirb.) and a Scots pine (Pinus sylvestris L.) stand receiving high loads of nitrogen in throughfall (>50 kg N ha⁻¹ year⁻¹). In the Douglas fir stand δ¹⁵N values of the vegetation ranged between −5.7 and −4.2‰ with little variation between different compartments. The vegetation of the Scots pine stand was less depleted in ¹⁵N and varied from −3.3 to −1.2‰δ¹⁵N. At both sites δ¹⁵N values increased with soil depth, from −5.7‰ and −1.2‰ in the organic layer to +4.1‰ and +4.7‰ at 70 cm soil depth in the Douglas fir and Scots pine stand, respectively. The δ¹⁵N values of inorganic nitrogen in bulk precipitation showed a seasonal variation with a mean in NH₄ ⁺-N of −0.6‰ at the Douglas fir stand and +10.8‰ at the Scots pine stand. In soil water below the organic layer NH₄ ⁺-N was enriched and NO₃ ⁻-N depleted in ¹⁵N, which was interpreted as being caused by isotope fractionation accompanying high nitrification rates in the organic layers. Mean δ¹⁵N values of NH₄ ⁺ and NO₃ ⁻ were very similar in the drainage water at 90 cm soil depth at both sites (−7.1 to −3.8‰). A dynamic N cycling model was used to test the sensitivity of the natural abundance values for the amount of N deposition, the ¹⁵N ratio of atmospheric N deposited and for the intrinsic isotope discrimination factors associated with N transformation processes. Simulated δ¹⁵N values for the N saturated ecosystems appeared particularly sensitive to the ¹⁵N ratio of atmospheric N inputs and discrimination factors during nitrification and mineralization. The N-saturated coniferous forest ecosystems studied were not characterized by elevated natural ¹⁵N abundance values. The results indicated that the natural ¹⁵N abundance values can only be used as indicators for the stage of nitrogen saturation of an ecosystem if the δ¹⁵N values of the deposited N and isotope fractionation factors are taken into consideration. Combining dynamic isotope models and natural ¹⁵N abundance values seems a promising technique for interpreting natural ¹⁵N abundance values found in these forest ecosystems. Received: 5 May 1996 / Accepted: 10 April 1997     

Influence of functional feeding groups and spatiotemporal variables on the δ<Superscript>15</Superscript>N signature of littoral macroinvertebrates

The δ¹⁵N trophic enrichment in littoral food webs is not well known despite the importance of macroinvertebrates in lacustrine energy fluxes. We wanted to assess the influence of functional feeding group (grazer, collector, shredder, predator, predator–hematophagous, predator–sucker) and spatiotemporal variables (year, month, station of sampling) on littoral macroinvertebrate δ¹⁵N signatures. For 2 years, during the plant growth period phytophilous littoral macroinvertebrates were sampled in Lake St. Pierre, a large fluvial lake of the St. Lawrence River, Québec, Canada. The δ¹⁵N analyses showed that station was the most important factor for explaining δ¹⁵N variation, followed by sampling month and functional feeding group. The organisms sampled in the stations of the south shore, which experienced greater macrophyte abundance, slower currents, and stronger NO₃ depletion exhibited higher δ¹⁵N values than those sampled on the north shore. Grazer-to-predator δ¹⁵N enrichment valued 1.6‰, which is inferior to the 3.4‰ generally admitted in food-web research. Shredders exhibited the lowest δ¹⁵N values and predators–hematophagous the highest. δ¹⁵N signature of invertebrates increased 3‰ through the summer between May and September. Only samples collected within a short period should be pooled to avoid an error value equivalent to one trophic level (1.6) enrichment. Furthermore, it is recommended not to pool macroinvertebrate samples collected at stations with differing watershed land uses.     

Age-Dependent Changes in Ecosystem Carbon Fluxes in Managed Forests in Northern Wisconsin,USA

The age-dependent variability of ecosystem carbon (C) fluxes was assessed by measuring the net ecosystem exchange of C (NEE) in five managed forest stands in northern Wisconsin, USA. The study sites ranged in age from 3-year-old clearcut to mature stands (65 years). All stands, except the clearcut, accumulated C over the study period from May to October 2002. Seasonal NEE estimates were −655 ± 17.5 g C m^–2 in the mature hardwood (MHW), −648 ± 16.8 in the mature red pine (MRP), −195 ± 15.6 in the pine barrens (PB), +128 ± 17.1 in the young hardwood clearcut (YHW), and −313 ± 14.6 in the young red pine (YRP). The age-dependent differences were similar in the hardwood and conifer forests. Even though PB was not part of either the hardwood or conifer chronosequence, and had a different disturbance agent, it still fits the same general age relationship. Higher ecosystem respiration (ER) in the young than in the mature stands was the combined result of earlier soil warming in spring, and higher temperature and greater biological activity in summer, as indicated by temperature-normalized respiration rates. The fire-generated PB had lower ER than the harvest-generated YHW and YRP, where high ER was sustained partly on account of logging residue. During the main growing season, the equivalent of 31 (MHW), 48 (MRP), 68 (PB), 114 (YHW) and 71% (YRP) of daily gross ecosystem production (GEP) was released in ER during the same day. The lower ER:GEP ratio in the mature stands was driven by greater age-dependent changes in ER than GEP. The magnitude of the increase in ER:GEP ratio in spring and fall was interpreted as the extent of the decoupling of ER and GEP. Decoupling (sustained high ER despite decreasing GEP) was observed in YHW, PB and MHW, whereas in coniferous stands (MRP and YRP) the stable ER:GEP ratio suggested preferential use of new photosynthates in ER. The results indicate that a great part of the variation in landscape-level C fluxes can be accounted for by mean stand age and associated parameters, which highlights the need to consider this source of heterogeneity in regional C balance estimates.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

Assessment of the effectiveness of best management practices for streams draining agricultural landscapes using diatoms and macroinvertebrates

In this study, a bioassessment was conducted to determine the effectiveness of best management practices (BMPs) implemented in farms in the Upper Delaware River watershed, NY (USA). Diatom and macroinvertebrate communities were analyzed across 17 low-order streams, designated as reference, BMP, or non-BMP. Streams lacking improvements (non-BMP) had significantly greater specific conductance, pH, TDP, NH₄ ⁺-N, and NO₃ ⁻-N than did reference streams. Diatom model affinity (DMA) values were significantly greater in reference and BMP streams than in non-BMP streams; non-BMP streams bordered on a “severely impacted” rating. The Trophic Diatom Index (TDI) varied two-fold among stream classes, with non-BMP > BMP > reference. TDI and DMA values were highly correlated, and both varied significantly with conductance, TDP, NH₄ ⁺-N, and NO₃ ⁻-N. Macroinvertebrate taxa, EPT richness, and Simpson’s diversity did not differ significantly among stream classes. Macroinvertebrate metrics (HBI, Bioassessment Profile, Percent Model Affinity) varied by stream class, but none indicated greater water quality in BMP sites. Nonetheless, each correlated significantly with conductance and TDP in the directions predicted by each model. Our data suggest that diatoms are more sensitive to moderate increases in nutrients, conductivity, and pH in high-gradient agricultural streams, and may be more useful in assessing stream management practices.     

Ecosystem N Distribution and δ<Superscript>15</Superscript>N during a Century of Forest Regrowth after Agricultural Abandonment

Abstract Stable isotope ratios of terrestrial ecosystem nitrogen (N) pools reflect internal processes and input–output balances. Disturbance generally increases N cycling and loss, yet few studies have examined ecosystem δ¹⁵N over a disturbance-recovery sequence. We used a chronosequence approach to examine N distribution and δ¹⁵N during forest regrowth after agricultural abandonment. Site ages ranged from 10 to 115 years, with similar soils, climate, land-use history, and overstory vegetation (white pine Pinus strobus). Foliar N and δ¹⁵N decreased as stands aged, consistent with a progressive tightening of the N cycle during forest regrowth on agricultural lands. Over time, foliar δ¹⁵N became more negative, indicating increased fractionation along the mineralization–mycorrhizal–plant uptake pathway. Total ecosystem N was constant across the chronosequence, but substantial internal N redistribution occurred from the mineral soil to plants and litter over 115 years (>25% of ecosystem N or 1,610 kg ha⁻¹). Temporal trends in soil δ¹⁵N generally reflected a redistribution of depleted N from the mineral soil to the developing O horizon. Although plants and soil δ¹⁵N are coupled over millennial time scales of ecosystem development, our observed divergence between plants and soil suggests that they can be uncoupled during the disturbance-regrowth sequence. The approximate 2‰ decrease in ecosystem δ¹⁵N over the century scale suggests significant incorporation of atmospheric N, which was not detected by traditional ecosystem N accounting. Consideration of temporal trends and disturbance legacies can improve our understanding of the influence of broader factors such as climate or N deposition on ecosystem N balances and δ¹⁵N. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population structure and growth-stress relationship of Pinus taeda in rock outcrop habitats

Abstract. On the granite outcrops of the southeastern United States, soil accumulates in shallow depressions on the rock surface. A specific sequence of vascular plants characterizes the temporal development of these systems. The edaphic end point of the succession is apparently attained with a herb-shrub-tree stage with Pinus taeda as the dominant tree. We studied the characteristics of this stage and the population structure of P. taeda on outcrop islands in order to specify the successional status of the species in this habitat. We compared the radial growth pattern of outcrop and Piedmont populations of P. taeda, and of two outcrop sub-populations. We checked whether trees on outcrops experience more limiting conditions than trees on the Piedmont, and studied the recent change in the relationship between growth and stress (e.g. drought and atmospheric deposition) reported for loblolly pine in the southeastern United States. We also attempted to identify the climatic variables most critical for tree growth on outcrops. On outcrop soil islands, P. taeda maintains populations that are of irregular age distribution, possibly in response to irregular recruitment and survival. There are no signs of loblolly pine replacement by hardwood species on any of the islands studied, although an understory of shrubby oak appears to characterize larger and deeper-soil islands. Although trees on the Piedmont were growing at a higher rate than those on the outcrop in the first part of the 1950–1988 period, their growth declined at a higher rate than that of trees on the outcrop. As a result, at the end of the period considered (1950–1988), the differences in radial growth between outcrop and Piedmont populations were relatively small. On the outcrop, trees < 22 yr old in 1989 were growing at a rate somewhat lower than that of trees of similar age, 40 yr ago. Differences were apparent in the initial growth patterns between the two outcrop subpopulations (1989 and 1949 stems), and these could have resulted from differences in competition regime, stress or climatic conditions, or a combination thereof. However, a repeated-measures ANOVA failed to reveal a significant recent decrease in the radial growth of loblolly pine in the system studied. The identification of numerous signature years (years with ≥ 80 % of the trees with similar increasing or decreasing trend in their radial growth) suggests that similar environmental variables control the growth of loblolly pine in both outcrop and Piedmont habitats. High temperature and low precipitation in the first part of the summer (June-July) seem to limit radial growth. Those pines growing on outcrop soil islands, however, appear more sensitive to climatic fluctuations.     

Stable isotopes as indicators of wastewater effects on the macroinvertebrates of urban rivers

Rivers in urban locations frequently receive contaminated wastewater and particulate waste either directly from storm overflows or from sewage treatment facilities. Although many urban streams are now recovering from wide-scale historic pollution, lower-level effects on water chemistry, nutrients and biotic composition are still widespread. We aimed to determine whether such effects could be detected using stable isotope ratios (??¹⁵N, ??¹³C and ??³⁴S) in macroinvertebrates alone or in conjunction with traditional biomonitoring. Macroinvertebrates were collected upstream and downstream of 11 different secondary wastewater treatment works (WwTW) in South Wales and the Welsh borders (United Kingdom). Overall, mean invertebrate ??¹⁵N signatures downstream of the WwTW were significantly enriched despite variation amongst sites. Moreover, changes between upstream and downstream macroinvertebrate ??¹⁵N values were highly correlated with patterns in macroinvertebrate community composition, increased total macroinvertebrate abundance, and reduced Shannon Diversity and other biomonitoring indices (% EPT, % shredders and ASPT scores). Changes in invertebrate ??¹⁵N values also paralleled the consented discharge volumes and population equivalents from each WwTW. In contrast, isotopic ratios of ??¹³C and ??³⁴S were unable to distinguish or quantify wastewater input into the rivers but differences were apparent amongst study streams. Overall, these results suggest that macroinvertebrate ??¹⁵N signatures can detect and quantify the effects of secondary sewage treatment inputs to riverine ecosystems. Moreover, the method potentially provides a sensitive means for tracing sewage-derived nutrients into food webs while inferring effects on aquatic communities where sewage-loads are subtle or confounded by other stressors.     

Plant and soil natural abundance <Emphasis Type="Italic">δ</Emphasis><Superscript>15</Superscript>N: indicators of relative rates of nitrogen cycling in temperate forest ecosystems

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of δ ¹⁵N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the δ ¹⁵N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil δ ¹⁵N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil δ ¹⁵N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, δ ¹⁵N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground δ ¹⁵N values are a better indicator of relative rates of soil N cycling than are aboveground δ ¹⁵N values.