Bruce L. Haines

Fluxes of Dissolved Organic Nutrients and Humic Substances in a Deciduous Forest

We evaluated the importance of dissolved organic matter as a vehicle for the movement of N and P from the canopy and the forest floor into the mineral soil of a deciduous forest. We also examined the origin and nature of dissolved organic matter from the forest floor to see whether it was simply soluble plant material or highly humified matter. The average annual output from the forest floor in the form of dissolved organic matter was 18, 28, and 14% of the input in solid litterfall for C, N, and P, respectively. In throughfall, about half of the dissolved N and P was organic. But, in solution percolating from the forest floor, 94% of the N and 64% of the P was organic. Leaching from the forest floor was not a source of inorganic N and P for the mineral soil. Instead, the forest floor was a sink for the removal of these inorganic nutrients delivered in throughfall. Microbial immobilization was the most likely explanation for much of the inorganic nutrient removal. In contrast, the forest floor was an abundant contributor of N and P to the mineral soil in the form of dissolved, and possibly particulate, organic matter. Much of the dissolved organic matter entering the A horizon originated from the upper (Oa and Oe horizon) forest floor, but it was modified in several respects compared to the original soluble material. The solution percolating from the forest floor over most of the year was much richer in nitrogen, contained a much larger proportion of hydrophilic acids, and contained a much smaller proportion of carbohydrate-rich hydrophilic neutrals, than did the original water- extractable material in autumn litter. However, the fresh autumn litter did contain a similar proportion of soluble hydrophobic acids that resembled dissolved humic substances in several respects. Most of the flux of nitrogen from the forest floor to the A horizon was carried by humic substances and highly colored hydrophilic acids.

SOIL MICROARTHROPOD CONTRIBUTIONS TO DECOMPOSITION DYNAMICS: TROPICAL–TEMPERATE COMPARISONS OF A SINGLE SUBSTRATE

This study examined the effect of soil microarthropods on the decomposition of a single substrate (Quercus prinus L.) at two humid tropical forests (La Selva, Costa Rica (LAS), and Luquillo Experimental Forest, Puerto Rico (LUQ)) and one temperate forest (Coweeta Hydrologic Station, North Carolina, USA (CWT)). In this litterbag ex- periment, naphthalene was applied to reduce the microarthropod population density from half of three replicate plots established at each site. This enabled us to quantify the mass loss contributed by the fauna (MLCF) at each site and permitted an analysis of the influence of site-specific differences in the composition of the microarthropod assemblages on de- composition rates. We hypothesized that microarthropod regulation of the microbial pop- ulations involved in leaf litter decomposition would be stronger in humid tropical forests, which experience conditions of low climatic variability. In these conditions, there can be an enhanced degree of biotic interactions between microarthropods and their microbial food sources. The elevated extent of these interactions should be expressed as a greater influence of microarthropo ds at the tropical sites and could result in a site-specific effect of faunal assemblages on decomposition . Decomposition of the oak litter proceeded faster in Puerto Rican and Costa Rican forests than in a temperate forest in North Carolina, USA. Microarthropods had little effect on decomposition in the temperate forest, whereas their influence was pronounced at tropical sites. Mass loss of litter from plots with reduced microarthropod populations was similar at the tropical sites. When plots with intact faunal communities were compared, differences in the tropical sites were apparent, suggesting that there was a site-specific faunal contri- bution to decomposition at these sites. Oribatid mites constituted a dominant component (41-64%) at each of the sites. Species richness of oribatids and Fishers alpha diversity were similar in each of the three sites. The Shannon index revealed a lower diversity at LUQ. Abundance of microarthropods was lowest at LAS. Species accumulation curves for each site, though similar in form, were distinctive, as were diversity accumulation patterns in samples of increasing size. There was a positive relationship between species richness and the contribution of the fauna to litter mass loss within each site. Thus, species diversity of decomposer fauna may have important ecosystem consequences, particularly in warm moist tropical forests.

Retention of soluble organic nutrients by a forested ecosystem

We document an example of a forested watershed at the Coweeta HydrologicLaboratory with an extraordinary tendency to retain dissolved organic matter(DOM) generated in large quantities within the ecosystem. Our objectives weretodetermine fluxes of dissolved organic C, N, and P (DOC, DON, DOP,respectively),in water draining through each stratum of the ecosystem and synthesizeinformation on the physicochemical, biological and hydrologic factors leadingtoretention of dissolved organic nutrients in this ecosystem. The ecosystemretained 99.3, 97.3, and 99.0% of water soluble organic C, N and P,respectively, produced in litterfall, throughfall, and root exudates. Exportsinstreamwater were 4.1 kg ha−1yr−1of DOC, 0.191 kg ha−1 yr−1 ofDON, and 0.011 kg ha−1 yr−1 ofDOP. Fluxes of DON were greater than those of inorganic N in all strata. MostDOC, DON, and DOP was removed from solution in the A and B horizons, with DOCbeing rapidly adsorbed to Fe and Al oxyhydroxides, most likely by ligandexchange. DON and DOC were released gradually from the forest floor over theyear. Water soluble organic C produced in litterfall and throughfall had adisjoint distribution of half-decay times with very labile and veryrefractory fractions so that most labile DOC was decomposed before beingleachedinto the mineral soil and refractory fractions dominated the DOC transportedthrough the ecosystem. We hypothesize that this watershed retained solubleorganic nutrients to an extraordinary degree because the soils have very highcontents of Fe and Al oxyhydroxides with high adsorption capacities and becausethe predominant hydrologic pathway is downwards as unsaturated flow through astrongly adsorbing A and B horizon. The well recognized retention mechanismsforinorganic nutrients combine with adsorption of DOM and hydrologic pathway toefficiently prevent leaching of both soluble inorganic andorganic nutrients in this watershed.

Acid rain: Threshold of leaf damage in eight plant species from a Southern Appalachian forest succession

Eight plant species were subjected to artifical acid rains of pH 2.5, 2.0, 1.5, 1.0 and 0.5 in order to determine the threshold for and symptoms of damage. The plants were Erechtites, Robinia, Pinus, Quercus, Carya, Liriodendron, Acer and Corpus from the Coweeta Hydrologic Laboratory near Franklin, North Carolina Droplets of pH 2.0 produced brown necrotic spots on all species except Pinus while droplets of pH 1.0 produced necroses on leaves of all species examined. The size of necrotic spots increased with increasing acidity. Comparison of these results with the literature suggests that developing leaves are more easily damaged than are the mature leaves used in this study. The volume weighted average rainfall pH for Coweeta is 4.6 with observations ranging from 3.2 to 5.9. Results of this study suggest that a 10-fold increase in acidity from pH 3.2 to 2.2 in a single spring or summer storm could bring damage or death to mature leaves of dominant flowering plants in the Southern Appalachians.

Nitrification potentials in early successional black locust and in mixed hardwood forest stands in the southern Appalachians, USA

Soil nitrogen mineralisation and nitrification potentials, and soil solution chemistry were measured in black locust (Robinia pseudo-acacia L.), in pine-mixed hardwood stands on an early successional watershed (WS6), and in an older growth oak-hickory forest located on an adjacent, mixed hardwood watershed (WS14) at Coweeta Hydrologic laboratory, in the southern Appalachian mountains, U.S.A. Nitrification potentials were higher in black locust and pine-mixed hardwood early successional stands than in the oak-hickory forest of the older growth watershed. Ammonification rates were the main factor controlling nitrification in the early successional stands. There was no evidence of inhibition of nitrification in soils from the older growth oak-hickory forest site.Within the early successional watershed, black locust sites had net mineralisation and nitrification rates at least twice as high as those in the pine mixed-hardwood stands. Concentrations of exchangeable nitrate in the soil of black locust stands were higher than in pine-mixed hardwoods at 0–15 cm in March and they were also higher at 0–15, 16–30 and 31–45 cm depth in the black locust dominated sites in July. Soil solution nitrate concentrations were higher under black locust than under pine-mixed hardwoods. Areas dominated by the nitrogen fixing black locust had greater nitrogen mineralisation and nitrification rates, resulting in higher potential for leaching losses of nitrate from the soil column in the early successional watershed.

Hillslope nutrient dynamics following upland riparian vegetation disturbance

AbstractWe investigated the effects of removing near-stream Rhododendron and of the natural blowdown of canopy trees on nutrient export to streams in the southern Appalachians. Transects were instrumented on adjacent hillslopes in a first-order watershed at the Coweeta Hydrologic Laboratory (35°03′N, 83°25′W). Dissolved organic carbon (DOC), K+, Na+, Ca2+, Mg2+, NO3−-N, NH4+-N, PO43−-P, and SO42− were measured for 2 years prior to disturbance. In August 1995, riparian Rhododendron on one hillslope was cut, removing 30% of total woody biomass. In October 1995, Hurricane Opal uprooted nine canopy trees on the other hillslope, downing 81% of the total woody biomass. Over the 3 years following the disturbance, soilwater concentrations of NO3−-N tripled on the cut hillslope. There were also small changes in soilwater DOC, SO42−, Ca2+, and Mg2+. However, no significant changes occurred in groundwater nutrient concentrations following Rhododendron removal. In contrast, soilwater NO3−-N on the storm-affected hillslope showed persistent 500-fold increases, groundwater NO3−-N increased four fold, and streamwater NO3−-N doubled. Significant changes also occurred in soilwater pH, DOC, SO42−, Ca2+, and Mg2+. There were no significant changes in microbial immobilization of soil nutrients or water outflow on the storm-affected hillslope. Our results suggest that Rhododendron thickets play a relatively minor role in controlling nutrient export to headwater streams. They further suggest that nutrient uptake by canopy trees is a key control on NO3−-N export in upland riparian zones, and that disruption of the root–soil connection in canopy trees via uprooting promotes significant nutrient loss to streams.

Factors Controlling Nitrification in Soils of Early Successional and Oak/Hickory Forests in the Southern Appalachians

Abstract Factors regulating nitrification were examined in three forests of contrasting nitrifying activity in the southern Appalachians of North Carolina, U.S.A. NH4N availability was the main factor regulating nitrification in pine/mixed-hardwood and black locust (Robinia pseudo-acacia L.)-dominated early successional forests. Litter leachate solutions from black locust had high concentrations of N and other nutrients, but their influence upon nitrification as estimated in laboratory-amended soil incubations was relatively small. In a mature oak/hickory forest, nitrification was not stimulated by NH4N amendments, nor by amendments of black-locust litter leachate solutions. Amendments with CaCO3 and CaCl2 stimulated ammonification but did not stimulate nitrification in the soils of this forest. Laboratory incubations of soils amended with oak/hickory live leaves, litter, and forest-floor extracts suggested a possible inhibitory action on nitrification from oak leaves. Low nitrification was also found in glucose-amended laboratory incubations of black-locust soils, suggesting that an increase of the C:N ratio of the soil following amendment with extracts could be responsible for low nitrification rates.

Fine root dynamics along an elevational gradient in the southern Appalachian Mountains, USA

Attributes of fine roots (<2.0 mm diameter) were quantified in five southern Appalachian plant communities along an elevational gradient. These attributes include the seasonal dynamics of fine root mass and length, the depth distribution of fine roots, fine root width and, most importantly, the annual appearance and disappearance of fine roots. The principal objectives of this study were two-fold: (1) to compare these attributes of fine roots between plant communities and (2) to compare the results of the two methods used to quantify the attributes: (1) harvesting roots from forest soil with soil cores and (2) photographing roots growing against the windows of minirhizotron boxes. The plant communities that were sampled are characteristic of the region and are designated as follows from lowest elevation (782 m) to highest elevation (1347 m): (1) xeric ridge, (2) cove hardwoods, (3) low elevation mixed oak, (4) high elevation mixed oak, and (5) northern hardwoods. Fine root mass varies seasonally in this temperate region with lowest and highest mass in the spring and autumn, respectively. Fine root mass and fine root mass appearance were lowest in the cove hardwood community and highest in the low elevation mixed oak community. The total length of fine roots was highest in the xeric ridge community and lowest in the low elevation mixed oak community. The high total root length in the xeric ridge community was due to the presence of an exceptionally dense mat of very fine roots found there. The width of these roots was significantly less than that of roots on all other plots. Subsequent regression illustrates two strong patterns in the data. First, fine root mass, fine root mass appearance and leaf production were positively correlated. Second, fine root length and soil moisture were negatively correlated. The accumulation of root mass in these communities was linked to overall site productivity and the development of root length in response to moisture stress. Only the timing of root growth initiation was related to elevation and the associated parameter of soil temperature. The best estimates of fine root appearance and disappearance were generated by harvesting roots rather than photographing them. Some methodological problems with root photography implemented in this study are addressed. # 2003 Elsevier B.V. All rights reserved.

Seed rain during initial colonization of abandoned pastures in the premontane wet forest zone of southern Costa Rica

Understanding tropical succession requires insight into propagule availability, which constrains possible woody plant recruitment, yet seed rain composition in multiple post-agricultural sites has seldom been examined. We monitored seed rain for 60 wk in five abandoned pastures in southern Costa Rica, collecting a total of 1 140 688 seeds of 165 morphospecies. Most seeds (80.1%) arrived during the wet season. Species richness was highest in the wet season and greater in forest than in pasture. Seed rain density was greatest at the forest/pasture edge and decreased drastically just a few metres into pastures. In and near the forest, animal-dispersed seeds were more abundant than seeds dispersed by other means, while wind-dispersed taxa increased in relative importance at greater distances from the forest. Total seed input to pastures did not reflect size of adjacent forest fragments, although seed rain density varied more than threefold among sites. Among-site variation in density of regenerating woody seedlings was roughly proportional to among-site variation in seed rain. Morphospecies composition differed significantly among sites. Also, seed rain and woody plant colonists were rather dissimilar in composition, suggesting that while propagule availability is necessary for early woody plant establishment, it is a poor predictor of successional trajectory.

Soil-solution chemistry in black locust, pine/mixed-hardwoods and oak/hickory forest stands in the southern Appalachians, U.S.A.

Abstract Soil-solution chemistry was measured over a 15-month period in three forest stands of contrasting nitrogen mineralization and nitrification rates in the southern Appalachians of North Carolina, U.S.A., using porous-cup lysimeters. In a black-locust-dominated stand, soil solution NO3N was 3.73 and 5.04 mg l−1 at 30- and 60-cm depth respectively, and dissolved organic N ( don ) was 0.718 and 0.582 mg l−1 respectively. Values at 30 and 60 cm for a pine/mixed-hardwood stand were 0.032 and 0.058 mg l−1 NO3N, and 0.201 and 0.168 mg l−1 don (values are means over the whole duration of the study). At both depths, soil solution conductivity, pH, Ca, Mg, K and PO4P were higher in black locust than in pine/mixed-hardwoods, and there were no differences in soil solution Na. In an oak/hickory stand, soil solution NO3N at 30-cm depth was 0.008 mg l−1, and don was 0.357 mg l−1. At 30-cm depth, soil-solution conductivity, Ca, Mg and PO4P were higher in black locust than in oak-hickory, with no differences in pH, K and Na; don , pH and K were higher in oak/hickory than in pine/mixed-hardwoods. In the oak/hickory and pine/mixed-hardwoods forest stands, with relatively lower soil N turnover rates, don was a major portion of soil solution N.