Long-term Carbon and Nitrogen, and Phosphorus Dynamics of Leaf and Fine Root Litter in North American Ecosytems.

LIDET-Long-term Intersite Decomposition Experiment Team

INTRODUCTION

Decomposition and the release of nitrogen from litter is one of the most basic of all ecosystem processes. Despite the fact that the formation of "stable" soil organic matter and nitrogen are long-term processes, the majority of past studies on this process have been short-term. The primary objective of this study is therefore to examine the control that substrate quality and climate have on patterns of long- term decomposition and nitrogen accumulation in above- and below-ground fine litter. Of particular interest is the degree these two factors control the formation of stable organic matter and nitrogen after extensive decay.

EXPERIMENTAL DESIGN

The major factors considered in this experiment are site, species of and type of litter (leaves, vs. roots vs dowels), and time. Twenty-eight sites (Table 1), representing a wide array of biomes (Table 2), moisture and temperature (Table 3) conditions, are used for litter incubations. Ten types of "standard" litters were sent to each site (Table 4). These included three types of fine roots (graminoid, hardwood, and conifer), six types of leaf litter (which ranged in lignin/nitrogen ratio from 5 to 75), and wooden dowels . Samples are to be collected ten times; the time between samples will be one year for all sites except tropical sites which will collect samples every three to six months. There are four replicates for each species, site and time.

In addition to the standard litters, each site is represented by a "wildcard" litter which appears at one randomly selected site for each sample collection. The purpose of the wildcard species is to verify the predictions from the standard species. There are four replicates for each wildcard species, site and time.

METHODS

Litter Collection

Each site was responsible for collecting the litter used in the experiments. For most sites, the leaf litter was collected directly from senesecent plants or as freshly fallen litter. Green leaves were collected from the Jornada, San Diego, Luquillo, and LaSelva sites. All leaf litter except, Drypetes glauca which was oven dried at 40 C to prevent decay, was air dried prior to shipment to Oregon. Pinus elliotii fine roots were collected by excavating surface roots and washing. Graminoid roots were collected from material exposed along stream banks. Graminoid and pine roots were air dried, whereas the tropical hardwood roots were oven dried at 40 C to prevent decomposition.

In the case of the LaSelva site, the litter was sterilized after the bags were filled to kill all invertebrates, fungi, and virus prior to shipment. Sterilization was conducted at the Battelle National Laboratory by exposing the litter to 20 hours of gamma rays with 60Co as the source. The total exposure was 2 Mrad.

Bag Design

All bags were 20- by 20-cm and filled with 10 g leaves or 5- 7 g of fine roots. Each bag was identified with a unique number embossed on an aluminum tag. The bag openings were sealed with six monel staples. The initial air dry weight, calculated oven dry weight, species, site, replicate number for each litterbag were recorded prior to placement in the field. Subsamples of litter material were taken to determine the air dry to oven dry conversion factor and the initial chemistry of the litter. Moisture content of the air dried litter ranged from 2-10%.

Three types of bags were used in this experiment (photo). For the long-term leaf litter experiment the bags had a top mesh of 1 mm and a bottom of 55 micron mesh. The bags used for fine roots were entirely of 55 micron mesh. The bags used in the mesh size effects study had a top of 7 mm mesh and a bottom of 55 micron mesh.

Dowels

The wooden dowels used in the experiment are made of ramin (Gonystlylus bancannus). This species is a tropical hardwood from southeast Asia. It is not resistant to decay and rated as perishable. The dowels are 13 mm in diameter and 61 cm in length. One half of the dowel is embedded vertically into the soil and the other half is exposed to aerial conditions. The air dry weight of each dowel was recorded, and a subsample of dowels was measured for diameter, density, air dry moisture content, nitrogen content, and carbon chemistry.

Recovery of the initial dowels, particularly the below- ground portion, proved difficult. In 1994 a second group of wooden dowels was sent out to all LIDET sites. These dowels have a sleeve of 1 mm mesh on the lower 30.5 cm with an attached aluminum tag identical to the tag attached to the upper half of the dowel. A 20 cm section of orange plastic flagging was sewn into the top corner of the mesh sleeve to improve recovery.

Sample Placement

Samples were placed in the field during fall of 1990, or 1991, by each of the participating sites. Locations were near climatic stations and in areas protected from disturbances that could destroy the litter bags. The locations were also selected to be typical of areas that other intersite decomposition experiments might be conducted.

The exact method for placement varied from site to site, but the following standards were applied:

1) Four separate locations were selected to avoid pseudo- replication problems.

2) Each set of bags to be collected was connected by a cord; these sets of bags were laid out in parallel lines in a random order (photo).

3) Leaf litterbags were placed so that contact with the underlying litter layer is made (photo 3578 #28). Fine root litterbags were inserted 10-20 cm into the upper mineral soil or humus layer for histosols (photo).

4. Dowels were installed at the end of the string opposite the fine root bags. The dowels were placed so that 30 cm is exposed to the air and 30 cm is embedded in the soil (photo).

Sample Processing

Once the litter or dowels are collected they are oven dried in paper bags at 55oC until the mass is stable. Any mosses, lichens, fine roots, or other plant parts that had grown into the bags or dowels were removed prior to harvesting. Samples were be pooled by species, site, and time for grinding and archiving. A set of unpooled samples has also be saved to determine the internal variability of pooled samples. Chemical analyses are performed using two methods. Each sample is analyzed for total nitrogen, ash, lignin, and cellulose using near infrared reflectance spectoscopy. Approximately 25% the pooled samples are sampled for ash, Kjeldahl nitrogen, lignin, cellulose, water extractive, non- polar extractive, and ash content using wet chemical. These samples are then used to calibrate the near infrared reflectance spectoscopy methods. Finally, the cation concentration of the pooled samples is determined using ICAP.

REFERENCES

Bolster, K. L., M. E. Martin, and J. D. Aber. 1996. Determination of carbon fraction and nitrogen concentration in tree foliage by near infrared reflectance: a comparison of statistical methods. Canadian Journal of Forest Research 26:590-600.

Currie, W. S. 1995. Forest Floor Leachate Biogeochemistry and Decomposition Dynamics. Ph.D. Dissertation. Institute for the Study of Earth, Oceans, and Space, University of New Hampshire, Durham, NH. 184 pp.

Currie, W. S. and J. D. Aber. 1997. Modeling leaching as a decomposition process in humid, montane forests. Ecology 78(6): 1844-1860.

Currie, W. S., K. Nadelhoffer, and J. D. Aber. In review. Soil detrital processes controlling the movement of 15N tracers to forest vegetation. Ecological Applications.

Gholz, H. L., D. Wedin. M. E. Harmon and S. Smitherman. in prep. Long-term dynamics of pine and heartwood litter in contrasting environments:toward a global model of decomposition. Ecology.

Harmon, M. E., K. J. Nadelhoffer, and J. Blair. in press. Measuring Decomposition, Nutrient Turnover, and Stores in Plant Litter. Pages XX-XX in Robertson, G. P., C. S. Beldsoe, D. C. Coleman, and P Sollins, editors. Standard Soil Methods for Long-term Ecological Research. Oxford University Press. Harmon, M. E. and K. Lajtha. in press. Analysis of Detritus and Organic Horizons for Mineral and Organic Constituents. Pages XX-XX in Robertson, G. P., C. S. Beldsoe, D. C. Coleman, and P Sollins, editors. Standard Soil Methods for Long-term Ecological Research. Oxford University Press.

Long-term Intersite Decomposition Experiment Team (LIDET). 1995. Meeting the challenge of long-term, broad-scale ecological experiments. Publication No. 19. US. LTER Network Office: Seattle, WA, USA. 23 p.

Moorhead, D. L., W. S. Currie, E. B. Rastetter, W. J. Parton, and M. E. Harmon. in review. Climate and litter quality controls on decomposition: an analysis of modeling approaches. Ecosystems

O'Lear, H. A., T. R. Seastedt, J. M. Briggs, J. M. Blair, and R. A. Ramundo. 1996. Fire and topographic effects on decomposition rates and N dynamics of buried wood in tallgrass prairie. Soil Biol. Biochem. 28:323-329.

Parton, W. J., D. S. Ojima, C. V. Cole, and D. S. Shimel. 1994. A general model for soil organic matter dynamics: sensitivity to litter chemistry, texture, and management. Soil Science Society of America Special Publication 39:147-167.

Parton, W. J. and M. E. Harmon. in prep. Abiotic controls on decomposition of wood at the global scale. Ecological Applications.


Table 1. Twenty one sites are involved as of 10/90. These include the 17 LTER sites and 4 non-LTER sites. Seven additional sites were added in 1991 bringing the total to twenty-eight.

Site LATITUDE LONGITUDE
Andrews 44o14'N 122o11'W
Arctic Lakes 68o38'N 149o43'W
Barro Colorado 9o10'N 79o51'W
Bonanza Creek 64o45'N 148o00'W
Blodgett 38o52'N 120o38'W
Cedar Creek 45o24'N 93o12'W
Central Plains 40o49'N 104o46'W
Coweeta 35o00'N 83o30'W
Florida 29o45'N 82o30'W
Guanica 17o57'N 65o52'W
Hubbard Brook 43o56'N 71o45'W
Harvard Forest 42o40'N 72o15'W
Jornada 32o30'N 106o45'W
Juneau 58o00'N 134o00'W
Kellogg 42o24'N 85o24'W
Konza 39o05'N 96o35'W
LaSelva 10o00'N 83o00'W
Loch Vale 40o17'N 105o39'W
Luquillo 18o19'N 65o49'W
Monte Verde 10o18'N 84o48'W
Niwot Ridge 40o03'N 105o37'W
North Inlet 33o30'N 79o13'W
Northern Lakes 46o00'N 89o40'W
Olympics 47o50'N 123o53'W
Santa Margarita 33o29'N 117o09'W
Sevilleta 34o29'N 106o40'N
Virginia Coast 37o30'N 75o40'W


Table 2. The study includes all major terrestrial biomes in North America. Twenty-eight sites are involved. Individual site descriptions are stored in the TD2306 file.

Site                   Biome

Andrews               cool temperate conifer
Arctic Lakes          tussock tundra
Barro Colorado Island moist tropical
Bonanza Creek         boreal conifer
Blodgett State Forest temperate conifer
Cedar Creek           tall grass/ forest transition
Short Grass Steppe    shortgrass steppe
Coweeta               warm temperate hardwood
Curley Valley         cold desert
Florida               warm temperate conifer
Guanica               dry tropical forest
Hubbard Brook         cool temperate hardwood
Harvard Forest        cool temperate hardwood
Jornada               warm desert
Juneau                cool temperate conifer
Kellogg               agriculture
Konza                 tallgrass prairie
LaSelva               wet tropical
Luquillo              wet tropical
Loch Vale             cold alpine conifer 
Monte Verde           tropical cloud forest
Niwot Ridge           alpine tundra
North Inlet           coastal marine
Northern Lakes        cool temperate mixed
Olympics              cool temperate conifer   
Santa Margarita       chaparral
Sevilleta             desert
Virginia Coast        coastal marine

Table 3. Climatic parameters for sites. Note these values should be checked against the most recent data for the sites.

Site  
Mean Annual
Temp C       Precip cm
 

AET cm
 

PET cm
 

Elev. m
Andrews   8.6 230.9   76.4   98.2   500
Arctic Lakes   -7.0 32.7   28.4   42.3   760
Barro Colorado   25.6 269.2   136.8   151.7   30
Bonanza Creek   -5.0 40.3   36.0   57.6   300
Blodgett   14 .4 124.4   75.3   109.7   1300
Cedar Creek   5.5 82.3   73.3   102.6   230
Central Plains   8.9 44.0   43.0   120.2   1650
Coweeta   12.5 190.6   117.3   135.3   700
Florida   21 .0 123.8   116.6   162.1   35
Guanica   26.3 50.8   50.2   142.2   80
Hubbard Brook   5.0 139.6   71.2   81.7   300
Harvard Forest   7.1 115.2   85.1   104.1   335
Jornada   14.6 29.8   29.2   166.6   1410
Juneau   4.4 287.8   49.5   54.4   100
Kellogg   9.0 81.1   70.6   100.7   288
Konza   12.8 79.1   74.7   125.0   366
LaSelva   25.0 409.9   169.9   177.3   35
Loch Vale   1.6 109.6   85.1   108.3   3160
Luquillo   23.0 336.3   123.4   125.9   350
Monte Verde   17.7 268.5   108.4   116.6   1550
Niwot Ridge   -3.7 124.9   64.7   75.6   3650
North Inlet   18.1 149.1   120.6   145.6   2
Northern Lakes   4.4 67.7   64.9   88.4   500
Olympics   10.0 153.1   79.4   104.4   150
Santa Margarita   16.4 24.0   23.6   186.0   500
Sevilleta   16.0 25.4   25.2   160.2   1572
Virginia Coast   15.0 113.8   99.3   121.5   0

Table 4. Species being tested in LIDET experiments. Those with a * are the "standard" species sent to each site each time.

ASPEN (Populus tremuloides)
BEACH GRASS (Ammophila breviligulata)
BEECH (Fagus grandifolia)
BIG BLUE STEM (Schizachyrium gerardi)
BLACK GRAMA (Bouteloua eriopoda)
BLACK LOCUST (Robinea pseudoacacia)
BLUE GRAMA (Bouteloua gracilis)
CEANOTHUS (Ceanothus greggii)
*CHESTNUT OAK (Quercus prinus)
CREOSOTE BUSH (Larrea tridentata)
DOUGLAS-FIR (Pseudotsuga menzesii)
*DRYPETES (Drypetes glauca)
EASTERN WHITE PINE (Pinus strobus)
GYMNANTHES (Gymnanthes lucida)
KOBRESIA (Kobresia myosuroides)
LITTLE BLUE STEM (Schizachyrium scoparium)
PACIFIC RHODODENDRON (Rhododendron macrophyllum)
PACIFIC DOGWOOD (Cornus nuttalii)
RAMIN (Gonystylus bancanus)
*RED PINE (Pinus resinosa)
*SLASH PINE (Pinus elliottii)
SPARTINA (Spartina alternaifolia)
SUBALPINE FIR (Abies lasiocarpa)
*SUGAR MAPLE (Acer saccharum)
VOCHYSIA (Vochysia ferragenea)
WAX MYRTLE  (Myrica cerifer)
*WESTERN REDCEDAR (Thuja plicata)
*WHEAT (Triticum aestivum)
YELLOW BIRCH (Betula lutea)
WHITE FIR (Abies concolor)