uid=AND,o=lter,dc=ecoinformatics,dc=org all public read SP016 Robert P. Griffiths

Oregon State University;Dept. of Forest Science;321 Richardson Hall Corvallis OR 97331-5752 USA

(541) 737-6559 bbgriff@peak.org griff@for.orst.edu Robert P. Griffiths

Oregon State University;Dept. of Forest Science;321 Richardson Hall Corvallis OR 97331-5752 USA

(541) 737-6559 bbgriff@peak.org griff@for.orst.edu Principal Investigator Robert P. Griffiths

Oregon State University;Dept. of Forest Science;321 Richardson Hall Corvallis OR 97331-5752 USA

(541) 737-6559 bbgriff@peak.org griff@for.orst.edu Creator 2011-09-14 Measurements were made along transects running from mountain meadows, into transition zones where trees were invading meadows and then into mature forest to determine if the invasion of high central Oregon Cascade Mountain meadows by the surrounding forest altered soil properties. Prior studies shown that meadow soil chemical and biological characteristic change when they are invaded by surrounding trees. For instance meadow litter has been shown to be enriched in nitrogen when compared with tree litter. In this study, differences in nitrogen pools and cycling were observed supporting the view that nitrogen is more available in meadow soils than in forests and that these differences change rapidly when tree invade mountain meadows. As trees invade meadows, ß-glucosidase activity is also rapidly reduced suggesting that qualitative changes are taking place in microbial populations as microorganisms adjust to changes in litter quality. High correlations between litter depth and most variables suggest that meadow litter may control other aspects of biogeochemical cycling; a relationship not observed in the transition zone or the mature forest. With one exception, the values observed in the transition zone were intermediate between those in meadows and those in forest soils. In most cases, the values found in this zone were closer to those found in mature forests than in meadow soils suggesting that when trees invade meadows, soil properties are rapidly shifted toward those found in forests. These rapid changes may alter soils so that they are more likely to support trees than grass. This may partially explain why, areas where trees had been cut after they became established as small islands within the meadow are rapidly recolonized by trees rather than grass. soil properties disturbance inorganic nutrients meadows LTER controlled vocabulary Long-Term Ecological Research (LTER) Andrews Experimental Forest site thesaurus disturbance inorganic nutrients LTER core research areas References Franklin, J.F., Moir, W.H.., Douglas, G.W., and Wiberg, C. (1971) Invasion of subalpine meadows by trees in the Cascade Range, Washington and Oregon. Arctic Alpine Research 3, 215-224. Göceoðlu, M. (1988) Nitrogen mineralization in volcanic soil under grassland, shrub and forest vegetation in the Aegean region of Turkey. Oecologia 77, 242-249. Griffiths, R. P., Bradshaw, G. A., Marks, B., and G. W. Lienkaemper. 1996. Spatial distribution of ectomycorrhizal mats in coniferous forests of the Pacific Northwest, USA. Plant and Soil 180:147-158. Griffiths, R.P., Homann, P.S., and Riley, R. (1998) Denitrification enzyme activity of Douglas-fir and red alder forest soils of the Pacific Northwest. Soil Biology and Biochemistry 30, 1147-1157. Groffman, P.M., and Tiedje, J.M. 1989. Denitrification in north temperate forest soils: relationships between denitrification and environmental factors at the landscape scale. Soil Biology & Biochemistry 21, 621-626. Hickman, J.C. (1976) Non-forest vegetation of the central western Cascade Mountains of Oregon. Northwest Scientist 50, 145-155. Hunt, H.W., Ingham, E.R., Coleman, D.C., Elliott, E.T., and Reid, C.P.P. (1988) Nitrogen limitation of production and decomposition in prairie, mountain meadow, and pine forest. Ecology 69, 1009-1016. Ingham, E R., Coleman, D.C., and Moore, J.C. (1989) An analysis of food-web structure and function in a shortgrass prairie, a mountain meadow, and a lodgepole pine forest. Biology and Fertility of Soils 8, 29-37. Johnson, N.C., and Wedin, D.A. (1997) Soil carbon, nutrients, and mycorrhizae during conversion of dry tropical forest to grassland. Ecological Applications 7, 171-182. Keeney, D.R., and Bremner, J.M. 1966. Comparison and evaluation of laboratory methods of obtaining an index of soil nitrogen availability. Agronomical Journal 58, 498-503. Keeney, D.R., and Bremner, J.M. 1966. Comparison and evaluation of laboratory methods of obtaining an index of soil nitrogen availability. Agronomical Journal 58, 498-503. Keeney, D.R., and Nelson, D.W. 1982. Nitrogen-inorganic forms. In Methods of soil analysis. Edited by A.L. Page, R.H. Miller, and D.R. Keeney. American Society of Agronomy, Madison, Wis. pp. 643-698. Köchy, M. and Wilson, S.D. (1997) Litter decomposition and nitrogen dynamics in aspen forest and mixed-grass prairie. Ecology 78, 732-739. Magee, T.K. and Antos, J.A. (1992) Tree invasion into a mountain-top meadow in the Oregon Coast Range, USA. Journal of Vegetative Science 3, 485-494 Miller, E.A., and Halpern C.B. (1998) Effects of environment and grazing disturbance on tree establishment in meadows of the western Cascade Range, Oregon, USA. Journal of Vegetative Science 9, 265-282. Perry, D.A., Amaranthus, M.P., Borchers, J.G., Borchers, S.L. and Brainerd, R.E. (1989) Bootstrapping in ecosystems Biological Science 39, 230-237. Popenoe, J.H., Bevis, K.A., Gordon, B.R., Sturhan, N.K., and Hauxwell, D.L. (1992) Soil - vegetation relationships in Franciscan terrain of Northwestern California. Soil Science 56, 1951-1952. Ross, D.J., Tate, K.R., and Feltham, C.W. (1996) Microbial biomass, and C and N mineralization, in litter and mineral oils of adjacent mountain ecosystems in the southern beech (Nothofagus) forest and a tussock grassland. Tabatabai, M. A., and J. A. Bremner. (1969) Use of p-nitrophenyl phosphate for assay of soil phosphatase activity. Soil Biology & Biochemistry 1,301 - 307. Yakimenko, E.Y. (1997) Soil comparative evolution under grasslands and woodlands in the forest zone of Russia. In Management of Carbon Sequestration in Soil, eds. R. Lal, J. K. Kimble, and B.A. Stewart, pp. 391-404. CRC Press, New York. Zou, X., D. Binkley, and K. G. Doxtader. (1992) A new method for estimating gross phosphorus mineralization and immobilization rates in soils. Plant & Soil 147,243 - 25 The National Science Foundation provided financial support from grants .Students:Michael Madritch, Brian Mitchell, Edwin Price, Ronald Slangen, Cedar Johnson ,Amy Holcomb.John Phillips of the US Forest Service ChCharles Halpern of the Univ of Washington Data Use Agreement: The re-use of scientific data has the potential to greatly increase communication, collaboration and synthesis within and among disciplines, and thus is fostered, supported and encouraged. This Data Set is released under the Creative Commons license CC BY "Attribution" (see: https://creativecommons.org/licenses/by/4.0/). Creative Commons license CC BY - Attribution is a license that allows others to distribute, remix, tweak, and build upon your work (even commercially), as long as you are credited for the original creation. This license accommodates maximum dissemination and use of licensed materials. It is considered professional conduct and an ethical obligation to acknowledge the work of other scientists. The Data User is asked to provide attribution of the original work if this data package is shared in whole or by individual parts or used in the derivation of other products. A recommended citation is provided for each Data Set in the Andrews LTER data catalog (see: http://andlter.forestry.oregonstate.edu/data/catalog/datacatalog.aspx). A generic citation is also provided for this Data Set on the website https://portal.edirepository.org in the summary metadata page. Data Users are thus strongly encouraged to consider consultation, collaboration and/or co-authorship with the Data Set Creator. While substantial efforts are made to ensure the accuracy of data and associated documentation, complete accuracy of data sets cannot be guaranteed and all data are made available "as is." The Data User should be aware, however, that data are updated periodically and it is the responsibility of the Data User to check for new versions of the data. The data authors and the repository where these data were obtained shall not be liable for damages resulting from any use or misinterpretation of the data. General acknowledgement: Data were provided by the HJ Andrews Experimental Forest research program, funded by the National Science Foundation's Long-Term Ecological Research Program (DEB 2025755), US Forest Service Pacific Northwest Research Station, and Oregon State University. If data used in publication, the PI will be listed as a coauthor. Whenever these data are presented in whatever form, the PI will be acknowledged. http://andlter.forestry.oregonstate.edu/data/abstract.aspx?dbcode=SP016 1998-01-07 1998-01-09 All Organisms All Organisms Highest common category (ca. kingdom) Fungi Division or Phylum Basidiomycota Division or Phylum Basidiomycetes Order Phallales Family Hysterangiaceae Genus Hysterangium Family Gomphaceae Genus Gautieria Although forest meadows in the Central Oregon Cascade Mountains make up a relatively small fraction of total area, they contain a large variety of plant species that greatly enrich biodiversity over the landscape (Hickman, 1976). Under present climatic and forest management conditions, many high-dry mountain meadows of Pacific Northwest are being invaded by the surrounding forest providing an opportunity to study changes in soil properties in response to vegetative succession (Franklin et al., 1971: Magee and Antos, 1992; Yakimenko, 1997; Miller and Halpern, 1998). It is likely that these meadows were originally established and maintained by aboriginal burning (Miller and Halpern, 1998). However, factors responsible for the current invasion are not known with certainty but climate change, fire suppression and termination of sheep grazing may all have played a role (Popenoe et al., 1992; Miller and Halpern, 1998). In a comprehensive study of mountain tree invasion in the Central Oregon Cascade Mountains, Miller and Halpern (1998) noted that in addition to the allegoric factors mentioned above, autogenic factors may also control this process (i.e. local influence of trees on the establishment of seedlings by altering microclimate). In addition to controlling moisture, the pioneer trees could also be altering soil properties resulting in increased seedling establishment. Vegetation can profoundly influence both the chemistry and biology of soils; altering soils in a way that enhances plant community resiliency to perturbation (Perry et al.1989). To a large degree, this finding explains why forests that have been disturbed by fire, disease, wind-throw, harvesting or other factors typically return to the same vegetative assemblage that was present before the disturbance. Similar mechanisms may explain why invading vegetation alter soils to favor trees rather than the original meadow vegetation. Comparative studies between grasslands and forests have shown large differences in soil chemistry (Göceoðlu, 1988; Hart et al., 1992; Popenoe et al., 1992; Ross et al., 1996; Yakimenko, 1997); litter decomposition rates (Hunt et al., 1988; Köchy and Wilson, 1997) and food web compositions (Hunt et al., 1987; Ingham at al., 1989). Our study was designed to measure changes in both the chemical and biological characteristics of high-elevation mountain meadows as adjacent forests invade them. This includes an analysis of the transition zone to obtain a rough idea of which soil properties are most rapidly altered in response to the tree invasion. Within the context maintaining biodiversity and habitat diversity, forest managers are looking techniques to reverse invasion high elevation mountain meadows by surrounding trees (Popenoe et al., 1992). One of the objectives of this study was to provide basic information about biogeochemical transformations associated with tree invasion that could be used to monitor the effectiveness of different treatments.

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Andrews Forest LTER Program US Forest Service Pacific Northwest Research Station 3200 SW Jefferson Way Corvallis OR 97331

hjaweb@fsl.orst.edu http://andrewsforest.oregonstate.edu/ Andrews Forest LTER Site

Forest Ecosystems and Society Department in Forestry Oregon State University 201K Richardson Hall Corvallis OR 97331-5752

(541) 737-8480 lterweb@fsl.orst.edu http://andrewsforest.oregonstate.edu/ Field Methods - SP016 At each of these positions a 4.7 x 10 cm soil core was taken for subsequent analysis. The samples were transported to the laboratory in an ice chest and subsequently stored at 15 degrees C until the initiation of analyses, usually within 16 h of their receipt. The following measurements were made in the field: litter depth, mineral soil respiration, soil temperature and ectomycorrhizal mat characteristics. Field (forest floor) respiration rates were measured with a nondispersive, infrared CO2 analyzer (Li-Cor, LI-6200). Measurements were made over a period of 1 min after the chamber gas reached ambient CO2 concentration. The instrument was calibrated on site against a known standard at each location. A Q10 adjustment was made for ambient soil temperature. Soil temperature was measured by electronic thermometers calibrated at 0 degrees C with ice water. The temperature probes were inserted into the mineral soil to a depth of 10 cm. The distribution of ectomycorrhizal mats was determined visually in the field by inspecting the relative abundance of mats in 4.7 x 10 cm cores. Two distinct mat types were scored: (1) mats similar to those of the genus Hysterangium and (2) mats similar to those of the genus Gautieria. This approach has been used successfully in the past to document ectomycorrhizal mat distribution patterns in coniferous forests of the Pacific Northwest (Griffiths et al. 1996). Laboratory Methods - SP016 In preparation for laboratory analyses, all soils were sieved through a 2-mm sieve. Soil moisture was determined by drying duplicate 10 g field-moist sieved soils at 100 degrees C for at least 8 h. The percent soil moisture was calculated by dividing the difference between wet and dry samples and dividing that number by the dry wt., which was then multiplied by 100. Soil pH was measured in 1:10 (soil:distilled water) slurries of oven-dried (100 degrees C) soil. These slurries were shaken for 1 h prior to reading pH values with a Sigma model E4753 electrode. Soil organic matter was measured by loss-on-ignition at 550 degrees C for 6 h after oven drying at 100 degrees C. Denitrification potential was measured using a method by Groffman and Tiedje (1989) as modified by us (Griffiths et al., 1998). Each reaction vessel (25-mL Erlenmeyer flask) contained 5 g of less than 2 mm, field-moist soil. Flasks were sealed with rubber serum bottle stoppers and purged with Ar to displace O2 in the headspace gas. After purging with Ar, 2 mL of a 1 mM solution of glucose and NO3- was added to each flask. Flasks were subsequently incubated at 25 degrees C for 1 h. This preincubation period was used because previous time-series experiments showed a lag in N2O production during this period. The same experiments have shown linear N2O production rates during the following 2-4 h (unpublished data). After the preincubation period, 0.5 mL of headspace gas was removed from the reaction vessel and injected into a gas chromatograph (GC) fitted with an electron capture detector (Hewlett Packard model 5890 GC, connected to a Hewlett Packard model 3396 integrator). The integrator was calibrated by the external calibration method with known gas standards. A second headspace N2O analysis was made after an additional 2-h incubation at 25 degrees C. The net N2O released over this 2-h period was used to estimate N2O production rates. Laboratory respiration measurements were made on field-moist, sieved soils (4 g dry weight). These rates represent the basal respiration rate for soil microorganisms. Soils were brought to 75% moisture content by the addition of enough sterile deionized water to equal 3 g water per 25-mL Erlenmeyer flask. Once sealed with serum bottle stoppers, the flasks were incubated at 24 degrees C for 14 days after which headspace CO2 concentrations were measured using gas chromatography. This was a measure of labile soil carbon. The same GC and integrator as were used for this assay as that used to measure N2O, but in this case a flame ionization detector and a methanizer in series were used. Substrate induced respiration (SIR) was also measured in these soils. The reaction vessels were prepared as before except 0.1 mL of 1M glucose solution was added to the reaction vessel and the assay for CO2 evolution rates were calculated from the difference between the headspace CO2 concentrations after the first h incubation and the concentration 2 h later. SIR was calculated by subtracting CO2 evolution rates without the glucose amendment from the rates in the presence of glucose. Extractable ammonium was determined by shaking 10 g of field-moist soil with 50 mL 2 M KCl for 1 h (Keeney and Nelson 1982), adding 0.3 mL 10 M NaOH to the slurry, and measuring ammonium concentration with an Orion model 95-12 ammonium electrode (Orion Research Inc., Boston, MA). Mineralizable N was measured by the water-logged technique of Keeney and Bremner (1966). For each analysis, 10 g of field-moist soil were added to 53 mL of distilled water in a 20 x 125 mm screw-cap test tube, and incubated at 40 degrees C for 7 d. Then 53 mL of 4 M KCl were added to the slurry, and ammonium concentration was determined with the ammonium electrode. Mineralizable N was calculated as the difference between initial and final ammonium concentrations. Beta-glucosidase activity was determined by the spectrophotometric assay of Tabatabai and Bremmer (1969), as modified by Zou et al. (1992). One mL of 10 mM p-nitrophenyl b-D glucopyranoside substrate was added to duplicate 1-mL subsamples containing a soil slurry (1 gdw in 1 mL deionized H2O). The tubes were shaken and then placed with duplicate controls without substrate in a 30°C water bath for 2 h. After incubating, 1 mL of 10 mM p-nitrophenyl b-D glucopyranoside was added to the controls, and all reactions were immediately stopped by the addition of 2 mL of 0.1 M tris[hydroxymethyl]aminomethane at pH 12.0. The mixtures were centrifuged for 5 min at 500 x g. From the supernatant, 0.2 mL was diluted with 2.0 mL deionized water. The optical density was measured at 410 nm, and a standard curve was prepared from 0.02 to 1.0 micro-mol/mL p-nitrophenol (pNP). Sampling frequency: 1 set measurements for each transect at each time Sampling transects ran from forest meadows into transition zones, where conifers were becoming established in forest meadows, and then into old-growth forests with relatively little understory vegetation. Each transect was made up of three 75 meter segments. Soil samples were taken and field observations made at 5 meter-intervals along these segments. Five sample locations were use as independent sample plots; each containing a single transect which, in turn, had segments in a meadow, transition zone and an old-growth forest. Andrews Experimental Forest (HJA) -122.26172200 -122.10084700 44.28196400 44.19770400 1631 1631 meter Cascade Head Experimental Forest -123.99172777 -123.89730000 45.06476948 45.03130000 Sherri L. Johnson

US Forest Service ;Pacific NW Research Station ;3200 SW Jefferson Way Corvallis OR 97331 USA

541-758-7771 sherri.johnson2@usda.gov sherri.johnson@oregonstate.edu https://www.fs.fed.us/research/people/profile.php?alias=sherrijohnson Principal Investigator Julia A. Jones

Oregon State University;Department of Geosciences; Wilkinson Hall 104 Corvallis OR 97331-5506 USA

(541) 737-1224 Julia.Jones@oregonstate.edu geojulia@comcast.net http://ceoas.oregonstate.edu/profile/jones/ http://orcid.org/0000-0001-9429-8925 Principal Investigator Matthew G Betts

Department of Forest Ecosystems and Society; 201E Richardson Hall; College of Forestry; Oregon State University Corvallis OR 97331

(541) 737-3841 matt.betts@oregonstate.edu http://www.fsl.orst.edu/flel/index.htm Principal Investigator Michael P. Nelson

Department of Forest Ecosystems and Society; 201K Richarson Hall; College of Forestry; Oregon State University Corvallis OR 97331

541-737-9221 mpnelson@oregonstate.edu http://www.michaelpnelson.com http://orcid.org/0000-0001-6917-4752 Principal Investigator David Bell david.bell@usda.gov david.bell@oregonstate.edu https://lemma.forestry.oregonstate.edu/about/david-bell Principal Investigator The H.J. Andrews Experimental Forest is a living laboratory that provides unparalleled opportunities for the study of forest and stream ecosystems in the central Cascade Range of Oregon. Since 1980, as a part of the National Science Foundation Long Term Ecological Research (NSF-LTER) program, the Andrews Experimental Forest has become a leader in the analysis of forest and stream ecosystem dynamics. Long-term field experiments and measurement programs have focused on climate dynamics, streamflow, water quality, and vegetation succession. Currently researchers are working to develop concepts and tools needed to predict effects of natural disturbance, land use, and climate change on ecosystem structure, function, and species composition. The Andrews Experimental Forest is administered cooperatively by the USDA Forest Service Pacific Northwest Research Station, Oregon State University and the Willamette National Forest. Funding for the research program comes from the National Science Foundation (NSF), US Forest Service Pacific Northwest Research Station, Oregon State University, and other sources. Data were provided by the HJ Andrews Experimental Forest research program, funded by the National Science Foundation's Long-Term Ecological Research Program (DEB 2025755), US Forest Service Pacific Northwest Research Station, and Oregon State University. National Science Foundation: DEB1440409 The Andrews Forest is situated in the western Cascade Range of Oregon, and covers the entire 15,800-acre (6400-ha) drainage basin of Lookout Creek. Elevation ranges from 1350 to 5340 feet (410 to 1630 m). Broadly representative of the rugged mountainous landscape of the Pacific Northwest, the Andrews Forest contains excellent examples of the region's conifer forests and associated wildlife and stream ecosystems. These forests are among the tallest and most productive in the world, with tree heights of often greater than 250 ft (75 m). Streams are steep, cold and clean, providing habitat for numerous aquatic organisms. 1 SP01601 Influence of coniferous tree invasion on forest meadow soil properties: SP01601.csv 21973 c44c59d7e03a03aa25d565885885af20 1 \r\n column , " http://andlter.forestry.oregonstate.edu/ltermeta/ltersearch/dataaccess.aspx?docid=SP01601_v1.csv TRANSLOC Location of sample site along transect starting within og numeric(2,0) number 1.000000 natural 1.0000 48.0000 SITE_NO Site designator numeric(2,0) number 1.000000 natural 53.0000 57.0000 TYPECODE Type code: 1=grass; 2=transition; 3=old-growth numeric(1,0) number 1.000000 natural 1.0000 3.0000 HYSTER Percentage of core containing mycorrhizal mats like those of the genus Hysterangium numeric(3,0) percent 1.000000 whole 0.0000 100.0000 GAUT Percentage of core containing mycorrhizal mats like those of the genus Gautieria numeric(3,0) percent 1.000000 whole 0.0000 100.0000 LITTER Litter depth numeric(2,0) centimeters 1.000000 whole 0.0000 20.0000 MOIST Percent moisture numeric(5,1) percent 0.100000 real 0.0000 110.0000 SOILTEMP Soil temperature measured with Licor numeric(4,1) degrees Celsius 0.100000 real 0.0000 30.0000 AIRTEMP Air temperature measured with Licor numeric(4,1) degrees Celsius 0.100000 real 0.0000 30.0000 LIGHT Amount of light measured with Licor numeric(6,1) micromoles per square meter per second 0.100000 real 0.0000 2000.0000 SOM Soil organic matter numeric(5,2) percent 0.010000 real 10.0000 110.0000 PH pH numeric(4,2) pH units 0.010000 real 4.0000 7.0000 EXAMM Extractable ammonium (dry weight basis) numeric(4,2) micromoles per gram 0.010000 real 0.0000 3.0000 MINN Mineralizable nitrogen (dry weight basis) numeric(5,2) micromoles per gram 0.010000 real 0.0000 50.0000 DENIT Denitrification potential (dry weight basis) numeric(4,1) nanograms per gram per hour 0.100000 real 0.0000 100.0000 B_GLUC Beta-glucosidase activity (dry weight basis) numeric(6,3) micromoles per gram per hour 0.001000 real 0.0000 1.0000 LABRESP Laboratory respiration rates (dry weight basis, as C) numeric(4,1) micrograms per gram per hour 0.100000 real 0.0000 50.0000 FLDRESP Field respiration rates (dry weight basis) numeric(5,2) grams per square meter per day 0.010000 real 0.0000 100.0000 SIR_H2O Substrate-induced respiration with only H2O; no substrate added (dry weight basis, as C) numeric(4,2) micrograms per gram per hour 0.010000 real -3.0000 10.0000 SIR_1NM Substrate-induced respiration with 1 nm glucose added (dry weight basis, as C) numeric(6,2) micrograms per gram per hour 0.010000 real -8.0000 10.0000 SIR_1M Substrate-induced respiration with 1 m glucose added (dry weight basis, as C ) numeric(6,2) micrograms per gram per hour 0.010000 real -9.0000 20.0000 NOTNULL SP01601.AIRTEMP SP01601.DENIT SP01601.FLDRESP SP01601.GAUT SP01601.HYSTER SP01601.LABRESP SP01601.LIGHT SP01601.LITTER SP01601.MINN SP01601.MOIST SP01601.SIR_1M SP01601.SIR_1NM SP01601.SITE_NO SP01601.SOILTEMP SP01601.TRANSLOC SP01601.TYPECODE 235 micromoles per g per hour nanograms/gram*hour grams per square meter per day micromoles per square meter per second centimeters; .01 meters micromoles per gram Scale used for pH measurements micrograms per gram per hour Degrees Celsius; a common unit of temperature; constantToSI=273.18 percent; a number dimensionless number, i.e., ratio, count