Sie sind auf Seite 1von 14

ecological indicators 9 (2009) 462475

available at www.sciencedirect.com

journal homepage: www.elsevier.com/locate/ecolind

Riparian forest indicators of potential future stream condition


Paul L. Ringold a,*, John Van Sickle a, Mike Bollman b,1, Jeff Welty c, Jerry Barker b,2
U.S. Environmental Protection Agency, Ofce of Research and Development, Western Ecology Division, 200 SW 35th Street, Corvallis, OR 97330, United States b Dynamac Corporation, 200 SW 35th Street, Corvallis, OR 97330, United States c Weyerhaeuser Company, PO Box 9777, Federal Way, WA 98063, United States
a

article info
Article history: Received 21 April 2007 Received in revised form 26 June 2008 Accepted 27 June 2008

abstract
Large wood in streams can play an extraordinarily important role in inuencing the physical structure of streams and in providing habitat for aquatic organisms. Since wood is continually lost from streams, predicting the future input of wood to streams from riparian forests is crucial to assessing or managing stream ecosystems. Unfortunately, regional monitoring protocols have no established capacity to provide this information. The goal of this research is to propose one or more methods that could meet this need. This goal is pursued by using stream wood delivery models to aid in the design of a monitoring method. Two questions are asked. First,

Keywords: Riparian forest indicator Stream wood Stream assessment Coarse woody debris Anticipatory indicator

does simpler data change model predictions of future contributions of wood from riparian ecosystems to the stream? The answers to this rst question enable monitoring design to be tailored to details affecting estimates of future stream condition. These answers are important, because more detailed data is typically more costly. Second, which metrics, if any, correlate well with model predictions? If such metrics can be identied, then these measures can serve as effective indicators of ecosystem function directly, without using ecosystem models. These questions were addressed by collecting highly detailed eld observations of riparian forests from 109 forested riparian sites in the Coast Range, Willamette Valley, and western Cascades of northwestern Oregon. Detailed and simplied versions of these data were used in models that forecast the potential of riparian forests to provide wood to the stream. Model predictions with less detailed data typically provided answers different than did predictions made with more detailed data. Thus, ecosystem assessments requiring these types of model predictions would benet from more detailed data. In contrast, riparian metrics easily observed in the eld (e.g. number of basal area of trees) or derived from remotely sensed imagery (e.g. number or height of canopy trees) were well correlated with model predictions of potential stream wood recruitment. When direct model predictions or model scenario analyses are not required, these metrics can serve as effective indicators of the potential of riparian forests to provide wood to the future stream network. Published by Elsevier Ltd.

The information in this document has been funded wholly (or in part) by the U.S. Environmental Protection Agency. It has been subjected to review by the National Health and Environmental Effects Research Laboratorys Western Ecology Division and approved for publication. Approval does not signify that the contents reect the views of the Agency, nor does mention of trade names or commercial products constitute endorsement or recommendation for use. * Corresponding author. E-mail addresses: ringold.paul@epa.gov (P.L. Ringold), vansickle.john@epa.gov (J. Van Sickle), bollman.mike@epa.gov (M. Bollman), weltyj@wdni.com (J. Welty), jbarker@walshenv.com (J. Barker). 1 Current address: U.S. Environmental Protection Agency, Ofce of Research and Development, Western Ecology Division, 200 SW 35th Street, Corvallis, OR 97330, United States. 2 Current address: Walsh Environmental Scientists and Engineers LLC, 4888 Pearl E. Circle, Boulder, CO 80301, United States. 1470-160X/$ see front matter . Published by Elsevier Ltd. doi:10.1016/j.ecolind.2008.06.009

ecological indicators 9 (2009) 462475

463

1.

Introduction

Large wood in streams can play an extraordinarily important role in creating the physical structure of streams and providing habitat for aquatic organisms (Gregory et al., 2003a). Wood currently in streams contributes to stream complexity by creating pools, trapping sediments, and affecting bank erosion and bar formation. Stream wood also alters river ow patterns and affects the relationship between a river and its oodplain (Montgomery et al., 2003a). Not surprisingly, larger pieces of wood can play a more prominent and extensive role in inuencing stream structure than can smaller pieces of wood (Bilby and Ward, 1991). While wood can reside in stream channels for centuries, the residence time of most wood is typically much shorter. Hyatt and Naiman (2001) found wood in the Queets River in northwest Washington that had been in the stream for 1400 years, but also estimated the half-life of a piece of stream wood at 20 years. Given that wood is lost from stream channels by uvial transport, fragmentation, and decomposition (Keller and Swanson, 1979; Harmon et al., 1986; Gurnell and Gregory, 1995; Bilby and Bisson, 1998; Hyatt and Naiman, 2001; Van Der Nat et al., 2003), an understanding of the future input of wood is crucial to assessing and managing stream ecosystems. Future stream wood originates in current and future riparian and upland forests. Trees from these forests become stream wood by a variety of episodic and chronic mechanisms. Trees die from competitive suppression, pests and pathogens, gradual erosion of root systems, bank erosion, blowdown, re, or massive failure of banks or hillslopes. When tree death does not directly place wood in the stream channel, transport processes such as debris ows, landslides, oods, and downhill creep can move wood into the stream channel (Harmon et al., 1986; Sedell et al., 1988; Benke and Wallace, 1990; Gregory et al., 2003a). A growing literature is quantifying these processes, the sources of stream wood, and the patterns of variability in the delivery of stream wood (Murphy and Koski, 1989; Benke and Wallace, 1990; Naiman et al., 2000; Burnett, 2001; Martin and Benda, 2001; Benda et al., 2002; May and Gresswell, 2003; Montgomery et al., 2003a; Montgomery et al., 2003b; Reeves et al., 2003; Allan, 2004; Balian and Naiman, 2005; Bragg, 2000). The picture that emerges is that the relative roles of these processes vary across the stream network and landscape, as does the relative contribution of wood from upland as opposed to riparian sources. Consistent with the important roles that wood plays in streams, national and regional stream monitoring protocols quantify wood in streams. While these data provide information on the current status of streams and insight on the effectiveness of current and past regional management, they do not provide information on the potential future contribution of wood to streams. Assessment of the potential for future contributions of wood to the stream network requires evaluation of riparian and upland systems. The analysis reported here focuses only on the riparian portion of the landscape which is appropriate to provide an accounting for this function required by Federal, regional and state authorities (USDA/FS and DOI/BLM, 1994; Oregon Department of Forestry, 1996; Phillips et al., 2000; Young, 2000).

Existing regional monitoring protocols for riparian ecosystems reect a general recognition that riparian ecosystems provide wood to the stream network, rather than a specic measurement explicitly based on a specic understanding of how riparian forests contribute wood to the stream network. For example, the Northwest Forest Plans aquatic and riparian effectiveness monitoring plan (Reeves et al., 2004) notes that wood delivery is a key process. The plan proposes to acquire information on this process by securing estimates of the proportions of watersheds in various seral stages. This information would be provided by remotely sensed imagery. EPAs Environmental Monitoring and Assessment Program (EMAP) acquires eld observations on the presence and characteristics (size class and deciduous or conifer) of the vegetation within 10 m of the stream (Kaufmann, 2006). The USGS National Water Quality Assessment Program (NAWQA) develops information on stem density of all woody species using eld observationsspecically the point-centered quarter (PCQ) method (Fitzpatrick et al., 1998). As for any ecological indicator, the development of an indicator of future stream wood contributions should reect ecological processes (e.g. Cairns et al., 1993; Soule, 1995; Jackson et al., 2000; Dale and Beyeler, 2001). Because ecological models are an explicit representation of our understanding of ecological processes, their use in indicator design enables the linkage of ecological processes with indicator development. Gregory et al. (2003b) lists 12 models which forecast future contributions of wood from riparian ecosystems to stream channels. Each of these models requires a characterization of the riparian forest on which model representations of ecosystem processes operate in order to make predictions of potential future contributions of wood to the stream. The riparian forest data required to support these models typically includes information on the taxonomic composition of the trees, the distance of each tree from the stream bank, the height of each tree, and the number trees in the stand. While model predictions provide insight into potential future contributions of wood to streams, these models can require large amounts of detailed eld data requiring considerable time to collect. Riparian forest data can be collected from direct eld observation or from extensive data bases. Direct eld observation offers the most exibility and certainty for site specic observations and many sources describe eld methods for specic attributes and the levels of certainty that can be expected with these methods (e.g. Barker et al., 2002a; Mueller-Dombois and Ellenberg, 2002; Gray and Azuma, 2005). However, eld crews are expensive in terms of both direct and indirect or opportunity costs. Extensive data, typically including remotely sensed information, allows for the assessment of large areas, but has less exibility than does eld data collection in the attributes and spatial character of the data. The number of options available to provide riparian characterization from remotely sensed data is growing rapidly (e.g. Fassnacht et al., 2006). One widely used source for extensive landcover data is Thematic Mapper (a multispectral sensor TM). Information from this sensor has been used to provide landcover classications for millions of hectares in 30 m pixels. Others have used data from TM and from SPOT (a sensor similar to TM, but with 10 m resolution) to develop regressions between remotely sensed data and

464

ecological indicators 9 (2009) 462475

attributes (e.g. stand height) of Douglas-Fir/Western Hemlock stands in the western Oregon Cascades (Cohen and Spies, 1992). Although the information provided by these sensors has a specic spatial resolution, developers of its products typically suggest using it at a coarser level of resolution, e.g. for a 3 3 block of pixels (Cohen and Spies, 1992; Congalton and Green, 1999; Ohmann and Gregory, 2002). Spatial accuracy, in addition to spatial resolution, is a consideration in the use of extensive geographic data. For example, IKONOS imagery (4 m resolution multispectral imagery) can be purchased with four different levels of spatial accuracy ranging from RMSE of 11.8 to 0.9 m (Space Imaging, 2006) with an acquisition cost inversely proportional to its spatial accuracy. Given that there are divergent methods and a wide range of options for collecting riparian forest data that could be used in stream wood contribution models, we pose two questions. First: does simpler data change model predictions? If simpler data provides the same prediction as more detailed data, then perhaps model predictions can be made with less effort. Second, which metrics are well correlated with best model predictions? If such metrics can be identied, and if these measures are simpler to collect than the full set of data required for model predictions, then these parsimonious measures, although not allowing for the models to be operated, can serve as an indicator of the potential of a riparian stand to provide wood to the stream network. The approach to addressing these two questions is shown in Fig. 1. To respond to the rst question, best and simplied model input data sets were created and used as input for two different models (Van Sickle and Gregory, 1990; Welty et al., 2002). The models forecast the potential of riparian forests to provide wood to the stream and compare the predictions

arising from different data. Data were simplied by representing stands with class averages, by reducing spatial resolution, or by adding spatial error. To respond to the second question, correlations between simulated metrics, features that could easily be observed by eld crews or in remotely sensed imagery, and best model predictions were evaluated.

2.
2.1.

Methods
Detailed eld observations

Detailed eld observations were made at 109 forested riparian plots in northwestern Oregon. The description of the sampling and the sites is described in (Barker et al., 2002a; Barker et al., 2002b); therefore only an abbreviated description is provided here. Sample plots were selected on a stratied random basis from the stream network of this region. Strata were dened by three ecoregions (Coast Range, Willamette Valley, and West Cascades) (Clarke and Bryce, 1997) and four vegetation classes (Broadleaf and other, Small Conifer/Mixed, Medium Conifer/ Mixed, and Large or Very Large Conifer/Mixed). Measurements in each plot (0.16 ha 40 m 40 m) included the size (in diameter at breast height or DBH), location (including true horizontal distance from the streams bankfull edge), and taxonomic identity of each tree greater than 10 cm DBH. One to 4 h was required to collect this information for each plot; this excludes travel time to the plots; more time was required for plots with more trees. Sampling on 109 plots identied 6645 trees. The distribution of trees within 40 m of the stream is consistent with other reports (e.g. Gregory et al., 1991; Nierenberg and Hibbs, 2000; Russell and McBride, 2001; Wimberly and Spies, 2001) noting

Fig. 1 Approach to the analysis. Tables 713 are available in the Appendix.

ecological indicators 9 (2009) 462475

465

Fig. 2 Basal area (A) and tree stem density (B) as a function of distance from stream edge.

that basal area density (m2 per ha) of trees increases with distance from the stream, while the density of trees (number per ha) is stable. Deciduous trees (D) were more prevalent (in numbers and in basal area) closer to the stream; conifers (C) were more prevalent further from the stream. Large conifer (C5 those with dbh > 50 cm) were few in number, but evenly distributed as a function of distance from the stream. Deciduous trees were, on average, narrower and shorter than conifers. These observations are summarized in Fig. 2 and Table 1. The median bankfull width of the streams at the study sites was 6.7 m and ranged from 1.7 to 64 m.

These eld observations were used to estimate metrics that could be derived either by eld observation or remote sensing, and to provide model input as shown in Fig. 1.

2.2.

Simulated metrics

Two categories of candidate metrics were derived from the eld data. One reects metrics that could be developed by eld measurements, while the other is constructed around metrics that could be developed by remotely sensed imagery. These metrics were developed to support the second question: Which metrics are well correlated with best model predictions?

466

ecological indicators 9 (2009) 462475

Table 1 Characteristics of trees observed in riparian field plots Individual taxa


Douglas Fir (Pseudotsuga menziesii) Alder (Alnus spp.) Western Hemlock (Tsuga heterophylla) Maple (Acer macrophyllum) Red Cedar (Thuja plicata) California Laurel (Umbellularia californica) Silver Fir (Abies amabilis) Ash (Fraxinus latifolia) Yew (Taxus brevifolia) Mt. Hemlock (Tsuga mertensiana) Oak (Quercus garryana) Grand Fir (Abies grandis) Cottonwood (Populus balsamifera ssp. trichocarpa) Incense Cedar (Calocedrus decurrens) Noble Fir (Abies procera) Sitka Spruce (Picea sitchensis) Cherry (Prunus emarginata) Englemann Spruce (Picea engelmannii) Other Total

Number of individuals
2137 1812 911 466 327 223 166 105 75 71 50 39 33 32 28 27 23 2 118 6645

Number of plots
88 84 53 42 41 6 11 11 19 3 5 9 6 3 5 10 5 1 24 110

Mean diameter at breast height (cm)


35 25 30 30 43 22 21 35 19 43 23 44 24 25 33 71 19 60 17

Mean estimated height (m)


26 21 22 18 23 15 15 20 8 24 15 27 21 13 23 36 20 36 13

VSG group
3 1 5 2 4 2 3 1 4 5 2 3 1 4 3 6 1 6

RAIS group
1 3 2 4 2 4 1 3 2 2 4 1 3 2 1 1 3 1

Groups of trees
All trees All deciduous trees All conifer trees All trees >50 cm dbh Conifers >50 cm dbh

Number of individuals
6645 2811 3834 862 696

Number of plots

Mean diameter at breast height (cm)


30 26 34 77 82

Mean estimated height (m)


22 19 24 43 46

Effective tree height VSG (m)


17 14 19 38 41

The category Other includes 13 smaller species which were not included in the analyses.

2.2.1.

Simulated eld metrics

Metrics of tree number (N) and basal area (B) were derived from the detailed eld observations. These metrics described all trees (N or B) or large conifers (those with dbh = > 50 cm and noted as NC50 or BC50) over the 40 m breadth of the plots for varying cumulative distances from the stream bank in 5 m increments, i.e. 05, 010. . . through 040 m from the stream. Candidate eld metrics were derived by simulating two different eld methods one based on a count-plot (or xed area plot or FAP) method, and the second based on the PCQ method (Mueller-Dombois and Ellenberg, 2002). While the FAP method is considered the most robust method, the PCQ method is an efcient eld method providing results well correlated with the count-plot method if the trees are randomly distributed (Cottam and Curtis, 1956; MuellerDombois and Ellenberg, 2002). The PCQ method was simulated not only because of its efciency as a eld method, but also because of its use by the USGS to quantify riparian vegetation (Fitzpatrick et al., 1998). To simulate measures that could be derived using the count-plot method, eld observations for each plot were simply summarized for the appropriate set of trees and location. To estimate metrics that could be derived if eld crews were to have used the PCQ method, the placement of PCQ sampling points was simulated in the plots. This procedure is illustrated in Fig. 3. For example, to develop a PCQ estimate of riparian forest attributes in the 20 m closest to

the stream, two sample points, each 10 m from the stream, were dened. One was 10 m from the downstream plot edge, and the second was 10 m from the upstream plot edge. At each simulated PCQ sample point, the surrounding space was divided into four quadrants (A, B, C, and D). The quadrants were dened by two lines, one perpendicular to the stream and a second parallel to the stream. The lines intersected at the sample point. In each quadrant the distance to the nearest tree was calculated based on the xy coordinates in the eld data. Only trees within the plots radius of the sample point were used. The distances from the sample point to each tree were used to calculate tree number or basal area in the dened area as described by (Mueller-Dombois and Ellenberg, 2002). Correction factors developed by Warde and Petranka (1981) were used for sample points with no trees within one or more sample quadrants.

2.2.2.

Simulated remotely sensed metrics

Three forest attributes were derived from our eld data to simulate metrics that could be derived from remotely sensed imagery: the mean diameter at breast height, mean height (H), and tree density (N) (Cohen and Spies, 1992). These attributes were derived for two sets of trees: canopy (c) and canopy and subcanopy trees (c + s) for a total of six metrics. These estimates were developed only for the coarsest resolution allowed by the data sets 40 m 40 m. This dimension is roughly the size of the useful scale of a 3 3 pixel SPOT

ecological indicators 9 (2009) 462475

467

simulator describes the growth of the trees and simulates rates of mortality over time, particularly as a function of stand density. Thus, while the two models appear to predict the same thing, they are predictions that reect very different processes and are not directly or simply comparable to one another. The VSG model is used to describe the potential stream wood contribution given complete mortality of a current stand, while RAIS describes the contribution of stream wood that would result from chronic mortality over a specied period as the trees in the stand grow. Both models have been widely cited and used. The RAIS model, for example, is the foundation of much of the analysis of riparian management in Washington state (Washington Department of Natural Resources, 2001)

2.3.2.

Predictions

Fig. 3 Illustration of PCQ simulation.

imagery, and smaller than the 3 3 pixel TM imagery. We simulated these metrics by simple summaries of the eld data. We calculated tree height from DBH using regression models (Garman et al., 1995, Wang and Hann, 1988, or Larsen and Hann, 1987). c + s trees were dened using Kings (1966) methods. Because these metrics were simulated from eld data rather than from remotely sensed imagery, our analysis excludes errors that would be associated with the processing of imagery.

2.3. 2.3.1.

Modeling methods The models

The predictions from the VSG model were all wood, or conifer wood > 50 cm dbh, and were expressed in terms of the number (per 100 m of riparian forest) or the volume of pieces (m3 per 100 m of riparian forest) of pieces in each category. Volume described the wood volume within the bankfull channel. Predictions for RAIS were the cumulative number of potentially contributed pieces (per 100 m of riparian forest) of all wood or functional wood accumulated over 10, 50, and 100 year periods. Functional wood was dened by Bilby and Wards (1991) formulation which, for an 11 m channel our best case assumption required that a piece of functional wood must be at least 50 cm in diameter. The VSG and RAIS models predict potential stream wood contributions from riparian forests. The predictions are potential contributions because they are based on general assumptions not necessarily true in any specic location or time. In a reach or watershed assessment, these predictions could be modied by reach-specic information to provide more realistic estimates of the potential contribution of wood to the stream.

The VSG model (Van Sickle and Gregory, 1990) estimates the delivery of tree boles to the stream from sources of tree fall such as windthrow, decomposition, and stream bank erosion. Trees were assumed to fall independently of each other; multiple-tree input events such as debris ows and landslides were not considered. Estimates of wood input (bole number and volume) were modeled as functions of a riparian stands species mix, effective tree height, stem density, and of the assumed probabilities that trees fall (the assumption is made that all trees fall, thus simulating an episode of complete mortality of the current stand) and upon falling land in a userspecied direction. Tree height was modeled from DBH as described above. Effective tree height is 5 m less than tree height to account for small pieces of wood at the tops of trees that would not function in streams (Robison and Beschta, 1990; Van Sickle and Gregory, 1990). The model sums expected inputs from all trees in the stand to yield a mean and variance of total input from the stand; here results were presented based only on mean inputs. The RAIS model (Riparian Aquatic Interaction Simulator) (Welty et al., 2002) was chosen to evaluate the implications of stand growth and future mortality. This model combines the mechanisms of tree fall incorporated in the VSG model with a tree growth model, ORGANON (Hann et al., 1995). This

2.3.3. Model assumptions and variations in model assumptions


Model stands were assumed to have a width of 40 m, divided into four equal 10 m wide subplots as illustrated in Fig. 4A. Streams were assumed to have a bankfull channel width of 11 m with trees falling in any direction with equal probability. The VSG model tracks seven taxonomic groups of trees. The RAIS model tracks four taxonomic groups. The reassignments from observed taxa to these seven or four groups are provided in Table 1. Sensitivity of the predictions to variations in these model structural assumptions was evaluated by examining predictions for different channel widths (2 m, 30 m, and observed width), for a different subplot structure of the models (four subplots of unequal width5 m, 5 m, 10 m and 20 m width with the smaller subplots closer to the stream), and for the assumption that trees fell towards the stream rather than at random.

2.4.

Model input data

Input for the models was developed from the original highly detailed eld data in six versions: the best case, and ve

468

ecological indicators 9 (2009) 462475

Fig. 4 Illustration of the Data Simplification Cases.

simplications of the best case. Comparisons were made between model predictions based on different input data for six pairs of the cases shown in Table 2. The specic rules generating each of these cases from the original highly detailed eld data

are described in the following sections along with an illustration of the application of these rules provided for one plot in Fig. 4. In this gure D is a deciduous tree, C is a conifer. The size of the letter denotes the size of the tree.

ecological indicators 9 (2009) 462475

469

Table 2 Listing of the cases compared and the information loss evaluated in each comparison Case with less information
40 m 40 m 10 m 40 m 40 m 40 m class class class class offset 10 m

Case with more information


10 m 10 m 10 m 10 m class 40 m 10 m

Information loss evaluated


Spatial resolution Spatial resolution and stand classication Stand classication Spatial resolution Stand classication Geo-registration error

2.4.1.

Generating the best input data

In the best, or 10 m case, trees were assigned to 10 m 40 m model subplots based on their observed presence in those subplots. The plot is properly registered with regard to the location of the stream. The best case input data is depicted in Fig. 4A.

provides the maximum difference from a properly registered plot. If the displacement were parallel to the stream rather than perpendicular to it, there would be no meaningful error.

2.5.

Analytical methods

2.4.2.

Generating input data with coarser spatial resolution

2.5.1. Linkages between model predictions and simulated metrics


Spearman correlations, r, were calculated to quantify associations between simulated metrics and best case model predictions. To evaluate the generality of the results the correlation analysis was conducted not only for all sites, but also for subsets of the plots associated with specic ecoregions or assigned to different forest stand classes. To be included in the analysis subsets of the data were required to contain at least 25 plots. The generality of the results was also assessed by calculating r for model predictions based on variations in model structure.

In the coarser resolution, or 40 m case, the plot average values for numbers, types, and sizes of trees were assigned to each 10 m 40 m subplot. In the example plot depicted in Fig. 4 there were 40 trees in the plot. The number of trees in the subplots ranged from 5 to 18. When the information for these four subplots is simplied to the 40 m case, the average number of trees per subplot, 10, is allocated to each 10 m subplot as shown in Fig. 4C.

2.4.3.

Generating two cases based on forest stand classes

To provide descriptions of forest classes, each 40 m 40 m plot and each 40 m 10 m subplot was assigned to one of 13 forest classes. Twelve classes were dened by the combination of the size of the dominant and co-dominant trees (small <25 cm dbh; medium 2550 cm; large >5075 cm; and very large >75 cm) and the stand composition (conifer >70% conifer basal area, deciduous >70% deciduous, and mixed all other stands). Dominant and co-dominant trees were identied following the method of King (1966) a 13th class had no trees. Average tree density, tree size, and stand composition were calculated for each class in the class cases. Those average attributes were assigned to each plot or subplot of the class. Descriptions were generated at two spatial resolutions10 m and 40 m. Specic examples of these simplications are shown in Fig. 4. The stand classication assignment for each subplot is shown in Fig. 4B. One subplot is classied as very large mixed. In the full dataset 13 of the 109 plots were members of the very large mixed class; the average plot in this class has 37 trees; 10 were conifers. In creating a 10 m class dataset, that density of trees is assigned to subplots classied as very large mixed. The same process is used to transform the 40 m case to the 40 m class case (Fig. 4D), except that the accounting is done at the plot level rather than at the subplot level.

2.5.2.

Evaluation of the effects of data simplication

We established three criteria for deciding if predictions from a simplied data set are equivalent to predictions using the best case data. Two criteria are established for the reach scale and one for the network scale. The reach scale is the individual stream reach adjacent to a riparian stand this is the scale at which eld observations are typically made and reported. The network scale is an important scale because it may be the total amount of wood in a stream network that inuences stream function rather, than the wood at any one point within the stream network. At the reach scale, less detailed data may be appropriately used, so long as two conditions are met. First, if the prediction error is less than the error of measuring in-stream wood, and second, if r, the correlation between the two predictions, is greater than 0.7 (i.e. if half the variability in predictions from one set of assumptions is accounted for by predictions using another set of assumptions). Prediction error for potential stream wood uses the following formulation: ! P i1 to n log10 Xi log10 Y i 2 n

RMSEp SQRT

(1)

2.4.4. Generating cases with simulated errors in georegistration


To simulate the effect of a geo-registration error, a data set was created in which the position of the stream was displaced by 40 m. The result of this simulated displacement was that trees actually 40 m from the stream were listed as on the stream bank, and trees that were actually on the stream bank were listed as 40 m from the stream. This case is illustrated in Fig. 4E. This simulated 40 m displacement is in the direction that

Xi is the predicted contribution of stream wood from plot i in the case when the model is provided with less information, Yi is the predicted contribution of stream wood from plot i when the model is provided with more information, and n is the number of the eld plots. Measurement error for observations of stream wood is documented in this region (Kaufmann et al., 1999). These values serve as a benchmark and were designated as RMSEK. Predictions and measurements values of 0 were

470

ecological indicators 9 (2009) 462475

Table 3 Model predictions of potential stream wood contributions for the best case Amount Mean (per 100 m of stream length)
VSG predictions Total wood (number of pieces) Large conifer (number of pieces) Total wood volume (m3) Large conifer volume (m3) RAIS predictions Total wood (number of pieces) 10 years Total wood (number of pieces) 50 years Total wood (number of pieces) 100 years Functional wood (number of pieces) 10 years Functional wood (number of pieces) 50 years Functional wood (number of pieces) 100 years 23.3 5.1 10.1 6.9

S.D.
13.7 5.9 9.3 9.7

14.2 74.9 128.4 0.00 0.01 0.25

22.3 91.9 133.6 0.00 0.06 0.92

replaced with a value equal to 0.1 times the smallest nonzero measurement so that log values could be calculated for all cases. For the network scale, the simulation simply summed predictions across all plots. A bootstrap approach generated a network scale benchmark using paired visit data from Kaufmanns (1999) repeat visits, the data underlying RMSEK calculations. These pairs were sampled with replacement 109 times (matching the number of plots for which there were eld observations) at random. This sampling was repeated 1000 times and a benchmark for network relative error, NREK as dened in Eq. (2), was calculated. For each sample, one member of the pair of visits was assigned to measurement 1, M1 and the other to measurement 2, M2. For each set of 109 samples, we calculated the network relative error and averaged the result over the 1000 sets to give NREk NREK Average over 1000 sets of 109 samples of   jSM1 SM2 j AverageSM1 ; SM2

NREP 100%

! jSY j SX j j AverageSY j ; SX j

(3)

NREP is the network relative error for a particular comparison of cases; SYj and SXj were the total amounts of predicted stream wood summed over all plots given cases with respectively more and less information; j designates the case of interest. The analysis is based on a comparison of NREP to NREK. Our evaluation is an approximation of contributions to a stream network because the riparian data were collected from multiple ecoregions and watersheds rather than from the stream network of a single watershed.

3.
3.1.

Results
Model predictions

(2)

Network relative error was calculated for each comparison of data simplication cases as shown in Eq. (3).

Best model predictions for all plots are summarized in Table 3. The standard deviation describes the variability in the predictions across the 109 plots. Since best case RAIS predictions for functional wood for virtually all plots were zero for 10 and 50 years, analysis of RAIS functional wood

Table 4 Summary of model predictions in comparison to three evaluation criteria ( X) a (Y ) Model


VSG
b

40 ma 10 m
b

40 m classa 10 m
b

10 m classa 10 m
b

40 m classa 10 m class
b

40 m classa 40 m
b

40 m offset 10 ma 10 mb

Indicator type
All wood Large conifer Number Volume Number Volume 10 Years 50 Years 100 Years 100 Years Pass l and 3 Pass 3 l and 3 l and 3 3 All 2 l and 2 l and 3 1 All All 2 and 3 All 2 l and 3 l and 3 l and 3 All All All All

Pass or fail
2 3 3 3 All All l and 2 All 2 Pass l and 3 1 All All l and 2 All l and 2 All Pass 3 All All All l and 2

RAIS

Total wood

Functional wood

The use of simpler data can fail in comparison to any of three criteria: 1 Reach scale prediction error > reach scale measurement error; 2 low correlation between predictions; and 3 Network scale prediction error > network scale measurement error. a Case with less information. b Case with more information.

ecological indicators 9 (2009) 462475

471

Table 5 Maximum correlations between field observations of riparian status and model prediction metrics of riparian function Model predictions Simulated eld observations Correlation between eld metric and model prediction Fixed area plot
0.75 0.81 0.88 0.89 0.98 0.85 0.94

Metric
Total wood (number of pieces) Total wood (number of pieces) Total wood (number of pieces) Total wood (number of pieces) Large conifer (number of pieces) Total wood volume Large conifer volume

Time period
10 50 100

Model
RAIS RAIS RAIS VSG VSG VSG VSG

Metric
N N N N NC50 B BC50

Cumulative extent (m from stream)


10 10 20 15 40 20 30

PCQ
0.65 0.68 0.68 0.78 0.87 0.66 0.80

predictions were restricted to accumulations over 100 years. Predictions for variations on the best case assumptions are summarized in Appendix Table 7 for simplied data and Appendix Table 8 for variations in model structure.

3.1.1. Model prediction sensitivity to the level of detail in the input data
Only 4 of the 48 comparisons passed all three of our evaluation criteria (Table 4), i.e. network and reach prediction errors were

smaller than our benchmarks, and reach scale predictions were reasonably well correlated with best case predictions. Of the 44 cases failing, 27 failed both at reach and network scales, 11 failed only at the reach scale, and 6 failed only at the network scale. As at the reach scale the RAIS model was more sensitive to the loss of information than was the VSG model with all RAIS cases failing to meet one or more criteria while 83% of the VSG cases failed to meet the criteria. At the reach scale, 79% (38 of 48) of the comparisons did not meet the criteria, i.e. variations in the level of detail in the input data led to model predictions that were different from or poorly correlated with model predictions from the best case data (Table 4 and Appendix Table 9). In most instances (23), the comparisons failed to meet both the RMSE and r criteria. In ten cases, the failure was solely a failure to meet the RMSE criterion, i.e. RMSEP > RMSEK, and ve additional cases failed because the r was less than 0.7 ranging from 0.42 to 0.50 (see Appendix Table 10). Overall, information loss with the VSG model was more likely to result in predictions that met the criteria than the RAIS model (37% meeting the criteria vs. 4%). The best category of predictions was VSG Large Conifer, where half of the comparisons met the criterion. Examination of the columns of Table 4 and Appendix Table 9 showed that some types of information loss led to more frequently acceptable results than others. For example, half of the 10 m vs. 40 m resolution cases had acceptable results; none of the 10 m to 10 m class did. At the network scale, 69% (33 of 48) of the comparisons did not meet the criteria set for them (Table 4 and Appendix Table 11), i.e. variations in the level of detail in the input data led to model predictions that were different from model predictions from the best case data. Overall, information loss with the VSG model was more likely to result in predictions that met the criterion than the RAIS model (50% meeting the criteria vs. 13%). The best category of predictions was VSG All Wood number, where all cases met the criterion.

3.2. Indicators of the potential of a riparian forest to contribute wood to the stream network 3.2.1.
Fig. 5 Spearman correlation coefficients between best case model predictions and simulated fixed area plot field observations for six different subsets of the plots and two model predictions.

Simulated eld measures as indicators


observed in the eld using xed area plot well correlated with best-case model potential stream wood recruitment (see aspects of this result are noteworthy: rst,

Features easily methods were predictions of Table 5). Three

472

ecological indicators 9 (2009) 462475

the specic eld metrics that correlated best with the model predictions, second, the cumulative spatial extents where these correlations were maximized, and third the robust nature of these results across subsets of the data and across varying assumptions of model structure. Observations that could be collected by using the point-center quarter (PCQ) method also correlated well with model predictions, but the magnitude of the correlation was on average 14 percentage points less than for xed area plot methods. Field metrics with highest r were logical matches to the model prediction metrics (Table 5): e.g. number of all trees in the riparian plots correlated well with the number of trees potentially contributed to the stream, and basal area of trees observed in the riparian plots paired with the volume of wood potentially contributed to the stream. The cumulative extent at which these correlations were maximized corresponded well to the effective height of the trees in the group of trees of interest. For example, the average estimated effective height of all trees is 17 m (Table 1); r was at a maximum when the model prediction of all wood ranges was correlated with eld observations that ranged from 0 to 10 through 0 to 20 m from the stream bank (Figs. 5 and 6). Similarly, the average estimated effective height of large conifers is 41 m (Table 1); r was at a maximum when model prediction of potential conifer contributions were correlated with eld observations that ranged from 0 to 30 through 0 to 40 m from the stream bank (Figs. 5 and 6). There was no substantial sensible correlation between eld observations and the three functional wood predictions, in part because so many of the plots were predicted to contribute no functional wood even at 100 years. As a result, there was little variability in model predictions to associate with the variability in eld observations. These results were robust in several ways, as illustrated in Figs. 5 and 6. First, variations in the cumulative distance of these eld observations resulted in correlations almost as high as the maximum correlations and changing the cumulative distance of the eld observation by 10 m usually led to small changes in r. Second, variations in model assumptions (different spatial structure, different stream widths, or different assumptions about tree fall direction) showed the same patterns in correlations as the best estimates did with the eld metrics see two examples of this in Fig. 6 and Appendix Table 12. Third, the pattern of maximum correlations was also robust across subsets of the data for ecoregion and vegetation class subsets of the data see two examples of this in Fig. 5.

Fig. 6 Spearman correlation coefficients between best case model predictions and field observations for five variations in model structure and two model predictions.

3.2.2.

Simulated remotely sensed measures

Features observable in remotely sensed imagery were well correlated with model prediction metrics of potential stream wood recruitment (Table 6). Two aspects of this result are noteworthy: rst, the specic metrics best correlated with the model predictions and second, the robust nature of these results across subsets of the data and across varying assumptions of model structure. The remotely sensed metrics

Table 6 Maximum correlations between observations of riparian status derived from simulations of metrics that could be derived from remotely sensed data and model prediction metrics Model prediction Metric
Number of pieces of wood Number of pieces of wood Number of pieces of wood Number of pieces of wood Number of pieces of large conifer wood Volume of all pieces of wood Volume of large conifer wood

Simulated remotely sensed metric Time period


10 50 100

Model
RAIS RAIS RAIS VSG VSG VSG VSG Nc + s Nc + s Nc + s Nc + s Hc + s Hcan Hc + s

Correlation between simulated metric and best model prediction


0.57 0.70 0.77 0.82 0.81 0.69 0.77

ecological indicators 9 (2009) 462475

473

that correlated with the best model estimates were either average tree height or stand density for canopy and subcanopy trees or just canopy trees. Simulated remotely sensed measures were well correlated with model predictions across ecoregion and vegetation subsets of the plots (the correlations for these subsets average 93% of the value of the correlations shown in Table 6 see Appendix Table 13), as was the case for the simulated eld measures.

4.

Discussion and conclusions

In general, model predictions derived from simplied data did not compare well with the best model predictions (Table 4). The important exceptions were the VSG model predictions for the number of pieces of wood and for number of pieces of large conifer wood. In these instances, a loss in spatial resolution (to 40 m from 10 m) led to VSG model predictions similar to (at both the reach and network scale) and well correlated with model predictions that had ner spatial resolution (see Table 4 and Appendix Tables 911). The implication is that where the processes embodied in the VSG model are appropriate, and when numbers of pieces of wood or of large conifer wood are an indicator useful for addressing assessment or management questions, then information based on less expensive efforts can provide useful results. For other types of information loss or simplication, or for other indicators, the resulting model predictions were different from or not well correlated with model predictions using more rened data. Of special note is that forest stand classication always leads to model predictions that were different from or not well correlated with the model predictions based on the direct enumeration of the characteristics of the forest stands. Simple eld measures were strongly correlated with model prediction metrics of the potential for riparian forests to provide wood to the stream network (see Table 5, Appendix Table 13, and Figs. 5 and 6). The measures were counts of the cumulative number of trees or basal area within specied distances of the stream edge. Metrics that can be derived from remotely sensed imagery were also well correlated with model predictions of the potential for riparian forests to provide wood to the stream (see Table 6 and Appendix Table 13). Thus metrics that can be obtained either from eld observations or from remotely sensed imagery can serve as an effective indicator of the potential of a riparian stand to provide wood to the stream network. An indicator of the potential of a riparian forest to provide wood to the stream network, whether derived from eld measures (and listed in Table 5) or by remotely sensed imagery (and listed in Table 6), would enable managers to track changes in the potential for a specic riparian stand or class of stands to provide wood to the stream network over time. Within a class of riparian stands, it would provide the capacity to rank the potential of sites to provide wood to the stream. Class denitions or boundaries would need to be delineated by classes of mortality and transport processes that could be associated with a stand. For example, in the best case, it is assumed that trees fall in a random direction. The sensitivity analysis shows that if all trees were to fall towards the stream, the indicator selected could effectively remain the same

(Fig. 6). However, the number of pieces (or the volume of wood) would more than double see Appendix Table 8. So, the indicator would not change, but the assessment of the stand might. A number of attributes, or context information, including hillslope, rooting condition, and species inuence tree fall direction (Sobota et al., 2006). As knowledge about the inuence of these factors improves, the usefulness of this indicator will increase. The use of additional, or context information, to support the interpretation of a metric is common. One example of this requirement is in the use of the Observed/Expected index used to evaluate the health of macroinvertebrate assemblages in streams (e.g. (Hawkins et al., 2000; Stoddard et al., 2005). The denominator for this index is developed on the basis of a set of calculations using candidate variables that describe the climate and geomorphic settings of stream reaches. In the case of potential stream wood contributions, the context variables in the models provide one way to approach the delineation of this additional information. These conclusions have some implications for the three riparian monitoring protocols used within this study region. Clearly, the PCQ method used by the NAWQA program could be easily adapted either to provide an indicator that would be correlated with the potential of a site to provide wood to the stream, or to provide input to a model, such as the VSG model, which provides predictions of the potential contribution of wood to the stream network. While remotely sensed imagery is capable of providing extensive information on forest cover with known certainty (Cohen et al., 1995; Stehman et al., 2003) the seral stage information to be derived from Northwest Forest Plan monitoring does not appear to be useful for driving models such the VSG or RAIS models. However, counts of trees and their size can be derived from remotely sensed imagery (e.g. Cohen and Spies, 1992; Ohmann and Gregory, 2002; Fassnacht et al., 2006) and this information is of more direct value in supporting assessments of riparian areas based on the processes reected in the VSG model. Last, the EMAP protocol (Peck et al., 2006) does not provide direct support for developing the eld metrics identied in this analysis.

Acknowledgements
This research would have been impossible without the efforts of the eld crew members who collected the eld data upon which this effort is based. In addition to authors (M.B. and J.B.): Brooke Abbruzzese, Mary Barczac, Chris Brugato, Adrien Elseroad, Andy Herstrom, Christy Larson, Lynn McAllister, Kouya Nester, Jennifer Sackinger, Greg Verret, and Ann Versluis. This paper has also beneted by comments from John Faustini, Phil Kaufmann, Kelly Burnett, and several anonymous reviewers as well as from the editorial assistance of Sarah L. Ringold.

Appendix A. Supplementary data


Supplementary data associated with this article can be found, in the online version, at doi:10.1016/j.ecolind.2008.06.009.

474

ecological indicators 9 (2009) 462475

references

Allan, J.D., 2004. Landscapes and riverscapes: the inuence of land use on stream ecosystems. Annual Review of Ecology, Evolution, and Systematics 35, 257284. Balian, E., Naiman, R., 2005. Abundance and production of riparian trees in the lowland oodplain of the Queets river, Washington. Ecosystems 8, 841861. Barker, J., Bollman, M., Ringold, P., Sackinger, J., Cline, S., 2002a. Evaluation of metric precision for a riparian forest survey. Environmental Monitoring and Assessment 75, 5172. Barker, J.R., Ringold, P., Bollman, M., 2002b. Patterns of tree dominance in coniferous riparian forests. Forest Ecology and Management 166, 311329. Benda, L.E., Bigelow, P., Worsley, T., 2002. Recruitment of wood to streams in old-growth and second-growth redwood forests, northern California, USA. Canadian Journal of Forest Research 32, 14601477. Benke, A.C., Wallace, J.B., 1990. Wood dynamics in coastal plain blackwater streams. Canadian Journal of Fisheries and Aquatic Sciences 47, 9299. Bilby, R.E., Ward, J.W., 1991. Characteristics and functions of large woody debris in streams draining old-growth, clearcut, and second-growth forests. Canadian Journal of Fisheries and Aquatic Sciences 48, 24992508. Bilby, R.E., Bisson, P.A., 1998. Function and distribution of large woody debris (Pages 324346). In: Naiman, R.J., Bilby, R.E. (Eds.), River Ecology and Management. Springer, New York, p. 705. Bragg, D.C., 2000. Simulating catastrophic and individualistic large woody debris recruitment for a small riparian system. Ecology 8, 13831394. Burnett, K. 2001. Relationships among juvenile anadromous salmonids, their freshwater habitat and landscape characteristics over multiple years and spatial scales in the elk river, Oregon. Ph.D. Dissertation thesis, Oregon State University, Corvallis, OR. Cairns, J., McCormick, P., Niederlehner, B., 1993. A proposed framework for developing indicators of ecosystem health. Hydrobiologia 263, 144. Clarke, S.E., Bryce, S.A., 1997. Hierarchical subdivisions of the columbia plateau and blue mountains ecoregions, Oregon and Washington, USDA Forest Service, Portland, OR, General Technical Report PNW-GTR-395, 114 pp. Cohen, W.B., Spies, T.A., 1992. Estimating structural attributes of Douglas-Fir/Western Hemlock forest stands from Landsat and SPOT imagery. Remote Sensing and the Environment 41, 117. Cohen, W.B., Spies, T.A., Fiorella, M., 1995. Estimating the age and structure of forests in a multi-ownership landscape of western Oregon, U.S.A. International Journal of Remote Sensing 16, 721746. Congalton, R.G., Green, K., 1999. Assessing the Accuracy of Remotely Sensed Data: Principles and Practices: Mapping Sciences Series. Lewis Publishers, Boca Raton, FL, 137 pp. Cottam, G., Curtis, J.T., 1956. The use of distance measures in phytosociological sampling. Ecology 37, 451460. Dale, V.H., Beyeler, S.C., 2001. Challenges in the development and use of ecological indicators. Ecological Indicators 1, 3 10. Fassnacht, K.S., Cohen, W.B., Spies, T.A., 2006. Key issues in making and using satellite-based maps in ecology: a primer. Forest Ecology and Management 222, 167181. Fitzpatrick, F.A., Waite, I., Darconte, P.J., Meador, M.R., Maupin, M.A., Gurtz, M.E., 1998. Revised Methods for Characterizing Stream Habitat in the National Water-Quality Assessment Program, US Department of the Interior, US Geological

Survey, Water Resources Division, Raleigh, North Carolina, Pages 67 + vii. Garman, S.L., Acker, S.A., Ohmann, J.L., Spies, T.A., 1995. Asymptotic height-diameter equations for twenty-four tree species in western Oregon. In: Forest Research Laboratory, Oregon State University, Corvallis, OR, 22 pp. Gray, A.N., Azuma, D.L., 2005. Repeatability and implementation of a forest vegetation indicator. Ecological Indicators 5, 5771. Gregory, S.V., Boyer, K.L., Gurnell, A.M., 2003a. The Ecology and Management of Wood in World Rivers. Symosium 37. American Fisheries Society, Bethesda, MD, 431 pp. Gregory, S.V., Meleason, M.A., Sobota, D.J., 2003b. Modeling the dynamics of wood in streams and rivers (Pages 315335). In: Gregory, S.V., Boyer, K.L., Gurnell, A.M. (Eds.), The Ecology and Management of Wood in World Rivers, Symposium 37. American Fisheries Society, Bethesda, MD, 431 pp. Gregory, S.V., Swanson, F.J., McKee, W.A., Cummins, K.W., 1991. An ecosystem perspective of riparian zones. BioScience 41, 540551. Gurnell, A.M., Gregory, K.J., 1995. Interactions between seminatural vegetation and hydrogeomorphological processes. Geomorphology 13, 4969. Hann, D.W., Hester, A.S., Olsen, C.L., 1995. ORGANON Users Manual, Ed.5.0. Department of Forest Resources, Oregon State University, Corvallis, OR, 127 pp. Harmon, R.E., Franklin, J., Swanson, F.J., Sollins, P., Gregory, S.V., Lattin, J.D., Anderson, N.H., Cline, S.P., Aumen, N.G., Sedell, J.R., Lienkaemper, G.W., Cromack Jr., K., Cummins, K.W., 1986. Ecology of coarse woody debris in temperate ecosystems. In: Macfadyen, A., Ford, E.D. (Eds.), Advances in Ecological Research. Academic Press, Orlando, FL, pp. 133302. Hawkins, C.P., Norris, R.H., Hogue, J.N., Feminella, J.W., 2000. Development and evaluation of predictive models for measuring the biological integrity of streams. Ecological Applications 10, 14561477. Hyatt, T.L., Naiman, R.J., 2001. The residence time of large woody debris in the Queets river, Washington, USA. Ecological Applications 11, 191202. Jackson, L.E., Kurtz, J.C., Fisher, W.S., 2000. Evaluation Guidelines for Ecological Indicators. EPA/620/R-99/005, U.S. Environmental Protection Agency, Ofce of Research and Development, Research Triangle Park, NC, 107 pp. Kaufmann, P.R., 2006. Physical habitat characterization (Pages 107164). In: Peck, D.V., Herlihy, A.T., Hill, B.H., Hughes, R.M., Kaufmann, P.R., Klemm, D.J., Lazorchak, J.M. (Eds.), EMAP Surface Waters: Western Pilot Study Field Operations Manual for Wadeable Streams. Washington, D.C., U.S. Environmental Protection Agency, Ofce of Research and Development. Kaufmann, P.R., Levine, P., Robison, E.G., Seeliger, C., Peck, D.V., 1999. Quantifying Physical Habitat in Wadeable Streams EPA/620/R-99/003. U.S. Environmental Protection Agency, Washington, D.C., 125 pp. + Appendices. Keller, E.A., Swanson, F.J., 1979. Effects of large organic material on channel form and uvial processes. Earth Surface Processes 4, 361380. King, J.E., 1966. Site Index Curves for Douglas-Fir in the Pacic Northwest. Weyerhaueser Forestry Research Center, Centralia, WA, 49 pp. Larsen, D.R., Hann, D.W., 1987. Height-Diameter Equations for Seventeen Tree Species in Southwest Oregon. Oregon State University, College of Forestry, Corvallis, Oregon, 16 pp. Martin, D.J., Benda, L.E., 2001. Patterns of Instream Wood Recruitment and Transport at the Watershed Scale, 130. Transactions of the American Fisheries Society, pp. 940958. May, C.L., Gresswell, R.E., 2003. Large wood recruitment and redistribution in headwater streams in the southern Oregon

ecological indicators 9 (2009) 462475

475

coast range, U.S.A. Canadian Journal of Forest Research 33, 13521362. Montgomery, D.R., Collins, B.D., Bufngton, J.M., Abbe, T.B., 2003a. Geomorphic effects of wood in rivers (Pages 2148). In: Gregory, S.V., Boyer, K.L., Gurnell, A.M. (Eds.), The Ecology and Management of Wood in World Rivers. Symposium 37. American Fisheries Society, Bethesda, MD, 431 pp. Montgomery, D.R., Massong, T.M., Hawley, S.C.S., 2003b. Inuence of debris ows and log jams on the location of pools and alluvial channel reaches, Oregon coast range. Geological Society of American Bulletin 115, 7888. Mueller-Dombois, D., Ellenberg, H., 2002. Aims and Methods of Vegetation Ecology. The Blackburn Press, Caldwell, New Jersey, 547 pp. Murphy, M.S., Koski, K.V., 1989. Input and depletion of woody debris in Alaska streams and implications for streamside management. North American Journal of Fisheries Management 9, 427436. Naiman, R.J., Bilby, R.E., Bisson, P.A., 2000. Riparian ecology and management in the Pacic coastal rain forest. BioScience 50, 9961011. Nierenberg, T.R., Hibbs, D.E., 2000. A characterization of unmanaged riparian areas in the central coast range of western Oregon. Forest Ecology and Management 129, 195206. Ohmann, J.L., Gregory, M.J., 2002. Predictive mapping of forest composition and structure with direct gradient analysis and nearestneighbor imputation in coastal Oregon, U.S.A. Canadian Journal of Forest Research 32, 725741. Oregon Department of Forestry. 1996. Forest practices administrative rules, Salem, Oregon. Peck, D.V., Herlihy, A.T., Hill, B.H., Hughes, R.M., Kaufmann, P.R., Klemm, D.J., Lazorchak, J., McCormick, F.H., Peterson, S.A., Ringold, P.L., Magee, T.K., Cappaert, M., 2006. Environmental Monitoring and Assessment Program Surface Waters: Western Pilot Study: Field operations Manual for Wadeable Streams, EPA/620/R-06/003, 257 + Appendices. Phillips, M.J., Swift Jr., L.W., Blinn, C.R., 2000. Best management practices for riparian areas (Pages 273286). In: Verry, E., Hornbeck, J.W., Dolloff, C.A. (Eds.), Riparian Management in Forests of the Continental Eastern United States. Lewis, Boca Raton, FL, 402 pp. Reeves, G.H., Burnett, K., McGarry, E.V., 2003. Sources of large wood in the mainstem of a fourth order watershed in coastal Oregon. Canadian Journal of Forestry 33, 13631370. Reeves, G.H., Hohler, D.B., Larsen, D.P., Busch, D.E., Kratz, K., Reynolds, K., Stein, K.F., Atzet, T., Hays, P., Tehan, M., 2004. Effectiveness Monitoring for the Aquatic and Riparian Component of the Northwest Forest Plan: Conceptual Framework and Options. U.S. Department of Agriculture, Forest Service, Pacic Northwest Research Station, Corvallis, OR, 71 pp. Robison, E.G., Beschta, R.L., 1990. Identifying trees in riparian areas that can provide coarse woody debris to streams. Forest Science 36, 790801. Russell, W.H., McBride, J.R., 2001. The relative importance of re and watercourse proximity in determining stand

composition in mixed conifer riparian forests. Forest Ecology and Management 150, 259265. Sedell, J.R., Bisson, P.A., Swanson, F.J., Gregory, S.V., 1988. What we know about large trees that fall into streams and rivers (Pages 4782). In: Maser, C., Tarrant, R.F., Trappe, J.M., Franklin, J. (Eds.), From the Forest to the Sea: A Story of Fallen Trees. USDA FS GTR PNW-229, Portland, OR, 153 pp. Sobota, D.J., Gregory, S.V., Van Sickle, J., 2006. Riparian tree fall directionality and modeling large wood recruitment to streams. Canadian Journal of Forest Research 36, 12431254. Soule, M.E., 1995. Biodiversity indicators in California: Taking natures temperature. California Agriculture 49, 4044. Space Imaging. 2006. Imagery products. http:// www.spaceimaging.com/products/imagery.htm. Stehman, S.V., Wickham, J.D., Smith, J.H., Yang, L., 2003. Thematic accuracy of the 1992 national land-cover data for the eastern United States: statistical methodology and regional results. Remote Sensing of Environment 86, 500516. Stoddard, J.L., Peck, D.V., Olsen, A.R., Larsen, D.P., Van Sickle, J., Hawkins, C.P., Hughes, R.M., Whittier, T.R., Lomnicky, G., Herlihy, A.T., Kaufmann, P.R., Peterson, S.A., Ringold, P.L., Paulsen, S.G., and Blair, R., 2005. Environmental montoring and assessment program (EMAP): Western streams and rivers statistical summary, EPA-620/R-05/006, 1762. USDA/FS, and DOI/BLM. 1994. Record of decision for amendments to forest service and bureau of land management planning documents within the range of the northern spotted owl and standards and guidelines for management of habitat for late-successional and oldgrowth forest related species within the range of the northern spotted owl.74 pp. + Appendices. Van Der Nat, D., Tockner, K., Edwards, P., Ward, J., 2003. Large wood dynamics of complex alpine river oodplains. Journal of the North American Benthological Society 22, 3550. Van Sickle, J., Gregory, S.V., 1990. Modeling inputs of large woody debris to streams from falling trees. Canadian Journal of Fisheries and Aquatic Sciences 20, 15931601. Wang, C., Hann, D.W., 1988. Height-Diameter Equations for Sixteen Tree Species in the Central Western Willamette Valley of Oregon. Oregon State University, College of Forestry, Corvallis, OR, 7 pp. Warde, W., Petranka, J.W., 1981. A correction factor table for missing point-center quarter data. Ecology 62, 491494. Washington Department of Natural Resources, 2001. Final EIS for forest practices rule changes: Chapter 3.4 riparian habitats. Olympia, WA, 53 pp. Welty, J.J., Beechie, T., Sullivan, K., Hyink, D.M., Bilby, R.E., Andrus, C., Pess, G., 2002. Riparian aquatic interaction simulator (RAIS): a model of riparian forest dynamics for the generation of large woody debris and shade. Forest Ecology and Management 162, 299318. Wimberly, M., Spies, T., 2001. Inuences of environment and disturbance on forest patterns in coastal Oregon watersheds. Ecology 82, 14431459. Young, K.A., 2000. Riparian zone management in the Pacic Northwest: whos cutting what? Environmental Management 26, 131144.

Das könnte Ihnen auch gefallen