United States

Department of

Agriculture

Forest

Service

Northern

Research Station

General Technical

Report NRS-29

Woody Detritus Density and 

Density Reduction Factors for 

Tree Species in the United States: 

A Synthesis

Mark E. Harmon, Christopher W. Woodall, 

Becky Fasth, and Jay Sexton
Abstract

Woody detritus biomass estimates are most often achieved by determining the 

volume of dead wood and then converting to mass by use of density values. 

There have been few studies on how density (mass/volume) of this material 

changes during the decay process. The goal of this study was to synthesize 

both published and unpublished data on woody detritus density so as to improve 

estimates of coarse woody detritus (CWD) and fi ne woody detritus (FWD) 

biomass across the diverse forests of the United States. In the case of CWD, a 

total of 88 species were found to have data on densities for fi ve decay classes 

that had been published and/or collected in North America from the boreal to 

the tropical zones. In general, density declined from decay class 1 to 5 with at 

least fi ve density reduction patterns observed. For FWD, our search indicated 

approximately 25 species had been sampled. FWD density was a function of 

piece diameter and the general state of decay. We determined that by sampling 

representative species within a genus, the uncertainty of CWD estimates could 

be reduced by up to 50 percent over not having sampled a genus. Our analysis 

indicated that the uncertainty of FWD mass estimation ranged from 12 to 19 

percent when FWD relative density was estimated. We conclude that a more 

systematic sampling of CWD and FWD density is needed for major species if 

uncertainties of mass estimates nationwide are to be reduced.

The Authors

MARK E. HARMON, Richardson Chair and Professor, Department of Forest 

Science, Oregon State University 

CHRISTOPHER W. WOODALL, Research Forester, U.S. Forest Service, 

Northern Research Station

BECKY FASTH, Research Assistant, Department of Forest Science, Oregon 

State University

JAY SEXTON, Senior Research Assistant, Department of Forest Science, 

Oregon State University

Manuscript received for publication December 2007

Visit our homepage at: http://www.nrs.fs.fed.us/

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June 2008 Fax: 740-368-0152
CONTENTS

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

 Importance of Woody Detritus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

 Measurement of Woody Detritus . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

 Objectives. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Data, Methods, and Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 Data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

 Field Methods Common to Most Studies. . . . . . . . . . . . . . . . . . . . . 4

 Steps in Estimating Dead Wood Mass . . . . . . . . . . . . . . . . . . . . 4

 Assessing the State of Decay of Woody Detritus . . . . . . . . . . . . 4

 Density Determination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

 Mass Determination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

 Data Processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 CWD Predictions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 FWD Predictions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

 Analysis of Uncertainty on CWD Mass Estimates . . . . . . . . . . . 9

 Analysis of Uncertainty on FWD Mass Estimates. . . . . . . . . . . 10

 Biomass to Carbon Conversion Factors. . . . . . . . . . . . . . . . . . .11

Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

 Qualitative Changes in CWD Decay Classes. . . . . . . . . . . . . . . . . 12

 Density of Decomposing CWD . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

 CWD Density Reduction Patterns . . . . . . . . . . . . . . . . . . . . . . . . . 14

 Fine Woody Debris Density . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

 Uncertainties of CWD Mass Estimates. . . . . . . . . . . . . . . . . . . . . . 23

 Uncertainties of FWD Mass Estimates . . . . . . . . . . . . . . . . . . . . . . 24

 Biomass to Carbon Conversion Factors. . . . . . . . . . . . . . . . . . . . . 25

Databases of Density Estimates. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27

Examples of Use . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

 Within a Decay or Size Class for a Species and Site. . . . . . . . . . . 29

 Within a Species and Site. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

 Within a Species for All Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

 Within a Site . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

 All Sites. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

(Contents continued on next page)
Future Needs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Literature Cited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Appendices. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

 Appendix 1 Literature sources. . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

 Appendix 2 CWD absolute density . . . . . . . . . . . . . . . . . . . . . . . . 39

 Appendix 3 CWD relative density . . . . . . . . . . . . . . . . . . . . . . . . . 46

 Appendix 4 FWD absolute and relative density. . . . . . . . . . . . . . . 54

 Appendix 5 Examples of estimates 1 . . . . . . . . . . . . . . . . . . . . . . 81

 Appendix 6 Examples of estimates 2 . . . . . . . . . . . . . . . . . . . . . . 83

CONTENTS (continued)
1

INTRODUCTION

IMPORTANCE OF WOODY DETRITUS

Woody detritus or dead wood is an important part of forest ecosystems. Although 

woody detritus is associated with many ecological benefi ts (plant and animal 

habitat, carbon, nutrient and water storage, as well as soil formation), the 

magnitude of these benefi ts varies with the species of log, the environment 

(i.e., climate, soil characteristics, topography), and the amount of dead wood on 

a site (Franklin et al. 1987, Harmon et al. 1986, Triska and Cromack 1980). The 

same is true for possible negative aspects of this material such as a source of 

fuel infl uencing fi re behavior and smoke emissions. Therefore, this material is 

increasingly being included in ecological studies and environmental assessments, 

and the number of publications on this subject is rapidly growing. 

Woody detritus takes many forms. It may range from sound to highly decayed 

materials. Woody detritus is present as roots, stumps, branches (including 

attached dead branches), standing dead (i.e., snags), downed material, and buried 

wood in litter, duff, and mineral soil. Very few inventories measure all these 

forms and size classes—snags and downed material being the most commonly 

measured. For example, the Forest Inventory and Analysis (FIA) program of the 

U.S. Forest Service conducts a national inventory of woody material including 

standing dead, FWD (fi ne woody detritus), and CWD (coarse woody detritus) 

(for more details, see Woodall and Monleon 2008). FWD with the exception 

of roots is typically less than 7.6 to 10 cm, the former size limit being based on 

lag-times of fuels (i.e., the time to equilibrate with the atmosphere in terms of 

moisture content; see Burgan 1988). For woody roots the size break is usually 

2 mm, which is based on conventions on the maximum size of live fi ne roots. 

CWD exceeds these diameters but also typically must exceed a length of 1 m. 

It can either be standing (e.g., snags) or downed (e.g., logs). 

MEASUREMENT OF WOODY DETRITUS 

Measurements of woody detritus abundance include the number of pieces per 

area, volume, mass, and cover (typically either basal area for snags or projected 

cover for logs). Many ecosystem functions, such as a fuel for fi re, depend on 
2

mass. However, there is a basic problem because the mass of woody detritus is 

not easy to directly measure unless the material is very small. Volume is easier 

to measure, and most methods, be they transect- or plot-based, measure piece 

dimensions to estimate volume (Brown 1974, Harmon and Sexton 1996). Mass 

can be derived from volume if the density (i.e., mass/volume) of this material 

is known. A similar problem exists with live trees because these are also rarely 

weighed directly. Rather, equations to predict volume from more easily measured 

dimensions such as diameter at breast height and/or height are established. 

Volume can then be converted to mass using the density of wood and bark. 

Unfortunately, estimation of woody detritus amounts is not as straightforward 

as for live trees. Woody detritus is often irregular in shape relative to live trees 

and therefore volume is estimated directly for each piece by techniques such as 

line intercepts or dimensional measures in plots. Moreover, rather than a single 

density value per species, the density of woody detritus pieces also depends 

on the extent of decomposition. This change can be dealt with by recognizing 

various decay classes of material and determining the density of each decay class. 

When decay class-specifi c density and elemental concentrations are coupled with 

the volume in each decay class, one can estimate the mass, carbon, and nutrient 

stores of woody detritus. 

The downed woody material (DWM) sampling design of the FIA program uses 

line-intersect transects to estimate the volume of downed CWD and FWD, but 

not their biomass (Woodall and Williams 2005). Biomass conversion constants 

published in the literature are currently used to convert volume to biomass. 

Two constants are used: (1) the initial density (i.e., specifi c gravity) and (2) a 

decay reduction factor that accounts for the decline in density as this form of 

detritus decomposes. Unfortunately, there are very few published values of the 

decay class-specifi c density or decay reduction factors; these have typically 

been available only for a few western tree species (Sollins et al. 1987). The 

little work that has been done has shown that species have different patterns of 

density decline (Yatskov et al. 1993). Thus, uncertainty in biomass estimates can 

be introduced if the incorrect pattern is assumed for a species that has not been 

studied. FIA’s own simulation studies have shown that a 15-percent variation in 

biomass conversion constants could affect a plot’s total DWM biomass estimate 

by 5 percent (Woodall and Lutes 2004). However, because FIA uses decay 

reduction factors based on a few western tree species for the entire Nation, 

conversion constants may vary in excess of 50 percent causing substantial error 

in biomass estimates.
3

OBJECTIVES

Given the fact woody detritus inventories are rapidly becoming a common 

component of forest resource inventories around the world, there is great need 

to refi ne the constants used in biomass estimation procedures for both CWD and 

FWD. While the current datasets on changes in wood density with decomposition 

are far from ideal, a substantial amount of published and unpublished data 

exists. Therefore, the goal of this study was to synthesize both published and 

unpublished data on woody detritus density and carbon content to improve 

estimates of CWD and FWD biomass and carbon across the diverse forests of the 

United States. The specifi c objectives of this study are to (1) estimate CWD and 

FWD density across decay classes for the tree species currently considered by 

the FIA inventory, (2) develop a methodology to estimate the uncertainty in these 

estimates, (3) provide examples of how these estimates can be used, (4) suggest 

future study directions to reduce uncertainty in these estimates, and (5) provide 

preliminary biomass to carbon conversion factors.
4

DATA, METHODS, AND ANALYSIS

DATA 

We assembled data from two sources. Over the years a substantial quantity 

of decayed wood density data has been generated by the Andrews Long-term 

Ecological Research (LTER) program. Some of this data has been published, but 

much of it has not, at least in terms of densities for decay classes. This program 

has also produced a large dataset on undecayed branch density as part of branch 

decomposition studies that have been conducted over the years. The other source 

was the published literature, which, although not as extensive as the fi rst source, 

does provide information for many systems not sampled (Appendix 1).

FIELD METHODS COMMON TO MOST STUDIES

Steps in Estimating Dead Wood Mass

Estimating the mass of decomposing wood involves several steps. Ideally one 

would weigh the mass of woody debris in an area, but realistically this would be 

time consuming and physically diffi cult. The fi rst step is to measure the density 

of the wood to be inventoried by sampling wood and bark in a range of decay 

states. The second step is to determine the volume of the woody detritus in the 

inventory area by decay classes. The third step is to convert the volume to mass 

using the decay class-specifi c density values. Finally, total mass can be converted 

to specifi c elemental stores by knowing the concentration of elements such as 

carbon and nitrogen. Unfortunately, volume inventories are often conducted 

without knowledge of how density or nutrient contents change with decay class. 

While some substitution of these conversion factors is inevitable, it has been 

common to use western U.S. conifer values for many temperate biomes and 

in some cases tropical ones. This introduces substantial uncertainty in woody 

detritus mass and other related estimates (e.g., carbon stocks, fuel loadings, 

smoke emissions).

Assessing the State of Decay of Woody Detritus 

Decay classes are a subjective way to break the decomposition process, which 

is a continuum, into recognizable stages. While continuous methods using 

techniques such as ordination exist (Harmon et al. 1987), they are very awkward 
5

to implement in the fi eld. Classes can be defi ned by the presence of criteria that 

tend to be associated with certain decay classes. However, there can be a great 

deal of variation within and between pieces of decomposing wood, and this may 

cause misclassifi cation of some individuals. By averaging over a large number of 

pieces, these errors tend to cancel out. 

The number of decay classes used varies from study to study. For CWD, as many 

as 13 classes and as few as 2 classes have been used in the past (Harmon et al. 

1986). One of the more common systems developed in the Pacifi c Northwest uses 

fi ve decay classes (Sollins 1982). For FWD, a single average number is often 

used, although a separation into undecayed or fresh versus decayed has also been 

used especially after recent disturbance (Brown 1974). The density of decayed 

FWD tends to be highly variable, and it is highly dependent on whether pieces 

are added in a pulse or continuously. When added as a pulse, many FWD pieces 

have similar densities that decrease through time. When continuously added, the 

average density of FWD is likely to be fairly consistent; however, there is a great 

range in densities depending on when a particular piece was added. 

It has been demonstrated that wood density generally declines as decay advances 

with some exceptions. For CWD, some species have very decay resistant 

heartwood (e.g., black locust, Robinia pseudoacacia). If the outer decayed 

sapwood and bark layers disappear via respiration or fragmentation and the 

residual heartwood has little decay, then more advanced decay classes can 

theoretically have an increase in density. The same can be true for FWD such as 

branches, which often lose decomposed outer wood as decomposition proceeds. 

This often results in an undecayed, resin-impregnated core of wood being left; as 

the proportion of this resin-impregnated wood increases the density can increase. 

Density Determination

Density is expressed as a dry mass divided by green volume in most cases, 

although density can be determined in alternative ways. For example, recently 

resistance drill systems have been used, although these are usually calibrated to 

traditional methods. For FWD it is typical to collect a random sample within an 

area or along transects. For CWD a number of logs, snags, or stumps of each 

decay class are located and then subsampled with a chainsaw to remove crosssections along the stem. Alternatively, a coring device is used to remove samples, 

although this can be used only for relatively sound wood. Key characteristics 

such as presence of leaves, twigs, branches, bark, cross-sectional shape, wood 

hardness, and strength are typically recorded. Volume is determined either by 

displacement in water or a particulate solid (e.g., millet seed), taking a known 

volume using a core, or by measuring external dimensions. While determining 

volume using water displacement is probably the most accurate method for solid 
6

samples, it is relatively slow and works best for small volumes. However, it is 

very diffi cult to use this method for highly decomposed wood and bark, which 

means either decay classes with this material are not included or only the more 

solid pieces are examined leading to an uncertainty or outright bias for more 

advanced stages of decay. Coring suffers from the same problem regarding the 

extent of decay present. While volumes determined by external dimensions are 

less precise, these measurements can be taken quite rapidly in the fi eld on very 

large volumes. This eliminates bias and leads to a better averaging of density 

within a piece. 

Mass Determination

Mass of samples is typically determined by drying in an oven, often at 

temperatures ranging between 55 and 75 oC until mass remains constant. This 

can take weeks for even small cross-sections. To speed up drying times, smaller 

subsamples are often used. This entails weighing the entire sample and then 

subsampling for moisture determination. The ratio of oven dried mass to the fresh 

mass of the subsample is used to convert the fresh mass to dry mass of the entire 

sample. 

As noted above, for FWD, decay classes are not usually noted, although green 

versus decayed material generally may be recorded separately. Subsamples are 

sometimes taken to determine density for a particular ecosystem, but this is rarely 

done and the default values presented in Brown (1974) are often used. When 

sampled, volume is determined by measuring dimensions of pieces or by volume 

displacement. Mass is determined via oven drying using approaches similar to 

that of CWD. 

ANALYSIS

Woody detritus density was expressed in two ways: absolute density (mass/green 

volume) and relative density (decayed density/undecayed density). Relative 

density is alternatively called the density reduction factor in the FIA system. We 

used the existing data to estimate both variables for all the species inventoried by 

the FIA. In addition to estimating the mean values, we estimated the uncertainty 

associated with these estimates of density, with the least uncertainty for species 

that had been sampled and the most for genera that had not been sampled. To 

estimate densities for species that had not been sampled, we examined the pattern 

of density reduction for related species and genera that had been sampled. In 

the case of CWD, we compared absolute and relative density among fi ve decay 

classes. For FWD, we were able to use only two decay classes (i.e., undecayed 

versus decayed), although we did this for three size classes. 
7

Data Processing 

When several sources of data for a species, genus, decay class, or size class 

were available, we combined the values to estimate an average and standard 

error. When sample sizes were listed, we used those to weight the average as 

well as the standard error of the samples. When sample sizes were not listed, we 

calculated a simple average and used the highest observed standard error as an 

estimate of uncertainty. For undecayed wood density of CWD, we used estimates 

provided by the FIA database, which is largely derived from the Wood Handbook 

(U.S. Department of Agriculture 1999). Although not all FWD studies reported 

species-specifi c values, we assumed that they represented the values of the 

dominant species in the ecosystem from which they had been sampled. 

CWD Predictions

We used the available information to estimate the CWD density (absolute and 

relative) of each species currently encountered in the national FIA inventory. 

While it is important to estimate the mean density of decayed wood for all 

species, it is even more important to estimate the uncertainty introduced in this 

process. We devised a system in which the uncertainty would increase as the 

degree of extrapolation increased. Minimal extrapolation was involved when 

a species had been sampled and maximum extrapolation was involved when a 

genus had not been sampled. The uncertainty was expressed as the standard error 

of the mean. There were three levels of uncertainty:

A. Species that had been sampled. In this case the mean was the average 

of all the observed values and the uncertainty was represented by the 

standard error of the mean. The uncertainty in relative density was 

calculated as the standard error of the mean absolute density divided by 

the undecayed density. In this case the uncertainty term only involved 

sampling error.

B. Genera that had been sampled. If a species had not been sampled, but 

others in its genus had been sampled then there was some question as to 

where the mean would lie. The mean density was estimated by averaging 

the minimum and maximum relative density for the species in the 

genus that had been sampled. The absolute density was determined by 

multiplying the undecayed density of that species by the mean relative 

density. In this case the uncertainty involved not only sampling error, but 

also uncertainty about the mean itself. We used the maximum difference 

of mean relative density observed in the genus for each decay class to 

provide an estimate of uncertainty associated with the genus mean. This 

was then increased by two standard errors of the mean of the relative 

density for observed species to account for sampling errors. We then 

assumed that 95 percent of the estimates of the mean would fall within 
8

that range and divided it by 4 to rescale it to something akin to a standard 

error. The uncertainty of the absolute density was then calculated by 

multiplying the undecayed density of the species by the uncertainty in the 

relative density. 

C. Species and genus not sampled. In this case the mean could lie 

anywhere between the minimum and maximum of the observed values. 

We therefore estimated the mean as the average of the minimum and 

maximum observed values of relative density. The absolute density was 

the product of the mean relative density and the undecayed density. The 

uncertainty was calculated as when only genera were sampled (section 

B above); however, we used the maximum difference in relative density 

means of all the species that had been examined. 

FWD Predictions

We also estimated green and decayed density of FWD of the species encountered 

in the FIA inventory. For undecayed density, we used actual measures or in most 

cases derived this from undecayed branch density to undecayed bole density 

ratios (branch to bole ratios). For decayed density, we used either means of 

observed values or derived them from decayed versus undecayed FWD density 

ratios. As with CWD, in addition to estimating the mean value, we also estimated 

the uncertainty in FWD estimates based on the level of information available:

A. Species that had been sampled for FWD density. We used the mean 

and standard error of the observed values of undecayed and decayed 

density. We calculated the relative density by dividing the mean by the 

undecayed density. We calculated the uncertainty of the relative density 

by dividing the standard error of decayed density by the mean undecayed 

density. When the standard error had not been reported for a species, we 

multiplied the maximum relative variation of species where this statistic 

had been reported by the mean density of the species in question to 

provide some estimate of uncertainty. 

B. Species lacking undecayed FWD density values. We used the mean bole 

density multiplied by the mean ratio of branch to bole density to estimate 

undecayed FWD density. The uncertainty in undecayed FWD density 

using this method was determined by multiplying the standard error 

of this ratio for all species with observations by the bole density of the 

species that was being estimated. 

C. Species lacking decayed FWD density. Estimates of decayed FWD 

density for species that have not been sampled were computed from the 

product of the mean ratio of decayed to green branch density for species 

and the green density of the species to be estimated. The uncertainty in 

the decayed FWD density for this set of species was estimated by: 
9

UDecayed FWD = sqrt (ID2

*UID

2

 +DGR2*UDGR

2

)

where UDecayed FWD is the uncertainty in decayed FWD density, ID is the 

mean initial density, UID is the uncertainty in initial density, DGR is the 

decay to undecayed ratio, and UDGR is the uncertainty in the undecayed 

ratio. This formula accounts for the fact that uncertainty for decayed 

FWD density is a function of two uncertainties. Our formula assumed no 

correlation between the uncertainties. 

Analysis of Uncertainty on CWD Mass Estimates

We analyzed the uncertainty of CWD mass estimates caused by using current 

knowledge about relative density of decay classes. This was achieved by 

applying various density reduction patterns that are commonly observed to the 

following likely decay class volume distributions:

A. Normal distribution. The distribution of decay classes with respect to 

volume depends on the time interval that the decay class represents as 

well as the nature of the inputs. For the fi ve decay class system we used, 

the tendency is for the time interval represented to increase geometrically 

as decay classes advance. For example, decay class 2 lasts about twice 

as long as decay class 1, and decay class 3 lasts roughly twice as long 

as decay class 2, etc. If the input of dead trees to the forest is uniform 

over time, this tends to result in a peaked volume distribution. Decay 

class 3 has the most volume because it includes a relatively long period 

relative to decay classes 1 and 2 and has not lost a great deal of volume 

to decomposition relative to decay classes 4 and 5. A steady input of 

CWD, therefore, leads to a normal distribution of decay classes in terms 

of volume (Harmon et al. 1986). 

B. Exponential distributions. If there is a pulse in dead tree inputs, one can 

also have a peak in volume that advances from decay class to decay class 

as the forest ages. To assess a situation in which there had been a recent 

pulse of input, we used a negative exponential volume distribution. To 

look at a pulse that occurred in the distant past, we used the complement 

of this distribution (i.e., classes 1 and 5 were switched, etc). 

C. Uniform distribution. Although not common, a uniform distribution was 

used where the volume of each decay class was equal to determine an 

“average” uncertainty. 

D. Observed distribution. This was based on large-scale summaries from 

the FIA databases for Maine and gives a sense of the actual operational 

uncertainty likely to be encountered. The data were taken from 200 plots 

and involved approximately 2,400 CWD pieces. 
10

To calculate the uncertainty in CWD biomass estimates, the relative volume in 

each decay class was multiplied by a range of relative density reduction patterns 

to assess the range of mass estimates that would occur. The relative density 

reduction patterns that were investigated included (1) a steady decrease from 

decay class to decay class, (2) an asymptotic pattern with decay classes 4 and 

5 similar, (3) a mid-plateau in density decline with decay classes 2 and 3 being 

similar, and (4) the pattern for Douglas-fi r (the most commonly used pattern in 

previous studies). We also assessed the uncertainty for a well-sampled species 

(Douglas-fi r) and a well-sampled genus (pines) as well as the minimum and 

maximum relative values observed. The latter two patterns place upper and lower 

uncertainty bounds on species or genera that have not been sampled. For most 

cases, the products of relative volume and relative density for each decay class 

for each relative density reduction pattern were summed and then compared 

to the value for Douglas-fi r, which serves as a useful reference given that this 

pattern of density change has been used frequently. The exception was that for 

the overall minimum and maximum relative densities we used the mean of all 

species as the reference. 

Analysis of Uncertainty on FWD Mass Estimates

We assessed the importance of two facets of uncertainty for estimates of FWD 

mass. The fi rst aspect was the effect of having directly determined the relative 

density of decayed FWD; this was assessed by comparing the uncertainty in 

relative density of species that had actual samples versus those that did not. The 

second aspect involved the fact that the current system estimates an average 

FWD relative density, but does not account for the effect of pulses of input. 

Given that many sound branches and tops are left after disturbances such as 

harvests, pulses of FWD input are common. Immediately after a disturbance, 

FWD density is likely to be close to the undecayed density. As the time since 

the disturbance increases, the overall density of FWD is likely to decline at least 

until new material replaces it. To mimic this situation, we tracked the abundance 

and relative density of two sources of wood: a pulse and that due to regular 

mortality processes. For the pulse, we assumed the abundance of this FWD pool 

would follow a negative exponential decline. We assumed the density of the 

pulse would also decline, but that density would asymptote to refl ect the presence 

of decay resistant portions of branches (i.e., knots). For the FWD generated by 

regular mortality processes, we assumed that pool would gradually accumulate 

and that the density would decline to a lesser degree given that undecayed wood 

is being added regularly. This asymptote was assumed to equal the average value 

we found in our analysis of the FWD dataset. We assumed the rate the pulse 

FWD was lost was the same as the rate the new FWD accumulated. Given that 

the accumulation rate is often close to the disappearance rate, this assumption 

is reasonable (Olson 1963). We explored the effect of not knowing the decay 
11

state of FWD by varying the size of the pulse from 5, 10, and 20 times the size 

of the regular FWD pool. We also varied the asymptotic density of the pulse of 

FWD from a relative density of 0.1 to 0.4 as well as explored the effect of the 

decomposition rate of FWD. We then noted the difference between the minimum 

and the average relative density because this indicated the uncertainty that might 

be introduced by not noting the decay state of FWD. 

Biomass to Carbon Conversion Factors 

We reviewed the literature to fi nd estimates of the carbon concentration of dead 

wood in various states of decomposition. Given the paucity of data, we averaged 

across all species within CWD decay classes to derive a mean estimate. For FWD 

we averaged across all species for undecayed and decayed wood. We used all 

available data to derive a standard error of carbon concentration, weighting these 

estimates by the sample sizes reported in the publications. 
12

RESULTS

QUALITATIVE CHANGES IN CWD DECAY CLASSES

The following describes the general qualitative changes observed as logs 

decompose. While not all species move through this exact progression, many 

do; major exceptions are noted. Decay class 1 logs are the least decomposed, 

with most having leaves still attached and all having intact bark, fi ne twigs, and 

branches. Logs originating from cutting may not have branches and twigs, but 

the cuts appear fresh and have not yet turned gray due to sun bleaching. Decay 

class 2 logs are ones starting to decompose, leaves largely are absent, and many 

of the fi ne twigs have fallen off the larger branches. Bark is typically loose, but 

only starting to fall off the log. An exception would be for the genera Betula and 

Prunus, which tend to retain bark throughout decomposition. For all species, 

there is evidence the surface layers of the wood are decomposing, but the inner, 

central region of the wood is undecayed unless previously infected with heart 

rots. For logs originating from cutting, the ends are gray from sun bleaching. 

Decay class 3 logs have only a few large branches remaining, often in the form 

of stubs, the bark is falling off in large patches, and evidence of sloughing of 

sapwood is also evident. The outer wood is easily crushed by hand, although the 

inner portions can appear completely sound. Despite the large amount of decay, 

decay class 3 logs are able to support their own weight along most of their length. 

For certain genera with decay resistant heartwoods, such as Calocedrus, Quercus, 

and Thuja, decayed sapwood may fall off to the extent that relatively sound 

heartwood may form the outer surface. Decay class 4 logs cannot support their 

own weight and most of their length conforms to the contours of the underlying 

ground. Although circular cross-sections can remain, much of the log forms an 

elliptical cross-section. Branches, if present, are short stubs, which move when 

pulled. This indicates decay has spread to the innermost portions of the log and 

has weakened the wood considerably. Bark, if present, is in small loose patches 

on the log and found in piles alongside or under the log. In the case of the genera 

Betula and Prunus, the bark loosely surrounds the inner, highly decomposed 

wood. Decay class 5 logs are the most decomposed, of elliptical shape (the 

long axis is often many times that of the short axis), and are beginning to be 

incorporated into the forest fl oor. The wood is extremely decayed, usually in the 
13

form of cubical brown rot that can be easily crushed by hand. Bark is not evident 

from the surface (except for the genera Betula and Prunus) and in most cases 

underlies the extremely decomposed wood. 

DENSITY OF DECOMPOSING CWD

A total of 88 species were found to have data on CWD decay class densities 

that had been published and/or collected from North America from the boreal 

to the tropical zones. Most species (53) were from either boreal or temperate 

ecosystems. A total of 49 genera had some data about CWD decay class density, 

most of which were represented by one species. Of the species compiled, 60 were 

hardwood and 28 were softwood species. 

Examining all the species that have been sampled reveals that wood density of 

CWD decay classes declines as expected, but that it is highly variable (Fig. 1). 

Much of this variation is associated with variation in the initial density, which 

ranges from 0.25 to 0.95 g/cm3

. Expressed as a percent, minimum and maximum 

initial densities are 52 percent lower and 95 percent higher than the mean value, 

respectively. The relative variation appears to increase through decay class 

3 and then declines for decay classes 4 and 5. This pattern may be caused by 

decomposition and wood’s lack of structural integrity below a certain density. 

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

012345

Decay Class

Mean Density (g/cubic cm)

Figure 1.—Density of 

all species sampled by 

decay class. Each point 

represents the mean of 

a species that has been 

sampled for each decay 

class.
14

Comparison of hardwood and softwood mean density indicates that softwoods 

have a slightly lower density than hardwoods for decay classes 1 to 3 (Fig. 2). 

However, the mean densities of decay class 4 are quite similar for softwoods and 

hardwoods. Hardwood density, at least for undecayed wood and decay classes 1 

to 3, appears more variable than softwood density. The variation, expressed as a 

range, within these groups is much higher than the variation between these two 

groups. This classifi cation would therefore not seem to be a useful way to stratify 

unsampled species, although the uncertainty for softwood density is considerably 

lower than that for hardwoods. 

CWD DENSITY REDUCTION PATTERNS

When all species that have been sampled are considered, there is a clear decrease 

in relative density as CWD decay class advances (Fig. 3). Although the mean 

relative density exhibits a steady decline, the maximum and minimum relative 

density can be as much as 60 percent higher or 40 percent lower than the mean, 

respectively. The greatest differences between the minimum and maximum 

appear in decay classes 2 and 3, but even for decay class 1 there is considerable 

variation. Despite the high level of variation observed, relative density is less 

variable than absolute density. This indicates that removing the effect of initial 

density can help reduce uncertainty when estimating density of unsampled 

species.

As with absolute density, dividing the species into hardwoods and softwoods 

does not appear to help reduce uncertainty in estimating relative density. 

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

012345

Decay Class

Mean Density (g/cubic cm)

hardwood softwood

hardwood max hardwood min

softwood max softwood min

Figure 2.—Density 

of hardwood versus 

softwood by decay 

class. The mean as well 

as the minimum and 

maximum for each class 

are displayed.
15

Although the mean relative density of hardwoods is slightly lower than that of 

softwoods (Fig. 4), these differences are very small compared to the range in 

values observed. For example, the relative density of decay class 3 hardwoods 

is 0.1 relative density units lower than that for softwoods. However, this is 

approximately one-third the difference between the minimum and the mean 

for hardwoods. The range for softwoods is lower than that for hardwoods, 

but as with absolute density it completely overlaps that of hardwoods. This 

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean relative density

max relative density

min relative density

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

12345

Decay Class

Relative Density

hardwood

softwood

HW max

SW max

HW min

SW min

Figure 3.—Relative 

density of all species by 

decay class. Each point 

represents a species 

that has been sampled 

for each decay class. 

The relative density is 

the observed density 

divided by the initial or 

green density.

Figure 4.—Relative 

density of hardwood 

versus softwood by 

decay class. The mean 

as well as the minimum 

and maximum for each 

class are displayed.
16

indicates that separating species by hardwood versus softwood classes will not 

substantially decrease the uncertainty in prediction of mean relative density. 

However, separation into these two classes might reduce uncertainty below using 

the maximum and minimum of all species because the uncertainty range for 

softwoods is lower than that for hardwoods. 

At least 18 temperate genera have been sampled, about half of which contain 

multiple species. Several patterns of relative density decline are evident in the 

mean for each genus, which suggests that perhaps stratifying by genus would 

reduce uncertainty (Fig. 5). However, plotting the minimum and maximum 

relative density for well-sampled genera indicates there is considerable variation 

(as much as 0.2 relative density units) within each genus (Fig. 6). This variation 

is quite high relative to the typical standard error of the mean for a species, which 

generally ranges from 0.02 to 0.05 relative density units. This indicates that while 

genus may be used to predict unsampled species, the uncertainty would be as 

much as tenfold higher than for species that have been sampled. 

Examination of the mean of each species indicates a number of repeated patterns 

in relative density declines (Fig. 7). The simplest pattern, a steady decline in 

density (hereafter referred to as the S pattern), was observed for 15 species 

(Fig. 7a). In this case, density appeared to decline by approximately an equal 

amount between decay classes 1 to 4 and then to a lesser degree from decay 

class 4 to 5. The most common pattern, a lag followed by steady decline (the 

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

012345

Decay Class

Relative Density

abies acer

alnus betula

calocedrus carya

castanea fagus

fraxinus liquidambar

liriodendron picea

pinus populus

pseudotsuga quercus

thuja tsuga

all

All genera

Pseudotsuga

Figure 5.—Mean 

relative density for 

common temperate 

genera.
17

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

maximum

mean relative density

minimum

n=92

sp=7

n=502

sp=6

n=699

sp=7

n=466

sp=6 n=317

sp=4 n = # of samples

sp = # of species

LS pattern), occurred in 30 species (Fig. 7b). In this pattern, decay class 1 is 

similar in density to fresh wood and then there is a steady decline in density. 

A lag that lasted into decay class 2 was evident in fi ve species, and this was 

typically followed by a steady decline of density in decay classes 3 to 5 (the SLS 

pattern; Fig. 7c). For six species, it appears that a steady decline is followed by 

an asymptote in decay classes 4 and 5 (the A pattern; Fig. 7d). This pattern is the 

one followed by Douglas-fi r, the most common species used to represent density 

reduction patterns. The most complex pattern involved an initial decline in 

density to decay class 1, followed by a mid-plateau, and fi nally a steady decline 

in density (the MP pattern; Fig. 7e). This pattern was followed by seven species. 

Understanding the causes of these patterns might be helpful in predicting 

unsampled species. Unfortunately, we can only hypothesize about possible 

controls of these patterns. It is possible that the lags we observed in the LS 

and SLS patterns are associated with the progression of exterior indicators of 

decay versus the loss of wood via decomposition. In species or environments 

in which the exterior indicators develop quickly relative to the rate of wood 

decomposition, the interior wood does not have “enough time” to lose density. 

Thus, a “lag” in density decline appears. Conversely, if exterior indicators take 

longer to develop, then the wood can decompose to a greater degree and a steady 

density decline appears. This difference might not be caused by the species per 

se, but the environment. In cold and wet environments, bole decomposition 

rates can be very slow due to waterlogging (Harmon et al. 1986). The same 

limitation would not be evident for leaves, twigs, and branches; therefore, their 

Figure 6.—Examples 

of the mean, minimum, 

and maximum relative 

density for genera 

that have been well 

sampled. The number 

of species and total 

number of samples for 

each decay class are 

noted.
18

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean steady max steady min steady

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean lag-steady max lag-steady min lag-steady

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean super-lag-steady max super-lag-steady min super-lag-steady

Figure 7.—Relative 

density reduction 

patterns. The mean, 

minimum, and 

maximum relative 

densities are presented 

for each pattern. 

A: steady decline (S); 

B: lag followed by 

steady decline (LS); 

C: super lag followed by 

steady decline (SLS).

A

B

C
19

decomposition would proceed ahead of the bole’s decomposition. The cause of 

the asymptotic pattern (A) might be related to the presence of a relatively small 

resistant core of wood, although this is unlikely in the case of Douglas-fi r. The 

most complex pattern, the MP, is possibly associated with a proportionally large 

amount of highly resistant wood. This would be typifi ed by western redcedar. 

In this case, sapwood decomposition is quite rapid leading to a decline to 

decay class 1. The remaining heartwood is, however, very resistant to decay 

and although certain indicators of decay class 3 develop (e.g., the sloughing of 

sapwood) the underlying wood remains relatively sound. Eventually this wood 

begins to decompose and weaken leading to the characteristics typical of decay 

class 4. 

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean asymptote max asymptote min asymptote

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0.80

0.90

1.00

12345

Decay Class

Relative Density

mean mid-plateau max mid-plateau min mid-plateau

Figure 7 (continued).—

Relative density 

reduction patterns. 

The mean, minimum, 

and maximum relative 

densities are presented 

for each pattern. 

D: asymptotic (A); 

E: mid-plateau in 

density (MP).

D

E
20

Although these density reduction patterns are useful for descriptive purposes, at 

this time they cannot be used for predictive purposes. First, as discussed above, 

the mechanisms causing the differences are not known. Second, determining 

the degree to which these patterns are statistically different is problematical. In 

some cases, patterns might differ for only one decay class (e.g., S versus the LS 

patterns) and therefore for most decay classes they are statistically the same. 

In others cases, the mean relative density of two patterns are quite different, 

yet the range for both patterns overlap considerably. Third, while it is logical 

to hypothesize density reduction patterns would be similar within a genus, we 

saw little evidence to support that hypothesis in the genera that had been well 

sampled. 

FINE WOODY DEBRIS DENSITY

Relatively few species have had FWD density determined. Our literature search 

indicated approximately 25 species have been sampled, although some of these 

“species” represent mixtures named after a dominant species. Most studies 

appear to have been from northern regions. 

The density of undecayed or green FWD decreased as diameter increased 

(Fig. 8). For example, in the smallest size class (0-8 mm diameter) the mean 

density of undecayed FWD was 0.61 g/cm3. In contrast, for the largest size class 

(25 to 76 mm diameter) the mean undecayed density was 0.50 g/cm3. Regardless 

of diameter, undecayed density was highly variable with values ranging between 

0.1 and 0.95 g/cm3. Given the few species in which undecayed FWD has been 

determined, the ratio of undecayed branch density to bole wood density is a 

useful variable from which to predict the undecayed FWD density of species 

that have not yet been studied. These ratios also decline as diameter size class 

increases, refl ecting the decrease in FWD density as the size class increases 

(Fig. 9). 

Density of decayed FWD also decreases as diameter size class increases 

(Fig. 10) from 0.44 to 0.38 g/cm3

 from the smallest to largest size classes. 

Relative density of FWD as indicated by ratios of decayed to undecayed pieces 

increases as size class increases (Fig. 11). This indicates that large diameter 

pieces are less decomposed than smaller ones. Specifi cally, the largest pieces 

have a mean relative density of 0.88, whereas the smallest ones have a mean 

relative density of 0.61. This might be caused by the increased surface area to 

volume ratio of the smaller pieces, which might allow faster colonization by 

agents of decomposition. 
21

0

5

10

15

20

25

30

0.01-0.20 0.21-0.30 0.31-0.40 0.41-0.50 0.51-0.60 0.61-0.70 0.71-0.80 0.81-0.90 0.91-1.00

Density Classes (g/cubic cm)

Number of samples

mean density and

standard error

N=51

0

5

10

15

20

25

30

0.01-0.20 0.21-0.30 0.31-0.40 0.41-0.50 0.51-0.60 0.61-0.70 0.71-0.80 0.81-0.90 0.91-1.00

Density Classes (g/cubic cm)

Number of samples

mean density and

standard error

N=70

0

5

10

15

20

25

30

0.01-0.20 0.21-0.30 0.31-0.40 0.41-0.50 0.51-0.60 0.61-0.70 0.71-0.80 0.81-0.90 0.91-1.00

Density Classes (g/cubic cm)

Number of samples

mean density and

standard error

N=56

Figure 8.—Distribution 

of density of undecayed 

pieces of FWD for three 

size classes. 

(A) 0-1 cm, (B) 1-5 cm, 

(C) 5-12 cm.

A

B

C
22

Figure 9.—Observed 

ratios of green or 

undecayed bole to 

branch density for three 

size classes (1=0-1 cm, 

2=1-5 cm, 3=5-12 cm).

Figure 10.—Density of 

decayed FWD pieces 

for three size classes 

(1=0-1 cm, 2=1-5 cm, 

3=5-12 cm).

Figure 11.—Relative 

density of decayed 

FWD for three size 

classes (1=0-1 cm, 

2=1-5 cm, 3=5-12 cm).

0.00

0.50

1.00

1.50

2.00

2.50

0123

Size Class

Ratio

0.00

0.10

0.20

0.30

0.40

0.50

0.60

0.70

0123

Size Class

Density (g/cubic cm)

mean

density

0.00

0.20

0.40

0.60

0.80

1.00

0123

Size Class

Relative Density

mean
23

UNCERTAINTIES OF CWD MASS ESTIMATES

Douglas-fi r has been the most commonly used species to estimate decay class 

densities of unsampled species based on the assumption that all species density 

reduction patterns are similar. Using this as a standard to assess the differences 

with other possible density reduction patterns indicates that considerable error 

might have been introduced into CWD biomass estimates (Fig. 12). The size 

of the error depends on both the density reduction pattern and the volume 

distribution. For example, if the steady density decline pattern (S) actually 

occurred, then depending on the distribution of volume for the decay classes, 

the CWD mass might have been 4 to 12 percent higher than calculated using the 

Douglas-fi r density reduction pattern. While many of these volume distributions 

are theoretical, it is important to note that the observed volume distribution 

indicated using Douglas-fi r density reduction patterns would have led to 12 

percent less CWD biomass than was actually there if the S density reduction 

pattern was followed. Likewise for the LS pattern (lag-steady), there might have 

been 20 to 35 percent more CWD biomass present than calculated using Douglasfi r density reduction patterns. Douglas-fi r density reduction patterns seem to 

underestimate for the MP (mid-plateau) pattern as well; there might have been 14 

to 44 percent more CWD biomass if the MP pattern was followed. As with the S 

pattern, the largest errors for the LS and MP patterns occurred for the observed 

distribution. The overall conclusion is that use of Douglas-fi r density reduction 

patterns has probably underestimated CWD biomass in FIA inventories. 

0

5

10

15

20

25

30

35

40

45

50

uniform peaked neg exp old neg exp observed

Maine

Volume Distribution

Possible Error (%)

steady decline pattern

lag- steady decline pattern

mid plateau

Figure 12.—Uncertainty 

in CWD mass 

estimates using 

various relative density 

reduction patterns. 

The differences in 

two common density 

reduction patterns 

relative to that of 

Douglas-fi r in estimated 

biomass assuming 

various volume 

distributions (uniform, 

peaked, negative 

exponential, and 

observed).
24

Comparison of the uncertainty for a well-sampled species (Douglas-fi r) to that 

when a genus has not been sampled indicates that uncertainty of CWD biomass 

estimates could be reduced fourfold to tenfold by actually sampling species 

(Fig. 13). By sampling representative species within a genus, the uncertainty 

could be reduced by up to 50 percent over not having sampled a genus. Sampling 

species within a genus could result in a fi vefold to sevenfold reduction in 

uncertainty if values for well-sampled genera (e.g., pines) are typical. Depending 

on the volume distribution used, uncertainty in CWD biomass for a well-sampled 

species ranged from ±4 to 7 percent. For a well-sampled genus, the uncertainty 

ranged from ±21 to 38 percent. In contrast, if a species followed the maximum 

density reduction pattern, then the estimate of CWD biomass would be 38 to 

56 percent higher than if it followed the mean for all species. Similarly, if a 

species followed the minimum density reduction pattern observed, then the 

CWD biomass estimated would be 39 to 54 percent lower than if it followed the 

mean for all species. This indicates that if the density reduction pattern was not 

known for a genus or species, then there is considerable uncertainty in estimates 

of CWD mass. Although these uncertainties are a function of the volume 

distribution assumed, it is important to note that some of the highest uncertainties 

(±55 percent) were for the observed volume distribution. 

UNCERTAINTIES OF FWD MASS ESTIMATES

Our analysis indicated that when FWD relative density is actually measured, 

the uncertainty is 0.01 to 0.03. Expressed as a percent, this would mean an 

uncertainty of mass of 1 to 3 percent when species are sampled. In contrast, when 

-60

-40

-20

0

20

40

60

80

uniform peaked neg exp old neg

exp

observed

Maine

Volume Distribution

Error Relative to Mean

overall maximum overall minimum species maximum

species minimum genus maximum genus minimum

Figure 13.—Uncertainty 

in CWD mass estimates 

based on the ranges 

expected for a wellstudied species, a 

well-studied genus, 

and all the species 

sampled. The volume 

distributions included 

uniform, peaked, 

negative exponential, 

and observed.
25

relative density has to be estimated, the uncertainty of FWD mass estimation 

ranges from 12 to 19 percent. Sampling FWD density for species, therefore, 

would decrease uncertainty by up to an order of magnitude. 

These conclusions need to be tempered with the realization that density is likely 

to vary considerably when pulses of FWD occur. Regardless of the size of the 

FWD pulse, there was a dip in the FWD relative density (Fig. 14). This was 

caused by the changing proportion between the pulse-related FWD and that 

added by regular mortality. Varying the decomposition rate of the pulse of FWD 

did not infl uence the range of relative densities that occurred. The larger the 

size of the pulse, the greater the dip in density as time since the pulse increased. 

Moreover, as difference between the average and the asymptotic density of the 

pulse increased, the uncertainty in the FWD relative density also increased. This 

sensitivity analysis indicates that even if the mean FWD density is known, there 

are times after disturbances when density could be at least 20 percent higher or 

lower than the average. For a less conservative set of parameter values, there are 

times after a disturbance that relative density could be less than half the average 

relative density. This indicates that the uncertainty introduced by not knowing the 

history of the stand could introduce as much error as not sampling a species. 

BIOMASS TO CARBON CONVERSION FACTORS

There appears to be no consistent trend in carbon concentration as wood 

decomposes (Tables 1 and 2). The mean carbon concentration of CWD across 

all decay classes was 49.8 percent; the lowest mean concentration for a decay 

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

1

0 20 40 60 80 100

Time Since Pulse

Mean Relative Density

10X

5X

20X

Figure 14.—Possible 

changes in FWD 

relative density caused 

by pulses of undecayed 

FWD of various sizes 

relative to background 

levels. In this case 

the pulse of FWD was 

assumed to disappear 

at a rate of 10 percent a 

year to a relative density 

of 0.2. The averaged 

relative density for 

FWD from regular 

mortality processes was 

assumed to be 0.8. The 

size of the pulse relative 

to regular mortality 

varies between lines.

Pulse size
26

class was 48.6 percent and the highest was 51.8 percent. For FWD the mean 

carbon concentration was 50.8 percent, with a range of 49.1 to 51.6 percent. 

Although the range of values within decay and size classes was up to 10 percent 

of the mean, the uncertainty in carbon concentrations used to convert biomass to 

carbon is very low relative to densities. As a fi rst approximation, using a mean 

concentration of 50 percent with an uncertainty of 2 percent would be adequate. 

Table 1.—Biomass to carbon conversion factors for CWD based on 

percent carbon of samples

Decay class Mean Standard error Minimum Maximum

 percent carbon

 1 49.9 0.7 47.8 51.5

 2 48.8 0.7 42.4 53.6

 3 48.6 1.5 42.4 51.6

 4 51.8 0.9 48.2 55.2

 5 50.1 1.3 41.8 55.2

Source of data: Busse (1994); Currie and Nadelhoffer (2002), Lang and Forman (1978), Harmon et al. 

(1987), Lambert et al. (1980). 

Table 2.—Biomass to carbon conversion factors for FWD based on 

percent carbon of samples 

Decay class Size class Mean Standard error Minimum Maximum

 cm percent carbon

Undecayed 1.0-2.5 50.8 2.0 51.5 50.3

Undecayed 2.5-5.0 49.8 1.2 50.0 49.5

Undecayed 5.0-10.0 49.1 0.1 49.1 49.1

Decayed 1.0-2.5 51.3 1.9 52.0 50.6

Decayed 2.5-5.0 51.6 2.9 52.4 51.0

Decayed 5.0-10.0 50.5 2.9 50.9 49.8

Source of data: Currie and Nadelhoffer (2002). 
27

DATABASES OF DENSITY ESTIMATES

We have created three databases, two CWD and one FWD, that can be 

combined with woody detritus volume data to calculate the mass of this material 

(Appendices 2-4). For both FWD and CWD, we present the density in either 

absolute or relative terms. In the case of FWD, we report the estimates for 

absolute and relative density for each of three size classes in one database. For 

CWD, we report the estimates for the fi ve decay classes in separate absolute 

and relative density databases. For each estimate, except undecayed CWD, we 

present a code for the method of estimation, which, as described in the methods, 

determines the uncertainty of the estimate. While the absolute density values are 

probably the easiest to use, the relative density values help one understand how 

the absolute densities were calculated. In addition to an estimate of the most 

likely value (i.e., the mean or median value depending on how the estimate was 

made), we also present an uncertainty value. This is analogous to one standard 

error of the mean; therefore, to approximate the 95 percent confi dence intervals, 

the estimated uncertainty needs to be multiplied by 2 and then added and 

subtracted from the most likely value. 

For both FWD and CWD, most species have not been sampled and therefore the 

uncertainty is generally high. For CWD, of the 260 species considered by the FIA 

inventory, 53 (20 percent) have been sampled for at least some decay classes. 

While sampling more species and genera would improve estimates, it should be 

noted that many major genera have been sampled. For FWD, very few speciessize class combinations have been sampled, with only 25 of the possible 1,046 

combinations with actual data. The low proportion of combinations sampled 

(2 percent) has led to a very high uncertainty in FWD density estimates. 
28

EXAMPLES OF USE

Estimating the most likely value of mass of FWD or CWD is straightforward. 

If using the relative density values, then one needs to multiply the undecayed 

density by the relative density for each decay or size class, and then multiply that 

number by the volume in each decay or size class. Summing the masses for each 

decay or size class gives the total mass. If using the absolute density database, 

one multiplies the decay or size class density by the volume in each class and 

then sums to get the total mass. 

Estimating the uncertainty in mass is complex with only an approximation of 

uncertainty. However, it is better to have some sense of this uncertainty than 

to not report it. Given that uncertainty can only be approximated, one needs to 

adopt several guiding principles. The fi rst is that it is better to overestimate than 

to underestimate uncertainty. Therefore, when there is a choice between using 

calculation methods that can give either higher versus lower estimates, the higher 

one should be selected as long as it represents a reasonable set of assumptions. 

Second, uncertainty estimation in a complex set of calculations can be broken 

down hierarchically: there is no reason to calculate all uncertainty terms at once. 

Third, understanding the correlation between variables is important to estimating 

overall uncertainty because as uncertainty terms for the parts are combined 

the degree of correlation determines whether the uncertainty is increased or 

reduced. For example, if all the uncertainties for each decay class are perfectly 

and positively correlated, then the total uncertainly will be maximized (i.e., the 

masses are either all the minimums or all the maximum masses). If, on the other 

hand, the uncertainties are perfectly and negatively correlated, then the total 

uncertainty will be minimized (i.e., some maximum masses are being offset by 

minimum masses). When there is no correlation between uncertainties in the 

decay classes, then the total uncertainty is intermediate. In the case of woody 

detritus mass estimates, the least likely case would be for all the uncertainties 

to be negatively correlated because there is no specifi c mechanism to cause this 

pattern. 

Given the complexity of the problem there is no single formula to estimate 

uncertainty. One method would be to perform a Monte Carlo analysis drawing 
29

from the distributions for the decay- and size-class databases. As an alternative, 

we combine uncertainties using calculations based on summary statistics. In the 

following sections, we present a series of calculations to provide examples, but 

because each project is likely to have data structured in different ways, it is best 

to use the examples only as guidance and not as a substitute for a project-specifi c 

analysis. To provide a hierarchical framework, we start at the fi nest level of detail 

(e.g., sampled species) and work toward more assumptions (e.g., unsampled 

genera). In this particular set of examples, we use subscripts to indicate the level 

within the hierarchy being considered (DC=decay class; Sp=species; S=site). Our 

example considers CWD, but a similar set of calculations could be performed for 

FWD if size classes were also considered. Specifi c examples for each level in the 

hierarchy are given in Appendices 5 and 6. 

WITHIN A DECAY OR SIZE CLASS FOR A SPECIES AND SITE

To calculate the uncertainty for a decay class for a species at a particular site (or 

plot), the uncertainty can be estimated by multiplying the volume by the absolute 

density uncertainty values:

Uncertainty Mass DC-Sp-S = VolumeDC-Sp-S*Uncertainty DensityDC-Sp-S

WITHIN A SPECIES AND SITE

To calculate the uncertainty for species with estimated densities at a particular 

site (or plot), this term can be approximated by adding uncertainty in mass for all 

the decay classes:

Uncertainty Mass Sp-S = Σ Uncertainty MassDC-Sp-S

This assumes there is a perfect positive correlation of uncertainties for the decay 

classes, which refl ects the fact that an unsampled species is systematically 

underestimated or overestimated in terms of decay class specifi c densities. 

In other words, there is a consistent pattern for a species relative to the mean 

estimates. However, for species that have been sampled, it is likely that there 

are few systematic differences, and a random relationship would be more 

appropriate:

Uncertainty Mass Sp-S = sqrt Σ (Uncertainty MassDC-Sp-S

2

)

where the uncertainty of each decay class is squared and then summed. By taking 

the square root of this term, the total uncertainty for a species within a site can 

be estimated. This is based on the notion that the square of uncertainties are 

additive, but the uncertainties are not (i.e., variances are additive). 
30

WITHIN A SPECIES FOR ALL SITES

To calculate the uncertainty for species that have not had density estimated when 

multiple sites (or plots) are combined, we assume the total uncertainty in mass 

can be estimated by adding uncertainty in mass for all the sites or plots:

Uncertainty Mass Sp = Σ Uncertainty MassSp-S

This assumes a perfect positive correlation in uncertainty, which is the most 

likely case for species that have not been sampled. In this case, all decay classes 

for a species at all places were likely either underestimated or overestimated 

to the same degree. For species that have been sampled for density, it is more 

realistic to assume that the uncertainties are not correlated and the uncertainty 

represents random variation in the population:

Uncertainty Mass Sp = sqrt Σ (Uncertainty MassSp-S

2

)

where the uncertainty of a species mass for each plot is squared and then 

summed. By taking the square root of this term, the total uncertainty for the 

species for all sites can be estimated. 

WITHIN A SITE

The uncertainty in mass for a site would combine the mass of all the species at 

a site. Because there is no reason to believe that uncertainty in one species is 

correlated with another, the overall uncertainty would be approximated by:

Uncertainty MassS

= sqrt Σ (Uncertainty MassSp-S

2

)

where the uncertainty of each species is squared and then summed. By taking the 

square root of this term, the total uncertainty within a site can be estimated. 

ALL SITES

Assuming that the uncertainty in mass for each species for all sites has been 

estimated, the uncertainty for all sites would be computed by assuming there is 

no correlation between sites:

Uncertainty MassT= sqrt Σ (Uncertainty MassSp

2

)

where the uncertainty of each species for all sites is squared and then summed 

over all species. Taking the square root of this term gives the total uncertainty at 

all levels. 
31

FUTURE NEEDS

While the databases we created will be helpful for improving FWD and CWD 

mass estimates, they have several notable problems that limit their usefulness. 

In the case of CWD, it is essential to separate the pieces inventoried into the 

decay classes. To be effective, the decay classes used in inventories need to 

be better defi ned so that they are repeatable. Most decay class systems are 

based on Douglas-fi r decay patterns; however, species can differ in substantial 

ways. While adjustments inevitably are made in the fi eld, these are not always 

documented, and therefore uncertainty about their meaning is introduced. This 

uncertainty could be reduced by consistently describing decay classes so that 

actual differences between species and locations can be documented. 

A more systematic sampling of CWD for species and locations is needed. Past 

sampling of CWD density has been ad hoc or related to studies of decomposition 

rates, the main focus of ecological studies. Ideally, all the major species would be 

studied, the decay classes defi ned in objective and systematic ways, and the wood 

density as well as carbon content determined. Currently many major species 

have not been sampled or decay classes within species have been inadequately 

sampled (< 5 samples). By sampling the most abundant species, uncertainty 

of mass estimates could be reduced to approximately 5 percent, a considerable 

improvement over the current uncertainty, which could be as high as 50 percent. 

FWD densities have rarely been sampled and although the current database is a 

logical starting place, it could be signifi cantly improved. Current publications 

often do not report sample numbers or variation. Mixtures of species are often 

reported, and while this makes sense when working within a uniform forest 

type, it is not effi cient when mixtures of species change from place to place. A 

more fl exible system might be derived if species values were reported because 

mixtures could be created continuously from this data structure. The separation of 

undecayed versus decayed FWD is helpful in estimating density, but separation 

of FWD into decay classes would reduce uncertainty. It would probably not 

be reasonable to classify each piece of FWD into a decay class, but a system 

to characterize decay state quickly would greatly reduce uncertainties in mass 

estimation of this pool. 
32

Despite the lack of data on carbon concentrations in decaying wood, our review 

indicated that this variable does not change greatly with size or degree of decay. 

Compared to the uncertainty related to wood density, carbon concentrations are 

relatively certain. While additional sampling would be desirable, it would not be 

a priority relative to wood density. 

Finally, it should be noted that most of these improvements are static in nature: 

they can be applied to a set of inventory-based volumes, but they do not allow 

one to incorporate decomposition induced density reductions over time. To make 

dynamic adjustments, it would be extremely valuable to know the decomposition 

rates of FWD and CWD. Knowledge of these rates would also allow one to 

directly estimate fl uxes from a single inventory rather than a series of inventories. 

This information would also allow one to better interpret why inventory estimates 

are changing over time. 
33

ACKNOWLEDGMENTS

This research was supported by the Forest Inventory and Analysis program and 

the Pacifi c Northwest Research Station of the U.S. Forest Service, the Bullard 

Fellowship of Harvard University, the Oregon State University Richardson 

Endowment, and the Long-term Ecological Studies of the National Science 

Foundation. 
34

LITERATURE CITED

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36

APPENDICES

APPENDIX 1.—Literature sources of raw density and carbon 

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some central Appalachian hardwood forest species. Res. Pap. NE-716. 

Newtown Square, PA: U.S. Department of Agriculture, Forest Service, 

Northeastern Research Station. 4 p.

Brown, J.K. 1974. Handbook for inventorying downed woody material. Gen. 

Tech. Rep. INT-16. Ogden, UT: U.S. Department of Agriculture, Forest 

Service, Intermountain Forest and Range Experiment Station. 24 p.

Busse, M.D. 1994. Downed bole-wood decomposition in lodgepole pine forests 

of central Oregon. Soil Science Society of America Journal. 58: 221-227. 

Chojnacky, D.C.; Mickler, R.A.; Heath, L.S.; Woodall, C.W. 2004. Estimates 

of down woody materials in eastern U.S. forests. Environmental 

Management. 33: (Supplement 1) S44-S55.

Currie, W.S.; Nadelhoffer, K.J. 2002. The imprint of land-use history: patterns 

of carbon and nitrogen in downed woody debris at the Harvard Forest. 

Ecosystems. 5: 446-460.

Duvall, M.D.; Grigal, D.F. 1999. Effects of timber harvesting on coarse woody 

debris in red pine forests across the Great Lakes states, U.S.A. Canadian 

Journal of Forest Research. 29: 1926-1934.

Erickson, H.E.; Edmonds, R.L.; Peterson, C.E. 1985. Decomposition of logging 

residues in Douglas-fi r western hemlock, Pacifi c silver fi r, and ponderosa 

pine systems. Canadian Journal of Forest Research. 15: 914-921.

Fahey, T.J. 1983. Nutrient dynamics of aboveground detritus in lodgepole 

pine (pinus contorta ssp. latifolia) ecosystems, southeastern Wyoming. 

Ecological Monographs. 53(1): 51-72.
37

Forrester, J.A.; Runkle, J.R. 2000. Mortality and replacement patterns of an oldgrowth Acer-Fagus woods in the Holden Arboretum, northeastern Ohio. 

American Midland Naturalist. 144(2): 227-242.

Fraver, S.; Wagner, R.G.; Day, M. 2002. Dynamics of coarse woody debris 

following gap harvesting in the Acadian forest of Central Maine, U.S.A. 

Canadian Journal of Forest Research. 32: 2094-2105.

Gerald, E.L.; Forman, R.T.T. 1978. Detrital dynamics in a mature oak forest: 

Hutcheson Memorial Forest, New Jersey. Ecology. 59(3): 580-595.

Graham, R.L.; Cromack, K., Jr. 1982. Mass, nutrient content, and decay state of 

dead boles in rain forests of Olympic National Park. Canadian Journal of 

Forest Research. 12: 511-521.

Hale, C.M.; Pastor, J. 1998. Nitrogen content, decay rates, and decompositional 

dynamics of hollow versus solid hardwood logs in hardwood forests of 

Minnesota, U.S.A. Canadian Journal of Forest Research. 28: 1276-1285.

Harmon, M.E.; Sexton, J. 1996. Guidelines for measurements of woody detritus 

in forest ecosystems. Pub. 20. Seattle, WA: University of Washington, 

U.S. LTER Network Offi ce. 73 p.

Harmon, M.E.; Cromack, K., Jr.; Smith, B.G. 1987. Coarse woody debris in 

mixed-conifer forests, Sequoia National Park, California. Canadian Journal 

of Forest Research. 17: 1265-1272.

Harmon, M.E.; Hennessy, T.; Silsbee, D.G. 1980. Woody fuel dimensions within 

Great Smoky Mountains National Park. Res./Resour. Manage. Rep. 31. 

Gatlinburg, TN: U.S. Department of the Interior, National Park Service, 

Uplands Field Research Lab. 28 p.

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and stores of woody detritus in the dry tropical forests of the Northeastern 

Yucatan Peninsula, Mexico. Biotropica. 27: 305-316

Hicks, W.T. 1995. Tree mortality, coarse woody debris and the equilibrium status 

of an old-growth fagus-acer forest in southwestern Ohio. Oxford, OH: 

Miami University. 47 p. M.S. thesis.
38

Lambert, R.L.; Lang, G.E.; Reiners, W.A. 1980. Loss of mass and chemical 

change in decaying boles of a subalpine balsam fi r forest. Ecology. 61(6): 

1460-1473.

Lang, G.E.; Forman, R.T. 1978. Detrital dynamics in a mature oak forest: 

Hutcheson Memorial Forest, New Jersey. Ecology. 59(3): 580-595. 

MacMillan, P.C. 1981. Log decomposition in Donaldson’s woods, Spring Mill 

State Park, Indiana. American Midland Naturalist. 106(2): 335-344.

Nalder, I.A.; Wein, R.W.; Alexander, M.E.; de Groot, W.J. 1997. Physical 

properties of dead and downed round-wood fuels in the boreal forests of 

Alberta and Northwest Territories. Canadian Journal of Forest Research. 

27: 1513-1517.

Nalder, I.A.; Wein, R.W.; Alexander, M.E.; de Groot, W.J. 2000. Physical 

properties of dead and downed round-wood fuels in the boreal forests of 

western and northern Canada. International Journal of Wildland Fire. 

9: 85-99. 

Pittman, J.R. 2005. Coarse woody debris in industrially managed Pinus taeda

plantations of the southeastern United States. Blacksburg, VA: Virginia 

Polytechnic Institute and State University. M.S. thesis. 114 p.

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Lake States tree species. Res. Pap. NC-88. St. Paul, MN: U.S. Department 

of Agriculture, Forest Service, North Central Forest Experiment Station. 

8 p. 

Ryan, K.C.; Pickford, S.G. 1978. Physical properties of woody fuels in the Blue 

Mountains of Oregon and Washington. Res. Note PNW-315. Portland, OR: 

U.S. Department of Agriculture, Forest Service, Pacifi c Northwest Forest 

and Range Experiment Station. 9 p.

Sollins P.; Cline, S.P.; Verhoeven, T.; Sachs, D.; Spycher, G. 1987. Patterns of 

log decay in old-growth Douglas-fi r forest. Canadian Journal of Forest 

Research. 17: 1585-1595.

Wendy, H.L.; Bryant, D.M.; Hutyra, L.R.; Saleska, S.R.; Hammond-Pyle, E.; 

Curran, D.; Wofsy, S.C. 2006. Woody debris contribution to the carbon 

budget of selectively logged and maturing mid-latitude forests. Oecologia. 

148: 108-117.
39

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5Abies amabilis ABAM 0.400 0.360 0.047 A 0.332 0.022 A 0.212 0.090 B 0.142 0.058 B 0.106 0.025 BAbies balsamea ABBA 0.340 0.360 0.061 A 0.360 0.020 A 0.290 0.042 A 0.170 0.100 A 0.100 0.030 AAbies bracteata ABBR 0.360 0.371 0.058 B 0.308 0.087 B 0.212 0.090 B 0.142 0.058 B 0.106 0.025 BAbies concolor ABCO 0.370 0.340 0.019 A 0.277 0.018 A 0.121 0.014 A 0.138 0.016 A 0.122 0.020 AAbies fraseri ABFR 0.340 0.371 0.058 B 0.308 0.087 B 0.212 0.090 B 0.142 0.058 B 0.106 0.025 BAbies grandis ABGR 0.350 0.341 0.003 A 0.294 0.021 A 0.225 0.018 A 0.142 0.058 B 0.106 0.025 BAbies lasiocarpa ABLA 0.310 0.371 0.028 A 0.288 0.078 A 0.233 0.023 A 0.152 0.030 A 0.117 0.011 AAbies magnifi ca ABMA 0.360 0.478 0.013 A 0.378 0.053 A 0.150 0.019 A 0.143 0.014 A 0.084 0.010 A

Abies magnifi ca var. shastensis ABMA2 0.360 0.371 0.058 B 0.308 0.087 B 0.212 0.090 B 0.142 0.058 B 0.106 0.025 B

Abies procera ABPR 0.370 0.368 0.008 A 0.269 0.009 A 0.235 0.013 A 0.150 0.027 A 0.106 0.025 BAbies species ABIE 0.340 0.371 0.058 B 0.308 0.087 B 0.212 0.090 B 0.142 0.058 B 0.106 0.025 BAcacia species ACSP2 0.600 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAcer barbatum ACBA 0.540 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer glabrum ACGL 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer grandidentatum ACGR 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer macrophyllum ACMA 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer negundo ACNE 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer nigrum ACNI 0.520 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer pensylvanicum ACPE 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer rubrum ACRU 0.490 0.436 0.050 A 0.260 0.064 A 0.198 0.075 A 0.177 0.033 B 0.135 0.077 BAcer saccharinum ACSA2 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer saccharum ACSA 0.560 0.679 0.028 A 0.452 0.063 A 0.324 0.043 A 0.182 0.014 A 0.150 0.015 AAcer species ACER 0.490 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAcer spicatum ACSP 0.440 0.536 0.081 B 0.377 0.078 B 0.281 0.075 B 0.177 0.033 B 0.135 0.077 BAesculus californica AECA 0.380 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAesculus species AESP 0.330 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAilanthus altissima AIAL 0.370 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAleurites fordii ALFO 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAlnus rubra ALRU 0.370 0.386 0.008 A 0.326 0.047 A 0.197 0.011 A 0.108 0.008 A 0.117 0.049 AAlnus species ALSP 0.370 0.386 0.233 B 0.326 0.263 B 0.197 0.261 B 0.108 0.175 B 0.117 0.077 BAmelanchier species AMSP 0.660 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CArbutus menziesii ARME 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CAsimina triloba ASTR 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CBetula alleghaniensis BEAL 0.550 0.580 0.076 B 0.403 0.263 B 0.190 0.030 A 0.170 0.020 A 0.110 0.010 ABetula lenta BELE 0.600 0.635 0.069 A 0.420 0.099 A 0.283 0.085 A 0.170 0.175 B 0.110 0.077 BBetula lutea BELU 0.600 0.636 0.043 A 0.385 0.058 A 0.234 0.045 A 0.170 0.175 B 0.110 0.077 BBetula nigra BENI 0.560 0.580 0.076 B 0.403 0.263 B 0.265 0.105 B 0.170 0.175 B 0.110 0.077 BBetula occidentalis BEOC 0.530 0.580 0.076 B 0.403 0.263 B 0.265 0.105 B 0.170 0.175 B 0.110 0.077 BBetula papyrifera BEPA 0.480 0.469 0.052 A 0.403 0.263 B 0.352 0.128 A 0.170 0.175 B 0.110 0.077 BBetula papyrifera var. commutata BEPA2 0.480 0.580 0.076 B 0.403 0.263 B 0.265 0.105 B 0.170 0.175 B 0.110 0.077 B

Betula populifolia BEPO 0.450 0.580 0.076 B 0.403 0.263 B 0.265 0.105 B 0.170 0.175 B 0.110 0.077 BBetula species BESP 0.480 0.580 0.076 B 0.403 0.263 B 0.265 0.105 B 0.170 0.175 B 0.110 0.077 B

APPENDIX 2.—CWD absolute density and its uncertainty for each decay class by species present in the U.S. FIA system

(Appendix 2 continued on next page)
40

Bumelia lanuginosa BULA 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCalocedrus decurrens CADE3 0.370 0.425 0.105 A 0.269 0.055 A 0.231 0.043 A 0.156 0.091 A 0.143 0.106 CCarpinus caroliniana CACA 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCarya aquatica CAAQ 0.610 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya cordiformis CACO 0.600 0.610 0.074 A 0.367 0.074 A 0.249 0.050 A 0.173 0.175 B 0.151 0.077 BCarya glabra CAGL 0.660 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya illinoensis CAIL 0.600 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya laciniosa CALA 0.620 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya myristicaeformis CAMY 0.560 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya ovata CAOV 0.640 0.551 0.071 A 0.479 0.087 A 0.308 0.065 A 0.173 0.175 B 0.151 0.077 BCarya species CASP 0.620 0.633 0.031 A 0.417 0.021 A 0.195 0.017 A 0.173 0.027 A 0.151 0.059 ACarya texana CATE 0.540 0.599 0.058 B 0.409 0.072 B 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCarya tomentosa CATO 0.640 0.601 0.059 A 0.372 0.033 A 0.238 0.061 B 0.173 0.175 B 0.151 0.077 BCastanea dentata CADE 0.400 0.360 0.046 A 0.348 0.038 A 0.255 0.056 A 0.212 0.175 C 0.158 0.077 CCastanea ozarkensis CAOZ 0.400 0.360 0.233 B 0.348 0.263 B 0.255 0.261 B 0.212 0.175 C 0.158 0.077 CCastanea pumila CAPU 0.400 0.360 0.233 B 0.348 0.263 B 0.255 0.261 B 0.212 0.175 C 0.158 0.077 CCastanopsis species CAST 0.420 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCatalpa species CATA 0.380 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCeltis laevigata CELA 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCeltis occidentalis CEOC 0.490 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCeltis species CESP 0.490 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCercis canadensis CECA 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCercocarpus intricatus CEIN 1.000 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCercocarpus ledifolius CELE 1.000 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCercocarpus montanus CEMO 1.000 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CChamaecyparis lawsoniana CHLA 0.390 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CChamaecyparis nootkatensis CHNO 0.420 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CChamaecyparis thyoides CHTH 0.310 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CCladrastis lutea CLLU 0.520 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCornus fl orida COFL 0.640 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 C

Cornus nuttallii CONU 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCornus species COSP 0.640 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCotinus obovaus COOB 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCrataegus species CRSP 0.620 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CCupressus species CUSP 0.440 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CDiospyros virginiana DIVI 0.640 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CEucalyptus species EUSP 0.670 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CFagus grandifolia FAGR 0.560 0.570 0.056 A 0.300 0.042 A 0.167 0.032 A 0.240 0.028 A 0.160 0.012 AFraxinus americana FRAM 0.550 0.475 0.070 A 0.286 0.070 A 0.317 0.048 A 0.212 0.175 C 0.100 0.030 AFraxinus latifolia FRLA 0.500 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 BFraxinus nigra FRNI 0.450 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 BFraxinus pennsylvanica FRPE 0.530 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 BFraxinus profunda FRPR 0.540 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 BFraxinus quadrangulata FRQU 0.530 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 B

(Appendix 2 continued)

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5
41

Fraxinus species FRSP 0.540 0.475 0.233 B 0.317 0.263 B 0.298 0.261 B 0.212 0.175 C 0.100 0.077 BGleditsia aquatica GLAQ 0.600 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CGleditsia triacanthos GLTR 0.600 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CGordonia lasianthus GOLA 0.370 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CGymnocladus dioicus GYDI 0.500 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CHalesia species HASP 0.320 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CIlex opaca ILOP 0.500 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CJuglans cinerea JUCI 0.360 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CJuglans species JUSP 0.510 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CJuglans nigra JUNI 0.510 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CJuniperus monosperma JUMO 0.450 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CJuniperus occidentalis JUOC 0.440 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CJuniperus species JUSP2 0.440 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CLarix laricina LALA 0.490 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CLarix lyallii LALY 0.480 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CLarix occidentalis LAOC 0.480 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CLarix species LASP 0.440 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CLiquidambar styracifl ua LIST 0.460 0.450 0.008 A 0.331 0.031 A 0.228 0.025 A 0.212 0.175 C 0.158 0.077 CLiriodendron tulipifera LITU 0.400 0.352 0.042 A 0.275 0.041 A 0.203 0.026 A 0.141 0.055 A 0.158 0.077 CLithocarpus densifl orus LIDE 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 C

Maclura pomifera MAPO 0.760 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CMagnolia acuminata MAAC 0.450 0.425 0.028 A 0.399 0.084 A 0.250 0.059 A 0.212 0.175 C 0.158 0.077 CMagnolia species MASP 0.450 0.425 0.233 B 0.399 0.263 B 0.250 0.261 B 0.212 0.175 C 0.158 0.077 CMalus species MASP2 0.610 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CMelia azedarach MEAZ 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CMorus alba MOAL 0.590 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CMorus rubra MORU 0.590 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CMorus species MOSP 0.590 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CNyssa aquatica NYAQ 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CNyssa ogeche NYOG 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CNyssa sylvatica NYSY 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 COlneya tesota OLTE 1.000 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 COstrya virginiana OSVI 0.630 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 COxydendrum arboreum OXAR 0.500 0.524 0.103 A 0.406 0.034 A 0.337 0.057 A 0.212 0.175 C 0.158 0.077 CPaulownia tomentosa PATO 0.380 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CPersea borbonia PEBO 0.510 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CPicea abies PIAB 0.380 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPicea breweriana PIBR 0.330 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPicea engelmannii PIEN 0.330 0.393 0.057 B 0.258 0.023 A 0.280 0.112 B 0.117 0.026 A 0.129 0.106 BPicea glauca PIGL2 0.370 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPicea lutzii PILU 0.370 0.376 0.034 A 0.347 0.035 A 0.255 0.067 A 0.151 0.027 A 0.103 0.018 APicea mariana PIMA 0.380 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPicea pungens PIPU 0.380 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPicea rubens PIRU 0.380 0.406 0.057 A 0.312 0.040 B 0.316 0.112 A 0.155 0.035 B 0.129 0.106 B

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5

(Appendix 2 continued on next page)
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Picea sitchensis PISI 0.370 0.410 0.008 A 0.341 0.021 A 0.263 0.021 A 0.192 0.029 A 0.137 0.008 APicea species PICE 0.380 0.393 0.057 B 0.312 0.040 B 0.280 0.112 B 0.155 0.035 B 0.129 0.106 BPinus albicaulis PIAL 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus aristata PIAR 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus arizonica PIAR2 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus attenuata PIAT 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus balfouriana PIBA 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus banksiana PIBA2 0.400 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus clausa PICL 0.460 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus contorta PICO 0.380 0.378 0.020 A 0.367 0.013 A 0.276 0.022 A 0.169 0.019 A 0.164 0.009 APinus coulteri PICO2 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus discolor PIDI 0.500 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus echinata PIEC 0.470 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus edulis PIED 0.500 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus elliotti PIEL 0.540 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus engelmannii PIEN2 0.370 0.368 0.031 B 0.324 0.038 B 0.280 0.071 B 0.169 0.055 B 0.129 0.018 BPinus fl exilis PIFL 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus glabra PIGL 0.410 0.393 0.046 B 0.312 0.040 B 0.280 0.071 B 0.155 0.035 B 0.129 0.018 BPinus jeffreyi PIJE 0.370 0.365 0.026 A 0.358 0.042 A 0.217 0.047 A 0.205 0.039 A 0.171 0.096 BPinus lambertiana PILA 0.340 0.369 0.037 A 0.267 0.011 A 0.155 0.020 A 0.122 0.025 A 0.171 0.096 BPinus leiophylla PILE 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus monophylla PIMO 0.500 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus monticola PIMO3 0.350 0.344 0.005 A 0.303 0.012 A 0.308 0.015 A 0.169 0.055 B 0.171 0.096 BPinus muricata PIMU 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus nigra PINI 0.410 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus palustris PIPA 0.540 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus ponderosa PIPO 0.380 0.338 0.005 A 0.333 0.018 A 0.330 0.035 A 0.129 0.008 A 0.188 0.134 APinus pungens PIPU2 0.490 0.393 0.046 B 0.312 0.040 B 0.280 0.071 B 0.155 0.035 B 0.129 0.018 BPinus radiata PIRA 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus resinosa PIRE 0.410 0.340 0.046 A 0.324 0.038 B 0.270 0.070 A 0.150 0.055 A 0.110 0.134 APinus rigida PIRI 0.470 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus sabiniana PISA 0.370 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus serotina PISE 0.510 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus strobiformis PIST2 0.350 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus strobus PIST 0.340 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus sylvestris PISY 0.410 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPinus taeda PITA 0.470 0.386 0.012 A 0.306 0.005 A 0.229 0.002 A 0.193 0.004 A 0.224 0.134 APinus virginiana PIVI 0.450 0.368 0.031 B 0.324 0.038 B 0.273 0.070 B 0.169 0.055 B 0.171 0.096 BPlanera aquatica PLAQ 0.530 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CPlatanus occidentalis PLOC 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CPopulus alba POAL 0.370 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus angustifolia POAN 0.340 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus balsamifera POBA 0.310 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus deltoides PODE 0.370 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 B

(Appendix 2 continued)

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5
43

Populus fremontii POFR 0.340 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus grandidentata POGR 0.360 0.336 0.067 A 0.422 0.263 C 0.298 0.066 A 0.160 0.175 B 0.110 0.077 BPopulus heterophylla POHE 0.370 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus sargentii POSA 0.370 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BPopulus species POSP 0.370 0.370 0.067 A 0.422 0.263 C 0.300 0.066 A 0.160 0.027 A 0.110 0.027 APopulus tremuloides POTR 0.350 0.353 0.233 B 0.422 0.263 C 0.299 0.261 B 0.160 0.175 B 0.110 0.077 BProsopis species PRSP 0.580 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CPrunus americana PRAM 0.470 0.489 0.080 B 0.418 0.065 B 0.281 0.261 B 0.212 0.175 C 0.158 0.077 CPrunus nigra PRNI 0.470 0.489 0.080 B 0.418 0.065 B 0.281 0.261 B 0.212 0.175 C 0.158 0.077 CPrunus pensylvanica PRPE 0.360 0.401 0.029 A 0.337 0.026 A 0.216 0.026 A 0.212 0.175 C 0.158 0.077 CPrunus serotina PRSE 0.470 0.577 0.072 A 0.499 0.049 A 0.346 0.097 A 0.212 0.175 C 0.158 0.077 CPrunus species PRSP2 0.470 0.489 0.080 B 0.418 0.065 B 0.281 0.261 B 0.212 0.175 C 0.158 0.077 CPrunus virginiana PRVI 0.360 0.489 0.080 B 0.418 0.065 B 0.281 0.261 B 0.212 0.175 C 0.158 0.077 CPseudotsuga menziesii PSME 0.450 0.386 0.013 A 0.308 0.017 A 0.152 0.022 A 0.123 0.011 A 0.148 0.110 AQuercus agrifolia QUAG 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus alba QUAL 0.600 0.567 0.031 A 0.387 0.019 A 0.335 0.045 A 0.168 0.071 A 0.248 0.011 BQuercus arizonica, grisea QUAR 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus bicolor QUBI 0.640 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus chrysolepis QUCH 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus coccinea QUCO 0.600 0.571 0.053 A 0.502 0.041 A 0.442 0.033 A 0.241 0.052 B 0.248 0.011 BQuercus douglasii QUDO 0.510 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus durandii QUDU 0.600 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus ellipsoidalis QUEL 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus emoryi QUEM 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus engelmannii QUEN 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus falcata var. falcata QUFA 0.520 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 B

Quercus falcata var. pagodifolia QUFA2 0.610 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 B

Quercus gambelii QUGA 0.640 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus garryana QUGA2 0.640 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus hypoleucoides QUHY 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus ilicifolia QUIL 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus imbricaria QUIM 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus incana QUIN 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus kelloggii QUKE 0.510 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus laevis QULA 0.520 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus laurifolia QULA2 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus lobata QULO 0.640 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus lyrata QULY 0.570 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus macrocarpa QUMA 0.580 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus marilandica QUMA2 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus michauxii QUMI 0.600 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus muehlenbergii QUMU 0.600 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus nigra QUNI 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus nuttalli QUNU 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 B

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5

(Appendix 2 continued on next page)
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Quercus oblongifolia QUOB 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus palustris QUPA 0.580 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus phellos QUPH 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus prinus QUPR 0.570 0.729 0.078 A 0.489 0.093 A 0.294 0.042 A 0.241 0.052 B 0.248 0.011 BQuercus rubra QURU 0.560 0.545 0.043 A 0.303 0.083 A 0.387 0.081 A 0.234 0.052 A 0.248 0.011 BQuercus shumardii QUSH 0.560 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus species QUSP 0.580 0.565 0.019 A 0.462 0.039 A 0.318 0.043 A 0.196 0.020 A 0.248 0.021 AQuercus stellata QUST 0.600 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus velutina QUVE 0.560 0.611 0.085 B 0.363 0.096 A 0.453 0.097 A 0.241 0.052 B 0.248 0.011 BQuercus virginiana QUVI 0.800 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BQuercus wislizeni QUWI 0.700 0.611 0.085 B 0.450 0.096 B 0.382 0.097 B 0.241 0.052 B 0.248 0.011 BRobinia neomexicana RONE 0.660 0.725 0.233 B 0.560 0.263 B 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CRobinia pseudoacacia ROPS 0.660 0.725 0.031 A 0.560 0.053 A 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CSalix species SASP 0.360 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CSapium sebiferum SASE 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CSassafras albidum SAAL 0.420 0.432 0.028 A 0.388 0.042 A 0.338 0.025 A 0.212 0.175 C 0.158 0.077 CSequoia sempervirens SESE 0.340 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CSequoiadendron giganteum SEGI 0.340 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CSorbus americana SOAM 0.420 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CTamarix species TASP 0.400 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CTaxodium distichum var. nutans TADI 0.420 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 C

Taxus brevifolia TABR 0.600 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CThuja occidentalis THOC 0.290 0.329 0.038 A 0.259 0.390 B 0.315 0.053 A 0.143 0.091 B 0.143 0.110 BThuja plicata THPL 0.310 0.318 0.038 A 0.259 0.390 A 0.248 0.030 A 0.132 0.002 A 0.143 0.110 ATilia americana TIAM 0.320 0.406 0.024 A 0.333 0.032 A 0.256 0.019 A 0.212 0.175 C 0.158 0.077 CTilia heterophylla TIHE 0.320 0.406 0.233 B 0.333 0.263 B 0.256 0.261 B 0.212 0.175 C 0.158 0.077 CTilia species TISP 0.320 0.406 0.233 B 0.333 0.263 B 0.256 0.261 B 0.212 0.175 C 0.158 0.077 CTorreya californica TOCA 0.340 0.381 0.105 C 0.318 0.245 C 0.257 0.122 C 0.162 0.091 C 0.143 0.106 CTsuga canadensis TSCA 0.400 0.398 0.059 A 0.322 0.245 B 0.325 0.084 A 0.178 0.091 B 0.140 0.106 BTsuga heterophylla TSHE 0.420 0.399 0.022 A 0.346 0.023 A 0.258 0.019 A 0.166 0.025 A 0.108 0.106 ATsuga mertensiana TSME 0.420 0.380 0.059 B 0.322 0.245 B 0.282 0.122 B 0.178 0.091 B 0.140 0.106 BTsuga species TSSP 0.380 0.380 0.059 B 0.322 0.245 B 0.282 0.122 B 0.178 0.091 B 0.140 0.106 BUlmus alata ULAL 0.570 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus americana ULAM 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus crassifolia ULCR 0.570 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus pumila ULPU 0.460 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus rubra ULRU 0.480 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus serotina ULSE 0.570 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus species ULSP 0.500 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUlmus thomasii ULTH 0.570 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CUmbellularia californica UMCA 0.510 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 CVaccinium arboreum VAAR 0.470 0.533 0.233 C 0.422 0.263 C 0.325 0.261 C 0.212 0.175 C 0.158 0.077 C

(Appendix 2 continued)

Genus Species code den0 den1 unc1 cod1 den2 unc2 cod2 den3 unc3 cod3 den4 unc4 cod4 den5 unc5 cod5
45

CWD Density Predictions Metadata (Appendix 2) Fieldname Defi nition Codes/units Method

genus Sample genusspecies Sample speciescode Genus-species code den0 Green density (decay class “0”) g/cm3

den1 Density of decay class 1 g/cm3unc1 Uncertainty of den1 g/cm3cod1 Uncertainty code for den1 (A, B, C) A Species sampled B Genera sampled C Species and genus not sampledden2 Density of decay class 2 g/cm3unc2 Uncertainty of den2 g/cm3cod2 Uncertainty code for den2 (A, B, C) A Species sampled B Genera sampled C Species and genus not sampledden3 Density of decay class 3 g/cm3unc3 Uncertainty of den3 g/cm3cod3 Uncertainty code for den3 (A, B, C) A Species sampled B Genera sampled C Species and genus not sampledden4 Density of decay class 4 g/cm3unc4 Uncertainty of den4 g/cm3cod4 Uncertainty code for den4 (A, B, C) A Species sampled B Genera sampled C Species and genus not sampledden5 Density of decay class 5 g/cm3unc5 Uncertainty of den5 g/cm3cod5 Uncertainty code for den5 (A, B, C) A Species sampled B Genera sampled C Species and genus not sampled
46

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5Abies amabilis ABAM 0.400 0.900 0.047 A 0.830 0.022 A 0.604 0.154 B 0.404 0.093 B 0.309 0.051 BAbies balsamea ABBA 0.340 1.000 0.061 A 1.000 0.020 A 0.828 0.042 A 0.500 0.100 A 0.294 0.030 AAbies bracteata ABBR 0.360 0.979 0.056 B 0.848 0.107 B 0.604 0.154 B 0.404 0.093 B 0.309 0.051 BAbies concolor ABCO 0.370 0.958 0.019 A 0.759 0.018 A 0.470 0.014 A 0.381 0.016 A 0.330 0.020 AAbies fraseri ABFR 0.340 0.979 0.056 B 0.848 0.107 B 0.604 0.154 B 0.404 0.093 B 0.309 0.051 BAbies grandis ABGR 0.350 0.974 0.003 A 0.840 0.021 A 0.643 0.018 A 0.404 0.093 B 0.309 0.051 BAbies lasiocarpa ABLA 0.310 1.000 0.028 A 0.929 0.078 A 0.752 0.023 A 0.490 0.030 A 0.377 0.011 AAbies magnifi ca ABMA 0.360 1.000 0.013 A 0.939 0.053 A 0.471 0.019 A 0.329 0.014 A 0.233 0.010 A

Abies magnifi ca var. shastensis ABMA2 0.360 0.979 0.056 B 0.848 0.107 B 0.604 0.154 B 0.404 0.093 B 0.309 0.051 B

Abies procera ABPR 0.370 0.995 0.008 A 0.727 0.009 A 0.635 0.013 A 0.405 0.027 A 0.309 0.051 BAbies species ABIE 0.340 0.979 0.056 B 0.848 0.107 B 0.604 0.154 B 0.438 0.093 A 0.309 0.051 BAcacia species ACSP 0.600 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAcer barbatum ACBA 0.540 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer glabrum ACGL 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer grandidentatum ACGR 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer macrophyllum ACMA 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer negundo ACNE 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer nigrum ACNI 0.520 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer pensylvanicum ACPE 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer rubrum ACRU 0.490 0.963 0.050 A 0.716 0.064 A 0.583 0.075 A 0.317 0.180 B 0.241 0.094 BAcer saccharinum ACSA2 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer saccharum ACSA 0.560 0.919 0.028 A 0.705 0.063 A 0.496 0.043 A 0.317 0.014 A 0.241 0.015 AAcer species ACSP2 0.600 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAcer spicatum ACSP 0.440 0.941 0.036 B 0.709 0.270 B 0.533 0.265 B 0.317 0.180 B 0.241 0.094 BAesculus californica AECA 0.380 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAesculus species AESP 0.330 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAilanthus altissima AIAL 0.370 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAleurites fordii ALFO 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAlnus rubra ALRU 0.370 0.990 0.008 A 0.836 0.047 A 0.505 0.011 A 0.277 0.008 A 0.300 0.049 AAlnus species ALSP 0.370 0.990 0.136 B 0.836 0.270 B 0.505 0.265 B 0.277 0.180 B 0.300 0.094 BAmelanchier species AMSP 0.660 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CArbutus menziesii ARME 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CAsimina triloba ASTR 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CBetula alleghaniensis BEAL 0.550 0.992 0.136 B 0.660 0.270 B 0.317 0.030 A 0.283 0.020 A 0.183 0.010 ABetula lenta BELE 0.600 1.000 0.069 A 0.700 0.099 A 0.472 0.085 A 0.309 0.180 B 0.200 0.094 BBetula lutea BELU 0.600 1.000 0.043 A 0.621 0.058 A 0.377 0.045 A 0.309 0.180 B 0.200 0.094 BBetula nigra BENI 0.560 0.992 0.136 B 0.660 0.270 B 0.482 0.265 B 0.309 0.180 B 0.200 0.094 BBetula occidentalis BEOC 0.530 0.992 0.136 B 0.660 0.270 B 0.482 0.265 B 0.309 0.180 B 0.200 0.094 BBetula papyrifera BEPA 0.480 0.977 0.052 A 0.660 0.270 B 0.733 0.128 A 0.309 0.180 B 0.200 0.094 BBetula papyrifera var. commutata BEPA2 0.480 0.992 0.136 B 0.660 0.270 B 0.482 0.265 B 0.309 0.180 B 0.200 0.094 B

Betula populifolia BEPO 0.450 0.992 0.136 B 0.660 0.270 B 0.482 0.265 B 0.309 0.180 B 0.200 0.094 BBetula species BESP 0.480 0.992 0.136 B 0.660 0.270 B 0.482 0.265 B 0.309 0.180 B 0.200 0.094 B

APPENDIX 3.—CWD relative density and its uncertainty for each decay class by species present in the U.S. FIA system
47

Brosimum alcastrum BRAL 0.740 0.836 0.010 A 0.491 0.052 A 0.335 0.053 A 0.193 0.018 A 0.270 0.094 ABumelia lanuginosa BULA 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CBursera simaruba BUSI 0.330 0.973 0.026 A 0.643 0.029 A 0.387 0.017 A 0.385 0.060 A 0.352 0.094 ACalocedrus decurrens CADE3 0.370 1.000 0.073 A 0.845 0.039 A 0.760 0.035 A 0.420 0.105 C 0.366 0.140 CCarpinus caroliniana CACA 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCarya aquatica CAAQ 0.610 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya cordiformis CACO 0.600 1.000 0.074 A 0.612 0.074 A 0.415 0.050 A 0.279 0.180 B 0.244 0.094 BCarya glabra CAGL 0.660 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya illinoensis CAIL 0.600 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya laciniosa CALA 0.620 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya myristicaeformis CAMY 0.560 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya ovata CAOV 0.640 0.861 0.071 A 0.748 0.087 A 0.481 0.065 A 0.279 0.180 B 0.244 0.094 BCarya species CASP 0.620 1.000 0.031 A 0.673 0.021 A 0.315 0.017 A 0.279 0.027 A 0.244 0.059 ACarya texana CATE 0.540 0.950 0.072 B 0.653 0.085 B 0.381 0.075 B 0.279 0.180 B 0.244 0.094 BCarya tomentosa CATO 0.640 0.939 0.059 A 0.581 0.033 A 0.313 0.075 A 0.279 0.180 B 0.244 0.094 BCastanea dentata CADE 0.400 0.900 0.046 A 0.870 0.038 A 0.630 0.056 A 0.370 0.180 C 0.366 0.094 CCastanea ozarkensis CAOZ 0.400 0.900 0.136 B 0.870 0.270 B 0.638 0.265 B 0.370 0.180 C 0.282 0.094 CCastanea pumila CAPU 0.400 0.900 0.136 B 0.870 0.270 B 0.638 0.265 B 0.370 0.180 C 0.282 0.094 CCastanopsis species CAST 0.420 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCatalpa species CATA 0.380 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCecropia obtusifolia CEOB 0.300 0.900 0.048 A 0.687 0.049 A 0.460 0.023 A 0.320 0.034 A 0.282 0.094 CCeltis laevigata CELA 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCeltis occidentalis CEOC 0.490 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCeltis species CESP 0.490 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCercis canadensis CECA 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCercocarpus intricatus CEIN 1.000 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCercocarpus ledifolius CELE 1.000 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCercocarpus montanus CEMO 1.000 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CChamaecyparis lawsoniana CHLA 0.390 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CChamaecyparis nootkatensis CHNO 0.420 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CChamaecyparis thyoides CHTH 0.310 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CCladrastis lutea CLLU 0.520 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCornus fl orida COFL 0.640 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 C

Cornus nuttallii CONU 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCornus species COSP 0.640 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCotinus obovaus COOB 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCrataegus species CRSP 0.620 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CCupressus species CUSP 0.440 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CDiospyros virginiana DIVI 0.640 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CEucalyptus species EUSP 0.670 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CFagus grandifolia FAGR 0.560 0.976 0.056 A 0.614 0.042 A 0.479 0.032 A 0.375 0.028 A 0.250 0.012 AFraxinus americana FRAM 0.550 0.864 0.070 A 0.520 0.070 A 0.570 0.048 A 0.370 0.180 C 0.182 0.030 AFraxinus latifolia FRLA 0.500 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 BFraxinus nigra FRNI 0.450 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 B

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5

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Fraxinus pennsylvanica FRPE 0.530 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 BFraxinus profunda FRPR 0.540 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 BFraxinus quadrangulata FRQU 0.530 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 BFraxinus species FRSP 0.540 0.864 0.136 B 0.576 0.270 B 0.542 0.265 B 0.370 0.180 C 0.182 0.094 BGleditsia aquatica GLAQ 0.600 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CGleditsia triacanthos GLTR 0.600 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CGordonia lasianthus GOLA 0.370 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CGymnocladus dioicus GYDI 0.500 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CHalesia species HASP 0.320 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CHeliocarpos appendiculatus HEAP 0.250 0.716 0.041 A 0.604 0.035 A 0.536 0.008 A 0.452 0.020 A 0.282 0.094 CIlex opaca ILOP 0.500 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CJuglans cinerea JUCI 0.360 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CJuglans species JUSP 0.510 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CJulglans nigra JUNI 0.510 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CJuniperus monosperma JUMO 0.450 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CJuniperus occidentalis JUOC 0.440 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CJuniperus species JUSP2 0.440 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CLarix laricina LALA 0.490 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CLarix lyallii LALY 0.480 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CLarix occidentalis LAOC 0.480 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CLarix species LASP 0.440 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CLiquidambar styracifl ua LIST 0.460 0.978 0.008 A 0.720 0.031 A 0.496 0.025 A 0.370 0.180 C 0.282 0.094 CLiriodendron tulipifera LITU 0.400 0.940 0.042 A 0.786 0.041 A 0.523 0.026 A 0.353 0.055 A 0.282 0.094 CLithocarpus densifl orus LIDE 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 C

Maclura pomifera MAPO 0.760 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CMagnolia acuminata MAAC 0.450 0.850 0.028 A 0.798 0.084 A 0.500 0.059 A 0.370 0.180 C 0.282 0.094 CMagnolia species MASP 0.450 0.850 0.136 B 0.798 0.270 B 0.500 0.265 B 0.370 0.180 C 0.282 0.094 CMalus species MASP2 0.610 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CManilkara zapota MAZA 0.950 0.790 0.040 A 0.787 0.040 A 0.740 0.138 A 0.566 0.050 A 0.282 0.094 CMelia azedarach MEAZ 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CMorus alba MOAL 0.590 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CMorus rubra MORU 0.590 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CMorus species MOSP 0.590 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CMyrcianthes fragans MYFR 0.750 0.968 0.136 A 0.872 0.047 A 0.745 0.095 A 0.400 0.180 A 0.282 0.094 CNyssa aquatica NYAQ 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CNyssa ogeche NYOG 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CNyssa sylvatica NYSY 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 COlneya tesota OLTE 1.000 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 COstrya virginiana OSVI 0.630 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 COxydendrum arboreum OXAR 0.500 1.000 0.103 A 0.812 0.034 A 0.674 0.057 A 0.370 0.180 C 0.282 0.094 CPaulownia tomentosa PATO 0.380 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CPersea borbonia PEBO 0.510 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CPicea abies PIAB 0.380 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 BPicea breweriana PIBR 0.330 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 B

(Appendix 3 continued)

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5
49

Picea engelmannii PIEN 0.330 0.916 0.073 B 0.782 0.023 A 0.745 0.201 B 0.355 0.026 A 0.347 0.140 BPicea glauca PIGL 0.410 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 BPicea lutzii PILU 0.370 0.989 0.034 A 0.913 0.035 A 0.671 0.067 A 0.397 0.027 A 0.271 0.018 APicea mariana PIMA 0.380 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 BPicea pungens PIPU 0.380 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 BPicea rubens PIRU 0.380 1.000 0.057 A 0.858 0.314 B 0.832 0.112 A 0.426 0.105 B 0.347 0.140 BPicea sitchensis PISI 0.370 1.000 0.008 A 0.868 0.021 A 0.740 0.021 A 0.476 0.029 A 0.385 0.008 APicea species PICE 0.380 0.998 0.073 B 0.858 0.314 B 0.745 0.201 B 0.426 0.105 B 0.347 0.140 BPinus albicaulis PIAL 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus aristata PIAR 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus arizonica PIAR2 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus attenuata PIAT 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus balfouriana PIBA 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus banksiana PIBA2 0.400 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus clausa PICL 0.460 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus contorta PICO 0.380 0.977 0.020 A 0.956 0.013 A 0.809 0.022 A 0.512 0.019 A 0.432 0.009 APinus coulteri PICO2 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus discolor PIDI 0.500 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus echinata PIEC 0.470 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus edulis PIED 0.500 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus elliottii PIEL 0.540 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus engelmannii PIEN2 0.370 0.916 0.061 B 0.826 0.090 B 0.745 0.096 B 0.421 0.063 B 0.347 0.038 BPinus fl exilis PIFL 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus glabra PIGL2 0.370 0.998 0.061 B 0.858 0.090 B 0.745 0.106 B 0.426 0.063 B 0.347 0.124 BPinus jefferyi PIJE 0.370 0.869 0.026 A 0.835 0.042 A 0.664 0.047 A 0.454 0.039 A 0.407 0.124 BPinus lambertiana PILA 0.340 1.000 0.037 A 0.788 0.011 A 0.553 0.020 A 0.364 0.025 A 0.407 0.124 BPinus leiophylla PILE 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus monophylla PIMO 0.500 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus monticola PIMO3 0.350 0.983 0.005 A 0.866 0.012 A 0.880 0.015 A 0.421 0.063 B 0.407 0.124 BPinus muricata PIMU 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus nigra PINI 0.410 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus palustris PIPA 0.540 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus ponderosa PIPO 0.380 0.889 0.005 A 0.876 0.018 A 0.868 0.035 A 0.339 0.008 A 0.495 0.134 APinus pungens PIPU2 0.490 0.998 0.061 B 0.858 0.090 B 0.745 0.106 B 0.426 0.063 B 0.347 0.124 BPinus radiata PIRA 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus resinosa PIRE 0.410 0.829 0.061 A 0.826 0.090 B 0.659 0.106 A 0.366 0.063 A 0.268 0.124 APinus rigida PIRI 0.470 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus sabiniana PISA 0.370 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus serotina PISE 0.510 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus strobiformis PIST2 0.350 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus strobus PIST 0.340 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus sylvestris PISY 0.410 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BPinus taeda PITA 0.470 0.829 0.012 A 0.682 0.005 A 0.550 0.002 A 0.423 0.004 A 0.421 0.124 APinus virginiana PIVI 0.450 0.916 0.061 B 0.826 0.090 B 0.697 0.106 B 0.421 0.063 B 0.407 0.124 BGenus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5

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50

Planera aquatica PLAQ 0.530 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CPlatanus occidentalis PLOC 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CPopulus alba POAL 0.370 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus angustifolia POAN 0.340 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus balsamifera POBA 0.310 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus deltoides PODE 0.370 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus fremontii POFR 0.340 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus grandidentata POGR 0.360 0.933 0.067 A 0.734 0.270 C 0.828 0.066 A 0.432 0.180 B 0.297 0.094 BPopulus heterophylla POHE 0.370 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus sargentii POSA 0.370 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BPopulus species POSP 0.370 1.000 0.136 A 0.734 0.270 C 0.811 0.265 A 0.432 0.180 A 0.297 0.094 APopulus tremuloides POTR 0.350 0.967 0.136 B 0.734 0.270 C 0.819 0.265 B 0.432 0.180 B 0.297 0.094 BProsopis species PRSP 0.580 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CPrunus americana PRAM 0.470 1.000 0.136 B 0.968 0.270 B 0.668 0.265 B 0.370 0.180 C 0.282 0.094 CPrunus nigra PRNI 0.470 1.000 0.136 B 0.968 0.270 B 0.668 0.265 B 0.370 0.180 C 0.282 0.094 CPrunus pensylvanica PRPE 0.360 1.000 0.029 A 0.936 0.026 A 0.600 0.026 A 0.370 0.180 C 0.282 0.094 CPrunus serotina PRSE 0.470 1.000 0.072 A 1.000 0.049 A 0.736 0.097 A 0.370 0.180 C 0.282 0.094 CPrunus species PRSP2 0.470 1.000 0.136 B 0.968 0.270 B 0.668 0.265 B 0.370 0.180 C 0.282 0.094 CPrunus virginiana PRVI 0.360 1.000 0.136 B 0.968 0.270 B 0.668 0.265 B 0.370 0.180 C 0.282 0.094 CPseudotsuga menziesii PSME 0.450 0.858 0.013 A 0.723 0.017 A 0.444 0.022 A 0.305 0.011 A 0.329 0.110 AQuercus agrifolia QUAG 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus alba QUAL 0.600 0.973 0.031 A 0.746 0.019 A 0.558 0.045 A 0.280 0.071 A 0.428 0.011 BQuercus arizonica, grisea QUAR 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus bicolor QUBI 0.640 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus chrysolepis QUCH 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus coccinea QUCO 0.600 0.952 0.053 A 0.837 0.041 A 0.737 0.033 A 0.416 0.086 B 0.428 0.011 BQuercus douglasii QUDO 0.510 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus durandii QUDU 0.600 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus ellipsoidalis QUEL 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus emoryi QUEM 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus engelmannii QUEN 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus falcata var. falcata QUFA 0.520 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 B

Quercus falcata var. pagodaefolia QUFA2 0.610 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus gambelii QUGA 0.640 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus garryana QUGA2 0.640 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus hypoleucoides QUHY 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus ilicifolia QUIL 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus imbricaria QUIM 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus incana QUIN 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus kelloggii QUKE 0.510 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus laevis QULA 0.520 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus laurifolia QULA2 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus lobata QULO 0.640 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus lyrata QULY 0.570 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 B

(Appendix 3 continued)

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5
51

Quercus macrocarpa QUMA 0.580 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus marilandica QUMA2 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus michauxii QUMI 0.600 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus muehlenbergii QUMU 0.600 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus nigra QUNI 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus nuttallii QUNU 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus oblongifolia QUOB 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus palustris QUPA 0.580 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus phellos QUPH 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus prinus QUPR 0.570 1.000 0.078 A 0.858 0.093 A 0.516 0.042 A 0.416 0.086 B 0.428 0.011 BQuercus rubra QURU 0.560 0.991 0.043 A 0.795 0.083 A 0.690 0.081 A 0.416 0.086 B 0.428 0.011 BQuercus shumardii QUSH 0.560 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus species QUSP 0.580 0.974 0.019 A 0.797 0.039 A 0.649 0.043 A 0.483 0.020 A 0.428 0.021 AQuercus stellata QUST 0.600 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus velutina QUVE 0.560 0.981 0.051 B 0.648 0.099 A 0.809 0.114 A 0.416 0.086 B 0.428 0.011 BQuercus virginiana QUVI 0.800 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BQuercus wislizeni QUWI 0.700 0.981 0.051 B 0.779 0.099 B 0.665 0.114 B 0.416 0.086 B 0.428 0.011 BRobinia neomexicana RONE 0.660 1.000 0.136 B 0.848 0.270 B 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CRobinia pseudoacacia ROPS 0.660 1.000 0.031 A 0.848 0.053 A 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CSabal species SASP 0.550 0.927 0.082 A 0.744 0.091 A 0.615 0.074 A 0.265 0.112 A 0.282 0.094 CSalix species SASP 0.360 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CSapium sebiferum SASE 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CSassafras albidum SAAL 0.420 1.000 0.028 A 0.924 0.042 A 0.805 0.025 A 0.370 0.180 C 0.282 0.094 CSequoia sempervirens SESE 0.340 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CSequoiadendron giganteum SEGI 0.340 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CSorbus americana SOAM 0.420 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CTamarix species TASP 0.400 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CTaxodium distichum var. nutans TADI 0.420 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 C

Taxus brevifolia TABR 0.600 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CThuja occidentalis THOC 0.290 1.000 0.038 A 0.835 0.195 B 1.000 0.053 A 0.461 0.001 B 0.461 0.055 BThuja plicata THPL 0.310 1.000 0.073 A 0.835 0.390 A 0.800 0.030 A 0.462 0.002 A 0.461 0.110 ATilia americana TIAM 0.320 1.000 0.024 A 1.000 0.032 A 0.800 0.019 A 0.370 0.180 C 0.282 0.094 CTilia heterophylla TIHE 0.320 1.000 0.136 B 1.000 0.270 B 0.800 0.265 B 0.370 0.180 C 0.282 0.094 CTilia species TISP 0.320 1.000 0.136 B 1.000 0.270 B 0.800 0.265 B 0.370 0.180 C 0.282 0.094 CTorreya californica TOCA 0.340 0.956 0.073 C 0.827 0.314 C 0.678 0.201 C 0.426 0.105 C 0.366 0.140 CTsuga canadensis TSCA 0.400 0.995 0.059 A 0.767 0.314 B 0.813 0.084 A 0.425 0.105 B 0.333 0.140 BTsuga heterophylla TSHE 0.420 0.890 0.022 A 0.767 0.023 A 0.636 0.019 A 0.425 0.025 A 0.332 0.106 ATsuga mertensiana TSME 0.420 0.916 0.056 B 0.767 0.314 B 0.681 0.086 B 0.425 0.105 B 0.333 0.140 BTsuga species TSSP 0.380 0.916 0.056 B 0.767 0.314 B 0.681 0.086 B 0.425 0.105 B 0.333 0.140 BUlmus alata ULAL 0.570 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus americana ULAM 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus crassifolia ULCR 0.570 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus pumila ULPU 0.460 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus rubra ULRU 0.480 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 C

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5

(Appendix 3 continued on next page)
52

Ulmus serotina ULSE 0.570 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus species ULSP 0.500 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUlmus thomasii ULTH 0.570 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CUmbellularia californica UMCA 0.510 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 CVaccinium arboreum VAAR 0.470 0.944 0.136 C 0.734 0.270 C 0.583 0.265 C 0.370 0.180 C 0.282 0.094 C

(Appendix 3 continued)

Genus Species code den0 rel1 unc1 cod1 rel2 unc2 cod2 rel3 unc3 cod3 rel4 unc4 cod4 rel5 unc5 cod5
53

CWD Relative Density Predictions Metadata (Appendix 3) Fieldname Defi nition Code/units Methodgenus genus species species code genus-species code den0 green density g/cm3

unc0 uncertainty green density rel1 relative density decay class 1 dimensionless unc1 uncertainty of rel1 dimensionless cod1 uncertainty code for rel1 A Species sampled B Genera sampled C Species and genus not sampledrel2 relative density decay class 2 dimensionless unc2 uncertainty of rel2 dimensionless cod2 uncertainty code for rel2 A Species sampled B Genera sampled C Species and genus not sampledrel3 relative density decay class 3 dimensionless unc3 uncertainty of rel3 dimensionless cod3 uncertainty code for rel3 A Species sampled B Genera sampled C Species and genus not sampledrel4 relative density decay class 4 dimensionless unc4 uncertainty of rel4 dimensionless cod4 uncertainty code for rel4 A Species sampled B Genera sampled C Species and genus not sampledrel5 relative density decay class 5 dimensionless unc5 uncertainty of rel5 dimensionless cod5 uncertainty code for rel5 A Species sampled B Genera sampled C Species and genus not sampled
54Appendix 4.—FWD absolute and relative density and their uncertainty for each size class by species present in 

 the U.S. FIA systemSpecies Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeABAM 1 0.40 1.29 0.02 B 0.51 0.01 A 0.82 0.19 C 0.42 0.04 CABAM 2 0.40 1.38 0.02 A 0.55 0.01 A 0.96 0.01 A 0.53 0.01 AABAM 3 0.40 1.12 0.03 A 0.45 0.01 A 0.99 0.03 A 0.44 0.01 AABAM 4 0.40 1.35 0.01 B 0.54 0.00 A 0.80 0.12 C 0.43 0.06 CABBA 1 0.34 2.04 0.09 B 0.70 0.03 A 0.64 0.06 B 0.45 0.04 BABBA 2 0.34 1.19 0.21 A 0.41 0.03 A 1.00 0.01 A 0.41 0.04 BABBA 3 0.34 1.11 0.23 C 0.38 0.08 B 1.00 0.09 B 0.38 0.04 BABBA 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CABBR 1 0.36 1.47 0.31 C 0.53 0.11 B 0.82 0.19 C 0.44 0.07 CABBR 2 0.36 1.22 0.18 C 0.44 0.07 B 1.03 0.13 C 0.45 0.04 CABBR 3 0.36 1.11 0.23 C 0.40 0.08 B 0.95 0.12 C 0.38 0.05 CABBR 4 0.36 1.25 0.17 C 0.45 0.06 B 0.80 0.12 C 0.36 0.06 CABCO 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CABCO 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CABCO 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CABCO 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CABFR 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 CABFR 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CABFR 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CABFR 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CABGR 1 0.35 1.47 0.31 C 0.51 0.11 B 1.14 0.11 B 0.58 0.05 BABGR 2 0.35 1.22 0.18 C 0.43 0.06 B 1.16 0.11 B 0.50 0.05 BABGR 3 0.35 1.11 0.23 C 0.39 0.08 B 0.95 0.12 C 0.37 0.04 CABGR 4 0.35 1.25 0.17 C 0.44 0.06 B 0.80 0.12 C 0.35 0.06 CABLA 1 0.31 2.02 0.05 B 0.63 0.02 A 0.65 0.06 B 0.41 0.04 BABLA 2 0.31 1.38 0.04 B 0.43 0.01 A 1.03 0.13 C 0.44 0.03 CABLA 3 0.31 1.21 0.03 B 0.38 0.01 A 0.95 0.12 C 0.36 0.03 CABLA 4 0.31 1.50 0.05 B 0.47 0.02 A 0.85 0.08 B 0.40 0.04 BABMA 1 0.36 1.47 0.31 C 0.53 0.11 B 0.82 0.19 C 0.44 0.07 CABMA 2 0.36 1.22 0.18 C 0.44 0.07 B 1.03 0.13 C 0.45 0.04 CABMA 3 0.36 1.11 0.23 C 0.40 0.08 B 0.95 0.12 C 0.38 0.05 CABMA 4 0.36 1.25 0.17 C 0.45 0.06 B 0.80 0.12 C 0.36 0.06 CABMA2 1 0.36 1.47 0.31 C 0.53 0.11 B 0.82 0.19 C 0.44 0.07 CABMA2 2 0.36 1.22 0.18 C 0.44 0.07 B 1.03 0.13 C 0.45 0.04 CABMA2 3 0.36 1.11 0.23 C 0.40 0.08 B 0.95 0.12 C 0.38 0.05 CABMA2 4 0.36 1.25 0.17 C 0.45 0.06 B 0.80 0.12 C 0.36 0.06 CABPR 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CABPR 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CABPR 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CABPR 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 C
55

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeABIE 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 CABIE 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CABIE 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CABIE 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CACSP2 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CACSP2 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CACSP2 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CACSP2 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CACBA 1 0.54 1.47 0.31 C 0.79 0.16 B 0.83 0.10 C 0.65 0.13 CACBA 2 0.54 1.22 0.18 C 0.66 0.10 B 0.85 0.15 C 0.56 0.07 CACBA 3 0.54 1.11 0.23 C 0.60 0.12 B 0.92 0.14 C 0.55 0.08 CACBA 4 0.54 1.25 0.17 C 0.68 0.09 B 0.71 0.12 C 0.49 0.07 CACGL 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CACGL 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CACGL 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACGL 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CACGR 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CACGR 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CACGR 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACGR 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CACMA 1 0.44 0.96 0.01 B 0.42 0.01 A 0.83 0.10 C 0.35 0.03 CACMA 2 0.44 1.13 0.01 B 0.50 0.00 A 0.85 0.15 C 0.43 0.03 CACMA 3 0.44 1.05 0.01 B 0.46 0.01 A 0.92 0.14 C 0.42 0.04 CACMA 4 0.44 1.07 0.01 B 0.47 0.00 A 0.71 0.12 C 0.34 0.04 CACNE 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CACNE 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CACNE 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACNE 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CACNI 1 0.52 1.47 0.31 C 0.76 0.16 B 0.83 0.10 C 0.63 0.13 CACNI 2 0.52 1.22 0.18 C 0.63 0.10 B 0.85 0.15 C 0.54 0.07 CACNI 3 0.52 1.11 0.23 C 0.58 0.12 B 0.92 0.14 C 0.53 0.08 CACNI 4 0.52 1.25 0.17 C 0.65 0.09 B 0.71 0.12 C 0.46 0.07 CACPE 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CACPE 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CACPE 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACPE 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CACRU 1 0.49 1.19 0.03 B 0.58 0.02 A 0.83 0.10 C 0.48 0.03 CACRU 2 0.49 1.13 0.01 B 0.55 0.01 A 0.85 0.15 C 0.47 0.03 CACRU 3 0.49 1.03 0.02 B 0.51 0.01 A 0.92 0.14 C 0.47 0.04 CACRU 4 0.49 1.12 0.01 B 0.55 0.01 A 0.71 0.12 C 0.39 0.04 CACSA2 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 C

(Appendix 4 continued on next page)
56

(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeACSA2 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CACSA2 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACSA2 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CACSA 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CACSA 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CACSA 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CACSA 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CACER 1 0.49 1.47 0.31 C 0.72 0.15 B 0.83 0.10 C 0.60 0.11 CACER 2 0.49 1.22 0.18 C 0.60 0.09 B 0.85 0.15 C 0.51 0.06 CACER 3 0.49 1.11 0.23 C 0.54 0.11 B 0.92 0.14 C 0.50 0.07 CACER 4 0.49 1.25 0.17 C 0.61 0.08 B 0.71 0.12 C 0.44 0.07 CACSP 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CACSP 2 0.44 1.22 0.18 C 0.54 0.08 B 0.87 0.05 B 0.47 0.03 BACSP 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CACSP 4 0.44 1.25 0.17 C 0.55 0.08 B 0.81 0.07 B 0.44 0.04 BAECA 1 0.38 1.47 0.31 C 0.56 0.12 B 0.83 0.10 C 0.46 0.07 CAECA 2 0.38 1.22 0.18 C 0.46 0.07 B 0.85 0.15 C 0.39 0.04 CAECA 3 0.38 1.11 0.23 C 0.42 0.09 B 0.92 0.14 C 0.39 0.05 CAECA 4 0.38 1.25 0.17 C 0.48 0.07 B 0.71 0.12 C 0.34 0.05 CAESP 1 0.33 1.47 0.31 C 0.49 0.10 B 0.83 0.10 C 0.41 0.06 CAESP 2 0.33 1.22 0.18 C 0.40 0.06 B 0.85 0.15 C 0.34 0.04 CAESP 3 0.33 1.11 0.23 C 0.37 0.08 B 0.92 0.14 C 0.34 0.05 CAESP 4 0.33 1.25 0.17 C 0.41 0.06 B 0.71 0.12 C 0.29 0.05 CAIAL 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CAIAL 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CAIAL 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CAIAL 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CALFO 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CALFO 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CALFO 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CALFO 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CALRU 1 0.40 1.86 0.03 B 0.74 0.01 A 0.83 0.10 C 0.61 0.03 CALRU 2 0.40 1.27 0.01 B 0.51 0.00 A 0.85 0.15 C 0.43 0.03 CALRU 3 0.40 1.15 0.01 B 0.46 0.00 A 0.92 0.14 C 0.42 0.03 CALRU 4 0.40 1.39 0.02 B 0.56 0.01 A 0.71 0.12 C 0.40 0.04 CALSP 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CALSP 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CALSP 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CALSP 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CALTE 1 0.40 1.27 0.03 B 0.51 0.01 A 0.83 0.10 C 0.42 0.03 CALTE 2 0.40 0.97 0.02 B 0.39 0.01 A 0.85 0.15 C 0.33 0.03 C
57

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeALTE 3 0.40 1.11 0.23 C 0.44 0.09 B 0.92 0.14 C 0.40 0.05 CALTE 4 0.40 1.07 0.03 B 0.43 0.01 A 0.71 0.12 C 0.31 0.04 CAMSP 1 0.66 1.47 0.31 C 0.97 0.20 B 0.83 0.10 C 0.80 0.20 CAMSP 2 0.66 1.22 0.18 C 0.80 0.12 B 0.85 0.15 C 0.68 0.10 CAMSP 3 0.66 1.11 0.23 C 0.73 0.15 B 0.92 0.14 C 0.67 0.11 CAMSP 4 0.66 1.25 0.17 C 0.83 0.11 B 0.71 0.12 C 0.59 0.10 CARME 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CARME 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CARME 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CARME 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CASTR 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CASTR 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CASTR 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CASTR 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CBEAL 1 0.55 1.47 0.31 C 0.81 0.17 B 0.83 0.10 C 0.67 0.14 CBEAL 2 0.55 1.22 0.18 C 0.67 0.10 B 0.85 0.15 C 0.57 0.07 CBEAL 3 0.55 1.11 0.23 C 0.61 0.13 B 0.92 0.14 C 0.56 0.09 CBEAL 4 0.55 1.25 0.17 C 0.69 0.10 B 0.71 0.12 C 0.49 0.08 CBELE 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CBELE 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CBELE 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CBELE 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CBENI 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CBENI 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CBENI 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CBENI 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CBEOC 1 0.53 1.47 0.31 C 0.78 0.16 B 0.83 0.10 C 0.65 0.13 CBEOC 2 0.53 1.22 0.18 C 0.65 0.10 B 0.85 0.15 C 0.55 0.07 CBEOC 3 0.53 1.11 0.23 C 0.59 0.12 B 0.92 0.14 C 0.54 0.08 CBEOC 4 0.53 1.25 0.17 C 0.66 0.09 B 0.71 0.12 C 0.47 0.07 CBEPA 1 0.48 1.43 0.04 B 0.69 0.02 A 0.83 0.10 C 0.57 0.03 CBEPA 2 0.48 1.07 0.05 A 0.52 0.01 A 0.85 0.15 C 0.44 0.03 CBEPA 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 CBEPA 4 0.48 1.25 0.17 C 0.60 0.08 B 0.71 0.12 C 0.43 0.06 CBEPAC 1 0.48 1.47 0.31 C 0.71 0.15 B 0.83 0.10 C 0.59 0.11 CBEPAC 2 0.48 1.22 0.18 C 0.58 0.09 B 0.85 0.15 C 0.49 0.06 CBEPAC 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 CBEPAC 4 0.48 1.25 0.17 C 0.60 0.08 B 0.71 0.12 C 0.43 0.06 CBEPO 1 0.45 1.47 0.31 C 0.66 0.14 B 0.83 0.10 C 0.55 0.10 CBEPO 2 0.45 1.22 0.18 C 0.55 0.08 B 0.85 0.15 C 0.47 0.05 CBEPO 3 0.45 1.11 0.23 C 0.50 0.10 B 0.92 0.14 C 0.46 0.06 C

(Appendix 4 continued on next page)
58

(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeBEPO 4 0.45 1.25 0.17 C 0.56 0.08 B 0.71 0.12 C 0.40 0.06 CBESP 1 0.48 1.47 0.31 C 0.71 0.15 B 0.83 0.10 C 0.59 0.11 CBESP 2 0.48 1.22 0.18 C 0.58 0.09 B 0.68 0.04 B 0.40 0.03 ABESP 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 CBESP 4 0.48 1.25 0.17 C 0.60 0.08 B 0.63 0.08 B 0.38 0.05 ABEPL 1 0.45 1.47 0.31 C 0.66 0.14 B 0.83 0.10 C 0.55 0.10 CBEPL 2 0.45 0.40 0.02 B 0.18 0.01 A 0.85 0.15 C 0.15 0.03 CBEPL 3 0.45 1.11 0.23 C 0.50 0.10 B 0.92 0.14 C 0.46 0.06 CBEPL 4 0.45 0.40 0.02 B 0.18 0.01 A 0.71 0.12 C 0.13 0.04 CBULA 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CBULA 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CBULA 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CBULA 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CCADE3 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CCADE3 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CCADE3 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CCADE3 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CCACA 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CCACA 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CCACA 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CCACA 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CCAAQ 1 0.61 1.47 0.31 C 0.90 0.19 B 0.83 0.10 C 0.75 0.17 CCAAQ 2 0.61 1.22 0.18 C 0.74 0.11 B 0.85 0.15 C 0.63 0.09 CCAAQ 3 0.61 1.11 0.23 C 0.68 0.14 B 0.92 0.14 C 0.62 0.10 CCAAQ 4 0.61 1.25 0.17 C 0.76 0.11 B 0.71 0.12 C 0.54 0.09 CCACO 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CCACO 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CCACO 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CCACO 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CCAGL 1 0.66 1.47 0.31 C 0.97 0.20 B 0.83 0.10 C 0.80 0.20 CCAGL 2 0.66 1.22 0.18 C 0.80 0.12 B 0.85 0.15 C 0.68 0.10 CCAGL 3 0.66 1.11 0.23 C 0.73 0.15 B 0.92 0.14 C 0.67 0.11 CCAGL 4 0.66 1.25 0.17 C 0.83 0.11 B 0.71 0.12 C 0.59 0.10 CCAIL 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CCAIL 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CCAIL 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CCAIL 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CCALA 1 0.62 1.47 0.31 C 0.91 0.19 B 0.83 0.10 C 0.75 0.18 CCALA 2 0.62 1.22 0.18 C 0.76 0.11 B 0.85 0.15 C 0.65 0.09 CCALA 3 0.62 1.11 0.23 C 0.69 0.14 B 0.92 0.14 C 0.63 0.10 CCALA 4 0.62 1.25 0.17 C 0.78 0.11 B 0.71 0.12 C 0.56 0.10 C
59

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeCAMY 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CCAMY 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CCAMY 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CCAMY 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CCAOV 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CCAOV 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CCAOV 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CCAOV 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CCASP 1 0.62 1.47 0.31 C 0.91 0.19 B 0.83 0.10 C 0.75 0.18 CCASP 2 0.62 1.22 0.18 C 0.76 0.11 B 0.85 0.15 C 0.65 0.09 CCASP 3 0.62 1.11 0.23 C 0.69 0.14 B 0.92 0.14 C 0.63 0.10 CCASP 4 0.62 1.25 0.17 C 0.78 0.11 B 0.71 0.12 C 0.56 0.10 CCATE 1 0.54 1.47 0.31 C 0.79 0.16 B 0.83 0.10 C 0.65 0.13 CCATE 2 0.54 1.22 0.18 C 0.66 0.10 B 0.85 0.15 C 0.56 0.07 CCATE 3 0.54 1.11 0.23 C 0.60 0.12 B 0.92 0.14 C 0.55 0.08 CCATE 4 0.54 1.25 0.17 C 0.68 0.09 B 0.71 0.12 C 0.49 0.07 CCATO 1 0.64 1.24 0.02 B 0.79 0.01 A 0.83 0.10 C 0.65 0.03 CCATO 2 0.64 1.15 0.01 B 0.73 0.01 A 0.85 0.15 C 0.62 0.03 CCATO 3 0.64 1.12 0.01 B 0.72 0.01 A 0.92 0.14 C 0.66 0.04 CCATO 4 0.64 1.16 0.01 B 0.74 0.01 A 0.71 0.12 C 0.53 0.04 CCADE 1 0.40 1.34 0.02 B 0.54 0.01 A 0.83 0.10 C 0.45 0.03 CCADE 2 0.40 1.30 0.01 B 0.52 0.01 A 0.84 0.06 B 0.44 0.03 ACADE 3 0.40 0.83 0.01 B 0.33 0.00 A 0.92 0.14 C 0.30 0.03 CCADE 4 0.40 1.19 0.02 B 0.48 0.01 A 0.71 0.12 C 0.34 0.04 CCAOZ 1 0.40 1.47 0.31 C 0.59 0.12 B 0.83 0.10 C 0.49 0.08 CCAOZ 2 0.40 1.22 0.18 C 0.49 0.07 B 0.85 0.15 C 0.42 0.05 CCAOZ 3 0.40 1.11 0.23 C 0.44 0.09 B 0.92 0.14 C 0.40 0.05 CCAOZ 4 0.40 1.25 0.17 C 0.50 0.07 B 0.71 0.12 C 0.36 0.06 CCAPU 1 0.40 1.47 0.31 C 0.59 0.12 B 0.83 0.10 C 0.49 0.08 CCAPU 2 0.40 1.22 0.18 C 0.49 0.07 B 0.85 0.15 C 0.42 0.05 CCAPU 3 0.40 1.11 0.23 C 0.44 0.09 B 0.92 0.14 C 0.40 0.05 CCAPU 4 0.40 1.25 0.17 C 0.50 0.07 B 0.71 0.12 C 0.36 0.06 CCASTA 1 0.42 1.47 0.31 C 0.62 0.13 B 0.83 0.10 C 0.51 0.09 CCASTA 2 0.42 1.22 0.18 C 0.51 0.08 B 0.85 0.15 C 0.43 0.05 CCASTA 3 0.42 1.11 0.23 C 0.47 0.10 B 0.92 0.14 C 0.43 0.06 CCASTA 4 0.42 1.25 0.17 C 0.53 0.07 B 0.71 0.12 C 0.38 0.06 CCATAL 1 0.38 1.47 0.31 C 0.56 0.12 B 0.83 0.10 C 0.46 0.07 CCATAL 2 0.38 1.22 0.18 C 0.46 0.07 B 0.85 0.15 C 0.39 0.04 CCATAL 3 0.38 1.11 0.23 C 0.42 0.09 B 0.92 0.14 C 0.39 0.05 CCATAL 4 0.38 1.25 0.17 C 0.48 0.07 B 0.71 0.12 C 0.34 0.05 CCELA 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 C

(Appendix 4 continued on next page)
60

(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeCELA 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CCELA 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CCELA 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CCEOC 1 0.49 1.47 0.31 C 0.72 0.15 B 0.83 0.10 C 0.60 0.11 CCEOC 2 0.49 1.22 0.18 C 0.60 0.09 B 0.85 0.15 C 0.51 0.06 CCEOC 3 0.49 1.11 0.23 C 0.54 0.11 B 0.92 0.14 C 0.50 0.07 CCEOC 4 0.49 1.25 0.17 C 0.61 0.08 B 0.71 0.12 C 0.44 0.07 CCESP 1 0.49 1.47 0.31 C 0.72 0.15 B 0.83 0.10 C 0.60 0.11 CCESP 2 0.49 1.22 0.18 C 0.60 0.09 B 0.85 0.15 C 0.51 0.06 CCESP 3 0.49 1.11 0.23 C 0.54 0.11 B 0.92 0.14 C 0.50 0.07 CCESP 4 0.49 1.25 0.17 C 0.61 0.08 B 0.71 0.12 C 0.44 0.07 CCECA 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CCECA 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CCECA 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CCECA 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CCEIN 1 1.00 1.47 0.31 C 1.47 0.31 B 0.83 0.10 C 1.22 0.46 CCEIN 2 1.00 1.22 0.18 C 1.22 0.18 B 0.85 0.15 C 1.04 0.22 CCEIN 3 1.00 1.11 0.23 C 1.11 0.23 B 0.92 0.14 C 1.02 0.26 CCEIN 4 1.00 1.25 0.17 C 1.25 0.17 B 0.71 0.12 C 0.89 0.22 CCELE 1 1.00 1.47 0.31 C 1.47 0.31 B 0.83 0.10 C 1.22 0.46 CCELE 2 1.00 1.22 0.18 C 1.22 0.18 B 0.85 0.15 C 1.04 0.22 CCELE 3 1.00 1.11 0.23 C 1.11 0.23 B 0.92 0.14 C 1.02 0.26 CCELE 4 1.00 1.25 0.17 C 1.25 0.17 B 0.71 0.12 C 0.89 0.22 CCEMO 1 1.00 1.47 0.31 C 1.47 0.31 B 0.83 0.10 C 1.22 0.46 CCEMO 2 1.00 1.22 0.18 C 1.22 0.18 B 0.85 0.15 C 1.04 0.22 CCEMO 3 1.00 1.11 0.23 C 1.11 0.23 B 0.92 0.14 C 1.02 0.26 CCEMO 4 1.00 1.25 0.17 C 1.25 0.17 B 0.71 0.12 C 0.89 0.22 CCHLA 1 0.39 1.47 0.31 C 0.57 0.12 B 0.82 0.19 C 0.47 0.08 CCHLA 2 0.39 1.22 0.18 C 0.48 0.07 B 1.03 0.13 C 0.49 0.04 CCHLA 3 0.39 1.11 0.23 C 0.43 0.09 B 0.95 0.12 C 0.41 0.05 CCHLA 4 0.39 1.25 0.17 C 0.49 0.07 B 0.80 0.12 C 0.39 0.07 CCHNO 1 0.42 1.47 0.31 C 0.62 0.13 B 0.82 0.19 C 0.51 0.09 CCHNO 2 0.42 1.22 0.18 C 0.51 0.08 B 1.03 0.13 C 0.52 0.05 CCHNO 3 0.42 1.11 0.23 C 0.47 0.10 B 0.95 0.12 C 0.45 0.06 CCHNO 4 0.42 1.25 0.17 C 0.53 0.07 B 0.80 0.12 C 0.42 0.07 CCHTH 1 0.31 1.47 0.31 C 0.46 0.09 B 0.82 0.19 C 0.38 0.06 CCHTH 2 0.31 1.22 0.18 C 0.38 0.06 B 1.03 0.13 C 0.39 0.03 CCHTH 3 0.31 1.11 0.23 C 0.34 0.07 B 0.95 0.12 C 0.32 0.04 CCHTH 4 0.31 1.25 0.17 C 0.39 0.05 B 0.80 0.12 C 0.31 0.06 CCLLU 1 0.52 1.47 0.31 C 0.76 0.16 B 0.83 0.10 C 0.63 0.13 CCLLU 2 0.52 1.22 0.18 C 0.63 0.10 B 0.85 0.15 C 0.54 0.07 C
61

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeCLLU 3 0.52 1.11 0.23 C 0.58 0.12 B 0.92 0.14 C 0.53 0.08 CCLLU 4 0.52 1.25 0.17 C 0.65 0.09 B 0.71 0.12 C 0.46 0.07 CCOFL 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CCOFL 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CCOFL 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CCOFL 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CCONU 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CCONU 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CCONU 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CCONU 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CCOSP 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CCOSP 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CCOSP 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CCOSP 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CCOOB 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CCOOB 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CCOOB 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CCOOB 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CCRSP 1 0.62 1.47 0.31 C 0.91 0.19 B 0.83 0.10 C 0.75 0.18 CCRSP 2 0.62 1.22 0.18 C 0.76 0.11 B 0.85 0.15 C 0.65 0.09 CCRSP 3 0.62 1.11 0.23 C 0.69 0.14 B 0.92 0.14 C 0.63 0.10 CCRSP 4 0.62 1.25 0.17 C 0.78 0.11 B 0.71 0.12 C 0.56 0.10 CCUSP 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CCUSP 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CCUSP 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CCUSP 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CDIVI 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CDIVI 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CDIVI 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CDIVI 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CEUSP 1 0.67 1.47 0.31 C 0.98 0.20 B 0.83 0.10 C 0.81 0.20 CEUSP 2 0.67 1.22 0.18 C 0.82 0.12 B 0.85 0.15 C 0.70 0.10 CEUSP 3 0.67 1.11 0.23 C 0.74 0.15 B 0.92 0.14 C 0.68 0.12 CEUSP 4 0.67 1.25 0.17 C 0.84 0.12 B 0.71 0.12 C 0.60 0.11 CFAGR 1 0.56 1.26 0.02 B 0.70 0.01 A 0.83 0.10 C 0.58 0.03 CFAGR 2 0.56 1.06 0.02 B 0.60 0.01 A 0.77 0.04 B 0.46 0.03 AFAGR 3 0.56 1.04 0.01 B 0.58 0.00 A 0.92 0.14 C 0.53 0.03 CFAGR 4 0.56 1.11 0.02 B 0.62 0.01 A 0.66 0.04 B 0.41 0.03 AFRAM 1 0.55 1.47 0.31 C 0.81 0.17 B 0.83 0.10 C 0.67 0.14 CFRAM 2 0.55 1.22 0.18 C 0.67 0.10 B 0.85 0.15 C 0.57 0.07 CFRAM 3 0.55 1.11 0.23 C 0.61 0.13 B 0.92 0.14 C 0.56 0.09 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeFRAM 4 0.55 1.25 0.17 C 0.69 0.10 B 0.71 0.12 C 0.49 0.08 CFRLA 1 0.50 1.47 0.31 C 0.74 0.15 B 0.83 0.10 C 0.61 0.12 CFRLA 2 0.50 1.22 0.18 C 0.61 0.09 B 0.85 0.15 C 0.52 0.06 CFRLA 3 0.50 1.11 0.23 C 0.55 0.11 B 0.92 0.14 C 0.50 0.07 CFRLA 4 0.50 1.25 0.17 C 0.63 0.09 B 0.71 0.12 C 0.45 0.07 CFRNI 1 0.45 1.47 0.31 C 0.66 0.14 B 0.83 0.10 C 0.55 0.10 CFRNI 2 0.45 1.22 0.18 C 0.55 0.08 B 0.85 0.15 C 0.47 0.05 CFRNI 3 0.45 1.11 0.23 C 0.50 0.10 B 0.92 0.14 C 0.46 0.06 CFRNI 4 0.45 1.25 0.17 C 0.56 0.08 B 0.71 0.12 C 0.40 0.06 CFRPE 1 0.53 1.47 0.31 C 0.78 0.16 B 0.83 0.10 C 0.65 0.13 CFRPE 2 0.53 1.22 0.18 C 0.65 0.10 B 0.85 0.15 C 0.55 0.07 CFRPE 3 0.53 1.11 0.23 C 0.59 0.12 B 0.92 0.14 C 0.54 0.08 CFRPE 4 0.53 1.25 0.17 C 0.66 0.09 B 0.71 0.12 C 0.47 0.07 CFRPR 1 0.54 1.47 0.31 C 0.79 0.16 B 0.83 0.10 C 0.65 0.13 CFRPR 2 0.54 1.22 0.18 C 0.66 0.10 B 0.85 0.15 C 0.56 0.07 CFRPR 3 0.54 1.11 0.23 C 0.60 0.12 B 0.92 0.14 C 0.55 0.08 CFRPR 4 0.54 1.25 0.17 C 0.68 0.09 B 0.71 0.12 C 0.49 0.07 CFRQU 1 0.53 1.47 0.31 C 0.78 0.16 B 0.83 0.10 C 0.65 0.13 CFRQU 2 0.53 1.22 0.18 C 0.65 0.10 B 0.85 0.15 C 0.55 0.07 CFRQU 3 0.53 1.11 0.23 C 0.59 0.12 B 0.92 0.14 C 0.54 0.08 CFRQU 4 0.53 1.25 0.17 C 0.66 0.09 B 0.71 0.12 C 0.47 0.07 CFRSP 1 0.54 1.47 0.31 C 0.79 0.16 B 0.83 0.10 C 0.65 0.13 CFRSP 2 0.54 1.22 0.18 C 0.66 0.10 B 0.85 0.15 C 0.56 0.07 CFRSP 3 0.54 1.11 0.23 C 0.60 0.12 B 0.92 0.14 C 0.55 0.08 CFRSP 4 0.54 1.25 0.17 C 0.68 0.09 B 0.71 0.12 C 0.49 0.07 CGLAQ 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CGLAQ 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CGLAQ 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CGLAQ 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CGLTR 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CGLTR 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CGLTR 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CGLTR 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CGOLA 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CGOLA 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CGOLA 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CGOLA 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CGYDI 1 0.50 1.47 0.31 C 0.74 0.15 B 0.83 0.10 C 0.61 0.12 CGYDI 2 0.50 1.22 0.18 C 0.61 0.09 B 0.85 0.15 C 0.52 0.06 CGYDI 3 0.50 1.11 0.23 C 0.55 0.11 B 0.92 0.14 C 0.50 0.07 CGYDI 4 0.50 1.25 0.17 C 0.63 0.09 B 0.71 0.12 C 0.45 0.07 C
63

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeHASP 1 0.32 1.47 0.31 C 0.47 0.10 B 0.83 0.10 C 0.39 0.06 CHASP 2 0.32 1.22 0.18 C 0.39 0.06 B 1.09 0.06 B 0.43 0.02 AHASP 3 0.32 1.11 0.23 C 0.35 0.07 B 0.92 0.14 C 0.32 0.04 CHASP 4 0.32 1.25 0.17 C 0.40 0.06 B 0.89 0.08 B 0.36 0.03 AHARDW 1 0.51 1.28 0.07 B 0.65 0.03 A 0.83 0.10 C 0.54 0.04 CHARDW 2 0.51 1.13 0.03 B 0.58 0.01 A 0.63 0.02 B 0.37 0.01 AHARDW 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CHARDW 4 0.51 1.25 0.17 C 0.64 0.09 B 0.54 0.03 B 0.35 0.02 AILOP 1 0.50 1.47 0.31 C 0.74 0.15 B 0.83 0.10 C 0.61 0.12 CILOP 2 0.50 1.22 0.18 C 0.61 0.09 B 0.85 0.15 C 0.52 0.06 CILOP 3 0.50 1.11 0.23 C 0.55 0.11 B 0.92 0.14 C 0.50 0.07 CILOP 4 0.50 1.25 0.17 C 0.63 0.09 B 0.71 0.12 C 0.45 0.07 CJUCI 1 0.36 1.47 0.31 C 0.53 0.11 B 0.83 0.10 C 0.44 0.07 CJUCI 2 0.36 1.22 0.18 C 0.44 0.07 B 0.85 0.15 C 0.38 0.04 CJUCI 3 0.36 1.11 0.23 C 0.40 0.08 B 0.92 0.14 C 0.37 0.05 CJUCI 4 0.36 1.25 0.17 C 0.45 0.06 B 0.71 0.12 C 0.32 0.05 CJUSP 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CJUSP 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CJUSP 3 0.51 1.11 0.23 C 0.56 0.12 B 0.95 0.12 C 0.53 0.07 CJUSP 4 0.51 1.25 0.17 C 0.64 0.09 B 0.80 0.12 C 0.51 0.08 CJUNI 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CJUNI 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CJUNI 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CJUNI 4 0.51 1.25 0.17 C 0.64 0.09 B 0.71 0.12 C 0.46 0.07 CJUMO 1 0.45 1.47 0.31 C 0.66 0.14 B 0.82 0.19 C 0.54 0.10 CJUMO 2 0.45 1.22 0.18 C 0.55 0.08 B 1.03 0.13 C 0.56 0.05 CJUMO 3 0.45 1.11 0.23 C 0.50 0.10 B 0.95 0.12 C 0.48 0.06 CJUMO 4 0.45 1.25 0.17 C 0.56 0.08 B 0.80 0.12 C 0.45 0.07 CJUOC 1 0.44 1.47 0.31 C 0.65 0.13 B 0.82 0.19 C 0.54 0.10 CJUOC 2 0.44 1.22 0.18 C 0.54 0.08 B 1.03 0.13 C 0.55 0.05 CJUOC 3 0.44 1.11 0.23 C 0.49 0.10 B 0.95 0.12 C 0.47 0.06 CJUOC 4 0.44 1.25 0.17 C 0.55 0.08 B 0.80 0.12 C 0.44 0.07 CJUSP2 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CJUSP2 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CJUSP2 3 0.44 1.11 0.23 C 0.49 0.10 B 0.95 0.12 C 0.47 0.06 CJUSP2 4 0.44 1.25 0.17 C 0.55 0.08 B 0.80 0.12 C 0.44 0.07 CJUVI 1 0.37 1.94 0.07 B 0.72 0.03 A 0.82 0.19 C 0.59 0.05 CJUVI 2 0.37 1.38 0.03 B 0.51 0.01 A 1.03 0.13 C 0.52 0.03 CJUVI 3 0.37 1.24 0.02 B 0.46 0.01 A 0.95 0.12 C 0.44 0.03 CJUVI 4 0.37 1.48 0.05 B 0.55 0.02 A 0.80 0.12 C 0.44 0.06 CLALA 1 0.49 1.47 0.31 C 0.72 0.15 B 0.69 0.01 A 0.50 0.05 B

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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeLALA 2 0.49 1.22 0.18 C 0.60 0.09 B 0.86 0.01 A 0.52 0.05 BLALA 3 0.49 1.11 0.23 C 0.54 0.11 B 1.01 0.01 A 0.55 0.05 BLALA 4 0.49 1.25 0.17 C 0.61 0.08 B 0.71 0.12 C 0.44 0.07 CLALY 1 0.48 1.47 0.31 C 0.71 0.15 B 0.83 0.10 C 0.59 0.11 CLALY 2 0.48 1.22 0.18 C 0.58 0.09 B 0.85 0.15 C 0.49 0.06 CLALY 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 CLALY 4 0.48 1.25 0.17 C 0.60 0.08 B 0.71 0.12 C 0.43 0.06 CLAOC 1 0.48 1.47 0.31 C 0.71 0.15 B 0.70 0.06 A 0.50 0.05 BLAOC 2 0.48 1.22 0.18 C 0.58 0.09 B 0.93 0.09 B 0.54 0.05 BLAOC 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 CLAOC 4 0.48 1.25 0.17 C 0.60 0.08 B 0.92 0.09 B 0.55 0.05 BLASP 1 0.44 1.47 0.31 C 0.65 0.13 B 0.83 0.10 C 0.54 0.09 CLASP 2 0.44 1.22 0.18 C 0.54 0.08 B 0.85 0.15 C 0.46 0.05 CLASP 3 0.44 1.11 0.23 C 0.49 0.10 B 0.92 0.14 C 0.45 0.06 CLASP 4 0.44 1.25 0.17 C 0.55 0.08 B 0.71 0.12 C 0.39 0.06 CLIST 1 0.46 1.30 0.02 B 0.60 0.01 A 0.83 0.10 C 0.50 0.03 CLIST 2 0.46 1.16 0.03 B 0.53 0.01 A 0.85 0.15 C 0.45 0.03 CLIST 3 0.46 1.12 0.02 B 0.52 0.01 A 0.92 0.14 C 0.48 0.04 CLIST 4 0.46 1.18 0.02 B 0.54 0.01 A 0.71 0.12 C 0.39 0.04 CLITU 1 0.40 1.38 0.05 B 0.55 0.02 A 0.83 0.10 C 0.46 0.03 CLITU 2 0.40 1.11 0.02 B 0.44 0.01 A 0.93 0.03 B 0.41 0.01 ALITU 3 0.40 1.11 0.02 B 0.44 0.01 A 0.92 0.14 C 0.40 0.04 CLITU 4 0.40 1.18 0.02 B 0.47 0.01 A 0.71 0.05 B 0.33 0.03 ALIDE 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CLIDE 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CLIDE 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CLIDE 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CMAPO 1 0.76 1.47 0.31 C 1.12 0.23 B 0.83 0.10 C 0.93 0.26 CMAPO 2 0.76 1.22 0.18 C 0.93 0.14 B 0.85 0.15 C 0.79 0.13 CMAPO 3 0.76 1.11 0.23 C 0.84 0.17 B 0.92 0.14 C 0.77 0.15 CMAPO 4 0.76 1.25 0.17 C 0.95 0.13 B 0.71 0.12 C 0.68 0.13 CMASP 1 0.45 1.47 0.31 C 0.66 0.14 B 0.83 0.10 C 0.55 0.10 CMASP 2 0.45 1.22 0.18 C 0.55 0.08 B 0.85 0.15 C 0.47 0.05 CMASP 3 0.45 1.11 0.23 C 0.50 0.10 B 0.92 0.14 C 0.46 0.06 CMASP 4 0.45 1.25 0.17 C 0.56 0.08 B 0.71 0.12 C 0.40 0.06 CMASP2 1 0.61 1.47 0.31 C 0.90 0.19 B 0.83 0.10 C 0.75 0.17 CMASP2 2 0.61 1.22 0.18 C 0.74 0.11 B 0.85 0.15 C 0.63 0.09 CMASP2 3 0.61 1.11 0.23 C 0.68 0.14 B 0.92 0.14 C 0.62 0.10 CMASP2 4 0.61 1.25 0.17 C 0.76 0.11 B 0.71 0.12 C 0.54 0.09 CMEAZ 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CMEAZ 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 C
65

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeMEAZ 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CMEAZ 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CMOAL 1 0.59 1.47 0.31 C 0.87 0.18 B 0.83 0.10 C 0.72 0.16 CMOAL 2 0.59 1.22 0.18 C 0.72 0.11 B 0.85 0.15 C 0.61 0.09 CMOAL 3 0.59 1.11 0.23 C 0.65 0.13 B 0.92 0.14 C 0.60 0.09 CMOAL 4 0.59 1.25 0.17 C 0.74 0.10 B 0.71 0.12 C 0.53 0.09 CMORU 1 0.59 1.47 0.31 C 0.87 0.18 B 0.83 0.10 C 0.72 0.16 CMORU 2 0.59 1.22 0.18 C 0.72 0.11 B 0.85 0.15 C 0.61 0.09 CMORU 3 0.59 1.11 0.23 C 0.65 0.13 B 0.92 0.14 C 0.60 0.09 CMORU 4 0.59 1.25 0.17 C 0.74 0.10 B 0.71 0.12 C 0.53 0.09 CMOSP 1 0.59 1.47 0.31 C 0.87 0.18 B 0.83 0.10 C 0.72 0.16 CMOSP 2 0.59 1.22 0.18 C 0.72 0.11 B 0.85 0.15 C 0.61 0.09 CMOSP 3 0.59 1.11 0.23 C 0.65 0.13 B 0.92 0.14 C 0.60 0.09 CMOSP 4 0.59 1.25 0.17 C 0.74 0.10 B 0.71 0.12 C 0.53 0.09 CNYAQ 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CNYAQ 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CNYAQ 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CNYAQ 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 CNYOG 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CNYOG 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CNYOG 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CNYOG 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 CNYSY 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CNYSY 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CNYSY 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CNYSY 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 COLTE 1 1.00 1.47 0.31 C 1.47 0.31 B 0.83 0.10 C 1.22 0.46 COLTE 2 1.00 1.22 0.18 C 1.22 0.18 B 0.85 0.15 C 1.04 0.22 COLTE 3 1.00 1.11 0.23 C 1.11 0.23 B 0.92 0.14 C 1.02 0.26 COLTE 4 1.00 1.25 0.17 C 1.25 0.17 B 0.71 0.12 C 0.89 0.22 COSVI 1 0.63 1.47 0.31 C 0.93 0.19 B 0.83 0.10 C 0.77 0.18 COSVI 2 0.63 1.22 0.18 C 0.77 0.12 B 0.85 0.15 C 0.66 0.10 COSVI 3 0.63 1.11 0.23 C 0.70 0.14 B 0.92 0.14 C 0.64 0.10 COSVI 4 0.63 1.25 0.17 C 0.79 0.11 B 0.71 0.12 C 0.56 0.10 COXAR 1 0.50 1.47 0.31 C 0.74 0.15 B 0.83 0.10 C 0.61 0.12 COXAR 2 0.50 1.22 0.18 C 0.61 0.09 B 0.85 0.15 C 0.52 0.06 COXAR 3 0.50 1.11 0.23 C 0.55 0.11 B 0.92 0.14 C 0.50 0.07 COXAR 4 0.50 1.25 0.17 C 0.63 0.09 B 0.71 0.12 C 0.45 0.07 CPATO 1 0.38 1.47 0.31 C 0.56 0.12 B 0.83 0.10 C 0.46 0.07 CPATO 2 0.38 1.22 0.18 C 0.46 0.07 B 0.85 0.15 C 0.39 0.04 CPATO 3 0.38 1.11 0.23 C 0.42 0.09 B 0.92 0.14 C 0.39 0.05 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePATO 4 0.38 1.25 0.17 C 0.48 0.07 B 0.71 0.12 C 0.34 0.05 CPEBO 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CPEBO 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CPEBO 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CPEBO 4 0.51 1.25 0.17 C 0.64 0.09 B 0.71 0.12 C 0.46 0.07 CPIAB 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CPIAB 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CPIAB 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CPIAB 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CPIBR 1 0.33 1.47 0.31 C 0.49 0.10 B 0.82 0.19 C 0.40 0.07 CPIBR 2 0.33 1.22 0.18 C 0.40 0.06 B 1.03 0.13 C 0.41 0.04 CPIBR 3 0.33 1.11 0.23 C 0.37 0.08 B 0.95 0.12 C 0.35 0.04 CPIBR 4 0.33 1.25 0.17 C 0.41 0.06 B 0.80 0.12 C 0.33 0.06 CPIEN 1 0.33 2.09 0.07 B 0.69 0.02 A 0.49 0.05 B 0.34 0.03 BPIEN 2 0.33 1.31 0.02 B 0.43 0.01 A 1.03 0.13 C 0.44 0.03 CPIEN 3 0.33 1.38 0.03 B 0.45 0.01 A 0.95 0.12 C 0.43 0.03 CPIEN 4 0.33 1.52 0.05 B 0.50 0.02 A 0.68 0.06 B 0.34 0.03 BPIGL 1 0.37 1.47 0.31 C 0.54 0.11 B 0.92 0.02 A 0.50 0.05 BPIGL 2 0.37 1.22 0.18 C 0.45 0.07 B 1.20 0.01 A 0.54 0.05 BPIGL 3 0.37 1.11 0.23 C 0.41 0.08 B 1.07 0.01 A 0.44 0.04 BPIGL 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIMA 1 0.38 1.47 0.31 C 0.56 0.12 B 0.92 0.02 A 0.51 0.05 BPIMA 2 0.38 1.22 0.18 C 0.46 0.07 B 1.20 0.01 A 0.55 0.05 BPIMA 3 0.38 1.11 0.23 C 0.42 0.09 B 1.17 0.01 A 0.49 0.05 BPIMA 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CPIPU 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CPIPU 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CPIPU 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CPIPU 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CPIRU 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CPIRU 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CPIRU 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CPIRU 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CPISI 1 0.37 1.70 0.04 B 0.63 0.01 A 0.82 0.19 C 0.52 0.04 CPISI 2 0.37 1.59 0.02 B 0.59 0.01 A 1.03 0.13 C 0.61 0.03 CPISI 3 0.37 1.64 0.02 B 0.61 0.01 A 0.95 0.12 C 0.58 0.03 CPISI 4 0.37 1.63 0.01 B 0.60 0.01 A 0.80 0.12 C 0.48 0.06 CPICEA 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CPICEA 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CPICEA 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CPICEA 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 C
67

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePISP 4 0.39 1.32 0.17 C 0.52 0.07 B 0.80 0.12 C 0.41 0.07 CPIAL 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIAL 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIAL 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIAL 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIAR 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIAR 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIAR 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIAR 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIAR2 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIAR2 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIAR2 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIAR2 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIAT 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIAT 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIAT 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIAT 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIBA 1 0.37 1.50 0.07 B 0.55 0.03 A 0.82 0.19 C 0.45 0.05 CPIBA 2 0.37 1.29 0.18 A 0.48 0.03 A 1.03 0.13 C 0.49 0.03 CPIBA 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIBA 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIBA2 1 0.40 1.47 0.31 C 0.59 0.12 B 0.78 0.02 A 0.46 0.04 BPIBA2 2 0.40 1.22 0.18 C 0.49 0.07 B 0.98 0.01 A 0.48 0.04 BPIBA2 3 0.40 1.11 0.23 C 0.44 0.09 B 1.00 0.01 A 0.44 0.04 BPIBA2 4 0.40 1.25 0.17 C 0.50 0.07 B 0.80 0.12 C 0.40 0.07 CPICL 1 0.46 1.47 0.31 C 0.68 0.14 B 0.82 0.19 C 0.56 0.10 CPICL 2 0.46 1.22 0.18 C 0.56 0.08 B 1.03 0.13 C 0.57 0.05 CPICL 3 0.46 1.11 0.23 C 0.51 0.10 B 0.95 0.12 C 0.49 0.06 CPICL 4 0.46 1.25 0.17 C 0.58 0.08 B 0.80 0.12 C 0.46 0.07 CPICO 1 0.38 1.60 0.02 B 0.61 0.01 A 0.82 0.04 A 0.50 0.05 BPICO 2 0.38 1.23 0.01 B 0.47 0.01 A 1.07 0.03 A 0.51 0.05 BPICO 3 0.38 1.13 0.01 B 0.43 0.00 A 1.02 0.10 B 0.44 0.04 BPICO 4 0.38 1.30 0.01 B 0.49 0.01 A 0.84 0.08 B 0.41 0.04 BPICO3 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPICO3 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPICO3 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPICO3 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIDI 1 0.50 1.47 0.31 C 0.74 0.15 B 0.82 0.19 C 0.61 0.12 CPIDI 2 0.50 1.22 0.18 C 0.61 0.09 B 1.03 0.13 C 0.63 0.06 CPIDI 3 0.50 1.11 0.23 C 0.55 0.11 B 0.95 0.12 C 0.52 0.07 CPIDI 4 0.50 1.25 0.17 C 0.63 0.09 B 0.80 0.12 C 0.50 0.08 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePIEC 1 0.47 1.36 0.05 B 0.64 0.02 A 0.82 0.19 C 0.53 0.05 CPIEC 2 0.47 1.11 0.03 B 0.52 0.01 A 1.03 0.13 C 0.53 0.03 CPIEC 3 0.47 0.92 0.02 B 0.43 0.01 A 0.95 0.12 C 0.41 0.03 CPIEC 4 0.47 1.12 0.03 B 0.53 0.01 A 0.80 0.12 C 0.42 0.06 CPIED 1 0.50 1.47 0.31 C 0.74 0.15 B 0.82 0.19 C 0.61 0.12 CPIED 2 0.50 1.22 0.18 C 0.61 0.09 B 1.03 0.13 C 0.63 0.06 CPIED 3 0.50 1.11 0.23 C 0.55 0.11 B 0.95 0.12 C 0.52 0.07 CPIED 4 0.50 1.25 0.17 C 0.63 0.09 B 0.80 0.12 C 0.50 0.08 CPIEL 1 0.54 1.47 0.31 C 0.79 0.16 B 0.82 0.19 C 0.65 0.13 CPIEL 2 0.54 1.22 0.18 C 0.66 0.10 B 1.03 0.13 C 0.68 0.07 CPIEL 3 0.54 1.11 0.23 C 0.60 0.12 B 0.95 0.12 C 0.57 0.08 CPIEL 4 0.54 1.25 0.17 C 0.68 0.09 B 0.80 0.12 C 0.54 0.08 CPIEN2 1 0.37 1.86 0.06 B 0.69 0.02 A 0.82 0.19 C 0.57 0.05 CPIEN2 2 0.37 1.17 0.02 B 0.43 0.01 A 1.03 0.13 C 0.44 0.03 CPIEN2 3 0.37 1.23 0.02 B 0.45 0.01 A 0.95 0.12 C 0.43 0.03 CPIEN2 4 0.37 1.36 0.05 B 0.50 0.02 A 0.80 0.12 C 0.40 0.06 CPIFL 1 0.37 1.37 0.02 B 0.51 0.01 A 0.82 0.19 C 0.42 0.04 CPIFL 2 0.37 1.17 0.02 B 0.43 0.01 A 1.03 0.13 C 0.44 0.03 CPIFL 3 0.37 1.15 0.01 B 0.43 0.01 A 0.95 0.12 C 0.41 0.03 CPIFL 4 0.37 1.21 0.02 B 0.45 0.01 A 0.80 0.12 C 0.36 0.06 CPIGL2 1 0.41 1.47 0.31 C 0.60 0.13 B 0.82 0.19 C 0.49 0.09 CPIGL2 2 0.41 1.22 0.18 C 0.50 0.08 B 1.03 0.13 C 0.51 0.05 CPIGL2 3 0.41 1.11 0.23 C 0.45 0.09 B 0.95 0.12 C 0.43 0.05 CPIGL2 4 0.41 1.25 0.17 C 0.51 0.07 B 0.80 0.12 C 0.41 0.07 CPIJE 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIJE 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIJE 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIJE 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPILA 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 CPILA 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CPILA 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CPILA 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CPILE 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPILE 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPILE 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPILE 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIMOP 1 0.50 1.47 0.31 C 0.74 0.15 B 0.82 0.19 C 0.61 0.12 CPIMOP 2 0.50 1.22 0.18 C 0.61 0.09 B 1.03 0.13 C 0.63 0.06 CPIMOP 3 0.50 1.11 0.23 C 0.55 0.11 B 0.95 0.12 C 0.52 0.07 CPIMOP 4 0.50 1.25 0.17 C 0.63 0.09 B 0.80 0.12 C 0.50 0.08 CPIMO 1 0.35 1.47 0.31 C 0.51 0.11 B 0.82 0.19 C 0.42 0.07 C
69

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePIMO 2 0.35 1.22 0.18 C 0.43 0.06 B 1.03 0.13 C 0.44 0.04 CPIMO 3 0.35 1.11 0.23 C 0.39 0.08 B 0.95 0.12 C 0.37 0.04 CPIMO 4 0.35 1.25 0.17 C 0.44 0.06 B 0.80 0.12 C 0.35 0.06 CPIMU 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIMU 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIMU 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIMU 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPINI 1 0.41 1.47 0.31 C 0.60 0.13 B 0.82 0.19 C 0.49 0.09 CPINI 2 0.41 1.22 0.18 C 0.50 0.08 B 1.03 0.13 C 0.51 0.05 CPINI 3 0.41 1.11 0.23 C 0.45 0.09 B 0.95 0.12 C 0.43 0.05 CPINI 4 0.41 1.25 0.17 C 0.51 0.07 B 0.80 0.12 C 0.41 0.07 CPIPA 1 0.54 1.47 0.31 C 0.79 0.16 B 0.82 0.19 C 0.65 0.13 CPIPA 2 0.54 1.22 0.18 C 0.66 0.10 B 1.03 0.13 C 0.68 0.07 CPIPA 3 0.54 1.11 0.23 C 0.60 0.12 B 0.95 0.12 C 0.57 0.08 CPIPA 4 0.54 1.25 0.17 C 0.68 0.09 B 0.80 0.12 C 0.54 0.08 CPIPO 1 0.38 1.44 0.02 B 0.55 0.01 A 0.89 0.11 A 0.49 0.05 BPIPO 2 0.38 1.26 0.02 A 0.48 0.01 A 0.95 0.03 A 0.46 0.03 APIPO 3 0.38 1.08 0.03 A 0.41 0.01 A 0.82 0.01 A 0.34 0.02 APIPO 4 0.38 1.28 0.01 B 0.49 0.00 A 1.04 0.10 B 0.51 0.05 BPIPU2 1 0.49 1.47 0.31 C 0.72 0.15 B 0.82 0.19 C 0.59 0.12 CPIPU2 2 0.49 1.22 0.18 C 0.60 0.09 B 1.03 0.13 C 0.62 0.06 CPIPU2 3 0.49 1.11 0.23 C 0.54 0.11 B 0.95 0.12 C 0.51 0.07 CPIPU2 4 0.49 1.25 0.17 C 0.61 0.08 B 0.80 0.12 C 0.49 0.08 CPIRA 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPIRA 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPIRA 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPIRA 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPIRE 1 0.41 1.48 0.04 B 0.61 0.02 A 0.82 0.19 C 0.50 0.05 CPIRE 2 0.41 1.23 0.05 A 0.50 0.01 A 0.82 0.03 A 0.41 0.02 APIRE 3 0.41 1.11 0.23 C 0.45 0.09 B 0.85 0.02 A 0.38 0.06 APIRE 4 0.41 1.25 0.17 C 0.51 0.07 B 0.73 0.06 A 0.37 0.02 APIRI 1 0.47 1.47 0.31 C 0.69 0.14 B 0.82 0.19 C 0.57 0.11 CPIRI 2 0.47 1.22 0.18 C 0.57 0.09 B 1.03 0.13 C 0.59 0.06 CPIRI 3 0.47 1.11 0.23 C 0.52 0.11 B 0.95 0.12 C 0.50 0.07 CPIRI 4 0.47 1.25 0.17 C 0.59 0.08 B 0.80 0.12 C 0.47 0.08 CPISA 1 0.37 1.47 0.31 C 0.54 0.11 B 0.82 0.19 C 0.44 0.07 CPISA 2 0.37 1.22 0.18 C 0.45 0.07 B 1.03 0.13 C 0.46 0.04 CPISA 3 0.37 1.11 0.23 C 0.41 0.08 B 0.95 0.12 C 0.39 0.05 CPISA 4 0.37 1.25 0.17 C 0.46 0.06 B 0.80 0.12 C 0.37 0.06 CPISE 1 0.51 1.47 0.31 C 0.75 0.16 B 0.82 0.19 C 0.62 0.13 CPISE 2 0.51 1.22 0.18 C 0.62 0.09 B 1.03 0.13 C 0.64 0.06 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePISE 3 0.51 1.11 0.23 C 0.56 0.12 B 0.95 0.12 C 0.53 0.07 CPISE 4 0.51 1.25 0.17 C 0.64 0.09 B 0.80 0.12 C 0.51 0.08 CPIST2 1 0.35 1.47 0.31 C 0.51 0.11 B 0.82 0.19 C 0.42 0.07 CPIST2 2 0.35 1.22 0.18 C 0.43 0.06 B 1.03 0.13 C 0.44 0.04 CPIST2 3 0.35 1.11 0.23 C 0.39 0.08 B 0.95 0.12 C 0.37 0.04 CPIST2 4 0.35 1.25 0.17 C 0.44 0.06 B 0.80 0.12 C 0.35 0.06 CPIST 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 CPIST 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CPIST 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CPIST 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CPISY 1 0.41 1.47 0.31 C 0.60 0.13 B 0.82 0.19 C 0.49 0.09 CPISY 2 0.41 1.22 0.18 C 0.50 0.08 B 1.03 0.13 C 0.51 0.05 CPISY 3 0.41 1.11 0.23 C 0.45 0.09 B 0.95 0.12 C 0.43 0.05 CPISY 4 0.41 1.25 0.17 C 0.51 0.07 B 0.80 0.12 C 0.41 0.07 CPITA 1 0.47 1.07 0.03 B 0.51 0.02 A 0.82 0.19 C 0.42 0.05 CPITA 2 0.47 1.04 0.03 B 0.49 0.01 A 1.03 0.13 C 0.50 0.03 CPITA 3 0.47 1.00 0.01 B 0.47 0.00 A 0.95 0.12 C 0.45 0.03 CPITA 4 0.47 1.04 0.02 B 0.49 0.01 A 0.80 0.12 C 0.39 0.06 CPIVI 1 0.45 1.47 0.31 C 0.66 0.14 B 0.82 0.19 C 0.54 0.10 CPIVI 2 0.45 1.22 0.18 C 0.55 0.08 B 1.03 0.13 C 0.56 0.05 CPIVI 3 0.45 1.11 0.23 C 0.50 0.10 B 0.95 0.12 C 0.48 0.06 CPIVI 4 0.45 1.25 0.17 C 0.56 0.08 B 0.80 0.12 C 0.45 0.07 CPLAQ 1 0.53 1.47 0.31 C 0.78 0.16 B 0.83 0.10 C 0.65 0.13 CPLAQ 2 0.53 1.22 0.18 C 0.65 0.10 B 0.85 0.15 C 0.55 0.07 CPLAQ 3 0.53 1.11 0.23 C 0.59 0.12 B 0.92 0.14 C 0.54 0.08 CPLAQ 4 0.53 1.25 0.17 C 0.66 0.09 B 0.71 0.12 C 0.47 0.07 CPLOC 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CPLOC 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CPLOC 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CPLOC 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 CPOAL 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CPOAL 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CPOAL 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CPOAL 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CPOAN 1 0.34 1.47 0.31 C 0.50 0.10 B 0.83 0.10 C 0.41 0.06 CPOAN 2 0.34 1.22 0.18 C 0.41 0.06 B 0.85 0.15 C 0.35 0.04 CPOAN 3 0.34 1.11 0.23 C 0.38 0.08 B 0.92 0.14 C 0.35 0.05 CPOAN 4 0.34 1.25 0.17 C 0.43 0.06 B 0.71 0.12 C 0.31 0.05 CPOBA 1 0.31 1.47 0.31 C 0.46 0.09 B 0.98 0.03 A 0.45 0.04 BPOBA 2 0.31 1.22 0.18 C 0.38 0.06 B 1.04 0.01 A 0.39 0.04 BPOBA 3 0.31 1.11 0.23 C 0.34 0.07 B 1.04 0.01 A 0.36 0.03 B
71

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePOBA 4 0.31 1.25 0.17 C 0.39 0.05 B 0.71 0.12 C 0.28 0.05 CPODE 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CPODE 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CPODE 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CPODE 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CPOFR 1 0.34 1.47 0.31 C 0.50 0.10 B 0.83 0.10 C 0.41 0.06 CPOFR 2 0.34 1.22 0.18 C 0.41 0.06 B 0.85 0.15 C 0.35 0.04 CPOFR 3 0.34 1.11 0.23 C 0.38 0.08 B 0.92 0.14 C 0.35 0.05 CPOFR 4 0.34 1.25 0.17 C 0.43 0.06 B 0.71 0.12 C 0.31 0.05 CPOGR 1 0.36 1.47 0.31 C 0.53 0.11 B 0.83 0.10 C 0.44 0.07 CPOGR 2 0.36 1.22 0.18 C 0.44 0.07 B 0.85 0.15 C 0.38 0.04 CPOGR 3 0.36 1.11 0.23 C 0.40 0.08 B 0.92 0.14 C 0.37 0.05 CPOGR 4 0.36 1.25 0.17 C 0.45 0.06 B 0.71 0.12 C 0.32 0.05 CPOHE 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CPOHE 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CPOHE 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CPOHE 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CPOSA 1 0.37 1.47 0.31 C 0.54 0.11 B 0.83 0.10 C 0.45 0.07 CPOSA 2 0.37 1.22 0.18 C 0.45 0.07 B 0.85 0.15 C 0.38 0.04 CPOSA 3 0.37 1.11 0.23 C 0.41 0.08 B 0.92 0.14 C 0.38 0.05 CPOSA 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CPOSP 1 0.37 1.47 0.31 C 0.54 0.11 B 0.89 0.08 B 0.48 0.04 BPOSP 2 0.37 1.22 0.18 C 0.45 0.07 B 0.92 0.02 A 0.41 0.04 BPOSP 3 0.37 1.11 0.23 C 0.41 0.08 B 0.93 0.09 B 0.38 0.04 BPOSP 4 0.37 1.25 0.17 C 0.46 0.06 B 0.71 0.12 C 0.33 0.05 CPOTR 1 0.35 1.60 0.07 A 0.56 0.02 A 0.88 0.01 A 0.49 0.05 BPOTR 2 0.35 1.27 0.05 A 0.45 0.02 A 0.91 0.01 A 0.41 0.04 BPOTR 3 0.35 1.08 0.01 B 0.38 0.00 A 1.01 0.01 A 0.38 0.04 BPOTR 4 0.35 1.22 0.01 B 0.43 0.00 A 0.71 0.12 C 0.31 0.04 CPRSP 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CPRSP 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CPRSP 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CPRSP 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CPRAM 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CPRAM 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CPRAM 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CPRAM 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CPRNI 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CPRNI 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CPRNI 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CPRNI 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codePRPE 1 0.36 1.47 0.31 C 0.53 0.11 B 0.83 0.10 C 0.44 0.07 CPRPE 2 0.36 1.22 0.18 C 0.44 0.07 B 0.85 0.15 C 0.38 0.04 CPRPE 3 0.36 1.11 0.23 C 0.40 0.08 B 0.92 0.14 C 0.37 0.05 CPRPE 4 0.36 1.25 0.17 C 0.45 0.06 B 0.71 0.12 C 0.32 0.05 CPRSE 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CPRSE 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CPRSE 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CPRSE 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CPRSP2 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CPRSP2 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CPRSP2 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CPRSP2 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CPRVI 1 0.36 1.47 0.31 C 0.53 0.11 B 0.83 0.10 C 0.44 0.07 CPRVI 2 0.36 1.22 0.18 C 0.44 0.07 B 0.85 0.15 C 0.38 0.04 CPRVI 3 0.36 1.11 0.23 C 0.40 0.08 B 0.92 0.14 C 0.37 0.05 CPRVI 4 0.36 1.25 0.17 C 0.45 0.06 B 0.71 0.12 C 0.32 0.05 CPSME 1 0.45 1.41 0.02 B 0.64 0.01 A 0.90 0.02 A 0.57 0.05 BPSME 2 0.45 1.20 0.03 A 0.54 0.01 A 0.99 0.03 A 0.54 0.04 APSME 3 0.45 1.05 0.03 A 0.47 0.01 A 0.80 0.02 A 0.38 0.02 APSME 4 0.45 1.26 0.01 B 0.57 0.00 A 0.75 0.07 B 0.43 0.04 BQUAG 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUAG 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUAG 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUAG 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUAL 1 0.60 1.25 0.02 B 0.75 0.01 A 0.83 0.10 C 0.62 0.03 CQUAL 2 0.60 1.11 0.01 B 0.66 0.01 A 0.85 0.15 C 0.56 0.03 CQUAL 3 0.60 0.96 0.01 B 0.57 0.01 A 0.92 0.14 C 0.52 0.04 CQUAL 4 0.60 1.10 0.01 B 0.66 0.01 A 0.71 0.12 C 0.47 0.04 CQUAR 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUAR 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUAR 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUAR 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUBI 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CQUBI 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CQUBI 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CQUBI 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CQUCH 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUCH 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUCH 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUCH 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUCO 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 C
73

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeQUCO 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CQUCO 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CQUCO 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CQUDO 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CQUDO 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CQUDO 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CQUDO 4 0.51 1.25 0.17 C 0.64 0.09 B 0.71 0.12 C 0.46 0.07 CQUDU 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CQUDU 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CQUDU 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CQUDU 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CQUEL 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUEL 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUEL 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUEL 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUEM 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUEM 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUEM 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUEM 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUEN 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUEN 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUEN 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUEN 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUFA 1 0.52 1.47 0.31 C 0.76 0.16 B 0.83 0.10 C 0.63 0.13 CQUFA 2 0.52 1.22 0.18 C 0.63 0.10 B 0.85 0.15 C 0.54 0.07 CQUFA 3 0.52 1.11 0.23 C 0.58 0.12 B 0.92 0.14 C 0.53 0.08 CQUFA 4 0.52 1.25 0.17 C 0.65 0.09 B 0.71 0.12 C 0.46 0.07 CQUFA2 1 0.61 1.47 0.31 C 0.90 0.19 B 0.83 0.10 C 0.75 0.17 CQUFA2 2 0.61 1.22 0.18 C 0.74 0.11 B 0.85 0.15 C 0.63 0.09 CQUFA2 3 0.61 1.11 0.23 C 0.68 0.14 B 0.92 0.14 C 0.62 0.10 CQUFA2 4 0.61 1.25 0.17 C 0.76 0.11 B 0.71 0.12 C 0.54 0.09 CQUGA2 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CQUGA2 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CQUGA2 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CQUGA2 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CQUGA 1 0.64 1.31 0.04 B 0.84 0.03 A 0.83 0.10 C 0.70 0.04 CQUGA 2 0.64 1.07 0.02 B 0.69 0.01 A 0.85 0.15 C 0.59 0.03 CQUGA 3 0.64 0.93 0.02 B 0.59 0.01 A 0.92 0.14 C 0.54 0.04 CQUGA 4 0.64 1.09 0.02 B 0.70 0.01 A 0.71 0.12 C 0.50 0.04 CQUHY 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUHY 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 C

(Appendix 4 continued on next page)
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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeQUHY 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUHY 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUIL 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUIL 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUIL 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUIL 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUIM 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUIM 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUIM 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUIM 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUIN 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUIN 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUIN 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUIN 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUKE 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CQUKE 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CQUKE 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CQUKE 4 0.51 1.25 0.17 C 0.64 0.09 B 0.71 0.12 C 0.46 0.07 CQULA 1 0.52 1.47 0.31 C 0.76 0.16 B 0.83 0.10 C 0.63 0.13 CQULA 2 0.52 1.22 0.18 C 0.63 0.10 B 0.85 0.15 C 0.54 0.07 CQULA 3 0.52 1.11 0.23 C 0.58 0.12 B 0.92 0.14 C 0.53 0.08 CQULA 4 0.52 1.25 0.17 C 0.65 0.09 B 0.71 0.12 C 0.46 0.07 CQULA2 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQULA2 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQULA2 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQULA2 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQULO 1 0.64 1.47 0.31 C 0.94 0.20 B 0.83 0.10 C 0.78 0.19 CQULO 2 0.64 1.22 0.18 C 0.78 0.12 B 0.85 0.15 C 0.66 0.10 CQULO 3 0.64 1.11 0.23 C 0.71 0.15 B 0.92 0.14 C 0.65 0.11 CQULO 4 0.64 1.25 0.17 C 0.80 0.11 B 0.71 0.12 C 0.57 0.10 CQULY 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CQULY 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CQULY 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CQULY 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CQUMA 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CQUMA 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CQUMA 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CQUMA 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CQUMA2 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUMA2 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUMA2 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 C
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Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeQUMA2 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUMI 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CQUMI 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CQUMI 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CQUMI 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CQUMU 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CQUMU 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CQUMU 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CQUMU 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CQUNI 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUNI 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUNI 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUNI 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUNU 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUNU 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUNU 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUNU 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUOB 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUOB 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUOB 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUOB 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CQUPA 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CQUPA 2 0.58 1.22 0.18 C 0.71 0.11 B 0.85 0.15 C 0.61 0.08 CQUPA 3 0.58 1.11 0.23 C 0.64 0.13 B 0.92 0.14 C 0.59 0.09 CQUPA 4 0.58 1.25 0.17 C 0.73 0.10 B 0.71 0.12 C 0.52 0.08 CQUPH 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUPH 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUPH 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUPH 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUPR 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CQUPR 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CQUPR 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CQUPR 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CQURU 1 0.56 1.29 0.03 B 0.72 0.01 A 0.83 0.10 C 0.60 0.03 CQURU 2 0.56 1.14 0.03 B 0.64 0.02 A 0.85 0.15 C 0.55 0.03 CQURU 3 0.56 1.10 0.02 B 0.61 0.01 A 0.92 0.14 C 0.56 0.04 CQURU 4 0.56 1.16 0.02 B 0.65 0.01 A 0.71 0.12 C 0.46 0.04 CQUSH 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUSH 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUSH 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUSH 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 C

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(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeQUSP 1 0.58 1.47 0.31 C 0.85 0.18 B 0.83 0.10 C 0.70 0.16 CQUSP 2 0.58 1.22 0.18 C 0.71 0.11 B 0.61 0.04 A 0.43 0.02 AQUSP 3 0.58 1.11 0.23 C 0.64 0.13 B 0.59 0.07 A 0.38 0.04 AQUSP 4 0.58 1.25 0.17 C 0.73 0.10 B 0.56 0.04 A 0.41 0.02 AQUST 1 0.60 1.47 0.31 C 0.88 0.18 B 0.83 0.10 C 0.73 0.16 CQUST 2 0.60 1.22 0.18 C 0.73 0.11 B 0.85 0.15 C 0.62 0.09 CQUST 3 0.60 1.11 0.23 C 0.66 0.14 B 0.92 0.14 C 0.61 0.10 CQUST 4 0.60 1.25 0.17 C 0.75 0.10 B 0.71 0.12 C 0.54 0.09 CQUVE 1 0.56 1.47 0.31 C 0.82 0.17 B 0.83 0.10 C 0.68 0.14 CQUVE 2 0.56 1.22 0.18 C 0.68 0.10 B 0.85 0.15 C 0.58 0.07 CQUVE 3 0.56 1.11 0.23 C 0.62 0.13 B 0.92 0.14 C 0.57 0.09 CQUVE 4 0.56 1.25 0.17 C 0.70 0.10 B 0.71 0.12 C 0.50 0.08 CQUVI 1 0.80 1.47 0.31 C 1.18 0.24 B 0.83 0.10 C 0.98 0.28 CQUVI 2 0.80 1.22 0.18 C 0.97 0.15 B 0.85 0.15 C 0.83 0.15 CQUVI 3 0.80 1.11 0.23 C 0.89 0.18 B 0.92 0.14 C 0.82 0.16 CQUVI 4 0.80 1.25 0.17 C 1.00 0.14 B 0.71 0.12 C 0.71 0.15 CQUWI 1 0.70 1.47 0.31 C 1.03 0.21 B 0.83 0.10 C 0.85 0.22 CQUWI 2 0.70 1.22 0.18 C 0.85 0.13 B 0.85 0.15 C 0.72 0.11 CQUWI 3 0.70 1.11 0.23 C 0.78 0.16 B 0.92 0.14 C 0.72 0.13 CQUWI 4 0.70 1.25 0.17 C 0.88 0.12 B 0.71 0.12 C 0.63 0.11 CRONE 1 0.66 1.47 0.31 C 0.97 0.20 B 0.83 0.10 C 0.80 0.20 CRONE 2 0.66 1.22 0.18 C 0.80 0.12 B 0.85 0.15 C 0.68 0.10 CRONE 3 0.66 1.11 0.23 C 0.73 0.15 B 0.92 0.14 C 0.67 0.11 CRONE 4 0.66 1.25 0.17 C 0.83 0.11 B 0.71 0.12 C 0.59 0.10 CROPS 1 0.66 1.47 0.31 C 0.97 0.20 B 0.83 0.10 C 0.80 0.20 CROPS 2 0.66 1.22 0.18 C 0.80 0.12 B 0.85 0.15 C 0.68 0.10 CROPS 3 0.66 1.11 0.23 C 0.73 0.15 B 0.92 0.14 C 0.67 0.11 CROPS 4 0.66 1.25 0.17 C 0.83 0.11 B 0.71 0.12 C 0.59 0.10 CSASP2 1 0.36 1.47 0.31 C 0.53 0.11 B 0.83 0.10 C 0.44 0.07 CSASP2 2 0.36 1.22 0.18 C 0.44 0.07 B 0.85 0.15 C 0.38 0.04 CSASP2 3 0.36 1.11 0.23 C 0.40 0.08 B 0.92 0.14 C 0.37 0.05 CSASP2 4 0.36 1.25 0.17 C 0.45 0.06 B 0.71 0.12 C 0.32 0.05 CSASE 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CSASE 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CSASE 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CSASE 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 CSAAL 1 0.42 1.24 0.06 B 0.52 0.03 A 0.83 0.10 C 0.43 0.03 CSAAL 2 0.42 1.19 0.03 B 0.50 0.01 A 0.85 0.15 C 0.43 0.03 CSAAL 3 0.42 0.91 0.02 B 0.38 0.01 A 0.92 0.14 C 0.35 0.04 CSAAL 4 0.42 1.11 0.03 B 0.47 0.01 A 0.71 0.12 C 0.34 0.04 CSESE 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 C
77

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeSESE 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CSESE 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CSESE 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CSEGI 1 0.34 1.47 0.31 C 0.50 0.10 B 0.82 0.19 C 0.41 0.07 CSEGI 2 0.34 1.22 0.18 C 0.41 0.06 B 1.03 0.13 C 0.42 0.04 CSEGI 3 0.34 1.11 0.23 C 0.38 0.08 B 0.95 0.12 C 0.36 0.04 CSEGI 4 0.34 1.25 0.17 C 0.43 0.06 B 0.80 0.12 C 0.34 0.06 CSOFTW 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CSOFTW 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CSOFTW 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CSOFTW 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CSOAM 1 0.42 1.47 0.31 C 0.62 0.13 B 0.83 0.10 C 0.51 0.09 CSOAM 2 0.42 1.22 0.18 C 0.51 0.08 B 0.85 0.15 C 0.43 0.05 CSOAM 3 0.42 1.11 0.23 C 0.47 0.10 B 0.92 0.14 C 0.43 0.06 CSOAM 4 0.42 1.25 0.17 C 0.53 0.07 B 0.71 0.12 C 0.38 0.06 CTASP 1 0.40 1.47 0.31 C 0.59 0.12 B 0.83 0.10 C 0.49 0.08 CTASP 2 0.40 1.22 0.18 C 0.49 0.07 B 0.85 0.15 C 0.42 0.05 CTASP 3 0.40 1.11 0.23 C 0.44 0.09 B 0.92 0.14 C 0.40 0.05 CTASP 4 0.40 1.25 0.17 C 0.50 0.07 B 0.71 0.12 C 0.36 0.06 CTADI 1 0.42 1.47 0.31 C 0.62 0.13 B 0.83 0.10 C 0.51 0.09 CTADI 2 0.42 1.22 0.18 C 0.51 0.08 B 0.85 0.15 C 0.43 0.05 CTADI 3 0.42 1.11 0.23 C 0.47 0.10 B 0.92 0.14 C 0.43 0.06 CTADI 4 0.42 1.25 0.17 C 0.53 0.07 B 0.71 0.12 C 0.38 0.06 CTABR 1 0.60 1.47 0.31 C 0.88 0.18 B 0.82 0.19 C 0.73 0.16 CTABR 2 0.60 1.22 0.18 C 0.73 0.11 B 1.03 0.13 C 0.75 0.08 CTABR 3 0.60 1.11 0.23 C 0.66 0.14 B 0.95 0.12 C 0.63 0.10 CTABR 4 0.60 1.25 0.17 C 0.75 0.10 B 0.80 0.12 C 0.60 0.10 CTHOC 1 0.29 1.47 0.31 C 0.43 0.09 B 0.82 0.19 C 0.35 0.06 CTHOC 2 0.29 1.22 0.18 C 0.35 0.05 B 1.03 0.13 C 0.36 0.03 CTHOC 3 0.29 1.11 0.23 C 0.32 0.07 B 0.95 0.12 C 0.31 0.04 CTHOC 4 0.29 1.25 0.17 C 0.36 0.05 B 0.80 0.12 C 0.29 0.06 CTHPL 1 0.31 1.67 0.02 B 0.52 0.01 A 0.92 0.09 B 0.48 0.05 BTHPL 2 0.31 1.60 0.01 B 0.50 0.00 A 1.03 0.13 C 0.51 0.03 CTHPL 3 0.31 1.19 0.02 B 0.37 0.01 A 0.95 0.12 C 0.35 0.03 CTHPL 4 0.31 1.49 0.01 B 0.46 0.00 A 0.72 0.07 B 0.33 0.03 BTIAM 1 0.32 1.47 0.31 C 0.47 0.10 B 0.83 0.10 C 0.39 0.06 CTIAM 2 0.32 1.22 0.18 C 0.39 0.06 B 0.85 0.15 C 0.33 0.04 CTIAM 3 0.32 1.11 0.23 C 0.35 0.07 B 0.92 0.14 C 0.32 0.04 CTIAM 4 0.32 1.25 0.17 C 0.40 0.06 B 0.71 0.12 C 0.29 0.05 CTIHE 1 0.32 1.47 0.31 C 0.47 0.10 B 0.83 0.10 C 0.39 0.06 CTIHE 2 0.32 1.22 0.18 C 0.39 0.06 B 0.85 0.15 C 0.33 0.04 C

(Appendix 4 continued on next page)
78

(Appendix 4 continued)

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeTIHE 3 0.32 1.11 0.23 C 0.35 0.07 B 0.92 0.14 C 0.32 0.04 CTIHE 4 0.32 1.25 0.17 C 0.40 0.06 B 0.71 0.12 C 0.29 0.05 CTISP 1 0.32 1.47 0.31 C 0.47 0.10 B 0.83 0.10 C 0.39 0.06 CTISP 2 0.32 1.22 0.18 C 0.39 0.06 B 0.85 0.15 C 0.33 0.04 CTISP 3 0.32 1.11 0.23 C 0.35 0.07 B 0.92 0.14 C 0.32 0.04 CTISP 4 0.32 1.25 0.17 C 0.40 0.06 B 0.71 0.12 C 0.29 0.05 CTOCA 1 0.34 1.47 0.31 C 0.50 0.10 B 0.83 0.10 C 0.41 0.06 CTOCA 2 0.34 1.22 0.18 C 0.41 0.06 B 0.85 0.15 C 0.35 0.04 CTOCA 3 0.34 1.11 0.23 C 0.38 0.08 B 0.92 0.14 C 0.35 0.05 CTOCA 4 0.34 1.25 0.17 C 0.43 0.06 B 0.71 0.12 C 0.31 0.05 CTSHE 1 0.42 1.46 0.01 B 0.61 0.01 A 0.82 0.19 C 0.50 0.04 CTSHE 2 0.42 1.38 0.03 A 0.58 0.01 A 0.95 0.01 A 0.55 0.01 ATSHE 3 0.42 1.13 0.02 A 0.48 0.01 A 0.80 0.04 A 0.39 0.01 ATSHE 4 0.42 1.37 0.01 B 0.58 0.00 A 0.80 0.12 C 0.46 0.06 CTSME 1 0.42 1.47 0.31 C 0.62 0.13 B 0.82 0.19 C 0.51 0.09 CTSME 2 0.42 1.22 0.18 C 0.51 0.08 B 1.03 0.13 C 0.52 0.05 CTSME 3 0.42 1.11 0.23 C 0.47 0.10 B 0.95 0.12 C 0.45 0.06 CTSME 4 0.42 1.25 0.17 C 0.53 0.07 B 0.80 0.12 C 0.42 0.07 CTSSP 1 0.38 1.47 0.31 C 0.56 0.12 B 0.82 0.19 C 0.46 0.08 CTSSP 2 0.38 1.22 0.18 C 0.46 0.07 B 1.03 0.13 C 0.47 0.04 CTSSP 3 0.38 1.11 0.23 C 0.42 0.09 B 0.95 0.12 C 0.40 0.05 CTSSP 4 0.38 1.25 0.17 C 0.48 0.07 B 0.80 0.12 C 0.38 0.07 CULAL 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CULAL 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CULAL 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CULAL 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CULAM 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CULAM 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CULAM 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CULAM 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 CULCR 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CULCR 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CULCR 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CULCR 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CULPU 1 0.46 1.47 0.31 C 0.68 0.14 B 0.83 0.10 C 0.56 0.10 CULPU 2 0.46 1.22 0.18 C 0.56 0.08 B 0.85 0.15 C 0.48 0.05 CULPU 3 0.46 1.11 0.23 C 0.51 0.10 B 0.92 0.14 C 0.47 0.06 CULPU 4 0.46 1.25 0.17 C 0.58 0.08 B 0.71 0.12 C 0.41 0.06 CULRU 1 0.48 1.47 0.31 C 0.71 0.15 B 0.83 0.10 C 0.59 0.11 CULRU 2 0.48 1.22 0.18 C 0.58 0.09 B 0.85 0.15 C 0.49 0.06 CULRU 3 0.48 1.11 0.23 C 0.53 0.11 B 0.92 0.14 C 0.49 0.07 C
79

Species Size Bole init Branch- Uncert Uncert_ FWD Uncert Uncert Relative Uncert Uncert Decayed Uncert Uncert bole B-B bb ratio initial FWD FWD Rel Rel Den Decayed Decayedcode class density ratio ratio code density density code density density code density density codeULRU 4 0.48 1.25 0.17 C 0.60 0.08 B 0.71 0.12 C 0.43 0.06 CULSE 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CULSE 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CULSE 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CULSE 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CULSP 1 0.50 1.47 0.31 C 0.74 0.15 B 0.83 0.10 C 0.61 0.12 CULSP 2 0.50 1.22 0.18 C 0.61 0.09 B 0.85 0.15 C 0.52 0.06 CULSP 3 0.50 1.11 0.23 C 0.55 0.11 B 0.92 0.14 C 0.50 0.07 CULSP 4 0.50 1.25 0.17 C 0.63 0.09 B 0.71 0.12 C 0.45 0.07 CULTH 1 0.57 1.47 0.31 C 0.84 0.17 B 0.83 0.10 C 0.70 0.15 CULTH 2 0.57 1.22 0.18 C 0.69 0.10 B 0.85 0.15 C 0.59 0.08 CULTH 3 0.57 1.11 0.23 C 0.63 0.13 B 0.92 0.14 C 0.58 0.09 CULTH 4 0.57 1.25 0.17 C 0.71 0.10 B 0.71 0.12 C 0.51 0.08 CUMCA 1 0.51 1.47 0.31 C 0.75 0.16 B 0.83 0.10 C 0.62 0.12 CUMCA 2 0.51 1.22 0.18 C 0.62 0.09 B 0.85 0.15 C 0.53 0.06 CUMCA 3 0.51 1.11 0.23 C 0.56 0.12 B 0.92 0.14 C 0.51 0.08 CUMCA 4 0.51 1.25 0.17 C 0.64 0.09 B 0.71 0.12 C 0.46 0.07 CVAAR 1 0.47 1.47 0.31 C 0.69 0.14 B 0.83 0.10 C 0.57 0.10 CVAAR 2 0.47 1.22 0.18 C 0.57 0.09 B 0.85 0.15 C 0.49 0.06 CVAAR 3 0.47 1.11 0.23 C 0.52 0.11 B 0.92 0.14 C 0.48 0.07 CVAAR 4 0.47 1.25 0.17 C 0.59 0.08 B 0.71 0.12 C 0.42 0.06 C

(Appendix 4 continued on next page)
80FWD Absolute and Relative Density Metadata (for Appendix 4)

Fieldname Defi nition Code Units/explanationspecies_code genus-species code Refer to Appendices 2 and 3Size class size code 1 0-1 cm size code 2 1-5 cm size code 3 5-12 cmBole init density bole initial density g/cm3Branch bole ratio branch to bole ratio uncert_bb ratio uncertainty of branch to bole ratio uncert_bbr_code uncertainty code for bbr A sampled green branch density with standard error B sampled green branch density no standard error uncertainty code for bbr C unsampled green branch densityFWD_init density branch initial density g/cm3Uncert_init density uncertainty of initial density g/cm3uncert_init density code uncertainty code for fwd_iden A species initial branch density sampledUncert FWD init density code uncertainty code for fwd_iden B unsampled green, use BBR to obtain greenRel density fwd relative density DimensionlessUncert rel density uncertainty of fwd relative density DimensionlessUncert rel density code uncertainty code for relative density A decayed density sampled with standard error B decayed density sampled no standard error C decayed density calculated from meanDecayed density FWD decayed density g/cm3Uncert_decayed den uncertainty of decayed density g/cm3

(Appendix 4 continued)

Uncert decay density code uncertainty code for decayed density A species decayed density sampled with standard error B species decayed density sampled no standard error C unsampled decayed
81

APPENDIX 5.—Examples of estimates of the mean and 

uncertainty for various levels in the calculation hierarchy (red 

spruce, Picea rubens) on fi ve FIA sample plots in Maine with 

the assumption that density was not directly measured 

Within a Decay or Size Class for a Species and Site

In plot 407074 there is a total of 18.13 m3

/ha of decay class 3 CWD. 

The estimated mean density is 0.316 Mg/m3 and uncertainty in density is 

0.112 Mg/m3. The mean mass and the uncertainty are therefore:

Mass DC-Sp-S = 18.13m3

/ha*0.316Mg/m3 = 5.73 Mg/ha 

UncertaintyMass DC-Sp-S = 18.13m3

/ha*0.112Mg/m3 = 2.031 Mg/ha

Within a Species and Site

In plot 407074 there were pieces in decay classes 2-4 and none in decay classes 

1 and 5. Because the density of this species was not measured directly, density 

of all decay classes is likely either underestimated or overestimated to the same 

degree and we can assume the uncertainties in these decay classes are positively 

correlated. Therefore, the mean and uncertainty of mass for this plot are:

Mass Sp-S = 0 Mg/ha+0.519 Mg/ha+5.73 Mg/ha+1.39 Mg/ha+0 Mg/ha 

= 7.64 Mg/ha

Uncertainty Mass Sp-S = 0.07 Mg/ha+2.03 Mg/ha+0.31 Mg/ha = 2.41 Mg/ha

The relative uncertainty is therefore 31 percent. 

Within a Species for All Sites 

There are fi ve plots each with its own mass and uncertainty estimate. Assuming 

density in all plots is underestimated or overestimated to the same degree, the 

total mass and uncertainty are:

Mass Sp = 7.64 Mg/ha+15.15 Mg/ha+1.34 Mg/ha+1.45 Mg/ha+1.42 Mg/ha 

= 27.00 Mg/ha

Uncertainty Mass Sp = 2.41 Mg/ha+4.02 Mg/ha+0.21 Mg/ha+3.76 Mg/ha

+0.35 Mg/ha = 10.75 Mg/ha

The relative uncertainty is therefore 39.8 percent. 

(continued on next page)
82

APPENDIX 5 (continued)

Within a Site

The uncertainty in mass for a site combines the mass of all the species at a site. 

Because there is no reason to believe that uncertainty in one species is correlated 

with another, the mean and uncertainty would be:

MassS

 = 7.64 Mg/ha+5.99 Mg/ha = 13.64 Mg/ha

Uncertainty MassS

 = square root (2.412+0.612) = 2.49 Mg/ha

The relative uncertainty is therefore 18 percent.
83

APPENDIX 6.—Examples of estimates of the mean and 

uncertainty for various levels in the calculation hierarchy (red 

spruce, Picea rubens) on fi ve FIA sample plots in Maine with 

the assumption that density was directly measured 

Within a Decay or Size Class for a Species and Site 

In plot 407074 there is a total of 18.13 m3

/ha of decay class 3 CWD. 

The estimated mean density is 0.316 Mg/m3 and uncertainty in density is 

0.112 Mg/m3. The mean mass and the uncertainty are therefore:

Mass DC-Sp-S = 18.13m3

/ha*0.316Mg/m3 = 5.73 Mg/ha 

UncertaintyMass DC-Sp-S = 18.13m3

/ha*0.112Mg/m3 = 2.031 Mg/ha

Within a Species and Site 

In plot 407074 there were pieces in decay classes 2-4 and none in decay classes 

1 and 5). Because the density for all decay classes was measured, the uncertainty 

is related to random sampling error. Therefore, the mean and uncertainty of mass 

for this plot are:

Mass Sp-S = 0 Mg/ha+0.519 Mg/ha +5.73 Mg/ha+ 1.39 Mg/ha+0 Mg/ha 

= 7.64 Mg/ha

Uncertainty Mass Sp-S = Square root (0.072

 +2.032 + 0.312) = 2.06 Mg/ha

The relative uncertainty is therefore 27 percent. 

Within a Species for All Sites 

There are fi ve plots each with its own mass and uncertainty estimate. There is 

no reason to believe the uncertainty in the plots is correlated. Therefore, the total 

mass and uncertainty are:

Mass Sp = 7.64 Mg/ha+15.15 Mg/ha +1.34 Mg/ha+ 1.45 Mg/ha+1.42 Mg/ha 

= 27.00 Mg/ha

Uncertainty Mass Sp = square root (2.062

 +2.892 + 0.132 +0.272 + 0.282) 

= 3.57 Mg/ha

The relative uncertainty is therefore 13 percent. 

(continued on next page)
84

APPENDIX 6 (continued)

Within a Site 

The uncertainty in mass for a site combines the mass of all the species at a site. 

There is no reason to believe that uncertainty in one species is correlated with 

another. Therefore, the mean and uncertainty would be:

MassS = 7.64 Mg/ha + 5.99 Mg/ha = 13.64 Mg/ha

Uncertainty MassS = square root (2.062

+0.282) = 2.08 Mg/ha

The relative uncertainty is therefore 15 percent.
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Printed on recycled paper

Harmon, Mark E.; Woodall, Christopher W.; Fasth, Becky; Sexton, Jay. 2008. 

Woody detritus density and density reduction factors for tree species in 

the United States: a synthesis. Gen. Tech. Rep. NRS-29. Newtown Square, 

PA: U.S. Department of Agriculture, Forest Service, Northern Research 

Station. 84 p.

This report presents a synthesis of published and unpublished data on 

woody detritus density as a step toward improving estimates of coarse woody 

detritus (CWD) and fi ne woody detritus (FWD) biomass across the forests 

of the United States. In the case of CWD, 88 species were found to have 

data on densities for fi ve decay classes that had been published and/or 

collected in North America. For FWD, about 25 species had been sampled. 

We determined that by sampling representative species within a genus, the 

uncertainty of CWD estimates could be reduced by up to 50 percent over not 

having sampled a genus. Our analysis indicated that the uncertainty of FWD 

mass estimation ranged from 12 to 19 percent when FWD relative density 

was estimated. We conclude that a more systematic sampling of CWD and 

FWD density is needed for major species if uncertainties of mass estimates 

nationwide are to be reduced.

KEY WORDS: Dead wood, forest carbon stores, coarse woody debris, fi ne 

woody debris, decay classes, downed woody material.
Capitalizing on the strengths of existing science capacity 

in the Northeast and Midwest to attain a more integrated 

cohesive landscape scale research program