EARTH SURFACE PROCESSES AND LANDFORMS

Earth Surf. Process. Landforms 41, 130–142 (2016)

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
Published online 4 November 2015 in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/esp.3831

Tools for gauging the capacity of salmon spawning

substrates
Brandon T. Overstreet,1* Clifford S. Riebe,2* John K. Wooster,3 Leonard S. Sklar4 and Dino Bellugi5
1
Department of Geography, University of Wyoming, Laramie, WY, USA
2
Department of Geology and Geophysics, University of Wyoming, Laramie, WY, USA
3
NOAA – Fisheries, Habitat Conservation Division, Santa Rosa, CA, USA
4
Department of Earth and Climate Sciences, San Francisco State University, San Francisco, CA, USA
5
Department of Earth, Atmospheric and Planetary Sciences, Massachusetts Institute of Technology, Cambridge, MA, USA

Received 19 January 2015; Revised 29 July 2015; Accepted 18 August 2015

*Correspondence to: Brandon T. Overstreet, Department of Geography, University of Wyoming, Laramie, WY, USA. E-mail: boverstr@uwyo.edu or Clifford S. Riebe,
Department of Geology and Geophysics, University of Wyoming, Laramie, WY, USA. E-mail: criebe@uwyo.edu
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction
in any medium, provided the original work is properly cited and is not used for commercial purposes.

ABSTRACT: We present a set of river management tools based on a recently developed method for estimating the amount of
salmon spawning habitat in coarse-bedded rivers. The method, which was developed from a mechanistic model of redd building
by female salmon, combines empirical relationships between fish length, redd area, and the sizes of particles moved by fish during
spawning. Model inputs are the grain-size indices D50 and D84 and an estimate of female fish length, which is used to predict the size
of the redd that they will build and the size of the largest particle that they can move on the bed. Outputs include predictions of the
fraction of the bed that the fish can use for redd building and the number of redds that they can build within the useable area. We cast
the model into easy-to-use look-up tables, charts, an Excel worksheet, a JavaScript web applet, and a MATLAB user interface. We explain
how these tools can be used in a new, mechanistic approach to assessing spawning substrates and optimizing gravel augmentation
projects in coarse-bedded rivers. © 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
KEYWORDS: river restoration; Pacific salmon; salmon spawning habitat; fluvial geomorphology; aquatic ecology

Introduction
Salmon populations are in decline throughout much of their
historical range, reflecting natural and human-induced changes
in aquatic food webs and habitats, including degradation of
riverbed sediments where salmon spawn (e.g. Nehlsen et al.,
1991; Yoshiyama et al., 1998; Carlson and Satterthwaite, 2011).
To reverse losses in salmon populations, and thus maintain the
economic, cultural, and ecosystem services that salmon provide,
river managers spend millions of dollars every year on restoring
salmon spawning habitat (e.g. Kondolf et al., 2007). For example,
in 2013 alone, the United States Bureau of Reclamation
budgeted over $600,000 (USD) for improving spawning habitat
as part of the Central Valley Project Improvement Act (USBR
and USFWS, 2014), which covers just a few of the US rivers
targeted for habitat restoration that year. Despite expenditures
such as these, the benefits of restoration have remained unclear,
in part because of poor communication between scientists and
managers (Wohl et al., 2005; Bernhardt et al., 2007). To over-
come this limitation, scientists must develop tools that address
the questions that managers ask when designing and monitoring
restoration projects (Bernhardt et al., 2005, 2007; Wohl et al.,
2005; Beechie and Snover, 2014). Moreover, for the science to
confer benefits to restoration, the tools that are developed must
be both accessible to restoration managers and practical to use in
field settings (Bernhardt et al., 2007; Beechie and Snover, 2014).
In a recent study, we developed a new understanding of how
the amount of salmon spawning habitat in coarse riverbeds
depends on grain size and fish length (Riebe et al., 2014). These
findings, which are based on empirical relationships between
fish length, redd area, and the size of the largest particles that
fish can move, provide a scientific basis for designing spawning
substrate restoration projects and monitoring their benefits over
time. However, to be useful in restoration, our findings must
first be translated into practical tools and made available to
the managers who might use them.
Here we present a set of tools that managers can readily use to
gauge the amount of salmon spawning habitat in coarse-bedded
rivers. We begin by summarizing the conceptual and mathema-
tical basis of the approach. We then describe how we encoded
the math into a series of easy-to-use look-up charts and tables.
We also present computer-based applications of the approach,
including an Excel worksheet, a JavaScript web applet, and a
MATLAB user interface. This comprehensive set of tools, inclu-
ding charts and software, is publically available in the main
article, in Supplementary Material, and online. We discuss
 TOOLS FOR GAUGING THE CAPACITY OF SALMON SPAWNING SUBSTRATES
how managers can use these tools as part of a new, mechanistic
approach to assessing salmon spawning substrates and to
designing cost-effective spawning habitat restoration projects.
A New Approach to Assessing Salmon
Spawning Substrates
Particle sizes and redd building
Grain size can limit spawning by preventing redd construction if
particles are too big for female fish to move and thus use during
redd building (Kondolf, 1988; Kondolf and Wolman, 1993; Ligon
et al., 1995; Kondolf, 2000; Riebe et al., 2014). Female salmon
construct redds by turning on their sides, swatting at the bed with
their tails, and thus inducing lift forces that excavate particles from
the bed surface (Burner, 1951). In coarse-bedded rivers, some par-
ticles may be too big for fish to move, depending on the magni-
tude of the lift forces that the fish can generate with its tail. If the
bed harbours just a few of these immovable particles, they may
not hinder spawning because salmon may be able to simply build
their redds around them (Quinn et al., 1995). However, as immov-
able particles become more concentrated, the area of the bed that
can accommodate redd building decreases and should be roughly
equal to the total area of the bed minus the area occupied by par-
ticles that are too big to move. This relationship is illustrated con-
ceptually with bed-surface grain-size distributions in Figure 1. For
each of the two distributions in Figure 1B, the shaded area under
the curve corresponds to the fraction of the total bed-surface area
that is covered by particles with diameters bigger than DT, the
Figure 1. Relationship between grain-size distribution, threshold parti-
cle size, and area of the bed covered by particles that are too big for fish
to move. (a) Immovable particles on the bed limit ability to construct redds
by decreasing the amount of area useable for redd building. For each dis-
tribution in (b), the shaded area under the curve is the fractional coverage
by particles bigger than DT, illustrating how usable spawning area
changes with D84. As D84 increases for a given D50, the amount of usable
substrate decreases, because a greater fraction of the bed is covered by
particles with diameters greater than DT. This figure is available in colour
online at wileyonlinelibrary.com/journal/espl
© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
131
threshold size that differentiates what fish can and cannot move.
When the median particle size on the bed (D50) and DT are both
constant, coverage by immovable particles increases with increas-
ing variance in the grain-size distribution (Figure 1B), as reflected
in increasing D84 (i.e. the 84th percentile of grain diameters). As
coverage by immovable particles increases, less of the bed is suit-
able for redd building, leading to a decrease in the number of
redds that salmon can build per unit area of bed. Henceforth,
we use the term “spawning capacity” and the notation NREDDS
to refer to the maximum number of redds per unit area that a sub-
strate can physically accommodate for a species of interest. As we
show in sections that follow, NREDDS depends on the grain-size
distribution of the bed, the size of the largest particle that fish
can move, and the size of the redds they build.
Predicting the fractional coverage of movable
particles
If the size of the largest movable particle is known, the fractional
coverage of the bed by movable particles can be estimated from
the bed-surface grain-size distribution by simply integrating it from
zero to DT, and thus quantifying the non-shaded area under the
curve in Figure 1B (Riebe et al., 2014). Although salmonid
spawning substrates have been studied for decades (Burner,
1951; Chambers et al., 1955; McNeil and Anhell, 1964; Orcutt
et al., 1968; Crisp and Carling, 1989; Bjornn and Reiser, 1991;
Kondolf and Wolman, 1993; Kondolf, 2000), much of the work fo-
cused on understanding how fine sediment degrades salmon
spawning habitat by influencing fluvial scour (May et al., 2009),
preventing delivery of life-sustaining oxygen to eggs (McNeil
and Anhell, 1964; Chapman, 1988; Greig et al., 2005), and
entombing fry (Phillips et al., 1975). Meanwhile, the upper
particle-size limits on redd building in coarse-bedded rivers have
rarely been quantified (Kondolf, 1988; Kondolf and Wolman,
1993). Moreover, the relationship between fish length and DT
had not been quantified until our own recent study of redd build-
ing by Chinook, sockeye, and pink salmon (Oncorhynchus
tshawytscha, O. nerka, and O. gorbuscha), three different-sized
species of Pacific salmon (Riebe et al., 2014). According to data
from 73 redds examined in that work, the intermediate axis diam-
eter of the largest particle moved increased systematically with fish
length (L), in a power-law relationship expressed in Equation (1).
D T ¼ 115ðL=600Þ0:62 (1)
Here, DT and L are both expressed in millimetres, and the constant
in the denominator is a reference fish length (600 mm); when L is
equal to it, DT is equal to the power-law intercept (115 mm).
Equation (1) assumes that the intermediate-axis diameters of
the largest particles moved in redds can be used as a proxy for
DT (Riebe et al., 2014).
When applied at the reach scale (e.g. over hundreds of square
metres of spawning habitat) Equation (1) can be used to divide the
riverbed into the fraction of grains on the surface that fish of a
particular size can move and the fraction they cannot. All that
is needed is an estimate of the bed-surface grain-size distribution,
which can be readily quantified in a grid-based, Wolman-style
pebble count (Wolman, 1954; Bunte and Abt, 2001b). In these
pebble counts, the frequency of a particular-sized particle repre-
sents its fractional coverage on the bed (Wolman, 1954;
Kellerhals and Bray, 1971; Bunte and Abt, 2001b). Hence, the
fractional area of the bed that the fish can move (FM) can be quan-
tified graphically from the cumulative distribution function (CDF)
of grain size by simply reading the value of the CDF that corre-
sponds to DT (Figure 2; after Riebe et al., 2014). If the full grain-
size distribution is not available, but D50 and D84 are known,
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132 B. T. OVERSTREET ET AL.

data in many instances although grain-size measurement

Figure 2. A graphical approach to quantifying the suitability of
spawning substrates. Cumulative distribution functions (CDFs) of grain
size for two lognormal grain-size distributions with equal D50 and differ-
ing D84, illustrating how the fraction of the bed that the fish can move
(FM) can be quantified graphically when DT is known (after Riebe et al.,
2014); FM is equivalent to the fraction finer than DT, which can be read
from the vertical axis at the intersection of DT with the CDF. This figure
is available in colour online at wileyonlinelibrary.com/journal/espl.
methods and data quality should always be carefully assessed.
Moreover, the need for just three readily measured variables
and the fairly simple form of Equation (4) make it easier to apply
than the graphical approach. Nevertheless it is important to
quantify the scale of typical errors introduced by using Equation
(4) instead of the graphical approach. We did so here using
grain-size distributions from the Riebe et al. (2014) calibration
sites, which span a wide range of conditions; D50 ranged from
49 to 118 mm, D84 ranged from 86 to 385 mm, and mean fork
length of spawning female salmon ranged from 445 to 721 mm.
The agreement between predictions from Equation (4) and the
graphical approach is illustrated in Figure 3. All of the data plot
close to the 1:1 line of perfect correspondence between the
analytical and graphical approaches. Residuals relative to the 1:1
line (Figure 3B) show that the model agrees with observations to
within ~7% across a greater than four-fold range in fractional cov-
erage by movable particles. Moreover, the Nash–Sutcliffe statistic
is 0.92, indicating that the variance in the graphically measured
FM values is efficiently explained by the model. Hence, although
none of the grain-size distributions that we examined were strictly
lognormal, our analysis indicates that errors introduced in the as-
sumptions of Equation (4) may often be small enough to ignore.

FM can still be estimated using the analytical approximation

expressed in Equations (2)-(4) (after Riebe et al., 2014). Predicting the spawning capacity of riverbeds

-1 Once FM is quantified, whether by the graphical or analytical

F M ¼ 1þ e1:702z (2) approach, it can be recast in terms of spawning capacity,
defined as the number of redds that salmon can build per unit
Here, FM is the dimensionless fractional area of the bed that fish area and denoted NREDDS (e.g. in redds/100 m2) in Equation (5).
can move (e.g. in m2/m2). The right side of Equation (2) is a
one-parameter logistic function that approximates the cumula- N REDDS ¼ F M =AREDD 100 (5)
tive normal distribution to within 1% (Bowling et al., 2009). Here,
z is analogous to the Z statistic in the parlance of statisticians.
Recognizing that grain-size distributions are often more
lognormal than normal in shape (e.g. Bunte and Abt, 2001b),
we calculate z using Equation (3) (after Riebe et al., 2014).

logðDT =D 50 Þ
z¼ (3)
logðD84 =D 50 Þ

Here log(D50) and log(D84/D50) are the mean and standard

deviation of a lognormal grain-size distribution, and the grain
size indices D84 and D50 are expressed in consistent units of
length (e.g. millimetres). Assuming that the grain-size distribution
is approximately lognormal, Equations (2) and (3) can be used
together to estimate the fraction of the bed covered by grains
smaller than the threshold size DT, as shown in Equation (4) (after
Riebe et al., 2014).

8 2 h i39-1
< log 115ðL=600Þ0:62 =D 50 =
FM ¼ 1 þ exp4-1:702 5 (4)
: log ½D84 =D 50  ;

Here we express DT in terms of Equation (1). If a site-specific

estimate of DT is available then it could be used instead in the
numerator of the exponential term.
Equation (4) is easy to apply when estimates of fish length and
grain size are available, but it assumes that the grain-size distribu-
Figure 3. Comparison between Equation (4) and graphical approach to
tion is lognormal. Hence, it will generally be more accurate to
predicting FM. (top graph) Coverage by movable particles (FM) measured
use the graphical approach (e.g. Figure 2) for estimating FM
using graphical approach (see Figure 2) plotted against coverage by
because it does not require assumptions about the shape of the movable particles predicted from Equation (4) for Chinook salmon (grey
grain-size distribution. However, complete grain-size distribu- circles; L = 721 mm), sockeye salmon (open circles; L = 569 mm), and pink
tions are not often reported in studies of rivers, whereas the salmon (black circles; L = 445 mm) based on data from Riebe et al. (2014).
indices D50 and D84 typically are. Equation (4) should be a Residuals show that model-predicted values agree with graphical estimates
suitable option for assessing spawning substrates from existing to within ~7% across the three species of salmon (bottom graph).

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
 TOOLS FOR GAUGING THE CAPACITY OF SALMON SPAWNING SUBSTRATES 133

Here, AREDD is the area (in m2) of a typical redd constructed by
the species of interest. The factor of 100 is included so that
NREDDS is expressed in units of redds/100 m2, consistent with
the spatial dimensions of typical spawning patches. The value
of AREDD can be estimated from redd dimensions measured in
the field or from Equation (6), which is an empirical relation-
ship between redd area and female salmonid length (Riebe
et al., 2014).
AREDD ¼ 3:3½L=6002:3
Here, the power-law intercept and exponent are regression pa-
rameters based on a best fit to data from redds built by salmonids
spanning a wide range of sizes (after Riebe et al., 2014, based on
data from Crisp and Carling, 1989). As in Equation (1), the de-
nominator on the right side of Equation (6) is a reference value
for the regression; when L is equal to it (i.e. 600 mm), the pre-
dicted redd area is equal to the power-law intercept (i.e. 3.3
m2). Although Equation (6) provides a very good fit to Crisp and
Carling’s (1989) data on Atlantic salmon (Salmo salar) and trout
(S. trutta and Oncorhynchus mykiss), and also to Riebe et al.’s
(2014) data on Chinook, sockeye, and pink salmon (see Riebe
et al., 2014), site-specific values of redd area could be used to
translate values of FM into estimates of substrate spawning capac-
ity via Equation (4).
For a wide variety of species, including both Atlantic and
Pacific salmon, Equation (6) should be an acceptable predictor
of redd area. When it is, Equations (4)-(6) can be combined into
a single expression for estimating spawning capacity.
accommodate when fish are spawning en masse, in conditions
that are not limited by other factors such as flow velocity and
depth (Moir and Pasternack, 2010), and hyporheic flow
through redds (Tonina and Buffington, 2009). This is demon-
strated in Figure 4 for spawning by three Pacific salmon species
across the 31 sites of Riebe et al. (2014). In all but a few cases,
field-surveyed areal coverage by redds was less than or equal to
the fractional coverage by movable particles predicted from
(6) Equation (4) (Figure 4A). Likewise, spawning use, which was
estimated from the areal coverage of redds on the bed, was
less than or equal to spawning capacity, which can be
predicted from Equation (7) (Figure 4B). Data that plot near
the 1:1 line in both Figures 4A and 4B imply instances in
which fish were building redds in nearly all of the movable
substrate. The fact that some of the observations in Figures 4A
and 4B are close to the 1:1 line implies that FM and NREDDS are
indices of the maximum amount of spawning habitat in a
reach when other factors, such as flow conditions and fine
sediment content, are not limiting (Riebe et al., 2014).
Assessing and Monitoring Salmon Spawning
Substrates
Sensitivity to differences in grain size
Equations (4) and (7) show that FM and NREDDS can each be
expressed as a function of just three variables. In applying
them to the assessment of spawning substrates for a particular

    !#)1
2:3 logð115Þþ 0:62logðL=600Þ  logðD 50 Þ
N REDDS ¼ 100 3:3ðL=600Þ 1þ exp 1:702 (7)
logðD 84 Þ  logðD 50 Þ

Like the expression in Equation (4) for FM, Equation (7) for
NREDDS has just three variables: fish length and two grain-size
indices that are widely reported in the literature. Hence,
although Equations (4) and (7) are somewhat complicated in
form, they are both computationally easy to apply to data
available from the literature (Riebe et al., 2014).
Coverage by movable particles and the spawning capacity of
substrates predicted from Equations (4) and (7) are both
potentially useful in river management, because they constrain
the maximum amount of redd building a substrate can
run of salmon, it may be helpful to set fish length equal to the
mean fish length for the run and thus reduce the number of
variables to just two: D84 and D50. One can then explore the
sensitivity of substrate spawning capacity to differences in
grain size alone (e.g. from one reach to the next) by construct-
ing contour plots of FM and NREDDS in a plotting space that
includes D84 and D50. The most straightforward way to display
grain size in a two-dimensional plotting space is to plot the
standard deviation against the mean. Here, in the lognormal
approximations of Equations (4) and (7), this can be done by

Figure 4. (a) Spawning use observed in field versus FM predicted from Equation (4) for three species of salmon. The fractional area used for spawning
should be less than or equal to the fractional area covered by movable substrate (FM) – this is a theoretical cap on spawning use illustrated by the 1:1
line. (b) Spawning use observed in field versus spawning capacity predicted from Equation (7) for three species of salmon (after Riebe et al., 2014).
Error bars show ±1 standard error propagated from measurement and model uncertainties. Points that plot close to the 1:1 lines (a and b) are cases
of fish spawning en masse and building redds in nearly every patch of movable substrate. This shows that model predictions of FM and NREDDS yield
realistic estimates of substrate capacity for accommodating redds in riverbeds.

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
134 B. T. OVERSTREET ET AL.

plotting log(D50) (the mean) on the horizontal axis and log
(D84/D50) (the standard deviation) on the vertical axis.
Because D50 and D84 appear in exponential form in Equations
(4) and (7), it is appropriate to plot them using log scales. This
yields the patterns in FM and NREDDS shown in Figures 5A and
5B. Inspection of these panels shows that, for a given D84/D50
(i.e. for a constant spread in the grain-size distribution), FM and
NREDDS are both more sensitive to D50 when D84/D50 is low
than when D84/D50 is large. In addition, when D50 is less than
DT, increases in D84/D50 correspond to decreases in FM
(Figure 5A) and NREDDS (Figure 5B). This is consistent with
concepts illustrated in Figure 1: when the spread in the distribution
is bigger, more of the distribution lies above the threshold particle
size, such that there is less coverage by movable particles. The
converse is true when D50 is greater than DT, because increases
in the spread in the distribution increase the fraction of the bed
sediment with grain sizes smaller than the threshold particle size
(which means there is more useable substrate for redd building).
Thus, the plots shown in Figures 5A and 5B provide useful tools
for understanding how changes in grain size are likely to
influence the amount of spawning substrate in riverbeds.
Graphical assessment of substrate spawning
capacity
The plots in Figures 5A and 5B illustrate the sensitivity of FM and
NREDDS to differences in grain size for a fish with a length of 600
mm. Another way to show the results in grain-size space is to
simply plot D84 versus D50 (Figures 5C and 5D). This plotting
space is complicated by a ‘forbidden zone’, (the grey space in
Figures 5C and 5D), which arises because D84 can never be less
than D50. Nevertheless, plots like these are easier to use with
data from pebble counts, because they obviate the intermediate
step of calculating D84/D50; one can simply read the value of FM
or NREDDS directly from the plot at the intersection of D84 and
D50 (Figures 5C and 5D). Thus a field worker can readily quan-
tify the amount of spawning habitat in a study reach using a
graphical approach that obviates the somewhat complicated
calculations of Equations (4) and (7). This graphical approach
can be applied to salmon of any size after generating a size-
specific chart for the fish of interest. We illustrate this for salmon
ranging in length from 250 to 950 mm (in 50 mm increments) for
both FM in Figure 6 and NREDDS in Figure 7. The physical limita-
tions on redd building that are encapsulated in Equations (4)
and (7) should apply across all of the fish lengths represented
in Figures 6 and 7, but only Figures 6e–6h lie within the tested
range of the model (Riebe et al., 2014); more work is needed
to verify the model’s applicability to larger and smaller fish.
Applying the charts in Figures 6 and 7 to spawning habitat as-
sessment is straightforward. For example, if the goal is to assess
spawning substrate in a reach with D50 = 100 and D84 = 250
mm for a sockeye salmon with average length of 710 mm, FM
and NREDDS can be read from the intersection of D50 and D84 in
Figures 6j and 7j, which correspond to contour plots of FM and
NREDDS of a 700 mm fish. Because the plots do not correspond
exactly to the fish length of interest, the graphical estimates
(i.e. FM = 0.61 and NREDDS = 12.8 redds/100 m2) are not exact.
In the hypothetical case outlined here, the graphical approach
underestimates FM and overestimates NREDDS by less than 1%
relative to what one would calculate using Equations (4) and
(7). We suggest that errors introduced by rounding fish lengths
to the nearest 50 mm may often be small enough to ignore.

Figure 5. Variations in FM and NREDDS with grain size. Fractional coverage by movable particles (a) and spawning capacity, equal to the number of
2
redds per 100 m (b) plotted as a function of the standard deviation and the median grain size (equal to D84/D50 and D50, respectively, for a lognormal
bed-surface grain-size distribution) for a 600 mm long female salmon. Increases in the spread of the distribution lead to increases or decreases in FM
depending on whether D50 is greater than or less than DT (which is denoted by the bold vertical line in each plot). (c) and (d) recast the functions
shown in (a) and (b) in terms of D84 and D50. In this plotting space, there is a forbidden zone (grey space) encompassing impossible scenarios, in
which D50 > D84. These plots can be used in a graphical approach to assessing spawning substrates; managers can read the predicted value of FM
and NREDDS directly from these plots at the intersection of D84 and D50 for the reach of interest.

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
 TOOLS FOR GAUGING THE CAPACITY OF SALMON SPAWNING SUBSTRATES 135

Figure 6. Variations in FM with grain size for 15 different sizes of fish ranging from 250 mm (a) to 950 mm (o). Grey region encompasses forbidden
zone of impossible scenarios in which D84 is less than D50. These graphs are reproduced in Supplementary Material 1 in a series of full-page charts
that managers can print and use in the field to estimate the fractional coverage by movable particles using the following steps: (i) measure grain-size
distribution of the bed using a pebble count or other approach; (ii) select chart corresponding to fish length of interest; (iii) read FM from the intersec-
tion of D84 and D50. This figure is available in colour online at wileyonlinelibrary.com/journal/espl

Look-up charts and tables
To make the habitat assessment approach of Equations (4) and
(7) accessible to river managers, we have reproduced Figures 6
and 7 in a series of larger look-up charts (sized at one per page)
that could be printed and brought into the field for rapid assess-
ment of spawning substrates. These charts are included in
Supplementary Material 1 and 2, which are available online
and by request from the authors. The charts show contours of
FM and NREDDS, similar to those shown in Figures 6 and 7,
but without the colour, to conform to back-and-white printers
and thus facilitate use in the field. Each chart is specific to a
particular fish length ranging from 250 to 950 mm in
increments of 50 mm from one page to the next. The D50 and
D84 range from 10 to 300 mm and 10 to 500 mm, respectively,
in each chart, and thus cover a wide range of conditions that
might be encountered in the field. Managers can use them by
simply reading off FM and NREDDS at the intersection of a mea-
sured D50 and D84 for a fish of a particular length.
As an alternative to these charts, we have also encoded
Equations (4) and (7) into easy-to-use look-up tables, available
online as Supplementary Material 3 and 4. In the tables, D50
ranges from 20 to 300 mm in 10 mm increments, D84 ranges
from 40 to 500 mm in 20 mm increments, and L ranges from
100 to 1100 mm in 100 mm increments. To use these tables,
one simply finds the intersection of measured values of D50,

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
136 B. T. OVERSTREET ET AL.

Figure 7. Variations in NREDDS with grain size for 15 different sizes of fish ranging from 250 mm (a) to 950 mm (o). Grey region encompasses for-
bidden zone of impossible scenarios in which D84 is less than D50. These graphs are reproduced in Supplementary Material 2 in a series of full-page
charts that managers can print and use in the field to assess the capacity of spawning habitat using the following steps: (i) measure grain-size distri-
bution of the bed using a pebble count or other approach; (ii) select chart corresponding to fish length of interest; (iii) read NREDDS from the intersec-
tion of D84 and D50. This figure is available in colour online at wileyonlinelibrary.com/journal/espl

D84, and L, and reads of the value of FM (Supplementary Mate-
rial 3) or NREDDS (Supplementary Material 4). Values that lie be-
tween these increments can be approximated by interpolation.
With Supplementary Material 1 and 2 or 3 and 4 in hand, the
approach encapsulated in Equations (4) and (7) can be readily
implemented with grain-size data from pebble counts and with
fish lengths from carcass surveys and fish traps.
Computer-based tools for applying the approach
Recognizing that it may not always be convenient to use the
look-up tables or charts, we have also encoded Equations (4)
and (7) into a series of computer-based tools, which are avail-
able in Supplementary Material 5–7 and on the second author’s
website. One of the computer-based tools (Supplementary
Material 5) is an Excel workbook consisting of three worksheets.
Data are entered on the sheet labelled ‘Input data here.’ Model
outputs for the specified fish length and grain sizes are reported
in the sheet labelled ‘Model outputs.’ A third sheet titled ‘Read
me’ provides detailed instructions and an explanation of error
messages. Up to 500 sets of fish length and grain-size data can
be input at a time by copying and pasting input values into the
appropriate columns in the workbook.
We also generated a JavaScript web applet (Supplementary
Material 6), which accepts user-provided data in an easy-to-

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
 TOOLS FOR GAUGING THE CAPACITY OF SALMON SPAWNING SUBSTRATES 137

use graphical user interface (GUI) that can be opened in
JavaScript-enabled browsers (Figure 8). Data are entered in
the panel marked ‘Input’ (at right, under the ‘Data’ heading),
either by typing the values for fish length, D50, and D84, or by
clicking and dragging the slider bars. The input panel automat-
ically keeps the values within a realistic range that is appropri-
ate given the calibration data and ensures that D50 is never
mistakenly given a value greater than D84. When values are
changed, the button marked ‘Update Model’ must be clicked
to generate results for the specified fish length and grain size in-
dices in the panel marked ‘Output’. This also updates the con-
tour plots of fractional coverage by movable particles and
spawning capacity, which can be viewed by clicking the tables
under the ‘Plots’ heading in Figure 8. The bold red data point in
each plot corresponds to the specified D84 and D50. Inputs can
be reset to their default values (shown in Figure 8) by clicking
on the button marked ‘Reset Values’.
To cater to the wide audience of potential users, we have
replicated much of the functionality of the JavaScript web ap-
plet in a MATLAB-based GUI. Data are entered in the panel
marked ‘Enter data here’. Results for the specified fish length
and grain size indices are reported in the table in the middle
panel. The outputs are updated after new data are entered by
clicking within the GUI or pressing ‘Enter’ on the keyboard.
The user can select and update the desired plot (e.g. FM and
NREDDS) using buttons in the panel labelled ‘Update figure’. A
contour plot of the desired output is generated in the D84 versus
D50 plotting space within the GUI (at left) for the fish of interest,
with a data point corresponding to the specified D84 and D50.
Discussion
The strength of Equations (4) and (7) and the associated charts,
look-up tables, and computer-based applications is that they
are good predictors of FM (Figure 3), which in turn appears to
be a good reflector of NREDDS (Figure 4), the substrate’s capacity
for accommodating redds built by the fish of interest. The pre-
dictive power of the model stems from its ability to take a
bed-surface grain-size distribution (represented simply by D50
and D84) together with a measure of the largest movable parti-
cle on the bed (predicted from fish length) and arrive at a real-
istic estimate of the area covered by movable particles. Thus,
the toolkit we have derived from the model allows managers
to gauge the spawning capacity of a substrate for a given spe-
cies as a continuous function of substrate grain size. This is ap-
parently because Equation (4) provides a robust approximation
to the integral of the grain-size distribution from the smallest
grain on the bed to largest grain that fish can move (Figure 1).
Limitations of the tools
The tools presented here are only demonstrably robust over the
range of sizes represented by the calibration data  in this case,
where D50 is between 39 and 118 mm, D84 is between 85 and
385 mm, and L is between 445 and 721 mm (Riebe et al.,
2014). Outside these ranges, application of the relationship
for calculating DT is extrapolative. However, the calibration
data span a large range in all of the calibration variables. For
example, the dominant grain size on the bed ranged from me-
dium gravel to cobbles and boulders. Moreover, the 445–721
mm range in salmon length corresponds to a broad range in
the ability of salmon to move sediment, spanning a 0.4–2.6
kg range in the size of the largest particle moved in redds (Riebe
et al., 2014). Thus, the model should be applicable to salmon
and trout in general, to the extent that it captures interspecies
differences in ability to build redds using the cutting motion
typical of salmonids. More research is needed to test this
hypothesis.
Our tools do not account for the widely recognized effects of
fine sediment on egg survival to emergence (e.g. Everest et al.,
1987; Bjornn and Reiser, 1991; Greig et al., 2005, 2007).
Sediment finer than 10 mm can fill interstices of sediment and
trap fry in redds by blocking pathways for emergence (Bjornn,
1969; Phillips et al., 1975; McCuddin, 1977; Harshbarger and
Porter, 1982; Bennett et al., 2003). When sediment is even finer
– less than about 1 mm in diameter – it can impede intragravel
flow of water, thus depriving incubating eggs of oxygen-rich
water and preventing removal of toxic metabolic wastes

Figure 8. Screen capture of web-based JavaScript interface, which can be accessed online in Supplementary Material 6 and on the second author’s
website. The interface generates a plot (at left) and output data (at right) for a user-provided inputs of grain size and fish length (upper right). Users can
choose to display one of three indicators of the amount of spawning habitat in the D84 versus D50 plotting space by clicking on tabs in the plot win-
dow. This figure is available in colour online at wileyonlinelibrary.com/journal/espl

© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd. Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
138
(McNeil and Anhell, 1964; Greig et al., 2005). We refer the
reader to several thorough reviews of the effects of fine sedi-
ment on egg survival to emergence for details (e.g. Chapman,
1988; Bjornn and Reiser, 1991; Kondolf, 2000; Jensen et al.,
2009). In summary, egg-to-fry survival for Pacific salmon has
been shown to drop significantly when sediment with grain
sizes less than 0.85 mm exceeds 10% of the sediment
contained in the bed (Jensen et al., 2009). The tools presented
here apply to upper particle size limits on redd building and
we advise that they should be used in combination with
methods for assessing fine sediment suitability (e.g. Kondolf,
2000; Bunte, 2004; Jensen et al., 2009).
Particles on bed surfaces that have been winnowed of fine
sediment are often more difficult to move than the same sized
particles on a loose bed (Parker and Klingeman 1982; Dietrich
et al., 1989). On such armoured beds, estimates of DT from
Equation (1) may be larger than what a spawning salmon can
realistically move. For the most realistic estimates of DT, we
recommend site-specific measurements of the largest particles
moved in spawning redds, which could be used instead of
Equation (1) if armouring is expected to be important.
Our tools do not account for the effects of superimposition
(e.g. Fukushima et al., 1998), which occurs when female
salmon destroy existing redds by building new redds over
them. Somewhat paradoxically, superimposition can increase
the spawning capacity of a substrate by decreasing the average
spacing of egg pockets (van den Berghe and Gross, 1984).
Thus we suggest that the estimates of spawning capacity
(i.e. NREDDS in Figure 7) are minima, due to the reduction in
minimum area required per redd that may be caused by
superimposition. As a direction for future research, the effects
of superimposition on the spawning capacity of substrates
could be explored at the reach scale by coupling our
substrate model (Equations (4) and (7)) with an individually
based population model, and thus account for run timing,
guarded area, and other factors that influence superimposi-
tion (e.g. Bartholow, 1996).
Data requirements
Our tools for gauging the capacity of spawning substrates are
easy to use. Only fish length and a measurement of the
grain-size distribution are needed to apply it to a site. The
pebble count is a straightforward, widely used method for
characterizing the grain-size distribution of a riverbed.
However, care is needed to obtain accurate, repeatable
measurements. Operator bias (Hey and Thorne, 1983; Marcus
et al., 1995; Wohl et al., 1996), sample size (Rice and Church,
1996), particle selection methods (Bunte and Abt, 2001a,
2001b; Bunte et al., 2009), and measurement methods (Bunte
et al., 2009) all influence pebble count results. Likewise, fish
lengths can vary substantially both within a given reach of river
in any given year and also from one year to the next (Johnson
and Friesen, 2013). These factors lead to uncertainties in FM
and NREDDS that can be quantified by propagating estimated
errors in D50, D84, and L through Equations (4) and (7). Thus,
data quality and sampling methods should always be assessed
and accounted for prior to drawing conclusions about FM and
NREDDS from grain-size and fish length data reported in the
literature. For example, more exhaustive pebble counts may
be required to resolve smaller differences in D50 and D84
(Rice and Church, 1996; Larsen et al., 2004) and thus in
FM and NREDDS.
Pebble counts are often a poor reflector of fine particles in
the grain-size distribution due to the difficulty of sampling them
and measuring their dimensions (Rice, 1995). However, this
has few if any implications for application of the tools
© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
B. T. OVERSTREET ET AL.
presented here; in the conditions relevant to quantifying limits
on salmon spawning due to coarse sediment, D50 and D84
are typically much greater than the 2 mm cutoff for fine sedi-
ment that must be grouped into a single size class because it
is difficult to sample and measure via pebble counts (Wolman,
1954; Bunte and Abt, 2001b). Because the tools presented here
apply to coarse-bedded rivers, where large grains limit a
salmon’s ability to move sediment, our approach is well suited
to integration of image-based grain-size mapping, including
both ground-based (Butler et al., 2001; Sime and Ferguson,
2003; Graham et al., 2005; Graham et al., 2010; Buscombe
et al., 2010) and aerial-based techniques (Carbonneau et al.,
2004; Verdú et al., 2005; Black et al., 2014). However, conver-
sion between image- and pebble count-derived grain-size
distributions will be required (Graham et al., 2012).
In practice, grain size generally needs to be measured at as
many representative locations as possible, because it can vary
considerably, even over the confined scale of a spawning riffle.
Pebble counts should be conducted within carefully defined
textural facies (Buffington and Montgomery, 1999), where the
bed-surface grain-size distribution appears to be spatially in-
variant. The spatial extent of the grain-size measurement will
be determined by the variability of textural facies. For example,
in the study that calibrated Equations (4) and (7), pebble counts
were conducted in 100–200 m2 sampling areas, consistent
with the spatial extent of distinct textural facies within the study
reaches (Riebe et al., 2014). Smaller streams may have higher
spatial variability of textural facies and thus may need to have
smaller sampling areas. We suggest that our model can be ap-
plied to scales ranging from a few square metres to hundreds
and perhaps even thousands of square metres depending on
the redd size of the species of interest, the spatial variability
of grain size, and the required precision of a study.
In most cases application of this method will be limited to
wadeable streams – e.g. with flow less than 1.5 m/s and depth
less than 1 m (Abt et al., 1989) – where surface grain-size data
can be collected via pebble counts. However, image-based
grain size sampling or underwater sampling methods could al-
low sampling in larger rivers. Sampling in larger rivers could
also be possible where flow is seasonally reduced or diverted.
In any case, when sampling facies before or after a spawning
season, it will be vital to consider the flow stage and inundation
area that typically prevail during spawning runs to avoid
overestimating or underestimating available spawning habitat.
In addition, sample sites in a time series of monitoring measure-
ments would need to be located as closely in space as possible
during successive measurements if the goal is to track changes
in spawning habitat over time.
Estimating the reproductive potential of spawning
substrates
In attempting to gauge the viability of a prospective restoration
project, it may be useful to predict outcomes in terms of salmon
reproduction. For example, in our previous work, we defined
the reproductive potential of the substrate (NEGGS) as the num-
ber of eggs per unit area that can be accommodated within the
substrate (Riebe et al., 2014). This can be calculated from
Equation (8) when fecundity, E, the number of eggs produced
per fish (and thus per redd), is known for the species of interest.
N EGGS ¼ N REDDS E=1000 (8)
Here NEGGS is the number of eggs in redds within the reach in
units of thousands of eggs per 100 m2. Fecundity generally
increases with fish length both within and across salmonid
species (Beacham and Murray, 1993; Quinn, 2005). This is
Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
 TOOLS FOR GAUGING THE CAPACITY OF SALMON SPAWNING SUBSTRATES
illustrated in Equation (9), a best-fit least-squares relationship
between average fecundity and average fish length for a series
of measurements reported in Quinn (2005) (after Riebe et al.,
2014).
E ¼ 8:1L 1450 (9)
Thus, by combining Equations (7) and (9) in Equation (8), it is
easy to translate the contour plot of NREDDS (Figure 5B) into a
contour plot of NEGGS (Figure 9). Both the JavaScript and
MATLAB interfaces (Supplementary Material 6 and 7) include
plots and output data for NEGGS based on E calculated using
Equation (9) (see for example, Figure 8).
Equation (9) explains 70% of the variance in average
fecundity across 13 species of salmon, trout, and char spanning
a wide range in average fish sizes. However, it is important to
recognize that regional and intrapopulation differences in
fecundity–length relationships can be substantial (Healey and
Heard, 1984; Beacham and Murray, 1993). Hence, we stress
that estimates of NEGGS that incorporate Equation (9) can at best
provide a generalized understanding of how substrate
reproductive potential varies in nature. We recommend using
site-specific values of E whenever possible to translate estimates
of NREDDS from Equation (7) into estimates of NEGGS via
Equation (8).
Equations (4), (7), and (8) yield several potentially useful indi-
ces of ecosystem function (i.e. FM, NREDDS and NEGGS). They
provide the means for translating observed changes in grain
size into estimates of resulting changes in the salmon spawning
Figure 9. Variations in NEGGS with grain size. Number of eggs (in
2
thousands) per 100 m plotted as a function of the standard deviation
and the mean grain size (equal to D84/D50 and D50, respectively, for a
lognormal bed-surface grain-size distribution) for a 600 mm long fe-
male salmon. The value of NEGGS is the reproductive potential of the
substrate; it expresses the number of eggs the substrate could physically
accommodate and reflects limitations set by the size of the fish, which
determines fecundity, the number of redds that can be fit into a given
area, and the fractional coverage by immovable grains on bed. The
NEGGS reflects a space limitation and is distinct from survival to emer-
gence (i.e. successful reproduction), which is moderated by environ-
mental factors such as flow depth, velocity, scour, and contamination
of the bed by fine sediment. This figure is available in colour online
at wileyonlinelibrary.com/journal/espl
© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
139
capacity and reproductive potential of substrates. Thus, the
tools presented here should provide realistic estimates of how
spawning habitat degradation (or enhancement) contributes to
observed decreases (or increases) in salmon populations. Yet
it is crucial to recognize that Equations (4), (7), and (8) are con-
cerned with just one of the many potential limits on salmon
populations – i.e. the upper grain-size limit on redd building
in coarse riverbeds. Hence the tools presented here are only
likely to inform management of salmon populations when oth-
erwise confounding factors, such as the abundance of fine sed-
iment, ocean conditions (Nickelson, 1986), stream temperature
(Richter and Kolmes, 2005), disease (Krkošek and Hilborn,
2011), and predation (Peterman and Gatto, 1978; Quinn and
Kinnison, 1999) are taken into account. Moreover, care should
be taken in applying the tools presented here outside the
Pacific Northwest and California, where they were calibrated.
Although our toolkit should be widely applicable to coarse-
bedded rivers where salmonids spawn, it has limitations that
will need to be accounted for in some settings. For example,
in steep, step-pool reaches, where salmonid species spawn in
small isolated patches of movable substrate (Montgomery
et al., 1999; Sear, 2010), our tools would likely need to be
calibrated to apply to only the individual patches of movable
substrate.
Optimizing spawning habitat restoration projects
Equation (4) and the tools derived from it show that FM
decreases monotonically with increases in D50 for a given
spread in the distribution (i.e. for a given D84/D50). Hence, it
is apparently always better to have a finer D50 if the desire is
more movable substrate (and more spawning capacity).
However, other factors besides FM must be taken into account
to assess spawning substrates and optimize gravel augmenta-
tion projects. For example, in addition to the previously
mentioned effects on egg survival to emergence, a bed that is
too fine-grained might lack occasional large ‘centrum’ rocks,
which may be needed to stabilize redds and provide shelter
for egg deposition (Hobbs, 1937; Burner, 1951; Jones and Ball,
1954). Moreover any added sediment should be coarse enough
that it is stable against mobilization and rapid downstream
transport in the flow of the river (e.g. Buffington and
Montgomery, 1997). Hence, we suggest that FM should be used
together with sediment transport modelling (e.g. Bunte et al.,
2013; Heimann et al., 2014), known flow criteria for spawning
and incubation (e.g. Tonina and Buffington, 2009; Moir and
Pasternack, 2010), and known limits on fine sediment
accumulation in redds (Jensen et al., 2009) to engineer a
hydrodynamically stable substrate that accommodates as many
redds as possible without being so fine that it suffocates eggs or
entombs fry. Thus the physical limits on redd building that are
incorporated into Equations (4) and (7) can be used together
with conventional understanding of hydraulics and sediment
transport to create stable channels that support good salmon
spawning habitat (Wohl et al., 2015). With insight from
Equations (4), (7), and (9), the cost-benefit analysis of sediment
augmentation could be simplified to the cost per egg for
sediment added to the bed (Riebe et al., 2014). In this way,
our approach could help managers optimize selection of
restoration sites for minimal cost and maximum benefit to the
species of interest.
The tools presented here can also be readily used to quantify
how much bed-surface area needs to be restored to meet
established restoration objectives. As noted earlier, the sensitiv-
ity of FM to differences in D50 depends on the spread in the
grain-size distribution (Figures 5A and 5B); when the spread is
Earth Surf. Process. Landforms, Vol. 41, 130–142 (2016)
140 B. T. OVERSTREET ET AL.
large, bigger changes in D50 are needed for a desired change in
coverage by movable particles (and thus by extension, desired
changes in NREDDS and NEGGS). This suggests that initial condi-
tions must be considered in the assessment of how much resto-
ration is needed. The lookup charts in Figures 6 and 7 generally
suggest that bigger increases in FM and NREDDS (and thus bigger
increases in NEGGS) can be gained by adding movable sedi-
ment to reaches with relatively low D84/D50 (i.e. with rela-
tively narrow grain-size distributions). Moreover Figures 6
and 7 show that habitat can be improved by reducing D84/
D50 without changing D50 when D50 < DT. This implies that
habitat can be improved without gravel augmentation by re-
moving the coarsest sediment from the bed and thus reducing
D84/D50. Somewhat paradoxically, Figures 6 and 7 also sug-
gest that it may be more economical on a cost-per-egg basis
to focus restoration on reaches that already support consider-
able spawning. This presumes, of course, that hydraulic fac-
tors are suitable enough for fish to exploit new areas of
movable sediment in the reach. Such sites may often have
the added advantage of being hydrodynamically stable,
which would contribute to longer residence times for added
sediment. In any case, the tools presented here can help in al-
locating resources to the reaches that would yield the biggest
and most stable increases in reproductive potential of
spawning substrates.
Once augmentation projects have been initiated, our tools
for gauging spawning substrates can be readily used to monitor
changes in reproductive potential over time, thus enabling
assessment of whether restoration projects are meeting desired
goals (Riebe et al., 2014). When historical records are avail-
able, our tools can be used to quantify how measured changes
in grain size may have contributed to changes in populations
over time. Hence one could test hypotheses about cause–effect
connections between historical declines in spawning and
changes in spawning substrates, and thus build evidence for
(or rule out) gravel augmentation as a viable approach to revi-
talizing salmon populations in a reach of interest. Quantifying
historical changes in reproductive potential could also help
managers understand and possibly also predict impacts of land
use and climate change on the amount and quality of spawning
substrates (Riebe et al., 2014).
Conclusions
Our new tools for assessing spawning habitat quality employ
easy-to-measure inputs of D50, D84, and fish length to gauge
a substrate’s spawning capacity, defined as the number of redds
per unit area that can be accommodated by the riverbed.
Though the formulations of this new approach are somewhat
complex, they are derived from empirical relationships that
capture the mechanics of redd building and also fit data from
diverse species of salmon across riverbeds with diverse grain-
size distributions. Moreover, as demonstrated here, they are
readily encapsulated in a series of easy-to-use charts, tables,
and computer-based applications. The result is a management
toolkit that accounts for interspecies differences in ability to
move sediment for salmonids that build redds using a cutting
motion. All that is needed to apply this toolkit is a pebble count
and a carcass survey, or some other measurement of fish length.
With additional measurements of fecundity, the predictions of
spawning capacity can be translated to predictions of repro-
ductive potential, defined as the maximum number of eggs that
can be deposited per unit area for the fish of interest. Thus, the
tools presented here can be readily applied to address a wide
range of issues in the management of salmon spawning habitat.
These include:
© 2016 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.
• gauging the capacity of spawning substrates across sites and
salmonid species;
• optimizing selection of restoration sites for maximum benefit
to the species of interest;
• quantifying how much bed-surface area needs to be restored
to meet established restoration objectives;
• improving cost-benefit analysis in the allocation of river res-
toration budgets among prospective sites and activities;
• optimizing grain-size distributions in gravel augmentation
projects to create hydrodynamically stable substrates that
hold as many redds as possible;
• monitoring changes in the reproductive potential of
spawning substrates over time to assess the effectiveness of
restoration projects and to quantify impacts of land use and
climate change;
• contributing to more realistic models of salmon population
dynamics.
In summary, the tools presented here provide a multifaceted
and comprehensive new approach to gauging the spawning ca-
pacity and reproductive potential of riverbed substrates used by
female salmon for redd building when factors such as fine sed-
iment content, flow depth, and velocity are within suitable
ranges. Our toolkit also provides a mechanistic basis for
optimizing gravel augmentation, thus contributing to more
cost-effective, scientifically based restoration of salmon
spawning in coarse-bedded rivers.
Notation
AREDD Map area of redd (m2).
D50, D84 50th and 84th percentiles of grain diameters on river-
bed (mm).
DT Intermediate-axis diameter of largest sediment grain
that salmon of a particular size can move during redd
building (mm).
E Fecundity, which is the number of eggs female salmon
produce (eggs/fish) and thus also the maximum num-
ber they can deposit within an individual redd (eggs
/redd).
FM Fractional coverage of bed by particles that female
salmon can move during redd building.
L Fork length of female salmon (mm).
NREDDS Substrate spawning capacity, equal to the maximum
number of redds that can be built per unit area in a
given reach by a fish of a given size (redds/100 m2).
NEGGS Substrate reproductive potential, equal to the maximum
number of eggs that can be deposited per unit area in a
given reach by a fish of a given size (1000 eggs/100 m2).
z Dimensionless exponent analogous to Z statistic in one-
parameter logistic-function approximation of the cumu-
lative normal distribution.
Acknowledgements—This work was supported by the National Science
Foundation grant EAR-0956289 to Riebe. The authors thank Frank
Ligon for inspiration and Matt Sloat and Nick Brozović for helpful
discussions. The authors also thank two anonymous reviewers and
the associate editor for comments and suggestions that improved the
manuscript.
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