AN ABSTRACT OF THE DISSERTATION OF
Thomas H. Williams ifi for the degr e of Doctor of Philosophy in Fisheries Science
presented on April 21, 2004.
Title: Geographic Variation in Gene ic and Meristic Characters of Coastal Cutthroat
Trout (Oncorhynchus clarki clarki).
Abstract approved:
Redacted for privacy
H. Reeves
Understanding how populations
geographic and temporal scales is fu
conservation strategies. I examined t
characters of coastal cutthroat trout
approximately 1,400 fish sampled fr
range (northern California to Prince
based on general biogeographical be
ithin a species interact across various
idarnental to developing appropriate
ie geographic variation in genetic and meristic
)ncorhynchus clarki clarki) based on
m 54 populations spanning their distributional
Villiam Sound, Alaska). Four geographic areas,
ndaries largely driven by Pleistocene
glaciation, were used to examine re onal patterns of population structure. Across
their range, the population structure f coastal cutthroat trout was that of many
diverse local populations. Populatioi s exhibited extensive variation in meristic
characters across their range. Regiori 1 clustering of populations from the southern
portion of the range contrasted with opulations from the central and northern
portion of the range which did not e thibit geographic concordance. In addition to the
strong phenetic affinity of the southc n populations, the intra-regional differences
among populations in the southern n gion was greater than that observed in the other
regional areas. Analysis of genetic p pulation structure based on 30 enzyme
encoding loci revealed geographic c ncordance of populations in the northern and
southern regions of the range with ii t le geographic concordance in genetic structure
from populations in the central regic s of the range. Throughout the range, isolationby-distance (IBD) was detected at a r gional scale (<800 km) and was strongest in
the northern and southern regions. T1 e primary genetic structure of coastal cutthroat
trout populations occurred at the mdi ridual stream level, and there was genetic
affinity among populations at a regional scale. The strong geographic concordance
and inter-regional divergence of meristic characters exhibited by the southern
populations was consistent with other ecological studies that have found that
peripheral populations tend to be genetically and morphologically divergent from
central populations and morphological characters are expected to diverge more
rapidly in isolated populations than gene frequencies. These data suggest that
compared to other species of Pacific salmon and trout, coastal cutthroat trout are
characterized by many smaller, genetically diverse local populations that act in a
more independent, isolated nature over short time frames (<100 years).
© Copyright by Thomas H. Williams ifi
April 21, 2004
All Rights Reserved
Geographic Variation in Genetic and Meristic Characters of
Coastal Cutthroat Trout (Oncorhynchus clarki clarki)
by
Thomas H. Williams ifi
A DISSERTATION
submitted to
Oregon State University
in partial fulfillment of
the requirements for the
degree of
Doctor of Philosophy
Presented April 21, 2004
Commencement June 2004
Doctor of Philosophy dissertation of Thomas H. Williams ifi presented on April 21,
2004
APPROVED:
Redacted for privacy
Redacted for privacy
Redacted for privacy
I understand that my dissertation will become part of the permanent collection of
Oregon State University libraries. My signature below authorizes release of my
dissertation to any reader upon request.
Redacted for privacy
Thomas H. Williams ifi, Author
ACKNOWLEDGMENTS
I wish to acknowledge the support and patience of my graduate committee: Dr.
Gordon Reeves, Dr. Phillip Brownell, Dr. Paul Farber, Dr. Daniel Goodman, and Dr.
Carl Schreck. I am grateful to my major professor Dr. Reeves for continued
encouragement, guidance, and friendship. Special thanks to Dr. Ken Currens for
guidance throughout this research, particularly with the genetic analyses. Thanks to
the Oregon Cooperative Fishery Research Unit for the use of their facilities and their
collaboration throughout the project, Neil Ward was instrumental in the allozyme
laboratory analysis. A special thanks to Drs. C. Bond, J. Hall, and D. Markie for
guidance and encouragement. Robert Schick (NOAA) provided assistance with GIS
analysis and guided me through the spatial analyses. This study was made possible
with support and collaboration of many individuals. Individuals that I would like to
specifically acknowledge include: D. Aho, D. Anderson, D. Atagi, R. Billings, C.
Casipit, P. Caverhill, R. Cooper, J. Dunham, T. Down, R. Dunlap, F. Everest, E.
Gerstung, K. Griswold, J. Grunbaum, J. Hannon, B. Hansen, P. Harris, B. Hooten, S.
Johnson, T. Johnson, E. Johnston, D. Jones, G. Killinger, M. Knapp, P. Law, S.
Leider, M. Brennan Levesque, S. Levesque, E. Loudenslager, R. Medel, J.
McDonell, D. Moore, T. Noel, S. Pollard, R. Ptolemy, M. Ramsay, C. Riley, T.
Roelofs, D. Saiget, M. Scheiske, D. Schmid, A. Schmidt, V. Staroska, C. Thomas, R.
Troll, C. Wheeler, J. White, B. Wright, and C. Zimmerman. I am particularly
grateful for the support and encouragement I received from my colleagues and
supervisors at the NOAA Fisheries, Santa Cruz Laboratory, with a special thanks to
librarian K. Johnston.
My thanks, especially to my family, Susan for her never ending confidence and
encouragement, and Isabelle and Emma for patience far beyond their years. Thanks
also to my parents for their continued support.
Funding for this research was provided by the USDA Forest Service, Regions 6
and 10, and the Aquatic Land Interaction Program at the Pacific Northwest Research
Station. In addition, scholarship support was received from Oregon Council
Federation of Fly Fishers, Federation of Fly Fishers (Robert Marriott Scholarship),
Trout Unlimited (Lee Wuiff Scholarship), and the Oregon Chapter of the American
Fisheries Society (Graduate Scholarship Award).
TABLE OF CONTENTS
Page
1. INTRODUCTION
.
2. MERISTIC CHARACTERIZATION OF COASTAL CUTTHROAT TROUT
(Oncorhynchus clarki ciarki) ACROSS THEIR DISTRIBUTIONAL RANGE
.
1
5
.................................................. 5
2.2. INTRODUCTION ............................................. 6
2.3. METHODS ................................................. 12
2.3.1. Sample collection and analysis ............................. 12
2.3.2. Data analysis ........................................... 18
2.4. RESULTS .................................................. 24
2.4.1. Identification of steelhead ................................. 24
2.4.2. Differences in meristic counts between sex and among length groups
within sampling locations ................................ 26
2.4.3. Differences among sampling locations ....................... 27
2.5. DISCUSSION ............................................... 42
2.1.ABSTRACT
3. GENETIC POPULATION STRUCTURE OF COASTAL CUTTHROAT TROUT
(Oncorhynchus clarki clarki) IN WESTERN NORTH AMERICA
57
..........
................................................. 57
3.2. INTRODUCTION ............................................ 57
3.3. METHODS ................................................. 64
3.3.1. Sample collection and analysis ............................. 64
3.3.2. Data analysis ........................................... 71
3.4. RESULTS .................................................. 74
3.5. DISCUSSION ............................................... 92
4. CONCLUSION ................................................. 107
BIBLIOGRAPHY ................................................. 114
APPENDIX FIGURES ............................................. 129
APPENDIX TABLES .............................................. 137
3.1.ABSTRACT
LIST OF FIGURES
Figure
Page
2.1. Locations where samples were collected and regional groupings for analysis.. 13
2.2. Maxillary extension scores for coastal cutthroat trout
.................... 19
2.3. Mean (± 1 SE) counts of anal pterygiophores and dorsal pterygiophores for each
sampling location of coastal cutthroat trout
............................. 28
2.4. Mean (± 1 SE) counts of vertebrae and basibranchial teeth for each sampling
location of coastal cutthroat trout
..................................... 29
2.5. Mean (± 1 SE) counts of scales in lateral series and scales above lateral line for
each sampling location of coastal cutthroat trout
30
.........................
2.6. Mean (± 1 SE) counts of pelvic fin rays and pectoral fin rays for each sampling
location of coastal cutthroat trout
..................................... 31
2.7. Mean (± 1 SE) counts of branchiostegal rays and pyloric caeca for each
sampling location of coastal cutthroat trout
............................. 32
2.8. Mean (± 1 SE) counts of upper gill rakers and lower gill rakers for each
sampling location of coastal cutthroat trout
............................. 33
2.9. The mean (±1 SE) canonical variate I score for each sampling location based on
11 meristic characters
.............................................. 35
2.10. Mean canonical variate I versus latitude (UTM northern coordinate; X 106) for
coastal cutthroat trout from 54 locations
............................... 38
2.11. Phenogram from IJPGMA cluster analysis of Euclidian distance based on 11
meristic characters of coastal cutthroat trout from 54 locations
39
.............
2.12. Multidimensional scaling (dimensions 1,11 and ifi) of Euclidian distance based
on 11 meristic characters of coastal cutthroat trout from 54 locations ........ 41
2.13. Multidimensional scaling (dimensions 1,11 and ifi) of Euclidian distance based
on nine meristic characters of 54 samples of coastal cutthroat trout from this study
and other species of western trout
.................................... 48
3.1. Locations where samples were collected and regional groupings for analysis.
66
3.2. Consensus tree based on 1,000 replications of genetic relationships among 54
samples of coastal cutthroat trout from across distributional range and a steelhead
outgroup (Suntaheen Creek, Alaska, location 4)
......................... 79
3.3. Consensus tree based on 1,000 replications of genetic relationships among 54
samples of coastal cutthroat trout from across distributional range and a steelhead
outgroup (Suntaheen Creek, Alaska, location 4)
......................... 81
LIST OF FIGURES (continued)
Figure
Page
3.4. Multidimensional scaling (dimensions 1,11 and ifi) based on allele frequencies of
17 polymorphic loci (P095) of coastal cutthroat trout from 54 locations
83
.......
3.5. Multidimensional scaling (dimensions 1,11 and ifi) based on allele frequencies of
12 polymorphic loci (F095) of coastal cutthroat trout from 54 locations
84
.......
3.6. Isolation-by-distance analyses for all coastal cutthroat trout locations sampled
across distributional range
.......................................... 86
3.7. Isolation-by-distance analyses for all coastal cutthroat trout locations sampled
across distributional range
.......................................... 87
3.8. Isolation-by-distance analyses for coastal cutthroat trout locations in regional
groups A (upper plot) and B (lower plot)
............................... 88
3.9. Isolation-by-distance analyses for coastal cutthroat trout locations in regional
groups C (upper plot) and D (lower plot)
............................... 89
3.10. Multi-variate correlogram (non-linear Mantelogram) representing
autocorrelation statistics plotted against distance (shortest water distance).
...
90
3.11. Multi-variate correlogram (non-linear Mantelogram) representing
autocorrelation statistics plotted against distance (depth constrained water
distance)
........................................................ 93
4.1. A scatter plot of pairwise genetic distance (0) versus pairwise meristic distance
(Euclidian distance) for all locations of coastal cutthroat trout sampled
108
......
4.2. A simplified construct of the temporal scale (horizontal axis) and latitudinal
scale (vertical axis) of disturbances in coastal basins across the distributional
range of coastal cutthroat trout
...................................... 111
LIST OF TABLES
Table
Page
2.1. Collection sites, location, number of fish collected (N), and number of fish
determined to be coastal cutthroat trout based on laboratory examination of
basibrianchial teeth and maxillary extension (n)
.........................
14
2.2. Summary of meristic counts from all samples of 0. c. clarki and fish designated
as 0. mykiss from 54 sites
17
..........................................
2.3. Published characteristics of basibranchial teeth and maxillary extension used to
distinguish coastal cutthroat trout (0. c. clarki) and steelhead (0. mykiss).
.................................................. 21
2.4. Summary of meristic counts from all samples of 0. c. clarki and fish designated
as 0. mykiss or hybrids from 54 sites
25
..................................
2.5. Summary of mean meristic counts for various species of trout from western
North America reported in the literature and mean meristic counts for steelhead
from Suntaheen Creek, Alaska, collected in this study
47
....................
3.1. Collection sites, location, number of fish collected (N), and number of fish
determined to be coastal cutthroat trout based on laboratory examination of
basibrianchial teeth and maxillary extension (n)
.........................
65
3.2. Enzymes and electrophoretic conditions to resolve loci examined in this study.69
3.3. Location and frequency of private alleles detected from 54 samples of coastal
cutthroat trout. Location numbers correspond to numbers in Table 3.1 and Figure
3.1
............................................................. 76
3.4. Regional and overall F-statistics and correlation results from Mantel test for
isolation-by-distance based on 0(17 polymorphic loci) and geographic distance
(shortest water distance)
78
............................................
3.5. Hierarchical gene diversity analysis of 54 coastal cutthroat trout populations
from across their distributional range
..................................
78
3.6. Mantel correlogram values represented in Figure 3.13. Distances based on the
shortest water distance between a pair of locations
91
.......................
3.7. Mantel correlogram values represented in Figure 3.14. Distances are based on
depth constrained water distance
94
.....................................
3.8. A summary of genetic diversity analyses of selected salmonid species ...... 97
LIST OF APPENDIX FIGURES
Appendix Figure
APPENDIX FIGURES
Pge
............................................. 129
A2. 1. Mean (± 1 SE) counts of anal pterygiophores and dorsal pterygiophores for A:
fish identified as steelhead or coastal cutthroat trout and B: steelhead, suspected
hybrid, or coastal cutthroat trout based on basibranchial teeth and the length of
the maxillary extension
........................................... 130
A2.2. Mean (± I SE) counts of vertebrae and basibranchial teeth for A: fish
identified as steelhead or coastal cutthroat trout and B: steelhead, suspected
hybrid, or coastal cutthroat trout based on basibranchial teeth and the length of
the maxillary extension
............................................ 131
A2.3. Mean (± 1 SE) counts of scales in lateral series and scales above lateral line
for A: fish identified as steelhead or coastal cutthroat trout and B: steelhead,
suspected hybrid, or coastal cutthroat trout based on basibranchial teeth and the
length of the maxillary extension
.................................... 132
A2.4. Mean (± 1 SE) counts of pelvic fin rays and pectoral fin rays for A: fish
identified as steelhead or coastal cutthroat trout and B: steelhead, suspected
hybrid, or coastal cutthroat trout based on basibranchial teeth and the length of
the maxillary extension
............................................ 133
A2.5. Mean (± 1 SE) counts of branchiostegal rays and pyloric caeca for A: fish
identified as steelhead or coastal cutthroat trout and B: steelhead, suspected
hybrid, or coastal cutthroat trout based on basibranchial teeth and the length of
the maxillary extension
............................................ 134
A2.6. Mean (± 1 SE) counts of upper arch gill rakers and lower arch gill rakers for
A: fish identified as steelhead or coastal cutthroat trout and B: steelhead,
suspected hybrid, or coastal cutthroat trout based on basibranchial teeth and the
length of the maxillary extension
135
....................................
A3. 1. Percent polymorpic loci (P095; dashed line) and expected proportion of
heterozygotes (HE; Weir 1996; solid line) of coastal cutthroat trout populations
sampled across their distributional range
136
..............................
LIST OF APPENDLX TABLES
ApDendix Table
Page
.............................................. 137
A2.1. Meristic counts and collection information for all fish collected ......... 138
A2.2. Summary of meristic counts from each collection location .............. 209
APPENDIX TABLES
A2.3. Euclidian distances (lower left) and medium dissimilarities (upper right)
between pairs of 54 samples of 0. c. clarki
...........................
230
A3. 1. Summary of allele frequencies, sample sizes, polymorphic loci (F095), average
number of alleles per locus, and the expected proportion of heterozygotes (Weir
1996) at each location sampled
236
....................................
A3.2. Summary of loci that tested significant for deviations from Hardy-Weinberg
expectations based on exact test and x2 test for deficiency or excess of
heterozygotes
................................................... 250
GEOGRAPHICAL VARIATION IN GENETIC AND MERISTIC CHARACTERS
OF COASTAL CUTTHROAT TROUT (Oncorhynchus clarki clarki)
1P1Ii(S)1
Populations of a species are distributed in concert with both abiotic and biotic
conditions of the environment. Local populations (i.e., stocks) of Pacific salmon and
trout (Oncorhynchus spp.) are reproductively isolated in space and time and have
evolve4 in response to both regional and local environmental conditions. This
concept was formalized by Ricker (1972) to define a stock, his adaptation of the
biological species concept of Mayr (1942; 1963). Thompson (1965) recognized the
importance of local environmental conditions when he wrote "each stream or lake
has its own extremely complex characteristics, and if salmon live in one of them we
find these salmon are adapted in an equally complex way to that environment." More
recently, Taylor (1991b) found examples of local adaptation in salmonids on broad
and micro-geographic scales although most evidence was circumstantial. There is
now increasing evidence that local adaptation does occur in Pacific salmon and trout
(Wood 1995).
This stock concept is widely used in the management of Pacific salmon and trout.
Designation of a stock was initially based on morphological, behavioral, and
physiological features. Before the advent of techniques to describe molecular genetic
variation among populations, W. F. Thompson (1959; 1965) suggested that
conservation efforts for Pacific salmon should focus on local populations. More
recently, genetic data have been used (Utter 1991). Now stocks are being defined
more precisely as local populations with the use of genetic data (Booke 1981).
Interaction among populations of a particular species is a critical factor to
understand when developing management and conservation plans for Pacific salmon
2
and trout. The large intra- and inter-specific variability of life history characteristics
of Pacific salmon and trout (Groot and Margolis 1991) have evolved, in part,
because of diversity of environments inhabited by these fish. In North America
Pleistocene climates and landscapes influenced the distribution of freshwater biotas
(Avise 1992). The range of Pacific salmon and trout has been greatly influence by
large scale processes such as glaciation and volcanism (McPhail and Lindsey 1986)
and smaller scale processes such as fire and flood (Benda 1994). Bayley and Li
(1992) believe that species possessing polymorphic attributes (e.g., iteroparous and
semalparous life histories, anadromous and nonanadromous life histories) are
buffered against environmental uncertainty. Attempts to establish or reestablish
populations of Pacific salmon and trout, especially those with more variable life
history types such as sockeye salmon (0.
clarki),
nerka)
and coastal cutthroat trout (0.
clarki
have not been successful, suggesting that some adaptation for the local
environment has occurred.
The lack of success in efforts to establish or reestablish Pacific salmon and trout
to suitable habitat suggest that the plasticity observed in Pacific salmon and trout is
adaptive (Healey and Prince 1995). The diversity and availability of habitat may not
only be important to provide avenues for movement among populations, but may
also provide opportunities for the expression of phenotypic and genetic variation.
With an increasing amount of genetic data available to geneticists and resource
managers, the task of defining groups of populations became more difficult because
these data indicated that population structure was more complex than previous data
indicated (e.g., life history traits) and movement of individuals among populations
was occurring. The fundamental unit (e.g., demes, randomly mating populations,
local populations) identified by geneticists is no longer the same unit (e.g., stock)
that was defined prior to the use of genetic data and theory (Currens 1997).
The concept of "metapopulations" by Levins (1969) provides perspective for
examining spatial structure of populations. A metapopulation can be thought of as a
population consisting of many local populations that are connected by the movement
of individuals among the local populations. These local populations go through a
process of extinction and then recolonization from elsewhere within the
metapopulation (Gilpin 1991). The concept has much in common with dynamic
theory of island biogeography (MacArthur and Wilson 1963; MacArthur and Wilson
1967), although MacArthur and Wilson dealt with multi-species communities
(Hanski and Simberloff 1997). Pacific salmon and trout have some component of
each population that may stray to non-natal streams (Neave 1958; Quinn 1984;
Ricker 1989; Tailman and Healey 1994). This suggests that a metapopulation-type
spatial structuring of populations may be useful for conservation and management of
Pacific salmon and trout. Movement of individuals among populations (Wiens 1997)
depends on the availability of suitable habitat features at various spatial and temporal
scales.
In this dissertation, I will develop the argument that management and
conservation efforts for Pacific salmon and trout must recognize the importance of
both genetic and phenotypic variation and that this recognition should occur with an
appreciation and understanding of the influence environmental variation has in the
persistence of these fish. I present empirical evidence concerning the geographic
variation in population structure of coastal cutthroat trout from across their
distributional range. Chapter 2 examines the population structure using meristic
characters from populations sampled across the distributional range of this
subspecies. Chapter 3 examines the population structure of the same populations
using genetic characters allele frequencies. With this descriptive information
concerning the population structure of these populations, Chapter 4 synthesizes these
genetic and meristic data to describe the geographic variation observed in coastal
cutthroat trout and lays the conceptual framework for understanding how habitat
variability may interact at various spatial and temporal scales to shape the spatial
distribution and persistence of populations.
2. MERISTIC CHARACTERIZATION OF COASTAL CUTTHROAT TROUT
(Oncorhynchus clarki clarki) ACROSS THEIR DISTRIBUTIONAL RANGE
2.1. ABSTRACT
Conservation planning for any species requires an understanding of the
interaction among populations across different geographic scales. The use of
meristic measures to exam population structure of Pacific salmonids provides a
different perspective to recent efforts that have relied heavily on molecular
approaches. I examined meristic differences of 1,491 coastal cutthroat trout
(Oncorhynchus clarki clarki) from 54 sites across their distributional range using 13
meristic characters. Across their range, coastal cutthroat trout exhibited extensive
variation in the meristic characters examined. Populations at the southern end of the
distributional range exhibited phenetic affinity despite significant meristic
differences within this regional area. In other portions of the range, little geographic
concordance in meristic characters was observed. There were no latitudinal dines
detected for any of the meristic characters examined. Juvenile fish with intermediate
phenotypes consistent with those expected from decedents of coastal cutthroat trout
and steelhead hybridization were detected. Despite the large amount of meristic
variation observed across their distributional range, when compared to other western
trout species, the coastal cutthroat trout populations examined maintained a phenetic
affinity. The observation of unique meristic characteristics of southern populations
of coastal cutthroat trout at the peripherery of the range is consistent with
observations of other species at the margins of their distribution and suggest that
these populations may require special consideration in conservation planning.
L,-J
2.2. INTRODUCTION
Long-term conservation plans for any species are likely to depend on the
identification of distinct populations or populations groups and an understanding of
the spatial distribution and relationships among populations. This understanding is
critical with respect to defining units of conservation concern and for identifying
potential demographic constraints to recovery (Esler 2000), for it is these distinct
population segments that form the foundation upon which considerations of
extinction risk and recovery options are built (Waples 1991; Waples 1995). Fish
species typically consist of many geographically localized populations and, to
varying degrees, reproductively isolated populations (Taylor 199 ib). This is
particularly true for local populations of Pacific salmon and trout
(Oncorhynchus
spp.) that tend to become distinct from other local populations because of their high
fidelity for returning to their natal stream for spawning.
Current efforts to construct geographic boundaries of distinct population
segments of Pacific salmon and trout (Evolutionarily Significant Units) rely heavily
on molecular tools such as protein electrophoresis and DNA analysis (Waples 1991;
Waples 1995). These molecular tools provide data that are used to evaluate isolation
among populations since it is believed that they reflect levels of gene flow that
occurred over evolutionary time scales (Waples 1995).
More traditional methods for identifying populations that use morphology and
meristic characters for constructing species and population boundaries are used less
frequently because they usually require lethal sampling, the characteristics can be
substantially influenced by environmental factors, and the methods can be very time
consuming. However, these approaches may provide a different perspective on the
geographic distribution of populations and a context in which to interpret molecular
7
data. Meristic characters are body features such as vertebrae, fin rays, and scales,
that at one time in the evolutionary past correspond to body segmentation; other
characters that can be counted are sometimes referred to as meristic even though
they have no correspondence with the myomeres (i.e., myomerism) (Strauss and
Bond 1990).
Meristic characters have been used to examine interspecific differences and
intraspecific population structure of many fish species (i.e., species are usually
divided into subpopulations across their range). The systematics of western trout has
relied heavily on meristic and morphological characters (Behnke 1992). The
systematics of trout in the southern Sierra Nevada has been extensively studied with
the use of meristics (Evermann and Bryant 1919; Miller 1950; Schreck and Behnke
1971; Gold and Gall 1975; Gold 1977; Gold and Gall 1981). Legendre et al. (1972)
used meristic characters and coloring to indicate phenotypic similarities among
several species of western trout (Oncorhynchus spp.). Qadri (1959) used spotting
pattern, scale counts, and number of vertebrae to distinguish coastal cutthroat trout
(Oncorhynchus clarki clarki) and westslope cutthroat trout (0. c. lewisi).
Meristics have also been used to examine intraspecific variation in other fish
species. Populations of marine species such as black sea bass (Centropristis striata)
(Shepherd 1991), Atlantic cod (Gadus morhua) (Pepin and Can 1993; Swain et al.
2001), witch flounder (Glyptocephalus cynoglossus) (Bowering and Misra 1982),
American shad (Alosa sapidissima) (Melvin et al. 1992), and yellowfin tuna
(Thunnus albacares) (Royce 1964) have been differentiated by meristic characters.
Beacham (1985) found differences in vertebral and gill raker counts useful for
population identification of sockeye salmon (0. nerka) on a regional basis. Meristic
characters have been used to determine population structure among populations of
['I
iJ
Arctic charr (Salvelinus alpinus) across large regional areas (Dempson and Misra
1983), between different morphs of nonanadromous Arctic chan (Alekseyev et al.
2002; O'Connell and Dempson 2002), and between anadromous and nonanadromous
Arctic chan in the same basin (McCart and Craig 1971). No difference between
anadromous and nonanadromous Atlantic salmon (Salmo salar) parr were detected
within the same drainage based on dorsal, anal, and pectoral fin rays and vertebrae
(Riley et al. 1989). Many studies have examined differences among populations of
steelhead based on merjstjc characters (Winter et al. 1980; Schreck et al. 1985;
Currens et al. 1990).
Meristic characters in fish are generally considered to have a genetic basis
(Barlow 1961; Leary et al. 1985), but environmental factors can have substantial
influence on them (Strauss and Bond 1990) and may modify an individual's meristic
phenotype (Beacham 1985). Relatively minor changes in environmental factors
during ontogeny can result in significant within species differences in meristic
counts (TAning 1952). Efforts to identify populations of fish based on phenotypic
variation must consider that differences observed may be environmentally-induced
rather than genetically based (Swain and Foote 1999). Leary et al. (1991) found that
rearing density had a significant effect on vertebrae and pelvic fin ray counts in
rainbow trout, although the differences were minor compared to differences
associated with genetic and temperature effects. Blouw et al. (1988) examined the
variability of five meristic characters (vertebrae, gill rakers, pectoral fin rays, dorsal
fin pterygiophores, and anal fin pterygiophores) of Atlantic salmon over a 10-year
period to estimate within and among year variability. Although they found these
meristic traits to be remarkably homogenous within cohorts, the temporal variation
among years was extensive and persistent.
I,;
Even with environmental influence, if meristic differences among populations or
population groups exist, these measures may be meaningful for understanding
population structure (Shepherd 1991; Melvin et al. 1992). Differences may be very
useful if they relate to fitness and are subject to strong selection pressure (Swain and
Foote 1999). Selection could result in rapid genetic divergence in phenotypic traits
among populations faster than neutral genetic differences could accumulate (Swain
and Foote 1999). Leary et al. (1991) reported observations based on experiments
with rainbow trout, similar to findings of Ali and Lindsey (1974) with medaka
(Oryzias latipes), that the magnitude and direction of meristic variation resulting
from environmental factors appeared to have a genetic component.
Coastal cutthroat trout provide an opportunity to examine the usefulness of
meristic characters to understand the distribution and variation of populations in a
species of Oncorhynchus. Although few studies have investigated the genetic bases
for meristic counts in cutthroat trout (0. clarki spp.), a genetic basis has been shown
in steelheadlrainbow trout (0. mykiss spp.) for scale counts (Winter et al. 1980),
vertebrae (Winter et al. 1980; Leary et al. 1985), branchiostegals (MacGregor and
MacCrimmon 1977), gill rakers (Smith 1969; Leary et al. 1985), and fin rays
(MacGregor and MacCrimmon 1977; Leary et al. 1985).
Perhaps due to a lack of emphasis in research and management (National
Research Council 1996), few data exist concerning the population structure of
coastal cutthroat trout populations at regional scales and across its distributional
range. Nehisen et at. (1991) concluded that this prevented scientists from delineating
coastal cutthroat trout populations in a meaningful way (Nehisen et al. 1991). The
coastal cutthroat trout is the most widely distributed subspecies of cutthroat trout
(Behnke 1992; Behnke 2002). They occupy habitats ranging from small to large
10
rivers that drain the coastal rainforest between the Eel River, California, and Prince
William Sound, Alaska. The unique karyotype in 0. c. clarki indicates that the
subspecies has maintained unity across its range and maintained isolation from other
subspecies (Behnke 1992).
As with other Pacific salmonids, studies have shown that coastal cutthroat trout
form local populations. Applying various genetic techniques Campton and Utter
(1987) and Wenburg et al. (1998) have found that, within regional areas (<200 km),
coastal cutthroat trout show greater differences among populations than other species
of Pacific salmonids, In addition, coastal cutthroat trout exhibit a great amount of
variation in life-history types compared to other Pacific salmonids. In general, there
are two major life history forms, amphidromous' and potamodromous, although life
history traits (e.g., freshwater residence, age at sexual maturity) appear to be highly
variable between these two forms and among different populations of similar forms
(Sumner 1953; Giger 1972; Jones 1977; Johnston 1982).
The possibility of hybridization with steelhead exists for many populations of
coastal cutthroat trout. Behnke (1992) speculated that in small streams where two
species of trout are found, crossbreeding occurs if reproductive isolation (e.g.,
insufficient space to allow physical separation at spawning) breaks down. The
absence of basibranchial teeth in a coastal cutthroat has been thought to be indicative
of hybridization with steelhead (Behnke 1992). In addition, Behnke (1992) states
that hybrids may also differ in coloration, spotting pattern, and numbers of scales,
caeca, and vertebrae. Campton and Utter (1985) used allozymes to identify juvenile
fish with intermediate phenotypes consistent with those expected for decedents of
1
Myers (1949) described amphidromous fish migration as movement from
freshwater to the sea, or vice versa, for non reproductive purposes, such as feeding;
potamodromous fishes migrate wholly within freshwater.
11
coastal cutthroat trout and steelhead. Their results from two Puget Sound streams,
the first formal documented observation of hybrids in the wild, led them to suggest
that in streams where both species spawn "the production of hybrid offspring may
not be uncommon." More recently, many genetic studies have detected the
occurrence of hybrids between coastal cutthroat trout and steelhead in the wild
(Hawkins 1997; Wenburg et al. 1998; Young et al. 2001). Use of meristics to
delineate populations of coastal cutthroat trout may be complicated by the presence
of hybrids, however, meristic characters may provide a useful tool to examine the
distribution and prevalence of hybridization.
In this study, which included coastal cutthroat trout throughout the distributional
range of the subspecies, I present a comprehensive description of selected meristic
characters, determine the extent of within sample variation, and examine the
possibility of among sample differences. Based on these findings, I examine the
geographic variation in selected meristic characters and resulting population
structure (i.e., the structure of populations across the range). Previous descriptions of
the meristic characters of the subspecies have been limited to relatively small sample
sizes and populations from localized or regional geographic area. The basis for
Behnke's taxonomic description of the subspecies in his book "Native trout of
western North America" (Behnke 1992) was his review of several independent
regional surveys (Schultz 1936; Snyder 1940; DeWitt 1954; Qadri 1959) that were
not uniform in the meristic characters examined and Behnke's own examination of
277 fish from 22 locations (x = 13). Information based on meristic characters may
provide additional insight into geographic population structure based on molecular
markers and provide an additional tool for identifying the presence of individuals
12
with characters intermediate to those of coastal cutthroat trout and steelhead (e.g.,
hybrids).
2.3. METHODS
2.3.1. Sample collection and analysis
A total of 1,589 coastal cutthroat trout were collected from 55 sites from northern
California to Prince William Sound, Alaska (Figure 2.1, Table 2.1). For presentation
purposes the sites were numbered based on their location from north to south. Sites
were located in estuaries or streams downstream of migration barriers. A sample
consisted of individuals identified in the field as coastal cutthroat trout collected
from a common site (location). Samples were typically collected over a period of 1
d, although several sites were sampled in more than one year. The average sample
size was 29 fish (range: 13 to 56) per site (Table 2.1). Fish were collected by
minnow traps, angling, and downstream migrant traps. Attempts were made to
sample several size classes of fish (i.e., age classes) from each population. Two
samples from hatcheries, Humboldt State University Hatchery, California, and Fall
Creek Hatchery (Alsea River), Oregon, were also included in the analysis.
After capture, fish were euthanized (tricane methanesulfonate), weighed (g),
measured (fork length FL, mm), and a photograph (Ektachrome ASA 200) of the left
side was taken. Each fish was tagged with an identification number that was secured
to the lower jaw. Tissue samples for a genetic study done in conjunction with this
study were collected from large fish (> 200 mm) in the field while small fish were
placed directly on dry ice for transport to laboratory where tissues were removed.
Tissue samples were taken from right side of fish. Following tissue removal, fish
13
Figure 2.1. Locations where samples were collected and regional groupings for
analysis. Location numbers correspond to numbers in Table 2.1.
14
Table 2.1. Collection sites, location, number of fish collected (N), and number of fish
determined to be coastal cutthroat trout based on laboratory examination of
basibrianchial teeth and maxillary extension (n). Location numbers correspond to
numbers in Figure 2.1. If no value is given for n, all fish were determined to be
coastal cutthroat trout.
Location
River system/drainage
Collection site
Bosewell Bay, AK
Fish Creek
Martin River, AK
Martin Lake outlet
Location (UTM)
number Zone
E
N
N/n
1
6
544100 6697100
2
6
632050 6696900 25/23
Gines Creek, AK
3
7
643200 6568900
Suntaheen Creek, AK
4
8
493100 6434375 17/0
33
46
Freshwater Bay, AK
Bayhead Creek
5
8
486550 6423650 32/15
Kiag Bay, AK
West Chichagof Lake
6
8
436000 6394300
18
Portage Bay, AK
Portage Creek
7
8
606300 6310550
32
Duncan Salt Chuck, AK
8
8
603350 6304300 52/50
Kadake Creek, AK
9
8
561750 6293000
Eagle River, AK
10
9
339500 6226600 32/3 1
11
9
325150 6202750 50/49
Staney Creek, AK
12
8
620650 6186550
Vixen Inlet, AK
13
8
688000 6185700 34/22
14
9
335200 6174950 27/26
Throne River, AK
15
8
652000 6173850
Bakewell Creek, AK
16
9
394950 6128100 52/36
Wolverine Creek, AK
Traiters Cove, AK
McDonald Lake outlet
Margaret Creek
38
51
62
Skeena River, BC
Herman Creek
17
9
525500 6027000
25
Tiell River, BC
Survey Creek
18
9
290400 5916700
20
19
9
281100 5918600
18
25
Yakoun River, BC
Bella Coola River, BC
Molly Walker Creek
20
9
675550 5811900
Bella Coola River, BC
Fish Creek
21
9
665300 5806900 25/13
San Josef River, BC
22
9
558860 5613190
Willow Creek, BC
23
10
340140 5535680 25/9
25
Sakinaw Lake, BC
Mixal Creek
24
10
425560 5501100 25/14
FraserRiver,BC
Salmon River
25
10
531580 5440750
29
Ritherdon Creek, BC
26
10
358200 5424200
25
Sandhill Creek, BC
27
10
469760 5380850 25/23
15
Table 2.1. Continued
Location
Location (UTM)
River system/drainage
Collection site
Kirby Creek, BC
First tributary
28
10
433300 5360410 28/26
Stillaguamish River,
Church Creek
29
10
550240 5342720
56
Hoko River, WA
Bear Creek
30
10
401290 5332590
25
31
10
450800 5328760
25
Salt Creek, WA
number Zone
E
N
N/n
Quillayute River, WA
Dickey River
32
10
381435 5310896 13/12
Hoh River, WA
Alder Creek
33
10
407145 5299280
25
Hood Canal, WA
Big Beef Creek
34
10
516400 5277240
20
Humptulips River, WA
Stevens Creek
35
10
432330 5242280
25
McClane Creek, WA
West Fork
36
10
500180 5208400
32
Naselle River, WA
Alder Creek
37
10
450895 5147569 26/24
Nehalem River, OR
Foley Creek
38
10
432710 5054300
Trask River, OR
North Fork
39
10
453430 5033540 3 1/30
Sand Creek, OR
Andy Creek
40
10
429150 5017880
SchoonerCreek,OR
41
10
429762 4976525 19/10
Siletz River, OR
42
10
422939 4962940
22
30
21
Yaquina River, OR
Wolf Creek
43
10
443335 4935710
30
Alsea River, OR
Alsea Hatchery
44
10
440360 4916320
30
45
10
411570 4901795
27
Cummins Creek, OR
Umpqua River, OR
W. Br. N. Fk. Smith R.
46
10
429360 4860000
22
New River, OR
Davis Creek
47
10
383880 4758840
32
RogueRiver,OR
Iron Creek
48
10
401960 4717360
33
Winchuck River, OR
Wheeler Creek
49
10
408085 4654600
26
Lake Earl, CA
Jordon Creek
50
10
404300 4627560 28/26
Kiamath River, CA
Waukell Creek
51
10
413370 4594310 25/24
Redwood Creek, CA
May Creek
52
10
414200 4577870
Mill Creek, CA
53
10
403660 4546040 24/12
Widow White Creek,
54
10
406750 4534000 18/9
Humboldt St. Univ., CA Hatchery
55
10
408420 4525640
27
31
16
were fixed in 10% buffered formalin (Humason 1972; Markie 1984) for 1 week prior
to storage in 70% ethanol (ETOH).
Thirteen meristic counts were collected from each fish (Table 2.2). In general,
methods followed those of Hubbs and Lagler (1958). All counts were made on the
left side of fish. Vertebrae, dorsal fin ptergyiphores, and anal fin ptergyiphores were
counted from radiographs. Radiographs were taken with a cabinet x-ray machine
(Fixatron 804) on 20.3 cm X 25.4 cm x-ray film (Agfa Structurix CD4, daylight
wrapped, industrial x-ray film, ISO 9001) developed with a Kodak RP-OMAT
Processor (Model M6B). The shelf height (55.88 cm) and power (25.0 Kvp) were
held constant for all fish, exposure time varied with fish size ranging from 30
seconds for small fish (80 mm) to 5 minutes for larger fish (>300 mm). Radiographs
were placed on a light table for counting. Vertebrae counts included all vertebra
separated by sutures including the upturned or ural centra, but excluding the
urostyle. Dorsal and anal pterygiophores were scored as the number of distinct,
extended pterygiophores, excluding the anterior short pterygiophore when it was less
than half the length of the second most anterior pterygiophore.
Pyloric caeca were counted after fish had been transferred to 70% ETOH. All
caeca found posterior of stomach were counted after the intestine had been unwound
from the stomach and laid out (Hubbs and Lagler 1958; Behnke 1992). A dissecting
microscope was used to count caeca from fish < 150 mm. Sex determination was
done at the time caeca were counted.
Lateral line series counts were made from scales located two scale rows above
(dorsal) the lateral line starting at the first row immediately behind the pectoral
girdle and the last row being the one crossing the lateral line at the posterior edge of
the hypural plate (Strauss and Bond 1990). Scales above the lateral line were
Table 2.2. Summary of meristic counts from all samples of 0. c. clarki and fish designated as 0. mykiss from 54 sites.
0. C. clarki
Character
Fork length (mm)
N
1400
Mean
171.4
81.0
Weight(g)
1238
112.2
Anal pterygiophores
1430
Dorsal pterygiophores
Vertebrae
0. mykiss
52
Max
468
210.0
2
11.2
0.6
1429
11.5
1421
SD
N
SD
Mm
98
Mean
107.4
50.1
45
Max
409
2505
97
17.9
19.7
1
110
9
13
97
11.5
0.7
10
13
0.7
10
13
97
12.3
0.7
11
14
61.4
1.0
56
66
98
63.3
1.6
60
67
1429
144.9
10.9
114
187
97
124.6
7.7
104
150
1427
32.7
2.9
20
42
98
27.0
2.5
22
37
1422
9.1
0.4
8
10
98
9.3
0.5
9
11
1419
13.6
0.7
7
15
97
13.9
0.7
12
15
Gill rakers lower arch'
Gill rakers upper archa
1425
11.5
0.8
8
14
96
12.0
0.7
10
13
1426
6.7
0.8
3
12
96
7.0
0.8
5
9
Gill rakers total
1425
18.2
1.2
13
23
95
19.0
1.1
17
22
1417
11.0
0.7
9
13
93
11.3
0.8
9
13
43.3
10.8
6.9
7.5
19
74
48
77
98
44.2
0.0
7.2
0.0
24
0
59
0
Scales in lateral
seriesa
Scales above lateral
Pelvic fin rays'
Pectoral fin
Branchiostegal
raysa
raysa
linea
Pyloric caeca
1345
Basibranchial teeth
1431
a
Bilateral characters counted on left side of fish.
Mm
0
counted from the origin of the dorsal fin downward along one of the diagonal scale
rows to, but not including, the lateral line scale (Hubbs and Lagler 1958). Malachite
green stain (1 g stainll0O ml H20) was applied to the scales just prior to counting
and a dissecting microscope (6X 40X) was used. The length of the maxillary was
recorded on a scale from 0 to 2 (Figure 2.2). If the maxillary did not extend to the
anterior portion of the eye, it was recorded as a 0. If the maxillary extended to
somewhere between the anterior and posterior portion of the eye it was recorded as a
1, and if the maxillary extended beyond the posterior portion of the eye, it was
recorded as a 2.
Alizarin red stain (1 g stain/100 ml H20) was applied twelve hours prior to
counting paired fins, gill rakers, basibranchial teeth, and branchiostegal rays,. Fish
were placed in sealable plastic bags with paper towels saturated with 70% ETOH
until counts were made. Counts of paired fins (pelvic and pectoral), gill rakers,
basibranchial teeth, and branchiostegal rays were done with a dissecting microscope.
Counts of gill rakers were done on the first arch. Separate counts were made of the
lower and upper limbs of each arch with gill rakers straddling the angle of the arch
included in the count of the lower limb (Hubbs and Lagler 1958). Compressed air
directed through a large gauge hyper-dermic needle was used to remove tissue and
mucus that often obscured the basibranchial teeth and gill rakers.
2.3.2. Data analysis
During the initial stages of screening for data entry errors it became evident that
several samples contained 0. mykiss (steelhead or rainbow trout) misidentified as
19
Figure 2.2. Maxillary extension scores for coastal cutthroat trout.
20
coastal cutthroat trout2. In addition, many samples contained individuals with
meristic character counts intermediate to counts expected for steelhead and coastal
cutthroat trout that were reported in the literature. The observation of counts
intermediate to those of steelhead and coastal cutthroat trout would not be
inconsistent with hybridization (Behnke 1992). The two characters most commonly
used to differentiate between coastal cutthroat trout and steelhead are basibranchial
teeth and the maxillary extension (Table 2.3). In addition to these two characters, the
presence of a yellow, orange, or red line in the skin folds of the lower jaw is often
used as an identifying character of coastal cutthroat trout. Of these three characters,
the maxillary extension and the presence of the colored line on the lower jaw are
often used for field identification. Meristic counts most useful for distinguishing
between steelhead and coastal cutthroat trout are the number of vertebrae and
various scale row counts, but these counts are difficult to determine in the field. Prior
to analysis, a decision rule was established to identify steelhead individuals and
remove them from the analysis. The length of the maxillary and the absence of
basibranchial teeth were used to identify steelhead. Individuals with a maxillary
extension score of 0 or 1 (maxillary did not extend past the posterior portion of the
eye, see Figure 2.2) and an absence of basibranchial teeth were designated as a
steelhead. An individual with a maxillary extension score of 2 or the presence of
basibranchial teeth was designated as a coastal cutthroat trout. These two characters
were not used for subsequent analyses.
Since the meristic counts are discrete, analysis of frequency distributions was by
contingency j on untransformed data to determine if there were differences in
2
For purposes of this study, all 0. mykiss collected will be referred to as steelhead
since all samples were obtained downstream of migration barriers, although some
nonanadromous 0. mykiss (rainbow trout) may be included.
21
Table 2.3. Published characteristics of basibranchial teeth and maxillary extension
used to distinguish coastal cutthroat trout (0. c. clarki) and steelhead (0. mykiss).
Basibranchial teeth
Citation
0.
Maxillary extension
0. c. clarki
0. mykiss
"..extending to
point well behind
posterior margin
of eye.."
"..extending to
point below, or
slightly behind,
posterior margin
of eye.."
"..jaw usually
extends to well
behind a vertical
through the
posterior margin
of the eye"
in small fish
extends to or
slightly beyond
a vertical
through the
posterior margin
of the eye .."
absent
". .cutthroat may
be distinguished
by ... longer
maxillary..,
..reaching slightly
past orbit. ."
"..maxillary..
extending far
beyond orbit.."
usually present
absent
"..maxilla reaches
behind the hind
margin of the
eye."
Miller and Lea 1972
present
absent
Moyle 2002
present
absent
c. clarki
0. mykiss
Behnke 1992a1
always present
absent
Clemens and Wilby
present, "well
developed"
absent
1961
Dymond 1932
sometimes
absent, but,
whenever found,
serve to
distinguisth the
cut-throat from
either the
Kamloops or
steelhead."
Eddy and Underhill
"always present"
absent
present
absent
"..
"..
1982
Eschmeyer et al. 1983
Evermann and
Goldsborough 1 907b
McPhail and Lindsey
1970
"Cutthroat trout
in general have
larger mouths
(longer maxillary
bones).. .than
rainbow trout."
"..the maxillary
bone usually
extending
behind the
eye.."
22
Table 2.3. Continued
Basibranchial teeth
Citation
0.
c. clarki
0. mykiss
0. c. clarki
0. mykiss
Pollard et al. 1997
present
absent
"Maxillary
extends past rear
margin of eye on
fish >80mm."
"Maxillary does
not extend past
back margin of
eye of parr."
Scott and Crossman
usually present
absent
"..maxillary long,
at least to
posterior margin
of eye, often far
beyond.."
"..maxillary long,
usually passing
eye.."
usually present
absent
"Maxillary
extends well
beyond posterior
margin of eye.."
"Maxillary
extending
beyond posterior
margin of eye
only in large
specimens (>20
inches).."
1973
Wydoski and Whitney
1979
a
Maxillary extension
Behnke (1992) recognizes this the subspecies Oncorhynchus mykiss irideus as the
coastal rainbow trout from California to Alaska (both steelhead and nonanadromous
populations)
Evermann and Goisborough (1907) recognized both steelhead (Salmo gairdneri
Richardson) and Alaskan rainbow trout (Salmo irideus Gibbons).
23
merisitic counts within a sampling site between males and females and among fish
of various lengths within a sampling site. To avoid sparse cells in the j analysis,
fish within a sampling site were pooled into five size groups ( 100 mm, 101
200
mm, 201 300 mm, 301 400, > 400 mm). To test for differences in meristic counts
among size groups within a sampling site, a sample needed to consist of at least three
size groups with five or more individuals.
Phenetic affinities and relationships among samples were examined with several
multivariate statistical approaches using the eleven meristic characters described
previously (see Table 2.2). To determine if differences existed among sampling sites
a multivariate analysis of variance (MANOVA) was used with sample site as the
treatment variable. The null hypothesis of no significant differences among sampling
sites was tested with the Wilk's A. statistic. Data for the MANOVA were transformed
using Gower's method of ranging (Sneath and Sokal 1973) where the smallest value
for a character is subtracted from each value and the result is divided by the range.
Individual meristic characters were further examined with analysis of variance
(ANOVA) with each of the 11 meristic characters as a dependent variable.
Differences among sampling sites for the median of a given character were tested
with multiple comparisons with a Kruskal-Wallis' H test. Range-transformed data
were used for the Kruskal-Wallis' H tests with a Bonferoni experiment-wise multiple
comparison test to minimize the possibility of type I error.
Canonical variates were generated by discriminant analysis to remove correlation
among characters and provide a weighted combination of characters that maximized
distinction among locations. A canonical score was derived for each row (i.e.,
individual fish) in which all independent variables values (i.e., eleven meristic
characters) were counted. For each population, the mean of the first canonical variate
score was calculated, and these means were plotted to examine differences among
populations. I used a clustering and an ordination approach to further examine
patterns of geographical similarity. An Euclidian distance matrix created from the
range-transformed data was used to construct a dendrogram from cluster analyses of
pair-wise differences between samples using the unweighted pair-group method with
arithmetic averages (UPGMA) algorithm (Sneath and Sokal 1973). A non-metric
multidimensional scaling analysis (MDS) (Kruskal 1964a; Kruskal 1964b) was used
to create a plot of the locations in a three-dimensional configuration space of the
Euclidian distance matrix created from the range transformed data.
2.4. RESULTS
2.4.1. Identification of steelhead
Of the 1,589 fish examined, 98 steelhead were identified based on the absence of
basibranchical teeth and a short maxillary extension (Table 2.4; Appendix Table
A2. 1). In addition, fish with meristic character counts intermediate to that typically
found in coastal cutthroat trout and steelhead were also found in the samples. One
sample was determined to contain only steelhead (Suntaheen Creek, Alaska, location
4) and was not used for subsequent analyses except as an outgroup. Mean counts of
scales in lateral series, scales above the lateral line, and basibranchial teeth of coastal
cutthroat trout were greater than mean counts for steelhead (Table 2.4; Appendix
Figures A2. 1
2.6; Appendix Table A2.2). The absence of basibranchial teeth was
used as a character for steelhead therefore the counts were less by definition. All
other mean character counts were less for coastal cutthroat trout than for steelhead. If
fish that exhibited only one of the two characters that were used to designate coastal
Table 2.4. Summary of meristic counts from all samples of 0. c. clarki and fish designated as 0. mykiss or hybrids from 54
sites. Fish designated as hybrids did not have basibranchial teeth or had a short maxillary extension, for purposes of most
analyses in this study these fish were included with 0. c. clarki (including summary counts in Table 2.2).
0. c.
64
Max
468
"Hybrid"
Mean SD
Mm
226 116.4 62.4
52
3
2505
188
13
226
40.8
11.3
121.0
9
0.7
10
13
225
61.4
1.0
56
65
1205
146.1
10.6
120
Scales above lateral linea
1204
33.0
2.9
Pelvic fin rays'
1197
9.1
Pectoral fin raysa
1196
Gill ralcers lower arch°
clarki
Character
Fork length (mm)
Mean
1174 182.0
Weight(g)
Anal pterygiophores
1050 124.9 219.7
1204 11.2
0.6
Dorsal pterygiophores
1204
11.5
Vertebrae
1196
N
SD
Mm
N
0.
Max
mykiss
433
N Mean SD
98 107.4 50.1
2
926
97
17.9
0.6
10
13
97
11.6
0.7
10
13
225
61.4
1.1
59
187
224
138.3
9.7
20
42
223
31.5
0.4
8
10
225
13.6
0.7
7
15
1203
11.5
0.8
8
Gill rakers upper archa
1203
6.7
0.7
Gill rakers total
Branchiostegairaysa
1203
18.2
1193
11.0
45
Max
409
19.7
1
110
11.5
0.7
10
13
97
12.3
0.7
11
14
66
98
63.3
1.6
60
67
114
173
97
124.6
7.7
104
150
2.8
25
39
98
27.0
2.5
22
37
9
0.3
8
10
98
9.3
0.5
9
11
223
13.6
0.6
12
15
97
13.9
0.7
12
15
14
222
11.4
0.7
9
13
96
12.0
0.7
10
13
4
9
223
6.7
0.8
3
12
96
7.0
0.8
5
9
1.2
13
22
222
18.2
1.2
14
23
95
19.0
1.1
17
22
0.7
9
13
224
11.1
0.8
9
13
93
11.3
0.8
9
13
Pyloric caeca
1129 43.8
6.8
20
Basibranchial teeth
1205 11.6
7.5
1
a
Bilateral characters counted on left side of fish.
74
48
216
226
40.6
6.5
5.8
19
59
36
77
98
44.2
0.0
7.2
0.0
24
0
59
Scales in lateral
series1
79.8
6
0
Mm
0
c/I
26
cutthroat trout (i.e., the presence of basibranchial teeth or a maxillary extension past
the eye) were separated out from fish designated as either coastal cutthroat trout and
steelhead, all but one character were either similar to mean counts for coastal
cutthroat trout or intermediate to mean counts of coastal cutthroat trout and steelhead
(Table 2.4; Appendix Figures A2. 1 - 2.6). The mean count for pyloric caeca was the
exception, the mean count for fish with only one of the two discriminating characters
was less than the mean counts for both coastal cutthroat trout and steelhead (Table
2.4; Appendix Figure A2.5). Fish designated as steelhead were not used for
subsequent analyses. Fish with only one of the two characters used to designate
coastal cutthroat trout were kept in subsequent analyses as coastal cutthroat trout.
Although one might consider these individuals to be hybrids since all but one of the
mean counts were intermediate or similar to either coastal cutthroat trout or
steelhead, it is unclear if these fish are hybrids and sampling was not carried out in
such a way to adequately address this issue.
2.4.2. Differences in meristic counts between sex and among length groups within
sampling locations
Significant differences (P < 0.05) were detected for 14 of the 594 comparisons
(2.3%) between meristic character counts of males and females within a sampling
location. Given the number of comparison tests made (54 samples X 11 characters =
594 tests), the number of significant tests was far fewer than would be expected by
chance and therefore males and females within a sampling location were pooled into
a single sample. Six samples had sufficient numbers of fish in at least three size
groups to test for differences in meristic count among size groups (Fish Creek, AK,
location 1; Kadake Creek, AK, location 9; Wolverine Creek, AK, location 11;
27
Throne River, AK, location 15; Bakewell Creek, AK, location 16; and Siletz River,
OR, location 42). Only one significant difference (P <0.05) was detected among
size groups and meristic character counts within these six locations. Anal
pterygiophores (x2 = 19.21, df = 8, P
0.0 14) among size groups at Wolverine
Creek, Alaska (location 11), were significantly different. Given the number of
comparison tests made (six locations X 11 characters = 66 tests) I assumed that there
were no significant differences between size groups in the populations examined.
Based on these results, all sizes of fish within a sampling location were pooled into a
single sample.
2.4.3. Differences among sampling locations
There were significant differences in the meristic counts (MANOVA, Wilks' A =
0.0695, P < 0.001) among all locations (Figures 2.3 2.8). In addition, there were
significant differences detected among all locations at each of the 11 meristic
characters (ANOVA, P < 0.01). No linear clinal trends with latitude were apparent,
although there appears to be a undulating pattern with latitude for all of the mean
character counts, being most pronounced with the scale row counts (i.e., scales in
lateral series and scales above the lateral line - Figure 2.5). These patterns were not
consistent across character types in terms of increasing and decreasing at the same
geographical region nor were they consistent in the direction (i.e., greater or lesser
values) at the same geographical region. Anal pterygiophores, vertebrae, both scale
row counts, and pyloric caeca had counts that were generally higher for the northern
locations than for the southern locations. Branchiostegal ray counts were the only
character that exhibited greater mean counts at the southern locations. Samples from
contiguous locations were not necessarily more similar than samples from
12.0
11.5
I
r:I
I
1H I
I
1
1
'''1
Lht++{t
JJ'
I
i'H1
I
f
H?
H
I
10.5
10.0
12.0
rI
11.5
11.0
10.5
IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIITTTTTrtIIIIJIIIiijII
1357 91113151719212325272931333537394143454749515355
Location
Figure 2.3. Mean (± 1 SE) counts of anal pterygiophores and dorsal pterygiophores
for each sampling location of coastal cutthroat trout. Location 4 contained only
steelhead and therefore is not included in figure. Location numbers correspond to
numbers in Table 2.1.
29
631
t
4
+
62
61
f
f
60
59
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
I
4
20
f
is
+
f
5
If
f
W
f
0
III
111111111111111111
liii 11111111 liii
35 7 91113151719212325272931333537394143454749515355
I
1
I
I
Location
Figure 2.4. Mean (± 1 SE) counts of vertebrae and basibranchial teeth for each
sampling location of coastal cutthroat trout. Location 4 contained only steelhead and
therefore is not included in figure. Location numbers correspond to numbers in Table
2.1.
30
621ff
ff
61
f
15
10
I
0
I
1
I
I
I
I
I
I
I
I
I
35 7 9 1113 15 17 19 21 2325 2729 31 3335 373941 4345 4749 51 53 55
Location
Figure 2.5. Mean (± 1 SE) counts of scales in lateral series and scales above lateral
line for each sampling location of coastal cutthroat trout. Location 4 contained only
steelhead and therefore is not inëluded in figure. Location numbers correspond to
numbers in Table 2.1.
31
'
t
621ff
.
ti,
I
I
If
f
tff
11
61
15
11
+
10
+
1
ff
I
f
0
IIIIIIIII!IIIIlIJuIIIIIIIIJIjIIIII!IIIIIIIJIIIIIlIIu
1
35 7 9 1113 15 17 19 21 23252729 31 333537 3941 4345 4749 51 53 55
Location
Figure 2.6. Mean (± 1 SE) counts of pelvic fin rays and pectoral fin rays for each
sampling location of coastal cutthroat trout. Location 4 contained only steelhead and
therefore is not included in figure. Location numbers correspond to numbers in Table
2.1.
32
13
12
I
o
11-i
ft
I
f
I
io-I
+
50
f
ff
f
C.)
+C.)
0
o
+
+
+,
f
40
*
30
135791113151719212325272931333537394143454749515355
Location
Figure 2.7. Mean (± 1 SE) counts of branchiostegal rays and pyloric caeca for each
sampling location of coastal cutthroat trout. Location 4 contained only steelhead and
therefore is not included in figure. Location numbers correspond to numbers in Table
2.1.
33
+1
Q
V
1
35 7 9 1113 15 17 1921 23 25272931 33 35373941 43 4547 49 51 53 55
Location
Figure 2.8. Mean (± 1 SE) counts of upper gill rakers and lower gill rakers for each
sampling location of coastal cutthroat trout. Location 4 contained only steelhead and
therefore is not included in figure. Location numbers correspond to numbers in Table
2.1.
geographically discontinuous locations. Although, there were samples that were
most similar to samples from geographically adjacent locations for a specific
character. For example, vertebrae counts (Figure 2.4) were similar between locations
1 and 2; locations 6, 7, 8,and 10 were similar. Conversely, locations 1 and 2 were not
similar in their mean count of scales above the lateral line, and locations 6, 7, 8, and
10 were not similar in their mean count of scales above the lateral line (Figure 2.5).
The total variation accounted for by the first four variates in canonical analysis
was 71%. The first variate accounted for 28.1% of the variation and had an
eigenvalue of 0.91. The second, third, and fourth variates accounted for 16.0%,
15.6%, and 11.2% respectively (eigenvalues of 0.52. 0.51, and 0.37). The
eigenvaiues for these four variates were statistically significant (Wilks' i test). The
characters of primary importance in distinguishing between locations in the first
canonical variate were scales above the lateral line count (7.56) and pyloric caeca
count (-2.80). A plot of the mean of canonical variate I for each location is not unlike
several of the plots of mean character scores (Figure 2.9). Locations at the north of
the range tend to have higher values for these two characters than locations at the
south end of the range while values for the locations in the central part of the
distributional range are variable, with locations along the coast of Washington and
south to the Umpqua River having values similar to the most northerly locations
(locations 32 though 46). The California locations (locations 50 though 55) represent
six of the lowest eight values, the other low scores are for Staney Creek, Alaska
(west coast of Prince of Wales Island, location 12) and Salmon River (tributary to
the Fraser River, British Columbia, location 25). These low scores for the California
locations coincide with low counts for the scale row counts, conversely, locations at
the northern end of the range tended to have higher scale row counts (Figure 2.5).
35
-
I
I
0
0
-1
-2
-3
-4
1
3 5 7
9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53 55
Location
Figure 2.9. The mean (±1 SE) canonical variate I score for each sampling location
based on 11 meristic characters. Location 4 contained only steelhead and was
excluded. Location numbers correspond to numbers in Table 2.1.
36
There was a lack of agreement between the populations that tended to be very
different in terms of the number of significantly different medium counts in pairwise
comparisons for meristic characters (Appendix Table A23) and the pattern observed
in the plot of canonical variate I means (Figure 2.9). That is, the samples with the
largest number of significant differences did not standout as being very different
based on canonical variates generated by discriminant analysis that removed
correlation among the characters and provided a weighted combination of characters
that maximized distinction among locations. Gines Creek, Alaska (location 3) had
the most number of significant different mediums (168, 29% of the comparisons) yet
it was not dissimilar to other locations in its geographic area.
Since the canonical variate plot was only for variate I, differences in score were
heavily influenced by characters that were heavily weighted in the variate (scales
above the lateral line and pyloric caeca). Locations with many significant differences
in mediums for meristic characters are not captured by the canonical variate unless
those differences are for characters that strongly influence the variate (e.g., 106
differences Bakewell Creek, Alaska (16), 101 differences Kirby Creek, British
Columbia (28), 106 differences- Stillaguamish River, Washington (29), and 102
differences Winchuck River, Oregon (49); see also Appendix Table A2.3).
Locations at the southern portion of the range (locations 50 - 55) differed from
populations to the north as measure by canonical variate I resulting primarily from
the lower scale row counts for fish in southern populations (Figure 2.5). Conversely,
populations at the northern portion of the range tended to have higher scale row
counts and therefore higher canonical variate I scores. There was a general trend of
decreasing canonical variate I scores from north to south except for 10 of 14
populations that were between the Quillayute River, Washington (location 32) south
37
to the Umpqua basin (location 46) that tended to have canonical variate I scores
similar to locations at the north end of the range (locations 1
5 and 7 11). This
pattern where counts from locations from the Washington coast and the north and
mid-Oregon coast had counts similar to locations to the north was also evident in
several of the mean counts of the meristic characters (Figures 2.3 2.8). A regression
between mean canonical variate I and latitude (i.e., UTM northern coordinate)
exhibited a slight, but significant decrease in value with locations to the south
(r2 =
0.115; P = 0.012; Figure 2.10). With the southern locations removed there was no
significant relationship between latitude and canonical variate I
(r2
= 0.002; P =
0.93).
The phenetic affinities illustrated by the phenograin (Figure 2.11) capture the
relative distinctiveness of the California locations (locations 50 55) evident in the
canoncial variate pilot (Figure 2.9). Five of the seven samples from the Winchuck
River, OR, south to Widow White Creek, CA (locations 49 54), and including the
hatchery population from Humboldt State University (location 55) clustered together
along with Willow Creek, British Columbia (location 23), and formed a distinct
cluster from all the other samples (see shaded region of phenogram, Figure 2.11).
Moreover, the long branch lengths (i.e., large Euclidian distances) within the shaded
region of the phenogram (Figure 2.11) indicate large intra-regional differences
among populations at the southern extend of the species' distributional range. That
is, the southern populations tended to exhibit phenetic affinity corresponding to
geographic proximity in the cluster analysis, but within this regional grouping, the
differences among populations were greater than within other clusters. The Willow
Creek sample (location 23) exhibited meristic characters similar to those of steelhead
and may be a result of hybrids being included in the sample. Of the 25 fish sampled
3
2
46
Ij,
4j37
-
1
38.53
2
8
%4i
0
4Ø'24
Q
.)
----
.49
o
-.1-
.go
,.6S22
L
2
19
.0140
.17
.6
12
.52
2I
0
55
0
54
-34
4
7
UTM northern
Fiure 2.10. Mean canonical variate I versus latitude (UTM northern coordinate; X
10 ) for coastal cutthroat trout from 54 locations. The dashed regression line was
calculated with locations 50 to 55 included, the solid regression line was calculated
without locations 50 to 55. Location numbers correspond to numbers in Table 2.1.
Rgigs
A46
1H
u____
i21
____________________F
OO
O2
0.3
0L4
O.5
Eudidian distance
Figure 2.11. Phenogram from UPGMA cluster analysis of Euclidian distance based on 11 meristic characters of
coastal cutthroat trout from 54 locations, shaded region of the phenogram includes populations from southern
portion of range with large intra-regional differences. Location numbers correspond to numbers in Table 2.1.
from Willow Creek, 14 were misidentified (i.e., initially identified as coastal cutthroat
trout in the field, but lacking basibranchial teeth and having a short maxillary
extension), nine were designated as coastal cutthroat trout, and two could not be
assigned since they were missing a count of basibranchial teeth or a measure of the
maxillary extension (Appendix Tables A2. 1 and A2.2). The Willow Creek sample had
the second lowest count of scales in lateral series
(x
= 131.8, Figure 2.5, Appendix
Table A2.2) and the fifth lowest count of scales above the lateral line
(x
= 30.0, Figure
2.5, Appendix Table A2.2). These low scores are similar to values for locations at the
southern portion of the range, in addition to being similar to steelhead, and appear to
be the major feature of this distinct cluster. It is difficult to make many additional
inferences regarding phenetic affinities from the phenogram. There are a few instances
where geographically adjacent locations show phenetic affinity. A group of locations
in the greater Ketchikan, AK, area (Wolverine Creek location 11, Vixen Inlet
Margaret Creek (Traitors Cove)
13,
14, Throne River 15, and Bakewell Creek 16)
clustered together. The Siletz River (42) and Schooner Creek (41) clustered together as
did the Salmon River (25, Fraser River tributary) and Church Creek (29, Stillaguamish
River tributary).
As in the cluster analysis, with the exception of regional structuring at very large
spatial scales that separated out the southern populations, only weak geographic
concordance was evident in the MDS analysis. The phenetic affinity for the southern
locations evident in the canonical variate and cluster analyses was also evident in the
multidimensional scaling (Figure 2.12). The stress value of 0.082 of the MDS analysis
is considered good, a value of 0.05 would be considered excellent, 0.0 perfect, and 0.2
poor (Kruskal 1964a). This statistic measures the goodness of fit of of the and (Sneath
and Sokal 1973). All locatiOns from Jordon Creek, California (location 50) south were
41
0.97
0.41
[II
-0.1
-0.7
1.76
Figure 2.12. Multidimensional scaling (dimensions 1,11 and III) of Euclidian distance
based on 11 merjstic characters of coastal cutthroat trout from 54 locations. The stress
value for this clustering was 0.082. Location 4 contained only steelhead and was
excluded. Location numbers correspond to numbers in Table 2.1.
42
grouped together. As in the cluster analysis (Figure 2.11), Willow Creek, British
Columiba (location 23), had a phenetic affinity with locations in the southern portion
of the range. These southern locations along with Willow Creek formed the most
distinctive grouping in the MDS analysis. The first location south of the Rogue River,
the Winchuck River, Oregon (location 49) shared some phenetic affinity to southern
group to locations (i.e., is adjacent in MDS space). With the exception of the Rogue
River sample (Iron Creek, location 48), the distribution of these southern populations
is consistent with other salmonid studies that have found a zoogeographic break in the
area around Cape Blanco, Oregon (Hatch 1990; Weitkamp et al. 1995; Busby et al.
1996). The MDS analysis reveled phenetic distinctiveness of the Alsea hatchery
(location 44) sample and the Gines Creek (location 3) sample. The Alsea hatchery
sample exhibited the largest SE of the mean of canonical variate I of all the samples
(Figure 2.9).
2.5. DISCUSSION
Significant variation of meristic characters was found among 54 sample sites
consisting of 1,491 individual coastal cutthroat trout from across their distributional
range. This survey was based on samples collected over a five year period, providing a
snap shot of the meristic variation that existed from 1992 to 1996 and minimizing
temporal environmental effects that may have large influences on salmonid production
(Pearcy 1992) and are quite dynamic across the range of coastal cutthroat trout (Pearcy
1997). In the most comprehensive survey previous to this work, Behnke (Behnke
1992) suggested that anadromous coastal cutthroat trout were morphologicaily similar
throughout their range and exhibited no clinal variation in characters between northern
and southern populations. Behnke's review was based on his collections along with
3]
data from others (Schultz 1936; Snyder 1940; DeWitt 1954; Qadri 1959) that were
collected over many years. The number of meristic elements in the phenotype can be
influenced by the magnitude, timing, and duration of environmental factors (Beacham
1985), even minor changes in temperature, salinity, light, and dissolved gasses during
ontogeny can result in significant differences in meristic counts within the same
species (Tâning 1952).
The mean values for the various characters are similar to those reported by Behnke
(1992), although Behnke does report some values that were much different than
observed in my study. For instance, the highest number of basibranchial teeth observed
by Behnke was a mean of 29 from Lake Sutherland, Washington, about twice as many
as he found in any other sample. The highest mean count of basibranchial teeth from
my study were 20.4 from Throne River,'Alaska, 20.6 from Yakoun River, British
Columbia, and 22.4 from Staney Creek, Alaska. The mean count of 144.9 for scales in
lateral series is at the lower end of the range of 140 to 180 reported by Behnke (1992).
Although some of the individuals in my sample may have been hybrids based on the
absence of basibranchial teeth, the mean count with only individuals with
basibranchial teeth was 146.8, still on the low end of the range reported by Behnke.
There was no latitudinal dine detected for any of the meristic characters examined.
This is in contrast to the general pattern of an increasing number of elements for most
meristic characters in fish with increasing distance from the equator (Hubbs 1926;
Tâning 1952; Beacham 1985). The number of elements for a particular meristic
character is influenced by the development rate of the embryo with longer
developmental times resulting in a greater number of elements (Hubbs 1926), northern
areas of the coastal cutthroat trout range experience cooler rearing temperatures than
southern areas and therefore would he expected to have higher numbers of elements.
!YiI
Slower developmental rates in salmonids have been associated with higher numbers of
vertebrae in embryos (Kwain 1975; Beacham 1985).
Shepherd (1991) described meristic characters as phenotypic expressions of
environmental conditions during egg and larval development, adequate temporal and
spatial heterogeneity form the basis of population differentiation using these
characters. In my study, several of the individual characters exhibited a slight clime
across the range that was interrupted by counts from the Quillayute River, Washington
(location 32) south to the Umpqua basin (location 46). Counts from these locations
along the Washington coast south to the mid-Oregon coast were similar to the higher
mean counts found at the northern portion of the range.
Significant differences were found among all populations examined in this study,
although phenetic clustering and ordination exhibited very weak geographic
concordance. Meristic characters were not able to describe intra-specific population
structure at small spatial scales. With just a few exceptions such as the Ketchikan area
of southeast Alaska (locations 11,13 16), populations from the Winchuck River south
(locations 49 55), Schooner Creek and Siletz River (locations 41 and 42), and the
Salmon and Stillaguamish rivers (locations 25 and 29)), there was little phenetic
affinity even among adjacent locations based on the cluster analysis and MDS analysis
of Euclidian distances.
Others have found meristic characters to be useful for delineating population
structure in salmonids at relatively small scales (Smith 1969; Dempson and Misra
1983; Schreck et al. 1985). The general lack of geographic concordance in this study
may be a result of the large geographic range surveyed that encompassed a large range
of environmental conditions and gradients. My findings do suggest that caution should
be used when inferring population structure based on samples from spatially small,
45
local, or intra-regional studies to larger spatial scales. At the spatial scale surveyed in
this study, meristic characters provided information about population structure at only
large spatial scales. The populations at the southern portion of the range had a
relatively strong phenetic affinity. This did not appear to be a result of an overall
latitudinal dine across the range, but a result of coastal cutthroat trout at the southern
extent of the range having similar meristic characteristics. The clustering of locations
observed for the southern Oregon and Northern California locations was consistent
with findings from other studies and status reviews of anadromous salmonids that
detected a zoogeographic break in the area around Cape Blanco, Oregon (Hatch 1990;
Weitkamp et al. 1995; Busby et al. 1996; Johnson et al. 1999).
Although my results are not inconsistent with these previous studies, another
interpretation may be that there is a zoogeographic break for coastal cutthroat trout in
the region between the Rogue River and the Klarnath River. The separate clustering of
sampling locations south of the Winchuck River (locations 50 55) and the inclusion
of the Winchuck River (location 49) and Rogue River samples (location 48) with a
larger non-southern group in the MDS analysis supports this break (Figure 2.12). This
is a region of zoogreographic transition. The Rogue River is the southern limit of
distribution of the reticulate sculpin (Cottus perplexus), a species in virtually every
stream in western Oregon and Washington (Minckley et al. 1986; Moyle 2002), and
the Rogue River contains the same saltwater dispersant species that are found in the
lower Kiamath (Moyle 2002). Coastal Oregon streams south of the Rogue River
(Hunter Creek, Pistol Creek, Chetco River, Winchuck River) do not have inland fish
species like those found in the Rogue and Kiamath systems (Minckley et al. 1986).
The Kiamath drainage supports many unique fishes, particularly above Kiamath Falls
where the fish fauna is much different than that found in the lower Kiamath River, and
both the Sacramento and Columbia rivers (Minckley et al. 1986). No other patterns of
phenetic affinity at a regional scale were detected from my samples of coastal cutthroat
trout. The usefulness of meristic characters to delineate regional clusters has been
observed in other studies. Beacham (1985) found similar results for sockeye salmon
collected from North America; differences in vertebrae number and gill rakers were
only useful to identifying stocks of sockeye salmon on a broad regional basis.
For western trout species, meristics has been used for inter-specific comparisons
and systematics studies (Schreck and Behnke 1971; Gold and Gall 1975; Gold 1977;
Gold and Gall 1981). Although significant differences among locations for meristic
characters were detected in my study, coastal cutthroat trout maintained a phenetic
affinity when compared to other species of western trout based on nine meristic
characters that included anal pterygiophores (anal fin rays for historical counts, dorsal
pterygiophores (dorsal fin rays for historical counts), vertebrae, scales in lateral series,
pelvic fin rays, pectoral fin rays, branchiostegal rays, total gill rakers on the first arch,
and pyloric caeca; Table 2.5). Coastal cutthroat trout from this study grouped together
when the MDS scaling was used to compare them to steelhead from Alaska (Suntaheen
Creek, Alaska, this study), 0. mykiss from the Deschutes River (Currens 1987), 0. gila
(Gila trout (Miller 1950)), 0. chrysogaster (Mexican golden trout (Needham and Gard
1964)), 0. aguabonita (California golden trout (Gold and Gail 1975)), 0. apache
(Apache trout (Miller 1972)), 0. c. pleuriticus (Colorado cutthroat trout (Gold et al.
1978)(Gold et al. 1978)), and Redband (Gold 1977). The 0. mykiss samples grouped
together and exhibited some phenetic affinities with the Colorado cutthroat trout,
Apache trout, and California golden trout (Figure 2.13). The Gila trout, Mexican
golden, and Redband samples were separate from the other clusters and were much
different than the 0. mykiss, Colorado cutthroat trout, Apache trout, and California
Table 2.5. Summary of mean meristic counts for various species of trout from western North America reported in the
literature and mean meristic counts for steelhead from Suntaheen Creek, Alaska, collected in this study. Meristic characters
are: A: anal pterygiophores (fin rays for historical counts), B: dorsal pterygiophores (fin rays for historical counts), C:
vertebrae, D: scales in lateral series, E: pelvic fin rays, F: pectoral fin rays, G: branchiostegal rays, H: total gill rakers, first
arch, I: pyloric caeca (see text for description of character counts).
Meristic character (mean)
Species
Location
A
B
C
D
E
F
G
H
I
Source
0. mykiss
Suntaheen Creek, AK
11.4
12.4
64.5
125.3
9.5
14.5
11.8
20.4
46.3
this study
0. mykiss
Deschutes River, OW
12.5
13.0
63.4
140.2
9.8
142
11.8
19.1
49.7
Currens (1987)
0. mykiss
Deschutes River, ORb
12.5
12.9
64.1
145.5
9.9
14.0
11.2
19.9
40.4
Currens (1987)
Gila River, NM
10.3
10.7
60.2
141.2
9.6
13.8
10.2
19.0
34.9
Miller (1950)
Arroyo de la Rana,
Chihuahua, Mexico
10.1
10.7
56.9
142.0
9.0
13.3
8.9
17.4
20.2
Needham and Gard (1964)
0. aguabonita
South Fork Kern River,
CA
11.2
11.5
60.0 180.2
9.2
14.7
10.3
17.7
31.1
Gold and Gall (1975)
0. apache
Gila and Little Colorado
Rivers, AZ
10.8
11.4
59.5 156.0
9.5
15.2
10.8
19.8
30.8
Miller (1972)
0. c. pleuriticus
Williamson Lakes, CA
10.9
11.2 61.6
188.9
9.1
14.5
11.3
20.4
38.8
Gold et al. (1978)
Redband
Sheepheaven Creek, CA
10.5
11.4
61.4 162.0
9.5
13.3
9.8
15.6
36.0
Gold (1977)
0. gila
0. chryosgaster
a
b
Mainstem site from Currens (1987).
Bakeoven Creek, Deschutes River tributary, from Currens (1987).
A: 0. mykiss - Suntaheen Creek, AK
B: 0. mykiss mainstem, Deschutes River, OR
C: 0. mykiss - Bakeoven Creek, Deschutes River, OR
D: 0. gila Gila River, NM
E: 0. chryosgaster - Arroyo de a Rana, Mexico
F: 0. aguabonita - S. Fk. Kern River, CA
G: 0. apache - Gila and LiWe Rivers, AZ
H. 0. c. pleuriticus - Williamson Lakes, CA
I. Redband - Sheepheaven Creek, CA
2.06
1.36
III
D
0.66
F
'C
G
-0.05
B
II
-0.54
-1.28
-0.47
0.35
117
1
r
1.44
0.78
H0.12
.99
-0.54
-1.28
I
Figure 2.13. Multidimensional scaling (dimensions 1,11 and ifi) of Euclidian distance
based on nine meristic characters of 54 samples of coastal cutthroat trout (open circles)
from this study and other species of western trout (solid circles) based on published
mean values (Table 2.5) including Suntaheen Creek steelhead from this study. Both
figures are the same, but figure has been rotated. The stress value for this MDS was
0.067. Location 23 is a coastal cutthroat trout populations from Willow Creek, British
Columbia (this study).
golden trout. In summary, despite the large variation in meristic characters observed
within and among coastal cutthroat trout samples surveyed in this study, coastal
cutthroat trout remained a cohesive group when compared to other western trout
species in MDS analysis of nine meristic characters.
Locations sampled from regional areas that provided refuge during the Pleistocene
glaciation showed little phenetic affinity for geographically adjacent locations, but
areas that showed high pheneitc affinity for geographically adjacent locations (i.e.,
Ketchikan area; Wolverine Creek - location 11, Vixen Inlet 13, Margaret Creek
(Traitors Cove)
14, Throne River 15, and Bakewell Creek 16) may represent
similar recolonization events or reflect very similar environmental conditions resulting
in similar meristic characters. Gines Creek, Alaska (location 3), may represent a
relatively young population and the uniqueness of these fish based on the meristic
characteristics may be a result of a recent colonization. Gines Creek fish had the
highest number of significantly different medium counts (number of significantly
different medium counts = 168), the next most unique was the Humboldt State
University Hatchery (location 55) with 122 significant different medium counts. In
addition, Gines Creek clustered on the margins of the coastal cutthroat group when
compared to other western trout species with the MDS ordination (Figure 2.13).
Patterns found along the Washington coast south to the mid-Oregon coast may be a
result of Columbia River and its influence during Pleistocene glaciation or an artifact
of the many years of hatchery releases; including the introduction of hybrids, that has
occurred in this region (Johnson et al. 1999).
The two hatchery samples had the highest variation for canonical variate analysis
and the second (Humboldt State University Hatchery, location 55, 168 significant
differences) and seventh (Alsea Hatchery, location 44, 97 significant differences)
greater number of significantly different mean counts. The Humboldt State University
Hatchery samples had the highest phenetic affinity with Widow White Creek (location
54), a previous source for the hatchery's broodstock (E. Loudenslager, Humboldt State
University, personal communication). These two sites were similar to the sample from
Willow Creek, British Columbia (location 23), that included many individuals with
one of the two characters used to distinguish steelhead. These samples exhibited some
phenetic affinity for steelhead samples (Figure 2.13). Hatchery production of
anadromous coastal cutthroat trout has been focused on providing sport fishing
opportunities and has occurred primarily in Puget Sound, Grays Harbor, the Lower
Columbia River, and the Oregon coast (Johnson et al. 1999). Hartman and Gill (1968)
concluded that the introduction of hatchery coastal cutthroat trout in British Columbia
streams that had occurred as early as 1930, occurred in streams that where coastal
cutthroat trout were all ready present and were generally not successful and, therefore
were unlikely to have influenced the distributional characteristics of coastal cutthroat
trout in the areas they surveyed. As early as 1912 the State of Washington reported the
release of cutthroat trout and steelhead hybrids (cutthroat cross trout) in the Nisqually
River, Puget Sound, Washington (Riseland 1912). Crawford (1979) reviewed the
history of trout brood stocks in Washington and reported that returns from an
anadromous coastal cutthroat trout program started in the 1950s were poor and in
response, brought in Alsea River hatchery fish followed by crosses with steelhead in
the late 1 960s. It appears most of these efforts to establish high return rates with
hatchery fish were unsuccessful, it is unclear how the release of hybrid coastal
cutthroat trout and steelhead may have influenced the native populations.
For several of the meristic characters, the presence of hybrid fish, particularly of
hatchery origin, may explain the undulating pattern that disrupts what might be a clinal
51
change in counts that was observed for many of the locations along the Washington
coast south to the mid-Oregon coast. This would explain the increase in anal
pterygiophore counts and perhaps vertebrae counts in this region, but the direction of
the deviation of all other counts in this region are not consistent with this scenario.
Specifically, both scale row counts are higher in this region where I would suspect
lower counts if hybrids or steelhead were included in the samples.
The use of meristic characters proved useful for the identification of steelhead
misidentified as coastal cutthroat trout. In most cases, the steelhead that were
misidentified as coastal cutthroat trout were small, less than 75 mm in length.
Character traits used in the past (maxillary extension, basibranchial teeth, and presence
of a yellow, orange, or red line in the skin folds of the lower jaw) appear to be
reasonable for the field identification of these species when fish are greater than 75
mm in length. Misidentification of these species could be problematic in other studies
where non-invasive samples (e.g., fin clip) are taken for genetic analysis without the
collection of voucher specimens to confirm fish identification.
The presence in my samples of individual fish that were hybrids between coastal
cutthroat trout and 0. mykiss may have occurred since the absence of basibranchial
teeth and a short maxillary is likely not diagnostic. Behnke (1992) speculated that
although gene flow may occur between coastal cutthroat trout and coastal rainbow
trout (i.e., steelhead), reproductive isolation typically does not break down completely
and it is uncommon to find a hybrid swarm typically found when interior cutthroat
trout species hybridize. Campton and Utter (1985) concluded that a post zygotic
isolating mechanism may be contributing to the preservation of species integrity given
the infrequent observation of adult hybrids. The clearest example of a population that
may have contained hybrid individuals is Willow Creek, British Columbia (location
52
23). Fourteen of the 23 fish in the Willow Creek sample were identified as coastal
cutthroat trout in the field, but were latter determined to be steelhead based on the
absence of basibranchail teeth and a short maxillary extension. The mean length of
these 14 fish was 65.7 mm. Of the nine fish used for analyses, five had either no
basibranchial teeth or a short maxillary extension. In the cluster analysis (Figure 2.11),
fish from Willow Creek exhibits no geographic concordance with adjacent locations
and clusters with the southern populations (locations 49,50,53,54, and 55) that tended
to have several characteristics similar to those of 0.
mykiss
(e.g., lower scale row
counts, higher number of vertebrae). The Willow Creek sample had the highest mean
count for vertebrae
(x
= 63.1, Figure 2.4, Appendix Table A2.2) and the lowest mean
count of scales in lateral series
(x
= 131.8, Figure 2.5, Appendix Table A2.2), both
counts very similar to those fish identified as 0. mykiss in this study (X = 63.3
vertebrae, x = 124.6 scale in lateral series, Table 2.2). Tn the MDS analysis of coastal
cutthroat trout from this study (Figure 2.12), Willow Creek appears on the margins of
the cluster of coastal cutthroat trout populations. In the MDS analysis with other
western trout species (Figure 2.13), Willow Creek is the coastal cutthroat trout
population most closely associated with the group that included 0.
mykiss,
Colorado
cutthroat trout, Apache trout, and California golden trout.
The identification of juvenile fish with intermediate phenotypes consistent with
those expected from decedents of coastal cutthroat trout and steelhead reported by
Campton and Utter (1985) based on allozymes was the first formal documented
observation of hybrids in the wild. This led the authors to suggest that in streams
where both species spawn "the production of hybrid offspring may not be uncommon."
But the existence of hybrids, or at least fish with intermediated characteristics between
coastal cutthroat trout and steelhead, in the wild had been observed prior to Campton
53
and Utter's published results. The presence of hybrids between coastal cutthroat trout
and steelhead in the wild was suspected as early as 1940, when Professor Roland E.
Dimick at Oregon State University observed fish with meristic and morphological
characters intermediate to those of 0. c. clarki and 0. mykiss in coastal basins in
Oregon. He observed these fish with intermediate characters only where both 0. c.
ciarki and 0.
mykiss
were found, and in 1941 carried out paired matings in the Alsea
Fish 1-latchery, Oregon, to confirm his suspicion that these individuals with
intermediate characters were hybrids. Two of the key characters used by Dimick were
scales in the lateral series and the length of the maxillary extension. Unfortunately the
results were never published, and the only records of his work are his field notes with
his observations and the results from the hatchery experiment, and a four page
summary in the archives at the Department of Fisheries and Wildlife, Oregon State
University. Dimick's notes are extremely useful in that he developed a set of 10
"character quantities and their values", based on his field observations and hatcheiy
experiments, that he used to determine whether a fish in hand was 0. c. clarki or 0.
mykiss.
Recently, studies to examine hybridization between coastal cutthroat trout and
steelhead using molecular markers have been conducted (Hawk ins 1997; Wenburg et
al. 1998; Young et al. 2001; Baker et al. 2002). Genetic studies in conjunction with
meristic analysis should provide context to historical surveys and help with the
development of methods appropriate for use in the field to provide better identification
of coastal cutthroat trout, steelhead, and hybrids (Weigel et al. 2002; Baumsteiger
2002). it is not always possible nor appropriate to collect voucher specimens when
working with threatened or endangered species; but when possible, voucher specimens
should he taken to confirm field identification. This is especially true in areas where
54
coastal cutthroat trout and steelhead occur, the misidentification or the collection of
hybrids could lead to incorrect interpretation of the results.
In summary, coastal cutthroat trout exhibited extensive variation in meristic
characters across their distributional range with populations at the southern end of the
distributional range exhibiting phenetic affinity in spite of rather significant meristic
differences within this regional area. My findings are consistent with those of Behnke
(Behnke 1992) that there is no clinal variation across the range. The regional clustering
of locations in the southern portion of the range is in contrast to findings of Behnke's
that coastal cutthroat trout are morphologically similar throughout their range,
although coastal cutthroat trout populations formed a cohesive group in MDS analysis
when compared to other western trout species, there were significant differences in
ineristic characters among coastal cutthroat trout populations surveyed. The
populations examined from the northern periphery of the distributional range did not
show phenetic affinity, nor did the northern populations exhibit the large amount of
within region differences (based on branch length in the UPGMA cluster analysis) in
meristic characters that was observed from the locations sampled at the southern
periphery of the range.
Populations that occupy the periphery of a species' range may occupy areas that are
marginal in terms of habitat or ecological conditions in addition to being on the
margins of a species distributional range. Peripheral populations at both the northern
and southern extent of the range of coastal cutthroat trout are geographically marginal
based on spatial distance. The southern populations are also ecologically marginal
since they experience different biotic and abiotic environments (i.e., Kiamath
Province, etc.) than adjacent populations along the Oregon coast and throughout the
range of coastal cutthroat trout whereas the northern populations occupy areas that are
55
similar to conditions throughout a large part of the distributional range. This may
explain the unique meristic characters of populations in the southern end of the range
and the large within-region variation observed among these southern populations.
Coastal basins in Oregon south of, and including the Sixes River, drain the Klamath
Mountains and the unique geology that makes up the Kiamath Mountains Geologic
Province. Sampling locations from the region include locations 48 to 55. Lesica and
Allendorf (1995) reviewed various aspects of peripheral populations and their value to
species conservation with special attention to the distinction between marginal versus
peripheral habitat conditions. In general, peripheral populations tend to be genetically
and morphologicaliy divergent from central populations and that morphological
characters are expected to diverge more rapidly in isolated populations than gene
frequencies (Lesica and Allendorf 1995).
Recently, Channell and Lomolino (2000) found that in contrast to the conventional
thinking that predicts core populations persisting and range contraction as a species
becomes endangered, most species (of 245 examined) they considered persist in the
periphery of their historical geographical range. Although there is no indication that
the southern distribution of coastal cutthroat trout has changed over the last century,
long-term conservation plans are likely to depend on protection of distinct populations
(Lesica and Allendorf 1995). particularly peripheral populations in marginal habitats
that may contain high adaptive significance to the species as a whole (Scudder 1989).
Habitat that was suboptimal historically along the periphery of a species range may
provide valuable opportunities for conservation planning (Channell and Lomolino
2000), for it is in these areas one often finds divergence from central populations
(Lesica and Allendorf 1995). For coastal cutthroat trout, the unique meristic
characteristics of populations in the southern extent of the range suggest that these
populations may be essential to conserve the range of diversity in the sub-species.
57
3. GENETIC POPULATION STRUCTURE OF COASTAL CUTTHROAT TROUT
(Oncorhynchus clarki clarki) IN WESTERN NORTH AMERICA
3.1. ABSTRACT
I examined genetic differences among 1,413 coastal cutthroat trout (Oncorhynchus
clarki clarki) from 54 sites across their distributional range using 30 enzyme encoding
loci. Collection sites were located in estuaries or streams downstream of migration
barriers from northern California to Prince William Sound, Alaska. Genetic population
structure was primarily at the individual stream level. There were differences in the
distribution of genetic variation across the landscape although the amount of genetic
variation within regions was similar. Geographic concordance of populations at the
northern and southern extent of the subspecies range was observed and isolation by
distance was evident at regional scales (< 800 km). Compared to other species of
Pacific salmon and trout, coastal cutthroat trout are characterized by more diverse local
populations that act in an independent, isolated nature. The amount of sub-population
genetic structure in my study did not differ among regions, but the strength of isolation
by distance differed among regions with the strongest isolation by distance being
detected at the northern and southern extent of the range.
3.2. INTRODUCTION
An understanding of the spatial distribution of populations of a threatened or
endangered species across its distributional range is fundamental to developing
appropriate conservation and recovery strategies (Waples 1991; Waples 1995). Fish
species typically consist of many geographically localized populations that are, to
varying degrees, reproductively isolated (Taylor 1991a). This is especially true for
local populations of Pacific salmon and trout (Oncorhynchus spp.) that tend to become
distinct from other local populations because of their high fidelity for returning to their
natal stream for spawning. Although homing to natal streams appears to be very strong
in anadromous salmonids, straying among local populations (i.e., immigration) does
occur. The extent to which straying occurs is not well understood, but enough isolation
occurs so that differences exist and intraspecific differences may exist among closely
adjacent populations.
Little is known about the interaction (i.e., migration, gene flow) among populations
of coastal cutthroat trout
(Oncorhynchus clarki clarki)
and the resulting genetic
population structure of the subspecies across its range, perhaps due an under emphasis
of research and management (Nehisen et al. 1991). Moreover, although declining
numbers of coastal cutthroat trout in coastal basins have been suspected for many
years, little is known about the status of these populations. Snyder (1940) observed a
decline of coastal cutthroat trout in northern California from 1897 to 1940. As early as
1948 it was being suggested that harvest be reduced and programs be initiated to
restore and improve habitat in response to declining numbers of coastal cutthroat trout
in Oregon (Sumner 1948). Nehisen et al. (1991) stated that few data were available,
but existing data indicated that a major decline was occurring, mainly as a result of
habitat damage and over-fishing.
Coastal cutthroat trout are the most widely distributed subspecies of cutthroat trout
(Bebnke 1992; Behnke 1992; Behnke 2002). They occupy habitats ranging from small
to large rivers that drain the coastal rainforest between the Eel River, California, and
Prince William Sound, Alaska, with a large portion of their range falling within the
region often referred to as "Cascadia" (McPhail and Lindsey 1986; Behnke 1992). This
region is a highly dynamic environment and episodic changes in Pleistocene climates
and landscapes have influenced the distribution of freshwater biotas (Avise 1992).
59
Compared to other regions in North America, there are few native freshwater fish
species in this region. McPhail and Lindsey (1986) state the principal reasons for the
few number of species are historical and associated with glaciation; at times during the
Pleistocene, over three-quarters of the land area of Cascadia was covered with ice. In
addition, changes to the environment from logging, harvest, fire, debris removal,
barrier removal, out-of-channel water uses, hydroelectric development, and even
preservation of "pristine' environments may have affected fish communities (Michael
1983).
An understanding of the genetic population structure of coastal cutthroat trout must
be considered in the context of its environment for developing appropriate
conservation strategies. Catric et al. (2001) found contemporary genetic population
structure of brook charr (Salvelinusfontinalis) was influenced by landscape features,
although there was substantial variation. At smaller spatial scales (within basins) there
was evidence that the contemporary landscape shaped the pattern genetic diversity; but
at larger spatial scales (among basins) contemporary landscape features had a minor
influence on genetic structuring among basins.
The dynamic environment of the Pacific Northwest creates a situation in which
local populations may go extinct and habitats must be recolonized. The concept of
metapopulations is an useful starting point to consider how local populations interact at
various spacial scales to contribute to the persistence of a larger group of populations
(Hanski and Gilpin 1997). Although the metapopulation concept is widely invoked in a
range of studies and situations (Hanski and Simberloff 1997), its attractiveness for
considering how salmonids persist in a dynamic environment are it's two key premises
that are populations are spatially structured into assemblages of local breeding
populations and that migration among local populations effects local dynamics (Hanski
and Simberloff 1997). The position, distribution, and interaction among local
populations is considered, this is in contrast to standard approaches in population
biology that assume a panmictic population structure. Metapopulations spread the risk
of extinction among subpopulations, changes in local environmental conditions can
lead to local extinctions with the persistence of a group of populations (i.e.,
metapopulation) depending on how well the organism can track the shifting spatial
locations of suitable habitat (Wiens 1997). Some populations may operate as "sources"
and others as "sinks", with subpopulations assuming different roles different times
(Stacey and Taper 1992). However, all subpopulations can contribute to the overall
diversity (Poff and Ward 1990), stability, and persistence (Rieman and McIntyre
1993). Stray rates may not be static, but may vary spatially and temporally.
Environmental and geologic events (e.g., floods, droughts, glaciers) may limit access
to natal streams and spawning areas, perhaps resulting in increased straying in the
pursuit of other suitable spawning areas.
Collections of subpopulations (i.e., metapopulations) of coastal cutthroat trout are
connected through local exchange of individuals, and these metapopulations are
connected across the distributional range of the sub-species through exchange of
individuals. Genetic differentiation occurs when the exchange of individuals or entire
populations (i.e., gene flow) tends to produce genetic homogeneity and is sufficient to
overcome local differentiation that results from mutation, genetic drift, and natural
selection (Slatkin 1987). Wright (1943) introduced the term "isolation by distance" to
describe genetic differences resulting from geographically restricted dispersal (Slatkin
1993). In general, isolation by distance (IBD) suggest that genetic differentiation
increases with geographic distance. Slatkin (1993) examined IBD in populations that
were and were not in equilibrium under dispersal and genetic drift and found that with
61
reasonable sample sizes IBD can be detected. Hutchison and Templeton (1999)
summarize the above work and work by others that develop a stepping-stone model of
population structure where dispersal is constrained by geographic distance resulting in
closely situated populations tend to be more genetically similar, and based on this
foundation developed an approach to evaluate the relative influences of gene flow and
drift on population structure both within and between regional areas. Such a stepping
stone model is appropriate for anadromous salmonids since straying studies have
found that there is a decrease in the number of strays with increasing distance from
natal stream (Labelle 1992; Quinn 1993; Candy and Beacham 2000). The patterns of
genetic variability across the landscape is critical to establishing population units for
conservation and may prove more meaningful than clustering techniques and
ordination techniques that nnay not be appropriate for defining conservation units for
populations that are arranged continuously in space (Diniz-Filho and Telles 2002).
Coastal cutthroat trout exhibit a great amount of variation in life-history types
compared to other Pacific salmonids. In general, there are two major life history forms,
amphidromous3
and potamodromous, although life history traits (e.g., freshwater
residence, age at sexual maturity) appear to be highly variable between these two
forms and among different populations of similar forms (Sumner 1953; Giger 1972;
Jones 1977; Johnston 1982). Amphidromous coastal cutthroat trout, often referred to
as anadromous or sea-run, appear to have a high fidelity for spawning in their natal
stream (Johnston 1982; Campton and Utter 1987), although fish may enter non-natal
streams for reasons other than spawning (e.g., overwintering (Johnston 1982)).
Myers (1949) described amphidromous fish migration as movement from freshwater
to the sea, or vice versa, for non reproductive purposes, such as feeding;
potamodromous fishes migrate wholly within freshwater.
62
Straying (spawning in non-natal stream) is believed to occur at low levels (Giger
1972).
Patterns of allelic frequency divergence have been used to provide a description of
genetic exchange and the amount of isolation among local geographic populations of
salmonids (Allendorl and Phelps 1981). As with other Pacific salmonids, studies have
shown that coastal cutthroat trout form local populations across their range. Applying
various genetic techniques Campton and Utter (1987), Wenburg et al. (1998), and
Wenburg and Bentzen (2001) have found that, within regional areas (<200 km),
coastal cutthroat trout show greater differences among populations than other species
of Pacific salmonids. Campton and Utter (1987) found that anadromous populations of
coastal cutthroat trout within the Puget Sound area of Washington consisted of at least
three genetically divergent groups. In addition, Campton and Utter (1987) found
divergent allele frequencies for a nonanadromous population that suggested a
correspondence between population structure and gene flow. Wenburg et al. (1998)
concluded that coastal cutthroat trout were reproductively isolated at the individual
basin level and loosely structured at a regional scale that included the Puget Sound and
areas of coastal Washington. Moreover, Wenburg and Bentzen (2001) used both tagrecapture information in conjunction with molecular markers (microsatellites) to
conclude that independent streams sampled in Hood Canal, Washington, contained
distinct populations and that these independent populations formed the fundamental
units of coastal cutthroat trout genetic population structure. Their genetic data, that
included additional populations from the Puget Sound and Washington coast, were
consistent with their tag-recapture/genetic findings from the Hood Canal (Wenburg
and Bentzen 2001).
63
Wilson et al. (1985) reported two clonal lines of coastal cutthroat trout from the
lower Fraser River, British Columbia, but did not discuss if these represented two
different populations, either anadromous, resident, or sympatric populations. In the
Coquille River, Oregon, gene flow was found to be as restricted among local
populations of resident coastal cutthroat trout as it was between populations in distant
tributaries in the same drainage (Currens et al. 1992). They concluded that the
observed random genetic differentiation among local populations suggested a history
of genetic drift and small population sizes.
Besides Campton and Utter (1987), few data are available concerning genetic
differences between resident and anadromous forms of coastal cutthroat trout. Other
coastal cutthroat trout studies based on allozymes have reported results ranging from
mixing of anadromous and nonanadromous populations within a single basin
(Zimmerman 1995) to very limited gene flow among populations regardless of the
absence of migration barriers (Griswold 1996). Michael (1983) suggested that
sympatric populations of anadromous and resident coastal cutthroat trout may exist,
but this was based on trapping data and included no genetic analysis.
Studies of other salmonid species with anadromous and nonanadrornous life history
forms tend to find limited gene flow among life history forms. Birt et al. (1986) used
mtDNA to analyze anadromous and nonanadromous populations of Atlantic salmon
(Salmo salar) and detected very little genetic differentiation between the two forms.
Hinder et al. (1991) found no genetic differentiation between resident and anadromous
brown trout (Salmo trutta) from the same locality using electrophoretic techniques.
Costello et al. (2003) found that migration barriers were important in structuring
genetic variation of bull trout (Salvelinus confluentus) within and between basins.
In this study, I examine the geographic distribution of genetic variation of
anadromous coastal cutthroat trout across its distributional range to determine if
genetic diversity among populations differs regionally, perhaps reflecting historical
landscape/environmental features, and to determine the spatial distribution of genetic
diversity across the range of the subspecies.
3.3. METHODS
3.3.1. Sample collection and analysis
A total of 1,619 coastal cutthroat trout were collected from 55 sites in estuaries or
streams downstream of upstream migration barriers from northern California to Prince
William Sound, Alaska (Figure 3.1, Table 3.1). Samples from a location were typically
collected over a period of 1 d, and several sites were sampled in more than one year.
Sample. sizes ranged from 13 to 62 fish per site. Fish were collected by minnow traps,
angling, and downstream migrant traps. Attempts were made to collect several age
classes of fish by sampling different size classes at each sample location. Two samples
from hatcheries, Humboldt State University Hatchery, California, and Fall Creek
Hatchery (Alsea River), Oregon, were also included in the analysis (Table 3.1).
After capture, fish were euthanized (tricane methanesulfonate), weighed (g),
measured (fork length FL), and photographed (left side, Ektachrome ASA 200). Each
fish was tagged with an identification number that was secured to the lower jaw.
Sampled fish were either frozen whole on dry ice in the field or, in the case of larger
fish (>250 mm), tissue samples for allozyme analysis were dissected in the field and
placed on dry ice prior to transport. Liver, heart, eye, and skeletal muscle samples were
collected from each fish for allozyme analysis. Muscle samples were taken from right
DD
Figure 3.1. Locations where samples were collected and regional groupings for
analysis. Location numbers correspond to numbers in Table 3.1.
Table 3.1. Collection sites, location, number of fish collected (N), and number of fish
determined to be coastal cutthroat trout based on laboratory examination of
basibrianchial teeth and maxillary extension (n). Location numbers correspond to
numbers in Figure 2.1. If no value is given for n, all fish were determined to be coastal
cutthroat trout.
Location
River system/drainage
Collection site
Location (UTM)
number Zone
E
N
N/n
Bosewell Bay, AK
Fish Creek
1
6
544100 6697100
Martin River, AK
Martin Lake outlet
2
6
632050 6696900 25/23
Gines Creek, AK
3
7
643200 6568900 46/45
Suntaheen Creek, AK
4
8
493100 6434375
33
17/0
Freshwater Bay, AK
Bayhead Creek
5
8
486550 6423650 32/15
Kiag Bay, AK
West Chichagof Lake
6
8
436000 6394300 18/16
Portage Bay, AK
Portage Creek
7
8
606300 6310550 32/22
Duncan Salt Chuck, AK
8
8
Kadake Creek, AK
9
8
603350 6304300 52/50
561750 6293000
38
Eagle River, AK
10
9
339500 6226600 32/3 1
11
9
325150 6202750
50
Staney Creek, AK
12
8
620650 6186550
51
Vixen Inlet, AK
13
8
688000 6185700 34/12
14
9
335200 6174950 27/25
ThroneRiver,AK
15
8
652000 6173850 62/60
Bakewell Creek, AK
16
9
394950 6128100 52/35
Wolverine Creek, AK
Traiters Cove, AK
McDonald Lake outlet
Margaret Creek
Skeena River, BC
Herman Creek
17
9
525500 6027000
25
Tiell River, BC
Survey Creek
18
9
290400 5916700
20
19
9
281100 5918600 18/16
Yakoun River, BC
Bella Coola River, BC
Molly Walker Creek
20
9
675550 5811900
Bella Coola River, BC
Fish Creek
25
21
9
665300 5806900 25/13
SanJosefRiver,BC
22
9
558860 5613190
25
Willow Creek, BC
23
10
340140 5535680
25/9
SakinawLake,BC
MixalCreek
24
10
425560 5501100 25/14
Fraser River, BC
Salmon River
25
10
531580 5440750
29
Ritherdon Creek, BC
26
10
358200 5424200
25
Sandhill Creek, BC
27
10
469760 5380850 25/23
67
Table 3.1. Continued
Location
River system/drainage
Collection site
Location (UTM)
number Zone
E
N
N (n)
Kirby Creek, BC
First tributary
28
10
433300 5360410 28/26
Stillaguamish River,
Church Creek
29
10
550240 5342720 56/53
Hoko River, WA
BearCreek
30
10
401290 5332590 25/21
31
10
450800 5328760 25/24
Salt Creek, WA
Quillayute River, WA
Dickey River
32
10
381435 5310896
Hoh River, WA
Alder Creek
33
10
407145 5299280 25/13
Hood Canal, WA
Big Beef Creek
34
10
516400 5277240
20
Humptulips River, WA
Stevens Creek
35
10
432330 5242280
25
McClane Creek, WA
West Fork
36
10
500180 5208400
32
Naselle River, WA
Alder Creek
37
10
450895 5147569 26/24
Nehalem River, OR
Foley Creek
38
10
432710 5054300 30/29
Trask River, OR
North Fork
39
10
453430 5033540 3 1/30
Sand Creek, OR
Andy Creek
40
10
429150 5017880
SchoonerCreek,OR
41
10
429762 4976525 19/10
Siletz River, OR
42
10
422939 4962940
22
13/12
21
Yaquina River, OR
Wolf Creek
43
10
443335 4935710
30
Alsea River, OR
Alsea Hatchery
44
10
440360 4916320
30
45
10
411570 4901795 27/25
Cummins Creek, OR
Umpqua River, OR
W. Br. N. Fk. Smith R.
46
10
429360 4860000 22/2 1
New River, OR
Davis Creek
47
10
383880 4758840 32/28
Rogue River, OR
Iron Creek
48
10
401960 4717360
33
Winchuck River, OR
Wheeler Creek
49
10
408085 4654600
26
Lake Earl, CA
Jordon Creek
50
10
404300 4627560 28/26
Klamath River, CA
Waukell Creek
51
10
413370 4594310 25/24
Redwood Creek, CA
May Creek
52
10
414200 4577870
Mill Creek, CA
53
10
403660 4546040 24/12
Widow White Creek,
54
10
406750 4534000
18/9
Humboldt St. Univ., CA Hatchery
55
10
408420 4525640
31
27
side of fish. Tissue samples from each fish were placed in tubes labeled with the fish
identification number. Tissues from fish that had been frozen whole in the field were
dissected in the laboratory.
Coastal cutthroat trout and steelhead (0. mykiss) cooccur in many parts of their
ranges (Scott and Crossman 1973; Behnke 1992), are phenotypically very similar, and
share many alleles (Campton and Utter 1985). To avoid inadvertently including
steelhead4
in the genetic analysis, I developed an a priori decision rule to identify
steelhead individuals and remove them from the analysis based on two classical
taxonomic traits. Basibranchial teeth are absent in coastal 0. mykiss and the maxillary
extension extends beyond posterior portion of eye in 0. o. clarki (Scott and Crossman
1973; Scott and Crossman 1973; Behnke 1992; Moyle 2002). In addition, the presence
of a yellow, orange, or red line in the skin folds of the lower jaw often occurs and
gives "cutthroat" their name. Individuals with a maxillary extension score of 0 or 1
(maxillary did not extend past the posterior portion of the eye) and an absence of
basibranchial teeth were designated as a steelhead and not included in the analysis. An
individual with a maxillary extension score of 2 or the presence of basibranchial teeth
was designated as a coastal cutthroat trout.
Allozyme analysis followed methods of Aebersold et al. (1987). Thirteen enzymes
encoded by 30 loci were screened (Table 3.2). A known standard from rainbow trout
(0. mykiss) was placed on each gel as a reference for electrophoretic mobility for aid
in identifying alleles.
For purposes of this study, all 0. mykiss collected will be referred to as steelhead
since all samples were obtained downstream of migration barriers, although some
nonanadromous 0. mykiss (rainbow trout) may be included.
Table 3.2. Enzymes and electrophoretic conditions to resolve loci examined in this
study.
E.G
Enzyme name
Creatine kinase
Fructose-biphosphate aldolase
number a
2.7.3.2
4.1.1.13
Locus
1.1.1.8
Glyceraldehyde-3-phosphate
dehydrogenase
1.2.1.12
E
3
FBALD-] *
E
5
E
5
G3PDH-] *
G3PDH-2 *
M
M
1
GAPDH2*
E, H
E, H
1, 2
1, 2
GAPDH-3 *
GAPDH4*
GAPDH5*
Guanine deaminase
Glucose-6-phosphate isomerase
Isocitrate dehydrogenase
L-Lactate dehydrogenase
3.5.4.3
5.3.1.9
1.1 1.42
1.1.1.27
Buffere
CKB*
FBALD2*
Glycerol-3-phosphate
dehydrogenase
Tissueb
E,H
E,H
1
1,2
1,2
GDA-1
L
L
3
3
GPI-AI *
M
M
3, 5
GDA2*
GPIB1*
GPIB2*
mIDHp-1 *
mIDHp2*
sIDHp1,2*
LDH-A1 *
LDHA2*
LDHB1*
LDHB2*
LDHC1*
M
3,5
3,5
H, M
2, 1
H,M
2,1
L
1
M
M
3
3
E,H,
5,4,3
M
E,L
5,3
5
E
Malate dehydrogenase
1.1.1.37
sMDHA1,2*
sMDH-
H, L
2, 1
H,M
2,1
BJ,2*
Malate dehydrogenase (NADP+)
1.1.1.40
mMEP-] *
sMEP1*
sMEP2*
M
M
L,M
1
1
I
70
Table 3.2 Continued
Enzyme name
a
E.C.
number a
Locus
Tissueb
Bufferc
Peptidase Glycyl-L-leucine
34**
PEPA1*
E, H,
5, 4, 3
Phosphogluconate dehydrogenase
1.1.1.44
PGDH*
E, M
1
Superoxide dismutase
1.15.1.1
sSOD *
M
5
E.C. #: Ensyme Commission, International Union of Biochemistry Nomenclature
(1984).
b
Tissues are eye (E), heart (H), liver (L), muscle (M).
Buffers used: 1: ACE, an citrate-amine-EDTA gel and tray buffer pH 6.8; 2: ACE
with NAD; 3:TBCLE, a Tris-citrate gel buffer and lithium hydroxide, borate-EDTA
tray buffer pH 8.5; 4: TG, a Tris-glycine gel and tray buffer pH 8.5 using Trizma., 5:
TGIKG, a Tris-glycine tray buffer pH 8.4 using Tris.
71
3.3.2. Data analysis
The number of polymorphic loci, percentage of polymorphic loci, mean number of
alleles per loci, and expected proportion of heterozygotes
(He) were calculated for each
sample as measures of genetic variation within samples. Polymorphic loci were those
loci which at least one sample had a frequency of the most common allele less than or
equal to 0.95 (P095). The expected proportion of heterozygotes (i.e., average
heterozygosity) was calculated for each locus and averaged over all loci (Weir 1996).
Genotypic frequencies were examined for consistency with Hardy-Weinberg
expectations with exact probability tests, using BIOSYS-il (Swofford and Selander
1997) and Genetic Data Analysis (GDA) (Lewis and Zaykin 2002). Measures of within
population genetic variation did not include duplicated loci (sIDHp1,2*; sMDHA1,2*; sMDHB1,2*) since expected values could not be calculated for a pair of loci. A
sequential Bonferroni correction (Rice 1989) was used over the multiple tests carried
out(k= 864).
Population differentiation was examined by testing for homogeneity of allele
frequencies among all samples and between all pairs of samples with exact probability
tests using a Markov chain method by GENEPOP 3.3 (Raymond and Rousset 1995).
The combination of probabilities at all loci, a global significance over all loci, was
obtained using Fisher's method for test combination (Sokal and Rohlf 1981). Estimates
of subpopulation structure were obtained using Wright's F-statistics (Wright 1978)
calculated by Weir and Cockerham's 0 (Weir and Cockerham 1984), to correct for
sample size and number of populations allowing comparisons with different types of
molecular markers (Excoffier et al. 1992). A 95% confidence interval of U over all loci
was estimated by bootstrapping (1,000 replications) over loci. Hierarchical gene
diversity analysis was performed using HIERARCHY and WRIGHT78 in BIOSYS-il
72
(Swofford and Selander 1997) with population assigned to one of four regional areas
(see below).
Genetic distances between populations were estimated with Cavallis-Sforza and
Edwards chord distance, DCE (Cavalli-Sforza and Edwards 1967). Dendrograms based
on both neighbor-joining (Saitou and Nei 1987) and the unweighted pair-group method
with arithmetic averages (UPGMA) algorithm (Sneath and Sokal 1973) were
constructed using DCE. The robustness of the branches were evaluated by bootstrapping
the distance matrix (1,000 replications) and finding the consensus tree using the
programs NEIGHBOR and CONSENSE in PHYLIP 3.57c (Felseristein 1995).
Steelhead from Suntaheen Creek, Alaska, were included as an outgroup in the
dendrograms. Non-metric multidimensional scaling analysis (MDS) (Kruskal 1964a;
Kruskal 1964b) was used to create a plot of the locations in three dimensions based on
allele frequencies of 12 polymorphic loci (P095). The Suntaheen Creek steelhead
sample along with a sample of 0. mykiss from Bakeoven Creek, tributary to the
Deschutes River, Oregon (Currens 1997), were used as outgroups for the MDS
analysis. The reduced set of loci (GPI-A1
GPI-B1
GPIB2*, rnIDHp2*, sIDHp-
1,2*, LDHB2*, sMDHA1,A2*, sMDHB1,B2*, sMEP1*, sMEP2*, PEPA*,
PGDH*) corresponded to data from Currens (1997).
Mantel tests (Mantel 1967) were used to test for isolation by distance (IBD) by
comparing distance matrices describing geographic and genetic relationships among
sampling locations (Oden and Sokal 1986; Slatkin 1993). Pairwise O(i.e., pairwise
FST) values were calculated for all comparisons and were plotted against the pairwise
geographic distance (km). A product-moment correlation, r, and the Mantel test
statistic, Z, were computed to measure the degree of relationship between the two
matrices using NTSYS (Rohlf 2000). The significance of the correlations were tested
by randomly permuting (10,000 permutations) rows and columns of one of the
73
matrices while keeping the other constant resulting in the sampling distribution of the
test statistic under the null hypothesis that there is no correlation (Smouse et al. 1986).
Geographic distances were calculated using a geographical information system (GIS)
AML method developed by Bunn et al. (2000) that used least-cost modeling to
determine distances between sampling locations. Distances were calculated two ways.
One approach calculated the shortest water distance between each pair of locations.
The second approach calculated the shortest water distance with a water depth
constraint (i.e., crossing shallow water was less costly than crossing open, deep water).
Avoidance of deep water areas in the marine environment by coastal cutthroat trout
has been suggested (Jones and Seifert 1997). Paths between sampling locations was
restricted to depths less than 1500 m. Ocean depths less than 1500 m were partitioned
into four depth classes: < 100 m, 100 - 500 m, 500 - 1000 m, 1000
1500 m.
Spatial distribution of genetic variation was characterized with spatial
autocorrelation analysis. A nonlinear multivariate Mantel correlogram (S. Goslee,
USDA-ARS Pasture Systems and Watershed Management Research Unit, University
Park, PA; personal communication, 2003) was used to examine the relationships
between genetic variation and spatial structure. The Mantel test assumes a linear
correlation between the two distance matrices and is based on the general hypothesis
that samples close in space are similar genetically. Because the Mantel test averages
the correlation over all distance classes, a Mantel correlogram was used to examine the
spatial scale of the correlation. A Mantel correlogram assesses the Mantel correlation
between genetic distance and geographic distance in specific distance classes and
provides a picture of how the correlation changes over the spatial extent of these data
(Oden and Sokal 1986). The distance classes were selected so as to be of equal widths
following Sturge's rule (Legendre and Legendre 1998).
74
To determine if there were regional differences in genetic population structure and
IBD, sampling locations were partitioned into four regional groups (Table 3.1; Figure
3.1). The regional groups were based on general biogeographical boundaries largely
driven by Pleistocene glaciation. Region A represents the northern portion of the
current coastal cutthroat trout distribution and although subjected to glacial cover
during the Pleistocene, some refugia are considered to have existed in areas of the
Queen Charlotte Islands, west coast Prince of Wales Island, and north of the current
coastal cutthroat trout distribution (McPhail and Lindsey 1986). This region includes
the Skeena, Nass, and Stikine river systems. Region B represents the Puget Sound and
southern British Columbia coast that was completly covered by glaciers and includes
the Strait of Juan De Fuca, Strait of Georgia, and the Fraser River system (McPhail and
Lindsey 1986). Region C represented coastal locations that experienced some glacial
coverage in the northern portion, but even in the northern portion rufugia may have
existed (e.g., northern Vancouver Island). This region includes the Columbia,
Chehalis, and Umpqua river systems. Region D was partitioned out from region C
based on the slight genetic affinity detected and the stronger meristic affinity (Chapter
2) exhibited by samples south of Cape Blanco. This region includes the Rogue and
Kiamath systems.
3.4. RESULTS
Of the 1,619 fish initially collected, 98 steelhead were identified based on the
absence of basibranchical teeth and a short maxillary extension and were excluded
from the genetic analysis. Fish that only met one of the two criteria (i.e., absence of
basibranchical teeth or a short maxillary extension) were not excluded from the
analysis. One sample contained only steelhead (Suntaheen Creek, Alaska, location
number 4) and was only used as an outgroup.
Subsequent analyses were based on data from a total of 1,413 coastal cutthroat trout
from 54 locations (Table 3.1) with 95 alleles segregating at 30 loci. Seventeen of these
loci had an average frequency of the most common allele of 0.95 or less in at least one
sample: CKB*, GAPDH2*, GDA]*, GDA2*, GPIA1*, GPIB1*, GPIB2*,
mIDHp2*, sIDHp1,2*, LDHB2*, sMDHAl,A2*, sMDHB],B2*, sMEP1*, sMEP2*, PEP-A *, PGDH*, 5SOD* (Appendix Al 1). Nine loci were monomorphic in all
samples (FBALD2*, G3PDH2*, GAPDH3*, GAPDH4*, GAPDH5*, LDHA1*,
LDHA2*, LDH-B1 *, and mIDHp-1 *) and 11 private alleles were detected (Table 3.3).
Of particular note, the frequency of the private allele sMEP-2 * 90 was 0.13 in the
sample from Martin River, Alaska.
Average heterozygosity
(He) ranged from 0.036 (Siletz River, OR 43) to 0.101
(Widow White Creek, CA 54) with a mean of 0.062. The percentage of polymorphic
loci (P095) ranged from 7.4% to 25.9% with a mean of 17.4% (Appendix Table A3. 1).
The mean number of alleles loci ranged from 1.1 to 1.4 (Appendix Table A3. 1). No
latitudinal dines or regional patterns were detected in any of the measures of within
population genetic diversity (Appendix Figure A3.1). Samples generally conformed to
Hardy-Weinberg expectations. Only 20 (2.3%) of the 864 locus/sample combinations
were significant after application of the sequential Bonferroni correction for multiple
comparisons (Appendix Table A3.2). Three loci (GDA-1
GDA2*, and GPI-B/ *)
contributed disproportionately to the number of test not conforming to HardyWeinberg expectations. Seventeen of the 20 significant tests after sequential
Bonferroni correction were a result of a deficiency of heterozygotes at one of these
three loci.
At the scale sampled, coastal cutthroat trout from different locations across their
distributional range were composed of genetically distinct populaitons. Genetic
differentiation among the 54 samples was significant (P < 0.05) for 16 of the 17
Table 3.3. Location and frequency of private alleles detected from 54 samples of
coastal cutthroat trout. Location numbers correspond to numbers in Table 3.1 and
Figure 3.1.
Locus
Allelea
Frequency
Sample found in (location number)
FBALD-] *
84
0.0 15
Rogue River, Iron Creek (48)
G3PDH-1 *
-10
0.038
Bella Coola River, Fish Creek (21)
GPI-B1 *
160
0.017
Trask River, North Fork Trask River (39)
sIDHp1,2*
21
0.033
Freshwater Bay, Bayhead Creek (5)
sIDHp-1,2 *
52
0.033
Freshwater Bay, Bayhead Creek (5)
LDHB2*
119
0.040
Cummins Creek (45)
LDHB2*
85
0.083
Salt Creek (31)
LDH-C1 *
106
0.0 19
Winchuck River, Wheeler Creek (49)
sMDHA1,2*
39
0.071
Sakinaw Lake, Mixal Creek (24)
mMEP]*
62
0.010
Staney Creek (12)
sMEP2*
90
0.130
Martin River (2)
a
relative mobility based on 0. mykiss standard.
77
polymorphic loci analyzed and the global significance over all loci was also
significant (P<0.05). Significant heterogeneity in allele frequencies found in all
pairwise comparisons indicates that coastal cutthroat trout sampled in different
locations across their distributional range were composed of genetically distinct
populations. The global multilocus estimate for Owas 0.13 1 ± 0.045 (Table 3.4). The
95% confidence interval based on bootstrapping (0.176 - 0.086) was greater than zero
indicating a heterogenous distribution of the genetic variability of coastal cutthroat
trout over its distributional range. Regional estimates of 0 ranged from 0.084
(Washington and Oregon coast, region C) to 0.101 (Puget SoundlStrait of Juan De
Fuca/Strait of Georgia area, region B) (Table 3.4). There were no significant
differences in 0 among regions (i.e., the 95% confidence interval of Ofor each region
based on bootstrapping overlapped the 0 estimate for each region). These regional
95% confidence intervals were all greater than zero indicating a heterogenous
distribution of genetic variability within all regional groups. Hierarchical gene
diversity analysis found that allele frequency differences among populations was
10.9% with differences among populations within a regional area contributing more to
the variation than differences among regions (Table 3.5).
Genetic similarity of coastal cutthroat trout populations from, locations in
geographic proximity was most evident at large spatial scales, although some finer
scale genetic affinity corresponding to geographic proximity was observed. The
unrooted neighbor joining tree developed from DCE distances illustrates a slight
geographic affinity of coastal cutthroat trout populations at a regional scale and a
distinct separation of the 0. mykiss sample from the 54 coastal cutthroat trout samples
(Figure 3.2). Although there was weak bootstrap support for the neighbor joining tree
with only 19 of the branches having bootstrap support of greater than 30%, the
geographic affinity illustrated by the consensus tree should not be dismissed given the
Table 3.4. Regional and overall F-statistics and correlation results from Mantel test for
isolation-by-distance based on 0(17 polymorphic loci) and geographic distance
(shortest water distance). The upper and lower bounds of the 95% confidence interval
for F-statistics were derived from bootstrapping over loci (1,000 replications). Fvalues for Mantel r were based on permutation test of 10,000 runs. Regional groups are
those presented in Figure 3.1.
Grouping
F-statistics
Isolation by distance
0
95% confidence interval
Mantel r
P-value
Region A
0.095
0.125 0.059
0.238
0.079
Region B
0.101
0.165 0.059
0.236
0.172
Region C
0.084
0.125 0.040
0.159
0.179
Region D
0.100
0.185 0.050
0.436
0.099
Overall
0.131
0.176-0.086
0.415
0.0001
Table 3.5. Hierarchical gene diversity analysis of 54 coastal cutthroat trout populations
from across their distributional range.
Source of variation
Total
Percent of total
100.0
Within populations
88.9
Among populations
11.1
Among regions
2.5
Among populations within regions
8.6
79
1p
Regional groups
Figure 3.2. Consensus tree based on 1,000 replications of genetic relationships among
54 samples of coastal cutthroat trout from across distributional range and a steelhead
outgroup (Suntaheen Creek, Alaska, location 4) constructed with a neighbor-joining
algorithm from DCE chord distances shown as an unrooted tree. Neighbor-joining
bootstrap percentages based on 1,000 replications, only percentages greater than 30%
shown (e.g., 73), scale bar represents bootstrap values, not genetic distance. Location
numbers correspond to Table 3.1 and Figure 3.1.
number of populations in the analysis and the number of possible trees (1,000
replications). All but one of the Alaska samples are located in the "lower portion" of
the consensus tree (Figure 3.2) and only three non-Alaska samples are located in this
portion of the tree (locations 22 San Josef River, 30 Hoko River, and 33 Hoh
River). These sites are three of the five most northern sites in regional group C,
perhaps explaining their apparent affinity with the more northern Alaskan sites. The
general tree topology observed would not be inconsistent with a radiation from a
common ancestor. The long branch lengths suggest a distinctiveness of each
population and are consistent with the significant heterogeneity in allele frequencies
found in all pairwise comparisons. This adds further support that the coastal cutthroat
trout populations from different locations across their distributional range were
composed of genetically distinct populations. The steelhead sample (Suntaheen Creek,
Alaska) appears to be quite different based on the neighbor joining tree.
The UPGMA rooted dendrogram based on DcE distances also illustrates some
geographic affinity of coastal cutthroat trout populations at a regional scale and a
distinct separation of the 0. mykiss sample (Figure 3.3). Five of the eight samples from
southern Oregon/northern California (regional group D) grouped together, separately
from all other samples; included with these southern samples was the sample from the
Martin River, Alaska (location 2), near the northern extent of the distributional range
of 0. o. clarki. Although there was weak bootstrap support for the UPGMA
dendrogram with only 17 branches having greater than 30% support, some geographic
concordance was illustrated by the consensus dendrogram (Figure 3.3). There was
100% bootstrap support for separation of the 0.
mykiss
sample from the 54 coastal
cutthroat trout samples. Similar to the consensus neighbor joining tree, a large portion
(12 of 17) of the region A samples showed geographic affinity and grouped outside of
[3'
40
20
38 A
31
29
27
20U
34
39A
12 0
30
30 A
33 A
21 D
36
17 0
26A
51
35 A
60
70
50
22 A
37 A
hi
24
47 A
53 8
25
44A
10 0
42 A
23
54 .
28i
55 .
52
50
45
40
43
41
46
32
G
A
A
A
A
A
A
130
160
14 0
11 0
80
10
150
48
49
19 0
90
180
Figure 3.3. Consensus tree based on 1,000 replications of genetic relationships among
54 samples of coastal cutthroat trout from across distributional range and a steelhead
outgroup (Suntaheen Creek, Alaska, location 4) constructed with a UPGMA algorithm
from DCE chord distances shown as an unrooted rectangular cladogram. Bootstrap
percentages based on 1,000 replications, only percentages greater than 30% shown
(e.g.. 73). Location numbers correspond to Table 3.1 and Figure 3.1.
the major cluster that included samples from all regions. The sample from Martin
River, Alaska (location 2), appears to be even more distinct in the consensus
dendrogram (Figure 3.3). As with the neighbor joining tree, the relatively long branch
lengths in the dendro grams agree with the heterogenous allele frequency distributions
and suggest significant levels of differentiation among the samples collected.
The slight geographic concordance evident in the cluster analysis becomes more,
evident in the multidimensional scaling (Figure 3.4). The stress value of 0.111 of the
MDS analysis is considered good, a value of 0.05 would he considered excellent, 0.0
perfect, and 0.2 poor (Kruskal 1964a). In the multidimensional space illustrated in
Figure 3.4, samples from region A make up all of the samples in the lower left portion
of the plot (low axis I and axis H values) while only samples from region D are located
in the right hand portion of the plot (high axis I values and moderate to high axis II
values). Geographic affinity is also apparent at smaller spatial scales such as along the
northern Washington coast and Srait of Juan De Fuca where several adjacent sampling
locations grouped together in MDS space (Hoko River location 30, Salt Creek -
location 31, Quillayute River location 32, and Hoh River location 33).
As in the cluster analysis, the steelhead sample (Suntaheen Creek, Alaska) appears
to be quite different than all the coastal cutthroat trout samples when represented by
MDS (Figure 3.5). This difference does not appear to be unique to the Suntaheen
Creek sample, an 0. mykiss sample from Bakeoven Creek (Deschutes River, Oregon)
from Currens (1997) shows phenetic affinity for the Suntaheen Creek sample and both
are quite different from all the coastal cutthroat trout samples (Figure 3.5).
Across their distributional range, coastal cutthroat trout show a pattern of strong
isolation by distance with samples from geographically adjacent locations being more
similar than they are to populations from locations further away. Tests for isolation by
l.(
III -0.
I
1.32
Regional groups
Figure 3.4. Multidimensional scaling (dimensions 1,11 and III) based on allele
frequencies of 17 polymorphic loci (P095) of coastal cutthroat trout from 54 locations.
The stress value for this clustering was 0.111. Location numbers correspond to
numbers in Table 3.1.
0. mykiss
(Bakeoven Creek, Deschutes River, Oregon)
0. mykiss
(Suntahe" Cr1, A11rQ\
0.38
-0.06
III
-0.50
-0.94
-2.95
0.48
Figure 3.5. Multidimensional scaling (dimensions 1,11 and ifi) based on allele
frequencies of 12 polymorphic loci (P095) of coastal cutthroat trout from 54 locations.
The stress value for this clustering was 0.065. Two 0. mykiss samples (Suntaheen
Creek, Alaska, and Bakeoven Creek, Oregon) were included as outgroups. Location
numbers correspond to numbers in Table 3.1. B akeoven Creek data from Currens
(1997).
distance exhibit highly significant associations with genetic distance (pairwise F) and
geographic distance. Significant associations were detected using both the shortest
water distance (Figure 3.6; Mantel r = 0.415, P = 0.0001) and depth constrained water
distance (Figure 3.7; Mantel r = 0.395, P = 0.0001). The association of genetic
distance and geographic distance varied among regions (Figures 3.8 and 3.9) with
significant (P
0.10) associations being observed in region A (Mantel r = 0.23 8, P =
0.079) and region D (Mantel r
0.436, P = 0.099). This is in contrast to genetic
variation that was similar among the regions (Table 3.4). Based on plots of pairwise
FST and geographic distance (Figure 3.8 and 3.9), it is evident that the range in pairwise
distances are different among the regional groups, specially region D where no
pairwise distances greater than 229 km.
Isolation by distance appears to be significant at shorter distances regardless of the
regional area across the distributional range of coastal cutthroat trout. The Mantel
correlogram indicates that the association of genetic distance and geographic distance
(shortest water distance) is strongly positive and significant (P
0.05) at short
distances. In other words, fish from a location are more similar genetically to fish from
nearby locations than they are to fish from distant locations. The correlation becomes
not significantly positive at approximately 740 km (Figure 3.10 and Table 3.6).
Significant negative associations begin to occur at distances greater that 1,243 km.
That is, at distances greater than 1,243 km the genetic differences (i.e., statistically
significant differences) between fish from two sites increased with distance. For Figure
3.10, distances classes greater than distance class 19 (2,137 km) should be ignored
since they are based a small number of pairs and therefore have low statistical power
(Legendre and Legendre 1998) (see also Table 3.6). Regional Mantel correlograms
were not generated because of the low number of pairs in all distance classes.
r.1sI
0.35
S
Mantelr=0.415
S
S
0.30
S
0
.5S . .
0.25
P-value = 0.0001
.
S
.. S..Ss
.'L'...'. t.
%
S
S
..'j;W.\'
;
tp5
S
5
S.
0.20
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0.15
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S s.
S
S
.
.5.
,
S
S
S
S
S
S
S
0.00
0
500
1000
1500
2000
2500
3000
Distance (km)
Figure 3.6. Isolation-by-distance analyses for all coastal cutthroat trout locations
sampled across distributional range. Pairwise FST distances (y-axis) are plotted against
pairwise geographic distances (x-axis) based on the shortest water distance between a
pair of locations. P-values for Mantel test based on permutation test of 10,000 runs.
0.35
S
Mantelr=O.395
P-value=O.000l
S
0.30
S
S
S
S
?. S
0.25
S.
. :..
'.1
.
0.20-
ssIj
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5
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.
.
o
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0
1000
4
:.
S #5
¼1':
C'
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S
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:
S
.:
.ai..:4
.w1s..%
.
%SS
S45
S
S
S
I
1500
S
I
I
I
I
I
2000
2500
3000
Distance (km)
Figure 3.7. Isolation-by-distance analyses for all coastal cutthroat trout locations
sampled across distributional range. Pairwise FST distances (y-axis) are plotted against
pairwise geographic distances (x-axis) based on a depth constrained water distance
between a pair of locations. P-values for Mantel test based on permutation test of
10,000 runs.
A
Mantel r 0.23 8
P-value = 0.079
0.30
S
.
*
0.25-
0
S
0
0.20-
.
0
0
0
S
S.
0.15-
S
S.
0
.0
010-
0
5500
%
0.05-
S
S
.
S
S
S
0
S
S
5
.
.
S
0
S
S
S
S
S
S
.
S
0
S
S
.
0.00
Os
0
IS
1000
Is
500
S
1500
0.35
0.30
0.25
0.20
I
0.15
0.10
0.05
0.00
0
500
1000
1500
Distance (km)
Figure 3.8. Isolation-by-distance analyses for coastal cutthroat trout locations in
regional groups A (upper plot) and B (lower plot). Pairwise FST distances (y-axis) are
plotted against pairwise geographic distances (x-axis) based on the shortest water
distance between a pair of locations. P-values for Mantel test based on permutation
test of 10,000 runs. Regional groups are defined in Figure 3.1.
0.351
C
Mantel r= 0.159
P-value = 0.179
0.30
0.251
S
S
S
0.20.
:
015
S
S
5. si.5Ss
S
0.10-
S
S
S
S
;s'?.
, II
5
55
U.
S
555*
S
0,05- .
5
S
S
*
S
S
'S
0.00-
0
500
1000
1500
0.35
6
0.30
0,25
0.20
0.15
0.10
0.05
0.00
0
500
1000
1500
Distance (km)
Figure 3.9. Isolation-by-distance analyses for coastal cutthroat trout locations in
regional groups C (upper plot) and D (lower plot). Pairwise FST distances (y-axis) are
plotted against pairwise geographic distances (x-axis) based on the shortest water
distance between a pair of locations. P-values for Mantel test based on permutation
test of 10,000 runs. Regional groups are defined in Figure 3.1.
0.5
'-4
-0.5
-1.0
-1.5
0
500
1000
1500
Distance (1cm)
2000
2500
Figure 3.10. Multi-variate correlogram (non-linear Mantelogram) representing
autocorrelation statistics plotted against distance (shortest water distance). Dark
circles represent significant autocorrelation statistics (P 0.05) for a distance class. Pvalues, sample size, and distance class delineation presented in Table 3.7.
,J1
Table 3.6. Mantel correlogram values represented in Figure 3.13. Distances based on
the shortest water distance between a pair of locations.
Distance class
Mid-point
(km)
N
Mantel r
P-value
1
125
65
0.65
0.0002
2
237
100
0.67
0.0001
3
348
99
0.44
0.0001
4
460
98
0.33
0.0026
5
572
91
0.35
0.0020
6
684
71
0.26
0.0375
7
796
97
0.14
0.1828
8
907
71
-0.01
0.9115
9
1019
73
0.05
0.6899
10
1131
96
-0.03
0.7534
11
1243
87
-0.37
0.0016
12
1354
72
-0.41
0.0006
13
1466
71
-0.16
0.1887
14
1578
57
-021
0.1208
15
1690
54
-.0.37
0.0114
16
1802
46
-0.21
0.1637
17
1913
50
-0.57
0.0003
18
2025
37
-0.53
0.0026
19
2137
32
-0.55
0.0047
20
2249
20
-0.42
0.0747
21
2361
12
-1.26
0.0001
22
2472
12
-1.59
0.0001
23
2584
4
-1.03
0.0405
24
2696
5
-1.04
0.0271
25
2808
11
-1.30
0.0002
92
Twenty eight of 165 pairs of associations in the first two distance classes in the
Mantel correlogram were from locations in region D and suggest that the regional
differences in isolation by distance resulted from the disparity in the distribution of
distance classes among the regional groups. The use of depth constrained water
distance provided similar results (Figure 3.11 and Table 3.7). The correlation becomes
not significantly positive at distances of approximately 825 km.
3.5. DISCUSSION
Across the distributional range of coastal cutthroat trout, genetic population
structure was primarily at the individual stream level (i.e., sampling location in this
study). This survey was based on samples collected over a five year period, providing
a snap shot of genetic variation and the distribution of this variation that existed from
1992 to 1996. Significant genetic variation was found among 54 sample sites
consisting of 1,413 individual coastal cutthroat trout from across their distributional
range. In the only survey to date that has examined populations across their
distributional range, I found regional differences in how genetic variation was
distributed across the landscape even though the amount of genetic variation within
regions was similar. Geographic concordance of populations at the northern and
southern extent of the subspecies range was observed and isolation by distance was
evident at regional scales (<800 km).
My results based on samples collected across the range of the subspecies were
consistent with the hypothesis that coastal cutthroat trout from each sampling location
were from genetically distinct, isolated populations. When examining intra-regional
patterns of genetic population structure, my results were not inconsistent with those
reported by others using allozymes (Campton and Utter 1987) and microsatellites
93
-S
\.
0.5
S
S
0
0
0.0
0
0
\ /°\
0
/
.
-,
-0.5
/\
\
\/
S
S
\
-1.0
\/\
\
S
0
500
1000
1500
2000
2500
3000
Distance (km)
Figure 3.11. Multi-variate correlogram (non-linear Mantelo gram) representing
autocorrelation statistics plotted against distance (depth constrained water distance).
Dark circles represent significant autocorrelation statistics (P 0.05) for a distance
class. P-values, sample size, and distance class delineation presented in Table 3.8.
Table 3.7. Mantel correlogram values represented in Figure 3.14. Distances are based
on depth constrained water distance.
Distance class
Mid-point (km)
N
Mantel r
P-value
1
137
66
0.65
0.0001
2
262
88
0.69
0.0001
3
387
105
0.40
0.0001
4
512
114
0.28
0.0055
5
637
80
0.41
0.0008
6
762
97
0.26
0.0154
7
887
76
0.14
0.2260
8
1011
67
0.11
0.4239
9
1136
76
-0.04
0.7461
10
1261
78
-0.31
0.0094
11
1386
79
-0.18
0.1305
12
1511
70
-0.47
0.0001
13
1636
81
-0.20
0.0710
14
1761
51
-0.11
0.4373
15
1886
68
-0.30
0.0212
16
2010
60
-0.35
0.0096
17
2135
41
-0.41
0.0142
18
2260
31
-0.64
0.0011
19
2385
29
-0.28
0.1692
20
2510
17
-0.65
0.0115
21
2635
24
-0.83
0.0004
22
2760
13
-1.16
0.0002
23
2885
4
-1.03
0.0395
24
3009
4
-1.32
0.0104
25
3134
12
-1.19
0.0003
95
(Wenburg et al. 1998; Wenburg and Bentzen 2001). Wenburg and Bentzen (2001)
found significant heterogeneity among all populations and pairs of populations in tests
of genotypic differentiation from coastal cutthroat trout collections from the Hood
Canal, Washington.
The amount of variation observed within each regional area was significantly
different than zero and did not differ among regions. Genetic diversity as measured by
various measures of genetic diversity (e.g., heterozygosity, number of polymorphic
alleles) was similar across the range of coastal cutthroat trout. The contribution to the
total gene diversity was greater among populations within regions (8.6%) than among
regions (2.5%), while 88.9% of the total gene diversity was within populations.
Campton and Utter (1987) found allele frequency differences between the north Puget
Sound and Hood Canal regions contributed the most to the total genetic variation
among the coastal cutthroat trout populations they examined while differences among
basins within a region only contributed a small proportion to the total genetic variation
observed. The contrast of my results to those of Campton and Utter (1987) are a result
of the regional groupings made in both studies. All of the populations sampled by
Campton and Utter (1987) and placed into two regions would have been included in a
single geographical region in my study. Similarly, Beacham et al. (2003) delineated
samples within the Fraser River drainage into six regions and found that the genetic
population structure of Fraser drainage Chinook salmon had a strong geographic basis
with regional differences accounting for approximately twice the variation in allele
frequency as within regional variation among populations. Beacham et al. (2003)
sampled at a finer spatial scale than I did and the geographic extend of their six
regions would be represented by a single, larger regional group at that spatial scale
used in my analysis. Reisenbich1r et al. (1992) found regional differences between
northern and southern populations of steelhead sampled between the Columbia River
and Mad River, California, but not within basin or intra-regional genetic
differentiation (10 allozyme loci). The spatial extend of the regional scale used by
Campton and Utter (1987), Beacham et al. (2003), Reisenbichier et al. (1992) and in
other studies are smaller than the regional spatial scale used in my study and this helps
explain the relatively large contribution of among population gene diversity within a
region that I observed versus the low within region but high among region gene
diversity reported in other studies.
Compared to other species of anadromous salmonids, the amount of genetic
diversity attributed to differences among populations was high in coastal cutthroat
trout (Table 3.8). This is consistent with other studies that have been conducted at
smaller spatial scales and suggest a range-wide consistency in the population structure
of coastal cutthroat trout. Slight geographic concordance was observed in both the
neighbor-joining and UPGMA clustering approaches for samples located at the
northern and southern extend of the range. Although bootstrap support was weak, my
results suggest slight regional grouping of coastal cutthroat trout throughout their
range, particularly at the northern and southern extend of their range. Campton (1987)
found anadromous coastal cutthroat trout in the Puget Sound were primarily structured
at the between-region (North Puget Sound versus Hood Canal) and among-stream
levels (streams within a drainage). Wenburg and Bentzen (2001) proposed that the
strongest contributor to population structure of anadromous coastal cutthroat trout was
differentiation at the level of individual creeks in their survey that included 22 sites
from the Hood Canal, Puget Sound, and several coastal Washington sites.
The analysis of isolation by distance based on the Mantel correlograms indicated
that at regional scales (i.e., distances <800 km) populations of coastal cutthroat trout
exhibited a strong pattern of isolation by distance. The strength of IBD differed among
Table 3.8. A summary of genetic diversity analyses of selected salmonid species.
No. of
Species
O. populations
0. clarki clarki
Geographic range
throughout range
0.13 1
0. clarki clarki
Washington state
No. of
loci
Source
54
Marker
allozymes
30
this study
0.121
16
microsatellites
6
Wenburg et al. (1998)
0.058
0.095
21
allozymes
allozymes
31
7
41
Campton and Utter (1987)
Griswold (1996)
0. clarki clarki
0. clarki clarki
Puget Sound
Elk River,
0. clarki clarki
Vixen Inlet, Alaska'
0.016
4
aliozymes
41
Griswold (1996)
0. tshawytscha
Fraser River
0.039
13
microsatellites
52
Beacham et al. (2003)
Alaska
0.099
32
microsatellites
9
Olsen et al. (2003)
0.161
11
allozymes
7
Koljonen et al. (1999)
Indian Bay, Newfoundland
0.063
8
microsatellites
5
Adams and Hutchings (2003)
Denmark
0.032
5
microsatellites
8
Hansen et al. (2002)
Maine
0.020
14
microsatellites
11
Spidle et al. (2003)
Alaska Washington
0.023
13
microsatellites
5
Olsen et al. (1998)
Sainte-Marguerite River, Canada 0.034
14
microsatellites
5
Garant et al. (2000)
0. kisutch
Salmo salar
Salvelinus fontinalis
Salmo trutta
S. salar
0. gorbuscha
S. salar
Baltic
Oregona
Seac
Salvelinus confluentus
British Columbia
0.39
37
microsatellites
7
Costello et al. (2003)
Salvelinus confluentus
Northwest United States
0.659
65
microsatellites
4
Spruell et aT. (2003)
Maine and Quebec, Canada
0.2 16
30
microsatellites
6
Castric et al. (2001)
S. fontinalis
a included three sites that were upstream of migration barriers.
b three of the four sites located above a barrier, the three above barrier populations were not separated by barriers.
c Koijonen et al. (1999) analyzed both hatchery and wild populations, values in this table are for wild populations only.
regions and was strongest in the northern and southern regions. Wenberg and Bentzen
(2001) found genetic distances between populations of coastal cutthroat trout in the
Hood Canal, Washington, were positively correlated with geographic distance at
spatial scales smaller than that of this study; but partial reproductive isolation was still
evident with significant differences in genotype frequencies found between sampling
locations separated by as little as 2 km. At a spatial scale larger than the Hood Canal,
Wenburg et al. (1998) found significant correlation between genetic distance and
geographic distance among 13 coastal cutthroat trout populations throughout
Washington using DCE but not when using °ST and PST Stronger IBD at smaller spatial
scales is also suggested by Wenburg and Bentzen (2001) that found two of three
genetic distance measures significantly correlated with geographic distance within the
Hood Canal but only one of three genetic measures significantly correlated when
samples included sites throughout Washington.
Others have found similar patterns of population structure in salmonids. Recently
Olsen et al. <1898 Id> found patterns of coho salmon genetic population structure
similar to patterns found in my study of coastal cutthroat trout. Olsen et al. (2003)
reported significant intra-regional population differentiation, large among population
differentiation (FST= 0.099), phenetic clustering only exhibited weak geographic
concordance, and IBD was strong at small spatial scales. Fontaine et al. (1997) found
that the relative positions of Atlantic salmon populations in a neighbor-joining
phenogram did not correspond with their geographical proximity. They found an intraregional correspondence between genetic distance and geographic location that broke
down when interregional populations where included in their analysis (Fontaine et al.
1997). Estoup et al. (1998) found poor congruence between dendrogram topology and
geographic distance from 11 nonanadromous brown trout populations separated by 2.5
to 850 km (Vosges massif, France). Although samples from adjacent locations were
significantly different indicating limited gene flow (Estoup et al. 1998), strong IBD
and a significant tendency for errors in assignment tests to assign individuals to one of
the nearest populations indicated a high level of microgeographic structuring of these
brown trout populations. Castric et al. (2001) found their population phenogram
illustrated an overall lack of population grouping by drainage or any other hierarchical
clustering among 30 populations of brook charr in six major river drainages in Maine.
In addition, Castric et al. (2001) found reversed IBD (i.e, greater genetic similarity
with increased geographic distance). Costello et al. (2003) found significant IBD
among populations of bull trout in British Columbia and Alberta, Canada, but weak
bootstrap support for a neighbor-joining tree that tended to group populations by
physiogeographic region.
Patterns of genetic population structure found in this study along the Pacific coast
from Vancouver Island, British Columbia, to Cape Blanco, Oregon (regional area C in
this study) and non-coastal basins in Puget Sound, Strait of Juan de Fuca, and Strait of
Georgia (regional area B in this study) may be a result of Pleistocene glaciation or an
artifact of the many years of hatchery releases; including the introduction of hybrids,
that has occurred in this region (Johnson et al. 1999). These two regional areas (A and
B) had the weakest IBD yet still had significant genetic variation (0> 0). Cutthroat
trout and steelhead hybrids ("cutthroat cross trout") were released as early as 1912 by
the State of Washington in the Nisqually River, Puget Sound, Washington (Riseland
1912). Hatchery coastal cutthroat trout were introduced in British Columbia streams as
early as 1930 (Hartman and Gill 1968), although Hartman and Gill reported that
introductions were into areas already occupied by coastal cutthroat trout, were
generally not successful, and were unlikely to have influenced distribution of coastal
cutthroat trout in their survey area. In addition to the release of hybrids, out of basin
transfers have occurred. As a result of poor returns from an anadromous coastal
IDI
cutthroat trout program in Washington in the 1950s, hatchery coastal cutthroat trout
from the Alsea River in Oregon were brought in followed by crosses with steelhead in
the late 1 960s (Crawford 1979); these efforts to establish high return rates of hatchery
coastal cutthroat trout were not successful (Crawford 1979). Recent hatchery
production of anadromous coastal cutthroat trout has been focused on providing sport
fishing opportunities and has occurred primarily in Puget Sound, Grays Harbor, the
Lower Columbia River, and the Oregon coast (Johnson et al. 1999), although since
1997 no releases of hatchery coastal cutthroat trout have occurred in Oregon coastal
streams that contain anadromous coastal cutthroat trout (Hooton 1997). Recent genetic
surveys of Washington coastal cutthroat hatcheries found hybrids in the Aberdeen Fish
Hatchery and the Beaver Creek Fish Hatchery, Colombia River (Johnson et al. 1999).
The meristic rule used to identify 0. mykiss individuals resulted in very distinct
separation of steelhead from coastal cutthroat trout in the clustering and ordination
procedures. Other molecular markers have been developed (Wenburg et al. 1998;
Ostberg and Rodriguez 2002) that would provide greater resolution for species
identification than the allozymes used in this study and may provide additional insight
into 0. o. clarki and 0. mykiss hybrid issues in future analyses of my samples,
especially in studies designed specifically to examine hybridization.
Determining genetic structure can be complicated by hybridization. The possibility
of hybridization with steelhead exists for many populations of coastal cutthroat trout.
Wilson et al. (1985) used mtDNA sequencing to analyze populations of steelhead and
rainbow trout (anadromous and resident forms, respectively) and a population of
coastal cutthroat trout. With few exceptions, their results indicated detectable
divergence among all geographic populations of steelhead and rainbow trout.
hiterspecific nucleotide divergence between 0. mykiss and 0. c. clarki was two- to
four-fold larger than intraspecific divergence detected within 0. mykiss. Based on a
101
base substitution rate of 2% per million years (estimate based on mutation rate
calibration determined from primates therefore, used with caution), Wilson el al.
(1985) estimated that 0. mykiss
0. clarki clarki divergence occurred 1 to 2 million
years ago. Behnke (1992) speculated that in small streams where two species of trout
are found, crossbreeding occurs if reproductive isolation (e.g., limited niche diversity,
insufficient space to allow physical separation at spawning) breaks down. Campton
and Utter (1985) used allozymes to identify juvenile fish with intermediate phenotypes
consistent with those expected for decedents of coastal cutthroat trout and steelhead
and was the first formal documented observation of hybrids in the wild, and led the
authors to suggest that in streams where both species spawn "the production of hybrid
offspring may not be uncommon". More recently, many genetic studies have detected
the occurrence of hybrids between coastal cutthroat trout and steelhead in the wild
(Hawkins 1997; Wenburg et al. 1998; Young et al. 2001).
Surprisingly, no samples appeared to be intermediate to 0. mykiss and 0. clarki in
multidimensional space (Figure 3.5). The MDS ordination analysis of my samples that
included two outgroups of 0. mykiss did not exhibit any populations intermediate to
the two species. Based on reports in the literature that concluded that hybridization
between coastal cutthroat trout and 0. mykiss to be widespread throughout the range of
coastal cutthroat trout populations on the West Coast (Johnson et al. 1999), one might
expect some of my samples to include hybrids that would be intermediate to steelhead
and coastal cutthroat trout in the multidimensional space. This lack of intermediate
populations may be due to differences in the loci and number of loci used in the
different studies, my application of an independent morphological rule (short maxillary
extension and absence of basibranchial teeth) to exclude misidentified 0. mykiss from
the analyses, and the inclusion of older fish (fish >1+) in many of my samples.
Johnson et al. (1999) did observe hybrid individuals most often among age 0+ and 1+
102
fish and seldom in adults. In addition, Johnson et al. (1999) used a larger set of loci
and used three fixed or nearly fixed loci (ADA-2
mAH-2 *, and CK4-2 *) to identify
and exclude 0. mykiss individuals that had been inadvertently included in their
samples and used a decision rule based on eight loci (sAAT4*, ADA2*, mAH2*,
mAH-3 <, CKA 2*, IDDH-1
sIDH2*, and sMEP2 *) to attempt to identify and
exclude suspected hybrid individuals from their analyses. Consistent with my results
that did not exclude any suspected hybrids, Johnson et al. (1999) reported that coastal
cutthroat trout samples in their analysis were closely clustered together and were well
differentiated from steelhead after excluding suspected hybrids based on the presence
of steelhead like alleles.
In addition, surveys such as mine were not designed to estimate hybridization and
should be viewed with caution when inferring range-wide hybridization patterns. In
my study, no individuals other than 0. mykiss (based on, the absence of basibranchical
teeth and a short maxillary extension) were removed from the analysis. I did not use or
have results across all samples for many of the loci used by Johnston et al. (1999) to
identify hybrids. The reduced loci set I used may have limited my ability to detect the
extent of hybridization observed by others. My samples were collected by a limited
number of people, perhaps providing a bias against collecting hybrids. Recently
Baumsteiger (2002) found that expert evaluators were almost always able to
successfully identify coastal cutthroat trout in the field, but had moderate difficulty
identifying steelhead and were not able to correctly identify hybrids. In addition, the
morphological rule I used to identify 0. mykiss may have eliminated hybrids from the
analysis resulting in the lack of intermediate populations in the MDS analysis. I did not
perform analyses on individuals, rather allele frequencies of populations were used
perhaps masking the presence of hybrid individuals.
103
Recently, studies to examine hybridization between coastal cutthroat trout and
steelhead found that the occurrence of hybrid coastal cutthroat trout and steelhead was
not randomly distributed among populations they sampled in the Puget Sound and
Washington coast region (Young et al.
2001).
Campton and Utter (1985) concluded
that a post zygotic isolating mechanism may be contributing to the preservation of
species integrity given the infrequent observation of adult hybrids. Behnke
(1992)
speculated that although gene flow may occur between coastal cutthroat trout and
coastal rainbow trout (i.e., steelhead), reproductive isolation typically does not break
down completely and it is uncommon to find a hybrid swarm typically found when
interior cutthroat trout species hybridize. Hawkins and Foote
(1998)
did not find
reduced viability of hybrids between coastal cutthroat trout and steelhead in hatchery
experiments, but did report a mismatch between egg size and development rate relative
to pure species that may "play an important role in limiting introgression between the
species."
These data suggest that compared to other species of Pacific salmon and trout,
coastal cutthroat trout are characterized by many smaller, genetically more diverse
local populations that act in a more independent, isolated nature over short time frames
(<100 years). For example, on average approximately 11% of all genetic differences
we observed (total gene diversity) was attributed to differences among populations (0
= 0.131). This was much greater than that reported for other species of anadromous
salmonids in western North America. Consistent differences occurred among
populations from different regions (Alaska, Washington/Oregon, and California), but
within regions there was less geographical structure than that observed among
populations of other Pacific salmon and trout. Genetically, this can be explained by
genetic drift (common in small populations), by founder effect (common in small
104
populations in areas with dynamic habitat conditions or where species are colonizing
new habitat), and by periods of low dispersal.
I believe that differences in genetic population structure between coastal cutthroat
trout and other Pacific salmon and trout may reflect life-history differences among
species. The many migratory life-histories that coastal cutthroat trout have evolved, for
example, would be advantageous in unpredictable habitats where small populations
expand, decline, or become extinct with changing conditions. Dispersal, which would
be more likely with fish that migrate, might be rare or consist of only few individuals,
but they may be crucial for reestablishing populations in adjacent habitat following
localized extinctions or as new habitat becomes available.
Understanding genetic population structure across the landscape is critical to
developing appropriate conservation and management plans, but it will not paint a
complete picture of what is needed for development of appropriate conservation plans.
Knowledge of life history differences adds to this picture, but to bring it all into focus
one needs to also understand the habitat dynamics that coastal cutthroat trout have
adapted to and will be faced within the future. Connectivity of biological and physical
processes throughout the basin must be considered for management, conservation, and
restoration efforts (Reeves et aL 1995).
The slight geographic concordance of populations at the northern and southern
extend of the range and the regional separation of the southern populations suggest
coastal cutthroat trout populations at the periphery of the range have faced different
habitat conditions (both marine and/or freshwater) than populations located in the
central part of the distribution. Studies of other organisms have found that in general,
peripheral populations tend to be genetically and morphologically divergent from
central populations (Lesica and Allendorf 1995). Lesica and Allendorf (1995)
reviewed various aspects of peripheral populations and their value to species
105
conservation with special attention to the distinction between marginal versus
peripheral habitat conditions. The southern populations occupy habitat that
ecologically marginal in addition to being at the geographic margins of the
distributional range. Southern populations have experienced different habitat
conditions (i.e., Kiamath Province, etc.) than adjacent populations along the Oregon
coast and throughout the range of coastal cutthroat trout whereas the northern
populations occupy areas that are similar to conditions throughout a large part of the
distributional range. Coastal basins in Oregon south of, and including the Sixes River,
drain the Klamath Mountains and the unique geology that makes up the Klamath
Mountains Geologic Province (McKee 1972). Sampling locations from the region
include locations 48 to
55.
Conservation units for coastal cutthroat trout need to reflect appropriate geographic
areas based on the distribution of genetic diversity and life history variation among
populations. For instance, because small populations are typically at greater risk of
extinction than larger populations and because of the high gene diversity found in
coastal cutthroat trout, conservation measures designed for other Pacific salmon and
trout populations with larger, more connected populations may not be appropriate for
coastal cutthroat trout. The recent findings of Channell and Lomolino (2000) that
found as a species becomes endangered they tend to persist in the periphery of their
historical geographical range places a greater emphasis on these peripheral populations
of coastal cutthroat trout when conservation plans are developed. Although there is no
indication that there has been a contraction of the distribution of coastal cutthroat trout,
long-term conservation plans are likely to depend on protection of distinct populations
(Lesica and Allendorf
1995),
particularly peripheral populations in marginal habitats
that may contain high adaptive significance to the species as a whole (Scudder
1989).
The connections and interactions among local populations that are important for both
106
regional and local persistence are likely to be disrupted by management. Risks of local
extinction, both deterministic and stochastic, may be partially addressed through
management actions at the stream or watershed scale; long-term persistence will
require consideration at a much larger scale.
107
4. CONCLUSION
The results of my genetic and meristic survey of coastal cutthroat trout populations
across their distributional range found many diverse local populations with interregional differences in the distribution of that diversity across the landscape. These
data suggest that compared to other species of Pacific salmon and trout, coastal
cutthroat trout are characterized by more diverse local populations that act in a more
independent, isolated nature. The amount of sub-population genetic structure in my
study did not differ among regions, but the strength of isolation by distance differed
among regions with the strongest isolation by distance being detected at the northern
and southern extent of the range (Chapter 3). The northern and southern populations
exhibited geographic concordance based on allozymes (Chapter 3), but only the
southern populations exhibited meristic phenetic affinity at the regional spatial scale
(Chapter 2). The southern populations were relatively distinct in terms of their
phenetic affinity (i.e., the southern populations grouped together). In addition, the
among population differences in meristic characters observed in the southern region
were greater than those observed within the other three regional areas.
Although both meristics characters and allele frequencies exhibited significant
differences among populations, the pair-wise measures of between population
distances were not concordant (Figure 4.1). Lynch (1996) found such lack of
concordance not surprising for reasons related to statistical sampling and the weak
relationship between molecular and quantitative-genetic variation. Additionally, it has
been found that morphological and molecular evolution can occur at independent rates
(Avise 1994). Salducci et al. (2004) recently found a discordance in resolution between
population subdivision of the cyprinid
Leuciscus souffia soufJia
using aliozymes and
meristic characters; they attributed the lack of concordance to insufficient differences
Ip
0.05
e)
0
S
0.03
S
.
S
I
.
.
:.
I
:
0.02
: ..
S
0.01
S
:..'.
%*-4
S
I .
z
aS..
3.
.
:. S 3
S
I.
.
5
)
. 3
..
S
3
'. .4*
,
0.00
0.05
0.10
0.15
S
S
.
II
.
. ..
:
.
.
S
0.20
S
0.25
0.30
0.35
Pairwise genetic distance
Figure 4.1. A scatter plot of pairwise genetic distance (0) versus pairwise meristic
distance (Euclidian distance) for all locations of coastal cutthroat trout sampled.
109
in the environmental conditions to drive morphological adaptation or that population
subdivision was too recent for diversification of morphological traits to have occurred.
The lack of concordance I observed could be a result of many different scenarios,
including strong environmental influence on meristic characters or differential
selection acting on the characters and loci. Another factor contributing to the lack of
concordance between population structure based on meristic and allozyme characters
is that the meristic measures are phenotypic traits that are very likely polygenic traits
(i.e., the expression is a function of appropriate alleles at numerous loci (Avise 1994))
whereas the allozyme analyses used the frequency of various genotypes to characterize
a population.
The implications of this lack of concordance between these two measures of
population differentiation are not restricted to these meristic or allozyme derived
population studies. Conservation planning for any species requires an understanding of
the interaction among populations across different geographic scales and a
maintenance of historical genetic population structure. However, efforts might be
better directed if they focused on maintaining adaptive diversity rather than relying on
a measure that reflects genetic isolation (Crandall et al. 2000). The extant in which
either my meristic or allozyme measures captures a trait that enhances the potential for
coastal cutthroat to persist is unknown. Until further is known about the adaptive
significance of the population differences I detected, a conservative approach to the
use of these data would call for a recognition of the population subdivision suggested
by both the meristic and genetic measures (i.e., southern populations distinct,
geographic affinity of populations at the northern and southern portion of the range,
and a primary population structure at the individual stream level).
The strong geographic concordance and inter-regional divergence of meristic
characters exhibited by the southern populations (Chapter 2) was consistent with other
110
ecological studies that have found peripheral populations tend to be genetically and
morphologically divergent from central populations and morphological characters are
expected to diverge more rapidly in isolated populations than gene frequencies (Lesica
and Allendorf 1995).
The persistence of a population or group of populations (e.g., metapopulation) is
dependent on the presence of suitable ecosystem states and the ability of populations to
track and use these patches through time. My snap-shot of population structure across
the range of coastal cutthroat trout provides a glimpse of past landscape dynamics
since the current population structure is a reflection of dispersal and isolation that have
occurred over a range of time scales. The landscape within the distributional range of
coastal cutthroat trout is dynamic in space and time ranging from the geologic
processes of glaciation and volcanism that have shaped a large portion of the area
(McPhail and Lindsey 1986) to other events such as fire and flood have shaped the
landscape at smaller temporal
(101
-
102
years) and spatial scales (watersheds and
basins) (Benda 1994). These various disturbances create a shifting mosaic of abiotic
and biotic conditions (Reeves et al. 1995).
Across the range of coastal cutthroat trout, the pattern of temporal and spatial extent
(i.e., latitudinal extent) of disturbances can be thought of as being infrequent but
spatially large in the north and shifting to disturbances of a more frequent and smaller
spatial scale to the south (McPhail and Lindsey 1986; Benda et al. 1998; Naiman and
Bilby 1998). This is conceptually illustrated in Figure 4.2. In this simplified construct,
anthropogenic disturbances represent a range of processes from land management
activities to urbanization, and tectonics include processes such as coastal uplift and
events such as the 1964 Alaska earthquake. The disturbance regime that coastal
cutthroat trout encounter in the north is vastly different
111
North
Juneau
1
.i:L"'.:
.....
a
p.
BC/WA
tL..
.1
.
..
WA/OR
..
OR/CA
a
'.
.
...
South
Infrequent
Frequent
Temporal scale
Figure 4.2. A simplified construct of the temporal scale (horizontal axis) and
latitudinal scale (vertical axis) of disturbances in coastal basins across the
distributional range of coastal cutthroat trout.
112
from that in the south. Peripheral populations of coastal cutthroat trout at the northern
extent of the distribution (i.e., Alaska) occur in habitat that is marginal is terms of the
distributional range, but not condition, whereas coastal cutthroat trout populations at
the southern extent of the range (i.e., southern Oregon/Northern California) exist in
habitat that is marginal in terms of the distributional range and condition (i.e., Kiamath
Province, etc.). Coastal basins in Oregon south of, and including the Sixes River, drain
the Kiamath Mountains and the unique geology that makes up the Kiamath Mountains
Geologic Province. Tracking changes across the landscape in response to different
environmental conditions depends on distance and habitat conditions along the route of
dispersal. Coastal cutthroat trout populations at the southern extent of the range are
subject to smaller, but more frequent disturbances that I believe have resulted in less
exchange of individuals with populations to the north (i.e., Oregon coast) resulting in
regional distinctiveness and local extinction/recolonization events resulting in higher
levels of within region diversity.
Variations in habitat at various spatial and temporal scales will be reflected in any
snap shot of population structure. Taylor et al. (1993) used the term landscape
connectivity to express the degree to which the landscape facilitates or impedes
movement among areas. The opportunity for movement across the landscape during
various portions of its life cycle (e.g., spawning, over-wintering, ocean migration) is
critical for the survival of a local population and opportunities for individuals to move
among populations (i.e., stray) are critical for recolonization following local
extinctions. Movement in the marine environment can influence the observed
population structure of anadromous salmonids. Avoidance of deep water areas in the
marine environment by coastal cutthroat trout has been suggested (Jones and Seifert
1997) although test for isolation by distance were similar when either shortest water
distance or water-depth constrained distance were used (Chapter 3). At this scale,
113
limited dispersal over deep water did not appear to be infiucencing popu'ation
structure.
In summary, the persistence of a population or group of populations depends the
ability of populations to track changes in the environment through time and this
movement of individuals across the landscape depends on the availability of suitable
habitat features at various spatial and temporal scales. This suggests that understanding
the historical and current spatial structuring of populations is useful for conservation
and management of coastal cutthroat trout. Management and long-term conservation
plans are likely to depend on protection of distinct populations (Lesica and Allendorf
1995), particularly peripheral populations in marginal habitats that may contain high
adaptive significance to the species as a whole (Scudder 1989). In contrast to
conventional thinking that predicts core populations persisting and range contraction as
a species becomes endangered, most species (of 245 examined) persist in the periphery
of their historical geographic range (Channell and Lomolino 2000). Habitat that was
suboptimal historically along the periphery of a species range may provide valuable
opportunities for conservation planning (Channell and Lomolino 2000), for it is in
these areas one often finds divergence from central populations (Lesica and Allendorf
1995). For coastal cutthroat trout, the unique meristic characteristics of populations in
the southern extent of the range suggest that these populations may be essential to
conserve the range of diversity in the sub-species. But as this study has shown,
assuming similar population structure across regions would be inappropriate. Rangewide conservation planning must account for differences in landscape dynamics and
local population response.
114
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129
APPENDIX FIGURES
130
12.0
11.5
itO
0. mykiss
(n97)
0. c. clarki
(n
1430)
0. mykiss
(n97)
"hybrid" 0. c. clarki
(n226)
(n
1204)
13.0
I
12.0
0
11.0
0. mykiss
(n
97)
0. c. clarid
(n
1429)
0. mykiss "hybrid" 0. c. clarki
(n
97)
(n
225)
(n
1204)
Appendix Figure A2.1. Mean (± 1 SE) counts of anal pterygiophores and dorsal
pterygiophores for A: fish identified as steelhead or coastal cutthroat trout and B:
steelhead, suspected hybrid, or coastal cutthroat trout based on basibranchial teeth
and the length of the maxillary extension. Fish designated as steelhead had no
basibranchial teeth and a maxillary extension that did not extend beyond the posterior
portion of the eye, fish designated as suspected hybrids exhibited only one the two
attributes used to designate a steelhead.
131
64
63
62
61
60
0. myldss "hybrid" 0. c. clarki
(n=98)
(n=225)
(n1196)
0. mykiss
0. c. clarki
(n=98)
(n=1421)
0. mykiss
0. c. clarki
0. mykiss
(n=98)
(n=1431)
(n=98)
14
:E
12
10
6
4
2
0
"hybrid" 0. c. clarki
(n=226)
(n1205)
Appendix Figure A2.2. Mean (± 1 SE) counts of vertebrae and basibranchial teeth for
A: fish identified as steelhead or coastal cutthroat trout and B: steelhead, suspected
hybrid, or coastal cutthroat trout based on basibranchial teeth and the length of the
maxillary extension. Fish designated as steelhead had no basibranchial teeth and a
maxillary extension that did not extend beyond the posterior portion of the eye, fish
designated as suspected hybrids exhibited only one the two attributes used to
designate a steelhead.
132
150
145
140
135
130
125
120
0. mykiss
(n
97)
0. c. clarki
(n =
1429)
0. mylciss
(n = 97)
"hybrid" 0. c. clarki
(n =
224)
(n =
1205)
.1
33
31
0
.0
i
25
0. mykiss
(n
98)
0. c. clarki
(ii
1427)
0. myldss
(n = 98)
"hybrid" 0. c. clarki
(ii = 223)
(11 = 1204)
Appendix A2.3. Mean (± 1 SE) counts of scales in lateral series and scales above
lateral line for A: fish identified as steelhead or coastal cutthroat trout and B:
steelhead, suspected hybrid, or coastal cutthroat trout based on basibranchial teeth
and the length of the maxillary extension. Fish designated as steclhead had no
basibranchial teeth and a maxillary extension that did not extend beyond the posterior
portion of the eye, fish designated as suspected hybrids exhibited only one the two
attributes used to designate a steelhead.
133
10
0. mykiss
0. c. clarki
(n = 98)
(n = 1422)
0. mykiss
(n
98)
"hybrid" 0. c. clarki
(n 225)
(n
1197)
15
14
13
12
0. mykiss
0. c. clarki
0. mykiss
(n=97)
(n=1419)
(n97)
"hybrid" 0. c. clarki
(n=223)
(n=1196)
Appendix Figure A2.4. Mean (± 1 SE) counts of pelvic fin rays and pectoral fin rays
for A: fish identified as steelhead or coastal cutthroat trout and B: steelhead,
suspected hybrid, or coastal cutthroat trout based on basibranchial teeth and the
length of the maxillary extension. Fish designated as steelhead had no basibranchial
teeth and a maxillary extension that did not extend beyond the posterior portion of
the eye, fish designated as suspected hybrids exhibited only one the two attributes
used to designate a steelhead.
134
12
I
11
10
0. mykiss
0. c. clarki
0. rnyldss
(n=93)
(n='1417)
(n93)
0. mykiss
0. c. clarid
0. mykiss
(n=77)
(n=1345)
(n77)
"hybrid" 0. c. clarki
(n224) (ii'1196)
46
45
1
42
41
40
"hybrid" 0. c.
(n=216)
clarid
(n=1129)
Appendix Figure A2.5. Mean (± 1 SE) counts of branchiostegal rays and pyloric
caeca for A: fish identified as steelhead or coastal cutthroat trout and B: steelhead,
suspected hybrid, or coastal cutthroat trout based on basibranchial teeth and the
length of the maxillary extension. Fish designated as steelhead had no basibranchial
teeth and a maxillary extension that did not extend beyond the posterior portion of
the eye, fish designated as suspected hybrids exhibited only one the two attributes
used to designate a steelhead.
135
7
0.
0. mykiss
0. c. clarki
0. mykiss
(n=96)
(n=1426)
(n96)
0. mykiss
0. c. clark!
(n1425)
0. mykiss
"hybrid" 0. c. clarki
(n=223)
(nI2O3)
13
11
(n=96)
(n=96)
"hybrid" 0. c. clark!
(n=222)
(n1203)
Appendix Figure A2.6. Mean (± 1 SE) counts of upper arch gill rakers and lower
arch gill rakers for A: fish identified as steelhead or coastal cutthroat trout and B:
steelhead, suspected hybrid, or coastal cutthroat trout based on basibranchial teeth
and the length of the maxillary extension. Fish designated as steelhead had no
basibranchial teeth and a maxillary extension that did not extend beyond the posterior
portion of the eye, fish designated as suspected hybrids exhibited only one the two
attributes used to designate a steelhead.
136
30
0.200
0.175
0
0
0
0.150
0
0.125
15
0
0.100
00
0
0.075
0.050
0.025
1
5
8
11
14
17
20
24
28
34
26
33
38
41
44
47
50
53
Location
Appendix Figure A3. 1. Percent polymorpic loci (P095; dashed line) and expected
proportion of heterozygotes (HE; Weir 1996; solid line) of coastal cutthroat trout
populations sampled across their distributional range. Location numbers correspond
to numbers in Table 3.1.
137
APPENDIX TABLES
Appendix Table A2. 1. Meristic counts and collection information for all fish collected. Meristic characters are: A: anal
pterygiophores, B: dorsal pterygiophores, C: vertebrae, D: scales in lateral series, E: scales above lateral line, F: pelvic fin
rays, G: pectoral fin rays, H: gill rakers lower arch, I: gill rakers upper arch, J: branchiostegal rays, K: pyloric caeca, L:
basibranchial teeth, M: maxillary extension (see text for description of character counts). Sex: M = male, F = female;
locations are listed from north to south. Blanks represent missing data.
Fish
Location
number
931563
931564
931565
931566
931567
931568
931569
931570
931571
931572
931573
931574
931575
931576
931577
931578
931579
931580
931581
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
BoswellBay
Boswell Bay
BoswellBay
BoswellBay
BoswellBay
Boswell Bay
Boswell Bay
Boswell Bay
BoswellBay
BoswellBay
Boswell Bay
BoswellBay
Boswell Bay
BoswellBay
Boswell Bay
Date of
FL
Wt.
collection
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
9/13/93
(mm)
(g)
280
392
365
217
612
440
525
526
267
9113193
9/13/93
9/13/93
9/13/93
9/15/93
9/15/93
9/15/93
351
384
295
180
165
368
192
200
304
244
238
213
187
355
322
338
A
B
C
D
E
F
G
H
I
J
12
11
128
33
9
14
12
6
12
12
12
11
8
14
12
10
14
11
12
33
29
34
32
14
12
156
150
159
143
9
9
12
9
13
12
12
141
31
9
14
59
ii
12
134
14
11
12
154
26
32
9
43
505
84
80
308
62
63
63
63
58
63
63
62
9
12
12
65
183
35
11
12
63
139
10
12
61
12
11
118
151
11
11
12
12
62
63
62
94
54
10
11
61
11
11
64
12
12
61
126
150
154
153
146
125
156
30
27
33
35
12
12
62
161
12
11
61
142
462
387
334
13
L
M Sex
11
20
2
M
11
17
7
12
20
2
2
12
7
11
4
11
7
11
12
7
11
24
6
2
2
2
F
F
M
F
M
11
6
10
7
1
F
13
12
6
11
4
2
M
9
12
12
7
11
12
2
9
14
12
7
11
10
2
9
13
ii
7
Ii
6
2
9
13
12
7
11
6
2
F
M
M
F
M
K
9
14
12
7
11
2
31
9
14
14
7
11
11
34
29
33
34
29
9
14
11
7
11
14
9
14
12
7
12
17
2
2
2
2
9
14
12
7
11
16
2
9
13
11
6
12
6
2
9
14
12
7
11
4
2
M
F
F
F
M
F
-a
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
931582
942160
942161
942162
942163
942164
942165
942166
942167
942168
942169
942170
942171
942172
942173
942174
942175
942176
942177
942178
942179
942180
942181
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
Boswell Bay
9/15/93
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/6/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
196
76
743
489
213
362
12
11
61
126
12
63
12
13
63
154
146
12
12
12
12
181
12
11
62
62
62
159
149
124
11
11
61
131
60
242
11
10
158
11
12
64
62
185
12
13
61
154
112
155
224
11
12
62
10
12
61
154
143
25
36
34
34
34
37
34
35
32
35
33
34
9
12
12
12
61
136
33
462
734
340
926
677
1056
462
1010
670
320
12
13
61
163
12
13
65
11
12
12
12
12
12
11
12
ii
12
13
BoswellBay
Martin River
Martin River
Martin River
MartjnRiver
Martin River
Martin River
Martin River
Martin River
Martin River
425
383
288
338
269
224
186
299
270
223
249
289
363
435
337
433
415
454
362
468
410
320
G
H
I
J
K
L
M Sex
24
26
28
22
2
11
6
11
14
11
7
11
9
13
12
6
11
9
14
13
7
12
9
14
11
8
12
45
38
34
13
2
F
M
M
M
F
9
14
12
7
11
42
15
2
F
9
13
11
7
10
16
2
9
15
11
7
11
8
2
9
14
13
7
13
39
39
50
2
9
13
11
7
11
9
9
9
14
13
8
11
16
2
F
M
F
M
M
9
14
11
8
12
9
14
12
7
11
32
9
14
11
7
154
36
10
15
12
61
145
38
10
14
12
150
35
9
12
12
8
11
143
30
8
13
11
8
150
35
9
14
12
143
141
14
11
9
15
12
12
61
157
9
12
12
62
141
32
34
39
35
9
13
62
62
62
62
63
9
148
151
9
46
40
44
2
2
2
2
7
2
10
2
11
43
39
5
2
7
11
61
7
2
6
11
44
10
2
0
2
11
37
14
2
7
12
38
14
2
7
11
53
0
2
13
8
11
4
2
13
12
7
11
41
9
2
13
11
7
11
38
0
2
M
F
F
F
F
F
F
F
M
M
F
M
Appendix Table A2. 1. Continued
Fish
number
Location
942182
942183
942184
942185
942186
Martin River
Martin River
942 187
942188
942189
942190
942191
942192
942193
942194b
942l95'
953542
953543
931517
931518
931519
931520
931521
931522
931523
MartjnRjver
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Martin River
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
GinesCreek
Gines Creek
Date of
FL
Wt.
collection
(mm)
(g)
A
B
C
D
E
F
G
315
327
11
11
63
9
13
446
364
297
11
12
14
11
34
39
9
12
9
14
12
12
12
63
62
62
42
9
14
12
221
12
12
61
37
9
14
696
377
627
554
216
642
10
12
11
61
12
12
11
13
62
62
62
11
13
63
27
34
28
36
28
26
9
10
471
12
12
61
173
126
51
12
12
63
19
11
12
300
425
201
260
679
11
12
64
62
12
11
61
80
22
84
94
60
140
93
11
11
11
11
11
11
11
11
10
12
11
10
11
11
60
60
60
59
60
60
60
158
146
166
165
173
127
138
119
158
118
122
156
125
128
150
169
144
143
39
362
332
310
280
405
340
382
375
282
414
365
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/94
9/8/95
9/8/95
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
127
199
203
184
233
209
11
H
I
J
12
7
11
11
7
11
7
12
7
12
13
6
12
22
32
43
14
9
7
12
10
15
10
6
13
9
14
12
6
11
9
14
11
7
11
9
14
12
7
12
9
14
11
8
ii
37
9
14
12
7
11
27
28
10
15
10
8
11
14
12
15
12
7
6
11
41
9
9
36
10
14
13
35
37
9
14
10
14
141
35
9
14
11
7
11
143
35
9
14
11
6
11
149
34
40
9
14
12
8
11
9
14
12
7
10
33
9
14
11
7
11
156
144
K
L
M Sex
44
0
2
51
10
2
4
2
1
2
0
0
2
1
2
0
2
0
2
0
2
0
2
8
2
0
1
11
38
48
24
33
47
34
54
52
8
12
42
12
7
11
12
8
11
44
42
55
53
47
54
46
54
2
M
M
M
F
M
M
M
M
M
M
M
M
M
M
M
0
1
3
6
2
2
17
2
9
2
12
2
19
12
2
2
9
2
F
M
M
F
F
F
F
2
2
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
931524
931525
931526
931527
931528
931529
931530
931531
931532
931533
931534
931535
931536
931537
931538
931539
931540
931541
931542
931543
931544
931545
931546
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gjnes Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
198
87
58
42
26
69
42
11
11
11
11
11
11
11
11
11
11
11
11
GinesCreek
Gines Creek
Gines Creek
Gines Creek
GinesCreek
Gines Creek
Gines Creek
C
D
E
F
G
H
170
140
142
154
140
150
9
14
10
15
10
151
38
33
34
36
37
33
35
37
37
35
38
39
40
33
32
38
145
155
200
85
11
10
180
63
10
10
181
60
55
48
102
10
12
11
11
11
10
60
60
60
59
60
60
58
59
60
59
59
11
11
61
248
211
174
11
90
11
12
157
38
10
ii
166
46
47
45
136
116
112
12
12
11
10
10
11
11
11
11
11
11
11
59
62
60
59
59
60
60
58
60
87
11
13
61
158
61
10
11
60
155
183
163
136
186
166
177
9(10193
167
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
210
166
164
234
225
216
210
170
155
140
153
146
155
159
151
149
153
152
154
151
I
J
K
11
8
11
11
7
14
10
7
11
9
13
11
7
10
46
52
45
43
9
14
12
7
11
41
8
2
9
13
10
7
10
51
16
2
9
15
11
7
11
7
2
9
13
12
7
11
12
2
9
14
12
7
11
12
9
15
11
7
11
25
2
2
9
10
14
12
7
10
9
15
11
8
11
53
50
43
47
55
52
9
14
11
7
11
10
15
11
7
11
10
14
11
7
Ii
10
13
11
8
11
33
10
15
11
7
10
47
48
60
50
36
38
36
34
36
32
9
14
11
7
10
61
9
14
11
8
11
9
15
12
8
11
10
14
12
7
11
9
14
11
7
11
9
14
11
7
12
L
M Sex
9
2
F
10
2
7
2
9
2
2
M
M
M
F
F
F
M
F
M
M
3
2
F
10
2
13
2
14
2
10
2
10
2
17
2
F
F
M
F
F
M
54
6
55
17
44
47
44
14
2
2
2
5
2
13
2
F
F
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
931548
931549
931550
931551
931552
931553
931554
931555
931556
931557
931558
931559
931560
931561
931562
931963
931976b
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
Gines Creek
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
9/11f93
9/11/93
9/11/93
9/11/93
9/11/93
9/11/93
8/6/93
8/3/93
8/3/93
8/3/93
8/6/93
8/3/93
8/6/93
161
40
11
11
60
152
149
33
11
12
61
156
150
210
173
176
174
174
195
38
11
12
33
11
11
104
11
12
56
56
54
11
12
62
59
60
58
11
10
61
10
10
61
11
11
76
11
11
60
60
59
140
160
156
158
145
137
147
140
181
55
10
10
61
190
11
12
59
11
11
61
143
11
12
61
156
11
11
11
11
120
18
11
13
122
20
12
12
59
59
66
64
145
163
67
77
68
72
46
142
147
409
16
11
12
65
117
138
28
12
13
65
92
99
110
9
12
13
10
11
11
14
11
12
931977bb
931978b
931979b
931980b
931981b
931982h
GinesCreek
GinesCreek
GinesCreek
Gines Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
193
181
199
F
G
H
I
J
K
L
M Sex
36
9
13
11
7
11
2
9
13
12
8
10
8
2
38
9
14
12
7
10
34
38
32
34
35
35
33
34
36
9
13
12
8
10
9
13
12
7
11
9
14
11
7
11
9
14
12
7
11
9
13
11
7
11
44
43
44
48
46
50
53
39
6
35
9
14
12
7
11
9
13
12
8
10
9
13
11
7
11
9
14
11
8
11
37
37
9
14
ii
7
11
9
14
12
8
11
36
34
9
13
12
7
10
9
14
12
7
11
10
15
12
8
12
10
15
13
8
12
51
9
14
13
8
12
121
30
24
24
24
41
44
43
43
49
50
42
10
14
12
8
13
65
120
28
9
14
12
8
12
64
63
113
22
29
9
14
12
7
11
9
15
13
7
11
46
37
43
52
45
141
148
138
123
124
E
46
48
8
2
10
2
7
2
8
2
7
2
5
5
2
2
12
2
11
2
17
2
10
2
4
2
6
2
21
2
0
1
0
1
0
1
0
1
0
1
0
1
0
1
F
M
M
M
F
M
M
M
M
M
F
M
M
F
M
F
F
M
M
F
F
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
wt.
number
Location
collection
(mm)
A
B
931983"
931984"
931985"
931986"
93 1987'
931988"
931989"
931990"
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Suntaheen Creek
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
8/6/93
8/3/93
8/3/93
8/3/93
8/3/93
8/3/93
8/3/93
8/3/93
8/3/93
8/6/93
9/12/92
9/12/92
9/12/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
9/14/92
114
128
19
12
12
22
11
12
124
18
11
12
117
18
11
13
131
22
23
12
13
12
13
18
11
12
15
12
13
19
11
12
16
11
13
31
12
12
31
12
13
24
30
11
13
11
11
10
12
19
11
12
102
12
11
11
111
15
11
12
96
96
9
11
11
8
11
12
131
21
11
11
102
9
12
12
11
11
11
93 1991b
9319921)
920364"
920365"
920366"
920384
920386
920387
920388
920390
920391
920392
920393
920394
920395
126
117
109
118
112
135
138
135
147
144
122
106
C
64
64
64
64
64
64
64
65
65
66
63
64
65
63
62
63
62
62
62
63
63
62
63
D
E
F
G
HI
J
K
L
0
0
0
0
0
M Sex
120
26
10
14
13
7
133
25
10
14
13
8
11
131
27
28
10
15
13
7
12
45
43
50
9
14
13
8
12
41
28
27
28
26
29
25
28
27
27
32
34
36
33
37
36
33
32
33
34
10
15
13
9
12
47
9
15
12
8
11
9
15
12
8
12
46
54
9
14
12
7
12
9
10
15
13
8
12
14
13
8
12
9
13
12
7
11
9
14
12
6
9
14
13
9
14
12
51
8
2
13
11
6
6
11
10
11
3
2
9
13
11
6
11
5
2
9
14
12
6
11
32
50
47
4
2
9
14
11
6
11
2
2
9
13
11
6
10
5
2
9
14
11
6
11
7
1
6
2
3
2
4
2
125
124
131
130
121
132
127
126
127
126
137
143
149
137
146
148
151
143
140
145
12
49
50
47
11
9
14
11
6
11
9
14
12
5
11
52
40
49
46
45
9
14
11
6
11
51
0
0
0
0
0
0
0
0
1
M
M
M
1
F
1
1
1
1
1
1
1
1
1
1
1
M
M
F
M
F
M
M
M
M
M
M
F
F
F
F
F
M
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
920769
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
Freshwater Bay
FreshwaterBay
Freshwater Bay
FreshwaterBay
9/14/92
9/14/92
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
8/2/93
5/11/93
5/11/93
5/11/93
5/11/93
4/14/93
4/13/93
4/13/93
4/13/93
99
10
11
11
127
21
12
12
108
12
12
13
91
8
12
12
113
14
12
13
133
20
11
11
124
17
11
13
149
31
12
11
132
137
12
13
11
11
12
12
61
l2
13
12
13
13
12
130
22
23
23
26
24
20
22
63
66
65
63
66
64
65
63
66
60
11
11
64
66
67
62
107
12
12
13
65
119
16
12
12
154
31
10
10
120
130
17
12
13
11
11
11
12
66
63
66
62
62
10
ii
61
12
11
62
920770h
920902h
920903b
920904b
9209Q5
920906b
920907
920908b
920909
920910h
920911b
920912b
920913'
920914
931939b
931940b
931941
931942'
931136
931801
931802
931803
FreshwaterBay
KIag Bay
Kiag Bay
KlagBay
KlagBay
131
143
135
128
340
330
315
C
D
E
F
G
H
I
3
128
26
27
10
14
12
6
10
10
13
12
7
25
10
14
13
7
10
14
13
7
11
147
24
32
9
13
12
4
10
127
25
9
15
13
7
11
154
34
25
35
28
28
27
24
33
26
25
33
24
27
34
35
9
13
11
5
9
48
43
43
50
56
47
57
49
10
15
12
7
11
51
9
14
12
8
10
9
13
12
7
11
9
14
11
7
11
10
13
13
8
12
10
14
12
7
11
39
43
50
42
43
9
14
12
6
10
49
9
15
12
8
11
9
14
12
8
10
45
0
1
9
14
11
7
10
16
2
10
15
13
6
11
40
45
0
1
9
13
11
5
11
4
2
9
14
12
11
3
2
9
13
10
5
6
10
13
12
7
11
45
56
56
66
127
120
122
128
139
125
132
125
123
155
126
128
143
129
123
147
153
146
31
10
K
L
M
Sex
0
1
F
F
M
0
0
1
1
0
1
5
2
0
1
13
2
0
6
0
0
1
M
M
F
M
M
F
2
F
1
M
1
F
0
1
0
1
8
2
0
1
M
F
M
M
M
M
M
F
M
F
M
3
2
14
2
Appendix Table A21. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
931804
931805
931806
931807
931808
931809
931810
931811
931812
931813
931814
931816
931817
931818
931113b
Kiag Bay
Kiag Bay
4/13/93
4/13/93
4/13/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
4/14/93
5/27/93
5/27/93
5/27/93
5/27/93
5/27/93
5/27/93
5/27/93
5/27/93
5/27/93
291
271
288
116
91
931114
931115
93l116
93111D
931118
931119
931120
931121
KlagBay
KlagBay
KlagBay
KlagBay
Kiag Bay
Klag Bay
KlagBay
Kiag Bay
KIag Bay
Kiag Bay
Klag Bay
Kiag Bay
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
12
11
62
153
13
12
6
10
2
61
9
14
12
6
10
12
2
F
F
11
11
61
158
154
31
9
13
12
7
10
40
59
48
10
11
32
34
9
11
6
2
M
10
10
61
151
9
14
11
7
Il
51
5
2
12
11
61
131
12
6
11
6
2
12
61
13
10
6
10
6
2
12
10
9
13
12
7
11
12
2
13
12
62
62
31
9
13
11
7
11
7
2
11
12
61
31
9
13
12
7
10
9
2
111
11
11
62
142
147
150
136
139
29
9
13
11
7
11
213
11
11
61
35
9
13
11
7
10
143
12
11
30
9
13
10
5
9
315
295
12
12
140
31
9
13
12
7
11
12
11
62
60
62
150
128
151
31
9
13
12
8
11
43
42
66
57
53
63
49
49
50
52
F
F
11
9
9
13
243
335
256
267
30
30
34
33
12
65
130
26
9
14
12
7
10
33
65
61
187
41
9
14
11
7
11
48
10
111
9
13
12
7
12
9
14
12
6
11
9
13
12
48
36
42
76
62
5
3
11
12
118
19
11
12
59
68
102
116
112
2
11
12
4
11
13
65
64
11
12
11
61
149
16
ii
11
63
163
28
37
38
8
11
15
12
12
181
42
8
13
11
6
10
103
11
12
12
60
60
157
37
9
14
13
6
II
50
46
M
M
M
F
5
2
F
12
2
6
2
5
2
7
2
0
1
M
M
F
M
F
2
M
0
1
17
2
F
F
M
M
M
7
2
11
2
2
1
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
931904
931905
931906
931907
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
Portage Creek
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
6/9/93
7/8/94
7/8/94
7/8/94
7/8/94
7/8/94
7/8/94
7/8/94
7/8/94
97
8
11
11
60
13
12
7
11
16
12
11
61
9
15
12
7
11
114
15
10
11
9
13
13
7
10
114
15
12
12
35
34
39
39
29
39
36
9
117
136
140
179
152
118
177
164
9
14
11
7
11
43
40
39
44
9
14
12
7
9
13
12
6
11
9
13
12
7
145
35
9
11
112
106
24
24
23
29
28
9
13
11
9
14
9
931908b
931909
931910
931911
931912h
931913h
931914b
931915b
931916"
931917
931918
942049
942050
942051
942052
942053
942054
942055
942056
65
4
11
13
101
11
11
12
118
15
12
12
76
69
59
4
11
11
60
62
63
62
62
62
4
12
12
65
3
11
12
65
61
3
11
12
61
68
92
4
12
12
61
9
13
11
110
13
12
11
62
63
123
18
10
12
61
208
269
250
212
164
73
177
149
89
45
12
11
11
12
62
62
11
11
61
11
12
139
11
10
107
12
12
13
119
17
11
12
62
60
62
62
94
8
11
11
63
147
116
119
123
147
154
139
146
169
141
151
163
L
M Sex
7
2
13
2
9
2
9
2
0
1
4
2
11
44
50
9
2
6
11
34
5
2
7
12
35
12
6
11
0
0
13
11
7
11
37
0
1
9
14
12
7
11
1
14
12
8
11
33
9
14
12
6
12
37
42
56
0
9
38
9
14
12
7
11
33
9
14
10
6
11
38
9
13
13
7
11
33
9
14
12
6
11
33
32
35
33
33
10
15
12
8
11
9
14
11
7
11
9
13
13
7
11
9
13
12
7
11
9
14
11
7
11
1
1
M
M
F
M
M
M
M
M
M
M
0
1
6
2
41
5
2
41
21
2
32
56
50
46
46
48
39
5
2
5
2
5
2
M
M
F
F
M
F
M
M
M
6
10
2
F
2
F
9
2
4
2
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
WI
number
Location
collection
(mm)
(g)
A
B
C
D
931016
931017
931018
931019
931020
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan SaitChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
5/24/93
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
242
338
336
122
333
316
285
156
11
12
60
157
10
12
61
151
11
11
148
12
11
60
62
III
11
61
Il
ii
12
60
12
61
11
11
62
11
11
61
12
12
12
11
12
12
11
12
63
61
62
62
12
11
61
12
12
10
12
63
62
198
12
12
61
506
302
11
11
61
13
12
63
581
484
342
306
12
11
12
ii
12
11
931021
931022
931023
931041
931042
931043
931044
931045
931046
931047
931048
931049
942026
942027
942028k
942029
942030
942031
DuncanSaitChuck
DuncanSaitChuck
Duncan SaitChuck
Duncan Salt Chuck
DuncanSaltChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
291
268
259
285
371
294
167
193
295
314
360
339
279
274
351
312
390
351
325
304
185
224
435
268
45
76
333
345
472
438
205
129
134
160
138
145
135
129
157
12
61
61
152
152
154
164
140
163
154
160
140
143
140
12
62
150
F
G
H
I
J
K
L
M Sex
33 9
32 9
33 9
32 9
27
9
38 9
32 10
34 8
34 9
27 9
38 9
36 10
36 9
32 9
9
38
34 9
38
9
32 9
33 9
37 9
32 9
39 10
40 9
14
12
S
11
41
10
2
14
11
6
12
2
13
7
11
39
48
23
14
16
2
14
12
7
11
35
29
2
13
10
6
11
16
2
13
12
7
11
9
20
2
2
E
14
12
7
11
28
44
39
14
11
8
11
42
18
13
12
7
11
39
10
2
13
12
8
11
40
2
2
13
12
7
10
37
19
13
11
7
11
31
15
2
2
14
11
7
12
18
15
12
7
12
46
36
14
13
8
12
14
12
8
12
38
13
12
6
11
51
26
23
25
26
13
12
6
11
13
2
13
12
7
10
41
44
13
2
13
11
8
10
44
13
12
12
12
7
10
41
26
6
2
2
7
10
3
2
7
11
41
48
15
2
13
14
2
2
2
2
2
2
M
F
F
F
F
F
F
M
F
M
F
M
M
M
M
F
F
M
M
M
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
942032
942033
942034
942035
942036
942037
942038
942039
942040
942041
942042
942043
942044
942045
953388
953389
953390
953391
953392
953393
953394
953395
953396
Duncan Sail Chuck
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
7/6/94
8/9/95
8/9/95
335
355
352
332
279
315
251
148
360
375
209
289
333
407
10
12
130
13
12
154
32
33
9
471
392
374
9
11
12
62
62
62
161
35
12
12
61
140
155
169
163
164
154
147
144
152
152
162
165
163
159
159
160
147
153
30
36
34
36
DuncanSaltChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
DuncanSaitChuck
Duncan SaitChuck
Duncan SaitChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan SaitChuck
Duncan Salt Chuck
Duncan Salt Chuck
DuncanSaltChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan SaitChuck
DuncanSaltChuck
8/9195
8/9/95
8/9/95
8/9/95
8/9/95
8/9/95
8/9/95
217
346
11
11
11
11
62
62
138
12
12
61
34
525
11
ii
61
11
12
492
78
12
12
63
62
11
12
61
251
10
12
61
393
294
12
12
61
11
12
60
221
231
111
11
13
61
123
12
12
62
250
234
162
127
116
228
172
392
687
11
11
61
11
11
62
11
11
61
10
11
60
12
11
63
11
13
11
12
62
62
310
221
273
259
347
400
166
141
H
I
J
K
13
12
7
12
14
12
6
11
9
14
11
6
11
9
13
12
7
12
9
13
11
7
10
8
13
11
7
12
10
14
12
7
12
42
9
13
11
8
11
37
32
33
32
37
34
40
35
32
37
37
35
37
37
33
9
13
12
6
11
9
13
12
7
10
9
13
12
7
11
9
13
10
7
9
9
13
12
7
13
12
9
14
9
L
M Sex
41
9
2
M
35
18
2
F
51
6
2
F
17
2
F
10
2
M
15
2
2
F
F
F
11
44
42
40
46
40
29
33
43
50
11
38
7
12
11
7
12
14
11
6
12
9
13
11
7
11
10
14
13
7
12
9
13
12
9
11
9
13
12
6
11
9
13
12
6
11
9
13
11
7
11
9
14
12
7
11
25
21
2
2
2
2
13
2
12
2
43 14
42 10
44 18
34 20
40 8
40 8
48 14
50 10
30 28
2
8
10
5
2
2
2
2
2
2
2
2
2
2
M
M
M
M
F
M
M
F
F
F
M
M
M
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
953397
953398
953399
Duncan Salt Chuck
8/10/95
8/10/95
8/10/95
8/10/95
8/10/95
8/10/95
5/25/93
5/25/93
172
199
11
11
61
162
36
9
13
12
325
354
259
233
360
284
355
391
12
13
61
148
35
9
13
11
12
12
62
175
13
12
8
12
11
61
153
36
33
9
12
9
13
12
7
10
14
13
9
13
5/25193
320
9
13
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
251
953400
953501
953502
931050
931051
931052
931053
931054
931055
931056
931057
931058
931059
931060
931061
931062
931063
931064
931065
931066
DuncanSaltChuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Duncan Salt Chuck
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
KadakeCreek
KadakeCreek
KadakeCreek
Kadake Creek
KadakeCreek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
G
H
I
J
K
L
M Sex
7
10
2
11
44
40
4
6
8
2
42
44
7
2
11
16
2
7
10
44
17
2
12
7
10
38
18
2
12
6
10
37
24
16
2
2
14
2
482
174
123
F
F
495
211
12
11
62
167
459
12
11
61
325
149
102
322
328
303
251
316
315
327
223
11
11
11
12
169
155
154
12
7
11
12
155
31
9
9
13
11
14
11
6
11
12
12
62
62
62
63
36
39
32
38
159
40
10
14
12
7
11
12
11
61
135
35
14
11
7
10
11
11
64
144
31
9
9
43
43
39
42
13
11
6
10
11
11
61
151
34
9
13
12
6
11
12
12
14
11
6
10
9
14
12
7
11
12
12
63
9
14
11
6
11
12
12
61
9
13
12
7
10
450
236
10
11
9
14
11
8
10
11
12
62
62
139
157
139
9
13
12
7
11
231
129
11
11
61
168
9
13
11
6
11
45
20
40
47
282
123
12
12
62
147
37
38
34
34
30
38
37
35
9
12
64
62
155
11
9
13
12
6
10
41
224
320
317
336
297
324
319
363
290
365
288
M
F
167
151
10
2
M
F
M
F
M
M
5
2
F
31
8
2
42
37
43
10
2
10
2
10
14
2
2
13
2
F
M
M
F
M
F
17
2
18
2
9
2
20
2
F
F
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
931067
931068
931069
931070
931071
931072
942068
942069
942070
942071
942072
942073
942074
942075
942076
942077
942078
942079
942080
942081
942082
931859
931860
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
Kadake Creek
KadakeCreek
Kadake Creek
Kadake Creek
Eagle River
Eagle River
5/25/93
5/25/93
5/25/93
5/25/93
5/25/93
5/26/93
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
7/9/94
5/22/93
5/22/93
287
202
297
280
265
202
75
246
209
11
12
61
31
11
7
12
11
13
11
6
11
12
13
11
6
11
11
159
140
9
12
62
63
62
9
9
14
12
138
144
37
9
12
13
8
11
184
56
317
76
138
57
253
330
127
88
83
75
36
65
35
95
11
11
61
141
32
9
13
12
7
11
11
12
62
138
35
9
13
12
7
10
39
45
36
46
13
12
62
37
9
13
13
8
11
51
11
11
61
13
12
7
11
12
9
14
12
8
11
11
11
63
61
9
14
11
6
11
10
11
9
14
12
7
11
12
11
9
12
12
7
10
10
12
9
14
13
7
12
11
11
12
7
10
11
9
9
13
12
13
12
8
11
11
12
9
13
12
11
11
11
12
11
13
13
11
12
62
62
62
63
63
63
62
62
65
62
11
12
61
37
36
37
36
36
36
36
36
30
39
39
37
38
34
9
12
12
12
11
11
62
63
152
152
162
144
150
146
166
154
156
147
144
152
148
152
153
143
150
182
335
202
258
186
313
349
245
222
213
199
157
190
152
204
305
177
145
236
49
32
32
37
K
L
M Sex
37
9
23
2
F
2
17
2
21
6
2
2
16
2
9
2
41
13
2
14
2
M
F
F
F
M
F
M
F
16
2
4
2
F
16
2
M
17
2
5
16
2
2
F
M
21
2
42
9
13
13
7
7
12
48
39
48
46
46
45
53
38
43
3
2
9
14
12
8
11
41
24
2
9
13
11
6
11
2
13
13
8
11
17
2
9
13
11
7
11
30
2
35
9
14
12
5
12
33
9
14
11
7
11
50
50
47
54
39
12
9
9
2
13
2
F
M
M
M
F
F
M
F
F
C
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
931861
931862
931863
931864
931865
931866
931867
931868
931869
931870
931871
931872
931873
931874
931875h
Eagle River
EagleRiver
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
EagleRiver
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
EagleRiver
Eagle River
Eagle River
Eagle River
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
5/22/93
136
109
26
10
12
61
35
9
13
12
7
11
13
12
11
61
150
134
33
9
13
12
7
11
200
73
50
58
59
44
73
43
10
10
9
13
11
6
11
11
14
12
7
11
12
9
14
11
6
11
11
11
36
29
30
9
11
9
13
12
6
10
11
31
9
13
12
7
12
12
61
34
9
14
11
8
12
10
12
61
14
12
7
11
12
12
61
9
13
11
7
11
12
12
61
9
13
10
4
11
11
12
63
141
9
14
12
7
11
12
11
63
148
9
14
12
7
12
10
12
61
145
9
14
12
6
11
11
12
10
14
12
7
10
11
11
9
13
12
7
12
11
139
138
31
Il
31
9
13
11
6
11
12
12
134
13
12
7
Ii
11
9
14
11
195
69
11
12
61
140
143
32
30
9
12
64
62
61
62
60
131
239
246
77
116
77
30
40
72
72
128
144
30
35
35
30
32
32
26
9
134
154
150
142
146
147
143
145
140
154
31
11
60
62
62
62
59
43
59
38
44
9
14
12
246
208
153
11
12
61
139
33
32
9
14
85
10
11
62
163
33
9
179
49
12
11
61
151
34
9
931876
931877
931878
931879
931880
931881
931882
931883
5/22193
172
186
181
164
204
172
242
200
226
195
148
167
194
194
L
M
Sex
8
2
11
1
F
M
12
2
F
6
2
M
40
11
11
51
15
2
2
11
13
2
9
2
15
2
7
11
48
42
47
38
42
51
49
45
45
50
33
45
47
F
F
F
F
F
7
11
51
12
7
11
41
13
11
7
11
13
13
5
11
59
42
48
2
10
2
M
M
1
2
F
10
2
10
2
0
1
12
2
M
F
F
M
11
2
F
26
13
2
2
16
2
17
2
F
F
F
F
8
2
14
2
F
F
UI
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M
Sex
931884
931885
931886
931887
942046
942047
942048
931351
931352
931353
931354
931355
931356
931357
931358
Eagle River
5/22/93
5/22/93
5/22/93
5/22/93
7/7/94
7/7/94
7/7/94
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
7/28/93
175
48
154
438
11
11
61
13
8
5
11
47
12
2
12
62
9
13
13
7
11
38
21
2
11
12
61
142
32
30
34
9
12
140
146
9
13
12
7
11
18
2
139
31
9
14
12
7
11
155
33
13
13
7
11
141
9
9
13
11
7
11
9
12
12
8
12
93135,9
931360
93l36l'
931362
9313631
931364
931365
931366
EagleRiver
Eagle River
Eagle River
Eagle River
Eagle River
Eagle River
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
WolverineCreek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
266
371
6
2
F
F
F
M
11
2
M
10
2
13
2
17
2
3
2
M
F
F
F
33
19
2
M
16
2
16
2
F
F
12
2
M
18
2
F
17
2
8
2
F
F
M
M
192
71
11
11
229
104
13
12
199
180
304
69
55
234
195
415
267
344
56
312
412
192
12
13
62
62
62
11
11
61
135
12
11
63
135
36
34
30
9
13
11
6
11
11
12
61
141
34
10
13
11
6
11
36
32
59
63
43
42
44
11
11
60
148
34
9
13
12
5
11
11
10
61
139
35
9
13
10
5
11
Ii
12
60
148
34
9
13
11
6
11
11
11
61
147
35
9
13
11
7
11
11
11
60
144
9
14
10
6
12
11
11
61
161
11
12
61
121
11
11
61
149
177
110
12
13
64
123
294
346
257
11
12
61
12
11
159
145
10
11
60
149
119
11
11
61
140
39
12
12
61
163
36
40
34
38
26
39
38
34
33
36
279
366
307
330
184
323
355
282
241
216
325
340
301
236
163
9
14
11
8
11
9
14
12
8
11
40
45
46
50
48
48
9
13
11
4
11
38
14
2
10
15
12
7
11
0
1
9
13
9
5
11
12
2
9
14
12
6
12
9
14
10
5
12
9
12
12
6
12
9
13
11
6
11
37
47
48
41
47
45
7
2
21
2
15
2
11
2
F
F
F
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
931367
931368
931369
931370
931843
931844
931845
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
7/28/93
7/28/93
7/28/93
7/28/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
4/27/93
7/28/93
7/15/94
7/15/94
7/15/94
309
275
236
284
314
357
376
400
230
12
11
61
147
188
11
11
61
148
105
ii
11
61
158
208
267
11
12
63
147
11
11
61
138
34
35
33
33
34
436
11
12
61
533
11
11
672
745
373
383
205
462
339
364
277
12
11
61
10
11
ii
12
62
58
159
150
145
153
141
11
12
61
153
11
12
61
143
11
12
143
12
11
11
12
59
61
62
11
12
325
641
11
430
931846
931847
931848
931850
931851
931852
931853
931854
931855
931856
931857
931858
931960
942132
942133
942134
421
225
352
382
344
324
345
305
328
392
342
F
G
H
I
J
K
L
M
Sex
2
F
F
M
F
M
M
M
F
F
F
F
M
M
F
F
M
M
M
9
13
10
5
10
9
13
12
7
11
9
13
11
5
11
9
14
12
7
12
13
11
6
12
36
9
10
13
11
6
11
33
9
14
12
7
12
35
37
34
9
13
13
8
10
9
13
11
4
11
10
14
9
6
11
9
12
12
6
12
9
13
12
6
11
9
12
11
6
11
160
156
36
32
32
35
32
9
13
12
7
11
9
13
11
7
11
43 32
46 7
46 8
47 23
41 10
36 19
47
8
33 15
39
3
36 6
48 20
53
8
42 12
28 3
56 16
61
151
35
9
13
11
6
11
17
2
11
62
143
35
9
13
11
7
11
24
2
12
11
61
161
39
9
13
12
7
11
25
2
12
12
155
35
9
13
11
7
11
11
2
10
13
62
60
141
33
9
13
11
6
11
41
14
2
M
44
47
52
7
2
M
13
2
7
2
F
F
238
117
11
11
61
150
34
9
14
12
7
11
161
41
11
12
61
131
30
9
12
12
8
11
122
18
11
12
62
132
33
10
13
12
7
11
42
2
2
2
2
2
2
2
2
2
2
2
2
2
2
Appendix Table A2. 1. Continued
Fish
number
942135
942136
942137
942138
942139
942140
942141
942142
942143
942144
942145
942146
931226
931227
931228
931229
931230
931231
931232
931233
931234
931235
931301
Date of
FL
Wt.
Location
collection
(mm)
(g)
A
B
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
Wolverine Creek
WolverineCreek
Wolverine Creek
Wolverine Creek
Wolverine Creek
WolverineCreek
Staney Creek
Staney Creek
Staney Creek
StaneyCreek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
7/15/94
7/15/94
7/15/94
7/15/94
7/15/94
7/15/94
251
157
11
12
239
216
160
10
12
109
11
172
152
50
34
275
183
246
244
240
255
169
210
233
46
80
7115/94
7/15/94
7/15/94
7/15/94
7/15/94
7/15/94
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
C
D
E
F
G
H
I
J
K
149
31
9
13
12
7
11
62
144
9
14
10
7
11
12
61
147
9
13
12
5
11
12
11
62
140
30
32
30
9
13
12
7
12
12
12
61
140
33
10
13
12
6
12
12
61
12
10
6
11
11
12
61
37
38
10
145
9
14
12
5
12
142
140
142
11
12
61
31
9
13
11
6
10
11
11
61
7
11
61
9
9
11
12
13
12
6
10
11
12
9
13
10
7
10
12
12
10
14
11
6
11
106
12
12
155
31
9
13
11
7
10
184
60
11
11
60
62
62
60
38
38
30
33
13
11
146
138
144
155
150
135
143
140
27
9
14
12
6
10
46 10
44 15
52 17
42 15
46 10
62 25
49 20
211
87
12
11
62
140
23
9
14
12
7
12
45
33
2
174
51
11
11
136
27
9
14
12
7
12
41
16
2
143
26
60
69
42
50
90
140
12
11
9
13
11
7
11
45
13
2
12
149
129
33
11
62
62
62
27
9
13
11
6
12
38
ii
20
2
11
61
31
9
13
11
7
11
38
19
2
12
12
62
139
142
33
8
13
11
7
11
10
10
61
141
9
14
12
7
10
10
11
61
9
13
12
7
10
11
ii
62
144
144
30
27
31
9
14
12
7
11
182
156
157
175
218
249
L
M Sex
51
9
2
F
43
49
52
46
45
44
12
2
M
M
54
46
50
5
2
13
2
15
2
4
2
10
2
2
2
2
2
2
2
2
16
2
20
29
27
2
2
2
F
F
F
M
F
F
M
F
F
F
M
M
F
F
M
F
F
M
UI
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
931302
931303
931304
931305
931306
931307
931308
931309
931310
931311
931312
931313
931314
931315
931316
931317
931318
931319
931320
931321
931322
931323
931324
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
StaneyCreek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
7/27/93
263
288
192
184
11
11
11
6
11
12
9
14
12
7
10
12
12
61
185
55
11
11
63
224
105
12
11
197
67
100
93
12
11
63
64
11
11
60
11
10
61
105
10
12
13
12
207
246
45
80
12
ii
60
63
60
30
29
27
27
29
28
28
30
26
14
ii
150
148
130
147
158
156
134
143
134
143
144
9
216
68
62
62
160
11
11
61
158
35
11
10
168
47
92
55
49
37
48
300
75
50
12
227
217
234
171
217
175
176
154
173
364
232
172
162
40
L
M Sex
51
14
2
35
2
30
2
35
2
2
F
M
M
M
M
M
F
F
F
M
M
F
2
M
F
F
M
M
M
M
9
14
11
6
10
9
10
14
12
6
11
13
10
7
11
42
40
50
47
16
2
9
14
11
8
11
41
21
9
14
12
7
12
10
14
11
7
11
43
47
29
28
2
2
9
14
9
6
11
11
2
31
9
14
12
7
11
12
31
9
14
12
6
ii
2
2
141
29
9
13
12
7
11
62
135
27
9
14
12
7
ii
11
61
141
32
9
13
12
7
11
12
11
61
153
37
9
14
12
6
12
2
11
11
61
143
31
9
13
11
6
ii
11
11
61
158
9
14
11
7
11
18
2
12
11
60
9
14
10
6
12
41
27
2
11
11
61
130
147
30
27
40
45
43
49
42
45
44
44
43
31
10
14
13
6
10
41
19
2
11
10
62
149
35
9
14
12
7
11
3
2
12
11
61
14
12
7
11
23
2
10
9
14
12
7
11
15
2
11
11
61
61
27
30
9
11
128
133
140
31
9
14
11
7
11
50
44
40
43
19
2
27
20
20
17
2
15
2
25
2
F
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M
Sex
931325
942098
942099
942100
942101
942102
942103
942104
942105
942106
942107
942108
942109
942110
942111
942112
942113
920088
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Staney Creek
Vixen Inlet
Vixen Inlet
Vixen Inlet
7/27/93
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/17/92
7/17/92
7/17/92
7/17/92
7/17/92
7/17/92
199
73
122
11
11
62
137
31
9
14
ii
7
11
12
61
152
9
14
12
8
12
20
40
F
M
101
12
12
61
145
9
13
13
8
11
2
F
10
12
14
10
7
11
F
11
31
9
13
12
7
11
2
F
11
61
30
9
14
11
7
11
26
22
27
2
11
140
134
134
142
145
8
11
58
62
2
31
9
13
12
6
11
8
2
13
11
7
10
14
10
11
14
11
6
7
22
24
2
9
9
M
M
F
F
285
349
32
30
30
30
30
9
198
42
42
45
54
43
42
46
46
41
106
91
41
100
86
65
30
33
32
44
40
2
12
9
9
14
11
7
61
14
10
6
12
9
92OO89
920090k
920094
920095
920096
Vixenlnlet
Vixenlnlet
Vixen Inlet
240
240
219
220
164
119
12
11
61
11
12
11
12
217
453
11
ii
60
62
62
12
12
62
137
231
111
12
12
63
135
150
32
63
24
79
27
449
92
11
11
61
31
9
14
11
7
11
11
12
61
128
121
31
9
13
12
7
13
12
61
132
9
13
11
11
12
61
9
13
11
12
11
9
14
12
8
37
9
14
12
7
12
12
62
62
60
146
144
162
32
29
28
253
300
288
121
11
Ii
61
266
11
11
11
11
190
65
12
12
60
62
62
12
186
210
198
132
198
140
355
228
11
131
143
157
36
10
14
11
12
2
2
F
F
F
13
2
19
2
44
9
2
51
37
2
11
51
39
2
7
12
51
16
2
7
1!
1
F
12
55
24
36
16
2
11
40
49
2
56
48
45
12
2
M
M
F
F
F
F
7
67
52
M
M
M
F
L1
Appendix Table A2. 1. Continued
Fish
number
920097
920098a
920099
920100
Date of
FL
Wt.
Location
collection
(mm)
(g)
A
B
C
Vixenlnlet
7/17/92
7/17/92
7/17/92
7/17/92
7/17/92
7/17/92
7/20/92
7/20/92
7/20/92
7/20/92
145
11
12
191
25
74
11
11
62
62
233
130
11
11
61
143
26
48
35
377
42
10
11
61
11
11
59
11
11
61
10
11
62
143
35
9
14
11
6
11
10
11
61
144
33
9
13
11
6
11
150
141
32
37
33
33
9
14
12
5
12
9
14
12
7
11
9
13
12
7
10
Vixen Inlet
Vixen Inlet
Vixen Inlet
9201011
Vixenlnlet
920l02
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
Vixen Inlet
920257
920258
920259
920260
920261
920262
920263
920309
931371
931372
931373
931374
931375
931376
931377
931378
931379
Vixenlnlet
Vixenlnlet
Vixen Inlet
Vixenlnlet
Vixenlnlet
Vixen Inlet
Vixen Inlet
166
151
328
168
D
E
F
G
H
I
J
K
L
M Sex
46
48
44
54
48
42
128
116
17
12
12
15
12
11
62
62
7120/92
162
36
10
10
61
141
7/20/92
7/20/92
7/20/92
7/29/93
7/29/93
7/29/93
7/29/93
7/29/93
7/29/93
7/29/93
7/29/93
7/29/93
276
181
11
11
61
118
13
11
11
100
9
11
11
61
310
330
317
265
284
245
254
246
234
299
266
265
204
223
11
12
60
146
133
154
164
11
11
61
11
11
11
11
147
M
F
M
F
F
6
2
F
F
7
2
F
28
2
55
19
2
37
14
2
46
9
13
11
5
12
50
20
31
9
14
11
11
51
7
2
2
31
9
13
10
6
6
11
43
35
9
13
11
5
11
7
4
2
153
35
9
13
12
5
11
50
16
2
61
132
9
13
12
6
10
51
8
2
11
61
148
33
36
9
13
13
6
12
35
7
2
II
61
153
35
9
13
11
6
12
11
12
61
139
33
9
13
11
6
11
143
130
12
12
62
13
12
6
11
12
61
13
11
7
117
11
12
62
150
34
34
32
9
11
149
169
14
11
6
10
10
39
45
52
50
44
8
1
F
M
F
M
M
F
7
2
17
2
15
2
M
M
F
F
F
M
4
2
F
12
2
F
1l
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
931380
931381
931382
931383b
Vixen Inlet
7/29/93
7/29/93
7/29/93
7/29/93
4/26/93
7/16/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
7/20/92
4/22/93
7/16/94
7/16/94
7/16/94
7/16/94
224
113
12
11
231
111
ii
209
87
185
134
64
23
240
127
171
931842'
920071
920266
920267
920268
920269
920270
920271
920272
920273
920274
920275
920276
920315
931836b
942147
942148
942149
942150
Vixenlnlet
Vixenlnlet
Vixenlnlet
Vixen Inlet
MargaretCreek
MargaretCreek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
MargaretCreek
Margaret Creek
Margaret Creek
MargaretCreek
Margaret Creek
MargaretCreek
Margaret Creek
Margaret Creek
MargaretCreek
C
D
E
F
G
H
I
J
K
L
M Sex
62
62
158
145
9
9
13
6
11
51
11
2
12
12
6
11
12
61
137
9
13
12
7
11
12
12
130
10
14
13
7
11
11
12
62
62
11
11
61
152
32
9
14
10
6
12
46
20
11
11
61
159
35
9
13
11
7
12
12
12
61
29
9
14
12
11
11
31
9
13
11
10
12
12
60
63
9
13
11
15
11
11
61
9
13
11
7
11
19
11
12
61
32
30
30
7
6
6
11
19
9
13
12
7
11
27
89
45
11
12
31
9
13
11
6
10
11
12
62
62
152
142
154
133
145
148
163
9
13
11
8
12
Ii
12
63
161
35
38
9
14
10
6
12
18
11
12
131
13
11
7
11
11
11
9
14
11
11
12
9
13
11
6
7
11
86
3
12
13
121
32
30
33
29
9
11
11
6
10
22
11
12
143
31
108
12
10
12
138
142
23
12
12
63
63
61
64
62
64
62
40 1
40 8
44 0
45
42 15
54 13
37 24
47 12
39 14
45 13
53 24
44 14
52 15
44 18
43 12
48 20
54 23
2
12
30
33
35
28
14
11
106
ii
11
13
61
34
34
33
134
122
108
112
123
140
220
170
122
99
215
67
137
B
134
155
144
148
9
13
9
13
9
14
12
9
14
10
9
13
12
11
11
11
2
1
M
M
F
M
M
2
2
2
2
2
2
2
M
F
F
F
M
M
F
2
2
F
2
F
2
M
F
M
M
2
2
0
1
11
52
15
1
6
10
53
9
2
5
11
5
2
7
11
39
53
13
2
F
F
F
F
F
U'
00
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
942151
942152
942153
Margaret Creek
MargaretCreek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Margaret Creek
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
7/16/94
7/16/94
7/16/94
7/16/94
7/16/94
7/16/94
7/16/94
7/16/94
7/16/94
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
7/14/92
154
ii
11
61
148
35
29
12
11
61
261
147
10
12
178
41
11
12
127
19
12
12
138
25
11
12
122
17
11
12
111
13
11
11
231
333
334
11
13
13
12
11
11
61
11
11
61
11
11
60
166
148
11
11
63
152
11
11
61
145
11
11
61
130
11
12
62
147
312
96
332
348
244
57
47
59
29
30
293
62
62
62
62
60
63
62
12
11
61
125
18
11
11
143
29
11
10
31
12
12
7/14192
146
117
52
10
12
7/16/92
255
12
11
63
64
63
62
62
170
138
942154
942155
942156
942157
942158
942159
920001
920002
920003
920004
920005
920006
920007
920008
920009
920010
920011
920012
920013
920066
229
180
117
184
144
149
D
E
F
G
H
147
144
32
30
9
14
9
13
162
9
13
131
30
29
9
154
35
151
32
36
32
28
39
39
39
30
36
30
28
33
34
I
I
13
7
11
11
5
11
12
6
11
13
11
6
11
9
14
12
7
11
9
13
11
5
11
9
13
13
7
10
9
13
10
7
11
8
13
11
6
10
9
14
12
6
11
9
13
12
6
12
52 5
48 15
44 17
53 11
8
56
57 4
62 9
43 6
53 20
67 16
51 30
9
13
12
7
11
55
9
13
12
8
11
9
13
11
6
12
9
14
11
6
11
9
13
11
6
11
9
14
12
7
9
14
11
7
31
9
13
12
136
28
9
13
135
9
145
30
33
137
31
135
139
144
167
158
K
L
M Sex
2
2
2
2
2
2
2
M
M
F
M
F
M
M
2
2
F
2
2
M
M
17
2
M
23
2
25
2
F
M
24
2
10
2
10
50
50
44
46
54
23
2
13
51
17
2
6
10
11
6
11
14
10
6
12
9
14
11
6
11
48
42
46
42
9
14
13
7
12
52
F
M
M
9
2
F
M
F
11
2
M
19
2
32
25
2
F
F
2
M
Lu
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
920068
920069
920277
920278
920279
920280
920281
920282
920283
920284
920286
920287
920288
920289
920290
920301
920302
942083
942084
942085
942086
942087
942088
ThorneRiver
ThorneRiver
7/16/92
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorrie River
Thorne River
Thorne River
A
B
C
D
E
F
G
H
I
J
K
L
184
12
ii
14
12
6
12
2
11
9
14
11
5
11
19
2
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/22/92
7/16/92
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
283
345
344
185
11
12
8
13
11
6
10
425
345
46
11
11
33
9
13
ii
6
12
10
11
61
39
9
13
11
7
12
12
11
62
62
60
31
9
14
12
7
10
58
40
54
44
52
50
24
III
35
32
33
9
180
63
63
62
60
158
7116192
35
9
13
12
7
12
32
9
14
12
7
11
31
9
13
11
6
11
9
13
12
7
11
9
13
12
6
11
9
13
12
6
11
9
14
12
8
10
188
229
307
113
11
12
171
11
11
198
73
357
325
341
120
66
67
24
10
11
11
336
333
334
226
195
205
138
231
219
229
139
342
183
167
89
106
32
393
52
42
153
169
152
157
150
159
11
63
63
169
142
134
11
12
62
158
11
12
61
173
12
10
152
30
32
35
34
11
11
147
35
9
13
11
6
11
11
11
148
31
9
13
12
7
11
11
11
141
31
9
13
12
7
12
12
12
145
33
9
13
12
7
11
150
31
9
13
10
7
12
151
34
9
13
11
7
11
150
35
9
13
12
6
11
136
143
32
10
13
12
7
33
9
13
13
7
149
31
9
13
11
7
11
11
11
12
10
Ii
12
12
11
12
62
60
60
62
63
62
63
62
63
62
11
12
63
44
42
M Sex
15
2
19
2
26
2
33
2
2
22
22
2
F
M
F
F
F
F
F
F
10
2
2
44
52
50
11
2
M
8
2
8
2
F
M
12
2
23
2
F
36
2
19
2
36
27
26
2
M
M
M
2
12
52
52
52
43
45
44
47
1
2
11
41
10
2
11
46
30
2
19
2
F
M
F
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
942089
942090
942091
942092
942093
942094
942095
942096
942097
953346
953347
953348
953349
953350
953351
953352
953353
953354
953355
953356
953357
953358
953359
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
ThorneRjver
Thorne River
Thorne River
ThorneRiver
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
Thorne River
ThorneRiver
ThorneRiver
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
7/12/94
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/8/95
8/8/95
8/8/95
8/8/95
8/8/95
8/8/95
154
168
208
34
11
10
62
31
9
13
12
7
11
40
80
273
202
45
440
330
348
249
232
253
217
515
277
11
12
61
14
12
6
11
11
62
12
7
11
11
61
9
9
13
11
14
12
7
10
12
13
9
13
11
7
10
11
12
63
62
32
36
34
34
9
11
140
146
144
157
149
141
31
9
10
12
7
11
12
12
61
13
11
6
10
11
9
13
11
6
11
11
12
8
13
10
6
11
11
12
62
63
62
10
13
11
7
12
12
12
61
132
149
9
13
13
7
12
12
11
64
145
9
13
11
6
11
ii
12
61
9
13
13
7
11
10
11
61
154
148
9
14
12
6
12
11
11
9
14
11
6
11
116
54
11
12
9
14
11
7
11
12
12
62
62
63
35
37
32
33
34
30
36
32
33
33
32
9
12
140
147
27
45 13
43 26
56 11
42 13
39 32
52 23
54 22
42 17
46 30
50 33
52 30
40 16
50 16
54 28
74 18
9
13
11
6
11
60
17
2
121
12
11
61
33
9
13
11
6
11
15
2
395
260
289
118
264
11
11
61
33
9
14
12
8
10
Ii
12
61
36
9
13
12
6
11
11
11
61
38
9
14
13
7
12
58
52
48
42
11
12
32
32
13
12
6
11
42
12
63
62
9
12
9
14
11
7
11
48
230
283
175
365
333
332
308
291
350
298
393
314
237
189
243
365
314
320
240
303
151
156
159
147
150
143
152
176
148
146
K
L
61
M Sex
2
2
2
2
2
F
M
M
F
F
2
M
F
F
F
F
M
2
F
2
F
F
F
2
2
2
2
2
2
2
2
24
2
15
2
20
18
2
2
16
2
F
M
F
F
F
F
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
953360
953361
920039
920040
Thorne River
Thorne River
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
8/8/95
8/8/95
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
202
82
285
11
12
61
152
33
9
13
11
7
11
2
11
185
37
9
14
11
6
11
13
2
34
80
30
58
43
29
22
10
11
48
50
44
55
35
11
92004P
920042
920043
920044
92004&
920047
920048
920049k
920310
920311
920312
920313
920050
920051
920052
920053
920054
920055
920056
321
147
199
143
183
156
139
128
116
126
134
233
177
126
115
205
206
222
257
242
243
202
11
11
12
11
12
12
62
60
63
60
63
10
12
61
11
11
62
12
12
61
15
11
11
20
26
120
53
22
39
82
93
93
164
124
139
Ii
ii
11
11
11
12
12
12
11
12
63
62
60
64
62
61
11
12
61
160
11
11
61
11
12
61
152
138
11
12
61
141
11
10
61
163
12
12
63
60
148
153
61
156
85
11
Ii
ii
ii
51
M
55
54
56
49
37
47
M
M
F
F
F
51
M
M
M
M
42
57
36
34
33
30
33
34
37
36
7
11
12
6
12
11
6
13
13
7
12
14
12
7
11
9
14
12
6
12
9
14
11
6
12
38
47
43
47
54
45
50
48
9
14
12
7
11
51
9
14
9
14
9
14
9
13
9
13
M
F
M
M
F
F
F
10
2
8
2
0
2
22
2
M
M
14
2
F
16
2
2
2
9
2
F
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
920057
920058
920059
920061
920062
920063
920064
920065
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
BakeweliCreek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewel! Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
BakewellCreek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
7/15/92
4/23/93
4/23/93
4/23/93
4/23/93
4/23/93
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
278
113
113
114
119
117
200
100
226
290
309
197
398
146
309
187
194
198
192
216
199
203
205
184
11
11
61
162
35
9
25
10
11
138
37
9
24
30
64
10
11
60
60
151
35
11
11
61
149
11
11
143
58
11
11
62
60
80
11
12
61
158
12
11
12
61
147
36
33
36
36
34
164
11
12
232
227
63
11
11
12
12
11
11
571
33
11
11
12
11
227
66
11
12
11
11
64
60
62
62
62
62
63
60
71
11
12
61
75
73
89
87
83
12
12
61
11
12
12
11
12
13
10
11
91
11
11
60
62
64
63
60
9318371
931838
931839
931840
931841
942114
942115
942116
942117
942118
942119
942120
942121b
942122
942123
148
H
I
J
14
12
7
14
11
7
9
15
13
6
13
9
14
11
6
11
9
14
11
8
12
9
14
11
6
12
9
15
12
7
12
9
13
11
5
11
K
L
M Sex
12
51
11
2
12
49
55
46
44
43
56
44
17
7
2
2
8
2
1
2
7
2
8
2
5
1
51
163
175
36
9
13
12
6
11
51
6
2
33
9
14
11
7
11
19
2
158
35
9
14
10
6
10
2
2
146
30
30
32
34
37
36
9
14
10
6
11
16
2
9
15
11
7
12
49
49
49
56
62
1
2
9
14
11
6
11
55
6
2
9
13
11
6
11
11
2
9
14
11
6
11
47
54
52
50
43
48
72
10
2
145
154
165
164
163
9
14
12
6
11
165
131
35
9
13
11
7
11
25
10
15
11
7
12
162
150
36
9
14
11
7
12
31
8
14
12
7
11
F
M
F
F
F
M
F
F
F
M
M
F
F
F
M
M
M
F
0
1
8
2
M
F
M
M
4
2
F
5
2
17
2
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
942124
942125
942126
942127
942128
942129
942130
94213!
964178
964179
964180
964181
964182
964183
964184
964185
964186
964187
964188
964189
964190
964191
964192
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
Bakewell Creek
BakeweliCreek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
7/14/94
8)6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
8/6/96
159
204
206
43
11
12
62
38
9
14
88
12
12
61
35
9
14
77
344
271
106
12
11
61
37
9
14
11
12
63
31
9
12
12
61
33
12
12
61
59
66
11
12
61
164
170
158
150
154
146
159
10
11
61
101
11
11
95
125
92
113
140
11
11
11
12
11
12
62
62
62
62
12
11
61
12
12
87
11
129
8(6(96
8/6/96
I
J
K
L
M
Sex
13
7
11
2
7
10
8
2
13
8
12
2
2
14
11
7
12
6
2
9
14
10
6
12
11
2
31
9
14
12
7
11
7
2
37
9
13
12
11
16
2
151
31
9
14
12
6
6
53
48
44
53
47
46
50
10
12
11
41
7
2
126
141
31
9
14
11
7
11
38
6
2
32
9
14
11
7
11
2
31
10
13
11
7
10
7
2
29
9
13
10
6
11
34
9
13
12
6
11
61
149
146
9
13
12
6
10
12
61
136
9
14
11
6
10
38
5
1
12
12
61
138
10
13
12
7
11
38
10
2
!22
11
11
63
141
9
14
12
6
11
2
11
12
61
136
9
14
12
7
11
1
1
119
1!
11
60
150
9
14
11
7
10
8
2
142
125
11
12
61
144
31
9
13
12
6
11
6
2
11
12
61
31
11
6
11
12
2
12
12
61
31
9
9
13
128
147
13
12
6
10
5
2
12
12
62
140
154
139
31
9
14
11
7
11
40
42
48
36
48
42
48
0
93
30
30
30
29
30
32
44
40
42
44
36
3
143
138
M
F
F
F
F
M
F
M
M
F
F
F
M
M
7
2
330
303
228
175
187
9
1
10
2
5
2
M
M
F
F
M
M
M
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
964193
964194
964195
964196
964197
964198
964199
964200
964201
964202
953506
953507
953508
953509
953510
953511
953512
953513
953514
953515
953516
953517
953518
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Herman Creek
Survey Creek
Survey Creek
SurveyCreek
Survey Creek
SurveyCreek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
SurveyCreek
Survey Creek
8/6/96
8/6/96
8/7/96
8/7/96
8/7/96
8/7/96
8/7/96
8/7/96
8/7/96
8/7/96
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
A
B
C
D
E
F
G
H
I
121
12
12
145
13
11
11
11
126
9
14
11
160
10
12
9
14
121
11
12
61
136
9
13
156
155
12
12
62
159
32
30
28
28
29
10
83
62
62
62
9
10
11
61
138
33
143
130
10
11
11
62
62
134
10
122
129
145
178
159
11
12
61
11
12
28
54
12
12
11
12
37
10
236
131
195
72
73
J
K
L
M Sex
6
11
46
10
2
6
10
38
6
1
12
52
11
2
11
7
11
2
14
12
7
11
9
14
12
6
11
32
10
14
12
7
11
138
31
9
13
12
7
11
130
28
14
10
131
14
12
11
13
11
6
11
10
15
14
8
12
9
14
12
7
Ii
12
12
63
38
9
14
11
7
10
12
Ii
4
11
12
63
64
139
167
137
148
142
9
11
63
63
62
30
34
36
30
6
7
11
61
9
10
40 14
42 10
48 4
46 7
42 7
36 28
46 2
42 9
57 27
48 8
54 13
87
7
11
11
207
142
209
83
194
148
91
11
11
27
80
11
11
12
12
6
11
12
63
29
11
13
12
12
64
62
63
63
63
63
63
142
35
9
13
12
7
11
133
34
9
14
12
7
10
135
35
9
13
12
7
12
147
37
10
14
11
7
12
139
33
9
14
13
7
11
148
31
9
14
12
7
11
139
31
9
13
12
7
11
150
32
35
9
15
12
8
11
9
14
12
7
11
154
48
42
53
49
35
47
44
44
2
2
2
2
1
2
2
F
M
M
F
M
M
F
F
F
F
F
2
M
M
M
16
2
F
7
1
2
2
1
2
15
2
12
2
6
2
10
2
12
2
20
2
M
M
M
M
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
953519
953520
953521
953522
953523
953524
953525
953526
953527
Survey Creek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
Survey Creek
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/27/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/28/95
8/29/95
8/29/95
8/29/95
8/29/95
8/26/95
96
93
8
12
12
146
14
12
7
12
38
7
1
12
13
134
32
32
9
8
62
63
9
14
12
7
11
35
8
2
121
16
11
12
61
140
9
14
12
7
11
38
10
2
121
14
11
12
9
14
12
7
12
2
11
11
9
15
12
7
11
12
2
12
12
10
15
12
7
11
44
39
45
10
89
71
173
379
163
152
508
13
2
11
11
10
14
11
7
12
38
29
2
11
12
8
14
14
8
11
2
12
9
14
12
7
10
52
42
15
11
62
62
62
62
62
62
159
215
196
275
355
266
259
395
265
286
216
169
254
279
230
330
329
330
312
235
34
35
33
35
30
32
30
10
2
12
13
61
30
33
9
14
13
7
11
56
13
11
7
10
39
28
26
2
9
32
29
30
34
9
14
12
8
11
35
27
17
2
9
14
12
8
10
9
14
11
7
10
9
10
11
6
10
31
9
14
12
7
10
32
32
33
9
14
12
7
11
8
13
12
6
11
9
14
14
7
12
9535281
953529
953530
95353P
953532
953533
953534
953535
953536
953537
953538
953539
953540
953503
YakounRiver
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
YakounRivei
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
Yakoun River
151
153
156
148
145
ii
12
63
147
165
12
12
135
183
11
12
62
62
99
46
136
187
116
312
312
299
290
132
11
12
61
132
11
12
11
11
11
12
62
62
63
12
11
61
10
11
140
145
153
149
144
12
11
11
12
60
62
62
11
12
61
153
33
9
13
13
8
11
10
ii
61
154
34
9
14
12
7
ii
151
137
M Sex
2
11
2
40
18
2
12
2
25
48
38
56
40
34
24
2
37
2
39
26
2
13
2
22
2
2
F
M
M
M
M
M
F
F
M
M
M
F
F
M
M
M
M
F
M
F
F
M
Appendix Table A2.1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
953504
953505
964258
964259
964260
964261
964262
964263
964264
964265
964266
964267
964268
964269
964270
Yakoun River
Yakoun River
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly WalkerCreek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
8/26/95
8/26/95
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
206
83
11
11
120
16
11
12
149
34
11
12
118
19
11
12
163
48
43
10
11
11
12
964271
964272
964273
964274
964275
964276
964277
964278
10129196
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
C
D
E
F
140
132
149
31
G
H
9
14
9
14
10
I
J
K
L
M Sex
12
6
10
2
7
11
24
2
14
12
8
11
60
52
34
8
13
8
2
9
14
12
7
11
34
12
2
10
14
11
6
12
60
5
2
9
14
11
8
11
38
11
2
10
14
12
6
11
10
2
9
9
10
14
12
7
10
50
44
14
2
14
11
6
11
38
3
2
13
12
6
12
9
2
9
13
12
6
11
2
2
9
14
4
4
2
2
2
4
2
2
2
3
2
9
2
42
11
11
146
31
11
12
63
62
62
62
62
62
62
62
130
142
132
109
24
33
29
11
11
63
10
13
62
142
141
10
11
61
161
14
11
12
155
12
7
11
107
16
11
12
62
62
30
35
29
34
34
30
34
32
32
34
34
146
35
9
14
11
7
11
101
13
12
12
61
141
34
9
14
11
7
11
180
51
12
11
152
33
9
13
11
6
10
131
12
12
134
14
11
6
11
11
11
63
147
30
30
9
133
27
30
62
62
10
14
11
7
10
144
37
11
12
62
142
33
10
14
12
7
10
100
13
11
11
61
136
37
9
14
11
7
11
30
38
44
42
34
44
36
46
42
22
6
2
108
14
12
12
63
167
37
9
13
11
5
11
38
4
2
90
95
9
10
12
157
13
10
7
11
42
5
2
10
12
9
14
12
7
11
38
6
2
113
16
11
12
33
33
33
10
12
62
62
62
9
14
12
6
11
42
3
2
164
148
151
152
147
133
145
139
151
4
F
F
F
M
M
F
F
M
M
M
M
M
M
M
M
F
M
M
M
F
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
964279
964280
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Molly Walker Creek
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
110
16
11
12
91
8
11
11
86
107
143
132
122
113
104
95
103
97
68
108
105
85
174
208
124
137
133
103
109
9
11
12
964281
964282
964283h
964284
964285
964286
964287
964288
964289
964290'
96429 P
964292
964293
964294
Fish Creek(BellaCoolaRiver)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
FishCreek(BellaCoolaRiver)
Fish Creek (Bella Coola River)
964295 Fish Creek (Bella Coola River)
964296 Fish Creek (Bella Coola River)
964297 Fish Creek (Bella Coola River)
964298 FishCreek(BellaCoolaRiver)
964299 Fish Creek (Bella Coola River)
964300 Fish Creek (Bella Coola River)
96430P Fish Creek (Bella Coola River)
C
D
E
F
138
12
62
62
63
133
34
32
32
10
10
61
140
28
11
12
62
21
11
11
61
150
144
18
11
12
16
11
11
62
62
138
149
15
11
11
61
11
11
12
62
133
128
12
ii
12
61
10
10
12
62
G
H
I
9
14
11
10
14
12
10
13
11
7
11
33
9
13
12
6
11
37
9
13
12
7
10
33
32
9
14
12
6
11
9
13
12
6
10
32
9
13
12
6
11
31
9
13
11
7
12
9
9
13
11
7
11
146
28
35
13
12
7
11
135
J
K
6
10
6
11
52
36
26
32
48
44
48
42
48
46
40
L
M Sex
2
2
2
2
5
2
2
1
M
F
M
M
4
2
2
5
2
4
2
3
1
F
M
M
M
M
M
5
2
F
5
2
M
5
2
M
0
3
1
5
11
12
13
12
12
61
158
33
9
13
12
6
11
13
11
12
61
140
32
9
13
ii
6
12
8
11
13
62
62
110
24
11
ii
63
9
13
ii
5
10
50
5
2
11
31
9
13
11
7
12
2
M
M
11
132
9
14
11
7
10
1
1
M
21
12
12
9
13
11
6
11
10
2
27
10
10
63
30
33
30
42
48
46
5
12
62
63
63
140
144
34
11
9
13
11
6
11
36
3
2
F
M
14
11
11
13
12
11
62
63
135
143
42
44
Appendix Table A2. 1. Continued
Fish
number
Location
964302U Fish Creek (Bella Coola River)
964303
964304
964305
964306
964307
964000
964001
964002
964003
964004
964005
964006
964007
964008
964009
964010
964011
964012
964013
964014
964015
964016
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Befla Coola River)
Fish Creek (Bella Coola River)
Fish Creek (Bella Coola River)
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
SanJosefRiver
San Josef River
San Josef River
San Josef River
SanJosefRiver
SanJosefRiver
Date of
FL
Wt.
collection
(mm)
(g)
A
B
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
10/29/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
137
11
11
140
26
25
11
12
61
119
17
11
13
61
63
50
43
245
173
194
4
11
12
62
3
11
12
2
10
12
130
11
11
60
49
70
224
74
23
10
11
61
148
158
11
11
11
10
62
60
148
160
11
11
61
11
11
60
144
11
11
61
124
11
11
61
17
10
11
61
47
65
27
35
78
29
59
44
12
11
61
11
12
H
11
11
12
11
11
62
62
62
60
146
138
150
154
138
148
146
10
12
61
10
11
11
11
286
193
137
255
245
119
169
187
135
154
222
145
183
168
C
D
E
F
G
H
I
J
K
L
M
Sex
56
62
56
52
48
52
56
50
4
2
M
7
2
F
22
M
13
2
2
25
2
15
2
11
2
14
2
M
M
F
M
F
F
M
F
M
F
F
F
M
63
31
10
15
11
7
11
31
9
13
11
7
11
31
9
14
11
6
12
32
28
30
30
30
9
14
12
11
10
15
12
6
6
10
15
12
6
11
9
14
10
8
11
ii
7
10
9
12
34
9
14
11
9
14
11
6
6
11
31
30
34
30
10
14
11
7
11
10
14
12
7
12
9
13
11
5
11
9
14
12
6
147
33
29
9
13
12
61
159
33
10
14
64
170
33
10
14
151
147
147
11
2
36
2
10
6
2
2
2
11
46
44
48
48
42
13
2
6
11
58
12
2
13
7
11
5
2
12
7
11
46
54
7
2
11
14
F
-a
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
964017
964018
964019
964020
964021
964022
964023
964024
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
San Josef River
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/6/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
140
24
231
55
11
11
11
11
10
10
23
64
12
11
12
12
62
60
62
60
62
38
69
12
12
11
219
2
10
12
12
1
12
12
11
13
127
83
22
12
12
108
12
11
11
102
11
11
12
96
10
12
11
109
13
10
964026
964027
964028b
964029
964030
964031
964032
964033h
964034b
964035
964036h
964037
964038h
964039b
WillowCreek
Willow Creek
Willow Creek
WillowCreek
Willow Creek
292
177
132
189
155
192
296
50
45
203
D
E
F
G
H
152
9
13
13
9
14
10
9
14
12
7
11
10
14
12
7
11
148
30
33
33
30
29
9
14
11
7
12
63
131
31
10
14
11
6
11
12
61
9
10
14
14
12
12
6
11
7
12
9
14
12
7
12
11
6
10
12
60
62
63
63
63
62
64
64
62
144
154
117
118
132
125
122
124
119
32
11
159
143
153
126
45
54
54
1
11
12
61
2
11
11
63
113
118
1
11
11
64
116
51
1
ii
12
63
104
1
11
12
1
10
12
1
11
12
63
63
63
48
54
54
126
128
31
28
23
33
27
27
29
27
28
27
30
27
27
27
30
25
10
J
K
6
11
7
12
54
46
52
48
52
56
42
44
I
L
M Sex
F
F
1
2
10
2
12
2
F
18
2
M
M
M
F
M
21
2
1
2
25
16
2
2
0
1
1
9
14
12
8
12
9
15
13
7
12
10
14
12
7
12
9
14
11
7
11
9
14
11
7
12
9
13
11
6
10
9
13
13
6
9
15
12
9
13
12
10
14
ii
6
12
0
1
10
14
13
7
9
0
1
9
14
12
7
11
0
1
9
14
11
6
10
0
1
46
52
4
0
2
34
32
32
42
5
1
7
6
1
1
10
6
0
1
6
11
0
1
6
11
1
1
1
M
M
F
M
M
M
Appendix Table A2. 1. Continued
Fish
number
964041k'
964043b
964044
964045h
964046
964047
964048
964049b
964153
964154
964156
964157
964158
964159
964160
964161
964162h
964163
964164
964165
Date of
FL
Wt.
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
Willow Creek
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/7/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
50
1
11
12
63
119
9
13
1
11
12
123
9
1
11
12
1
12
112
96
12
11
151
68
13
12
133
37
20
12
12
131
11
12
117
15
11
11
59
54
62
48
75
2
11
11
62
62
63
62
64
66
62
63
63
27
26
26
25
2
12
12
61
2
11
11
61
1
11
12
61
4
13
12
83
6
12
12
60
62
78
62
5
11
12
61
2
11
12
7/1 1/96
47
1
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
61
2
11
58
2
69
WillowCreek
WillowCreek
MixaiCreek
Mixal Creek
Mixal Creek
Mixal Creek
Mixal Creek
Mixal Creek
MixaiCreek
Mixal Creek
Mixal Creek
Mixal Creek
MixaiCreek
MixaiCreek
Mixal Creek
47
49
46
221
186
150
H
I
J
11
0
1
13
12
6
11
0
1
9
13
12
7
12
9
13
12
5
11
31
9
15
12
7
12
0
0
4
2
10
14
13
8
12
0
1
137
32
30
9
14
13
8
11
0
2
140
33
9
14
12
8
12
5
2
135
10
14
12
7
12
8
2
9
14
12
9
14
11
6
7
12
138
32
26
34
126
29
9
14
11
7
10
118
123
12
K
38
48
50
40
44
32
34
24
L
0
4
M Sex
1
1
F
M
M
M
M
1
1
M
M
0
1
44
36
34
7
1
F
M
M
M
6
1
M
4
1
M
0
1
38
158
36
9
14
12
7
12
62
145
29
9
13
11
7
10
12
60
136
30
9
14
12
7
11
11
ii
62
141
31
9
13
12
7
11
42
34
4
11
11
61
142
34
9
14
11
7
11
38
65
3
11
12
61
141
31
9
14
12
7
11
62
3
11
12
60
140
30
9
13
11
7
11
32
5
1
2
1
2
1
M
F
M
Appendix Table A2.1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
964 166
Mixal Creek
Mixal Creek
Mjxal Creek
Mixal Creek
Mixal Creek
MixaiCreek
Mixal Creek
Mixal Creek
Mixal Creek
Mixal Creek
Mixal Creek
Mixal Creek
SalmonRiver
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
SalnionRiver
SalmonRiver
Salmon River
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
7/11/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
46
64
61
964167
964168
964169
964170h
964171
964172à
964173
964174
964175
964176
964177
964203
964204
964205
964206
964207
964208
964209
964210
964211
964212
964213
44
53
60
73
64
70
75
78
74
174
173
167
142
176
132
144
145
139
150
150
A
B
C
D
E
F
G
H
I
J
K
L
3
11
11
13
11
7
12
1
9
13
11
7
11
32
26
6
11
32
29
9
12
62
60
156
2
8
1
1
12
12
61
131
12
7
11
1
12
61
128
9
12
12
7
10
36
24
0
12
3
1
4
13
13
153
9
14
6
10
3
11
11
62
62
9
13
12
7
11
2
2
159
138
29
26
33
30
32
32
9
2
9
13
11
12
1
9
14
12
7
7
M
M
M
12
0
2
M
8
1
M Sex
1
135
M
M
1
4
11
12
61
5
11
12
62
5
11
12
61
4
48
48
38
25
44
11
12
11
11
11
11
11
142
31
8
13
11
8
11
61
160
144
31
9
14
12
7
11
130
136
146
13
12
7
10
13
9
6
10
8
13
11
6
9
10
11
61
13
11
7
11
11
60
127
147
9
11
9
13
11
7
10
18
ii
ii
61
139
Q
13
11
6
10
25
25
11
11
61
147
9
13
11
6
10
11
11
61
144
9
15
11
7
11
21
11
11
60
138
9
13
10
6
ii
28
26
12
11
61
9
14
10
7
11
11
12
61
154
136
30
29
29
29
30
30
33
32
30
35
30
9
9
12
59
60
62
9
13
11
6
11
34
42
32
38
38
32
48
44
38
42
54
44
42
34
52
42
50
1
5
1
23
2
3
2
3
2
M
M
M
M
M
4
2
F
F
M
4
2
F
1
2
3
2
10
2
13
17
2
2
M
F
M
M
5
2
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
964214
964215
964216
964217
964218
964219
964220
964221
964222
964223
964224
964225
964226
964227
964228
964229
964230
964231
964050
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Salmon River
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
6/6/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
147
29
12
11
61
134
35
9
13
11
7
11
2
14
11
Ii
62
141
29
9
14
13
7
11
128
16
11
11
61
141
32
9
13
11
7
10
44
44
42
13
123
11
38
12
964051
964052
964053
964054
2
2
6
2
1
2
M
F
M
M
44
36
54
0
2
F
1
2
2
2
40
42
5
2
F
M
F
3
2
13
1
3
5
2
2
4
2
2
2
5
2
2
2
6
2
130
16
11
12
62
145
31
9
14
11
124
14
11
10
61
137
31
9
13
11
6
6
150
27
10
11
136
31
9
13
11
6
11
144
21
10
12
153
31
9
13
11
8
11
130
134
125
130
114
123
124
18
12
11
60
62
62
14
11
6
11
11
12
61
9
12
12
7
11
15
11
11
61
30
29
30
9
20
132
139
143
9
13
11
6
12
17
11
11
61
121
31
9
13
10
7
10
12
12
12
9
13
11
7
11
10
12
29
9
13
10
7
11
16
11
10
31
9
14
11
7
12
141
22
11
10
130
134
132
144
31
15
13
10
7
10
14
11
11
9
13
11
7
10
133
19
10
11
61
125
32
29
33
9
120
62
60
62
59
62
9
13
ii
8
11
144
26
20
11
11
61
136
31
9
13
10
7
11
12
13
59
147
9
13
12
7
11
11
11
12
61
139
9
14
12
6
25
26
11
12
61
142
9
14
12
6
10
11
12
61
140
9
15
12
7
11
8
12
12
61
139
30
32
32
32
33
38
50
46
38
42
44
46
42
52
46
44
50
36
9
13
9
6
11
46
115
99
132
130
85
131
1
2
16
2
11
2
11
2
16
2
12
2
M
F
F
M
F
F
M
F
M
F
M
M
F
M
M
-a
Appendix Table A2. 1. Continued
Fish
number
Location
964055
964056
964057
964058
964059
964060
964061
964062
964063
964064
964065
964066
964067
964068
964069
964070
964071
964072
964073
964074
964075
964076
964077
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
RitherdonCreek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
RitherdonCreek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Sandhill Creek
Sandhill Creek
SandhillCreek
Date of
FL
Wt.
collection
(mm)
(g)
A
B
C
D
E
137
30
23
26
24
23
11
12
61
12
12
61
12
12
61
11
12
60
11
12
61
12
11
11
62
16
11
13
61
16
11
12
60
10
ii
12
61
8
11
12
61
144
132
144
132
152
127
150
140
142
136
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/8/96
7/10/96
7/10/96
7/10/96
128
135
130
125
108
116
119
94
88
94
F
U
31
9
30
10
H
I
J
K
L
15
10
7
11
2
12
8
10
40
44
42
44
50
44
58
44
46
42
44
42
38
42
40
50
44
48
48
32
42
44
36
13
14
6
2
33
9
14
11
5
11
28
10
14
11
6
11
31
9
15
11
5
10
29
32
9
15
11
6
11
9
14
12
6
11
31
9
14
12
7
10
9
14
12
6
10
9
13
10
6
11
145
29
34
32
9
15
11
6
11
132
31
9
14
12
6
11
132
140
32
30
9
14
12
5
11
9
14
10
6
11
127
31
9
14
11
7
11
135
31
9
14
10
6
11
153
128
30
32
9
13
11
6
11
9
7
13
7
128
31
9
14
11
6
10
128
29
32
32
34
9
13
12
7
10
9
14
13
8
11
9
14
11
7
10
9
14
11
7
11
9
12
12
90
9
11
12
93
10
12
12
89
7
12
12
85
7
11
11
83
89
88
80
80
148
119
150
7
11
11
60
60
60
62
60
60
8
12
12
61
9
11
12
6
12
12
5
12
12
37
22
11
11
60
60
62
60
11
10
61
146
37
11
10
61
137
138
M Sex
13
2
14
2
13
2
15
2
13
2
6
2
10
2
6
2
3
2
8
2
9
1
7
2
6
2
1
2
7
2
3
2
2
1
9
2
5
2
5
2
0
2
M
F
F
F
M
F
M
F
M
M
F
F
M
M
M
M
M
F
M
M
F
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
A
B
C
D
E
F
G
HI
51
12
11
60
9
14
13
47
12
11
61
37
9
13
10
112
16
12
11
31
9
13
11
8
10
181
67
10
11
14
11
7
19
12
11
12
12
11
61
9
9
14
26
13
11
16
11
11
62
9
14
11
7
10
20
22
27
35
12
11
61
38
37
34
33
30
9
122
130
116
59
62
62
9
13
11
7
12
12
10
60
33
9
13
10
7
11
12
11
61
150
150
147
148
154
134
144
142
140
137
35
163
9
13
10
6
10
11
10
9
13
12
7
10
17
11
11
131
24
11
10
60
60
60
121
19
12
10
61
125
24
12
11
61
32
36
32
33
38
33
29
34
32
36
32
36
33
number
Location
collection
(mm)
964078
964079
964080
964081
964082
964083
964084
964085h
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Saridhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhifl Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Sandhill Creek
Kirby Creek
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
159
964086
964087
964088
964089b
964090
964091
964092
964093
964094
964095
964096
964097
964098
964099
964 iooa
7/10196
wt.
124
126
137
138
108
146
139
138
146
134
140
148
143
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
105
13
12
11
61
115
17
12
12
114
16
10
11
63
61
128
25
11
12
61
113
18
10
11
61
154
135
179
142
63
11
11
61
136
32
11
11
60
140
7/10196
91
8
J
K
L M Sex
8
10
2
11
0
2
11
36
38
34
32
2
7
6
9
7
11
9
14
11
8
11
9
14
12
7
10
9
13
11
8
12
9
13
11
7
10
9
13
12
7
9
14
11
7
11
9
13
ii
7
ii
9
13
11
7
11
9
13
11
7
10
10
14
12
6
11
9
13
11
8
11
4
1
9
2
46
11
1
36
42
56
38
36
36
32
36
42
40
36
4
1
1
2
M
M
F
M
M
0
1
F
F
F
0
2
M
9
1
0
2
0
1
6
4
2
0
2
3
2
46
0
2
38
48
42
6
1
1
0
2
5
1
38
5
2
36
10
2
F
M
M
F
M
M
F
M
F
M
F
M
M
.J1
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
964101
964102
964103
964104
964105
964106
964107
964108
964109
964110
964111
964112
964113
964114
964115
964116
964117
964118
964119
964120
964121
964122
964123b
Kirby Creek
Kirby Creek
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
7/10/96
83
158
6
11
11
62
144
45
11
11
61
141
117
17
10
10
61
138
98
10
11
11
62
155
83
146
6
11
11
61
138
29
11
12
62
153
131
23
12
Ii
61
133
98
10
11
11
61
141
79
103
151
137
87
104
84
89
90
106
5
11
11
141
11
11
11
11
11
157
142
31
36
26
7
62
62
62
38
32
33
37
37
32
29
32
30
37
10
11
61
161
33
12
11
61
142
12
12
11
136
6
11
11
60
62
30
34
145
7
11
11
61
143
8
11
11
155
12
10
11
107
13
11
10
62
62
62
99
9
12
11
61
124
18
11
10
62
96
96
9
11
10
61
151
8
13
14
60
121
KirbyCreek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
140
142
156
147
F
G
H
I
J
K
L
M Sex
9
9
13
11
6
10
11
6
11
40
42
6
14
10
2
2
9
13
11
5
11
2
13
11
6
11
7
2
13
12
6
10
10
2
10
14
11
14
10
6
6
11
8
9
13
11
6
10
9
13
11
6
11
38
42
34
36
32
38
30
16
9
9
13
11
6
11
10
13
11
6
11
9
13
11
6
11
14
11
6
10
9
14
11
6
10
42
44
42
32
42
33
9
14
10
6
11
32
34
27
9
14
11
7
11
8
14
11
7
10
9
14
10
6
12
31
9
14
ii
6
10
34
33
36
26
9
13
9
6
10
9
14
11
7
11
9
14
12
7
11
10
14
12
7
11
12
38
32
44
48
40
38
36
38
32
1
2
F
M
M
F
M
M
3
2
M
12
2
5
1
7
2
5
2
5
2
8
1
9
2
7
2
15
2
8
2
3
2
13
2
7
2
13
2
12
2
0
1
F
M
M
M
F
M
M
M
M
M
M
M
M
M
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
964124
964125
964126
964127
953106
953107
953108
953109
953110
953111
953112
953113
953114
953115
953116
953117
953118
953119
953120
953121
953122
953123
953124
Kirby Creek
Kirby Creek
Kirby Creek
Kirby Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
ChurchCreek
Church Creek
Church Creek
ChurchCreek
ChurchCreek
Church Creek
ChurchCreek
ChurchCreek
Church Creek
ChurchCreek
Church Creek
ChurchCreek
Church Creek
7/10/96
84
6
11
11
61
144
30
10
14
10
6
10
7110/96
117
16
11
12
133
33
9
14
12
7
11
7/10/96
7/10/96
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
98
93
102
9
12
11
60
62
137
13
11
6
10
11
11
61
136
35
32
9
10
9
14
11
7
11
11
11
11
62
31
9
13
12
5
10
111
14
11
11
61
31
9
13
11
7
11
105
12
12
12
61
137
136
129
33
9
14
10
7
10
36
42
42
30
46
42
42
107
11
11
11
142
30
32
9
13
11
7
11
9
13
11
31
9
13
12
6
7
11
32
29
9
13
12
7
10
9
11
9
13
12
9
13
13
7
6
6
11
31
10
46
36
36
34
9
13
11
7
11
38
10
13
11
7
11
92
8
10
11
62
62
104
11
11
12
61
6/15195
105
11
10
12
62
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
87
99
8
10
11
61
9
12
11
62
105
11
11
11
61
132
143
127
136
137
114
14
11
11
61
131
98
9
11
11
125
87
7
10
11
62
60
30
29
30
121
28
9
13
11
6
11
135
10
48
42
36
L M Sex
11
1
12
2
2
2
10
9
2
2
1
2
4
2
11
2
3
2
2
3
7
2
2
2
3
2
12
2
1
6
M
M
M
M
M
M
M
M
F
M
M
F
120
15
12
13
61
154
33
9
13
11
7
11
34
4
2
112
103
14
11
11
62
131
31
9
12
11
7
10
2
11
11
61
133
31
9
13
12
7
12
7
2
115
14
10
11
61
30
9
14
12
7
10
3
2
F
105
12
10
10
61
134
137
31
9
14
11
6
11
50
46
44
34
5
11
M
M
F
F
F
M
F
F
3
1
96
9
11
11
62
138
29
9
13
11
6
11
40
8
2
F
M
48
34
1
1
6
1
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
953125
953126
953127
953128
953129
953130
953131
953132
953133
953134
953135
953136
953137
953138
953139
953140
953141
ChurchCreek
Church Creek
Church Creek
ChurchCreek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
ChurchCreek
ChurchCreek
6/15/95
6/15/95
6/15/95
135
11
ii
137
22
25
11
liii
13
6115/95
100
6/15/95
6/15/95
6/15/95
94
100
116
109
110
96
953142d
ChurchCreek
Church Creek
Church Creek
Church Creek
Church Creek
Church Creek
953143
953144
953145
953146
953147
Church Creek
Church Creek
6/15195
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
C
D
E
F
G
H
I
J
K
141
9
9
12
7
11
13
10
5
10
10
11
63
142
9
13
11
7
11
10
10
11
13
11
8
10
10
11
138
132
9
8
62
62
9
13
3
11
10
11
11
61
137
9
14
12
6
12
16
10
11
61
142
9
13
12
7
11
13
11
11
60
12S
9
12
11
7
10
13
10
12
61
9
13
12
8
11
36
36
46
44
42
50
34
48
64
9
11
11
60
147
126
9
12
11
7
10
118
17
11
11
61
143
9
13
12
6
10
98
10
10
11
61
127
33
24
35
32
30
34
34
32
29
30
32
32
13
12
60
62
9
13
12
7
11
103
ii
11
11
141
31
9
14
11
7
10
124
18
11
11
62
60
131
31
9
13
12
6
11
100
10
Ii
11
61
139
9
13
11
7
9
126
21
11
10
62
134
33
34
9
13
12
7
11
90
92
8
11
11
61
131
7
11
11
63
129
129
20
ii
11
61
131
95
148
94
104
8
11
11
61
128
26
11
11
61
147
8
12
12
61
128
30
32
28
30
30
30
11
11
12
62
138
31
131
9
13
11
6
9
9
13
ii
6
11
9
14
12
14
11
6
6
10
9
8
13
11
6
9
9
13
11
6
11
9
13
11
6
10
12
42
42
34
34
48
42
44
42
58
34
34
46
44
48
L
M Sex
15
2
10
2
1
2
5
2
11
1
4
2
2
2
6
2
5
2
3
1
12
2
7
2
5
2
11
2
1
2
8
2
4
2
2
3
2
8
2
0
2
2
8
2
1
F
M
M
M
M
M
F
M
F
M
F
F
M
M
M
M
F
M
F
F
F
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
953148
953149
953150
953151
953152
953153
953154
ChurchCreek
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/15/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
111
13
11
11
139
30
9
14
12
6
11
2
20
11
11
133
33
9
14
11
6
12
7
2
103
12
12
11
61
140
33
9
13
11
7
11
48
42
36
8
125
62
60
6
1
109
13
11
11
61
140
9
14
12
6
38
3
2
9
14
31
2
2
9
13
12
6
10
37
3
2
32
4
2
8
2
4
1
3
2
F
15
1
953155
953156
953157
953158
9531591
953160
9531611
953426
953427
953428
953429
953430
953431
953432
953433
953434
Church Creek
Church Creek
ChurchCreek
Church Creek
ChurchCreek
Church Creek
Church Creek
ChurchCreek
ChurchCreek
Church Creek
Church Creek
Church Creek
Church Creek
Bear Creek
BearCreek
Bear Creek
Bear Creek
BearCreek
Bear Creek
Bear Creek
Bear Creek
BearCreek
99
9
11
12
61
126
112
14
11
11
61
133
33
32
32
112
14
11
11
132
31
10
14
10
7
11
93
109
105
102
102
133
120
90
94
97
93
107
93
77
136
179
8
11
12
61
121
9
13
12
11
11
14
10
11
11
147
121
9
11
9
14
12
11
12
11
125
31
9
13
11
7
12
11
12
11
62
61
62
62
6
6
7
11
12
29
30
28
132
9
12
11
6
10
33
10
12
63
131
30
29
9
12
12
6
10
16
11
11
132
30
9
12
10
6
11
7
11
ii
138
30
9
14
10
6
11
8
12
12
62
63
63
141
31
8
14
11
6
11
9
12
13
61
140
32
9
14
12
6
12
9
11
11
134
35
9
14
11
7
10
12
11
12
120
14
9
6
11
12
12
9
14
11
7
12
4
25
54
10
11
8
13
11
12
14
11
11
11
9
9
5
7
12
12
29
32
34
30
32
9
9
62
62
62
63
62
62
14
11
7
10
141
134
131
144
11
11
12
42
36
33
7
1
M
M
M
F
M
21
2
M
4
2
F
12
M
15
2
2
15
2
2
36
33
38
38
34
44
46
44
44
44
M
F
M
F
F
M
M
M
M
8
2
2
8
1
15
2
18
2
F
F
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
number
Location
collection
(mm)
953435
953436
953437
953438
953439
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
Bear Creek
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/21/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
175
77
73
84
84
66
115
45
39
78
95
78
94
95344Ø
95344l
953442
953443
953444
953445
953446
953447
953448
953449
953450
953476
953477
953478
953479
953480
953481
953482
BearCreek
Bear Creek
Bear Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
wt.
A
B
57
11
12
5
12
13
4
13
12
8
11
12
6
10
12
12
12
5
12
12
8
11
12
HI
C
D
E
F
62
62
61
62
60
141
9
14
10
9
13
10
129
30
32
28
30
30
133
64
62
133
114
123
G
3
K
L
MSex
5
Il
2
11
36
32
16
7
4
1
9
1
3
1
12
2
12
2
9
14
11
6
11
9
14
11
7
10
9
14
11
7
10
31
9
13
11
5
11
30
30
9
13
12
4
11
9
14
11
7
11
136
130
31
9
14
11
6
11
32
9
14
10
11
143
31
9
14
11
6
6
134
31
9
14
11
5
9
130
3
15
1
44
34
40
42
52
M
F
M
F
M
F
M
M
1
137
42
42
7
1
14
1
7
1
20
2
11
17
2
2
M
F
4
8
11
12
111
13
10
12
99
11
13
13
127
20
11
11
62
62
62
62
105
13
11
12
61
153
33
9
13
11
6
10
123
19
12
12
61
132
31
9
13
11
6
11
104
14
12
11
63
145
8
13
11
6
10
137
27
11
12
61
152
30
30
9
13
11
7
11
106
12
12
12
61
142
31
8
13
10
4
11
38
5
1
106
106
10
11
11
62
13
12
6
10
34
6
2
13
12
63
32
30
9
9
144
136
10
15
13
6
11
54
2
1
10
40
42
42
36
44
36
5
2
10
2
38
7
2
42
4
2
M
M
F
F
F
F
M
M
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
953483
953484
953485
953486
953487
953488
953489
953490
953491
953492
953493
953494
953495
953496
953497
953498
953499
953500
942003h
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
Salt Creek
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
6/22/95
5/13/94
5/13/94
5/13/94
5/13/94
5/13/94
80
92
78
84
97
5
11
12
11
5
10
34
2
1
12
38
6
2
5
11
11
61
143
32
34
29
13
12
139
150
9
8
60
62
2
1
6
12
12
61
150
3
2
9
11
11
61
151
7
2
81
5
11
11
2
2
70
4
12
11
62
62
2
1
143
31
11
11
5
2
138
25
11
11
15
165
46
12
11
2
2
2
83
6
11
133
26
11
18
2
3
2
8
2
942004
942005
942006
942007
SaltCreek
Salt Creek
Salt Creek
Salt Creek
Dickey River
Dickey River
Dickey River
Dickey River
Dickey River
14
12
7
11
9
14
11
6
10
31
9
14
12
7
11
31
9
14
11
11
147
33
9
14
12
6
6
138
31
8
13
10
5
11
61
136
31
9
13
12
6
11
144
9
13
11
6
10
9
14
12
8
12
11
62
62
63
11
61
147
11
42
34
38
38
42
32
40
M Sex
94
9
12
11
62
149
30
30
32
29
32
8
12
11
7
11
109
14
10
11
61
153
33
9
13
11
6
11
126
105
99
76
106
186
20
11
11
63
148
34
9
14
10
6
12
40
40
44
9
2
11
11
11
61
31
9
13
11
4
11
35
6
2
11
11
11
62
142
156
35
9
14
11
7
11
2
11
12
61
151
36
8
14
12
5
11
42
44
12
4
6
2
13
12
64
117
23
10
15
13
6
10
51
0
1
25
2
14
2
12
2
12
2
154
145
9
13
11
6
11
10
14
11
5
11
11
11
63
157
37
9
14
12
6
10
185
12
12
144
30
9
14
11
6
10
221
11
12
62
61
168
31
9
14
12
7
10
45
69
47
228
12
11
61
157
32
9
13
12
6
11
41
1
M
F
M
F
F
M
F
M
M
F
F
F
F
F
F
F
M
F
F
M
F
00
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
942008
942011
942012
942013
942014
942015
942016
942017
953401
953402
953403
Dickey River
Dickey River
5/13/94
5/27/94
5/27/94
5/27/94
5/27/94
5/27/94
5/27/94
5/27/94
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
104
953404
9S3405
953406
95340T
953408
953409
95341 oa
95341P
953412
9534 13
9534 14
953415
DickeyRiver
Dickey River
Dickey River
Dickey River
Dickey River
Dickey River
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
A
B
C
D
E
F
G
H
I
J
9
14
11
7
9
14
12
6
9
14
12
6
11
10
14
12
6
10
K
L
12
51
13
2
11
8
2
11
11
62
139
163
35
11
12
61
147
159
35
12
12
62
147
36
32
34
154
31
12
13
61
136
31
160
150
35
12
12
61
32
9
13
12
6
10
11
11
62
31
9
14
12
6
10
149
29
27
140
158
12
12
61
137
32
9
14
12
7
10
124
17
11
12
136
31
9
14
12
8
9
124
23
21
12
12
146
37
9
13
11
7
10
11
12
63
63
62
44
44
36
44
37
44
47
52
155
34
9
13
11
7
11
11
12
62
142
32
9
13
12
7
11
128
46
44
46
43
45
45
45
42
44
44
48
43
108
M Sex
12
2
11
2
3
2
3
2
F
F
F
F
F
M
M
11
2
0
2
42
2
7
2
2
M
M
M
48
7
2
M
1
1
1
1
1
1
1
1
1
1
1
1
14
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
953416
953417
953418
953419
953420
953421
953422
953423
953424
953425
942000
942001
942002
942009
AIder Creek (Hoh River)
6/19/95
125
23
ii
12
AlderCreek(HohRiver)
6/19195
120
18
ii
11
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
Alder Creek (Hoh River)
6/19/95
128
19
11
12
6/19195
129
22
11
11
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
6/19/95
5/12/94
5/12/94
5/12/94
5/13/94
5/13/94
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
121
17
11
12
123
21
11
12
60
62
60
60
60
62
128
23
10
11
11
61
11
11
86
11
11
21
12
12
163
ii
12
143
12
ii
166
12
11
128
11
11
62
62
60
62
62
62
62
942010k'
953000
953001
953002
953003
953004
953005
953006
953007
BigBeefCreek
Big Beef Creek
Big Beef Creek
BigBeefCreek
Big Beef Creek
Big Beef Creek
Big Beef Creek
Big Beef Creek
Big Beef Creek
BigBeefCreek
BigBeefCreek
BigBeefCreek
Big Beef Creek
98
193
132
128
13
12
119
156
150
60
30
30
11
11
11
11
12
11
172
41
11
11
137
21
12
12
156
170
32
11
41
12
il
ii
144
26
12
10
G
H
I
9
13
11
6
10
9
13
12
7
10
35
9
13
12
7
12
34
35
9
13
12
6
11
9
13
12
8
11
44
42
32
5
2
32
36
33
9
14
11
6
10
4
2
9
13
11
6
10
7
2
9
13
11
7
10
35
9
14
11
7
10
36
9
13
12
7
11
35
9
13
11
'7
11
9
13
11
6
11
151
32
30
9
13
10
7
10
138
31
9
13
11
6
10
152
9
13
12
6
11
9
14
11
7
11
9
14
12
7
11
9
13
11
6
11
9
13
11
7
10
137
32
32
34
30
30
28
38
50
42
34
43
43
41
38
46
48
33
29
9
13
10
6
11
41
36
43
44
1
D
E
F
155
37
140
146
31
151
154
139
154
149
157
164
148
144
J
62
62
62
61
63
62
139
132
38
9
15
11
7
12
61
141
32
9
12
12
7
10
62
141
31
8
13
10
6
10
141
157
132
L
M
Sex
26
6
2
38
6
2
6
2
11
2
M
M
M
M
M
K
42
48
7
2
M
M
M
M
M
14
2
M
12
2
7
2
F
M
7
2
5
2
7
2
4
2
3
2
0
2
5
2
5
2
11
2
F
2
M
3
2
5
2
M
F
M
F
F
F
M
F
00
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
3
K
L
M Sex
953008
953009
953010
953011
953012
953013
953014
953200
BigBeefCreek
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
4/19/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
149
27
32
33
26
11
11
61
131
31
9
13
11
43
4
2
11
61
150
31
10
14
10
6
6
11
11
10
37
5
2
11
11
161
14
11
7
11
2
131
9
14
10
7
11
10
2
11
11
61
131
31
9
14
11
6
11
11
2
162
28
45
43
42
42
13
11
36
30
9
11
62
62
11
11
63
149
30
9
14
11
6
11
40
1
2
169
41
11
11
62
138
31
9
13
12
7
11
41
15
2
92
110
104
7
11
12
61
145
33
9
13
12
7
11
2
11
12
61
14
12
7
10
11
1
12
13
61
9
13
12
6
13
151
31
11
ii
9
13
12
6
10
114
14
10
12
63
62
9
13
12
6
10
121
18
11
11
61
153
9
13
12
7
10
102
12
11
12
62
9
13
12
6
11
38
24
2
117
17
11
12
61
9
14
13
6
11
2
12
11
12
61
9
13
12
7
11
2
105
11
12
11
9
13
13
7
10
15
2
2
96
96
10
11
12
9
13
12
6
10
7
2
9
12
12
62
62
62
9
14
12
6
105
13
11
12
63
9
13
13
7
12
10
169
109
83
43
11
11
61
9
13
12
ii
12
62
37
9
13
12
7
6
10
16
147
144
148
148
7
12
11
62
144
32
9
15
12
7
12
36
40
42
40
34
32
38
40
30
2
100
146
137
144
30
33
34
37
34
30
33
35
30
37
34
36
37
9
10
146
149
153
40
42
40
36
44
42
13
14
M
F
F
M
F
F
M
F
F
95320.1
953202
953203
953204
953205
953206
953207
953208
953209
953210
953211
953212
953213
953214
953215
Big Beef Creek
BigBeefCreek
BigBeefCreek
BigBeefCreek
Big Beef Creek
Big Beef Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
StevensCreek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
StevensCreek
StevensCreek
Stevens Creek
135
157
143
145
161
143
143
ii
2
2
F
18
2
10
2
F
F
17
2
5
2
10
2
7
2
10
2
7
1
M
F
F
F
M
F
M
F
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
953216
953217
953218
953219
953220
953221
953222
953223
953224
953314
953315
953316
953317
953318
953319
953320
953321
953322
953323
953324
953325
953326
953327
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
Stevens Creek
StevensCreek
Stevens Creek
StevensCreek
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
7/12/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
162
45
82
32
39
21
11
12
62
150
35
9
13
13
7
11
11
11
61
33
9
14
11
6
10
11
12
31
9
13
12
12
37
9
14
12
7
6
11
12
48
40
46
50
11
12
15
12
7
12
11
12
9
13
14
7
12
133
24
12
11
32
36
36
10
16
9
13
12
7
11
98
11
11
12
62
62
62
62
62
62
150
139
153
143
144
157
148
36
9
13
12
6
11
210
94
11
12
62
32
9
13
12
7
11
122
19
11
12
31
9
14
1!
6
10
10
2
66
3
11
12
62
60
166
143
139
9
13
10
6
11
1
1
133
22
32
12
11
61
142
9
14
12
7
12
11
138
33
36
32
29
35
9
14
11
7
9
12
10
9
14
31
9
33
34
30
34
35
36
W.Fk.McClane Creek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McC!ane Creek
W.Fk.McClaneCreek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
197
145
160
130
115
158
96
8
11
11
118
16
12
11
130
96
108
92
164
19
11
12
62
60
60
60
9
11
11
61
13
11
11
62
7
11
11
61
40
13
12
111
13
12
12
62
60
110
102
13
11
12
61
10
11
12
61
137
135
139
129
154
133
160
148
155
129
31
L
M Sex
8
2
11
2
M
M
M
6
2
12
2
50
4
2
16
2
12
2
6
2
22
2
10
44
54
44
50
38
40
39
11
2
11
38
13
6
10
35
3
2
2
12
7
11
38
8
2
13
11
7
11
11
2
9
13
13
7
11
38
34
4
2
9
14
11
7
11
7
2
10
13
10
6
42
40
3
2
9
13
12
8
11
6
2
9
13
11
6
11
38
41
0
2
9
14
12
7
11
38
8
2
M
M
F
F
M
F
F
F
M
M
F
9
14
11
7
11
34
4
2
F
10
F
M
M
F
M
F
M
00
UI
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
953328
953329
953330
953331
953332
953333
953334
953335
953336
953337
953338
953339
953340
953341
953342
953343
953344
953345
953225
953226
953227
953228
953229
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClane Creek
W.Fk.McClaneCreek
W.Fk.McClane Creek
W.FkMcClaneCreek
W.Fk.McClaneCreek
W.Fk.McC!ane Creek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClane Creek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McClaneCreek
W.Fk.McC!ane Creek
W.Fk.McCIa.ne Creek
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
8/1/95
160
37
12
12
61
139
9
13
10
7
11
2
11
12
12
130
9
13
11
5
11
3
2
84
122
112
112
6
11
12
62
60
9
13
12
2
11
12
61
9
14
11
6
6
6
17
11
13
14
12
11
31
9
14
11
7
11
2
2
14
12
12
102
11
11
11
62
64
62
42
32
36
48
38
18
102
34
28
28
33
133
21
12
11
100
107
17
11
13
104
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
129
142
129
134
10
12
M
F
F
F
M
30
9
14
12
7
12
38
3
1
138
31
9
J3
11
7
11
19
2
61
133
31
9
13
12
3
2
61
31
9
13
10
8
2
11
12
34
9
13
11
11
3
2
10
11
11
61
63
130
137
144
6
6
6
10
11
31
9
13
11
6
11
9
2
106
12
11
11
60
31
9
13
13
6
11
14
2
97
105
10
12
12
61
30
10
14
10
6
11
4
2
F
M
F
M
F
10
11
12
61
31
9
13
12
7
10
4
2
M
91
8
12
10
62
13
12
7
12
8
2
11
12
12
63
34
30
9
117
132
133
149
136
152
9
13
11
7
11
4
2
102
10
13
12
61
131
31
9
13
12
7
11
811/95
107
11
12
11
61
140
31
9
13
12
6
11
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
134
126
21
12
12
149
13
13
7
10
11
11
9
13
12
5
10
131
23
12
13
9
13
11
7
10
F
M
M
F
F
M
F
75
118
4
11
12
63
132
8
13
11
6
10
19
12
12
61
151
34
34
37
38
35
9
19
62
62
62
9
14
11
5
10
36
34
38
38
44
38
38
38
34
34
34
36
33
33
36
34
37
142
149
10
8
2
9
2
4
2
3
2
2
2
6
1
2
2
M
M
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
953230
953231
953232
953233
953234
953235
953236
953237
953238
953239
953240
95324 P
953242
AIder Creek (Naselle River)
Alder Creek (Naselle River)
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
110
157
16
12
12
37
9
13
11
12
12
167
31
9
14
12
145
28
11
12
62
62
62
147
35
174
9
15
112
14
11
12
61
153
9
122
129
18
10
12
61
149
9
19
11
12
62
143
36
32
34
30
9
155
39
11
12
61
137
31
9
123
21
12
11
61
149
114
18
12
12
61
141
121
17
11
12
61
136
206
74
84
11
11
62
171
5
10
12
61
132
129
156
132
22
11
11
61
18
11
12
62
139
166
149
32
36
30
36
37
37
12
11
63
23
11
11
219
109
11
117
14
115
953243
953244
953245
953246
953247
953248
953249
953250b
942396
942397
AlderCreek(NaselleRiver)
AlderCreek(NaselleRiver)
AIder Creek (Naselle River)
Alder Creek (Naselle River)
AIder Creek (Naselle River)
AIder Creek (Naselle River)
AlderCreek(NaselleRiver)
AIder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
AlderCreek(NaselleRiver)
AlderCreek(NaselleRiver)
AlderCreek(NaselleRiver)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Alder Creek (Naselle River)
Foley Creek
Foley Creek
7/ 13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
7/13/95
10/19/94
10/19/94
J
K
L
M Sex
7
11
7
22
F
11
2
12
7
11
35
36
37
1
7
8
2
14
10
6
11
41
10
2
13
12
7
11
36
2
2
13
11
6
10
35
3
2
15
13
4
10
4
2
9
13
12
7
10
32
32
M
M
F
F
M
M
10
2
9
13
12
7
10
31
14
2
10
14
10
6
11
34
6
1
9
13
11
7
11
11
2
9
14
12
6
10
9
13
12
'7
10
19
2
31
9
13
11
7
11
32
36
3
2
151
33
9
13
11
6
10
31
7
2
164
14
12
6
10
39
14
2
156
36
34
9
12
62
63
9
14
11
6
10
31
15
2
11
12
63
165
31
10
15
12
7
12
36
13
1
F
F
15
12
12
63
165
3
10
11
134
71
4
12
13
11
12
11
11
62
64
62
63
34
34
25
33
35
M
70
116
110
117
146
162
F
F
M
F
1
M
F
M
M
9
13
12
7
10
38
6
2
10
14
11
6
11
19
8
1
10
13
0
1
9
9
13
12
7
11
13
2
F
13
11
7
10
50
39
22
2
M
6
Appendix Table A2.1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
942398
942399
942400
942401
942402
942403
942404
942405
942406
942407
942408
942409
942410
942411
942412
942413
Foley Creek
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
942414
942415
942416
942417
942418
942419
942420
FoleyCreek
Foley Creek
FoleyCreek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
FoleyCreek
Foley Creek
FoleyCreek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
10119/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
10/19/94
A
B
C
D
120
12
12
61
156
110
11
12
61
143
94
110
108
117
110
125
138
133
128
149
11
12
11
12
62
60
138
146
11
12
61
151
13
12
62
144
11
11
61
140
11
12
61
12
12
12
11
11
11
11
155
E
F
G
H
I
J
K
L
33
9
13
12
7
11
2
9
14
11
8
10
14
2
33
9
14
12
7
11
33
9
14
11
7
11
9
13
12
10
9
14
11
7
7
9
14
12
7
10
150
32
33
34
35
9
13
11
7
12
61
151
37
9
13
11
6
11
41
48
29
44
30
43
39
44
45
29
33
61
153
34
10
14
12
7
10
141
35
9
13
12
7
11
12
60
60
159
36
9
15
12
6
11
12
61
171
37
9
14
11
7
165
11
12
61
161
31
9
14
12
80
12
12
61
161
37
9
13
12
53
106
10
12
61
9
14
12
33
9
13
14
7
11
120
65
66
11
11
37
9
14
12
12
9
14
11
7
6
12
12
11
12
131
14
12
6
12
13
12
7
10
11
13
9
9
13
79
34
36
34
9
81
14
10
7
71
11
11
63
59
64
62
62
61
62
126
147
147
127
36
12
35
9
14
12
7
141
150
150
M Sex
17
2
33
25
2
M
11
46
46
47
F
M
M
F
M
F
F
F
F
F
2
12
63
21
2
M
M
7
11
13
2
7
11
33
38
37
47
45
10
1
12
11
10
10
22
1
6
2
20
23
2
2
13
2
32
2
18
2
1
19
2
14
2
35
10
1
15
1
6
2
11
32
43
39
5
2
11
38
9
1
F
F
F
M
M
F
F
M
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
942421
942422
942423
942424
942425
931339
931340
931341h
Foley Creek
Foley Creek
Foley Creek
Foley Creek
Foley Creek
10/19/94
10/19/94
10/19/94
94
114
109
70
129
224
258
176
219
239
253
931342
931343
931344
931345
931346
931347
931348
953293
953294
953295
953296
953297
953298
953299
953300
N.Fk.TraskRiver
N.Fk.TraskRiver
N. Fk. Trask River
N. Fk. Trask River
N. Fk. Trask River
N. Fk. Trask River
N. Fk. TraskRiver
N.Fk.TraskRiver
N.Fk.TraskRiver
N.Fk.TraskRiver
N.Fk.TraskRiver
N. Fk. Trask River
N.Fk.TraskRiver
N.Fk.TraskRiver
N. Fk. Trask River
N. Fk. Trask River
N.Fk.TraskRiver
N. Fk. Trask River
10119194
10/19/94
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
9/10/93
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
11
13
61
161
9
14
13
7
12
2
12
63
149
9
15
13
7
12
44
44
6
12
14
2
12
12
154
9
14
12
6
10
41
12
9
14
11
6
10
1
M
F
M
M
12
12
61
169
9
13
12
6
13
37
41
20
4
2
12
63
64
7
2
M
104
12
11
61
163
10
14
13
12
159
9
14
11
57
12
12
11
14
12
7
12
0
1
M
M
M
103
10
11
9
14
12
7
10
10
2
M
125
11
11
142
10
14
12
7
11
12
2
162
11
11
62
64
62
59
62
22
20
2
11
7
7
10
166
36
36
33
34
34
34
34
26
34
32
141
35
9
14
12
7
Ii
7
2
30
167
105
116
52
67
330
59
98
11
11
61
142
31
10
14
12
7
10
6
2
11
12
135
10
14
11
10
2
11
10
15
11
6
7
12
12
10
18
2
11
12
22
2
12
13
11
2
12
13
11
12
ii
11
60
62
63
60
62
62
59
11
12
163
11
Ii
137
24
12
13
260
168
12
12
151
241
223
230
173
195
325
183
216
250
151
122
144
10
145
33
35
162
38
9
13
11
6
11
134
33
10
13
12
7
11
161
33
9
14
9
6
11
148
34
9
14
13
7
11
160
36
9
13
12
7
10
61
158
37
9
13
12
8
11
62
62
62
166
154
177
38
9
14
11
6
10
35
10
15
12
7
11
36
9
14
12
7
11
42
40
52
36
52
42
60
29
2
7
2
23
2
17
2
6
2
15
2
10
2
6
2
F
M
F
M
F
F
M
M
F
M
F
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
953301
953302
953303
953304
953305
953306
953307
953308
953309
953310
953311
953312
953313
953047
953048
953049
953050
953051
953052
953053
953054
953055
953056
N. Fk. TraskRiver
140
27
12
11
60
32
9
14
11
6
11
81
11
12
63
38
10
15
12
6
10
12
61
151
14
12
7
11
12
13
62
184
9
14
11
7
12
N. Fk. Trask River
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
7/25/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
175
50
11
11
61
167
9
14
11
6
10
122
18
11
12
61
145
9
14
11
6
11
40
52
42
54
32
40
139
27
10
13
9
15
11
6
10
51
12
13
210
234
12
12
11
12
61
11
12
12
11
62
63
10
11
61
12
12
ii
11
139
94
134
64
88
30
90
30
27
62
60
62
131
172
33
36
34
33
34
32
9
7/25195
116
119
11
N.Fk.TraskRiver
207
224
226
136
154
N. Fk. TraskRiver
7/25/95
7/25/95
7/25/95
11
13
178
51
12
13
62
62
62
62
120
104
18
11
12
61
11
11
12
61
101
11
ii
12
61
105
11
11
12
60
104
11
11
11
61
150
152
154
140
136
154
149
138
142
125
119
17
11
12
61
151
N.Fk.TraskRiver
N.Fk.TraskRiver
N.Fk.TraskRiver
N. Fk. Trask River
N. Fk. TraskRiver
N.Fk.TraskRiver
N. Fk. Trask River
N.Fk.TraskRiver
N.Fk.TraskRiver
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
188
201
145
201
145
156
153
143
147
167
58
42
34
9
14
12
7
10
33
9
14
11
6
11
35
36
34
37
30
9
14
11
6
11
9
14
11
7
10
9
14
12
7
12
9
13
11
7
11
42
54
42
9
14
ii
6
11
35
9
14
12
7
30
9
14
13
7
30
30
9
13
10
9
14
ii
32
9
14
36
32
9
L
M Sex
M
F
F
8
2
12
2
5
10
11
2
2
2
8
1
F
F
13
2
M
F
M
8
2
15
2
6
20
2
10
2
8
2
38
12
2
11
54
13
2
11
38
17
2
7
11
1
11
23
1
12
6
12
40
30
48
16
7
19
2
14
12
8
12
8
1
9
13
11
6
12
34
8
13
11
6
32
9
14
13
7
2
13
2
12
48
36
38
12
2
11
46
14
2
F
F
F
M
M
F
F
F
F
F
M
M
M
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
953057
953058
953059
953060
953061
953062
953063
953064
953065
953066
953067
931214
931613
9423491
Andy Creek
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
5/23/95
7/14/93
7/14/93
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
132
23
11
11
61
143
9
14
12
7
10
119
17
11
12
61
149
9
14
13
8
10
149
154
9
14
12
7
12
9
14
12
8
12
9
14
11
7
11
942350
9423511
942352
942353
942354
9423551
942356
942357
942358b
AndyCreek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
Andy Creek
SchoonerCreek
Schooner Creek
SchoonerCreek
SchoonerCreek
SchoonerCreek
SchoonerCreek
Schooner Creek
Schooner Creek
Schooner Creek
SchoonerCreek
Schooner Creek
Schooner Creek
11
11
11
61
12
12
13
ii
62
56
12
12
61
137
9
14
11
6
10
12
13
37
9
15
11
7
12
12
12
62
62
148
132
22
32
82
28
32
30
32
30
32
35
30
157
33
9
13
12
8
11
128
21
11
13
61
145
10
14
12
8
11
134
25
24
12
12
61
9
14
13
7
11
11
11
61
149
147
34
32
32
9
14
11
7
11
10
11
11
3
11
11
60
62
100
132
131
201
142
136
99
62
141
K
L
46 8
44 8
34 17
42 16
38 17
32 19
52 25
46 26
48 18
40 20
48 13
M Sex
2
F
1
M
M
M
2
2
2
2
2
2
2
2
2
61
3
11
12
61
145
11
12
11
61
152
62
3
12
11
62
154
51
1
11
53
2
ii
13
62
56
57
92
2
12
11
61
149
2
10
11
62
158
9
14
8
11
12
61
152
9
14
11
7
10
103
14
11
12
155
9
14
12
7
10
109
15
11
13
60
64
10
15
12
7
11
34
61
137
32
35
27
9
14
9
15
9
14
8
14
9
14
10
15
12
8
11
37
7
1
10
46
43
0
2
10
40
12
6
1
M
M
F
1
0
47
47
32
37
M
M
F
F
F
51
100
F
F
F
F
2
8
2
7
2
0
1
M
M
M
M
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
942359
942360
Schooner Creek
Schooner Creek
Schooner Creek
Schooner Creek
Schooner Creek
Schooner Creek
Schooner Creek
Siletz River
Siletz River
Siletz River
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
7/7/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
168
12
12
61
147
128
53
22
12
12
62
143
124
23
12
12
65
135
60
52
3
11
11
135
2
11
11
63
62
95
102
9
11
12
11
12
363
942361h
942362
942363
942364
942365
942230
942231
942232
942233
942234
942235
942236
942237
942238
942239
942240
942241
942242
942243
942244
942245
SiletzRiver
Siletz River
SiletzRiver
Siletz River
Siletz River
Siletz River
SiletzRiver
Siletz River
Siletz River
Siletz River
SiletzRiver
Sjletz River
Siletz River
10121/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
345
350
200
170
315
245
260
248
325
193
200
345
314
356
205
233
F
G
H
I
J
K
L
32
9
13
13
7
10
2
10
14
11
7
10
3
2
28
10
15
12
7
10
48
40
52
14
32
0
1
30
10
15
11
6
12
32
0
2
F
F
M
M
125
31
9
13
M
155
33
10
15
12
7
11
2
11
61
156
32
9
14
13
7
11
45
35
0
4
2
61
5
2
12
12
9
13
11
7
11
41
16
2
11
11
138
146
31
440
69
46
294
14
11
7
11
41
8
2
12
9
14
12
7
10
11
12
9
13
12
6
10
11
11
112
148
144
13
13
61
144
147
12
12
62
141
12
11
61
144
300
62
65
419
270
12
12
149
11
11
11
12
168
11
12
145
11
11
133
451
72
130
11
11
63
61
61
62
62
60
36
34
34
30
32
33
34
37
35
38
10
12
63
62
62
60
62
F
M
F
F
11
12
61
140
140
11
11
61
131
43
40
47
38
43
39
45
45
50
42
43
38
39
34
148
133
147
E
12
9
14
12
7
10
9
14
12
7
12
9
14
11
5
10
9
14
12
7
11
10
14
13
7
9
10
14
12
6
ii
8
14
14
7
11
31
9
13
12
7
11
37
34
33
34
9
10
13
13
7
10
14
13
8
11
10
14
11
6
11
8
14
13
6
11
M Sex
18
2
32
2
F
M
M
M
M
F
F
M
16
2
14
2
21
2
9
2
35
25
2
11
2
20
20
2
F
2
2
7
2
18
2
M
F
F
17
2
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
942246
942247
942248
942249
942250
942251
942366
942367
942368
942369
942370
942371
942372
942373
942374
942375
942376
942377
942378
942379
942380
942381
942382
Siletz River
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/21/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
239
115
11
11
62
138
31
9
14
12
7
11
34
21
188
62
418
126
170
189
11
11
61
143
9
14
12
7
12
12
l2
61
148
9
14
12
6
Ii
11
11
9
15
11
11
11
62
62
148
11
10
11
11
61
145
11
12
61
155
12
13
60
137
95
11
11
61
132
36
29
32
34
31
34
33
30
95
11
10
117
ii
11
136
159
73
11
12
59
62
60
96
115
11
11
61
10
12
85
11
103
SiletzRiver
SiletzRiver
SiletzRiver
Siletz River
Siletz River
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
lO/5194
10/5/94
M Sex
2
8
14
11
9
14
12
6
6
7
9
13
11
7
10
39 20
40 10
42 28
34 11
35 26
38 29
10
14
12
6
11
42
19
1
9
14
12
7
10
41
24
2
31
9
13
11
8
11
13
2
31
9
14
12
6
11
16
2
133
32
9
14
13
7
10
17
2
130
31
9
14
12
6
10
42
42
43
46
5
1
61
149
31
9
13
11
7
12
51
19
2
11
62
138
9
13
11
5
11
41
11
2
12
12
61
147
9
14
12
6
11
40
11
1
101
11
10
61
140
9
13
10
38
19
2
12
12
61
9
14
11
10
2
11
11
9
15
13
7
11
11
1
11
11
135
31
9
14
12
7
10
12
1
11
12
62
62
60
149
31
9
14
12
ii
5
13
12
61
135
9
14
11
6
12
11
12
61
142
35
32
9
14
12
6
11
39
42
41
43
43
47
8
99
94
100
112
96
156
135
7
6
11
108
29
35
33
33
30
339
229
251
274
94
96
148
12
30
23
8
2
2
2
2
2
1
1
2
2
M
M
F
M
F
F
M
F
F
M
M
M
F
M
F
M
M
F
F
M
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
942383
942384
942385
942386
942387
942388
942389
942390
942391
942392
942393
942394
942395
920317
920318
920319
920320
920321
920322
920323
920324
920325
920327
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
10/5/94
1015/94
WolfCreek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Wolf Creek
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
10/5/94
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
105
11
11
62
149
38
9
15
13
6
11
27
17
2
89
11
12
61
132
9
14
11
8
10
2
M
F
9
14
11
7
10
7
9
14
12
7
11
49
44
47
17
136
138
147
157
169
145
146
135
32
34
30
32
35
35
33
30
29
2
2
M
9
13
6
7
11
46
24
12
16
2
2
19
2
127
110
10
12
61
111
11
12
124
11
12
145
11
12
62
60
62
154
12
12
61
155
12
12
63
103
10
12
96
12
11
105
10
12
62
62
60
117
12
12
61
134
ii
12
11
4
F
9
13
10
12
10
15
12
6
12
9
14
12
7
11
43
41
40
39
2
9
14
12
6
12
41
21
1
10
15
11
6
11
15
1
31
8
12
10
7
11
3
2
F
141
31
9
14
11
5
11
14
131
9
14
11
6
10
21
11
62
131
9
10
11
7
10
4
2
11
11
61
137
9
13
12
7
10
7
2
10
10
12
11
7
10
8
11
11
6
10
26
12
2
12
1931
10
10
9
13
11
10
10
63
9
11
11
6
6
10
1653
2503
2505
1264
1375
11
11
63
33
8
12
12
7
10
11
11
61
130
148
133
142
138
145
9
12
60
60
62
30
32
35
30
35
34
30
2
2
F
61
44
42
40
52
M
M
M
F
M
F
8
11
12
6
10
11
11
11
7
Il
62
62
9
Ii
32
34
32
9
10
7
2005
1599
1430
1190
133
135
36
52
36
32
34
F
F
M
13
2
24
22
2
10
8
2
M
F
M
F
F
F
M
10
19
2
M
10
44
9
2
2
2
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
376
516
530
496
593
682
484
474
370
439
530
442
471
552
349
455
289
523
394
363
11
13
140
37
10
14
12
7
12
12
147
15
12
12
10
13
11
12
12
11
12
11
12
155
32
35
34
36
36
9
12
11
12
146
35
11
12
63
62
63
62
63
63
63
62
157
37
10
11
12
61
170
41
12
12
151
41
13
12
137
11
12
62
62
60
10
11
61
152
11
11
159
10
11
163
11
12
153
37
12
12
151
11
11
10
12
11
11
63
63
62
63
62
63
63
36
36
35
36
36
114
11
12
61
146
11
11
61
136
150
120
11
12
62
138
number
Location
collection
920328
920329
920330
920331
920332
920333
920334
920335
920336
920337
920338
920339
920340
920341
920342
920343
920344
920345
920346
920347
953591
953592
953593
Alsea Hatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
Alsea Hatchery
AlseaHatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
Alsea Hatchery
CumminsCreek
CumminsCreek
CumminsCreek
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
9/4/92
8/7/95
8/7/95
8/7/95
149
143
162
10
K
L
M Sex
10
15
2
7
10
11
7
10
14
2
2
M
M
M
12
6
10
14
2
10
15
12
7
10
16
2
9
14
12
8
10
20
2
11
8
10
17
2
14
12
7
10
16
2
M
M
10
14
11
7
10
18
2
M
10
14
12
7
11
8
2
9
14
11
7
9
15
2
10
14
12
8
10
14
2
10
15
12
6
10
11
2
9
14
11
7
10
11
2
10
14
11
7
10
9
14
12
7
10
37
8
14
11
7
10
160
39
10
13
12
6
10
131
33
12
7
10
154
34
36
37
33
145
M
M
39
45
36
9
13
12
8
11
9
15
Ii
8
11
9
13
11
7
11
42
54
9
14
11
6
11
38
M
M
M
M
M
9
2
10
2
11
2
15
2
2
2
7
2
19
2
16
2
M
M
F
4
2
F
M
M
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
953594
953595
953596
953597
953598
953599
953600
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
CumminsCreek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
CumminsCreek
Cummins Creek
Cummins Creek
Cummins Creek
CumminsCreek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
Cummins Creek
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/7/95
8/4/95
8/7/95
953601
953602
953603
953604
953605
953606
953607
953608
9536091
953610
953611
953612
953613
953614
953615
953616
A
B
C
D
E
F
G
H
I
J
K
195
12
13
62
156
9
14
12
6
11
155
10
11
61
144
35
34
9
14
12
8
11
180
11
12
62
137
31
8
14
11
5
182
86
10
11
61
142
9
15
12
11
12
62
138
36
33
13
11
161
11
11
63
156
33
9
13
124
10
11
148
37
8
14
132
11
12
137
33
112
114
12
12
155
11
12
62
62
63
63
34
39
98
10
11
61
146
74
76
11
12
62
137
11
12
61
138
114
11
12
63
150
127
11
12
62
152
34
37
34
35
34
119
12
11
83
11
12
62
60
159
145
75
59
173
194
206
12
11
61
135
12
13
61
152
11
11
62
11
11
61
151
L
M Sex
50
17
2
38
21
2
11
37
13
2
7
11
15
2
8
12
12
2
12
6
11
12
7
11
9
14
11
6
11
9
14
12
8
11
9
13
11
7
12
9
15
12
7
11
50
38
42
50
48
42
29
42
9
13
12
7
10
9
14
12
7
11
10
14
11
7
11
10
14
11
7
11
39
34
38
9
15
11
6
11
12
6
10
9
14
12
6
11
36
38
10
15
12
5
10
168
9
13
12
10
151
33
9
14
10
6
6
44
38
9
2
12
2
12
2
10
2
6
2
10
1
3
1
F
M
F
F
M
F
F
M
F
F
F
1
1
11
1
F
4
2
F
M
M
F
80
52
9
10
48
42
42
48
34
40
3
2
2
1
7
1
42
11
2
2
13
2
M
F
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
953617
931254
931255
931256
931257
931258
931261
931262
931264
Cummins Creek
W. Br. N. Fk. Smith River
W. Br. N. Fk. SmithRiver
W. Br. N. Fk. SmithRiver
8/7/95
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
9/14/93
228
111
11
12
14
11
36
37
9
12
9
14
158
11
12
11
11
38
34
9
212
62
62
63
60
167
111
9
122
132
164
120
12
11
61
38
12
12
61
157
148
11
12
61
13
12
61
162
149
143
101
12
11
61
156
39
35
36
38
12
11
61
150
11
11
154
12
12
161
12
12
123
11
11
171
11
11
137
11
13
128
12
ii
62
62
62
61
60
62
62
158
12
11
113
12
141
93l265
931266
931267
931268
931269
931271
931272
931273
931276
931277
931278
931279
931280
931281
W.Br.N.Fk.SmithRiver
W.Br.N.Fk.SmjthRiver
W. Br. N. Fk. Smith River
W. Br. N. Fk. Smith River
W. Br. N. Fk. SmithRiver
W. Br. N. Fk. Smith River
W. Br. N. Fk. SmithRiver
W. Br. N. Fk. Smith River
W.Br.N.Fk.SmithRiver
W. Br. N. Fk. SmithRiver
W. Br. N. Fk. Smith River
W. Br. N. Fk. Smith River
W. Br. N. Fk. Smith River
W. Br. N. Fk. SmithRiver
W. Br. N. Fk. Smith River
W.Br.N.Fk.SmithRiver
W. Br. N. Fk. SmithRiver
W.Br.N.Fk.SmithRiver
W. Br. N. Fk. Smith River
I
J
K
L
M Sex
10
7
11
46
13
2
12
7
11
9
1
14
11
6
12
14
12
8
12
9
14
12
7
10
9
14
12
7
10
9
13
13
7
11
9
14
12
7
11
9
14
12
7
11
32
33
9
14
11
7
11
9
14
12
7
11
38
9
14
11
7
11
39
39
39
38
42
46
28
39
35
53
45
28
22
33
32
9
14
12
6
10
32
9
14
10
5
11
156
35
9
14
11
6
10
154
34
9
14
13
7
61
170
38
9
14
12
7
11
39
35
46
43
12
62
141
39
9
14
12
7
11
41
11
11
61
158
36
9
14
11
ii
2
11
12
62
155
35
9
15
12
122
12
12
61
164
38
9
14
12
8
11
121
11
12
62
146
38
9
14
11
7
11
42
44
45
39
7
124
6
6
146
161
156
164
152
160
144
179
11
6
2
12
2
9
2
5
2
27
2
12
2
23
2
17
2
14
12
2
2
2
14
2
4
2
8
2
16
2
13
2
8
2
14
2
14
1
M
F
M
F
F
F
F
F
M
M
M
M
F
M
M
M
F
F
F
M
F
M
F
Appendix Table A2. 1. Continued
Fish
number
942198
942199
942200
942201
942202
942203
942204
942205
942206
942207
942208
942209
942210
942211
942212
942213
942214
942215
942216
94221T'
942218
942219
942220
Date of
FL
Wt.
Location
collection
(mm)
(g)
A
B
C
D
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
77
78
5
11
11
61
5
11
12
61
128
20
4
4
11
11
61
139
140
144
11
11
12
13
DavisCreek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
DavjsCreek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
10120194
10/20/94
10/20/94
10/20/94
74
82
109
147
11
11
10
62
62
63
30
12
12
61
123
16
12
12
62
66
74
70
3
11
11
61
4
11
11
61
3
12
13
130
21
11
11
59
74
79
2
12
12
4
11
11
62
60
62
60
6
11
12
61
E
F
G
H
13
13
166
132
140
157
156
120
30 8
37 9
37 9
34 9
33
9
26 9
35
9
33 9
32 9
33 9
30 10
33 9
32 9
133
31
138
130
132
166
I
I
K
11
7
11
12
5
11
15
1
5
1
L
M Sex
13
11
7
12
12
11
7
11
13
12
6
12
53
52
48
43
43
4
1
14
11
7
12
37
7
2
M
M
M
F
F
M
14
12
7
11
51
17
2
M
13
12
8
10
14
2
13
11
8
10
12
7
11
2
7
1
13
2
14
12
6
11
12
1
13
12
6
11
13
2
F
F
F
M
M
13
11
7
12
9
13
12
6
11
52
40
46
40
52
39
50
6
1
33
33
8
13
12
7
10
41
15
2
9
14
12
7
10
55
53
45
42
28
46
44
45
91
7
11
12
60
139
149
76
80
5
12
11
61
143
31
9
13
12
6
10
5
11
10
63
130
33
9
13
11
7
12
85
6
12
Ii
141
33
10
14
12
7
11
65
66
3
11
1
60
63
9
'i3
ii
7
12
3
11
Ii
61
126
32
30
9
13
11
6
11
85
6
12
11
62
141
35
9
13
12
7
11
77
5
10
11
61
135
31
9
13
11
6
11
1
2
1
6
1
11
1
12
1
6
2
4
1
9
1
6
1
4
1
M
M
F
F
F
F
F
F
M
M
I
Appendix Table A2. 1. Continued
Fish
Date of
FL
number
Location
collection
(mm)
94222P
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Davis Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
10/20/94
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
74
72
72
72
63
73
69
71
69
97
130
91
9422221
942223
942224
942225
942226
942227
942228
942229
953251
953252
953253
953254
953255
953256
953257
953258
953259
953260
953261
953262
953263
953264
wt.
A
B
C
D
E
F
G HI
J
K
133
30
9
12
6
11
9
12
7
10
46
42
37
52
29
32
34
4
4
11
11
61
11
12
61
10
1
5
1
M
4
2
F
6
2
M
5
2
11
48
46
50
47
45
48
F
F
F
F
M
F
11
38
38
3
1
7
2
6
1
M
M
F
F
8
2
M
9
1
F
5
11
11
63
129
32
9
12
12
8
12
4
11
11
60
137
31
9
14
12
6
11
3
11
12
61
127
31
9
13
11
6
4
11
12
60
132
13
11
7
10
1 1
LI
1 1
II
L'
131
30
30
9
3
9
12
12
6
11
4
11
11
61
134
31
9
13
11
6
10
3
11
12
9
13
10
7
11
11
10
30
9
14
11
7
11
21
11
12
61
130
132
144
159
147
138
35
10
62
60
35
9
14
12
8
11
33
9
14
12
11
31
9
14
12
7
6
31
9
15
12
8
13
162
9
15
13
7
11
9
13
12
7
11
9
14
12
7
9
14
11
7
9
14
12
7
9
14
12
6
11
12
7
11
43
38
56
9
11
11
60
13
12
12
61
85
7
12
11
141
29
20
9
11
12
10
12
12
12
11
11
11
60
60
60
60
60
101
12
ii
12
61
146
111
14
11
12
154
103
13
11
11
133
33
8
14
123
20
12
12
153
9
14
12
6
11
12
11
11
32
34
40
36
48
102
60
60
60
60
34
34
30
30
33
34
9
14
13
7
11
38
98
99
153
150
136
139
M Sex
F
F
M
M
M
M
F
F
105
126
L
11
1
4
1
1
1
11
2
4
1
5
2
5
1
4
1
3
2
8
2
8
2
8
1
4
1
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
I
K
953265
953266
953267
953268
953269
953270
953271
953272
953273
953274
953275
953276
953277
953278
953279
953280
953281
953282
953283
942196
942197
953040
953041
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
7/20/95
10/19/94
10/19/94
5/11/95
5/11/95
90
92
98
89
109
104
100
8
11
11
61
141
14
11
6
12
41
7
1
12
12
61
144
9
15
12
7
12
40
19
1
11
11
12
137
9
14
11
5
12
38
7
2
10
11
11
60
60
139
30
32
32
34
9
11
9
14
12
7
12
4
1
13
12
12
61
144
31
9
14
12
7
11
4
2
13
10
12
60
143
35
9
15
13
8
12
40
50
44
6
2
11
12
11
61
138
31
9
14
12
7
1!
38
6
1
111
17
11
12
60
140
30
9
15
11
7
11
35
6
2
109
112
13
12
12
61
31
9
14
10
7
11
1
12
ii
60
32
9
15
12
16
11
11
61
29
9
14
12
101
12
11
11
31
10
14
11
8
11
112
122
129
91
92
95
95
196
16
12
11
60
60
146
136
140
7
6
11
114
35
9
14
12
7
10
23
12
12
61
141
35
9
14
11
7
12
23
11
Ii
61
162
14
12
7
11
11
12
60
130
9
14
10
6
11
7
12
12
61
161
9
15
11
7
11
9
11
12
61
150
34
29
30
34
9
8
9
14
12
7
12
10
11
11
159
33
9
14
11
7
11
72
11
12
60
60
148
33
9
14
12
7
13
11
12
61
161
31
9
15
12
7
12
145
98
27
34
48
38
38
44
42
48
32
44
48
50
38
35
3
15
142
155
11
12
61
14
12
7
12
42
18
11
11
60
34
36
9
122
144
146
9
14
11
7
11
38
6
1
IronCreek
Iron Creek
IronCreek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Iron Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
209
12
L
M Sex
5
2
16
1
4
2
8
2
8
2
4
1
6
2
10
2
5
1
6
2
8
2
5
2
6
2
M
F
F
F
F
F
F
M
M
F
F
F
F
F
F
F
F
M
F
M
F
F
F
Appendix Table A2. 1. Continued
Fish
Date of
FL
C
D
E
F
G
HI
J
K
147
34
9
14
12
7
12
139
35
9
14
12
7
11
143
33
9
14
12
7
13
157
35
9
13
12
7
12
9
14
12
8
11
14
12
7
11
12
61
9
14
12
5
12
21
12
13
62
9
14
14
7
11
120
97
17
11
12
61
151
9
14
12
7
13
9
ii
12
140
9
15
12
6
12
136
23
9
10
9
14
11
6
12
117
16
11
12
10
14
12
6
11
130
20
12
12
62
62
59
60
32
32
32
36
32
33
35
32
10
11
136
144
132
144
31
12
60
60
62
60
62
60
9
14
13
8
13
258
155
12
11
61
31
8
13
11
6
12
151
35
10
12
61
159
158
36
9
15
12
7
12
46
45
45
43
48
40
46
44
46
42
48
40
46
44
42
8
1
147
33
12
12
61
133
34
9
14
12
7
12
41
6
2
163
143
39
11
12
61
142
9
14
12
8
12
42
9
2
25
11
12
61
151
32
34
9
14
12
6
13
7
140
167
33
36
9
14
11
7
13
9
14
11
7
12
36
8
14
12
7
12
35
9
14
10
6
12
28
9
15
12
7
13
45
42
45
43
36
52
number
Location
collection
(mm)
953042
953043
953044
953045
953046
953068
953069
953070
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Wheeler Creek
Jordon Creek
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/30/95
5/30/95
5/30/95
5/30/95
5/30/95
5/30/95
5/30/95
5/30/95
7/21/95
7/21/95
7/21/95
7/21/95
7/21/95
7/21/95
7/21/95
7/21/95
171
953071
953072
953073
953074
953075
953284
953285
953286
953287
953288
953289
953290
953291
953292
942270
7121/95
11/2/94
Wt.
A
B
46
10
12
147
28
10
12
169
148
120
142
42
32
11
12
11
13
17
11
11
28
10
127
21
131
144
149
133
161
42
12
12
59
191
68
11
12
61
146
28
223
11
12
61
12
13
61
150
134
28
12
11
62
142
301
142
L M Sex
13
2
10
2
6
2
6
2
F
F
F
M
1
2
F
12
2
9
2
10
2
M
M
F
11
1
M
5
2
F
12
2
0
2
6
2
M
M
F
19
2
5
2
2
17
2
5
2
8
2
13
1
F
F
M
M
F
M
M
M
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
942271
942272
942273
942274
942275
942276
942277
942278
942279
942280
942281
942282
942283
942284
942285
942286
942287b
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
JordonCreek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
Jordon Creek
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
127
23
11
11
110
15
11
10
145
30
29
35
12
942288
942289
942290
942291
942292
942293
D
F
F
G
61
134
31
9
14
63
145
9
14
12
62
142
9
13
11
11
61
138
9
12
11
61
18
12
11
137
25
11
10
62
62
134
158
36
33
34
30
34
142
31
8
145
29
24
24
11
12
61
32
11
12
61
ii
10
140
148
148
142
132
135
120
135
135
123
18
12
11
62
62
96
9
12
11
61
81
7
12
12
60
129
141
83
89
7
11
11
61
135
7
12
11
62
88
7
11
11
87
107
7
11
13
86
92
82
83
92
H
I
J
K
L
12
6
12
1
7
13
5
1
12
7
12
7
1
14
12
7
13
3
1
9
13
12
7
12
3
2
10
14
12
7
12
2
2
14
12
7
12
5
2
M
M
9
15
11
7
12
2
2
F
35
33
9
9
14
12
7
12
49
45
53
50
49
50
46
42
43
12
12
6
13
12
6
12
61
8
2
2
30
9
14
12
7
13
5
2
M Sex
2
2
4
2
4
1
F
33
9
14
12
6
12
3
2
10
13
12
7
12
49
3
1
9
14
12
7
11
41
7
1
145
34
32
32
9
14
ii
7
12
0
2
61
126
33
9
14
12
7
13
2
1
11
62
144
34
9
14
11
7
13
0
1
11
11
61
132
9
13
12
7
12
7
11
12
61
126
8
13
12
7
12
8
11
11
61
142
9
13
10
6
13
6
Ii
11
61
141
6
12
11
61
143
8
12
11
62
138
35
29
32
34
32
32
39
56
54
42
47
47
42
34
54
9
13
11
7
12
12
11
6
12
9
14
13
7
11
M
F
M
F
M
M
M
F
F
F
F
M
F
F
F
M
58
49
10
M
F
4
1
4
2
5
2
C
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
942294
942295
942296b
JordonCreek
1112/94
97
10
11
12
61
9
14
11/2/94
11/2/94
11/2/94
91
7
11
11
61
14
5
11
12
9
5
11
12
31
9
5111195
107
14
11
12
131
31
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
74
4
11
12
105
12
11
12
62
62
62
60
62
29
29
9
76
80
149
130
133
139
31
Jordon Creek
Jordon Creek
Jordon Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
WaukeliCreek
WaukeliCreek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
99
10
12
11
61
105
161
14
11
11
61
48
11
11
114
16
11
12
29
32
29
30
33
30
95
97
103
93
100
146
10
11
12
60
60
60
129
129
132
137
146
146
131
9
11
11
61
131
14
12
11
133
8
11
11
60
62
9
11
11
32
11
128
21
111
20
24
942297
953015
953016
953017
953018
953019
953020
953021
953022
953023
953024
953025
953026
953027
953028
953029
953030
953031
953032
953033b
125
99
86
80
L
M Sex
I
J
K
12
8
12
11
7
12
13
12
9
ii
13
13
7
12
9
15
11
7
12
9
13
10
7
10
9
13
13
7
12
4
1
9
13
12
7
11
3
1
9
14
11
7
ii
8
13
11
7
12
9
14
13
7
12
46
50
49
39
48
26
32
45
42
44
39
33
9
14
12
8
9
14
12
8
12
9
14
11
7
10
135
30
30
30
9
14
12
7
13
61
140
33
9
14
13
8
12
11
61
132
31
9
14
12
7
11
11
12
61
137
31
9
14
11
7
11
ii
12
61
141
34
9
13
12
7
10
10
11
63
32
9
14
12
7
11
10
12
12
60
156
136
39
45
50
40
29
9
13
11
7
11
8
11
12
61
137
31
9
13
11
7
11
4
11
11
60
128
33
9
13
12
6
10
2
2
3
2
0
4
2
7
2
M
M
M
M
F
4
1
F
1
4
1
1
2
4
1
38
2
2
32
33
39
32
3
1
8
1
7
1
4
1
8
2
F
M
M
M
M
M
M
M
M
F
3
2
5
1
F
M
M
5
1
M
40
3
1
37
3
1
M
F
36
0
1
F
C
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
953034
953035
953036
953037
953038
953039
942322
942323
942324
942325
942326
942327
942328
942329
942330
942331
942332
942333
942334
942335
942336
942337
942338
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
Waukell Creek
May Creek
May Creek
May Creek
May Creek
May Creek
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
5/11/95
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
96
10
11
11
62
91
9
12
12
61
94
95
86
8
10
12
61
11
11
12
30
30
29
29
7
11
Ii
62
62
91
8
11
12
61
130
127
135
123
134
134
124
136
MayCreek
May Creek
May Creek
MayCreek
May Creek
May Creek
MayCreek
May Creek
May Creek
May Creek
May Creek
May Creek
11/3/94
11/3/94
11/3/94
F
G
H
I
J
K
L
M Sex
7
7
11
38
4
2
12
36
3
1
2
2
9
14
12
9
14
11
9
12
11
7
11
9
14
11
7
12
31
9
14
11
7
33
9
14
12
27
28
33
27
27
28
28
26
9
14
9
9
3
1
12
42
40
5
1
8
12
41
3
1
11
6
10
11
2
14
10
6
1!
4
2
13
12
9
10
44
60
50
8
1
9
14
ii
7
11
41
1
2
9
14
11
7
10
36
7
2
9
14
10
42
5
2
14
11
7
6
12
9
9
11
2
13
10
7
11
7
16
2
F
F
F
F
F
F
F
F
M
M
115
15
11
12
61
160
126
120
99
119
113
125
174
120
135
149
38
11
13
19
11
12
17
11
13
62
62
62
10
11
11
61
134
124
17
11
12
61
131
13
11
11
61
140
21
11
13
62
133
46
11
11
61
162
31
9
13
10
7
11
18
11
11
61
125
9
14
11
8
11
12
13
138
9
13
10
6
11
11
11
60
62
9
13
11
6
12
156
28
34
34
11
12
61
136
29
30
30
30
45
47
40
45
42
43
9
14
11
7
11
106
14
12
12
61
131
31
9
13
11
6
10
134
23
11
11
61
143
29
9
14
10
7
129
26
11
12
61
151
31
9
14
11
6
119
16
11
11
61
155
20
9
14
12
6
148
156
F
F
F
M
13
2
M
17
2
F
12
2
6
2
41
18
2
M
M
F
17
1
12
40
50
7
2
10
45
15
2
11
49
4
2
M
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(nmi)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
M Sex
942339
942340
942341
942342
942343
942344
942345
942346
942347
942348
942298
942299
942300
MayCreek
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
138
26
11
11
62
142
9
14
10
7
12
2
21
11
11
61
9
14
11
8
12
10
2
113
15
11
11
60
150
137
9
14
12
7
11
3
2
121
18
12
12
61
9
13
11
7
11
16
2
110
114
15
12
12
61
138
129
9
14
Ii
6
12
48
48
41
46
47
17
126
21
1
14
11
12
61
139
8
14
11
7
12
41
6
2
102
11
11
12
62
142
9
14
12
8
11
38
8
1
F
F
F
F
109
14
12
12
61
130
9
13
11
7
11
44
5
1
M
116
114
14
12
12
145
9
14
11
7
11
ii
2
12
9
13
11
7
13
58
41
10
14
6
2
F
M
167
45
10
11
9
14
12
5
11
31
1
2
M
118
15
11
13
9
12
12
7
12
1
15
10
11
9
13
11
7
12
42
50
7
106
3
2
77
74
110
5
11
12
9
14
12
6
11
41
0
1
5
12
12
13
11
7
11
44
0
1
11
12
9
9
M
M
M
M
14
14
11
7
12
35
4
1
12
ii
M
103
12
62
60
62
62
60
62
62
59
61
8
14
12
8
12
2
M
24
10
12
61
9
13
12
6
12
12
2
79
5
11
12
62
12
6
12
12
11
12
61
9
9
14
105
13
11
6
12
39
40
45
34
11
136
65
70
70
3
11
12
61
9
13
12
7
12
38
0
1
F
M
F
M
4
11
12
9
14
11
7
12
1
10
11
9
13
11
7
12
48
45
0
4
62
62
29
32
29
29
29
29
29
27
30
27
26
30
30
29
27
33
32
33
27
29
28
29
26
0
1
May Creek
May Creek
May Creek
MayCreek
MayCreek
May Creek
May Creek
942303
942304
942305
MayCreek
May Creek
Miii Creek
Miii Creek
Mill Creek
Mill Creek
Mill Creek
Mill Creek
Mill Creek
Mill Creek
942306b
Mill Creek
942307
942308b
MiliCreek
942301b
942302b
942309"
942310"
Mill Creek
Mill Creek
Mill Creek
138
135
147
120
119
124
136
152
133
120
126
120
119
119
0
1
2
2
F
F
M
M
Appendix Table A2. 1. Continued
Fish
Date of
FL
Wt.
number
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
942311b
Mill Creek
Mill Creek
Mill Creek
Mill Creek
Mill Creek
Miii Creek
Mill Creek
MiliCreek
Mill Creek
Mill Creek
Mill Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/3/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
69
73
72
85
70
88
116
116
109
116
4
11
13
62
28
9
12
4
11
12
61
116
120
10
13
5
II
12
117
9
14
6
10
12
61
62
9
4
11
12
8
10
12
62
62
117
128
30
27
29
28
28
16
11
11
61
13
10
12
13
11
11
60
62
152
139
147
16
11
11
61
132
11
12
61
11
12
11
12
11
11
12
14
12
13
179
22
78
95
126
52
66
64
12
12
144
3.
11
13
182
76
ii
13
101
14
10
ii
123
27
11
12
92
11
10
11
62
63
63
64
62
62
64
63
59
63
60
109
15
11
12
61
942312"
942314"
942315b
942316b
942317
942318
942319
942320
942321
942252
942253
942254
942255"
942256b
942257"
942258"
942259
942260
942261"
942262
942263
131
198
205
225
159
170
H
I
J
K
12
7
11
11
6
12
10
8
11
14
12
6
12
9
14
12
7
11
14
12
6
12
0
0
1
9
31
9
13
12
12
17
32
9
14
11
6
6
6
2
2
M
M
M
M
M
M
F
F
33
9
Ii
5
12
4
1
M
9
13
12
6
12
137
30
29
9
13
12
6
12
127
25
9
13
12
7
11
126
25
9
13
12
8
13
42
46
38
49
42
42
33
36
36
36
42
48
40
130
132
129
124
125
25
9
14
12
6
12
26
26
24
24
32
30
23
30
27
10
14
12
7
12
9
14
11
7
12
9
13
11
6
11
9
14
12
12
9
14
12
6
7
9
13
11
8
9
13
11
9
13
9
14
128
131
124
127
121
127
11
L
M Sex
0
1
0
1
0
0
1
1
1
7
2
F
14
2
0
2
0
2
41
0
2
0
1
0
0
1
12
46
45
47
29
44
11
51
F
F
F
F
M
M
F
F
M
M
7
12
12
7
12
11
7
12
54
48
44
1
0
1
0
2
6
1
0
1
3
1
5
1
M
M
M
0
Appendix Table A2. 1. Continued
Fish
number
942264b
942265b
942266b
942267
942268b
942269
953560
953561
953562
953563
953564
953565
953566
953567
953568
953569
953570
953571
953572
953573
953574
953575
953576
Date of
FL
Wt.
Location
collection
(mm)
(g)
A
B
C
D
E
F
G
H
I
J
K
L
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
Widow White Creek
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
11/2/94
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
132
116
123
26
11
13
121
14
12
7
12
59
13
9
14
11
7
13
53
27
12
13
9
13
12
8
11
110
142
17
11
12
9
14
11
7
11
33
53
0
0
0
1
12
6
1
36
12
13
10
14
11
7
12
31
0
1
179
68
12
13
126
27
22
23
28
27
27
27
28
29
9
19
64
63
63
60
64
62
9
13
12
7
12
0
2
10
13
11
8
11
8
2
10
12
12
7
12
3
2
9
13
11
12
11
0
2
15
2
M
M
M
M
M
M
M
M
F
F
6
2
F
122
116
137
125
9
13
13
8
11
10
14
10
6
11
9
13
11
7
12
9
14
11
8
11
9
13
13
7
11
28
9
14
11
7
12
137
31
9
13
11
7
ii
39
37
46
46
50
44
54
48
36
36
50
48
46
130
129
133
150
129
27
10
12
12
7
11
31
9
14
11
8
12
29
32
9
13
11
7
Ii
9
14
10
6
11
27
10
14
12
7
12
165
11
12
61
128
179
11
12
62
126
200
203
12
13
61
11
11
61
123
138
31
10
15
12
8
12
180
11
12
124
27
9
13
12
8
11
203
205
11
11
62
62
12
13
61
128
133
175
10
11
61
135
173
12
12
62
135
204
210
11
12
61
123
30
29
30
32
30
12
12
61
129
189
11
12
61
193
11
1!
61
208
184
12
12
61
11
12
60
193
12
12
61
198
12
11
60
M Sex
1
1
6
2
F
10
2
14
2
5
2
16
2
6
2
6
1
41
0
2
50
42
46
50
9
2
0
2
3
2
20
2
M
F
F
F
F
M
F
F
M
F
M
C
Appendix Table A2. 1. Continued
Fish
a
Date of
FL
Wt.
(g)
number
Location
collection
(mm)
953577
953578
953579
953580
953581
953582
953583
953584
953585
953586
953587
953588
953589
953590
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
H.S.U. Hatchery
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
519/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
5/9/95
A
B
C
D
E
F
232
12
12
61
134
181
11
11
125
29
28
184
10
11
139
31
185
11
12
59
60
59
125
28
165
11
12
61
135
182
162
11
12
12
13
129
138
173
12
12
62
62
60
180
185
11
12
61
11
11
61
150
11
11
167
11
12
62
63
181
11
11
61
181
13
12
61
140
138
137
124
130
132
131
G
H
10
13
13
9
13
13
14
12
7
11
9
13
10
7
11
31
9
14
12
7
11
28
30
28
29
27
29
29
27
9
10
14
11
7
11
14
11
7
11
10
14
12
7
12
14
11
7
12
9
13
11
7
12
10
13
11
7
11
9
13
10
7
11
9
14
11
7
11
28_ 9
13
127
J
K
L
M
Sex
8
12
2
12
44
42
42
42
42
48
44
0
6
0
2
M
M
F
M
M
F
I
ii
46
48
44
42
50
44
2
2
9
2
13
2
6
1
16
2
17
2
7
2
0
2
8
2
F
F
M
M
F
9
1
F
14
2
M
3610
2
E
missing data for at least one of the following meristic counts: vertebrae, scales in lateral series, scales above lateral line,
basibranchial teeth, or maxillary extension.
individuals designated as 0. mykiss based on the absence of basibranchial teeth and a maxillary extension score of 0 or 1.
13
209
Appendix Table A2.2. Summary of meristic counts from each collection location.
Meristic characters are: A: anal pterygiophores, B: dorsal pterygiophores, C:
vertebrae, D: scales in lateral series, E: scales above lateral line, F: pelvic fin rays, G:
pectoral fin rays, H: gill rakers lower arch, I: gill rakers upper arch, J:
branchiostegal rays, K: pyloric caeca, L: basibranchial teeth (see text for description
of character counts). Locations are listed from north to south with location number in
parentheses; FL: fork length (mm), WT: weight (g), blanks represent missing data.
Character
0. c. clarki
N
Bosewell Bay, AK (1)
FL
33
WT
33
A
33
B
C
D
E
F
G
H
I
J
K
L
Martin River, AK (2)
FL
WT
A
B
C
D
E
F
G
H
I
J
K
L
Gines Creek, AK (3)
FL
WT
A
33
33
33
33
33
32
33
33
33
Mean
SD
284.1
269.5
73.7
189.72
11.5
11.8
62.1
147.6
067
32.4
9
13.7
11.7
6.9
11.2
12
41.6
33
12.5
23
23
23
370.5
537.9
22
22
23
12
23
23
23
23
23
23
11.5
62
147.3
34.7
9.1
13.9
11.7
7
11.4
0.7
1.27
12.57
2.82
0.17
0.58
0.76
0.56
0.58
4.29
7.18
55.71
245.54
0.73
0.62
0.95
15.35
4.43
21
23
41.6
4
0.46
0.73
0.93
0.67
0.58
9.25
4.79
46
46
46
184.7
69.5
10.8
25.84
31.54
0.42
0. mykiss
Mm
Max
165
43
10
10
58
125
25
9
425
743
12
11
15
14
6
10
13
34
2
50
28
280
216
10
N
Mean
SD
468
1056
2
2
13
2
149.5
35
11.5
11
13
2
12
61
118
2
2
8
65
173
42
10
12
15
2
63.5
126.5
27.5
9.5
14.5
9
13
2
11
1.41
6
8
2
7.5
26
Mm
Max
33.23
22.63
126
19
0.71
0
0.71
11
173
51
12
12
12
13
65
183
37
10
8
2
2
2.12
0.71
0.71
0.71
11
13
2
11
0.7]
0
22
0
61
14
2
44
0
14.14
0
127
22
10
248
174
12
2
12
63
125
27
9
14
10
7
64
128
28
10
15
12
8
11
11
34
0
54
0
210
Appendix Table A2.2. Continued
Character
B
0. c. clarki
N
Mean
45
46
46
46
46
46
46
46
45
44
46
11.1
C
D
E
F
G
H
I
J
K
L
Suntaheen Creek, AK (4)
FL
59.8
149
35.5
9.2
13.9
11.4
7.2
10.8
48.4
10.5
0. mykiss
SD
0.7
0.93
7.03
2.09
0.4
Mm
10
13
58
137
32
9
62
170
40
10
0.64
0.58
0.48
0.47
5.08
4.83
13
15
10
12
6
10
8
12
39
61
2
25
Max
N
Mean
SD
17
134.5
17.8
11.4
12.4
64.5
125.3
26.5
9.5
14.5
12.6
7.8
11.8
46.3
0
71.63
4.46
0.51
0.62
0.8
6.51
124.1
19.9
11.9
12.6
11
13.79
6.7
0.44
0.5
1.54
2.86
1.46
0.52
0.73
0.6
0.65
0.55
WT
17
A
17
B
17
C
D
17
E
17
F
17
17
G
H
I
J
K
L
Freshwater Bay, AK (5)
FL
15
WT
14
A
15
17
17
17
16
16
17
124
18.7
11
B
C
15
D
E
F
G
H
15
15
11.2
62.5
145.1
33.8
9.1
13.7
11.4
I
15
5.9
J
15
10.5
15
15
15
15
20.39
8.16
0.53
0.56
0.92
5.66
96
154
8
31
12
60
64
1.57
137
32
0.26
0.49
9
155
37
10
13
14
0.51
11
12
0.88
0.64
4
8
16
16
16
16
16
16
16
16
16
16
15
9
11
14
10
10
12
64.9
126.2
26
9.5
14
12.3
7
2.21
0.51
0.51
0.51
0.56
0.54
4.53
0
Mm
Max
9
409
28
11
12
92
11
13
63
113
22
66
9
14
12
7
11
138
30
10
15
13
9
13
37
0
54
91
8
143
31
11
13
12
13
61
67
132
28
10
120
24
9
0
13
15
ii
13
6
8
10
12
211
Appendix Table A2.2. Continued
Character
0. c. clarki
N
0. mykiss
K
L
Kiag Bay, AK (6)
15
Mean
45.8
15
6.3
SD
5.66
3.79
FL
WT
18
241.7
85.63
91
340
A
B
C
D
E
18
11.4
11.1
0.78
0.68
0.62
9.77
2.17
10
10
13
F
18
G
H
18
1
18
J
K
18
H
32
13
Mean
47.3
2
16
16
0
SD
5.09
0
42
Max
57
0
0
59
92
3
11
9
13
11
13
65
130
29
Mm
0
18
18
61.4
18
144.4
31.6
9.1
13.2
11.4
6.4
10.5
52.5
7.3
18
18
18
L
18
Portage Creek, AK (7)
FL
22
WT
22
A
22
B
22
C
22
D
22
E
22
F
22
G
N
Max
52
Mm
21
22
I
22
J
22
K
22
L
22
Duncan Salt Chuck, AK (8)
FL
50
WT
50
A
50
B
50
C
49
D
50
133.8
34.4
11.3
11.5
61.5
155.7
36
9
0.24
0.38
0.78
0.86
0.62
7.78
3.36
52.38
46.78
0.65
0.67
12
60
62
123
5
158
35
10
14
12
8
9
11
40
66
3
14
76
4
269
8
10
177
12
7
69.8
4.3
11.6
27
9
13
10
10
13
8
12.1
10.39
1.85
0.79
0.64
1.01
60
61
8
42
8
63.3
116.9
26.4
1.71
136
32
63
187
8
15.15
2.92
0.31
7.49
2.45
10
8
9
0
106
23
9
13.6
11.8
0.52
0.46
0.53
0.69
5.47
8
8
11
0.66
0.8
0.63
0.38
44.1
8.8
6.31
4.55
32
56
7
6
4
21
8
292.1
287.5
11.4
11.7
61.4
151.9
59.87
151.97
0.72
0.59
0.82
10.89
148
34
400
687
10
13
11
13
60
63
175
13.7
11.8
6.7
13
15
8
10
6
10
13
8
8
8
12
129
7
11.1
38.7
0
0
9
11
14
12
6
10
12
33
0
48
0
13
8
212
Appendix Table A2.2. Continued
Character
E
F
G
H
0. c. clarki
N
50
50
50
50
1
50
J
50
K
48
L
50
Kadake Creek, AK (9)
FL
38
WT
38
A
B
C
D
E
F
G
H
1
J
K
L
38
38
38
38
38
38
38
38
38
37
37
38
Eagle River, AK (10)
FL
31
WT
31
A
B
C
D
E
F
G
H
I
J
K
L
31
31
31
31
31
31
31
31
31
31
31
31
Wolverine Creek, AK (11)
Mean
34.7
SD
3.05
9.1
13.4
11.7
7
11.1
41
14.3
0.37
0.53
0.65
0.68
0.66
5.52
268.7
60.41
116.4
0.72
198.7
11.4
11.5
62.1
151.3
35.4
9.1
13.3
11.8
6.9
10.8
42.2
14.3
199
86
11.2
11.5
61.4
144.4
32.5
9
13.4
11.6
6.6
11.2
45.7
13.2
7.21
0.56
0.95
9.15
2.7
0.23
0.57
0.69
0.7
0.6
6.04
6.14
0. mykiss
27
Max
42
Mm
8
10
13
15
10
6
10
13
28
2
29
365
459
10
11
13
13
135
65
169
30
40
9
10
12
14
11
13
6
10
12
20
3
Mean
9
12
51
152
35
61
N
8
53
30
47.04
75.26
109
371
438
1
167
13
1
40
0.8
0.63
0.89
6.68
1.96
10
13
1
11
10
59
134
29
9
13
1
12
1
64
1
131
1
26
12
63
163
36
9
14
8
4
0
0.56
0.95
0.88
0.37
7.75
7.95
1
10
1
14
13
1
12
8
1
11
12
1
7
10
32
63
48
1
1
1
45
0
SD
Mm
Max
213
Appendix Table A2.2. Continued
Character
0. c. clarki
N
FL
49
WT
49
A
50
B
50
C
47
D
50
E
50
F
50
6
50
H
50
1
50
J
49
K
47
L
50
Staney Creek, AK (12)
FL
51
WT
51
A
51
B
C
D
E
F
51
51
51
51
51
51
Mean
282.4
252
11.2
11.5
61
147.6
34.3
9.1
13.1
11.2
6.2
11.1
44.4
13
203.5
91.7
Max
N
122
421
1
18
745
1
Mean
216
110
10
12
1
12
10
13
1
13
58
1
64
8.8
131
63
177
1
123
2.57
0.35
0.57
0.86
0.96
0.55
5.52
6.15
30
40
1
11.4
11.2
61.4
1
9
10
1
26
10
12
14
1
15
9
13
1
12
4
10
8
1
7
12
1
11
56
32
1
37
0
28
3
364
453
8.25
2.41
0.32
0.48
0.8
0.57
0.66
5.21
121
13
158
37
10
14
9
13
6
9
38
8
62
8.61
3
41
236.4
159.6
76.32
122.98
100
355
449
11.1
11.3
10
141
29.9
51
J
K
51
11
50
L
51
45.5
22.4
Vixen Inlet, AK (13)
FL
22
WT
22
A
21
B
22
Mm
119
24
10
10
58
I
51
SD
72.15
171.84
0.55
0.58
0.87
49.06
74.21
0.69
0.62
9
13.7
11.4
6.8
G
H
0.
23
8
1
13
12
64
12
1
185
1
64
1
12
10
12
12
60
62
1
132
1
1
28
10
1
14
9
C
D
E
21
61.3
22
22
1148.5
0.62
0.55
0.64
9.75
33.8
1.93
30
F
21
9
8
G
22
13.3
0.32
0.57
169
37
10
12
14
1
1
12
62
130
.'nykiss
SD
Mm
Max
214
Appendix Table A2.2. Continued
Character
0. c. clarki
N
22
22
22
H
I
J
K
19
L
22
Margaret Creek, AK (14)
FL
26
WT
26
A
26
B
26
C
25
D
26
E
26
F
26
G
26
H
25
I
25
J
26
K
26
L
26
Throne River, AK (15)
FL
62
WT
58
A
62
B
62
C
62
D
62
E
62
F
62
G
62
H
62
I
62
J
62
K
59
L
62
Bakewell Creek, AK (16)
FL
36
WT
36
A
36
Mean
11.5
6.1
11.1
SD
0.74
0.68
068
0. mykiss
Max
N
Mean
10
13
1
13
5
10
7
1
7
12
1
11
55
28
1
1
44
0
261
147
1
67
10
1
3
10
12
1
12
11
13
1
13
Mm
46.3
11.2
6.04
6.43
35
151.5
39.2
46.07
37.6
0.52
0.59
99
11.1
11.8
61.8
146.5
32.1
9
13.3
11.2
6.4
11
48.7
13.6
245.8
180.3
11.2
11.4
61.9
150.3
33.2
9
13.3
11.6
6.5
11.1
49.1
20.4
207.9
116.3
11.1
1
1.01
60
64
1
64
9.56
2.42
0.2
0.47
0.83
0.76
0.57
6.32
5.69
131
1
121
1
29
8
163
38
9
1
9
13
14
1
13
10
13
1
11
77.14
137.05
0.6
117
18
393
515
10
13
0.61
10
13
0.99
60
64
10.93
2.59
0.26
130
28
185
39
8
10
0.64
0.72
0.62
0.66
6.69
7.98
10
14
10
13
69.65
108.36
0.59
28
5
8
1
6
10
37
4
12
1
10
62
24
0
5
8
10
13
39
74
36
1
100
12
10
1
0
1
199
87
12
398
571
1
12
1
SD
Mm
Max
215
Appendix TaMe A2.2. Continued
Character
B
C
D
E
F
G
H
I
J
K
L
0. c. clarki
N
Mean
36
36
36
36
36
36
36
36
36
36
36
11.4
Herman Creek, BC (17)
FL
25
WT
0
A
25
B
C
D
E
F
G
H
I
J
K
25
25
25
25
25
25
24
24
25
25
25
L
Survey Creek, BC (18)
FL
20
WT
20
A
20
B
20
C
20
D
20
E
20
F
20
G
20
H
20
I
20
J
20
0. mykiss
Max
N
Mean
10
12
1
13
60
63
175
38
1
64
1
1
13
9
15
131
25
10
1
15
0.84
0.65
0.69
5.86
5.46
10
13
1
11
123.1
21.81
11.1
11.6
0,67
0.49
0.65
7.92
61.2
154.7
34.3
9
13.9
11.6
6.5
11.4
50
8.8
61.5
140
30.5
9.2
13.6
11.4
6.5
10.8
42.4
7.7
158.7
50.4
11.4
11.8
62.7
145.9
33.6
9.2
SD
0.56
0.91
8.97
2.32
0.17
0.5
Mm
1.53
0.41
0.51
138
30
8
5
8
1
7
10
13
1
12
41
1
0
72
22
43
0
83
160
10
12
11
12
60
126
28
63
159
34
9
10
10
14
12
7
12
13
0.65
0.51
0.5
4.47
5.44
10
36
0
52
28
57.01
46.11
0.59
73
4
275
10
12
0.64
0.75
9.14
2.23
14
0.41
0.65
12
7.1
11.2
0.69
0.39
0.62
1
6
173
11
13
61
64
133
30
167
38
9
10
13
15
11
14
6
8
10
12
1
SD
Mm
Max
216
Appendix Table A2.2. Continued
Character
0. c. clarki
N
K
L
19
20
Yakoun River, BC (19)
FL
16
WT
16
A
B
C
D
16
16
16
F
16
F
G
H
16
I
16
J
K
L
16
16
16
16
Mean
44.2
12.3
268.2
202.7
11.1
11.6
61.8
144.1
31.6
8.9
13.6
12.3
7.1
10.6
43.5
20.6
Molly Walker Creek, BC (20)
FL
25
123.9
WT
25
24
A
25
10.9
B
25
11.6
C
25
62
D
25
145.4
E
25
33.1
F
25
9.4
G
25
13.7
H
25
11.4
I
25
6.6
J
25
10.9
K
25
39.3
L
25
5.6
Fish Creek, BC (21)
FL
WT
A
B
C
D
14
16
13
13
13
13
13
13
127.5
27.8
11.2
11.4
61.9
140.8
SD
6.32
6.74
72.18
134.11
0.57
0.5
0.83
35
Max
57
1
29
Mm
120
16
10
11
60
395
508
12
12
7.36
132
63
154
1.54
29
34
0.34
8
9
1.03
10
14
0.93
0.68
0.62
10.85
9.28
11
14
6
8
10
12
25
60
8
39
86
180
51
25.77
12.96
0.64
0.64
0.58
8.96
133
8
10
10
12
61
13
1.98
29
63
167
37
0.49
0.46
0.58
9
10
13
14
10
12
0.71
0.53
5
10
12
8.1
22
2
60
95
208
110
3.47
31.8
27.9
0.55
0.65
0.86
7.96
11
10
10
61
128
N
0. mykiss
Mean
SD
8
14
12
12
63
158
1
143
1
27.8
1
11
1
12
1
62
150
1
Mm
Max
217
Appendix Table A2.2. Continued
Character
0. c. clarki
N
E
F
G
H
I
13
13
13
13
13
13
13
13
J
K
L
San Josef River, BC (22)
FL
25
WT
25
A
25
B
C
D
E
F
G
H
1
25
25
25
25
25
24
25
25
25
24
25
J
K
L
Willow Creek, BC (23)
FL
9
WT
9
A
9
B
C
9
9
D
E
F
9
G
9
H
9
9
9
I
J
9
9
K
9
L
9
Mixal Creek, BC (24)
FL
WT
14
14
Mean
31.8
SD
0. mykiss
Mm
Max
N
Mean
37
SD
Mm
Max
45
1
186
68
10
13
104
25
9
50
40.34
18.27
0.73
0.38
0.73
7.63
2.02
0.43
0.7
0.69
0.78
0.95
2.83
0
28
9
35
9
1
1
9
13
14
12
1
1
10
7
12
13
12
7
11
1.86
0
0.38
0.51
0.63
0.71
44.3
4.5
3.9
2.07
36
50
1
10
48
1
10
1
0
191.3
78.5
10.9
11.2
61.2
149.6
31.2
9.4
51.99
64.32
0.64
0,55
119
296
17
10
10
231
1.04
60
8.1
131
1.62
28
9
64
170
34
10
9
13.2
11.4
6.3
14
11.5
6.5
11.2
50.5
13.2
137,4
33
11.2
11.7
63.1
131.8
30
9.2
14
11.8
7.2
11.6
39.8
5
0.5
0.55
0.77
0.65
0.5
5.35
8.12
45.48
33.23
0.67
0.71
1.36
10.17
2.4
0.44
0.5
0,67
0.67
0.73
6.36
2.29
66.6
8.1
3.3
1.33
11
5
13
10
5
10
42
15
13
8
12
1
96
221
10
10
96
62
119
27
9
13
11
6
10
32
0
54
2
1
12
12
62
36
ii
1
12
13
66
151
33
10
15
13
8
12
50
8
83
6
14
14
14
13
14
13
14
14
14
13
13
14
2
14
65.7
7.5
11.3
11.8
62.7
119.9
27.4
9.2
13.8
12.2
6.5
11.1
11
12
61
64
133
32
13
10
15
11
13
5
8
9
48
12
0
0
0
52
3
53
5
48
58
3
1.3
0.58
1
2
218
Appendix Table A2.2. Continued
Character
A
B
C
D
E
F
G
H
I
J
K
0. c. clarki
0. mvkiss
N
Mean
SD
Mm
Max
N
Mean
11.3
11.7
61.1
143.1
31.1
8.9
13.4
11.5
7.1
0.47
0.47
0.77
9.16
2.5
0.27
0.65
0.52
0.27
0.66
6.09
2.44
il
12
12
3
3
11.3
11.7
61.3
132.7
SD
0.58
0.58
0.58
7.64
Mm
14
14
14
3
29.7
1.15
29
3
9
13.5
11.7
0
0.71
0.58
9
13
7
0
10.7
0.58
6.43
24
0
0
14
14
14
14
14
14
14
13
14
L
Salmon River, BC (25)
FL
29
WT
29
A
29
B
29
C
29
D
29
E
29
F
29
G
29
H
29
I
29
J
29
K
29
L
29
Ritherdon Creek, BC (26)
FL
25
WT
25
A
25
B
25
C
25
D
25
E
25
F
25
G
25
H
25
1
25
J
23
K
25
11.1
34.9
4.4
139.9
23.3
11
11.1
61
137.9
30.8
9
13.2
10.9
6.7
10.7
43.9
5.5
104.5
14.2
11.4
12
60.7
138.2
16.56
9.96
0.57
0.58
0.87
7.9
7
8
10
12
24
0
44
114
12
10
10
59
121
176
48
12
12
26
8
12
0.74
0.6
0.7
5.26
5.44
9
19.92
7.88
0.5
0.45
0.75
7.94
1.43
0.28
44.2
11
62
160
36
9
14
12
29
31
6.2
60
128
1.66
0.19
0.56
9.1
13.7
11.3
10.7
11
1.54
0.94
0.72
0.47
5.16
8
12
6
8
62
154
35
9
15
13
8
12
9
34
0
54
23
80
137
5
30
11
12
13
11
59
127
28
9
62
153
34
10
5
15
13
8
10
32
58
7
9
11
3
3
3
3
3
3
3
3
31.3
0
11
11
Max
12
12
61
62
126
11
141
31
9
14
12
7
10
11
7
36
0
219
Appendix Table A2.2. Continued
Character
0. c. cIa rki
N
25
Mean
9.2
4.43
Sandhill Creek, BC (27)
FL
23
WT
23
133.7
29.3
21.14
15.05
11.3
10,8
0.71
L
A
B
C
D
E
F
G
H
I
J
K
L
23
23
23
23
23
23
23
23
23
22
23
23
Kirby Creek, BC (28)
FL
26
WT
26
A
26
B
26
C
26
D
26
E
26
F
24
G
26
H
26
I
26
J
26
K
26
L
26
Church Creek, WA (29)
FL
53
WT
53
A
60.9
142.8
33.9
9
13.4
11.3
SD
0.58
0.87
6.24
2.35
6.2
10.7
38.4
8.6
107.9
13
16
105
13
10
10
181
2
67
2
12
12
2
116
18.5
11.5
2
11
59
134
29
63
154
38
10
2
60.5
140.5
2
31
1.41
30
9
13.5
0
0.71
0
0.71
0.71
16.97
9
13
9
14
11
11
1
1
0.56
0.48
0.64
7.82
2.69
0.46
0.5
0.65
0.49
0.62
4.7
3.78
10
10
12
12
1
60
62
1
60
133
161
1
121
27
38
1
26
8
1
10
14
12
7
10
10
14
12
7
12
1
11
30
48
1
32
1
16
1
0
87
148
33
12
13.08
5.16
0.59
0.52
13
9
5
7
0.72
7.16
63
121
E
F
53
53
61.3
134.5
30.9
60
D
52
53
1.91
24
9
G
52
0.24
0.56
12
154
35
10
14
10
10
8
13
1
1
1
12
11
20
61
45
1
11
142
32
158
12
124
60
5
8
108
17
139
79
10
6
9
14
13
Max
2.12
22.51
10.07
13
2.12
Mm
0.71
0
0.71
0
9
10.9
11.2
13.2
11.31
1
53
53
B
C
SD
11
11.1
13.6
10.9
Mean
48
106.2
14.1
9
N
32
39
3.9
10.9
61.4
144.3
32.9
Max
2
2
2
2
2
2
2
2
0.21
0.51
0.81
0.6
0.67
4.22
3.52
7.1
10.5
0. mykiss
Mm
11
7.5
11.5
44
0
96
8
13
14
0
11
7
8
11
12
32
0
56
0
220
Appendix Table A2.2. Continued
0. c. clarki
Character
N
H
51
I
52
50
52
53
J
K
L
Bear Creek, WA (30)
Mean
11.4
SD
10
3
8
9
31
12
40.8
0.8
0.75
6.38
64
5.5
3.64
0
15
73
4
179
57
10
13
13
21
21
103.7
A
21
21
11.4
11.9
62.1
133.6
29.23
14.69
0.86
0.62
0.83
7.66
31
8.9
13.8
10.8
6.1
10.9
20
G
21
21
21
21
H
21
I
21
21
19
21
F
J
K
L
Salt Creek, WA (31)
FL
WT
A
B
C
D
E
F
24
24
24
24
24
24
24
23
24
24
24
24
G
H
I
J
21
K
L
24
Dickey River, WA (32)
FL
WT
12
A
B
12
7
12
Max
0.66
6.5
10.6
FL
WT
B
C
D
E
0, mykiss
Mm
14.1
11
60
64
1.58
0.3
0.4
8
144
35
9
13
14
9
4
12
7
9
12
40.2
0.68
0.89
0.79
4.16
32
46
11.8
5.3
3
21
106.5
14.3
11.4
11.4
61.5
145.9
31.6
8.9
13.5
11.3
24.22
70
4
165
0.49
0.78
6.44
132
1.84
29
10.8
39.9
6.1
0.55
0.66
0.74
0.95
0.56
4.63
4.24
165.3
29.9
11.5
11.8
35.69
6.52
0.52
0.62
6
10
11
60
8
15
13
8
12
34
54
2
18
104
228
35
11
106
13
12
12
10
4
10
11
1
46
63
156
36
10
17
Mean
13
114
28
10.35
0.65
N
12
13
0
1
13
1
12
SD
Mm
Max
221
Appendix Table A2.2. Continued
Character
N
C
D
E
F
G
H
12
12
12
12
12
12
12
12
12
12
U. c. clarki
Mean
SD
61.7
0.78
147.2
32.4
9.1
13.8
11.8
6.4
J
10.3
K
45.8
L
10.3
Alder Creek, WA, Hoh River (33)
FL
13
127.5
WT
13
24.5
A
13
11.2
B
13
11.6
C
13
61.2
D
13
150.2
E
13
34.4
I
F
0
H
I
J
K
L
13
13
13
13
13
13
13
Big Beef Creek, WA (34)
FL
20
WI
15
A
20
20
B
C
D
E
F
G
H
I
1
K
L
19
20
20
20
20
20
20
20
20
20
9
13.2
11.5
6.8
10.5
41.8
5.8
149.6
34.2
11.4
11.1
61.8
142.2
31.8
9
13.4
11
6.5
10.8
40.4
7
0. mykiss
Mm
Max
61
63
168
37
10
10.56
2.15
0.29
0.39
136
0.39
0.67
0.78
11
8.41
36
6.5
0
21.8
18.89
0.38
0.51
1.09
7.34
1.94
98
10
193
11
12
12
0
0.38
0.52
0.6
0.66
7.23
2.23
30
9
13
6
9
11
60
139
31
9
13
14
12
8
12
69
25
86
63
164
37
9
11
14
12
6
8
10
12
26
52
2
11
15.24
9.88
0.6
0.45
0.6
9.04
2.36
0.32
0.68
0.69
0.51
0.55
119
21
172
11
13
12
4.61
4.51
29
0
10
61
131
28
8
12
10
6
10
60
63
161
38
10
15
12
7
12
48
15
N
1
1
1
1
1
1
Mean
64
117
23
10
15
13
1
6
10
1
51
1
0
1
SD
Mm
Max
222
Appendix Table A2.2. Continued
Character
0. c.
N
clarki
Mean
SD
125
33.26
22.14
0.5
0.52
0.6
6.51
2.38
0.2
0.64
0.58
0.51
0.85
5.89
5.94
0. mij kiss
Mm
Max
83
7
210
94
10
12
N
Mean
Stevens Creek, WA (35)
FL
WT
A
B
C
D
E
F
G
H
I
J
K
25
25
25
25
25
25
25
25
25
25
25
25
25
25
24.3
11.2
11.8
61.8
148
34
9
13.4
12.2
6.6
10.8
41.6
L
10.3
West Fork McClane Creek, WA (36)
FL
WT
I
32
32
32
32
32
32
32
32
32
32
32
J
31
K
L
32
32
A
B
C
D
E
F
G
H
111.5
14.6
11.5
11.5
61.3
138.7
31.9
9.1
13.3
11.3
6.5
10.8
37.8
7.4
11
13
61
137
30
63
166
37
9
10
13
11
6
15
14
7
10
13
30
2
54
24
21.12
66
164
8.21
3
40
0.62
0.57
0.98
8.37
2.14
0.25
0.54
0.85
0.62
0.52
3.3
4.71
11
13
10
12
60
64
160
36
129
28
9
12
10
5
10
10
14
13
8
12
32
0
48
70
219
1
3
1
10
109
12
11
13
1
61
1
8
63
174
38
10
1
64
117
25
10
13
15
1
13
19
Alder Creek, WA, Naselle River (37)
FL
WT
A
24
24
D
24
24
24
24
B
23
F
G
24
24
B
C
131.2
26.5
11.3
11.8
61.9
151.7
33.7
9.1
13.5
32.5
23.45
0.62
0.53
0.74
12.05
2.4
0.41
0.72
132
30
1
1
1
71
4
12
13
SD
Mm
Max
223
Appendix Table A2.2. Continued
Character
H
I
J
K
L
0. c. clarki
N
24
24
24
23
24
Mean
11.5
6.4
10.5
34.1
8.3
0. mykiss
SD
0.78
0.82
0.59
4.24
5.54
Mm
10
4
Max
13
N
0
Mean
1
6
10
19
7
12
41
0
2
22
1
27.8
53
165
0.62
0.55
10
13
13
0
0
Foley Creek, OR (38)
FL
WT
30
107.5
0
A
30
11.4
B
30
11.9
C
30
61.5
D
30
149.2
E
29
34.5
F
30
9
G
30
13.7
H
30
11.8
I
30
6.8
J
30
10.9
K
30
41.6
L
30
15.9
North Fork Trask River, OR (39)
FL
WT
A
B
C
D
E
F
G
H
I
J
K
L
Andy Creek, OR (40)
FL
WT
A
B
30
30
30
30
30
30
30
30
30
30
30
30
20
30
21
21
21
21
205.3
97.9
11.3
11.8
61.4
152.5
34.5
9.3
14
11.5
6.7
10.7
44.3
11.9
133.5
28.3
11.4
11.9
1.17
10.9
1.64
0.18
0.6
0.79
0,49
0.83
6.72
11
59
64
126
31
9
13
10
6
10
171
29
63
33
8
4
45.8
64.72
0.64
0.76
1.07
122
12.71
131
31
9
13
9
37
10
15
14
8
13
325
330
1
176
18
1
56.7
10
12
1
11
13
1
59
63
184
38
10
1
64
1
122
1
26
1
11
15
13
1
14
1
6
8
1
12
7
10
12
1
12
29
60
23
0
5
28.91
100
201
21.5
0.6
0.7
11
90
11
13
11
13
1.91
0.45
0.56
0.78
0.55
0.65
8.73
5.44
1
12
12
0
SD
Mm
Max
224
Appendix Table A2.2. Continued
Character
C
D
E
F
G
H
0. c. clarki
Mean
SD
21
21
21
61
1.28
7.7
2.17
125
032
048
8
13
0.85
13
54
26
21
9
13.9
11.7
J
K
L
21
Schooner Creek, OR (41)
FL
10
WT
10
A
10
B
C
D
B
F
G
H
I
J
K
L
145.7
32.3
21
21
21
21
21
1
0. mykiss
N
10
10
10
9
10
10
8
8
10
9
10
Mm
56
30
7
0.71
11.2
0.68
10
6
10
42.2
654
30
15.9
5.13
8
96.1
13.5
11.4
11.6
61.3
146.5
32.3
9.3
34.61
15.08
52
2
0.52
0.52
0.82
10.02
11
125
1.5
30
048
9
14.1
0.74
0.83
0.53
0.82
6.48
4.44
13
11.9
7
10.7
40.2
4.8
11
60
11
6
10
Max
62
A
B
C
D
E
F
G
H
I
J
K
L
22
22
22
22
22
22
22
22
22
22
22
22
22
264.7
203
22
11.4
11.5
61.5
143.8
33.5
61.12
140.06
0.58
0.6
0.8
7.6
2.42
168
53
12
12
63
156
35
10
15
13
8
12
48
170
46
356
11
13
13
14
451
60
63
131
0.61
8
13
15
11
14
5
8
9
34
12
40.5
0.47
0.82
0.66
0.75
4.22
18.3
7.61
7
6.6
10.8
29
Mm
Max
2
2
2
116.5
18.8
11.5
12.5
64.5
136
27.5
10
10.61
109
5.66
0.71
0.71
15
11
124
23
12
12
13
0.71
0.71
64
135
27
0
10
65
137
28
10
15
12
7
0
15
12
7
15
12
10
11
37
0
52
8
168
38
10
9.1
13.9
12
SD
12
32
0
11
Mean
157
37
10
15
Siletz River, OR (42)
FL
WT
N
50
35
2
2
2
2
2
2
2
2
2
2
2
10.5
44.5
0
141
0
0
0.71
10.61
0
7
0
225
Appendix Table A2.2. Continued
Character
0. c. clarki
N
Mean
SD
30
0
30
30
107.6
19.07
73
155
11.1
0.61
0.67
10
10
12
0.86
59
63
10.05
2.12
127
0.37
0.7
0.82
0.77
0.78
4.55
8
12
10
5
10
169
38
10
27
8.1
3
Mm
Max
Wolf Creek, OR (43)
FL
WT
A
B
C
D
E
30
30
30
F
30
G
30
H
30
I
30
J
30
K
30
L
30
Alsea Hatchery, OR (44)
FL
WT
A
B
C
D
E
F
G
H
0
30
30
30
30
30
30
28
25
30
30
30
I
J
K
9
L
30
Cummins Creek, OR (45)
FL
25
WT
2
A
25
B
C
D
E
F
G
25
25
25
25
24
24
11.6
61.1
142
32.1
9.1
13.8
11.5
6.4
10.9
42.5
16.4
892.8
11.1
11.5
62.1
146.6
35
9.3
13.2
11.5
6.9
10
39.3
13.3
135.5
81.5
II
11.7
61.7
147.5
35.2
9
14
672.92
0.74
0.73
0.98
29
13
15
13
8
13
52
39
289 2505
10
10
13
13
60
63
170
10.74
2.68
0.7
1.36
0.57
0.61
130
10
6
41
10
15
12
8
0.32
6.42
5.6
9
32
2
52
26
44.46
41.72
0.61
0.63
0.79
9.47
2.12
0.46
0.69
74
52
10
30
8
10
11
60
135
11
228
111
12
13
31
8
63
168
39
10
13
15
N
0. mykiss
SD
Mean
Mm
Max
226
Appendix Table A2.2. Continued
Character
0. c. clarki
0. m-ykiss
N
Mean
Max
H
I
J
K
L
25
25
25
23
25
11.4
6.6
10.9
42.7
11.4
12
FL
21
141.2
24.32
111
212
WT
0
A
21
21
21
21
21
21
21
21
21
11.6
11.6
0.6
0.6
0.75
9.05
2.32
0
0.32
0.72
0.7
0.55
5.97
5.99
11
13
13
SD
Mm
0.65
10
0.86
5
0.53
10
5.95
29
8.33
1
West Branch North Fork Smith River, OR (46)
B
C
D
E
F
G
H
I
J
20
K
21
L
21
Davis Creek, OR (47)
61.4
156.1
36
9
14
11.7
6.8
10.9
40.1
12.7
FL
28
83.6
WT
28
28
28
28
26
27
28
27
28
7
A
B
C
D
E
F
G
H
I
J
K
L
28
6.7
10.9
43.3
7.9
22.35
6.65
0.5
0.73
0.98
11.26
2.16
0.33
0.58
0.58
0.66
0.73
7.25
4.36
33
33
33
33
105.4
13.5
11.3
11.5
13.59
5.2
0.59
0.57
28
27
27
11.2
11.4
61.3
139.2
32.1
9
13.1
11.5
11
60
141
32
9
8
12
54
42
63
179
39
10
9
15
13
8
12
28
4
53
27
63
147
3
10
30
13
10
5
10
12
13
60
63
126
166
37
10
14
12
26
8
12
10
6
10
8
12
28
55
1
17
85
7
10
10
141
Iron Creek, OR (48)
FL
WT
A
B
29
12
12
N
Mean
SD
Mm
Max
227
Appendix TaMe A2.2. Continued
Character
0. c. clarki
N
C
D
E
F
G
H
I
33
33
33
33
33
33
33
J
32
K
33
L
33
Winchuck River, OR (49)
FL
WT
A
B
C
D
B
F
G
H
I
J
26
26
26
26
26
26
26
26
26
26
26
26
K
26
L
26
Jordon Creek, CA (50)
FL
26
WT
26
A
26
B
26
C
26
D
26
E
26
F
26
G
26
H
26
1
26
J
26
K
26
L
26
Mean
60.3
145.3
32.2
9
14.2
11.7
6.9
11.3
42.7
6.6
SD
0.54
9.05
1.89
0.25
0.46
0.73
0.65
0.58
5.7
3.39
157.2
45.5
44.55
11
0.75
0.63
0.86
11.9
60.8
145.8
33.6
9
14
11.8
6.8
12
42.8
8.1
4758
9.21
1.72
0.4
0.45
0.73
0.69
0.69
3.42
4.25
0. mlikiss
Mm
59
130
29
8
13
10
5
19
97
9
9
10
59
132
301
Max
81.5
7.78
76
6
1.41
5
87
7
11
0
0.71
0
7.78
11
62
62
133
144
3.54
29
34
0
0.71
0.71
9
13
9
14
12
223
12
13
62
167
36
10
15
14
11
13
35
48
0
19
145
35
12
12
63
158
2
2
2
2
36
2
10
15
2
13
2
2
11
8
13
34
61
2
0
13
2
80
0.61
10
61.4
139.6
0.64
7.46
32.2
1.99
0.45
0.69
0.63
0.46
0.54
6.25
2.97
47.4
4.5
Mm
10
5
23.63
9.65
0.5
13.7
11.8
6.8
12.2
SD
10
15
13
8
13
56
8
13
Mean
35
3
31
N
61
162
10
32
109
15.6
11.4
11.2
9
Max
5
11
60
126
28
8
12
10
6
8
11.5
2
62
2
138.5
31.5
9
13.5
11.5
2
2
8
12
1.41
1.41
51.5
0
3.54
0
11
11
11
12
7
9
11
13
49
54
0
0
Appendix Table A2.2. Continued
Character
N
Waukell Creek, CA (51)
FL
WT
A
B
C
D
E
F
G
H
I
J
K
L
24
24
24
24
24
24
24
24
24
24
24
23
23
24
0. c. clarki
Mean
SD
Mm
Max
N
0. mykiss
Mean
SD
104.2
1938
74
161
1
14
11.1
11.5
9.59
0.5
0.51
0.83
7.02
1.52
0.2
0.64
0.78
0.38
0.78
5.6
1.84
4
10
48
1
80
4
12
1
11
11
12
1
11
60
63
156
34
9
1
60
1
15
13
1
128
33
9
13
12
6
10
61.1
135.1
30.8
9
13.7
11.6
7.2
11.4
39
4.1
123
29
8
12
10
1
1
1
7
10
8
1
13
1
26
50
1
1
8
1
Mm
Max
65
88
3
10
11
8
12
13
61
116
62
128
26
30
9
12
10
14
36
0
May Creek, CA (52)
FL
WT
27
27
27
27
27
27
27
27
27
27
27
27
27
27
124.5
18
99
174
20.4
8.91
10
46
11.2
11.8
0.42
0.7
0.62
9.87
2.41
0.19
0.48
0.64
0.77
0.77
5.44
5.47
11
12
13
119.4
18
103
18.1
10.7
11.7
167
45
10
11
11
13
C
12
60.9
D
E
F
G
12
12
12
11
138
9.27
0.49
0.65
0.9
9.99
12
B
12
12
12
12
59
120
26
62
152
33
8
9
14
A
B
C
D
E
F
G
H
I
J
K
L
Mill Creek, CA (53)
FL
WT
A
61.2
139.1
28.7
9
13.7
10.9
6.9
11.1
44.9
10
30.7
8.9
13.3
2.1
0.29
0.65
11
60
124
20
62
162
33
8
9
13
14
10
12
9
13
6
10
36
60
1
21
12
12
12
12
12
12
12
12
12
12
74.3
4.8
10.8
12
61.8
120.6
28
9.1
13.5
6.8
1.29
0.58
0.43
0.45
4.01
1.13
0.29
0.67
229
Appendix Table A2.2. Continued
Character
H
1
J
K
L
0. c. clarki
N
Mean
12
12
12
12
12
11.6
6.3
11.8
37.8
7.3
Max
N
11
12
8
12
12
12
12
31
50
12
1
17
12
225
126
9
9
9
SD
0.51
0.87
0.39
5.15
5.09
Mm
92
ii
12
13
11
5
0. mykiss
Mean
SD
11.5
0.67
6.7
0.65
Mm
11
43.3
0
0.51
3.5
0
12
8
12
38
0
49
0
143.1
22.17
116
39.2
17.31
19
11.7
12.9
0.5
0.6
0.83
4.67
11
179
66
12
14
11.6
10
Max
6
Widow White Creek, CA (54)
FL
WT
A
B
C
D
E
F
G
H
9
9
9
9
155.7
10.9
11.9
51.93
42.48
0.6
0.78
9
61.4
1.51
59
63
9
9
9
9
9
127.7
27.7
4.58
2.55
0
0.53
0.5
121
137
9
25
32
9
9
13
9
14
12
8
13
9
53
6
9
55.6
9
13.4
9
11.7
1
9
7.1
J
9
11.8
K
9
45.3
L
9
2.2
Humboldt State University Hatchery,
FL
31
186.1
WT
0
A
31
11.3
B
C
D
E
F
G
H
1
J
K
L
9
31
31
31
31
29
31
31
31
31
31
31
11.8
61
131.8
29
9.3
13.4
11.4
7.3
11.4
44.6
7.7
11
10
11
0.6
6
0.67
11
4.95
39
2.77
0
CA (55)
17.01
150
232
0.65
0.62
0.87
6.13
1.56
0.48
0.67
0.89
1.04
0.49
4.56
5.79
10
13
11
13
59
63
150
32
123
27
9
12
10
6
11
36
0
10
15
13
12
12
54
20
9
9
9
9
9
63.2
124.6
24.7
9.2
13.7
11.4
6.9
11.9
44.1
0
1.87
0.44
0.5
0.53
0.6
0,6
10.81
0
12
62
116
22
9
13
11
6
11
29
0
64
132
27
10
14
12
8
13
59
0
Appendix Table A2.3. Euclidian distances (lower left) and medium dissimilarities (upper right) between pairs of 54 samples of 0.
c. clarki. Location numbers correspond to numbers in Table 2.1, values in pairwise comparison refer to number of characters with
significantly different mediums (Bonferroni test, z-value >4.1386).
Location
1
1
20.150
2
3
5
6
7
8
9
10
ii
12
13
14
15
16
17
18
19
0
4
1
1
1
1
1
0
2
1
1
0
0
2
0
0
0
3
1
4
0
0
0
0
3
2
1
0
1
1
1
0
1
2
4
0
1
3
3
2
3
3
2
4
3
3
1
2
0
1
1
0
1
3
0
0
2
1
1
1
2
2
2
0
2
1
0
0
4
0
0
2
1
2
1
0
0
0
2
0
2
0
1
1
0
1
1
1
1
2
1
2
1
1
2
2
1
1
1
2
2
1
2
2
3
2
0
1
0
1
0
0
0
0
1
0
1
0
0
1
2
1
3
2
1
0
2
2
0
2
0
0
0
0
0
0
1
2
1
0
0
1
1
3
1
0
3
1
1
2
0
-
30.4360.4595
0.332
0.399
0.337
6
7
0.372
0.467
0.333
0.230
0.249
0.266
0.308
0.289
0.359
8
0.154
0.178
0.346
0.314
0.372
0.135
9
0.212
0.251
0.350
0.246
0.331
0.125
0.131
10
0.169
0.255
0.330
0.251
0.260
0.238
0.184
0.225
11
0.241
0.265
0.284
0.256
0.300
0.214
0.171
0.216
0.153
12
0.268
0.383
0.369
0.280
0.219
0.365
0.319
0.337
0.175
0.277
-
13
0.267
0.330
0.273
0.202
0.244
0.203
0.214
0.222
0.136
0.139
0.237
14
0.211
0.278
0.387
0.270
0.280
0.283
0.242
0.268
0.141
0.194
0.248
0.207
15
0.224
0,294
0.318
0.227
0.235
0.201
0.203
0.208
0.123
0.174
0.234
0.112
0.159
16
0.281
0.304
0.308
0.329
0.347
0.200
0.232
0.281
0.210
0.221
0.324
0.178
0.243
0.155
-
17
0.236
0.319
0.390
0.276
0.298
0.363
0.288
0.311
0.194
0.238
0.194
0.273
0.218
0.276
0.367
18
0.146
0.151
0.424
0.317
0.389
0.263
0.203
0.220
0.218
0.267
0.318
0.291
0.258
0.241
0.294
0.265
19
0.234
0.338
0.409
0.286
0.318
0.301
0.269
0.255
0.209
0.313
0.262
0.276
0.242
0.259
0.347
0.243
0.274
-
20
0.256
0.275
0.370
0.27 1
0.384
0.3 14
0.247
0.263
0.256
0.224
0.3 19
0.286
0.285
0.296
0.360
0.186
0.224
0.31 1
21
0.232
0.327
0.370
0.202
0.235
0.309
0.260
0.263
0.118
0.197
0.154
0,168
0.176
0.178
0.297
0.173
0.270
0.229
22
0.377
0.426
0.298
0.322
0.303
0.376
0.367
0.389
0.287
0.291
0.282
0.270
0.333
0.270
0.294
0.302
0.343
0.402
23
0.313
0.354
0.582
0.452
0.502
0.510
0.419
0.453
0.344
0.430
0.341
0.445
0.395
0420 0488
0.303
0.289
0.388
24
0.200
0.289
0.458
0.390
0.426
0.352
0.246
0.317
0.229
C. 289
0.269
0.318
0.287
0.327
0.380
0.234
0.309
0.259
25
0.377
0.471
0.349
0.276
0.247
0.417
0.382
0.386
0.256
0,294
0.182
0.268
0.295
0.307
0.396
0.243
0.424
0.320
26
0.270
0.327
0.443
0,368
0,354
0.395
0.326
0.362
0.252
0.283
0.281
0.339
0.235
0.340
0.402
0.176
0.331
0.292
27
0.404
0.479
0.297
0.288
0.301
0.354
0.353
0.322
0.326
0.324
0.291
0.292
0.414
0.348
0.417
0.353
0.434
0,371
28
0.384
0.463
0.332
0.225
0.304
0.360
0.351
0.333
0.300
0.286
0.266
0.251
0.362
0.319
0.390
0.306
0.419
0.3ii.
2
0
t'.)
0
Appendix Table A2.3. Continued
Location
1
29
30
0.372
0.295
0.243
0.239
0.248
0.306
0.173
0.232
0.262
0.132
0.238
0.117
0.217
0.185
0.192
0.374
0.208
0.240
2
3
5
6
7
8
0.464 0.386 0.265 0.290 0.428 0.384
0.355 0.545 0.375 0.421 0.456 0.381
31
0.354 0.404 0.243 0.285 0.298 0.261
32
0.317 0.401 0.267 0.281 0.281 0.267
33
0.303 0.313 0.246 0.307 0.188 0.171
34
0.406 0.380 0.223 0.250 0.350 0.315
35
0.223 0,387 0.283 0.360 0.196 0.147
36
0.319 0.411 0.303 0.323 0.345 0.263
37
0.310 0.446 0.315 0.427 0.279 0.232
38
0.144 0.395 0.333 0.390 0.181 0.107
39
0.236 0.327 0.298 0.361 0.208 0.189
40
0.175 0.412 0.376 0.391 0.274 0.181
41
0.274 0.353 0.308 0.346 0.286 0.228
42
0.258 0.339 0.240 0.308 0.225 0.183
43
0.266 0.336 0.255 0.300 0.282 0.219
44
0.413 0.376 0.256 0.364 0.329 0.323
45
0.216 0.315 0.234 0.351 0.174 0.156
46
0.252 0.365 0.335 0.410 0.111 0,147
47
0.241 0.331 0.331 0.236 0.235 0.306 0.249
48 0.216 0.279 0.321 0.353 0.352 0.276 0.222
49 0.289 0.253 0.468 0.493 0.536 0.365 0,296
50 0.365 0.401 0.464 0.461 0.442 0.434 0.390
51 0.264 0.335 0.438 0.392 0.411 0.407 0.318
52 0.281 0.368 0.479 0.402 0.367 0.447 0.367
53 0.356 0.398 0.517 0.471 0.518 0.470 0.386
54 0.423 0.472 0.625 0.563 0.544 0.606 0.5 15
55
0.343 0.387 0.507 0.479 0.437 0.518 0.419
Total number of significant differences compared to all other
52
64
168
53
79
68
74
9
10
0.377
0.403
0.267
0.229
0.150
0.299
0.148
0.285
0.217
0.163
0.199
0.267
0.241
0.174
0.258
0.245
0.164
0.151
0.259
0.267
0.301
0.192
0.250
0.215
0.239
0.192
0.210
0.316
0.207
0.259
0.175
0.247
0.177
0.127
0.361
0.192
0.283
0.124
0.30! 0.165
0.404 0.290
0.449 0.294
0.381 0.219
0.423 0.245
0.469 0.302
0.579 0.387
0.477 0.321
populations
85
31
11
12
13
14
15
16
17
18
0.314
0.351
0.238
0.284
0.185
0.270
0.222
0.242
0.290
0.214
0.208
0.240
0.263
0.221
0.181
0.323
0.172
0.259
0.196
0.230
0.322
0.369
0.311
0.325
0.355
0.486
0.378
0.213
0.311
0.203
0.298
0.314
0.180
0.324
0.210
0.387
0.339
0.358
0.278
0.278
0.244
0.197
0.404
0.317
0.388
0.147
0,220
0,412
0.318
0.226
0.217
0.357
0.382
0.298
0.289
0.387
0.206
0.294
0.215
0.243
0.236
0.276
0.333
0.265
0.276
0.282
0.299
0.220
0.204
0.352
0.206
0.272
0.184
0.228
0.353
0.331
0.316
0.348
0.362
0.497
0.429
0.322
0.278
0.235
0.241
0.228
0.298
0.233
0.277
0.329
0.228
0.259
0.214
0.307
0.263
0.180
0.377
0.210
0.329
0.217
0.436
0.461
0.322
0.357
0.301
0.374
0.295
0.389
0.411
0.273
0.290
0.283
0.356
0.308
0.285
0.461
0.246
0.271
0.315
0.240
0.290
0.323
0.379
0.388
0.407
0.525
0.474
0.214
0.245
0.235
0.234
0.262
0.246
0.254
0.175
0.293
0.278
0.269
0.234
0.196
0.204
0.119
0.317
0.256
0.378
0.166
0.250
0.388
0.414
0.233
0.209
0.337
0.367
0.240
0.403
0.362
0.333
0.267
0.295
0.358
0.196
0.306
0.314
0.190
0.227
0.213
0.219
0.206
0.236
0.361
0.209
0.277
0.336
0.394
0.302
0.228
0.354
0.408
0.358
0.335
0.385
0.239
0.272
0.229
0.272
0.228
0.301
0.348
0.243
0.263
0.259
0.299
0.238
0.224
0.368
0.213
0.274
0.208
0.248
0.348
0.326
0.330
0.328
0.390
0.478
0.412
81
68
41
50
83
106
47
0.261
19
0.283
0.338
0.254
0.214
0.230
0.313
0.181
0.278
0.298
0.252
0.312
0.241
0.264
0.191
0.204
0.329
0.250
0.331
0.281 0.219
0.279 0.271
0.313 0.40!
0.370 0.449
0.321 0.283
0.358 0.320
0.410 0.388
0.45 1 0.438
0.356 0.405
68
38
LeJ
Appendix Table A2.3. Continued
Location
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
1
0
0
1
1
0
4
1
2
3
3
3
1
0
0
2
0
0
3
2
1
3
2
3
1
5
3
2
1
0
0
2
0
0
36
0
37
1
2
1
3
3
3
3
3
3
4
4
2
4
6
6
4
2
1
3
4
6
4
5
0
0
1
1
2
2
1
2
0
2
0
0
0
0
0
0
2
1
6
1
0
0
2
1
1
2
1
1
1
1
1
0
1
1
1
1
1
3
2
1
3
2
3
1
1
0
2
0
2
1
3
1
3
2
4
2
0
0
1
0
2
2
7
1
2
1
3
2
8
0
0
2
3
1
3
1
2
2
1
4
3
2
2
3
3
2
0
0
2
I
1
1
1
0
0
1
1
1
0
0
1
1
3
2
2
2
2
2
2
3
1
0
0
0
2
1
1
3
1
1
1
1
0
1
2
2
0
1
1
1
0
2
1
3
0
4
2
1
0
1
1
2
1
0
0
0
0
1
1
1
0
1
1
2
2
2
2
1
2
1
1
2
4
1
2
2
1
3
2
1
0
0
1
2
2
0
4
0
0
0
0
0
0
0
0
2
0
0
0
0
0
1
1
2
2
5
3
3
3
5
3
2
1
2
2
1
3
2
2
1
1
0
0
0
2
1
0
0
0
0
1
0
1
0
3
1
0
2
4
1
2
4
4
3
1
0
1
1
0
1
1
1
0
0
1
0
0
1
1
2
2
0
0
1
1
0
1
2
1
0
0
0
0
3
1
3
2
2
1
4
4
2
0
0
0
0
0
2
0
1
0
2
0
1
0
0
0
1
1
1
2
1
1
0
2
1
1
1
0
0
0
1
1
0
0
2
3
1
1
1
10
0
0
11
1
12
2
13
0
14
2
15
18
2
2
0
0
19
1
9
16
17
20
0
1
0
0
0
0
210.256220.3040.310
-
23
0.340
0.331
0.441
-
24
0.296
0.249
0.441
0.336
25
0.348
0.191
0.352
0.437
0.317
-
26
0.305
0.267
0.413
0.398
0.268
0.320
27
0.372
0.299
0.401
0.519
0.372
0.251
0.430
-
28
0.318
0.243
0.362
0.482
0.347
0.203
0.401
0.163
1
1
0
0
0
0
0
0
0
0
0
2
1
2
2
1
1
3
0
2
1
3
1
1
0
0
0
0
0
0
0
1
0
2
1
2
3
0
0
3
1
0
0
1
4
1
1
0
0
2
0
1
1
1
0
0
0
0
2
0
1
1
3
N.)
N.)
Appendix Table A2.3. Continued
20
21
22
23
24
25
26
29
0.313 0.188 0.374 0.393 0.301 0.127 0.316
30
0.349 0.265 0.500 0.346 0.267 0.328 0.210
31
0.307 0.160 0.385 0.417 0.221 0.232 0.279
32 0.285 0.273 0.386 0.449 0.33 1 0.365 0.229
33 0.254 0.242 0.389 0.484 0.277 0.312 0.291
34 0.308 0.170 0,361 0.406 0.286 0.175 0.334
35
0.236 0.245 0.388 0.400 0.260 0.384 0.293
36 0.262 0.184 0.397 0.346 0.179 0.242 0.219
37 0.242 0.321 0.467 0.474 0.275 0.405 0.329
38 0.259 0.282 0.408 0.415 0.250 0.410 0.273
39 0.204 0.315 0.333 0.462 0.344 0.409 0.292
40 0.283 0.260 0.388 0.350 0.181 0.371 0.216
41
0.184 0.282 0.317 0.368 0.274 0.361 0.267
42 0.215 0.207 0.343 0.362 0.238 0.315 0.264
43 0.199 0.156 0.305 0.334 0.198 0.246 0.177
44 0.256 0.340 0,440 0.517 0.403 0.380 0.389
45
0.205 0.236 0.356 0.412 0.282 0.342 0.293
46 0.326 0.347 0.431 0.514 0.334 0.449 0.394
47 0.257 0.091 0.314 0.342 0.229 0.177 0.253
48 0.305 0.248 0.317 0.382 0.217 0.310 0.270
49 0.383 0.378 0.431 0.398 0.321 0.487 0.397
50 0.454 0.334 0.364 0.368 0.404 0.423 0.479
51
0.321 0.228 0.398 0.264 0.170 0.277 0.292
52 0.354 0.245 0.391 0.337 0.245 0.273 0.212
53 0.387 0.314 0.451 0,355 0.267 0.378 0.388
54
0.480 0.391 0.504 0.293 0.382 0.443 0.396
55
0.348 0.330 0.382 0.265 0.333 0.375 0,286
Total number of significant differences compared to all other
40
20
89
81
42
89
66
27
Location
28
29
0.221
0.393 0.320
0.215 0.243
0.378 0.352
0.295 0.315
0.139 0.202
0.375 0.349
0.258 0.224
0.338 0.377
0.399 0.405
0.381 0.411
0.408 0.372
0.341 0.334
0.289 0.278
0.286 0.230
0.323 0.345
0.315 0.338
0.372 0.456
0.241 0.164
0.335 0.325
0.514 0.488
0.455 0.440
0.355 0.256
0.397 0.308
0.438 0.358
0.576 0.426
0.481 0.365
0.265
0.456
0.282
0.382
0.300
0.196
0.383
0.275
0.370
0.406
0.397
0.419
0.335
0.283
0.328
0.316
0.347
0.361
0.253
0.341
0.545
0.467
0.378
0.450
0.504
0.615
0.497
populations
80
101
106
30
31
32
37
1
34
0
36
1
33
2
35
1
3
1
3
1
0
2
1
3
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
0.266
-
0.357
0.309
0.357
0.230
0.355
0.338
0.395
0.290
0.366
0.325
0.249
0.437
0.328
0.443
0.289
0.350
0.436
0.488
0.283
0.220
0.371
0.384
0.349
0.267
0.227
0.151
0.263
0.171
0.260
0.274
0.324
0.266
0.292
0.215
0.192
0.351
0.255
0.298
0.184
0.248
0.415
0.410
0.278
0.279
0.357
0.480
0.419
0.208
0.317
0.202
0.270
0.247
0.213
0.201
0.252
0.200
0.187
0.219
0.272
0.238
0.284
0.268
0.305
0.459
0.513
0.387
0.347
0.497
0.534
0.416
92
61
24
0.33 1
0.275
0.170
0.249
0.187
0.183
0.194
0.253
0.244
1
-
0.214
0.227
0.279
0.411
0.490
0.353
0.364
0.434
0.552
0.453
0.336
0.177
0.319
0.349
0.361
0.342
0.303
0.249
0.244
0.327
0.300
0.355
0.176
0.301
0.485
0.409
0.299
0.313
0.416
0.504
0.407
0.267
0.215
0.128
0.205
0.197
0.215
0.142
0.195
0.288
0.165
0.221
0.238
0.260
0.336
0.437
0.320
0.371
0.387
0.491
0.416
31
39
49
0.191
0.209
0.217
0.157
-
1
0
1
0
1
0.263
0.267
0.313
0.238
0.231
0.186
0.173
0,324
0.271
0.330
0.150
0.243
0.408
0.406
0.223
0.261
0.353
0.217
0.233
0.294
0.239
0.223
0.267
0.230
0.228
0.239
0.319
0.349
0.460
0.565
0.397
0.415
0.469
0.431 0.599
0.312 0.492
58
82
Appendix Table A2.3. Continued
Location
38
39
40
41
42
43
44
45
46
49
50
51
52
53
54
55
0
0
0
0
1
2
1
1
47
0
48
0
0
1
2
1
1
2
0
2
3
0
0
0
0
1
1
0
0
1
1
1
2
2
1
1
2
2
3
3
2
3
1
2
4
4
1
3
4
2
3
4
4
4
4
4
4
5
0
0
2
0
0
1
1
0
0
0
2
2
1
2
3
2
3
5
6
2
3
1
1
1
1
2
2
2
1
4
4
1
2
2
2
3
4
7
0
0
1
0
1
2
1
0
0
2
2
1
3
2
3
2
2
2
8
0
0
0
0
0
2
1
0
0
2
3
2
4
2
3
2
2
2
9
0
1
2
0
0
2
1
0
0
2
3
3
3
3
3
3
3
3
10
0
0
0
0
0
0
2
1
2
0
2
1
1
0
1
0
2
2
0
1
2
1
1
2
3
1
1
0
3
2
1
3
1
1
2
3
3
2
1
0
1
1
2
1
2
1
2
3
2
1
0
0
0
I
0
0
1
0
0
0
2
0
0
0
2
1
1
3
1
0
2
3
14
1
0
0
0
1
0
2
1
2
2
1
2
1
1
2
3
1
1
1
1
1
1
2
2
3
0
2
2
15
3
4
2
3
2
2
2
5
16
1
1
1
1
2
2
3
1
1
3
2
1
2
3
2
3
2
4
17
1
2
0
0
1
0
2
1
2
0
1
2
1
0
0
1
1
1
1
0
1
1
1
0
1
2
1
2
2
1
3
1
0
2
3
1
1
1
0
1
1
1
0
1
1
3
2
1
0
1
0
1
2
2
0
2
1
2
2
2
1
2
0
2
2
1
0
0
0
1
1
2
4
1
3
1
1
1
2
1
2
11
12
13
18
19
0
0
0
0
0
20
0
0
0
0
21
0
0
0
0
0
0
0
0
0
2
1
1
1
3
2
2
2
22
3
1
1
23
3
3
1
1
1
1
3
2
3
1
1
1
1
1
1
1
0
1
24
1
2
0
0
0
0
2
1
1
1
0
0
2
0
1
0
0
1
4
1
0
2
0
3
2
3
0
2
5
2
0
0
1
1
1
0
0
0
0
2
2
2
0
0
1
2
1
0
1
1
1
1
0
0
1
1
1
1
0
3
2
2
2
2
1
4
4
1
0
1
1
0
1
2
0
2
3
3
1
2
1
3
4
25
26
27
28
4
2
1
2
2
1
1
Appendix Table A2.3. Continued
38
39
40
41
29
30
3
3
4
2
1
2
2
31
2
2
1
32
33
34
0
0
0
0
0
0
1
1
1
0
0
0
0
0
35
36
37
38
0
0
2
0
2
1
0
1
1
0
0
0
390.166-
400.1390.244
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
Total
0.219
0.181
0.209
0.330
0.153
0.168
0.276
0.238
0.304
0.438
0.332
0.350
0.408
0.505
0.412
number
58
0
0
43
44
45
1
1
0
0
0
3
3
3
2
3
1
0
0
0
0
1
1
1
0
0
2
0
1
2
0
0
0
1
0
0
0
0
0
0
0
1
1
0
1
0
0
0
0
2
1
0
0
0
0
0
0
0
2
1
1
0
0
-
41
1
2
0
0.182 0.227
0
0
0
0.216 0.204 0.141
0
1
0.231 0.167 0.192 0.148
2
0.268 0.406 0.272 0.265 0.323
0.165 0.218 0.235 0.176 0.175 0.274
0.215 0.268 0.272 0.226 0.300 0.337
0.308 0.246 0.249 0.177 0.144 0.320
0.288 0.159 0.240 0.205 0.162 0.427
0.384 0.241 0.398 0.366 0.310 0.565
0.493 0.374 0.449 0.398 0.372 0.606
0.403 0.235 0.318 0.271 0.201 0.460
0.389 0.249 0.348 0.341 0.223 0.484
0.480 0.325 0.440 0.390 0.297 0.563
0.566 0.391 0.499 0.480 0.383 0.651
0.426 0.317 0.339 0.372 0.297 0.513
of significant differences compared to all other
61
Location
46
47
4
0
2
0
42
2
16
34
-
43
81
1
0
0
0
48
49
4
5
4
2
3
3
2
1
1
2
2
50
2
51
2
52
0
53
54
55
1
1
3
3
3
1
1
0
2
1
0
2
1
3
4
1
1
1
1
3
3
1
2
1
1
3
2
1
0
1
1
1
1
2
2
3
5
2
2
2
0
2
0
0
2
2
1
1
2
2
2
0
2
2
2
3
3
3
3
3
2
2
2
1
1
1
3
3
1
1
1
2
2
0
2
1
2
2
1
4
2
2
3
3
0
1
0
1
1
2
2
2
1
2
2
0
0
1
0
0
1
1
0
1
1
1
1
1
0
0
1
2
1
1
0
1
1
1
3
1
2
0
4
1
1
3
2
3
2
2
3
4
0
2
2
1
2
1
2
2
2
2
3
2
2
1
3
2
4
2
2
2
2
1
0
1
1
1
1
1
2
0
2
1
3
1
3
3
0
2
2
1
1
0
0
0
2
0
-
0.200
0.242 0.339
0.244 0.284
0.324 0.386
0.429 0.471
0.319 0.418
0.351 0.458
0.390 0.495
0.508 0.632
0.427 0.530
populations
53
76
0.218
0.378
0.336
0,202
0.257
0.322
0.394
0.305
0.252
0.304
0.207
0.270
0.304
0.407
0.334
0.319
0.298
0.373
0.264
0.390
0.400
0.308
0.400
0.342
0.395
0.386
48
97
102
97
1
2
0.223
0
0
0.201 0.310
0
0.268 0.282 0.283
0.261 0.248 0.362 0.255
68
79
62
1
2
1
1
-
0
94
124
cM
Appendix Table A3. 1. Summary of allele frequencies, sample sizes, polymorphic loci (P095), average number of alleles per locus,
and the expected proportion of heterozygotes (Weir 1996) at each location sampled. Duplicated loci are not included.
CKB*
Map
no.
Samplename
N
Bosewell Bay, AK
33
FJ3ALD2*
FBALD-1 *
100
97
N
100
84
G3PDH-1 *
N
100
N
-100
G3PDH-2 *
-10
1.000 0.000
33
1.000 0.000
33
1.000
33
1.000 0.000
GAPDH3*
GAPDH-2 *
N
-100
N
100
N
100
33
0.000
33
1.000 0.000
33
1.000
76
2
MartinRiver,AK
23
1.000 0.000
21
1.000 0.000
21
1.000
23
1.000 0.000
23
0.000
23
1.000 0.000
23
1.000
3
Gines Creek, AK
45
0.956 0.044
45
1.000 0.000
45
1.000
45
1.000 0.000
45
0.000
45
1.000 0.000
45
1.000
4
Suntaheen Creek, AK
17
1.000 0.000
17
1.000 0.000
17
1.000
17
1.000 0.000
17
0.000
17
1.000 0.000
17
1.000
5
Freshwater Bay, AK
IS
1.000 0.000
15
1.000 0.000
15
1.000
14
1.000
0.000
14
0.000
15
1.000 0.000
15
1.000
6
KIag Bay, AK
16
1.000 0.000
16
1.000 0.000
16
1000
16
1.000 0.000
16
0.000
16
1.000
0000
16
1.000
7
Portage Bay, AK
22
1.000 0.000
17
1.000 0.000
17
1.000
22
1.000 0.000
22
0.000
22
1.000
0.000
22
1.000
8
Duncan Salt Chuck, AK
43
1.000 0.000
48
1.000 0.000
48
1.000
50
1.000 0.000
50
0.000
47
1.000
0.000
47
1,000
9
KadakeCreek,AK
38
1.000 0.000
38
1.000 0.000
38
1.000
38
1.000 0.000
38
0.000
38
1.000 0.000
38
1.000
10
Eagle River, AK
30
1.000 0.000
31
1.000 0.000
31
1.000
31
1.000 0.000
31
0.000
31
1.000 0.000
31
1.000
11
Wolverine Creek, AK
50
1.000 0.000
50
1.000 0.000
50
1.000
50
1.000 0.000
50
0.000
50
1.000 0.000
50
1.000
12
Staney Creek, AK
51
0.922 0.078
51
1.000 0.000
51
1.000
51
1.000 0.000
51
0.000
51
1.000 0.000
51
1.000
13
Vixen Inlet, AK
12
1.000 0.000
12
1.000 0.000
12
1.000
12
1.000 0.000
12
0.000
12
1.000 0.000
12
1.000
14
Traiters Cove, AK
25
1.000 0.000
25
1,000 0.000
25
1.000
20
1.000 0.000
20
0.000
25
1.000 0.000
25
1.000
IS
59
33
1.000 0.000
0.978 0.022
58
29
1.000 0.000
1.000 0.000
58
29
1.000
1.000
60
16
Throne River, AK
Bakewell Creek, AK
34
1.000 0.000
1.000 0.000
60
34
0.000
0.000
60
34
1.000 0.000
1.000 0.000
60
34
1.000
1.000
17
Skeena River, BC
25
1.000 0.000
25
1.000 0.000
25
1.000
25
1.000 0.000
25
0.000
25
1.000 0.000
25
1.000
IS
TIell River, BC
20
1.000 0.000
19
1.000 0.000
19
1.000
20
1.000 0.000
20
0.000
20
1.000 0.000
20
1.000
1.000
19
Yakoun River, BC
16
1.000 0.000
16
1.000 0.000
16
1.000
16
1.000 0.000
16
0.000
16
1.000 0.000
16
20
25
1.000 0.000
25
1.000 0.000
25
1.000
25
1.000 0.000
25
0.000
25
1.000 0.000
25
1.000
21
BellaCoolaRiver,BC
BellaCoolaRiver,BC
13
1.000 0.000
13
1.000 0.000
13
1.000
13
0.962 0.038
13
0.021
13
1.000 0.000
13
1.000
22
San Josef River, BC
25
1.000 0.000
25
1.000 0.000
25
1.000
25
1.000 0.000
25
0.000
25
1.000 0.000
25
1.000
23
WillowCreek,BC
9
1.000 0.000
9
1.000 0.000
9
1.000
8
1.000 0.000
8
0.000
9
1.000 0.000
9
1.000
24
SakinawLake,BC
14
1.000 0.000
14
1.000 0.000
14
1.000
13
1.000 0.000
13
0.000
14
1.000 0.000
14
1.000
25
Fraser River, BC
29
1.000 0.000
29
1.000 0.000
29
1.000
29
1.000 0.000
29
0.000
29
1.000 0.000
29
1.000
26
Ritherdon Creek, BC
25
1.000 0.000
25
1.000 0.000
25
1.000
25
1.000 0.000
25
0.000
20
1.000 0.000
20
1.000
Appendix Table A3.1. Continued
CKB*
Map
no.
FBALD2*
FBALD-J *
G3PDH-1 *
G3PDH-2 *
GAPDH2*
GAPDH3*
32
Samplename
SandhillCreek,BC
KirbyCreek,BC
StillaguamishRiver,WA
Hoko River, WA
Salt Creek, WA
Quillayute River, WA
33
FIoh River, WA
13
1.000 0.000
13
1.000 0.000
13
1.000
13
1.000 0.000
13
34
HoodCanal,WA
20
1.000 0.000
19
1.000 0.000
19
1.000
16
1.000 0.000
16
35
25
1.000 0.000
25
1.000 0.000
25
1.000
25
1.000 0.000
25
36
Humptulips River, WA
McClane Creek, WA
32
1.000 0.000
32
1.000 0.000
32
1.000
32
1.000
1.000 0.000
1.000 0.000
1.000
24
24
1.000
24
24
1.000
38
Nehalem River, OR
Trask River, OR
Sand Creek, OR
SchoonerCreek, OR
Siletz River, OR
1.000 0.000
28
1.000 0.000
28
1.000
28
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
32
24
29
30
30
24
1.000
NaselleRiver,WA
30
24
1.000 0.000
37
28
1.000
29
1.000 0.000
29
1.000
1.000 0.000
30
1.000 0.000
30
1.000
29
1.000 0.000
29
1.000
30
30
21
19
1.000
21
21
1.000
1.000
1.000
10
10
10
1.000
22
1.000 0.000
22
1.000 0.000
1.000
22
1.000 0.000
22
22
1.000 0.000
22
1.000
30
1.000 0.000
26
1000 0.000
22
26
1.000
1.000
1.000
21
10
1.000 0.000
1.000 0.000
21
10
1.000 0.000
1.000 0.000
19
10
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
.000
1.000 0.000
30
1.000
20
1.000
1.000 0.000
30
1.000 0.000
1.000
1.000 0.000
29
1.000
24
24
1.000
25
1.000 0.000
25
1.000 0.000
1.000
25
1.000 0.000
25
1.000
23
23
1.000
21
1.000 0.000
20
1.000 0.000
30
25
20
1.000
21
1.000 0.000
21
1.000
21
0.975 0.025
1.000 0.000
0.935 0.065
1.000 0.000
20
30
30
29
28
1.000 0.000
28
1.000 0.000
28
1.000
28
1.000 0.000
28
1.000
28
33
1.000 0.000
33
0.985 0.015
33
1.000
30
1.000
1.000 0.000
26
1.000 0.000
26
1.000
30
23
1.000 0.000
26
1.000 0.000
23
1.000
26
24
1.000 0.000
26
1.000 0.000
26
1.000
26
1.000 0.000
26
1.000 0.000
22
1.000 0.000
22
1.000
22
1.000 0.000
27
1.000 0.000
27
1.000 0.000
27
1.000
27
12
1.000 0.000
12
1.000 0.000
12
1.000
9
1.000 0.000
9
1.000 0.000
9
31
1.000 0.000
28
1.000 0.000
28
27
28
29
30
31
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
Yaquina River, OR
Alsea River, OR
Cummins Creek, OR
Umpqua River, OR
New River, OR
Rogue River, OR
Winchuck River, OR
Lake Earl, CA
Klamath River, CA
Redwood Creek, CA
MillCreek,CA
WidowWhiteCreek,CA
1-lumboldtSt.Univ.,CA
N
100
97
23
1.000 0.000
23
1.000 0.000
23
1.000
22
-100 -10
1.000 0.000
26
1.000 0.000
26
1.000 0.000
26
1.000
26
1.000 0.000
26
53
1.000 0.000
52
1.000 0.000
52
1.000
53
53
21
1.000 0.000
18
1.000 0.000
18
1.000
21
24
1.000 0.000
24
1.000 0.000
24
1.000
24
1.000 0.000
1.000 0.000
1.000 0.000
24
12
1.000 0.000
9
1.000 0.000
9
1.000
12
1.000 0.000
12
N
100
84
N
100
N
N
-100
N
N
100
22
0.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
14
1.000 0.000
14
1.000
26
1.000 0.000
26
1.000
51
1.000 0.000
1.000 0.000
1.000 0.000
51
1.000
21
1.000
22
1.000
12
1.000
13
1.000 0.000
1.000 0.000
13
1.000
20
1.000 0.000
22
1.000 0.000
20
22
1.000
21
10
21
22
12
100
76
1.000
21
1.000
1.000 0.000
28
1.000
33
1.000 0.000
33
1.000
26
1.000 0.000
26
1.000
1.000
26
1.000 0.000
26
1.000
22
1.000
24
1.000 0.000
24
1.000
27
1.000
27
1.000 0.000
27
1.000
12
1.000 0.000
1.000 0.000
12
1.000
12
1.000 0.000
12
1.000
1.000
9
1.000 0.000
9
1.000
9
1.000 0.000
9
1.000
1.000
31
1.000 0.000
31
1.000
29
1.000 0.000
29
1.000
Appendix Table A 3.1. Continued
GAPDH4*
Map
no.
GAPDH5*
GDA1*
GDA2*
GPIA*
GP1B1*
Sample name
N
100
N
100
N
100
N
100
Bosewell Bay, AK
33
1.000
33
1.000
31
1.000 0.000
31
0.548 0.452
33
1.000 0.000
29
0.741
2
Martin River, AK
23
1.000
23
1.000
23
1.000 0.000
23
0.478
0.522
23
1.000 0.000
22
0.591 0.000 0.000
3
Gines Creek, AK
45
1.000
45
1.000
45
0.867
0.133
45
0.800 0.200
45
1.000 0.000
42
0.405
0.000 0.000 0.595
4
Suntaheen Creek, AK
17
1.000
17
1.000
17
0.824 0.176
17
0.824 0.176
17
1.000 0.000
16
0.969
0.000 0.000 0.031
5
FreshwaterBay, AK
15
1.000
15
1.000
15
0.767
0.233
15
0.833 0.167
15
1.000 0.000
10
0.850 0.000 0.000 0.150
6
Kiag Bay, AK
16
1.000
16
1.000
16
0.625 0.375
16
0.906 0.094
16
1.000 0.000
16
0.594 0.000 0.000 0.406
7
Portage Bay, AK
22
1.000
22
1.000
22
0.727 0.273
22
0.909 0.09 1
22
1.000 0.000
21
0.571
8
Duncan Salt Chuck, AK
50
1.000
50
1.000
49
0.918 0.082
49
0.592
0.408
50
1.000 0.000
49
0.827 0.000
0.000 0.173
9
Kadake Creek, AK
38
1.000
38
1.000
38
0.947 0.053
38
0.632 0.368
38
1.000 0.000
38
0.632 0.000
0.000 0.368
10
Eagle River, AK
31
1.000
31
1.000
30
0.800 0.200
30
0.850 0.150
31
1.000 0.000
31
0.677
0.000 0.000 0.323
II
Wolverine Creek, AK
50
1.000
40
1.000
50
0.980
0.020
50
0.580 0.420
50
0.920 0.080
49
0.796
0.000 0.000 0.204
12
Staney Creek, AK
43
1.000
38
1.000
51
0.608
0.392
51
0.941
0.059
51
1.000 0.000
49
0.714
0000 0.000 0.286
13
Vixen Inlet, AK
12
1.000
12
1.000
12
0.917 0.083
12
0.625 0.375
12
1.000 0.000
II
1.000 0.000 0.000 0.000
14
Traiters Cove, AK
25
1.000
25
1.000
25
0.840
0.160
25
0.860 0.140
19
1.000 0.000
19
0.711
0.000 0.000 0.289
15
Throne River, AK
59
1.000
59
1.000
60
0.900 0.100
60
0.717 0.283
60
1.000
0.000
59
0.839
0.000 0.000
16
Bakewell Creek, AK
33
1.000
33
1.000
34
0.912 0.088
34
0.735 0.265
34
1.000 0.000
33
0,833
0.000 0.000 0.167
17
Skeena River, BC
25
1.000
25
1.000
24
0.583 0.417
24
1.000 0.000
25
1.000 0.000
25
0.400 0.000 0.000 0.600
18
TIell River, BC
20
1.000
20
1.000
20
0.675 0.325
20
0.950 0.050
20
1.000 0.000
20
0.775
19
YakounRiver,BC
16
1.000
16
1.000
15
1.000 0.000
15
0.633 0.367
16
1.000 0.000
16
0.812 0.000 0.000 0.188
20
BeIla Coola River, BC
25
1.000
25
1.000
25
0.960 0.040
25
0.900 0.100
25
1.000 0.000
24
0.313 0.000 0.021 0.667
21
BellaCoolaRiver,BC
13
1.000
13
1.000
13
0.731 0.269
13
1.000 0.000
13
1.000 0.000
11
0.591
22
San Josef River, BC
25
1.000
25
1.000
25
0.720 0.280
25
0.900 0.100
25
1.000 0.000
25
0.660 0.000 0.000 0.340
23
WillowCreek,BC
9
1.000
9
1.000
9
0.722 0.278
9
0.944 0.056
9
1.000 0.000
9
0.611 0.000
24
SakinawLake,BC
14
1.000
14
1.000
11
0.682 0.318
11
1.000 0.000
14
1.000 0.000
14
0.464 0.000 0.143 0.393
25
Fraser River, BC
29
1.000
29
1.000
24
0.729 0.271
24
0.938
0.063
29
1.000 0.000
29
0.466 0.000 0.000 0.534
26
Ritherdon Creek, BC
25
1.000
25
1.000
25
0.520 0.480
25
1.000 0.000
25
1.000 0.000
25
0.420 0.000 0.000 0.580
27
Sandhill Creek, BC
23
1.000
23
1.000
23
0.804 0.196
23
0.761 0.239
23
1.000 0.000
19
0.605 0.000 0.053 0.342
115
N
100
87
N
100
90
160
42
138
0.000 0.000 0.259
0.409
0.000 0.000 0.429
0.161
0.000 0.000 0.225
0.000
0.000 0.409
0.056 0.333
Appendix Table A3.1. Continued
GAPDH4*
Map
GAPDH.5*
GDA1*
GDA2*
48
Samplename
KirbyCreek,BC
Stillaguainish River, WA
HokoRiver,WA
SaltCreek,WA
Quillayute River, WA
HohRiver,WA
Hood Canal, WA
Humptulips River, WA
McClane Creek, WA
NaselleRiver,WA
Nehalem River, OR
Trask River, OR
Sand Creek, OR
Schooner Creek, OR
SiletzRiver,OR
YaquinaRiver, OR
Alsea River, OR
Cummins Creek, OR
UmpquaRiver,OR
New River, OR
Rogue River, OR
49
WinchuckRive,OR
50
51
Lake Earl, CA
Klamath River, CA
52
RedwoodCreek,CA
27
1.000
27
1.000
27
53
Mill Creek, CA
Widow White Creek, CA
Humboldt St. Univ., CA
12
1.000
12
1.000
12
0.500 0.500
0.500 0.500
9
1.000
9
0.643
0.357
7
1.000
31
1.000
1.000
7
31
22
0.614 0.386
22
no.
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
54
55
N
100
N
100
N
26
1.000
26
1.000
26
100
115
N
GPIA*
100
87
N
GPJB1*
100
90
N
100
0.923 0.077
0.953 0.047
0.929 0.071
26
1.000 0.000
26
53
1.000 0.000
48
21
1.000 0.000
18
0.978 0.022
0.958 0.042
24
1.000 0.000
24
12
1.000 0.000
12
13
1.000 0.000
11
20
1.000 0.000
160
42
138
51
1.000
51
1.000
53
21
1.000
21
1.000
21
24
1.000
24
1.000
23
0.596 0.404
0.613 0.387
0.667 0.333
0.543 0.457
12
1.000
12
1.000
12
0.625 0.375
12
13
1.000
13
1.000
13
13
20
1.000
20
1.000
20
0.769 0.231
0.825 0.175
20
0.846 0.154
0.775 0.225
19
0.500
0.375
0.111
0.312
0.583
0.227
0.579
25
1.000
25
1.000
25
0.500 0.500
25
0.980 0.020
25
0.980 0.020
25
0.420 0.000 0.000 0.580
32
1.000
32
1.000
32
32
1.000 0.000
20
1.000
24
1.000
24
1.000 0.000
0.958 0.042
32
24
29
24
1.000 0000
0.275 0.000 0.125 0.600
0.458 0.000 0.000 0.542
1.000
29
1.000
29
29
1.000 0.000
29
0.500 0.000 0.000 0.500
1.000
21
1.000
30
30
0.750 0.250
29
29
0.707
21
1.000
21
1.000
20
20
21
21
0.524
10
1.000
1.000
10
1.000
10
10
1.000 0.000
1.000 0.000
10
22
1.000
18
18
0.944 0.056
22
30
30
1.000
30
30
1.000
1.000
28
1.000
1.000
0.914
1.000
1.000
0.977
1.000
1.000
24
29
30
0.516 0.484
0.604 0.396
0.500 0.500
0.900 0.100
0.350 0.650
0.400 0.600
0.694 0.306
0.429 0.571
0.500 0.000 0.000 0.500
0.977 0.000 0.000 0.023
0.500 0.000 0.000 0.500
0.944 0.000 0.000 0.056
25
1.000
25
15
21
1.000
21
1.000
1.000
28
1.000
28
1.000
33
1.000
1.000
28
33
26
26
.000
26
1.000
26
24
1.000
22
26
53
21
23
24
10
0.000
0.086
0.000
0.000
0.023
0.000
0.000
22
0.000 0.000 0.500
0.000 0.000 0.625
0.000 0.028
0.861
0.000 0.063 0.625
0.000 0.000 0.417
0.000 0.000 0.773
0.000 0.158 0.263
0.017 0.000 0.276
0.000 0.000 0.476
28
1.000 0.000
30
0.767 0.233
0.567 0.433
0.595 0.405
30
0.850 0.150
30
15
1.000 0.000
25
21
0.952 0.048
21
0.500 0.500
0.652 0.348
28
1.000 0.000
28
1.000 0.000
19
0.632 0.000 0.000 0.368
33
33
32
0.547 0.453
33
1.000
26
0.788 0.212
26
0.924 0.076
0.904 0.096
26
0.981 0.019
26
1.000
25
25
0.827 0.173
1.000
24
1.000 0.000
1.000 0.000
26
24
0.500 0.500
0.521 0.479
24
0.854 0.146
1.000 0.000
1.000 0.000
27
1.000 0.000
1.000 0.000
26
24
22
0.500 0.000 0.000 0.500
0.596 0.000 0.000 0.404
0.481 0.000 0.000 0.519
30
21
24
27
12
0.929 0.071
0.909 0.091
12
9
31
0.960 0.040
1.000 0.000
1.000 0.000
1.000 0.000
29
27
25
20
12
9
26
0.580 0.000 0.000 0.420
0.525 0.000 0.000 0.475
0.583
0.000 0.000 0.417
0.000 0.000 0.523
0.500 0.000 0.000 0.500
0.722 0.000 0.000 0.278
0.712 0.000 0.000 0.288
0.477
(J
Appendix Table A3. 1. Continued
Map
GPI-B2 *
no.
Samplename
N
1
Bosewell Bay, AK
33
2
MartinRiver,AK
3
mIDHP-1 *
100
110
mIDHp-2 *
N
100
N
1.000 0.000
33
1.000
23
1.000 0.000
23
Gines Creek, AK
45
1.000 0.000
4
SuntaheenCreek,AK
17
5
Freshwater Bay, AK
6
100
sIDHp-1,2 *
140
N
33
1.000 0.000
33
1.000
23
1.000 0.000
45
1.000
45
1.000 0.000
17
1.000
15
1.000 0.000
15
Kiag Bay, AK
16
1.000 0.000
7
PortageBay,AK
22
100
LDH-A] *
46
69
21
29
128
52
N
100
33
1.000
23
0.250 0.000 0.750 0.000 0.000 0.000 0.000
0.337 0.011 0.587 0.000 0.065 0.000 0.000
23
1.000
1.000 0.000
45
0.117 0.000
0.883 0.000 0.000 0.000 0.000
45
1.000
17
1.000 0.000
17
0.735 0.000 0.147 0.044 0.000 0.074 0.000
17
1.000
1.000
15
1.000 0.000
15
0.283
0.000 0.683 0.017
0.000 0.000 0.017
15
1.000
16
1.000
16
1.000 0.000
16
0.250
0.000 0.750 0.000 0.000 0.000 0.000
16
1.000
1.000 0.000
22
1.000
22
1.000 0.000
22
22
1.000
8
Duncan Salt Chuck, AK
50
1.000 0.000
40
1.000
40
1.000 0.000
50
0.284 0.000 0.693 0.000 0.000 0.023 0.000
0.265 0.000 0.753 0.000 0.000 0.000 0.000
50
1.000
9
KadakeCreek,AK
38
1.000 0.000
38
1.000
38
1.000 0.000
38
0.257 0.000
0.743 0.000 0.000 0.000 0.000
38
1.000
10
Eagle River, AK
31
1.000 0.000
31
1.000
31
1.000 0.000
31
31
1.000
11
Wolverine Creek, AK
50
1.000 0.000
50
1.000
50
1.000 0.000
50
50
1.000
12
51
1.000 0.000
51
1.000
51
1.000 0.000
51
51
1.000
13
StaneyCreek,AK
Vixenlnlet,AK
0.242 0.000 0.758 0.000 0.000 0.000 0.000
0.235 0.000 0.760 0.000 0.000 0.005 0.000
0.235 0.000 0.765 0.000 0.000 0.000 0.000
12
1.000 0.000
12
1.000
12
1.000 0.000
12
0.229 0.000 0.771 0.000 0.000 0.000 0.000
11
1.000
14
Traiters Cove, AK
19
1.000 0.000
25
1.000
25
1.000 0.000
25
0.240
0.000 0.730 0.000 0.020 0.010 0.000
20
1.000
IS
Throne River, AK
59
1.000
0.000
44
1.000
44
1.000 0.000
60
0.254
0.000 0.746
0.000 0.000 0.000 0.000
60
1.000
16
Bakewe!! Creek, AK
34
1000 0.000
34
1.000
34
1.000
0000
35
0.279
0.000 0.72!
0.000
0.000
34
I 000
17
Skeena River, BC
25
1.000 0000
25
1.000
25
1.000 0.000
25
0.260 0.000 0.720 0.000 0,010 0.010 0.000
25
1.000
IS
TIe!! River, BC
20
1.000 0.000
19
1.000
19
1.000 0.000
20
0.250
19
YakounRiver,BC
16
1.000 0.000
16
1.000
16
1.000 0.000
16
0.266 0.000
20
BellaCoolaRiver,BC
24
1.000 0.000
25
1.000
25
1.000 0.000
25
21
BellaCoolaRiver,BC
11
1.000 0.000
13
1.000
13
1.000 0.000
13
22
San Josef River, BC
25
1.000 0.000
25
1.000
25
1.000 0.000
25
23
WillowCreek,BC
SakinawLake,BC
9
1.000 0.000
9
1.000
7
1.000 0.000
24
14
1.000 0.000
13
1.000
13
1.000 0.000
25
FraserRiver,BC
29
0.948 0.052
29
1.000
29
26
Ritherclon Creek, BC
25
1.000 0.000
20
1.000
27
Sandhill Creek, BC
23
1.000 0.000
14
1.000
0.000 0.738
0.000 0.000
0.000 0.013 0.000 0.000
18
1.000
0.734 0.000 0.000 0.000 0.000
16
1.000
0.250 0.000 0.730 0.000 0.020 0.000 0.000
0.327 0.000 0.673 0.000 0.000 0.000 0.000
25
1.000
13
1.000
25
1.000
9
0.260 0.000 0.740 0.000 0.000 0.000 0.000
0.472 0.000 0.500 0.000 0.025, 0.000 0.000
9
1.000
13
0.250 0.000 0.731 0.000 0.019 0.000 0.000
14
1.000
1.000 0.000
29
0.259 0.000 0.612
0.000 0.129 0.000 0.000
29
1.000
20
1.000 0.000
25
25
1.000
14
1.000 0.000
23
0.260 0.000 0.590 0.000 0.150 0.000 0.000
0.250 0.000 0.739 0.000 0.011 0.000 0.000
23
1.000
C
Appendix Table A3. 1. Continued
Map
GPI-B2 *
no.
Samplename
N
28
26
29
Kirby Creek, BC
Stillaguamish River, WA
53
30
HokoRiver,WA
19
31
Salt Creek, WA
Quillayute River, WA
Hoh River, WA
24
HoodCanal,WA
HumptulipsRiver,WA
20
32
37
McClane Creek, WA
Naselle River, WA
38
NehalemRiver,OR
39
niIDHP-1 *
100
110
rnIDHp-2 *
N
100
N
1.000 0.000
26
1.000
44
21
24
29
0.962
1.000
1.000
0.958
1.000
1.000
1.000
1.000
0.958
1.000
30
42
Trask River, OR
Sand Creek, OR
Schooner Creek, OR
Siletz River, OR
43
100
sIDHp-1,2 *
LDFf-Al *
46
140
N
N
100
26
1.000 0.000
26
0.279 0.000 0.654 0.000 0.067 0.000 0.000
26
1.000
1.000
44
0.955 0.045
53
1.000
20
0.050
21
21
24
0.042
24
12
1.000
1.000
0.208
12
1.000
13
0.000
0.000
12
13
13
20
1.000
20
0.000
20
25
1.000
25
29
32
0.250 0.096 0.654 0.000 0.000 0.010 0.000
0.250 0.000 0.650 0.000 0.100 0.000 0.000
0.250 0.040 0.690 0.000 0.000 0.020 0.000
0.258 0.000 0.516 0.000 0.227 0.000 0.000
24
1.000
1.000
24
0.260 0.000 0.719 0.000 0.000 0.021 0.000
24
29
1.000
24
29
0.000
0.017
0.000
25
29
0.260 0.000 0.740 0.000 0.000 0.000 0.000
26
30
1.000
30
30
1.000 0.000
2!
1.000
21
0.250
0.275
30
2!
10
1.000 0.000
10
1.000
0
10
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
1.000
22
1.000 0.000
22
1.000
22
22
1.000
Yaquina River, OR
30
1.000 0.000
30
1.000
30
0.000
0.000
0.000
0.000
0.000
0.000
29
1.000 0.000
0.950
0.958
1.000
1.000
1.000
1.000
0.983
1.000
1.000
1.000
1.000
1.000
1.000
1.000
0.259 0.000 0.575 0.000 0.165 0.000 0.000
0.275 0.000 0.725 0.000 0.000 0.000 0.000
0.281 0.240 0.439 0.000 0.040 0.000 0.000
53
1.000
44
AlseaRiver,OR
27
1.000 0.000
1.000
30
1.000 0.000
29
45
Cummins Creek, OR
25
1.000 0.000
30
25
1.000
25
1.000 0.000
25
46
UmpquaRiver,OR
21
21
1.000
21
New River, OR
28
28
1.000
28
1.000 0.000
1.000 0.000
21
47
1.000 0.000
1.000 0.000
48
Rogue River, OR
33
33
1.000
33
1.000 0.000
32
49
WinchuckRiver,OR
26
1.000
26
1.000 0.000
25
50
Lake Earl, CA
26
26
1.000 0.000
1.000 0.000
26
1.000
26
1.000 0.000
26
51
Klamath River, CA
Redwood Creek, CA
Mill Creek, CA
Widow White Creek, CA
Humboldt St. Univ., CA
24
1.000 0.000
1.000 0.000
23
1.000
23
1.000 0.000
24
27
1.000 0.000
27
1.000
27
1.000 0.000
27
12
1.000 0.000
12
1.000
12
1.000 0.000
12
9
1.000 0.000
9
1.000
9
1.000 0.000
9
31
1.000 0.000
24
1.000
24
1.000 0.000
31
32
33
34
35
36
40
4!
52
53
54
55
12
13
25
24
0.038
0.000
0.000
0.042
0.000
0.000
0.000
0.000
0.042
0.000
12
20
10
22
30
28
100
69
21
29
128
52
0.000 0.77 1 0.000 0.021 0.000 0.000
0.017 0.717 0.000 0.000 0.017 0.000
0.000 0.725 0.000 0.000 0.000 0.000
0.300 0.000 0.675 0.000 0.025 0.000 0.000
0.26! 0.000 0.716 0.000 0.023 0.000 0.000
0.250 0.008 0.700 0.000 0.042 0.000 0.000
0.336 0.000 0.638 0.000 0.026 0.000 0.000
0.280 0.000 0.650 0.000 0.070 0.000 0.000
0.226 0.000 0.714 0.000 0.060 0.000 0.000
0.321 0.000 0.679 0.000 0.000 0.000 0.000
0.250 0.000 0.742 0.000 0.008 0.000 0.000
0.280 0.010 0.700 0.000 0.000 0.010 0.000
0.308 0.000 0.673 0.000 0.010 0.010 0.000
0.417 0.000 0.573 0.000 0.010 0.000 0.000
0.278 0.000 0.685 0.000 0.037 0.000 0.000
0.271 0.000 0.729 0.000 0.000 0.000 0.000
0.361 0.000 0.611 0.000 0.028 0.000 0.000
0.298 0.000 0.677 0.000 0.024 0.000 0.000
24
13
20
25
32
21
23
1.000
30
1.000
25
1.000
21
1.000
28
1.000
33
1.000
26
1.000
1.000
1.000
1.000
1.000
26
24
27
12
9
31
1.000
1.000
Appendix Table A3. 1. Continued
LDHA2*
Map
LDHB1*
LDHB2*
LDHCI*
no.
Sample name
N
100
N
100
N
1
BosewellBay,AK
33
1.000
33
1.000
33
2
MartinRiver,AK
23
1.000
23
1.000
23
3
Gines Creek, AK
45
1.000
45
1.000
45
4
Suntaheen Creek, AK
17
1.000
17
1.000
17
5
Freshwater Bay, AK
15
1.000
15
1.000
15
1.000 0.000
6
16
1.000
16
1.000
16
0.969 0.031 0.000 0.000
0.000
22
1.000
22
1.000
22
0.977 0.023 0.000 0.000
8
KlagBay,AK
PortageBay,AK
DuncanSaltChuck,AK
50
1.000
51
1.000
50
1.000 0.000 0.000 0.000
9
Kadake Creek, AK
38
1.000
38
1.000
10
Eagle River, AK
31
1.000
31
11
Wolverine Creek, AK
50
1.000
12
Staney Creek, AK
51
13
\'ixen Inlet, AK
14
7
100
76
119
108
85
N
0.000
33
1.000 0.000
1.000 0.000 0.000 0.000 0.000
23
1.000 0.000
0.978 0.022
0.000 0.000 0.000
45
1.000 0.000
0.912 0.088 0.000 0.000 0.000
17
1.000 0.000
0.000 0.000 0.000
15
1.000 0.000
16
1.000 0.000
0.000
22
1.000 0.000
0.000
50
1.000 0.000
38
1.000 0.000 0.000 0.000 0.000
38
1.000 0.000
1.000
31
1.000 0.000
0.000 0.000 0.000
31
1.000 0.000
50
1.000
50
1.000 0.000 0.000 0.000 0.000
50
1.000 0.000
1.000
51
1,000
51
1.000 0.000 0.000 0.000 0.000
51
1.000 0.000
11
1.000
12
1.000
12
0.917 0.083 0.000 0.000 0.000
12
1.000 0.000
Traiters Cove, AK
20
1.000
25
1.000
25
1.000 0.000 0.000
25
1.000 0.000
1.000 0.000 0.000 0.000
0.000 0.000
100
106
15
Throne River, AK
60
1.000
57
1.000
60
1.000 0.000
0.000 0.000 0.000
58
1.000 0.000
16
Balcewell Creek, AK
34
1.000
33
1.000
34
0.985 0.015 0.000 0.000 0.000
33
1.000 0.000
17
Skeena River, BC
25
1.000
25
1.000
25
1.000 0.000 0.000 0.000 0.000
25
1.000 0.000
18
TlellRiver,BC
18
1.000
19
1.000
20
1.000 0.000 0.000 0.000 0.000
19
1.000 0.000
19
Yakoun River, BC
16
1.000
16
1.000
16
1.000 0.000 0.000
0.000 0.000
16
1.000 0.000
20
25
1.000
25
1.000
25
1.000 0.000 0.000
0.000 0.000
25
1.000 0.000
21
BellaCoolaRiver,BC
BellaCoolaRiver,BC
13
1.000
13
1.000
13
1.000 0.000 0.000 0.000
0.000
13
1.000 0.000
22
San Josef River, BC
25
1.000
24
1.000
25
1.000 0.000 0.000
0.000 0.000
24
1.000 0.000
23
Willow Creek, BC
9
1.000
9
1.000
9
1.000 0.000 0.000 0.000 0.000
9
1.000 0.000
24
SakinawLajce,BC
14
1.000
14
1.000
14
1.000
0.000 0.000 0.000 0.000
14
1.000 0.000
25
Fraser River, BC
29
1.000
29
1.000
29
1.000 0.000 0.000 0.000 0.000
29
1.000 0.000
26
Ritherdon Creek, BC
25
1.000
25
1.000
25
1.000 0.000 0.000
0.000 0.000
25
1.000 0.000
27
Sandhill Creek, BC
23
1.000
20
1.000
23
1.000 0.000 0.000 0.000 0.000
20
1.000 0.000
L'.)
Appendix Table A3. 1. Continued
LDHA2*
Map
LDHBi*
LDHB2*
N
100
N
100
N
28
Sample name
Kirby Creek, BC
26
1.000
26
1.000
26
29
StillaguamishRiver,WA
53
1.000
53
1.000
51
30
Hoko River, WA
21
1.000
17
1.000
31
SaltCreek,WA
24
1.000
17
32
Quillayute River, WA
12
1.000
no.
0.917
12
1.000
12
0.000 0.000 0.000 0.083
0.958 0.042 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
25
1.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000
0.905 0.095 0.000 0.000
24
13
1.000
13
1.000
13
1.000
19
1.000
20
25
1.000
25
1.000
25
32
1.000
32
1.000
32
37
24
1.000
24
1.000
24
38
NehalemRiver,OR
26
1.000
29
1.000
29
39
Trask River, OR
30
1.000
30
1.000
30
40
SandCreek,OR
21
1.000
21
1.000
21
41
Schooner Creek, OR
Siletz River, OR
10
1.000
10
1.000
10
22
1.000
22
1.000
22
23
1.000
30
1.000
28
44
Yaquina River, OR
Alsea River, OR
30
1.000
30
Cummins Creek, OR
30
25
1.000
45
1.000
25
1.000
25
46
UmpquaRiver,OR
21
1.000
21
1.000
21
47
28
1.000
28
1.000
33
1.000
1.000
28
33
26
26
24
1.000
1.000
26
1.000
1.000
26
26
24
1.000
26
24
52
New River, OR
Rogue River, OR
Winchuck River, OR
Lake Earl, CA
Kiamath River, CA
Redwood Creek, CA
27
1.000
27
1.000
27
53
MillCreek,CA
12
1.000
12
1.000
12
54
Widow White Creek, CA
Humboldt St. Univ., CA
9
1.000
9
1.000
9
31
1.000
31
1.000
31
49
50
SI
55
1.000
N
24
20
48
85
1.000
HohRiver,WA
43
108
21
Hood Canal, WA
Humptulips River, WA
McClane Creek, WA
Naselle River, WA
42
119
26
34
36
76
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000
33
35
LDHC1*
100
33
1.000 0.000 0.000 0.000
0.977 0.000 0.000 0.023
0.964 0.036 0.000 0.000
1.000 0.000 0.000 0.000
0.960
1.000
1.000
0.758
0.692
0.942
0.000
0.000
0.000
0.000
0.000
0.000
0979 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
0.040
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.242
0.308
0000 0.058
0.000 0,021
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
100
106
53
1.000 0.000
1.000 0.000
21
1.000 0.000
24
1.000 0.000
1.000 0.000
12
13
19
32
29
30
21
10
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
30
1.000 0.000
1.000 0.000
30
1.000 0.000
25
1.000 0.000
1.000 0.000
22
21
28
33
1.000 0.000
1.000 0.000
26
0.981
26
1.000 0.000
24
1.000 0.000
27
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
12
9
31
0.000
Appendix Table A3. 1. Continued
Map
MDH-A 1,2 *
no.
Samplename
N
1
BosewellBay,AK
33
100
MDH-B1,2 *
56
niMEP-1 *
2
Martin River, AK
23
3
GinesCreek,AK
45
115
84
39
140
20
0.879 0.121 0.000 0.000 0.000 0.000 0.000
0.728 0.272 0.000 0.000 0.000 0.000 0.000
0.706 0.294 0.000 0.000 0.000 0.000 0.000
4
Suntaheen Creek, AK
17
1.000 0.000 0.000 0.000
0.000 0.000 0.000
17
0.868 0.000 0.000 0,132
17
1.000 0.000 0.000
5
FreshwaterBay,AK
15
15
1.000 0.000 0.000 0,000
15
1.000 0.000 0.000
0.000 0.000
16
0.969
0.000 0.011 0.000
22
1.000 0.000 0.000
N
100
88
115
83
N
100
115
62
33
0.985 0.000 0.015 0.000
33
1.000 0.000 0.000
23
0.978 0.000 0.022 0.000
23
1.000 0.000 0.000
45
1.000 0.000 0.000 0.000
45
1.000 0.000 0.000
6
KIag Bay, AK
16
0.867 0.133 0.000 0.000 0.000 0.000 0.000
0.844 0.156 0,000 0.000 0.000 0.000 0.000
16
1.000 0.000
7
PortageBay,AK
22
0.909 0.091
0.000 0.000
22
0.989
8
Duncan Salt Chuck, AK
50
0.980 0.005
0.015 0.000
50
0.990 0.010 0.000
Kadake Creek, AK
38
38
1.000 0.000
0.000 0.000
38
1.000 0.000 0.000
10
EagleRiver,AK
31
0.840 0.160 0.000 0.000 0.000 0.000 0.000
0.895 0.105 0.000 0.000 0.000 0.000 0.000
0.935 0.065 0.000 0.000 0.000 0.000 0.000
50
9
31
0.984 0.000 0.016 0.000
31
1.000 0.000 0.000
11
Wolverine Creek, AK
49
0.842
0.158 0.000 0.000 0.000 0.000 0.000
46
1.000 0.000 0.000
0.000
50
1.000 0.000
51
0.725
0.275 0.000 0.000 0.000 0.000 0.000
0.938 0.063 0.000 0.000 0.000 0.000 0.000
51
0.956 0.000 0.015
0.029
51
0.980 0.010 0.010
12
1.000 0.000
0.000 0.000
12
1.000 0.000 0.000
0.880 0.120 0.000 0.000 0.000 0.000 0.000
0.758 0.242 0.000 0.000 0.000 0.000 0.000
0.879 0.121 0.000 0.000 0.000 0.000 0.000
25
0.980 0.000 0.020 0.000
20
1.000 0.000 0.000
60
0.992
35
1.000 0.000
25
0.950
12
Staney
Creek,
AK
0.000 0.000 0.000
13
Vixen Inlet, AK
12
14
Traiters Cove, AK
25
15
Throne River, AK
60
16
BakewellCreek,AK
35
17
SkeenaRiver,BC
25
18
TIellRiver,BC
20
19
Yakoun River, BC
16
0.680 0.320 0.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000 0.000 0.000
0.938 0.031 0.031 0.000 0.000 0.000 0.000
20
BellaCoolaRiver,BC
25
1.000 0.000
21
Bella Coola River, BC
13
1.000 0.000 0.000 0.000
22
San Josef River, BC
25
0.750 0.250
23
Willow Creek, BC
9
24
Sakinaw Lake, BC
14
25
FraserRiver,BC
26
27
0.000 0.000 0.008
0.031 0.000
0.000
59
1.000 0.000 0.000
0.000 0.000
34
1.000 0.000 0.000
0.040 0.010 0.000
24
1.000 0.000
19
1.000 0.000 0.000 0.000
15
1.000 0.000 0.000
16
1.000 0.000
0.000 0.000
16
1.000 0.000 0.000
23
0.913 0.087
0.000 0.000
25
1.000 0.000 0.000
13
0.808 0.192
0.000 0.000
13
1.000
0.000 0.000
0.000
25
1.000 0.000
0.000 0.000
25
1.000
0.000 0.000
9
0.9 17 0.083
0.000 0.000
9
1.000 0.000
0.000
14
1.000 0.000
0.000 0.000
9
1.000 0.000
0.000
29
0.972 0.028 0.000 0.000 0.000 0.000 0.000
0.821 0.143 0.000 0.000 0,036 0.000 0.000
1.000 0.000 0.000 0.000 0.000 0.000 0.000
29
0.897 0.043
0.060 0.000
26
1.000 0.000
0.000
Ritherdon Creek, BC
25
1.000 0.000 0.000 0.000 0.000 0.000 0.000
25
0.970 0.000 0.030 0.000
25
1.000 0.000
0.000
Sandhill Creek, BC
23
1.000 0.000 0.000 0.000 0.000 0.000 0.000
23
0.935
23
1.000 0.000 0.000
0.000 0.000 0.000 0.000 0.000
0.000 0.000 0.000
0.000 0.000 0.000 0.000
0.065 0.000 0.000
0.000
Appendix Table A3. 1. Continued
MDHAI.2*
Map
26
29
Sample name
Kirby Creek, BC
Stillaguamish River. WA
30
HokoRiver,WA
21
31
Salt Creek, WA
Quillayute River, WA
24
HohRiver,WA
HoodCanal,WA
HumptulipsRiver,WA
13
no:
28
32
33
34
35
36
N
53
12
20
25
32
40
McClane Creek, WA
Naselle River, WA
Nehalem River, OR
Trask River, OR
Sand Creek, OR
41
SchoonerCreek,OR
10
42
22
44
Siletz River, OR
Yaquina River, OR
Alsea River, OR
45
CumminsCreek,OR
25
46
21
49
Umpqua River, OR
New River, OR
Rogue River, OR
Winchuck River, OR
50
LakeEarl,CA
26
51
Kiamath River, CA
24
52
RedwoodCreek,CA
27
53
Mill Creek, CA
Widow WhiteCreek,CA
Humboldt St. Univ., CA
12
37
38
39
43
47
48
54
55
24
29
30
21
30
30
28
33
26
9
31
100
MDHBJ,2*
56
115
84
39
140
1.000 0.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000 0.000 0.000 0.000
0.881 0.119 0.000 0.000 0.000 0.000
0.990 0.010 0.000 0.000 0.000 0.000
0.938 0.042 0.021 0.000 0.000 0.000
0.808
1.000
1.000
1.000
1.000
0.192
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.957
0.043
1.000 0000 0.000
0.964 0.000 0.036
0.975 0.015 0.025
0.966 0.011 0.000
0.958 0.000 0.042
0.900 0.075 0.000
1.000 0.000 0.000
0.929 0.000 0.071
1.000 0.000 0.000
1.000 0.000 0.000
0.990 0.000 0.010
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
0.992 0.008 0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
N
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
26
1.000 0.000
53
0.934
0.867
0.979
0.958
1.000
0.813
0.000
0.000
0.000 0000
0.023 0.000
0.000 0.000
21
24
12
13
20
100
21
0.980
0.609
0.990
0.948
0.908
0.952
0.900
0.886
0.917
0.958
0.930
0.929
28
0.99 1
33
1.000
26
0.971
26
1.000
24
0.917
27
1.000
12
1.000
9
1.000
31
1.000
25
32
24
29
30
21
10
22
30
0.025 0.000
30
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
25
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
mMEPJ*
20
88
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.045
0.000
0.033
0.000
0.000
0.009
0.000
0.010
0.000
0.000
0.000
0.000
0.000
0.000
115
83
N
0.000
0.000
0.000
0.000
0.000
0.000
0.066
0.131
0.021
0.042
0.000
0.188
0.020
0.336
0.000
0.000
0.067
0.024
0.000
0.011
0.025
0.000
0.060
0.071
0.000
0.000
0.019
23
0.000
0.000
0.000
0.055
0.010
0.052
0.025
0.024
0.100
0.057
0.058
0.008
0.010
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.083
0.000
0.000
0.000 0.000
0.000 0.000
100
115
62
1.000 0.000 0.000
53
1.000 0.000 0.000
21
1.000 0.000
24
1.000 0.000 0.000
0.000
12
1.000 0.000 0.000
13
1.000 0.000 0.000
20
1.000 0.000 0.000
23
1.000 0.000 0.000
32
1.000 0.000 0.000
24
1.000 0.000 0.000
10
1 000 0.000
31)
1.000
21
10
1.000
1.000
22
1.000
30
1.000
30
1.000
20
12
1.000
0.952
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
1.000 0.000
9
1.000 0.000
31
1.000 0.000
21
28
28
23
26
24
27
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.048
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
Ui
Appendix Table A3.1. Continued
Map
sMEPI*
sMEP2*
no.
Sample name
N
I
Bosewell Bay, AK
33
0.970 0.030 0.000
2
Martin River, AK
23
0.913 0.087
3
Gines Creek, AK
45
4
Suntaheen Creek, AK
17
0.882
5
Freshwater Bay, AK
15
6
KIag Bay, AK
7
100
88
97
N
PEPA1*
100
116
90
N
PGDH*
100
112
N
100
110
83
94
90
33
0.000 1.000 0.000
33
0.030 0.970
33
1.000 0.000 0.000 0.000 0.000
0.000
23
0.087
0.783 0.130
23
0.174 0.826
23
0.913
1.000 0.000 0.000
43
0.000
1.000 0.000
45
0.067 0.933
45
1.000 0.000 0.000 0.000 0.000
0.087
0.000 0.000 0.000
0.118 0.000
17
1.000 0.000 0.000
17
1.000 0.000
17
1.000 0.000 0.000 0.000 0.000
1.000 0.000 0.000
IS
0.133 0.867 0.000
15
0.067 0.933
15
1.000 0.000 0.000 0.000 0.000
16
0.969
0.000
16
0.063 0.938 0.000
16
0.063 0.938
16
1.000 0.000
0.000
Portage Bay, AK
22
1.000 0.000 0.000
21
0,000
1.000 0.000
22
0.045 0.955
22
0,977 0.023
0.000 0.000 0.000
8
Duncan Salt Chuck, AK
45
1.000
0.000 0.000
50
0.020 0.980 0.000
48
0.010 0.990
50
0.970 0.020 0.010 0.000 0.000
9
Kadake Creek, AK
38
1.000 0.000 0.000
38
0.000 1.000 0.000
38
0.013
0.987
38
0.526 0.237 0.158 0.079 0.000
10
EagleRiver,AK
30
1.000 0.000 0.000
31
0.000
1.000
0.000
31
0.016 0.984
31
0.952
0.000 0.048
11
Wolverine Creek, AK
50
1.000 0.000 0.000
50
0.000
1.000 0.000
50
0.050
0.950
49
0.929
0.071
12
Staney Creek, AK
44
0.898 0.102 0.000
51
0.000
1.000 0.000
51
0.039
0.961
51
0.873
0.069 0.059
13
Vixenlnlet,AK
12
1.000 0.000 0.000
12
0.000
1.000 0.000
12
0.042 0.958
12
0.875
0125 0.000 0.000 0.000
14
Traiters Cove, AK
20
1.000 0.000 0.000
23
0.065 0.935 0.000
25
0.020 0.980
25
0.840 0.160 0.000 0.000 0.000
15
Throne River, AK
52
0.971
0.029 0.000
60
0.000
1.000 0.000
58
0.034 0.966
49
0.990 0.010 0.000 0.000 0.000
16
Bakewell Creek, AK
34
1.000 0.000 0.000
34
0.191 0.809 0.000
33
0.045 0.955
34
0.897 0.103 0.000 0.000 0.000
17
Skeena River, BC
25
1.000 0.000 0.000
22
0.023 0.977 0.000
25
0.060 0.940
25
1.000 0.000
18
Tie!! River, BC
13
1.000 0.000 0.000
20
0.000 1.000 0.000
20
0.050 0.950
19
0.447
19
Yalcoun River, BC
14
1.000 0.000 0.000
16
0.063 0.938 0.000
16
0.063 0.938
16
0.750 0.250 0.000 0.000 0.000
20
BelIa Coola River, BC
25
1.000 0.000 0.000
25
0.040 0.960 0.000
23
0.087 0.913
25
1.000 0.000 0.000 0.000 0.000
21
BellaCoolaRiver,BC
13
1.000 0.000 0.000
13
0.269 0.731 0.000
13
0.115 0.885
13
1.000 0.000 0.000 0.000 0.000
22
San Josef River, BC
25
1.000 0.000 0.000
25
0.020 0.980 0.000
25
0.020 0.980
17
1.000 0.000 0.000 0.000 0.000
23
WillowCreek,BC
9
1.000 0.000 0.000
9
0.389 0.611
9
0.167 0.833
4
1.000 0.000 0.000 0.000 0.000
24
SakinawLake,BC
11
1.000 0.000 0.000
12
0.042
0.958 0.000
14
0.143
0.857
14
1.000 0.000 0.000 0.000 0.000
25
29
1.000 0.000 0.000
25
0.000 1.000 0.000
29
0.000 1.000
29
0.897 0.069
26
FraserRiver,BC
RitherdonCreek,BC
25
1.000 0.000 0.000
25
0120 0.880 0.000
25
0.040 0.960
15
1.000 0.000 0.000 0.000 0.000
27
Sandhill Creek, BC
23
1.000 0.000
23
0.000
20
0.075
14
1.000 0.000 0.000 0.000 0.000
0.031
0.000
0.000
1.000 0.000
0.925
0.000 0.000
0.000 0.000
0.000 0.000 0.000
0.000 0.000
0.000 0.000 0.000
0.553 0.000 0.000 0.000
0.034 0.000 0.000
Appendix Table A3. 1. Continued
Map
no.
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
sMEP1*
Sample name
N
Kirby Creek, BC
Stillaguamish River, WA
26
Hoko River, WA
Salt Creek, WA
Quillayute River, WA
Hoh River, WA
Hood Canal, WA
Humptulips River, WA
McClane Creek, WA
Naselle River, WA
Nehalem River, OR
21
53
24
12
13
20
18
32
21
20
Trask River, OR
Sand Creek, OR
Schooner Creek, OR
Siletz River, OR
Yaquina River, OR
Alsea River, OR
Cummins Creek, OR
Umpqua River, OR
30
New River, OR
Rogue River, OR
Winchuck River, OR
Lake Earl, CA
Klamath River, CA
Redwood Creek, CA
Mill Creek, CA
Widow White Creek, CA
Humboldt St. Univ., CA
28
20
10
22
30
30
22
21
27
23
26
24
27
12
8
31
sMEP2*
100
88
97
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
100
26
0.058
0.038
0.000
0.042
0.000
0.000
0.000
0.060
0.000
0.042
0.000
0.000
0.000
0.150
0.000
0.033
53
21
1.000 0.000 0.000
1.000 0.000 0.000
24
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
1.000 0.000 0.000
13
12
20
25
32
24
0.825 0.000 0.175
0.983 0.017 0.000
0.900 0.100 0.000
1.000 0.000 0.000
24
0.000
0.000
0.000
0.045
0.000
0.000
0.000
0.000
0.000
0.067
0.000
0.000
0.000
0.000
0.000
0.000
0.673 0.327 0.000
1.000 0.000 0.000
0.685 0.3 15 0.000
1.000 0.000 0.000
22
0.813 0.188
0.548 0.452
9
1.000
0.933
1.000
0.955
1.000
1.000
1.000
1.000
0.000
0.000
PEPA1*
N
30
21
10
30
29
22
21
25
31
26
0.000
0.000
0.000
0.060
0.016
0.019
112
N
0.942 0.000
26
0.385
22
0.962 0.000
53
20
30
0.038
0.150
0.021
0.000
0.200
0.150
0.060
0.047
0.000
0.091
0.100
0.000
0.050
0.045
0.000
0.615
0.962
0.850
0.979
1.000
0.800
0.850
0.940
0.953
1.000
0.909
30
0.017
21
1.000 0.000
0.981
15
1.000
26
0.98 1
0.938
1.000
1.000
24
1.000
27
1.000
12
0.024
0.024
0.036
0.000
0.019
0.000
0.063
0.000
0.000
100
0.909
0.850
1.000
0.833
0.958
1.000
1.000
1.000
1.000
1.000
0.966
1.000
0.975
0.850
0.909
0.933
0.883
0.940
0.929
0.964
1.000
0.933
12
1.000
9
0.611 0.389
9
1.000
24
0.063
31
1.000
0.958 0.000
1.000 0.000
24
1.000 0.000
13
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
20
1.000
0.940
1.000
0.958
1.000
1.000
1.000
0.850
1.000
0.967
1.000
1.000
1.000
0.940
0.984
0.981
1.000
24
27
0.037
0.963
12
0.000 1.000
0.333 0.667
0.196 0.804
28
PGDH*
100
0.000
0.063
26
90
N
116
0.938
12
25
32
24
22
30
21
10
22
21
28
33
26
26
24
27
0.900
1.000
0.950
0.955
1.000
0.983
0.976
0.976
0.964
1.000
0.938
50
21
24
12
13
20
25
32
25
29
26
20
10
22
30
30
25
21
28
33
110
83
0.000 0.09 1
0.100 0.050
0.000 0.000
0.000 0.167
0.000 0.042
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.034
0.000 0.000
0.025 0.000
0.000 0.150
0.000 0.091
0.000 0.067
0.000 0.117
0.000 0.000
0.000 0.071
0.000 0.036
0.000 0.000
0.067 0.000
0.000 0.019
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.000
0.000 0.000
94
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.060
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
90
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
Appendix TaNe A3. 1. Continued
Map
no.
sSOD*
Polymorphic loci
0.95
Alleles/locus
Sample name
N
100
145
49
number
percent
BosewellBay,AK
33
0.773
0.227
0.000
3
11.1
1.2
0.051
2
Martin River, AK
23
0.543
0.457
0.000
7
25.9
1.3
0.093
3
Gines Creek, AK
45
1.000
0.000
0.000
4
14.8
1.2
0.048
4
Suntaheen Creek, AK
17
0.618
0.382
0.000
5
Freshwater Bay, AK
15
0.733
0.267
0.000
6
22.2
1.2
0.063
6
Klag Bay, AK
16
0.969
0.031
0.000
5
18.5
1.3
0.061
7
Portage Bay, AK
22
0.932
0.068
0.000
4
14.8
1.3
0.063
8
Duncan Salt Chuck, AK
50
0.710
0.290
0.000
4
14.8
1.3
0.054
9
Kadake Creek, AK
38
0.934
0.066
0.000
5
18.5
1.3
0.068
10
Eagle River, AK
31
0.935
0.065
0.000
4
14.8
1.2
0.047
11
Wolverine Creek, AK
50
0.890
0.110
0.000
6
22.2
1.3
0.053
12
Staney Creek, AK
51
0.951
0.049
0.000
6
22.2
1.4
0.066
13
Vixen Inlet, AK
12
0.792
0.208
0.000
5
18.5
1.3
0.054
14
Traiters Cove, AK
25
0.720
0.280
0.000
6
22.2
1.3
0.066
IS
Throne River, AK
60
0.675
0.325
0.000
4
14.8
1.3
0.054
16
Bakewell Creek, AK
34
0.647
0.353
0.000
6
72.2
1.3
0.075
17
Skeena River, BC
25
0.920
0.080
0.000
4
14.8
1.2
0.048
HE
18
Tiell River, BC
20
0.975
0.025
0.000
5
18.5
1.2
0.058
19
Yakoun River, BC
16
0.969
0.031
5
18.5
1.2
0.051
20
25
0.960
0.040
3
11.1
1.3
0.039
21
BellaCoolaRiver,BC
BellaCoolaRiver,BC
0.000
0.000
13
0.962
0.038
0.000
4
14.8
1.2
0.059
22
San Josef River, BC
25
0.860
0.140
0.000
4
14.8
1.2
0.051
23
Willow Creek, BC
9
0.778
0222
0.000
6
22.2
1.3
0.090
24
Sakinaw Lake, BC
14
0,536
0.464
0.000
4
14.8
1.2
0.070
25
Fraser River, BC
29
0.517
0.483
0.000
6
22.2
1.3
0.068
26
Ritherdon Creek, BC
25
0.940
0.060
0.000
4
14.8
1.2
0.052
27
Sandhill Creek, BC
23
0.804
0.196
0.000
5
18.5
1.2
0.062
Appendix Table A3. 1. Continued
Map
sSOD*
Polymorphic loci
095
Alleles/locus
no.
Sample name
N
100
145
49
number
percent
28
26
0.081
4
14.8
1.4
5
18.5
1.2
4
14.8
1.4
2
7.4
1.2
13
3
11.1
1.2
34
Hood Canal, WA
Humptulips River, WA
20
5
18.5
1.2
25
0.900
0.940
0.070
0.044
0.066
0.049
0.048
0.062
35
5
18.5
1.3
36
McClaneCreek,WA
32
0.891
3
11.1
1.2
37
24
0.792
3
11.1
1.2
29
0741
5
18.5
1.3
30
0.833
6
22.2
1,3
21
18.5
1.2
6
22.2
1.3
42
SiletzRiver,OR
22
0.905
0.850
0.977
5
41
Naselle River, WA
Nehalem River, OR
Trask River, OR
Sand Creek, OR
Schooner Creek, OR
3
11.1
1.3
43
Yaquina River, OR
Alsea River, OR
Cummins Creek, OR
Umpqua River, OR
30
1.000
4
14.8
1.3
30
0.617
0.979
5
18.5
1.2
4
14.8
1.3
3
11.1
1.3
New River, OR
Rogue River, OR
Winchuck River, OR
Lake Earl, CA
Klamath River, CA
Redwood Creek, CA
26
0.952
0.750
4
14.8
1.2
33
0.985
5
18.5
1.3
26
0.808
0.615
0.958
0.383
0.021
0.048
0.250
0.015
0.192
0.385
0.042
6
1.4
6
22.2
22.2
5
18.5
1.3
14.8
1.2
12
0.130
0.292
0.222
0.097
4
Miii Creek, CA
Widow White Creek, CA
Humboldt St. Univ., CA
0.870
0.708
0.778
0.903
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.017
0.000
0.000
0.100
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
0.000
1.3
HohRiver,WA
0.154
0.170
0.000
0.042
0.000
0.000
0.100
0.060
0.109
0.208
0.241
0.167
0.095
0.050
0.023
0.000
2.5.9
33
0.846
0.830
1.000
0.958
1.000
1.000
7
32
Kirby Creek, BC
Stillaguamish River, WA
Hoko River, WA
Salt Creek, WA
Quillayute River, WA
3
11.1
1.1
7
25.9
25.9
1.3
29
30
31
38
39
40
44
45
46
47
48
49
50
51
52
53
54
55
53
21
24
12
10
24
21
26
24
27
9
31
7
HE
1.3
1.3
0.053
0.052
0.058
0.072
0.061
0.058
0.067
0.036
0.054
0.054
0.061
0.056
0.059
0.077
0.076
0.089
0.060
0.065
0.055
0.101
0.081
250
Appendix Table A3.2. Summary of loci that tested significant for deviations from
Hardy-Weinberg expectations based on exact test and x2 test for deficiency or excess
of heterozygotes. Location names correspond to those listed on Table 3.1. Exact test
P-values with asterisk (*) indicated the 20 tests that were significant after correction
for multiple comparisons using the sequential Bonferroni correction.
Exact test
Location
Heterozygote deficiency/excess
Locus
P
0.000*
28
15.4
0.824 21.02 0.000
0.000
22
11.5
0.9 17
Martin River
GDA2*
GDA2*
sMEP2*
0.000
4
8.3
-0.521 10.47 0.001
Martin River
sSOD
0.001
5
11.4
-0.562 7.26 0.007
Gines Creek
0.000*
0
3.8
-1.000 45.00 0.000
0.034
0
1.9
-1.000 15.00 0.000
Freshwater Bay
CKB*
PEPA1*
sMEP2*
0.003
0
3.5
-1.000 15.00 0.000
Portage Creek
GPI-B1 *
16
10.3
0.556
6.48 0.011
Duncan Salt Chuck
GDA2*
sMEP2*
GDA2*
0.026
0.000*
38
23.8
0.605
17.95 0.000
0.009
0
2.0
-1.000 50.00 0.000
0.000
28
17.7
0.583
0.005
16
24.2
-0.338 10.31 0.001
0.003
0.000*
36
24.4
0.478
40
24.3
0.645 21.23 0.000
0.108
9
5.6
0.600
0.045
0.072a
0
1.8
-1.000 12.00 0.001
1
2.8
-0.643
9.52 0.002
0.002
34
24.4
0.395
9.38 0.002
0.017
0
2.0
-1.000 33.00 0.000
0.076'
18
13.2
0.360
0.000*
3
10.5
-0.7 15 17.37 0.000
0.000
18
13.2
0.360
0.002
0.06Y*
20
12.0
0.667 11.11 0.001
13
8.8
0.48 1
4.64 0.03 1
0.09Y'
o.057
11
7.0
0.579
5.03 0.025
2
4.9
-0.590
5.56 0.018
0.036
15
11.0
0.366
3.68 0.055
0.019
0
1.9
Bosewell Bay
Martin River
Freshwater Bay
Duncan Salt Chuck
Kadake Creek
Kadake Creek
Wolverine Creek
PGDH
GDA2*
Staney Creek
GDA1*
Vixen Inlet
GDA2*
LDHB2*
Vixen Inlet
Traiters Cove
Thorne River
Bakewell Creek
Bakewell Creek
Bakewell Creek
Skenna River
SkeenaRiver
Tiell River
Yakoun River
Yakoun River
BellaCoolaRiver
Bella Coola River
sMEP2*
GDA2*
CKB*
GDA2*
sMEP2*
GDA1*
GPIB1*
GDAJ*
GDA2*
GPIB1*
GPIB1*
sMEP2*
Observed Expecte
D
X2
P
19.33 0.000
12.93 0.000
11.42 0.001
4.32 0.038
4.41
4.41
0.036
0.036
-1.000 25.00 0.000
251
Appendix Table A3.2.
Continued
Exact test
Location
Heterozygote deficiency/excess
Locus
P
Observed
Expected
D
San Josef River
GPJB]*
0.024
17
11.2
0.515
6.63 0.010
Sakinaw Lake
GPI-B1 *
13
8.5
0.523
6.61
Fraser River
27
14.4
0.871 22.00 0.000
0.000*
24
12.5
0.923 21.30 0.000
0.013
19
12.2
0.560
0.024
0.000*
0
L9
-1.000 25.00 0.000
41
25.1
0.631 21.09 0.000
0.004
32
22.5
0.422
0.059k'
2
3.85
-0.480 12.23 0.001
0.045
14
9.3
0.500
0.002
19
11.4
0.665 10.16 0.001
Salt Creek
GPIB1*
GDA1*
GPIBJ*
PEPA1*
GDA1*
GPIB1*
sMEP2*
GDA1*
GDA1*
LDHB2*
0.000
0.000*
0.002
0
3.7
-1.000 24.00 0.000
Salt Creek
mIDHp-
0.018
0
1.9
-1.000 24.00 0.000
Salt Creek
0.019
0.087a
0
1.9
-1.000 24.00 0.000
9
5.6
0.600
0.000*
23
12.5
0.840 17.64 0.000
0.000*
31
16.0
0.939 28.24 0.000
0.002
0.000*
19
11.5
0.655 10.30 0.001
Naselle River
sMEP2*
GDA1*
GDA1*
GDA1*
GDA1*
GPIBJ*
22
11.9
0.846 17.18 0.000
Nehalem River
GDA-1
0.000*
29
14.5
1.000 29.00 0.000
Nehalem River
GPIB1*
0.001
23
13.5
0.704 13.31 0.000
Nehalem River
PEPA-1 *
0.002
0
3.6
-1.000 22.00 0.000
Sand Creek
GDA1*
0.043
14
9.1
0.538
5.80 0.016
Sand Creek
GPJ-BJ
0.036
16
10.5
0.527
5.84 0.016
Schooner Creek
0.181'
8
4.8
0.667
4.44 0.035
0.008
10
5.0
1.000 10.00 0.002
Yaquina River
GDA1*
GPIB1*
GDA1*
22
13.7
0.604 10.22 0.001
Yaquina River
GPI-B1
0.002
0.000*
29
14.5
1.000 29.00 0.000
Yaquina River
PGDH*
o.1o8
2
3.7
-0.464
6.47 0.011
Alsea River
PGDH*
0.028
3
6.2
-0.515
7.95
Cummins Creek
GDAI*
0.007
13
7.4
0.765
8.77 0.003
Ritherdon Creek
Ritherdon Creek
Ritherdon Creek
Stillaguamish River
Stillaguamish River
Stillaguamish River
Hoko River
Salt Creek
Quillayute River
Humptulips River
W.F. McClane Creek
Naselle River
Schooner Creek
*
*
X2
0.0 10
7.84 0.005
8.56 0.003
5.25
0.022
4.32 0.038
0.005
252
Appendix Table A3.2.
Continued
Exact test
Location
Cummins Creek
Cummins Creek
Umpqua River
Umpqua River
Heterozygote deficiency/excess
Locus
P
Observed
Expected
D
GpIB1*
LDHB2*
GDA]*
GPIB1*
0.000
21
12.2
0.724
0.020
0
1.9
0.006
0.000*
17
10.1
0.680
19
10.0
0.905 16.37 0.000
0.000*
26
14.0
0.857 20.57 0.000
0.042
14
8.8
0.583
6.47 0.011
0.002
23
15.0
0.535
9.44 0.002
0.167'
3
4.6
-0.35 1
4.06 0.044
0.000*
33
16.5
1.000 33.00 0.000
0.001
21
12.5
0.677 11.93 0.001
0.001
4
11.1
-0.639 10.61 0.001
x2
P
13.11 0.000
-1.000 25.00 0.000
9.71 0.002
New River
GDA-1
New River
Winchuck River
GPIB1*
GDA1*
GDA2*
GP1B1*
GPIB1*
LDHB2*
Winchuck River
PGDH*
0.038
0
1,9
-1.000 15.00 0.000
Lake Earl
GDA1*
12.5
1.000 25.00 0.000
Lake Earl
0.000
0.000*
25
GPI-B1 *
25
13.0
0.926 22.29 0.000
Kiamath River
GDA1*
GPIB1*
0.000*
23
12.0
0.920 20.31 0000
0.000
20
11.7
0.714 12.24 0.001
0.064a
1
2.8
-0.644 9.97 0.002
0.000*
27
13.5
1.000 27.00 0.000
0.000
21
11.0
0.913 18.34 0.000
0.072a
7
11.6
-0.399 4.30 0.038
0.002
12
6.0
1.000 12.00 0.001
Mill Creek
sMEP2*
GDA1*
GPJB1*
sMEP]*
GDA1*
GPJB1*
0.002
12
6.0
1.000 12.00 0.001
Mill Creek
sSOD*
1
5.0
-0.798
HSU Hatchery
GDA1*
Rogue River
Rogue River
Rogue River
Winchuck River
Kiamath River
Kiamath River
Redwood Creek
Redwood Creek
RedwoodCreek
Mill Creek
*
0.013
0.075a
7.65 0.006
0.438 4.22 0.040
These loci did not exhibit significant deviation from Hardy-Weinberg expectations but
did exhibit a significant deficiency or excess of heterozygotes.
b
This locus did not exhibit a significant deficiency or excess of heterozygotes but did
exhibit a significant departure from Hardy-Weinberg expectations (prior to correction for
multiple comparisons).
a
15
10.4