AN ABSTRACT OF THE THESIS OF Master of Science Geoffrey Lynn Dorsey

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AN ABSTRACT OF THE THESIS OF
in
Geoffrey Lynn Dorsey
for the degree of
Master of Science
Fisheries and Wildlife
presented on
June 6, 1982
Title:
An Analysis of Avian Coimnunities on Three Islands, Lower
Columbia River, Oregon
Abstract approved:
Redacted for privacy
Dj
John A. Crawfo, major professor
Avian communities were described for t habitats (marsh, upland,
beach, and tree-shrub) on 3 islands (West Sand, Miller Sands, and Jim
Crow Sands) in the Lower Columbia River.
June 1978 through May 1979.
Research was conducted from
The variable circular-plot census method
was used to determine avian commmunity characteristics (number of
species, density, H', and J').
Percent cover of various habitat
characteristics were measured to determine relationships with avian
community characteristics.
Marsh habitats differed in number of species
observed and vegetative cover.
Increased number of avian species
was associated with increased vegetative cover for marsh habitats.
Shorebird usage of marshes was related to the incidence of bare
ground.
Upland habitats were similar with regard to avian community
characteristics; however, a marked difference in species composition
was observed for barren versus vegetated upland habitats.
Tree-shrub
habitats were not different with respect to avian community charac-
teristics although the transition habitat was markedly different with
regard to structural complexity.
Beach habitats were similar in avian
and habitat characteristics.
Comparisons of avian community charac
teristics among habitats grouped by island indicated that the most
successionally advanced and structurally complex habitats tended to
differ from less advanced and less complex habitats.
An Analysis of Avian
Communities on Three islands,
Lower Columbia River, Oregon
by
Geoffrey L. Dorsey
A Thesis
submitted to
Oregon State University
in partial
fulfillment of
the requirements for
the degree of
Master of Science
June 6, 1982
APPROVED:
Redacted for privacy
essor of Wildli
Ecolo-
in chfilge of major
Redacted for privacy
Head of
Department bf
Fisheries
and
Wildlife
Redacted for privacy
an of Graduate
Date thesis is presented
Typed by
LaVon Mauer for
January
7, 1982
Geoffrey Lynn
Dorsey
ACKNOWLEDGEMENTS
Many individuals have contributed to the development of this thesis and to each I am grateful.
Initially, C. Wayne Eshelinan took an
interest of mine and nurtured a desire to learn.
and gratitude to him continue to this day.
took the baton as my major professor.
My admiration for
Dr. John A. Crawford then
I-U.s guidance, support, insight,
and dogged determination have brought me to this point where further
attainment lies fully with me.
I truly appreciate the leadership he
has provided me.
Many fellow graduate students provided assistance in various
aspects of this research.
Russ Oates, Dan Edwards, Dirk Van Vuren,
Mike Morrison, and Cecilia Noyes are just a few to whom I owe thanks.
Dan helped me initiate this study; Russ provided advice and support;
Dirk edited one draft and discussed various aspects of the research
and results; and Cece and Mike provided invaluable assistance on statistical analysis.
Larry Morrison taught me field identification of
plants on my study islands.
Phil Hamm assisted me in the field and
lent moral suport, but more importantly has been a friend.
Eric Beals
often led me through the morass of coinputerdom.
The Department of Fisheries and Wildlife faculty have provided
advise, discussion, and an education.
this thesis.
period.
Dr. Robert Anthony has reviewed
Dr. Richard Tubb provided monetary support at a critical
Thanks seems inadequate but I am truly thankful for their
contributions and hope one day I can repay them in full.
The U.S. Army Corps of Engineers, Portland District provided
funding and provided assistance and direction throughout the research.
Dr. Thotnas Morse faciitated the initiation of this research and pro-
vided frequent advice and support throughout the research.
What words appear on this page and thoughout this thesis are due
in one sense, to only one individual--LaVon Mauer.
Her assistance,
innovativeness and persistence are appreciated.
Finally, I owe a great deal of thanks and love to my wife, Joyce,
for her patience, understanding, and tolerance of my frequent absence
from home.
My parents were also supportive of this endeavor and I
thank them.
My sincere thanks to all who have contributed in any manner to this
research.
TABLE OF CONTENTS
Page
INTRODUCTION
I
STUDY AREA
5
METHODS
9
RESULTS
14
Marsh
15
Upland
28
Tree-Shrub
36
Beach
43
West Sand Island
43
Miller Sands
50
Jim Crow Sands
52
CONCLUSION
52
MANAGNENENT RECOMMENDATIONS
52
LITERATURE CITED
54
APPENDICES
62
LIST OF FIGURES
Page
Figure
1.
Habitat locations for West Sand Island,
7
Columbia River, Oregon.
2
Habitat locations for Miller Sands Island,
8
Columbia River, Oregon.
3
Habitat locations for Jim Crow Sands Island,
Columbia River, Oregon.
10
LIST OF TABLES
Page
Table
1
Number of species, and avian density, diversity, and
16
evenness values by season for each habitat group,
Lower Columbia River, 1978-79.
2
Discriminant function results (avian community
18
characteristics) for habitat groups for which habitat
and season effects were determined significant by
multivariate analysis of variance.
3
Habitat comparisons of species composition values
19
(MacArthur's Difference) for similar habitat groups,
Lower Columbia River, 1978-79.
4
Seasonal comparisons of species composition values
20
(MacArthur's Difference) for combined similar
habitats, Lower Columbia River, 1978-79.
5
Percent cover of litter, forbs, and grass/sedge
21
for marsh habitats, Lower Columbia River, 1978-79.
6
Results of discriininant function analysis (habitat
23
characteristics) for habitat and season effects
determined significant by multivariate analysis of
variance.
7
Percent cover for each habitat characteristic, West
30
Sand Island, Miller Sands, and Jim Crow Sands uplands.
8
Percent cover of tail grass/sedge (
15 cm) for
32
MSU, WSIDU, and WSIU, Lower Columbia River, 1978-79.
9
Diversity indices for "edge" of individual habitats,
Lower Columbia River, 1978-79.
34
Table
10
Pa&e
Percent cover values for each habitat characteristic,
38
tree-shrub habitats, Lower Columbia River, 1978-79.
11
Percent occurence of bare ground for 4 beach
44
habitats: MSSB, MSB, JCSB, and WSIB, Lower Columbia
River, 1978-79.
12
Mean values of each avian community characteristic
46
for habitats comprising each island, Lower Columbia
River, 1978-79.
13
Habitat comparisons of species composition values
48
(MacArthur's Difference) for habitats grouped by
island, Lower Columbia River, 1978-79.
14
Seasonal species composition values (MacArthur's
Difference) for combined island habitats, Lower
Columbia River, 1978-79.
49
An Analysis of Avian Communities on Three Islands,
Lover Columbia River, Oregon
INTRODUCT ION
Previous investigations of dredged material islands were principally directed toward determining plant succession (Barnes 1971,
Coastal Zone Resource Corp. 1977, Dunstan and Lewis 1974), nesting by
colonial species of waterbirds (Buckley and Buckley 1975, Buckley and
McCaffrey 1978, Chaney et al. 1978, DuPue 1974, Landin 1978, Morrison
and Shanley 1978, Olsen 1975, Peters et al. 1978, Scharf 1978, Soots
and Parnell 1975, and Thompson and Landin 1978) or animal colonization
(Cammen et al. 1974).
Several investigations of avian communities have been conducted in
the
Columbia River estuary.
Woodward-Clyde Consultants (1978)
investigated both floral and faunal aspects of Miller Sands between 23
April and 24 June 1975 and determined that bird species diversity was
highest in the tree-shrub association and lowest in upland and
sandspit habitats.
Crawford and Edwards (1978) conducted research on
the responses of bird, mammal, and macroinvertebrate populations to
plantings on Miller Sands between June 1976 and August 1977.
Avian
density and diversity were greatest in the tree-shrub habitat; number
of species was highest in the beach and marsh habitats (Crawford and
Edwards 1978).
Number and type of avian species present were occa-
sionally altered by plantings, but density and diversity were usually
unchanged (Crawford and Edwards 1978).
Edwards (1979) determined that
avian community characteristics (density, number of species, diversity
2
(H'), evenness (J') and standing crop bioinass) associated with upland
and tree-shrub habitats on Miller Sands were significantly different.
Avian community characteristics of upland and tree-shrub habitats on
Miller Sands were significantly different from marsh and beach habitats (Edwards 1979).
Seasonal differences also occurred among avian
communities (Edwards 1979).
Avian community characteristics for
spring differed from summer (1976), fall, and winter and those for
summer (1977) differed from fall and winter.
Avian community charac-
teristics and species composition were influenced by season, vegetative complexity, and the physical attributes associated with each
individual habitat (Edwards 1979).
The variation in avian species composition, density, and diversity
with respect to vegetative communities observed by investigators on
Miller Sands also may be a function of the soil moisture gradient.
Investigations conducted on Mott Island, Columbia River, Oregon, by
the Coastal Zone Resources Committee (1977) revealed that soil
moisture gradient, a function of topography and tidal influence, iiad
an important role in determination of vegetative communities.
Plantings also influenced the vegetative communities on ?lott Island
(Coastal Zone Resources Committee 1977).
Factors affecting vegetative
succession on dredged material islands in Florida included climate,
substrate, wave action, nearness of plant sources, and human use
(Schreiber and Schreiber 1978).
Scharf (1978) found little correla-
tion between age of islands and succession of vegetation on islands in
the Great Lakes.
He ascribed this lack of a relationship to variation
in plant succession and parent dredged material and the influence of
3
Retardation of succession on some dredged material islands in
ice..
the Great Lakes was attributed to parent material, bird usage effects,
erosion, and/or inundation (Scharf 1978).
Yeaton and Cody (1974) reported that the principal effect of the
addition of canopy cover was the addition of more bird species to the
community.
Edington and Edington (1972) noted that different species
forage in different vertical strata, thus the more vertical stratifi-
cation, the tre species opportunities (see also MacArthur 1958, Morse
1970).
Avian density generally increases with succession (Shugart et
al. 1975, Kendeigh 1948, Odum 1950, Johnston and Odum 1966, Haapanen
1965, Karr 1968, Karr and Roth 1971, and Shugart and James 1973).
Adams (1908) and Bond (1957) noted that species richness was higher in
intermediate successional stages.
Meents (1979) stated that addition
of a shrub layer led to greater species richness.
McElveen (1977)
reported that species density and avian density was greater in edge
situations than for either vegetative community forming the ecotone.
Stauffer and Best (1980) reported an increase in mean densities of
breeding birds from herbaceous habitats (uplands-this study) to upland
woodlands to floodplain woodlands.
Karr (1968) observed a general
increase in number of species, density, and diversity from bare ground
to bottomland forest habitats on former strip mine sites.
Wilison
(1974) determined that increased environmental patchiness, specif i-
cally the addition of trees, strongly influenced the addition of avian
species to a habitat.
The increase in number of species was greatest
when formation of a tree layer was initiated.
Des Granges (1980)
stated that equitability depended on physiognomic diversity of a
4
forest stand; high equitability was associated with more complex
stands.
An inventory of riparian habitats and associated wildlife for the
Columbia River was conducted by the Oregon Cooperative Wildlife
Research Unit (OCWRu) (1976).
Investigations were conducted in habi-
tats similar to and in the vicinity of present study islands; however
habitats on dredged material islands were not investigated.
West Sand
Is land was included in Segment 1; Miller Sands and Jim Crow Sands were
included in Segment 2 of the study.
Alder communities in Segment 1
possessed the highest number of species, avian density, and avian
biomass (OCWRU 1976).
Seasonal variation in maximum numbers of birds
and densities were observed in Segment 2 (0CwRU 1976).
Sitka spruce
(Picea sitchensis) habitat contained the greatest number of species
during all seasons except spring; cottonwood communities contained the
highest number of species during spring.
Avian density was greatest
during fall and winter in Sitka spruce habitats.
Cottonwood/willow
communities were represented by the highest avian density during
spring.
Avian density was highest during summer in cottonwood coni-
munities.
Since habitats on these islands are potentially subject to drastic
changes in structure from dredged material deposition, this study was
initiated.
Habitats that varied in vegetative complexity were
investigated.
The purpose of this research was to describe avian com-
munity and habitat characteristics associated with 4 gross habitats
(marsh, upland, beach, and tree-shrub) on 3 islands (Miller Sands, Jim
Crow Sands, and West Sand) representing varying successional stages
5
and determine relationships between avian community and habitat
characteristics among similar habitats and for island habitat complexes.
STUDY AREA
The study islands, West Sand, Miller Sands, and Jim Crow Sands,
were chosen for investigation because they are current disposal sites
for dredged material.
Each island was typified by a complex of habi-
tats (marsh, upland, beach, and tree-shrub).
Ages of specific habi-
tats varied among islands and were related to date of formation from
dredged material deposition.
The climate of the Lower Columbia River is typified by wet winters
and dry summers (U.S. Army Corps of Engineers, Portland District, 1975).
Astoria, located centrally among study sites received 143.8 cm of precipitation with average maximum and minimum temperatures of 14.7 C and
5.8 C, respectively, for the period June 1978-May 1979 (National
Oceanic and Atmospheric Administration 1978, 1979).
Severe cold
weather from December until early February resulted in the freezing of
surface layers of intertidal marshes and mudflats and a virtually
complete blockage of the main river by ice floes during two periods.
West Sand Island (257 ha) was located in Baker Bay, east of Cape
Disappointment, Washington.
Formally a natural bar, West Sand Island
was shif ted to its present position by channelization activities that
occurred at the mouth of the Columbia River (Chief of Engineers, U.S.
Army 1937; Oregon Historial Society 1980).
Dredged material deposi-
tion altered and added habitats to West Sand Island (e.g. diked
upland).
Avian censusing stations were established in 6 habitats on
West Sand Island (Fig. 1): marsh (WSIM), diked upland (WSIDtJ), upland
(WSIU), beach (WSIB), transition (WSIT), and tree-shrub (WSITS).
Grass/sedge, principally Carex sp. and creeping bentgrass (Agrotstis
alba), was the dominant ground cover of WSIM.
Red fescue (Festuca
rubra) and tall fescue (F. elatior) were the dominant species forming
WSIDIJ ground cover.
Red fescue, seashore lupine (Lupinus littoralis),
coast strawberry (Fragaria chiloensis), and black knotweed (Polygonum
paronychia) characterized WSIU.
of WSIB.
Bare ground was the dominant feature
Carex sp. was the dominant plant species in WSIT.
The
overstory of WSITS was composed of red alder (AJ.nus rubra) and willow
(Salix sp.).
Willow, twinberry (Lonicera involucrata), and salmon-
berry (Rubus spectabilis) were the dominant understory shrubs.
Litter
and grass/sedge, principally Carex sp., comprised the ground cover.
Miller Sands (228 ha), located between river miles 22 and 25, was
formed in 1932 by deposition of dredged material (Edwards 1979).
Creation of a spit began in 1948 and was completed in 1976.
Presently, deposition is limited to the spit area on Miller Sands.
Censusing on Miller Sands was conducted in 6 habitats (Fig. 2):
marsh
(MSN), spit upland (MSSU), upland (MSTJ), spit beach (MSSB), beach
(MSB), and tree-shrub (MSTS).
Carex sp. and bare ground typified MSM
during the summer. MSSU was characterized by bare ground.
Moss,
f orbs, litter, and grass/sedge constituted the primary ground cover of
MSU.
MSSB and MSB were devoid of vegetation.
Red alder, cottonwood
(Populus trichocarpa), and willow were the main constituents of NSTS
overstory.
Salmonberry and elderberry (Sambucus raceimosa) were the
Figure 1.
Habitat locations for West Sand Island, Lower Columbia River, Oregon.
7
I
caie - 1
Figure 2.
.
lEO
Habitat locations for Miller Sands Island, Lower Columbia
River, Oregon.
most abundant understory shrubs.
Ground cover of MSTS was typified by
litter, forbs, and grass/sedge.
Jim Crow Sands (86 ha), also formed from dredged material, was
located between river miles 26 and 28.
Deposition has occurred on Jim
Crow Sands since at least 1957 (U.S. Army Corp. of Engineers Pipeline
Disposal History, Portland District 1980).
sused on Jim Crow Sands (Pig. 3):
beach (JCSB).
Three habitats iere cen-
marsh (JCSM), upland (JCSU), and
Deposition on the eastern portion of JCSU had occurred
just prior to study initiation, hence, regarding succession, the
eastern portion of JCSU was chronologically younger than the western
segment.
Bare ground was the prominent feature of the west and east
ends of JCSU.
Grass and forb plantings during September 1978 on the
recent deposition site markedly reduced the amount of bare ground.
Bare ground typified JCSB.
Bare ground and grass/sedge were the domi-
nant features of JCSM.
METHODS
The variable circular-plot method (Reynolds at al. 1980) was used
for avian sampling.
This method was selected because variations in
detectability among species of birds and among seasons and habitats
for a species is accounted for when calculating species densities,
A total of 15 habitats on 3 islands were sampled monthly from June
1978 through May 1979.
Two habitats were sampled each day and 10 sta-
tions were censused in each habitat.
at each station.
Data were recorded for 10 nun.
Birds, detected visually and/or aurally within the
boundaries of a habitat and under 40 m in altitude, were recorded by
j
ficale - 1 cm
103 m
New upland
Marsh
Old Uplflfl(i
Figure 3.
Habitat locations for Jim Crow Sands Island, Lower Columbia River, Oregon.
11
species and distance from observer for each individual or group of
individuals.
Density estimates were based on an effective detection distance
determined for each species (Reynolds et al. 1980).
recorded in concentric 10
in
Birds were
bands from the observer to 150 in, in 25
bands from 151 in to 200 in, and in 100
in
in
bands for greater distances.
Numbers of individuals for each band were recorded for each habitat.
Similar habitats (e.g. MSM, JCSM, WSIM) were combined to increase
sample size for determination of the effective detection distance.
Seasonally, the effective detection distance and subsequent species
densities were calculated by the method of Reynolds et al.
(1980).
The percentage of paired singing males was not determined, thus no
correction factor for avian density during the breeding season was
utilized.
Species density estimates for each habitat were determined
from the formula:
D =
(10,000 m2ha)N
= birds/ha; where x
the
xir-
number of stations sampled (10); r
the radius of the circle deter-
mined by the inflection point; and N is the number of birds detected
within the radius of the circle.
yielded total
Summation of species densities
avian density for each habitat.
Densities for groups
of birds (e.g. lands, hirundinids) were apportioned according to the
density determined for individuals composing the group or by proportion of individuals observed during incidental counts.
Diversity indices (Shannon and Weaver 1949) were calculated from
the formula H' =
in the ith species.
p. logp; where p
= the proportion of individuals
Where density estimates for groups (e.g. lands,
hirundinids) occurred for a habitat, p. was calculated only from those
12
densities associated with identified species.
Evenness (J') was
determined by dividing H' by the logo of the number of species
observed in each habitat.
Species composition among similar habitats and between seasons for
individual and combined similar habitats was examined with MacArthur's
(1965) difference measure:
ik
e
(lit-H)
p.,+p; where
lo
difference.
ih
of the ith species of Hh and HK (where H,
for the first census and
and
ik
=
equal the proportion
equals the diversity index
for the second).
Differences ranged from
1.0 to 2.0; a value of 2.0 denotes 2 communities with no species in
common.
Coinniunities are more similar as the difference value
approaches 1.0.
Vegetation sampling was conducted seasonally.
Habitats were
characterized by 80 0.25u2 quadrats (1 m2 for beach habitats and
uplands devoid of vegetation); 2 quadrats were randomly distributed
along lines in each cardinal direction at each station.
Habitat
characteristics evaluated were percent cover of moss/lichen,
rock/litter, bare ground, grass/sedge, and f orbs.
Additionally, per-
cent overhead cover, canopy height, canopy base height, and circumference of trees were determined for MSTS, WSIT, and WSITS.
Canopy
and canopy base height, indices of foliage height diversity, were
determined for these habitats by determining the number of individuals
(p.) in each 1 ni increment above 2 in.
The Shannon-Weaver (1949) for-
mula for diversity was again used for calculation.
Percent overhead
cover was determined by visually estimating the area of a leveled
13
Brunton Pocket Transit mirror obstructed by overhead cover (leaves,
branches, and/or stems of trees).
This characteristic
was nasured
seasonally at 8 randomly selected sites for each avian census station;
2 sites were randomly located along a line extending in each cardinal
direction from the census station; with 4 measurements taken at each
site.
Habitat edge effect was quantified based on the formula deve-
loped by Patton (1975):
DI
TP
where DI = diversity index, TP
2I
total perimeter (meters), and A equals area (meters squared).
Multivariate analysis of variance (SPSS-MANOVA) (Cohen and Burns
1976) was used to test for differences, with respect to the avian
community and vegetative characteristics among:
habitats;
2) habitats grouped by island; and
1) combined similar
3) seasons.
Correlation analysis was utilized to eliminate those variables which
exhibited a high intercorrelation.
Interaction (season by habitat),
habitat, and season were the main effects tested by the multivariate
procedure.
A significant interaction effect normally precludes
further analysis of main effects (Kim and Kohout l975)e
of Statistics OSU, pers.
Ramsay (Dept
comm. 1980) indicated that if a biological
basis for the interaction effect was determined, then further analysis
of the main effects (habitat and season) was not necessarily
precluded.
Significant differences (P
0.05) detected by the
SPSS-MANOVA procedure (Wilk's lambda value) were further analyzed with
the Hotelling T-squared statistic (Morrison 1967) to determine which
habitats or seasons were significantly different from one another.
Univariate analysis of variance, a subunit of the SPSS-MANOVA
14
statistical package, was used to gain an indication of the specific
parameter(s) which led to significant differences among habitat cornparisons.
Inadequate sample size obviated separation of seasons for
avian community data.
Discriminant function analysis (Kiecka 1975) was used for separation of habitats and seasons by avian and vegetative characteristics
when MMOVA indicated a significant difference.
Discrim.inant function
analysis provided an indication as to which characteristic(s) was most
important for group (habitat or season) separation.
Canonical corre-
lation Cr) was utilized as a measure of the degree of group separation for each discriniinant function.
Canonical correlations that
exceeded 0.70 were considered strong indicators of habitat differences.
0.15 was used for determination of signif 1-
An alpha level of P
cance among multiple comparisons unless otherwise specified.
For
multiple comparison tests alpha (0.15) was divided by the total number
of possible comparisons.
This procedure established the actual alpha
level for entry into the appropriate F-table and thus determination of
significance for comparisons.
Hence, fifteen possible comparisons
existed among the 6 habitats present on Miller Sands; therefore, x
0.15/15
0.01, the alpha level with associated degrees of freedom for
critical F-values.
RESULTS AND DISCUSSION
Results are separated on the basis of major habitat groups (marsh,
upland, beach, and tree-shrub) and islands (West Sand, Miller Sands,
Jim Crow Sands).
Habitats were grouped on the basis of similarities
15
in vegetation, substrate, tidal influence, and general structure.
Combination of similar habitats based on these features was conducted
to determine if subtle differences influenced avian community characteristics and composition.
Grouping habitats by island allowed com-
parison of various habitats.
MARSH
Among marsh habitats, JCSM:WSIM was the only comparison for which
significance was determined (Table 1) and number of species was the
only characteristic that differed (P
0.05).
The observed difference
in number of species for JCSM:WSIM comparison was consistent for all
seasons except winter when similar counts of species occurred (Table
1).
Discrimination among marsh habitats also was accomplished on the
basis of number of species; density was of secondary importance (Table
2).
Based on this separation, JCSM was least similar to WSIM, and MSM
was intermediate to those habitats.
The first discrimiaant function
was not a strong separator of marsh habitats (rc = 0.55).
composition of all marsh habitats was similar (Table 3).
Species
Seasonally,
the contrast in species composition was greatest for the suminer:winter
comparison (Table 4).
Analysis of marsh habitat characteristics indicated a significant
season by habitat interaction effect (P < 0.05).
This effect was
attributed to the paucity of forbs in JCSM which obscured seasonal
changes in f orb cover as observed for MSM and WSIM (Table 5).
Comparisons among marsh habitats
and habitat characteristics asso-
ciated with each indicated that all comparisons were significant
Thbk 1. Nwtr of bpcie. a1 twlan dentty, dirfty aiii eveoiiw va1ic,i ti ucu br edi
AvLn Cotiiuntty
Nwgeb of
11thitt
grwp
ctes
FLIII
Ithter
SprirE
12.3
10.7
10.7
12.0
11.4
NfH
12.3
17.3
11.0
14.3
WSIM
16.0
17.0
11.3
16.3
c,4
.iL
1trutby
Spr1r
I4asi
Siaeer
hilt
WInt.r
3.4
18.7
12.5
Lit
1.50
[.52
9.9
1.4
11.1
1.2
1.72
1.88
5.5
6.6
6.2
10.4
1.2
135
l'aU
WInter
8.9
18.8
13.7
5.9
15.2
*u
grwp, 0r (1uthta K1ur 19il$-1J.
rjcL0rI1t1C
(b1nW1i)
urIwr
JCSH
lkl
Suwor
btan
iieer
hiLt
Witer
0.94
1.42
0.69
0.t3
0.05
039
0.59
1.69
1.66
1.79
0.00
0.00
0.79
0.58
u.00
1.87
1.00
1.82
1.01
0.03
0.00
0.41
0.0/
0.59
Spd
r1rE
I*iiwi
Up1as
6.0
8.3k
8.5
8.2
2.3
3.6
6.6k
5.2
1.40
1.82
0.88
138
1.31
0.58
0.85
o.57
0.tE5
43.66
WSIW
10.0
11.3
1.3
8.0
9.2
43.0
4.0
1.9
4.2
5.8
138
1.49
0.81
1.29
1.24
0.64)
0.61
0.41
0.liS
0.51
J4J
10.7
6.3
3.3
[4.3
83
4.8
4.8
0.8
1.9
4.6
1.40
0.73
43.34
2.01
1.12
0.59
0.41
0.22
0.19
0.50
MS9l
9.1
7.3
7.0
9.0
8.3
17.4
6,9
Li
18.1
10.9
1.21
1.11
1.58
1,38
1.25
0.9
0.56
0.14
0.64
0.64
WSIU
16.3
1.7
5.0
12.3
10.3
8.1
1.4
0.5
4.7
3.6
1.56
1.61)
0.448
1.66
1.43
0.50
0.19
u.62
43.00
0.06
12.3
9.7
8.7
11.7
10.6
19.8
12.9
1.8
19.6
13.5
1.23
1.41
1.36
1.4/
130
0.49
0.0/
0.63
0.61
0.64)
)H
10.3
9.0
11.0
15.0
11.3
6.1
2.3
2.6
17.0
1.0
4.70
1.37
1.78
1.53
1.62
0.79
0.04
0.10
0.5/
0.60
JtB
14.3
43.3
8.3
14.0
11.5
15.5
7.5
/.4
135.6
41.5
[.44
1.60
1.02
1.52
1.40
0.60
u.02
0.48
0.u4
0.38
4I8
40.7
13.0
10.7
10.3
41.2
12.2
9.0
4.6
7.6
44.4
1.4o
1.49
1.53
1.59
1.52
0.02
43.56
u.o
0.69
0.64
beade
flhl1a 1.
(conttiud)
Avtai ccu.auilty cractertatic
IR1tya
Nnxibar at
Hahttat
Spthi
haii
Suiaiar
Fall
Wlaer
Sr10
t*an
Sua&at
hiLl
Whiter
5irui
a.ia
9.1
10.1
12.5
1.68
1.75
1.90
2.03
1.84
0.10
0.70
0.18
0.79
0.)h$
4.8
10.0
14.0
10.4
2.28
1.42
1.67
2.24
1.90
0.88
0.72
0.81
0.81
0.80
6.5
9.9
14.3
12.3
2.32
1.52
1.94
1.93
1.93
0.80
0.68
0.744
0.72
0.15
t$aU
&.aler
t1t
Wthtec
13.0
10.9
6.0
24.1
8.0
16.3
11.6
12.9
12.3
15.0
14.0
18.6
Fall
Wtnter
7.0
12.0
11.1
tvr
14.7
7.3
%flS
48.3
40.3
&aaer
grwp
(bt.1/I&a)
pedeia
rtng
?rae-uub
g
data
re trawfonaid (aquare coot) print to uaiywl.w.
wraity data were not Included in analyaia at upland habluata becanaie at ttai htJl correlation (r
C8k
aerttca1 lbnca j68iete
tUcant dii nteacca (ii
0.440) a.iLUa e'&nncea.
0.05); cwaccted ltncs indicate no ditietence.
unlinrlaie uI t'Lences uuors lasbitata.
deticai ltraia tot habitat groop represent oulihurtate dl erencea; veLticat 1laea tar ascii asian ccusuutty tharaotetiUc repnn.ent
was algnittcan4' dttfetent F
SprlIUJ was a1gn1f1cant
0.025) iron fall aiai winter with roapect to najiber at species and dauaily.
diffcrent F 4 0.025) fran winter with respect
Lu
tuiter of spectai density, and evenness.
further tests weca coaui,ctu.t it tailtiwariate analysts tcvealad no slit1icant dittarences.
I-
Table
Dtscriiainant functinu results (avian coaaunity characteristics) for habitat groups for which habitat
2.
and season effects were deteri1ned signiticant by multtvariat analysis o variance.
Effect tested
and habitat
group
Kotated atandardted discriminant
function coefficients (each parameter)
biscrituinant
function
Percent of
variance
correlation
Species
1)enaity
Diversity
8veuness
1*
78.4
21.6
0.55
0.32
-1.39
-0.67
1.05
0.43
0.54
1.34
---
72.3
26.1
0.48
0.31
-0.45
-0.32
1.24
0.35
---
0.12
1.08
2*
56.0
39.3
0.62
0.56
1.40
0.52
-0.54
0.38
0.39
1.23
---
0.69
0.48
-0.51
-0.12
1.33
0.06
---
0.00
2*
65.0
21.5
1*
92.0
0.69
0.27
-0.13
1.08
0.16
1.06
-0.27
0.73
0.35
0.21
1.26
-0.27
-0.48
-----
Canonical
HABITAT
Harsh
2
Upland
j*
2
Miller Sands
1*
West Saud Island
1.00
SEASON
Upland
2
Tree-Shrub
8.0
1*
88.0
2
10.9
*Significant discriminatory information existed prior to formation
1
discritainant function (P
-0.11
1.04
0.04
0.05).
19
Table
3.
Habitat comparisons of species composition
valuesa (MacArthur's
Difference) for similar habitat groups, Lower Columbia River,
19 78-79.
Habitat
Group
Marsh
Habitats
MSM
JCSM
JCSN
1.22
WSIM
1.33
1.24
Upland
MSSU
NSU
JCSU
MSU
1.45
JCSU
1.46
1.39
WSIDU
1.75
1.44
1.60
WSIU
1.62
1.34
1.34
Beach
MSSB
MSB
JCSB
MSB
1.20
JCSB
1.19
1.23
WSIB
1.41
1.40
MSTS
WSITS
Tree-Shrub
WSITS
1.15
WSIT
1.22
aValoes range from 1.0 to 2.0.
WSIDU
1.37
1.44
1.13
Differences in species composition among
comparisons are greater as 2.0 is approached.
20
Table
4.
Seasonal comparisons of species composition values8
(MacArthur's Difference) for combined similar habitats,
Lower Columbia River, 1978-79.
Season
Habitat
group
Summer
Fall
Winter
Marshes
Fall
1.45
Winter
1.71
1.27
Spring
1.36
1.24
1.26
Uplands
Fall
1.44
Winter
1.69
1.31
Spring
1.23
1.34
1.42
Beaches
Fall
1.42
Winter
1.74
1.27
Spring
1.32
1.32
1.32
Tree-Shrub
Fall
1.24
Winter
1.33
1.14
Spring
1.13
1.32
aValues range from 1,0 to 2.0.
1.33
Differences in species composition among
comparisons are greater as Z.0 is approached.
3.9
10.5
32.8
6.6
17.3
49.5
8.7
46.5
56.4
2.9
3.2
8.1
JCSM
HSM
WSU4
Spr1ng
Winter4
50.5
73.5
2.4
2,3
1.1)
0.2
0.0
0.0
1.7
0.2
7.1
8./
36.8
linea indicate atguiticant difference.
were tranaforwed (wreath /11) for analywia,
21.11
19.4
O.l
0.1
0.0
5.5(
0.1
SueaerC
0.1
Mean
gpringt
Winter
iail
Suaar'
Mean
Forb
ercent cover of forha cad grawe/aedge doring apring ditfered frow winter.
cover of litter loting apring differed true tall.
4Vercent cover of litter, lorba, and graaa/wedge during winter d1fferd from tall.
CVerceot cover of litter, forb, and graae/wedge during anawer differed true tail, winter and apriug.
b8k
liabttwt
Fall
Litter
24.1
8.2
1.4
Fail
t*ercent coverU of litter, torba, and graau/aed6e for i&irai habttata, Lower Coluwbia Kivr, 19111-79.
Suiuwer
Table 5.
4.5
1.2
J.2
Winter
2.o
9.9
2.9
priug
11.4
Heau
22
0.03).
(P
Percent cover of litter and grass/sedge were the prin-
cipal habitat characteristics that separated marsh habitats (Table 6).
A continuum from low (JCSM) to high (WSIM) percent cover of litter and
grass/sedge was developed on this basis.
A high degree of separation
(rc = 0.79) existed among marsh habitats based on the first disciminat ion.
Identical placement of marsh habitats occurred for ordinations
developed on the basis of avian community and habitat characteristics
Jefferson (1977) stated that plant distribution in salt marshes was
related to elevation.
She developed a classification scheme wherein
salt marshes were assigned to six categories:
low silt marsh,
3) sedge marsh,
immature high marsh, and
1) low sand marsh,
4) bulrush and sedge marsh,
6) mature high marsh.
2)
5)
Although JCSM and MSM
are freshwater marshes, characteristics of these marshes are very
similar to the low silt (2) and bulrush and sedge (4) marsh categories, respectively.
West Sand Island marsh, a saltwater marsh,
appeared most similar to a mature high marsh (6), which Jefferson
(1977) considered a climax situation.
Thus, a successional gradient
from JCSM to WSIM apparently existed and coincided with the ordination
of marsh habitats on the basis of number of avian species and the
habitat parameters, litter and grass/sedge.
The increased elevation and concomittant increased occurrence and
coverage of vegetation associated with WSIM compared to JCSM appeared
influential with regard to number of species of birds.
WSIM had a greater amount of edge than JCSN (DI
respectively).
Additionally,
3.93 and 2.58,
Habitats abutting WSIM were open water, upland, and
Table 6.
kesulta of d1ecrIutnLInt fujicttod analysis (habItat characteitsttce) for hb1tat anU season etfecte
teretuen at ttitaut by
tivartate
analysis of variance.
hoIsted taiidardize4 diacrisinaut
iuncctou coefttclenta (acU parseeter)
lifted
tested
Discriiainant
function
Percent of
variance
coirelation
94.0
0.79
-
0.94
2*
6.0
0.3!
--
1*
60.0
0.95
2*
29.0
1*
Canouicsl
t4oss/
Hchcu
hock/
titter
bare
ground
Iass/
1ercent
overhead
IOLbS
sedge
cover
0.21
0.75
-
0.10
---
1.25
-0.18
--
1.15
0.111
0.37
0.24
(1.53
0.91
0.20
0.30
1.2?
0.24
0.31
---
79.7
0.92
0.26
0.19
-0.43
-0.17
1.04
2*
20.3
0.77
0.64
1.43
---
0.50
1.99
-0.04
1*
91.6
0.98
1.05
0.12
0.14
0.14
2*
6.6
0.79
0.08
1.06
--
0.19
0.09
---
1*
87.8
0.77
0.34
0.55
-0.10
0.10
0.75
2*
12.2
0.41
1.01
1.20
2.15
-0.36
1.91
--
1*
71.6
0.95
0.08
0.12
2.12
0.17
0.15
2*
16.0
0.82
0.66
4.01
2.66
1.09
3.65
---
tlabt tat
Marsh
Upland
Tree-Shrub
Miller Sands
Jiia Crow Sands
West Saud Island
1
I
Table 6.
(contInued)
kotated ataudardized ala iuiinent
cocUicleitta (each peraateter)
lunction
ierceat
Effect teated
Diecrtaiiiant
function
Percent ot
variance
Canonical
correlation
Hoab/
Rockj
Craaa/
overheen
Iorba
sedge
cover
Uare
lichen
litter
ground
Season
1*
94.8
0.85
--
1.19
--
0.02
-0.43
--
2*
5.2
0.35
--
-0.34
--
-0.07
-1.19
--
Upland
1*
99.6
0.74
1.80
2.63
3.40
0.02
1.60
--
Tree-Shrub
j*
90.4
0.82
0.69
1.50
--
-0.49
-0.90
2*
7.8
0.39
0.17
0.51
--
1.31
0.85
Marali
*Stgntficeot (P
0.59
0.05) dlacr1utnatory Intor.ivation existed pilot to ioraat1on of diacrlwlnant function.
r',)
tree-shrub, whereas JCSN was abutted by open water and beach habitats.
Species that typically foraged in upland sites on West Sand frequented
WSIM during summer [e.g. marsh hawk (Circus cyaneus), long-billed marsh
wren (Cistothorus palustris), starling (Sturnis vulgaris), common
yellowthroat (Geothlypis trichas), house finch (Carpodacus mexicanus),
and song sparrow (Melospiza melodla)].
The high elevation of WSIM and
the immediate proximity of upland habitat facilitated use of WSIM by
upland-associated species.
Few species associated with upland habi-
tats were detected In JCSM during summer.
Number of species differed between JCSM and WSIM during fall
(Table 1).
This difference was apparently related to edge effect and
availability of foraging sites.
Some species that typically foraged
in upland sites [e.g. rough-legged hawk (Buteo lagopus), marsh hawk,
and starlingJ were observed foraging in WSIM.
Thus the higher eleva-
tion of WSIM and its juxtaposition to WSILJ influenced the number of
species present.
Additionally, species that typified an open water
habitat tended to occur more often in WSIM than JCSM.
Representative
open water species present in WSIM included common loons
(Gavia
immer), greater scaup (Aythya mania), bufflehead (Bucephala albeola),
and Bonaparte's gull (Larus philadelphia).
Other species that foraged
in shallow water or uiudflats [e.g. great blue heron (Andea herodias),
black-bellied plovers (Piuvialis squatarola), and dowitchers
(Limnodromus sp.) also were observed In WSIM but not JCSM during fall.
The juxtaposition of WSIM to Baker Bay, a large, shallow bay with
extensive amounts of intertidal
concentrations of
cormorants,
mudflat
at low tide which attracted
waterfowl, gulls, shorebirds, and other
26
aquatic associated species helped account for the increased number of
species observed in WSIM.
Hence at low tides, species typically pre-
sent in open water habitat often frequented WSIM whereas during low
tide species that foraged In shallow water or mudflats were present.
No concentrations of birds as large and diverse were observed near Jim
Crow Sands.
Severe winter weather probably contributed to the consistently
lower number of species observed in winter for marsh habitats (Table
1).
More species were recorded in WSIM than JCSM during spring.
The
elevated nature of WSIM was possibly an important element for foraging
by water pipits (Anthus spinoletta), starlings, marsh hawks, and
possibly some shorebirds.
More species of shorebirds were present in
WSIN than JCSM during spring (Appendix 2 and 3).
The extensive
amounts of mudflats in Baker Bay adjoining WSIM attracted large concentrations of shorebirds during spring.
During high tide phases
WSIM represented suitable habitat for shorebirds where foraging sites
were exposed for a longer duration.
Jim Crow Sands marsh was exposed
for less time than WSIM and did not abut an area that offered as
extensive array of foraging substrate as Baker Bay did for WSIM.
The lack of seasonal differences in avian community characteristics was ascribed to the numerical replacement of neotropic
migrants by nearctic migrants.
Thus the exodus of hirundinids and
other passerines was counterbalanced by the arrival of shorebirds and
waterfowl.
The considerable change in species composition observed
from summer to winter emphasized this turnover (Table 4).
Avian use, particularly shorebird, of the 3 marsh habitats exhibited a relationship to habitat characteristics.
Western sandpipers
27
(Calidris mauri) preferred marshes with extensive amounts of mudflat
(JCSM) during migration; peak migration periods for this species
occurred in late summer and spring.
Concentrations of dunlins (C. alpina) were greatest in JCSM during
fall and spring migration and lowest in WSIM; MSM was represented by
intermediate numbers of dunlins.
Dunlins selected marshes with the
greatest incidence of bare ground (mudflat).
Thus vegetation probably
influenced spatial distribution of dunlins and western sartdpipers
among marsh habitats.
Recher (1966) reported that the size and den
sity of migrant shorebird populations was limited by available space,
a function of tidal inundation, rather than by available food.
Substrate composition, an important determinant of abundance and
availability of food organisms for shorebirds, also influenced distri-
bution and density of shorebirds (Recher 1966).
Page et al. (1979)
reported that dunlins and western sandpipers preferred foraging sites
consisting of moderately sorted fine sand with little or no organic
debris.
Jim Crow Sands marsh was probably a more optimum foraging
site for dunlins and western sandpipers, because its low elevation
precluded dense stands of vegetation yet more niudflats were available
at low tide for foraging shorebirds than WSIM provided.
During winter, dunlins were most numerous in WSIM possibly because
marsh sites at Jim Crow Sands and Miller Sands were covered by ice
twice during winter.
Reduced availability of estuarine tidal flats
caused by freshets resulted in reductions in number of dunlins and
least sandpipers at Bolinas Lagoon, California (Page ec al. 1979).
During spring, the differences in density of dunlins among marsh
habitats was not as apparent and probably resulted from the phenology
of marsh vegetation; e.g. decreased incidence of litter, increased
amount of bare ground, and short growth forms of most vegetation which
allowed easier access and foraging for shorebirds.
UPLANDS
The significant season by habitat interaction effect (P
0.05)
among avian community characteristics was ascribed to the variable
vegetative attributes (bare ground to dense grass/sedge cover) which
seasonally attracted different species complexes and differences in
associated habitats which influenced species present for some uplands.
Among upland habitats only WSIU:MSU and WSIU:WSIDU exhibited signif i-
cant differences (P
0.01) in avian community characteristics.
No
differences in individual characteristics were detected for these comparisons (Table 1)
. Density maximized separation of upland habitats
(Table 2), but the separation (r
= 0.48) attained on this discrimina-
tion was not strong.
Seasonally, summer differed from fall and winter; number of species and density differed (P
0.05) in each comparison (Table 1).
Winter and spring were different (P
teristics (Table 1).
0.05) with regard to all charac-
Density was the variable which contributed most
to seasonal separation Cr
= 0.69) of uplands (Table 2).
Seasonal
placement on the continuum from low to high density was winter, fall,
spring, and summer.
The migration of neotropic migrants, the prin-
cipal species comprising the summer avifauna of most uplands, probably
accounted for this pattern.
Thus the pattern established was of
migrants returning in spring, attaining peak numbers during summer
residency (breeding), declining with the advent of fall migration and
with winter representing a season of minimum use by birds.
No winter
influx of neararctic species to upland habitats comparable in number to
summer levels of neotropic species was observed.
Several species of
raptors, northern shrikes (Lanius excubitor), western meadowlarks
(Sturnella neglecta), and some permanent resident species typified
winter avifauna of most uplands.
Low densities are characteristic of
some of these species, particularly raptors.
The counterbalancing
effect observed for marsh habitats wherein passerine migrants were
replaced by waterfowl and shorebirds in a numerical sense was not
observed for upland habitats with the exception of Y1SSU.
Species com-
position of MSSU least resembled WSIU and WSID(J avian communities
(Table 3).
Avian communities of JCSU and WSIDU were quite different
also (Table 3).
Seasonal differences in species composition were
st
apparent for suinmer:winter comparisons (Table 4).
The significant season by habitat interaction effect (P c 005)
determined for upland habitat characteristics was probably attributable to the variable vegetative features of the uplands.
parison of upland habitats were significantly different (P
All coni-
0.01).
Each habitat characteristic with some exceptions, differed for each
habitat comparison.
Exceptions were percent cover of grass/sedge on
JCS1J which was similar to amounts for MSU and WSIU and covereage of
litter on NSU and WSIU (Table 7).
differed (P
Percent cover of tall grass/sedge
0.05) among all comparisons of MSIJ, WSIU, and WSIDU.
IlihIe
1.
1*rceut cor for ead IiaUtae
racrertatic kOat baud lalaid, H1112r SuL, and Jiw (5i.w Sar
IItb I t
Hatitat
8iswair
fall Water Spring tiiari
e
HS
e
(Jia rae i era It Ic"
Li L
Utter&
Ibsa/lidieii
ugbai.
Bite
1wwwr fall Whter Spring ftiau
Iaiier 1131 Ui,ter Spdng f.ui
Saitier friU WInter
Spring taii
&aIer IatLL WinUx Spring ttain
1.0
2.9
0.0
0.2
1.0$
0.5
IL!
5.2
1.4
3.9
5.4$
lo.9
42.5
f
2.o$
1.2
4.8
5.7
6.e
4.8
s.s$
92.8
91.2
93.4
93.4
92.1$
(LU
t
r
(J.o
0.3
13.1
20.5
38.2
112.5
Iu.5
30.0
13.43
1
0.0
0.0
0.0
0.0
o.o$
11.8
52.5
85.5
15.2
56.2$
0.0
(3.0
jJ
t
0.5
1.0
0.5
0.5$
33.8
112.5
23.8
iLl
14.8J
72.6
63.3
55.4
41.5
58.2$
4.3
3.0
0.6
2.9
ti
15.1
40.5
49.5
41.5
43.3$
14.5
20.5
36.6
26.4
24.5
6.1
2.4
0.8
0.2
2.1$
23.8
7
5.2
13.8
1).O
IlLS
9.9
5.2
6.1
1.9
1.9
3.9
12.1
8.2
6.3$
15.9
2/.3
41.5
31.9
2.t
1/.4
22.3
21.3
25.6
21.4$
34.7
15.9
ILL
11.4
20.5$
25.6
38.9
5.2
3.9
14.1
wsitt4
1u
813.ta Were toeifoawaI (artn
for analyate.
jrce.it cowir of litter dtfferul for all eaenal eiparIenna eaeq.t fall:apring aid wtiter;apring.
CVetce,t wait of fnra differed fOr alt senarsial c&riarieona exCpt a&aia,irIu11 ad taLpring.
cor of graiwt/aedi differed for alt weaaonal creparlerne except uu,.,er:ti1l
cor of ,aiet/Ltrieu waw nat cawildered for
aierepring. aol tati:eprtiig.
W1I3J coipailacIn.
o.os iurcenL.
(J3
C
31
Tall grass/sedge was the dominant cover of WSIDU; percent cover of
tall grass/sedge was less on WSIU and MSU (Table 8).
0.95) among upland habitats
A high degree of discrimination (r
was attained on the basis of percent cover of moss/lichen. A second
discriminated
discrinjinant function, based on the variable bare
grounds
0.91) between upland habitats. MSSU, JCSU, and WSIDU had a low
incidence of moss/lichen (Table 7) and were distinctly separated from
(r
WSIU and MSU on the first discritniriant function.
Separation of MSSU,
JCStJ, and WSID[J was based on percent occurrence of bare ground; WSIDU
(low incidence) and MSSU (high incidence) represented the extremes on
this gradient.
Observed differences in upland habitats, with respect to avian
community characteristics, were associated with those uplands exhi-
biting the most dense vegetative cover.
differed (WSIU:MSU, WSIU:WSIDU) did so
Further, those uplands that
only in a tnultivariate sense;
avian characteristics were not different univarlately.
The influence of habitat characteristics on avian communities;
although not readily apparent with respect to barren (MSSU) versus
vegetated (MSU, WSIU, WSIDU) habitats, was quite apparent in species
composition.
Species observed in MSSU primarily were associated with
aquatic (gulls, terns, waterfowl) or intertidal (shorebirds) substra-
tes. Few passerines other than swallows, horned larks (Ereniophila
alpestris), and common crows (Corvus brachyrhynchos) were observed on
NSSU although passerines comprised a major proportion of the avifauna
of other upland habitats.
32
Table
8.
Percent cover of tall grass/sedgea
(
15 cm) for NSU, WSIDU,
and WSIU, Lower Columbia River, 1978-79.
Percent cover tall grass/sedge
Sunimer
b
MSU
WSIU
WSIDU
aD
I
Fall
Winter
Spring
Mean
5.2
0.1
t
t
1.5
11.8
7.6
1.4
0.1
5.3
76.1
37.6
8.7
12.4
33.7
were transformed (arcsin fiT).
bBroken vertical lines indicate significance (P
CTrace (< 0.05 percent).
0.05).
33
The importance of MSSU to various avian species appeared to stem
from its juxtaposition to open water and marsh habitats and its lack
of vegetative cover.
The diversity index for MSSU "edge effect"
(3.74) was higher than for any other upland habitat (Table 9).
Western sandpipers, dunlins, gulls, particularly California gulls
(Larus californicus), Caspian terns, and pintails (Anas acuta) were
observed either roosting, loafing, and/or foraging on MSSU during
fall, winter, and/or spring.
Page et al. (1979), Gerstenberg (1979),
and Gill and Jorgenson (1979) reported that shorebirds utilized a
variety of habitats--barren sand dunes, sand and rock beaches,
salt
marshes, uplands, pastures, and freshwater marshes for roosting,
loafing, and/or foraging during high tide cycles.
These activities
corresponded to the observed activities of western sandpipers and
dunlins on MSSU.
Birds normally associated with upland habitats were
absent from MSSU which indicated that vegetative structure strongly
influenced avian species composition.
Use of ?ISSU by species not nor-
mally associated with upland habitats accounted for the similarity of
MSSU to other uplands with regard to avian community ctxaracteristics.
For those upland habitats that differed from each other (WSIU:MSU,
WSIU:WSIDU) individual avian community characteristics were not different.
Density however, provided the most information with regard to
discrimination among these upland habitats.
Typically, 3 to 4 species
were most abundant each season in WSIDU whereas WSIU and NSU had 2-3
numerically dominant species during summer and spring and none during
fall and winter.
Although several species present in MSU during
winter were very common, their presence was attributed to the
34
Table
9.
Diversity indicesa for "edge" of
individual
habitats,
Lower Columbia River.
Habitat
WSIM
WSIDU
WSIU
WSLB
WSITS
WSIT
Total
perimeter (meters)
10,296
1,260
7,213
6,444
7,038
6,462
Hectares
DI
54.7
3.93
11.3
87.2
1.06
2.18
19.0
4.17
48.8
33.0
2.84
3.17
38.7
1.22
21.3
3.74
19.9
13.4
MSB
2,700
6,120
5,310
5,868
2,862
MSTS
9,468
34.1
3.36
4.52
2.48
4.57
JCSM
JCSU
JCSB
4,986
3,474
4,374
29.8
19.6
MSM
MSSU
MSU
MSSB
aDiversity = total perimeter/2i
10.6
27.2
(area)
2.58
2.21
2.37
35
proximity of other habitats or to transient species passing over the
habitat.
Elimination of these species from the analysis might have
increased similarity of MSU to '1SIU.
The difference observed for WSIU:WSIDU appeared to be habitat
related.
Tall grasses typified WSIDU whereas WSIU was represented by
a short grass/sedge community.
The height and structural integrity of
WSIDU tall grasses were suitable for long-billed marsh wrens
(Cistothorus palustris) which were abundant during all seasons whereas
this species was not observed in WSIU.
Verner (1965) and Verner and
Engelsen (1970) reported that long-billed marsh wrens typically
occurred and nested in dense stands of emergent vegetation.
Song
sparrows (Melospiza inelodia), relatively abundant during fall and
winter in WSIDU compared to WSIU, were probably attracted by the pre-
sence of tall grass, dead Sitka spruce trees and invading willow and
alder trees.
Further, American goldf inches (Carduelis tristis) were
observed foraging in WSIDU during late summer on achenes of
Composites
and concentrations of goldf inches caused a relatively high
density estimate for this species.
Densities of savannah sparrows
(Passerculus sandwichensis) were remarkably similar for WSIU, WSIDU,
and MSU (Appendices 5, 7, and 8).
Starlings were quite abundant in
WSIDIJ during winter and were observed foraging on the densely matted
residual vegetation.
The greater occurrence of organic debris asso-
ciated with the densely vegetated diked upland compared to WSIU probably resulted in a greater availability of invertebrates and thus was
more attractive to starlings.
36
Edaphic features of WSIDU may account for the dense vegetative
cover present.
Deposition of silt from the tiwaco Channel dredging
operation in 1973 probably resulted in a more nutrient laden substrate
for plant growth (N. Ellifrit, USFWS, pers. comm. 1981) than other
upland habitats which were sandy in nature.
Succession appeared to be
more rapid for WSIDU as red alder and willow were pioneering the site
during the study period and by 1981 considerable growth of these trees
had occurred.
TREE-SHRUB
A significant (P
'
0.05) season by habitat interaction effect was
determined and was attributed to seasonal differences in avian community characteristics among habitats.
Since the seasonal differences
contibutitig to the interaction effect were considered explainable
biologically (see discussion, p. 40) and apparently were not attributable to random sampling error, I proceeded to test season and habitat
effects.
Tree-shrub habitats were not different with respect to avian
community characteristics (Table 1).
Species composition of all tree-
shrub habitats was quite similar (Table 3) and exhibited considerable
seasonal similarity (Table 4).
Seasonal comparisons that exhibited differences were fall:surmner
and fall:spring (Table 1).
Evenness was different for both signifi-
cant seasonal comparisons; number of species differed for the fall:
spring comparison (Table 1).
Discrimination among seasons for tree-
shrub habitats was based on species equitability (Table 2).
The
degree of resolution (rc = 0.73) among seasons on this basis was high,
37
A gradient from low (fall) to high equitability (summer) was determined on this basis.
Season placement on this gradient was fall,
Fall and winter represented seasons when
winter-spring, and summer.
flocks of birds were encountered during censusing.
Typically one or
two species were numerically dominant, hence equitability was
Breeding birds during spring and summer were dispersed,
decreased.
hence a higher equitability.
A significant (P
0.05) season by habitat interaction effect was
determined for tree-shrub habitats and was ascribed to the differing
seasonal incidence of moss/lichen.
This difference in pattern for
percent occurrence of moss/lichen was apparent for WSIT where natural
degeneration of forbs and grass/sedge led to increased exposure of the
moss/lichen layer (Table 10).
ignored.
Consequently the interaction effect was
Among tree-shrub habitats, MSTS differed from WSIT (P
for all habitat characteristics measured.
0.05)
Percent f orb and overhead
cover of MSTS were similar to WSITS, otherwise habitat characteristics
of these two habitats differed (P
0.05).
Habitat characteristics of
tree-shrub habitats on West Sand Island were different (P
except for occurrence of litter and grass/sedge.
0.05)
Canopy height (1.90
vs. 1.22) and canopy base height diversity (1.54 vs 0.58) and mean
tree circumference (0.58 vs.
than WS1TS.
analysis.
0.40) of MSTS were greater (P c 0.05)
Paucity of trees in WSIT precluded its inclusion in this
The difference in tree characteristics for HSTS:WS1TS
comparison was attributed to the presence of black cottoawoods In
MSTS.
Cottonwood typically exceeded heights and circumferences
iSbIe 10. Innt ora va1ue tot eath hatd tat
C
oi.i,riat1c, tne-ahwb h.thlt&a, 1eE thioaubia &1wr, 1918-19.
Hbitnt C6nrncter1stc
Utter
Mrf1iden
Tree-6hnib
awtat
6
wsfl
WSIT
I
1C Fall
Wtltcrd SI)rtn4
1&un
&I1u0r
}rceuL or&
Jlir
18U Winter Spring
ttnO
10.5
10.5
9.3
ILl
1231 45.5
65.2
75.2
57.4
60.81
8.2
6.6
9.3
8.2
8.11 37.6
50.5
64.3
44.5
0.4
1.0
3.2
3.9
2.11 36.6
52.5
10.8
5/.4
Saaiere
inaat
10.5
1.6
0.4
15.1
1.4
8.2
5.2
2.9
3.9
4'J.2
4.8
6.6
2.9
53
5.0
40.5
31.9
21.3
31.6
49.3
23.0
16.0
1.1
33.1
Io.Uf 50.4
I.2
15.9
21.3
re tnasfonanh arcain
baiy perwit ccr of ead hahttar parater dittenal tot alt cu1parttaia except wtCe ot .aitsa notnd.
Ch±rUnt cor of iitas/lldlen t5.rIilg awaar did not difter frtu tall and w1ztr
Iniali.
4Lt C0lt of uuas/Iithen during wli&teti did not dilfer truu fall and aprtng lea1.
cower of forb dudng aIsjIar did act differ ira fall nix! aprhig levels.
coer of gras/aod dudng wliter did not ditler fran apring law3a.
o'ir6ead cover was a
lt for a
nrapr1ng and faLleiuter cparaia.
68rdn tlnoi indiante aignificait ditferenan (F < 0.115).
iu WUILet 8pu
11.1
45.5
41.3
iu.a
si.i
32.8
64.3
343
31.9
no.4
41.8
10.4
o.8
5.9
11.8
5.2
5.01
39
normally attained by willow and red alder, which were abundant in both
tree-shrub habitats.
Percent overhead cover accounted for most separation (rc = 0.92)
of tree-shrub habitats on the first discriminant function (Table 6).
Percent cover of grass/sedge was the habitat characteristic primarily
responsible for habitat separation (r
(Table 6).
= 0.77) for the second function
Habitats ordered along the first gradient emphasized the
upland nature of WSIT (few trees hence low overhead cover).
West Sand
Island tree-shrub was distinctly separate from MSTS on the second
function because of the higher percent cover of grass than NSTS.
Although no difference among tree-shrub habitats with regard to
avian community characteristics was found, considerable differences in
habitat characteristics existed.
The presence of a tree-shrub coni
ponent in WSIT attracted species associated with forested sites and
emphasized the importance of the tree-shrub component on avian communities.
Willson (1974) determined that the addition of trees was
instrumental in attracting additional species and attributed the
increase in number of species to increased environmental patchiness.
The greater vertical diversity of NSTS compared to WSITS did not produce a significant increase in number of species.
Wilison (1974)
determined that few species were added after initial development of a
canopy layer.
Seasonal differences in pattern (season by habitat interaction)
among tree-shrub habitats for number of species, density, and diversity were attributed to:
1) ecotonal nature of WSIT,
2) probable
differential availability of cavities for nest sites (MSTS, WSITS),
40
3) associated habitats,
4) passage of neotropical migrants
(particularly parulids), and
5) density of winter residents.
The contrasting seasonal patterns (interaction)
in avian conimunity
characteristics observed for tree-shrub habitats indicated that MSTS
differed from WSIT and WSITS at least during summer and fall.
Differences were noted in number of species, density, and diversity
during these seasons for MSTS contrasted to WSITS and JSIT.
The eco-
tonal nature of WSIT, which provided habitat suitable to species
characteristic of both upland and tree-shrub habitats, probably
accounted for the high number of species recorded there and the high
density and diversity values.
Thus only the contrast between the most
similar tree-shrub habitats (structurally) remained unexplained.
The mean number of species observed in MSTS during summer was
similar to that recorded by Crawford and Edwards (1978) (7 and 8.7
respectively).
Their estimated avian density for ?ISTS exceeded mine
(8.5 versus 6.0 birds/ha, respectively) during summer yet remained
well below the level recorded during this research for WSITS.
Their
estimated density for an unidentified tyrannid was excluded from the
comparison as I attributed it to sampling anomaly.
Consequently,
the recalculated avian density for their summer data complemented the
density estimate derived during this research and solidified my contention that MSTS differed from WSITS.
More cavity nesting species were present (7 vs. 3, respectively)
and they attained a higher density (4.2 vs.
respectively) in WSITS than NSTS.
2.1 birds/ha
Density of cavity nesting species
in MSTS was 2.5 birds/ha during research by Crawford and Edwards (1978).
41
Miller Sands, approximately 50 years old (Crawford and Edwards 1978),
was characterized by relatively young trees that probably have not
attained a suitable size and/or stage of decadence suitable for pro-
duction of cavities, hence fewer opportunities for cavity nesting species.
Apparently more snags and use of cavities by birds occurred in
WSITS than MSTS.
A 1939 aerial photo of West Sand Island confirmed
the presence of trees there, while Miller Sands, visible in the
background, appeared barren.
Although larger trees were present ía
MSTS, these were cottonwoods which were still growing vigorously and
thus provided few cavities.
Large cottonwoods in MSTS seemed more
susceptible to blowdown with increasing size which may effectively
remove them as potential snags.
The presence of 3 fringillids (American goldfinch, house finch,
and savannah sparrow) in WSITS that did not occur in MSTS was possibly
attributed to the greater suitability of WSIT and WSIDU for foraging.
These species utilized WSITS primarily for perch and loafing sites and
foraged along the edges or in upland sites.
Number of species present during fall in MSTS exceeded that of
WSITS primarily because
re parulids were observed.
warblers observed in MSTS were not observed in WSITS.
Five species of
Warbiers were
not observed during fall in FISTS by Crawford and Edwards (1978).
The
difference in number of warbler species may have reflected a difference in migratory patterns between FISTS and WSITS.
Bagg (1969)
reported that chickadees tended to funnel along shorelines upon
encountering large water bodies until a shorter crossing point or land
bridges were encountered.
The decreased number of species during fall
42
in WSITS and WSIT was attributed to the seasonal exodus of neotropical
migrants.
Density was the characteristic which exhibited the most contrast
between MSTS and West Sand Island tree-shrub habitats (WSIT, WSITS)
during fall.
The higher density of black-capped chickadees and ruby-
and golden-crowned kinglecs in MSTS compared to WSIT and WSITS appeared
most responsible for this observed difference.
Black-capped chicka-
dees were considerably more numerous during fall than summer in 1STS.
Crawford and Edwards (1978) also reported higher densities of blackcapped chickadees during fall than summer.
High estimated densities
for both species of kinglets during fall as determined here were not
reported by Crawford and Edwards (1978).
Their fall sampling period
occurred during an unseasonably dry period which may have delayed the
migration of kinglets to lower elevation areas such as MSTS; however
their data indicated a considerable increase in ruby-crowned kinglets
with the onset of winter.
The habitat factor responsible for these
differing densities was not apparent.
Differences in equitability among seasonal comparisons were
attributed to the very high densities attained by a few species during
fall.
Black-capped chickadees and ruby- and golden-crowned kinglets
flocked together during fall and typically were the most numerous
birds.
Consequently, equitability during fall was lower in contrast
to spring and summer when flocking was less prevalent; reduction in
flocking behavior was probably attributable to territoriality and
absence of some species (e.g. kinglet spp.) that occurred with tue
advent of spring and the breeding season.
43
BEACH
Avian community characteristics of beach habitats were similar for
habitat, season, and interaction effects (Table 1).
Species com-
position of beach habitats was quite similar although WSL3 exhibited a
consistent difference (20 percent) from all other beach nabitats
(Table 3).
This difference probably reflected the occurrence of
pelagic species at WSIB.
re
Seasonally, species composition of beach
habitats was most different between summer and winter (Table 4).
Although species composition exhibited a marked seasonal turnover,
avian community characteristics were not significantly affected.
As
in marsh habitats, the migration of neotropical migrants was negated
by migrant nearctic species.
Bare ground, the only characteristic considered in comparisons of
0.025) on WSIB than on
beach habitats, was significantly lower (P
other beach habitats (Table 11).
Accumulation of debris on WSIB was
probably enhanced by its exposure to the main channel of the Columbia
River; other beach habitats were not exposed to the main channel.
This difference apparently was of minimal importance to birds.
WEST SAND ISLAND
The significant interaction effect (P
'
0.05) associated with ana-
lysis of avian community characteristics was attributed to the diverse
array of habitats considered and varying seasonal utilization of these
habitats by birds.
Avian community characteristics differed for vir-
tually all habitat comparisons (P < 0.05) except WSIM:WSIU, WSIDU:WSIB,
44
Percent occurence of bare ground for 4 beach habitats:
Table 11.
MSSB, MSB, JCSB, and WSIB, Lower Columbia River, 1978-79.
Percent bare grounda
Summer
Fall
winter
Spring
Mean
97.1
97.4
94.8
98.5
96.9
MSB
99.2
93.8
91.8
94.8
94.9
JCSB
99.3
87.5
98.0
96.4
95.3
WSIB
92.3
93.4
76.1
95.2
89.2
Habitat
MSSB
b
aData were transformed (arcsin JiI).
bverticai lines connecting habitats indicate similarity; broken lines
indicate significant difference.
45
WSIT:WSIB and WSIT:WSITS.
Typically, species equitability and density
were higher f or the tree-shrub habitats than for other habitats (Table
12).
A continuum from low to high density formed the primary basis for
separation (r
= 0.69) of West Sand Island habitats (Table 2).
Habitats were ordered along the first function beginning with tISIU,
then WSIDU and WSIM, WSIB, and terminating with tree-shrub habitats
(WSITS and WSIT).
Equitability formed the basis for separation on the
second discriminant function.
This separation (r
0.48) enhanced
the distinctiveness of the tree-shrub habitats from all other West Sand
Island habitats.
Species composition of West Sand Island habitats was relatively
different for most comparisons (Table 13).
Species composition of
tree-shrub habitats differed from other West Sand Island habitats most,
although species composition of WSIT was similar to uplands (Table
13).
Considering that WSIT retained many similarities structurally of
an upland habitat, some similarity in species composition to uplands
was expected.
Species composition of WSIM differed considerably from
other West Sand Island habitats (Table 13).
Seasonally, species composition during winter was quite different
from summer; other seasonal comparisons were similar (Table 14).
The
changeover from summer to winter reflected the impact of migration on
species composition.
Tree-shrub habitats, the most advanced successional stage on West
Sand, were readily distinguishable from more intermediate stages.
Evenness and density were the characteristics that best separated
T1
Table 12.
Mean values of each avian community characteristics for
habitats comprising each island, Lower Columbia River,
1978-79.
Habitat
complex
Number of
species
Densitya
(birds/ha)
Diversity
Evenness
Miller Sands
175d
MSMb,c
137b,c
MSSUb
82b,c
1.28
MSUb
35b
1.36
7.2
13.5
068b,c
064C
o.00c
1.38
O.6O'
1.62
0.68
MSSB
10.6
MSB
11.3
MSTSb,c
109b
12.4
1.84
JCSM
11.4
12.4
1.42
0.59
JCS1J
8.7
4.6
1.12
0.50
JCSB
11.5
41.5
1.39
0.58
161d
059e,f
7.Oc
078b,c
Jim Crow Sand
West Sand Island
152e,f,g
92e,f
10.3
112e
WSITe
11.6
14.0
7.2
5,8,g
1.24
36f,g
1.43
8.4
1.52
lO.4f
1.90
12.3
1.93
0,66g
063g
074e
47
Table 12. (continued)
aDensity data were transformed (square root).
bMSM differed from MSSU, MSU, and MSTS.
Number of species differed in
each comparison; evenness differed for MSM:MSTS comparison.
CMSTS differed from all other Miller Sands habitats.
MSTS charac
teristics which differed among habitat comparisons were number of
species (MSM, MSSIJ), density (MSU, MSB), and evenness (MSM, MSSU,
MSU, MSSB).
dDiversity data were not utilized in analysis because of high
correlation with evenness:
Miller Sands (r = 0.74), West Sand Island
(r = 0.76).
eWSIM differed from WSIDU, WSIB, WSIT, and WSITS.
Number of species
was different from WSIDU and WSIB; evenness differed from WSIT and
WSITS.
WSIT differed from WSIDiJ, WSIU, and WSIM.
from WSIDU and WSIM:
Number of species differed
density differed from WSIDU and WSIEJ; and
evenness differed from WSIM and WSIDtJ.
differed from WSIM (number of species), WSIDEJ (density,
evenness), WSIU (densisty, evenness), and WSIB (evenness).
48
Table 13.
Habitat comparisons of species composition values
(MacArthur's Difference) for habitats grouped by island,
Lower Columbia River, 1978-79.
Habitats
Island/habitat
Miller Sands
MSM
MSSU
{SU
MSSB
MSB
MSSU
1.16
MSU
1.40
1.45
MSSB
1.16
1.10
1.42
MSB
1.16
1.23
1.37
1.20
MSTS
1.82
1.87
1.52
1.89
1.70
JCSM
JCSU
WSIU
WSIB
WSITS
Jim Crow Sands
JCSU
1.57
JCSB
1.16
1.32
WSIM
WSIDU
West Sand Island
WSIDU
1.49
WSIU
1.52
1.37
WSIB
1.53
1.63
1.45
WSITS
1.69
1.59
1.65
1.75
WSIT
1.57
1.37
1.39
1.62
1.13
49
Table 14.
Seasonal species composition values (MacArthur's
Difference) for combined island habitats, Lower Columbia
River, 1978-79.
Habitats group by island
Season
Habitat
complex
Summer
Fall
Winter
Miller Sands
Fall
1.40
Winter
1.58
1.34
Spring
1.27
1.36
1.31
Jim Crow Sands
Fall
1.64
Winter
1.84
1.32
Spring
1.44
1.39
1.37
West Sand Island
Fall
1.35
Winter
1,64
1.22
Spring
1.15
1.26
1.45
50
tree-shrub habitats from other West Sand Island habitats.
Concentra-
tions of waterfowl, shorebirds, or gulls commonly occurred in JSIM or
WSIB which tended to depress equitability.
A tendency for more spe-
cies to occur in WSIM than other West Sand Island habitats, except
tree-shrub habitats, was probably associated with edge effect.
The
high elevation, which precluded inundation and resulted in habitat
similar to uplands, was probably attractive to upland species
MILLER SANDS ISLAND
Terrestrial habitats on Miller Sands (MSsU, MSU, MSTS) were dif-
ferent from MSM, whereas avian community characteristics for beach
habitats were similar to MSM (Table 12).
Another pattern that deve-
loped concerned MSTS, which differed from all other habitats on Miller
Sands (Table 12).
Number of species differed for each significant
comparison except beaches (MSB and MSSB):MSTS and MSIJ:MSTS (Table 12).
Avian density for MSTS differed from that determined for MSU and MSJ3
(Table 12).
Evenness was higher for MSTS for all comparisons except
MSB:MSTS (Table 12).
Number of species best described habitat separation (r
for the first discrinilnant function (Table 2).
= 0.62)
A habitat continuum
was developed with upland habitats (fewest species) and MSM (most
species) most distant from each other.
Habitats were scaled from low
diversity to high, i.e. uplands, beaches, marsh, and tree-shrub along
the second function.
Species composition of MSTS differed considerably from that
recorded for other Miller Sands habitats (Table 13).
All habitats
51
subject to tidal inundation and MSSU and MSSB were quite similar in
species composition (Table 13).
Seasonally, species composition
during winter was most different from summer (Table 14) and reflected
the impact of migration on composition.
Two patterns were evident from comparisons among Miller Sands
habitats:
1) MSTS differed from all other habitats, and
2) MSM dif-
fered from terrestrial habitats on Miller Sands but not beach habitats
which were subject to the same tidal influence.
An increase in avian
community characteristics associated with a more advanced successional
stage seemed apparent for Miller Sands habitats.
Species equitability
for MSTS was greater in most significant comparisons (4 of 5) than for
other Miller Sands habitats.
Large, often monotypic flocks, of either
waterfowl, gulls, or shorebirds commonly occurred in marsh and beach
habitats.
Such aggregations tended to depress equitability.
Miller
Sands spit upland resembled beach and marsh habitats regarding aggregations of birds.
Low densities of birds during fall and winter in
MSU often were dominated by one species, hence equitability was low.
Miller Sands marsh differed from terrestrial habitats associated
with Miller Sands apparently because of its attractiveness to aquatic
associated species and certain terrestrial species.
Shorebirds,
gulls, waterfowl, and swallows, species which utilized MSM at varying
stages of tidal inundation and seasons, formed the basis for the
higher species richness components recorded for MSM than was recorded
for terrestrial habitats.
These species complexes also frequented
beach habitats which were similar to MSM with regard to avian com
munity parameters.
52
JIM CROW SANDS
Habitats on Jim Crow Sands were similar in avian community characteristics and only JSCM:JCSU comparison exhibited any notable difference in species composition (Table 13).
Species composition
changed from summer to winter reflecting the influence of migration on
community composition (Table 14).
The structural simplicity of Jim
Crow Sands habitats, related to their early successional stages and
age possibly accounted for the lack of observed differences among
these habitats.
CONCLLTS ION
Structural nature of the vegetation for the habitats investigated
influenced avian species composition and community parameters.
Differences were apparent for comparisons of various habitats and
among habitats.
Later successional stages, typified by greater
structural complexity, tended to support more species and attain
higher densities and equitability.
Seasonal changes in avian com-
munity parameters associated with the influx and exodus of neotropic
and neararctic migrants were also of importance.
MANAGEMENT RECOMMENDATIONS
Management decisions, with respect to these islands, should be
based not only on gross habitats present, but the structural nature of
each habitat and its juxtaposition to other habitats.
Changes in spe-
cies composition and parameters were quite apparent as vegetative
53
structural qualities changed.
Thus a manager has the capacity through
vegetative manipulation to alter avian species composition and parameters on these islands.
iiowledge of the species present and their
associated community parameters can aid in predicting the outcome for
events, such as dredged material deposition, that alter these habitats.
Differential seasonal use of habitats by birds points out the
necessity for managers to consider these habitats on a seasonal
basis.
Thus management decisions can be made apriori with regard to
species composition and community parameters desired.
54
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National perspective of colonial waterbirds
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A
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Breeding success of great
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Bird use and vegetation
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Shugart, H.H., Jr. and D. James.
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343 pp.
Ecological succession of
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1965.
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Breeding biology of the long-billed marsh wren.
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61
Woodward-Clyde Consultants.
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1976.
Terrestrial ecology of the Miller
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1975.
Competitive release in island song
Theor. Pop. Eiol.
5:42-58.
S aDIaNdcW
1.
Dowttcher app.
Ijeatern sandpIper
Sanderling
Northern phularope
Glaucous gull
Glaucous-winged gull
Western gull
flerrtog gnU
California gull
hung-billed gull
Hew gull
Bonaparte's gull
Casplan tecu
Belted kingfisher
Veux'a swift
black-b11ted plover
Spotted sandpiper
Greater yellowlege
Lesser yelluwlega
Least sandpiper
Dunlin
K1tt&ieer
Scmipaltaated plover
Anerican coot
Utd eagle
Coiwaon mergauaer
Red-breasted
rganser
24.5
4.0
39.8
43.3
4.8
10.9
6.6
158.6
21.3
4.2
4.8
July
15.1
5.3
12.4
34.2
2.6
2.8
June
1.9
101.1
0.8
1.6
30.2
33.4
3.2
15.9
235.8
23.7
5.1
6.5
40.0
11.0
2.8
11.4
14.6
59.7
9.9
6.4
0.7
1.9
Septewber
230.6
5.3
August
1.8
29.0
4.9
1.0
14.6
2.11
34.7
3.9
28.9
6.4
24.7
5.0
4.6
0.7
1.9
22.4
1,409.5
397.9
5.0
2.8
9.5
0.1
5.0
94.4
9.9
109.0
20.1
86.1
5.0
43.11
6.4
October Nuvewber
8.8
1.4
5.6
5.6
3.8
5.0
5.0
15.1
130.7
20.2
13.2
35.4
35.8
13.0
1.6
29.3
9.3
5.0
2.4
25.5
143.11
7.3
19.0
132.7
0.4
3.2
3.0
3.2
1.6
3.9
0.6
March
38.2
8.9
1.6
6.4
2.11
10
ebreary
0.2
2.5
1.4
3.2
bceaber January
Mouth
Species den1ty (blrde/l00 ha), June 19711-Hay 1979, mIter Saw.Iu Harsh, Cotuebte tUver, Oregon.
Uorned grebe
Western grehe
Døuble-crcted coraoraut
Crest blue heron
Whistling swan
Canada goose
Mallard
Gedeell
Ptntati
Green-winged teal
American wigeon
Greeter eceup
Lesaer ecaup
BufCLehead
Species
Appendix
00.1
35.7
13.2
20.2
i.s
10.3
135.4
752.5
9.5
0.4
4.0
Apr.L
2.6
4.5
4.5
12.0
21.9
106.1
430.6
3.11
25.5
11.8
5.3
19
tay
cy
1. (contInued)
Rveoneaa
In
250.3
2.06
0.14
19
486.8
2.29
0.78
11
651.2
1.48
0.62
13
499.8
2.03
0.79
13
621.1
1.65
0.64
Llen8ity (birds/ha)
Olveratty
35.4
Number of species
3.2
25.5
4.8
47.1
4.8
0itober
12.7
Septuwber
159.2
August
33.9
July
111.9
319.1
9.1
June
85.8
31.0
70.7
Violet-green wa1low
Tree swallow
Ram swallow
Cliii swallow
Common crow
Rrowa-headed cowbird
Song sparrow
SpecIes
Appendli
(1.46
1.31
2,235.9
17
7.2
Novesbet
40.2
2.09
0.91
It)
Decewber
Month
301.2
1.95
0.74
14
20.1
Januaty
79.8
1.62
0.74
9
February
405.5
1.56
0.69
10
254.6
54.1
March
In
721.2
4.57
0.57
17
4.8
80.4
2n.0
flay
2,392.9
1.85
0.48
16.1
72.7
Q75.0
104.8
Apr11
Appendix
2.
lpeclea density (birda/lOO ha), June 1978-Muy 1919, Jtiu Crow Sands Harsh, Coluwbta River, Oregon.
Houth
Species
June
July
August
SepLewbur
October
Novewber
Conenu loon
Oecaber January
1.6
Double-crested coranrant
Great blue heron
0.4
Whistling swan
Canada goose
tbtury
7.1
Piutati
Green-winged teal
Macrican wigeon
Northern hoveier
11.7
5.6
50.7
12.9
31.5
93.9
1.6
6.5
3.2
31.8
163.5
1.6
Surf acoter
3,9
0.8
47.4
0.8
18.4
17.7
15.8
12.7
31.8
3.2
35.4
3.2
70.3
462.6
4.4
260.7
681.8
83.9
4.4
4.9
12.6
1.6
6.5
7.1
2.4
5.4
Bonaparte's gull
69.7
241.1
Casptau tern
Violet-green swallow
120.5
11.8
2.4
35.9
18.0
20.2
0.8
60.5
9.0
6.6
3,351.9
61.7
19.9
33.6
3.0
6.5
70.5
1,274.5
8.8
329.4
44.2
13.2
2.8
2.8
3.9
8.8
6.6
3.9
246.2
13.1
8.1
7.3
5.1
66.2
14.7
63.9
1.8
5.4
4.4
225.8
6.4
6.5
2.4
26.9
42.1
4.0
49.7
12.7
15.1
21.4
39.0
39.8
20.i
Lb
11.1
327.2
0.8
hO.!
2,931.2
8.8
24.3
1,340.5
40.7
2.6
2.2
52.0
4.2
Cons&on crow
3.2
52.4
6.5
Western sandpiper
Sauderitag
Northern phalarope
Glaucous-winged gull
Western gull
Cliff swallow
0.6
7.9
2.5
27.5
I3unlin
Tree swallow
Barn swallow
5.6
4.0
5.4)
Coajan iaerganaer
Red-breasted merganser
Merlin
Semtpslinated plover
Cossion tern
8.9
0.6
24.5
Greater acaup
Duitleheati
New gull
Hay
21.2
62.9
Mallard
herring gull
California gull
King-billed gull
Aptil
13.5
32.5
Snow goose
KilIdeer
Spotted sandpiper
Greater yellowlegs
Least sandpiper
Hatch
9.0
2. (continued)
Evenness
Nuather of species
Denolty (birds/ha)
Dtveretty
Long-billed tharab wren
Water pipit
Aiaerican goldfinch
Savannah sparrow
Song aperrow
Spectea
Appendix
259.4
1.89
0.82
10
June
14
19.9
Auguat
543.2 1869.2
1.64
1.59
0.62
0.62
13
JuLy
263.0
2.09
0.79
14
38.2
SeLember
0.71
1.411
303.1
8
35.4
12.7
5.068.L
0.94
0.41
10
October Novebr
13
719.5
1.42
0.56
9
January
134.2
1.30
0.59
19.9
79
becewbet
Month
0.80
1.84
12.o
1(1
Iebcuaty
10
359.8
1.24
0.54
Match
3,981.0
1.01
0.43
12
25.5
April
0.21)
32.0
0.52
14
May
th
Appendix
3.
Species density (birds/lOU ha), June 1978-May 1919, WeSt Sand tslsnd flaroh, Coluabia Flyer, Oregon.
liontim
Species
Common Loon
Double-crested cormorant
Pelagic corworatit
June
July
1.4
5.3
2.2
19.9
1.6
Great blue heron
Mallard
l'intail
4.2
AuguSt
1.4
September
1.0
1.9
0.4
22.2
0.8
Green-winged teal
EuropeSu wigeon
&uiartcan wigeon
Northern shoveler
Greater aceup
Buff lehead
Red-breasted mnerganser
Rough-legged hawk
Marsh hawk
Seamipalamated plover
Kilideer
Black-bellied plover
2.2
October Novewber
1.8
22.2
3.2
0.8
12.4
6.4
23.9
51.7
January
February
March
April
4.8
31.4
46.9
0.9
1.7
1.3
0.7
17.1
1.0
0.6
0.8
17.5
10.3
1.4
16.1
0.8
66.5
24.1
2.1
2.8
24.5
1.1
19.9
38.2
0.4
4.4
2.2
0.2
0.8
6.4
3.2
9.5
6.4
0.8
12.7
8.8
6.8
2.6
65.1
3.2
2.6
155.9
417.8
234.7
16.2
6.4
811.3
406.8
44.2
1.4
Greater yellowlega
Least sandpiper
2.2
11.2
204.1
135.5
Dumilin
Dc,wttcher app.
Western aandplper
Herring gull
California gull
Ring-billed gull
41.7
12.9
213.7
32.5
23.0
17.4
1.6
11.9
Rufous hwaaiogbtrd
Violet-green swallow
Tree swallow
97.3
8.6
161.1
378.8
481.9
4.3
258.6
1.5
0.7
1o3.9
15.6
32.3
287 .6
245.2
3.2
2.4
21.1
2.8
0.8
4.2
3.2
15.3
1.8
0.2
9.1
19.6
318.3
10.8
60.0
1.4
21.5
28.7
1.4
23.4
1.4
4.3
4.1)
7.9
Mew gull
Bonaparte's gull
Caspian teru
Belted kingfIsher
1.4
ho
Whimbrel
Sanderling
GIaucou-winged gull
Western gull
Hay
1.4
1.4
113.9
21.5
1.6
2.8
Oecmbec
1.9
21.0
5.3
8.0
3.2
1.9
318.3
70.9
53.3
318.3
04.5
40.3
(contInued)
Eve nueaa
Number o spectea
Density (birds/ha)
Diversity
American goldfinch
Savaunab sparrow
Song sparrow
flouae finch
Red-winged blackbird
Common ylowtbroat
St ant eg
Long-billed uiarah wren
Water pipit
Comun crow
barn awallow
Cliff wmiIow
Spectea
3.
-
AppendLX
S59.5
1.58
0.56
19.9
5.7
6.4
39.8
8.8
June
11.9
4.8
20
868.1
16
329.3
2.09
0.75
11
517.6
2.03
0.72
14
564.1
1.64
0.62
0.67
1.99
i2.7
37.4
7.2
0ctobr
101.9
35.4
19.9
Seplember
28.7
August
39.8
79.6
25.5
9.9
39.8
39.8
5.7
30.9
0.4
4.8
19.9
July
1.53
0.57
1.55
0.59
7
161.6
0.77
0.40
13
43ó.3
0.66
0.26
14
655.4
15
ebruary
834.4
3.2
January
12.7
286.5
beceaber
35.4
9.7
November
Hoth
U.3
1.46
10
sOh.5
12.7
March
20
2,415.9
2.14
0.00
50.9
k2.7
3.2
17.7
april
1.ng
0.73
.5
13
0.5
10.1
2.4
9d.5
May
0'
-4
Appendix 4.
Species density (birds/100 ha), June 1978-May 1979, Miller Sands Spit Upland. Guheubta 1(iver, Oregon.
Month
Species
June
July
Doubl-creuted cormorant
August
1.4
Canads goose
September
October
November
1.4
6.0
Snow goose
0.4
3.9
2.2
35.4
Common iaerganaer
Bald eagle
Semipalmated plover
1,170.3
280.5
110.6
13.9
308.2
28.3
139.5
631.4
Water pipit
Diversity
Evenness
31.2
43.6
9.5
11.9
6.8
1.0
8.8
242.1
18.1
4.2
4.2
42.7
103.7
113.2
7
1,317.5
1.44
0.74
3.9
67.')
1.0
62.2
11.2
44.8
1.9
12.7
1.4
35.4
2o .1)
9.9
6.4
39.2
11.0
9.9
24.9
15.9
263.6
1.5
4.2
4.2
I.9
9
12
496.6 3,357.8
1.46
0.88
0.66
0.35
9
10.9
1.48
0.68
1.8
4.4
6.3
33.4
14.4
8.4
59.1
51.3
119.4
3,565.8
370.4
2.1
1.6
Common crow
Density (birds/ha)
0.1
11.1
2654.4
Common raven
Number of Species
3.9
8.8
6.4
139.7
Mew gull
Cliff swallow
1.0
91.3
Dun liii
Morned lark
Violet-green swallow
Tree awallow
barn swallow
May
0.4
Least sandpiper
Caspian tern
AprIl
15.11
79.6
Whimbrel
California gull
tng-Stlied gull
larch
5.7
50.9
0.1
0.1
Marsh hawk
Herring gull
ebivary
331.4
Crecu-winged teal
Greater scaup
buff 1ehed
Western sandpiper
Sauderling
Northern phalarope
Glaucous-winged gull
Western gull
3.8
January
0.3
Pintali
Kilideer
black-befli ad plover
December
3.2
3.1
0.4
0.1
0.8
245.6
145.6
109.2
56.6
18.2
36.3
11.7
35.4
9
4
142.7
0.83
0,60
1,844.3
1.01
0.46
1
10
4
8
21.8
1.23
0.63
213.8
1.69
0.74
67.5
1.21
0.87
100.2
1.65
0.79
9
10
651.6 4,u91.b
1.63
0.8/
0.74
0.38
S.
brown-hedd cowbird
Starling
Yellow-romped warbler
Townsend's warbler
Wileon'a warbler
Western weadowlark
Cedar wawtng
Violet-green swallow
Tree swallow
Rant swallow
Cliff swallow
Common crow
block-capped chickadee
American robin
Golden-crowned kiuglet
Ruby-crowned kinglet
Rufous huiai ngbird
Common f licker
Common nighthawk
CaSpiaft tecn
flew gull
Piniall
American wtgeoD
Cooper's hawk
Red-tailed hawk
bald eadle
Peregrine falcon
Merlin
American kestrel
Common snipe
Dunlin
Dowiichter app.
Western sandpiper
Sanderling
Glaucous-winged gull
Western gull
California gull
Ring-billed gull
Mal lard
26.5
8.0
12.7
19.1.
9.5
318.3
12.1
1.6
June
8.8
74.7
19.9
62.9
19.1
0.4
10.5
July
Species density (birds/LOU ha)
Crest blue heron
Species
Appeadtc
79.6
2.6
8.8
143.2
11.0
32.6
10.4
2.6
AuguSt
42.6
114.0
19.9
41.4
31.6
Septesber
1.0
25.5
12.1
1.0
12.7
25.5
[(.1
6.6
3.2
5.0
Noveabnr
12.7
22.0
65.9
25.5
12.1
3.9
2.2
13 0
October
Month
3.9
1.4
12.1
151.2
2.4
Oeceldber
2.6
13.0
3.1
12.19
1.5
144.4
211.1
0.8
13.0
January
15.7
3.1
February
tune 1918-May 1919, Miller Sands Uici. Coinebia lUver, Oregon.
5.3
2.6
8.8
5.3
15.6
81.5
March
623.3
140.8
bO.3
25.5
59. 1
3.9
35.4
35.4
19.5
April
21.1
35.4
bl.2
218.5
5.3
3.2
u.S
Nay
Appendix
5. (contInued)
tIe at
Species
June
July
Aiserican goldfinch
Savanoah sparrow
Dark-eyed Junco
White-crowned sparrow
Song sparrow
7.9
448.2
358.1
Number of utpeetea
Density (birds/ha)
Diversity
Evenness
60.3
August
3.9
89.1
6.5
537.1
38.2
IL
12
10
924.2
1.33
616.2
1.58
0.56
0.63
872.1
1.28
0.56
Septeather
35.0
October Nuveutber
beceinber
January
Februaty
Harct
33.7
6.4
toe.o
Li.?
11.7
Hay
211.o
39.8
392.6
1.15
U
180.5
1.91
0.90
0.80
1
April
8
117.0
1.80
0.87
6
195.3
0.74
0.41
10
811.0
1.47
0.64
6
18.8
0.45
0.65
129.8
1,08
0.60
11
1211.7
1.65
0.69
8
630.8
1.36
0.ob
6.
Cliff swallow
Common raven
Common crow
Ruby-crowned kinglet
Water piptt
Starling
Orange-crowned warbler
Red-winged blackbird
brown-headed cowbird
House finch
AmerIcan goldfinch
Rufous-sided towhee
White-crowned sparrow
Song sparrow
bgtii swallow
Tree awllow
American kestrel
glildeer
Spotted sandpiper
Dunlin
Dowitcher app.
Claucous-witiged gull
Western gull
Herring gull
California gull
Ring-billed gull
Mew gull
Ceaptan tern
Rufous hutsalngbtrd
Downy woodpecker
Western flycatcher
Horned Lark
Violet-green swallow
Karb hawk
0.8
7.9
20.5
13.1
0.8
59.1
25.5
12.1
127.3
38.2
6.5
6.4
3.2
9.5
July
6.4
35.4
12.1
136.9
12.7
32.6
15.6
19.6
June
39.8
14.8
1.9
12.7
6.4
684.4
3.8
5.0
3.9
August
3.2
5.0
46.2
26.5
9.4
62.4
2.2
10.1
19.9
3.9
2.6
3.9
Septesiber
8.8
6.5
76.3
99.5
October
Houth
636.6
'2.0
3.2
9.9
Oeceaber
5.0
13.0
2.8
*2.1
0.4
January
'
144.2
5.0
1.4
0.4
Lebruaty
Upland. Coluwbia giver, Oregon.
Novesber
SpecIes density (birds/tOO ha), June 1918-Hay 1919, JIsi Crow Saad
Canada gooae
Snow goose
Mallard
Pintati
Greater ucaup
Red-breasted isergauser
Species
AppendIx
u.S
5.1
19.9
5.9
'23.0
3.?
3.?
ltarcli
6.5
lob.0
119.4
5.3
8.8
2.8
8.8
13.2
14.9
114.3
45.1
2.7
11.8
24.4
13.0
15.9
April
39.8
05.0
179.0
122.5
59.1
0.1
1.4
3.8
110.8
19.9
41.0
41.0
00.9
41.0
35.4
19.9
Hay
A1ped1x
6. (coiittnud)
Mouth
July
Steabur
Spectea
June
Number of species
Density (bIrds/he)
DiversIty
Evenness
it
11
10
12
417.9
1.95
0.81
246.1
1.65
0.69
184.3
0.62
0.21
otl.9
1.18
0.48
AuMuat
October
4
191.9
0.96
0.70
November
3
642.4
0.06
0.05
Decewber
January
1
5
9.9
0
83.3
0.82
0.51
I)
FeD&uary
-4
bI.0
0.22
0.16
March
1
69.1
1.bS
0,86
Att1
ay
lb
20
573.4
113e.b
2.Uo
2.30
0.74
0.77
N.)
Appendix
7.
Spccies density (blrds/l00 he), June 1978-May 1919, West Saud Island Uled Upland. Coluwbia
1ver, Oregon.
Month
Spedee
Double-crested cormorant
Great blue heron
Mallard
Red-tailed hawk
Marsh hawk
Bald eagle
Merlin
ICilideer
June
Rufous hu.waingbicd
0.8
Septeuber
October
Number of specIes
Density (birds/ha)
Diversity
Evenness
1.6
Decesber
3.2
1.6
January
February
March
april
Play
2.4
6.5
2.4
15.1
1.6
3.2
13.0
1.6
2.4
0.8
5.0
8.8
16.4
1.4
7.1
61.3
2.8
19.9
3.2
4.7
0.8
12.7
35.4
5.9
2.8
17.9
2.8
10.0
6.5
10.0
88.4
3.2
79.6
636.6
19.9
1.0
19.9
2.6
1.3
1.3
11.9
19.9
36.2
22.3
114.6
7.9
4.8
159.2
3.2
79.6
10.3
238.7
3.2
2.2
59.7
19.9
1.3
39.8
8.8
389.0
i41.
75.2
84.4
2.o
8.8
4.2
3.9
0.8
19.7
59.1
9.9
58.5
5.3
19.8
71.5
3.1
233.9
6.2
2.6
2.6
75.2
IIue finch
AmerIcan goldfinch
Savannah sparrow
.White-crowned sparrow
Song sparrow
Novewber
0.8
0.8
A,aerlcan robin
Nottbern shrike
Starling
Western meadowlark
Red-winged blackbird
August
0.8
flack-bellied plover
Glaucouw-winged gull
Western gull
California gull
Ring-billed gui1
Mew gull
Short-eared owl
Coamon flicker
Downy woodpecker
horned lark
Violet-green swallow
Tree swallow
Barn swallow
ClIff swallow
Purple martin
Common crow
Long-billed marsh wren
July
3.2
62.9
397.9
12
314.4
601.9
12.8
10
721.0
14.8
239.7
212.2
3.2
37.4
39.0
8
1,415.9
1,113.8
1,368.7
1.50
1.24
1.32
0.60
0.5
0.63
12
:193.6
1.54
0.62
ii
272.8
1.88
0.78
298.1
32.5
11
520.6
(.06
0.44 3
34.8
7
123.1
1.08
0.55
14.9
8
267.6
0.54
0.26
39.8
7
193.6
0.81
0.42
20.0
7
212.9
1.25
0.64
119.0
229.2
39.0
11
449.1
1.11
0.49
6
592.1
1.45
0.81
8.
Yellow-ruisped warbler
Common yetlowthroat
Starling
Cedar waxwing
Townsend's solitaire
black-capped chickadee
American robin
Common crow
Purple matttn
Horned lark
Violet-green awallew
Tree swallow
barn swallow
Common fLicker
Caspian tCrn
bonaparte's gull
Mew gull
California gull
Ring-billed gull
Lesser ycilowlegs
Sandarling
Ciaucous-wlnged gull
Western gull
Herring gull
Common snipe
Whimbrel
bald eagle
Asierican keatrel
Kilideer
Marah hawk
Sliarp-ahiiuied hawk
Cooper's hawk
Red-tailed hawk
bufflehaad
Pintail
10.1
21.5
38.6
15.9
44.6
11.8
5.6
19,9
35.4
17.0
2.8
31.4
June
17.7
4.0
31.8
19.1
19.1
9.5
1.3
10.5
14.0
2.8
0.2
26.0
1.6
July
19.9
2.6
3.2
251.1
6.4
2.2
11.3
48.1
6.9
1.0
3.9
1.9
3.6
Auguat
31.4
8.1
3.2
0.3
32.2
9.4
Sepi.iaber
3.9
6.6
0.4
9.0
6.4
19.9
13.0
3.2
5.3
5.0
October Nuvewber
Month
9.4
4.2
u.S
0.8
1.4
5.0
12.5
0.8
I.!
0.2
Deceabet January iebruary
Species density (blrd/I00 h.i), June 1978--May 1979, Weat Sand laland UplCnd, Coluwbia River, OreRwi.
Double-crested cormorant
Pelagic cormorant
Mallard
Common loon
Species
Appendix
b.8
63.1
21.5
34.8
1.1
4.6
0.8
March
2.6
12.1
10.5
3.0
2.8
34.8
39.0
19.9
8.8
13.0
ApriL
23.1
34.2
4.2
29.6
86.7
1.9
35.4
6.4
ay
8. (cont:tnucd)
Nwuber of species
Density (birds/ha)
Diversity
Evenness
Wilson's warbler
Western meadowlarit
House finch
American goldfinch
Savannah sparrow
White-crowned sparrow
Song sparrow
Lapland iongspur
SPeCIeS
Appendix
0.50
1.42
16
765.8
1.38
0.50
17
688.0
381.1
16
2.1
15.7
184.1
1.70
0.77
9
6.5
0.10
36.2
1.13
S
3.2
October
55.0
457.6
5.0
54.5
Sepiewber
21.1
6.5
August
15.7
July
965.9
1.93
0.70
19,5
159.2
86.5
437.7
June
94.2
1.98
0.90
9
6.5
26.0
Novesber
12.0
0.52
0.75
2
2.6
Uccuwbct
Month
0.83
0.47
6
70.2
2.6
54.7
January
9
16s.O
1.70
0.11
1
19.9
.5
hatch
no.0
1.27
0.65
5.0
31.6
february
1.31
0.49
591.
15
2.7
6.5
4W.0
6.5
April
1.91
0.71
bTh.6
Ii
Lj.0
59.1
193.6
May
cormorant
House finch
Water pipit
Conimon crow
Cliff swallow
Rough-winged swat low
8am swallow
Horned lark
Violet-green swallow
Tree swallow
Ronaparte'a gull
CapJan tern
Hew gull
Herring gull
California gull
Ring-billed gull
Northern phalarope
Claucous-wtnged gnU
Western gull
Sande tl.tng
Western sandpiper
Llunitn
DowttchCr ap.
Kilideer
illack-beltfed plover
Ratrd'a aan4ptpar
Least sandpiper
Semipaluasted plover
Raid eagle
HarIi hawk
White-fronted goose
Canada goose
Great blue heron
I)ouble-creutsd
80.8
222.0
5.7
643.0
2.6
79.6
6.6
14.1
26.0
39.8
0.5
June
123.8
12.7
198.9
318.3
1.5
22.6
8.5
19.5
14.9
14.4
134.4
July
22.6
3,481.5
8.8
169.8
2.6
18.5
32.5
11.9
6.5
50.9
0.4
August
2.8
11.1
35.0
12.8
7.6
1.9
5.3
Septeauber
1.1
9.4
11.4
5.0
5.0
272.8
35.4
59.7
35.0
19.9
b.6
253.1
7.5
3.2
8.11
October
0.5
11:15.3
1,144.1
54.1
43.1
November
Motit Ii
5.3
4.2
3.2
23.7
140.11
2.6
15.0
10.4
4.1
3.6
20.4
20.3
103.1
0.1
0.5
Veccsber January
5.1
3.2
13.5
12.n
13.2
79.t
0.1
1.9
February
9. SpecieS density (his/1011 ha), June 19711-Hay 1919, Hitter Sand Spit Iteah, C1utsh1a Rivet, Oon.
Red-throsted loon
Species
Appendix
100.1
377.5
14.4
29.0
9.7
36.1
5.0
Hard,
82.0
100.5
620.1
26.3
210.1
5.0
14.13
03.1
209.9
1.6
0.4
April
1.4
191.0
23.e
2.6
2.2
19.9
13.0
1,021.0
75.0
691.4
1.0111.5
0.5
21.0
May
-3
Appendix
9. (conLinued)
HonLIi
Species
Number of Species
Density (birds/ha)
Diversity
Evenness
June
It
1,120.7
1.35
0.56
Juiy
13
August
13
885.4 3,921.3
0.55
1.18
0.69
0.22
Septeisber
1
83.1
1.58
0.81
October
15
930.7
1.81
0.67
November
1
2,860.4
1.02
0.52
December
1
198.9
1.02
0.53
Jauuary
9
184.7
1.42
0.64
Februsry
1k)
112.2
f.ôi
0.11
Hatch
1
b18.0
1.42
0.51
April
13
Hay
lb
2,148.4
3,loi.o
1.12
1.Sb
(J.b7
0.b9
10.
rganaer
Rufous hummingbird
belted kingfisher
Bonaparte'S gull
Caspian tern
Mew gufl
California gufl
King-billed gull
Herring gull
Spotted sandpiper
Least sandpiper
Dunlin
Dowitcher app.
Western sandpiper
Marbled godwit
Sonderling
Northern phalarope
Glaucous-winged gull
Western gull
Iitildeer
Semipalmated plover
American kestrel
Merlin
Red-breasted urganser
Bald eagle
Coawon
Ruddy duck
American wtgeon
Lesser scaup
Greater aesop
Bufflehead
Green-winged teal
Pintati
Haltatd
Wlntte-ttonted goose
Canada goose
Whistling swan
Horned grebe
Western grebe
Double-created corworaint
70.7
18.4
56.6
105.8
69.4
June
35.4
25.5
11.7
29.1
241.3
5.0
3.2
7.1
July
Species density (blrds/100 ha)
ked-throated loon
Spectea
Appendix
181.1
21.8
21.8
ÔJ.4
49.4
2.6
3.9
35.4
19.1
August
3.2
8.0
5.0
5.0
1.8
3.8
8.8
Septesber
June 1978--May
19/9k
23.7
3.5
22.0
42.1
U.3
3.9
8.8
October
1.4
3.8
8.1
316.5
22.6
7.9
5.0
Novewber
Mouth
8.1
1.4
168.4
6.7
26.0
1.0
24.3
20.7
6.5
6.4
19.9
Uecewbcr
29.5
27.3
27.4
11.0
1.6
42.1
12.7
3.9
o.4
15.7
39
4.9
January
87.0
5.8
29.3
23.9
8.8
13.0
lebruary
Millet Sands ticach, Coluahia River, 0reon.
318.3
58.7
13.5
4.7
14.9
15.7
9.9
24.9
19.9
March
3.9
No.7
14.8
23.2
37.9
2.6
129.9
n.5
7.3
22.3
7.2
730.1
l24j.1
32.5
U.)
39.8
May
9o3.8
08.2
19.9
5.0
5.0
2.6
19.9
39.8
April
Appendix
10. (continued)
Mouth
June
Specie.s
Violet-green swallow
Tree wa11ow
barn swallow
Clitf swallow
Camta crow
66.3
26.4
July
35.4
437.1
99.5
18.8
August
Septewbet
59.1
11.1
1.5
1.4
October
Noveabet
Dece4aber
2.8
January
1.4
February
9.9
M.rch
11.3
AprIL
144.7
38.4
34.8
21.1
19.8
79.6
21.0
35.4
35.4
19.6
Savannah sparrow
Song sparrow
(bttds/ha)
Btverity
Eveirnema
1.1
3.9
Amrtean goldfinch
Density
12.1
127.3
79.6
hewick's wren
Starling
Brown-headed cowbird
House finch
Number of specIes
May
7
413.6
1.83
0.94
13
11
LI
953.5
1.63
0.63
463.8
1.89
0.79
io9.1
1.67
0.10
12.8
9
154.6
1.85
0.84
1
365.3
0.59
0.30
12
14
323.1
1.ö6
0.67
261.3
2.15
0.8!
7
177.7
1.53
(1.79
10
491.1
1.34
0.58
19
2I)23.0
1.n4
0.56
In
260t.1)
1.59
0.57
II.
Water ptpit
Common crow
Cliii swallow
Casplan tern
Rufous huaidagbtrd
Horned lark
Violet-green swallow
Tree swallow
Rough-winged awllw
Barn swallow
Co,sn tern
Bonaparte's gull
Mew gull
California gull
Ring-billed gull
Herring gull
278.5
59.1
5.7
39.8
8.9
92.2
6.1
3.0
15.1
1.9
59.7
13.0
28.3
1.9
318.1)
159.2
2.2
6.5
80.6
8.5
269.7
123.0
54.6
5.0
6.5
17.1
12.1
2.4
46.3
11.0
14.1
103.9
2.1
4.4
10.4
44.4
2,217.2
23.9
168.5
5.3
12.3
247.6
5.7
114.0
9.4
26.0
1.7
IU.4
0ctbcr
6.0
0.9
13.2
50.8
0.4
Sep esber
126.3
79.0
August
Northern phalarope
Glaucous-winged gull
Western gull
12.1
141.5
July
Sander1ing
Spotted sandpiper
Baird's sandptpet
Lcaat sandpiper
Dunli
Western sandpiper
KIlideer
Black-bellied plover
Coman nrganser
Scaipalmated plover
Green-winged teaL
American wigeon
l'lntatl
White-fronted goose
Mallard
Canada goose
9.9
June
2.8
2.1
124.3
105.0
1.9
1.0
00.5
3.9
3.8
1.9
39.8
November
Month
14.1
3.6
3.3
8.1
233.6
5.3
1.5
7.1
33.2
Deceabet
7.1
28.1
35.4
17.5
115.18
1,340.8
3.2
January
1.4
141.5
1.8
3.4
15.5
20.0
25.5
February
Species density (birda/i00 ha), June 1978-N..y 1979, JIm Crow Snd5 beach, Columbia River, Oregon.
Great blue heron
Horned grebe
Common loon
Species
Appendix
).1
5.0
17.0
7.4
5.0
7.3
hi,rcli
4.2
12.)
43.2
18.4
27.4
10.2
21.0
32,506.2
b,43o.0
21.0
12.7
58.5
39.8
AFEII
7.1
191.0
t.9
)J.8
35.4
il.)
25.5
19.9
0.0
42.1
318.5
232.9
63.2
13.0
50.9
19.9
2.0
Hay
ii. (coiiiuueJ)
Nuwbar of apecieö
Denatty (birda/ha)
Diveraily
Evenueaa
Savannah aparrow
Whtte-crowned aparrow
Snow bunting
Aertcan goldfinch
Houae finch
Specea
Appendix
13
9.5
July
613.0 3,033.5
1.02
1.13
0.40
0.70
12
June
1.56
0.71
9
9511.1
2311.7
59.7
Augual
2.04
0.72
Il
8611.4
11
OcLobur
441.4
1.51
0.63
241.6
Sptewber
(1.50
937.7
1.24
12
54.7
Nuvuwbur
Mouth
328.2
1.13
0.49
10
l.4
Ueeeaber
1,554.3
0.57
0.29
7
January
0.66
1.31
334.5
8
119,4
Februat.y
14
Ajrii
19
141.5
59.7
May
41.9
39,327.1
1,341.6
1.67
0.51
2.38
0.93
0.19
(1.81
March
Appendix
12.
Species denaity (blcda/100 ha), June i918hay 1919, West Sand latand ileach, Colusbia kiver, Oregon.
Species
June
August
Steuther
October
Noveiuber
Deceabet
January
1ebruary
March
10.5
PelagIc coraorant
Great blue heron
2.6
42.6
8.6
14.9
0.1
35.4
24.2
176.8
44.2
8.11
9.9
8.8
5.0
17.1
99.5
5.0
50.9
$allard
3.9
Greater seaup
Surf scoter
Black scoter
llldeer
8.8
2.6
4.6
12.7
18.8
1.9
9.s
13.0
Black-bellied plover
11.5
kuddy turnstone
2.8
Whirubrel
19.6
22.1
9.9
0.2
141,4
65.8
10.4
6.5
9.0
6.0
13.7
93.8
15.3
21.6
37.2
26.2
146.6
2.0
5.3
319.5
503.1
899.6
64.8
419.9
2.3
3.4
259.0
446.9
50.9
70.8
Hew gull
Bonaparte's gull
Heermann's gull
Black-legged kittiwake
Barn awallow
11.1
3.2
3.9
3.8
44.2
3.2
Harsh hawk
Caspian tern
Belted kIngfisher
Violet-green swallow
3.1
8.8
7.9
White-winged acoter
king-billed gull
12.7
14.7
Coeison goldeneye
Bufflelread
Herring gall
California gull
5.0
50.9
39,11
Aruerican wigeon
Western gull
5.0
141.5
Brant
Glaucous-winged guLl
flay
3.8
aed-throeted loon
bowitcher app.
Western sandpiper
Sanderltng
Northern phalarope
iprtt
0.4
Coiswon loon
Western grabs
Doubie-eteated corworaut
July
13.2
s8.5
1.4
5.2
6.6
41.1
2.3
4.11
63.2
201.4
£19.5
415.9
509./
5.1
470.4
362.3
240.9
119.4
6.0
126.3
1.4
4.1
2.2
278.5
159.2
5.2
5.3
0.8
3.2
190.1
11.8
255
12. (continued)
Number of species
Density (birds/ha)
DIversity
Evenness
Purple martin
Cowman crow
American robin
Ruby-crowned kiuglet
Water plpit
Starting
Rouse finch
American goldfinch
Savannah sparrow
White-crowned sparrow
Song sparrow
Speclee
Appendix
13
79.6
19.9
6.5
July
9
3.8
Auguat
1,047.5
1,325.2
1,299.2
1.55
1.10
1.15
0.76
0.60
0.50
10
14.3
23.6
35.4
June
1,033.5
1.20
0.47
13
2.1
2.8
September
0.74
1.99
977.5
15
39.0
203.7
4.2
October
681.6
1.29
0.54
11
2.8
Nuvewber
Month
674.4
1.45
0.56
13
159.2
131.5
318.3
December
0.62
1.341
403.0
9
79.6
165.7
January
301.o
1.76
0.11
10
February
1/7.2
1.56
0.15
8
3.9
2.8
March
12
23.6
4.2
Hay
647.6 2,451.0
1.bZ
1.60
0.08
0.64
11
119.1
7.9
Apr11
13.
Eveuneaa
DiversIty
l)eueity (birds/ha)
Number of species
Fox sparrow
Song aparrew
kutous-ided towhee
Awericau goldfinch
Brown-headed cowbird
Black-throated gray warbler
Cheataut-sided warbler
Wjlaon's warbler
Orange-crowned warbler
Yellow warbler
Starling
Golden-crowned klnglet
Ruby-crowned kinglet
Varied thrush
Swainon'a thrush
Black-capped chickadee
Winter wren
bawick's wren
American robin
Coaaan crow
Tree swallow
Barn swallow
Western flycatcher
Downy woodpecker
0.74
1
467.1
1.44
1
676.6
1.84
0.95
119.4
1
159.2
19.9
12.1
219.4
59.7
127.3
12.1
179.1
3.9
August
135.8
59.1
99.5
35.4
July
649.5
1.71
0.91
119.4
114.6
42.4
99.5
70.)
99.5
3.9
June
1.32
0.55
1,1133.3
11
99.5
40.4
79.6
79.6
79.6
70.7
19.9
86.4
14.9
1,237.9
19.9
September
2.01
0.79
2,174.5
14
35.4
109.4
119.4
159.2
324.9
108.3
59.1
701.4
378.0
16.4
58.5
13.0
19.9
5.0
1.84
0.77
3,214.0
11
238./
70.7
318.3
1,034.5
813.5
6.5
318.3
278.5
89.1
26.0
19.9
13
0.19
1.71
0.14
1.98
0.11
/19.0
10
617./
10
136,9
1.83
15
1,544.6
2.17
0.80
278.5
79.6
170.2
59./
59.7
26.0
9.9
11.7
8.8
49.5
106.1
76.4
25.5
35.4
214.1
198.9
3.2
10.5
35.4
216.5
167.1
39.8
2.2
u.4
19.5
53.1
3.8
March
Febru,ry
99.5
106.1
35.4
6.4
218.8
59.7
2.2
35.4
3.2
January
39/.9
6.4
164.7
2/.9
22.3
99.5
159.2
79.6
2.6
35.4
31.6
3.2
Oscesaber
Month
free/Shrub. Columbia giver, Oreuzi.
October Novewber
Species density (birda/IQO ha), June l9/8-Mny 1979, Miller SaUd
Bald eagle
Merlin
Glaucous-winged gull
Western gull
Rufous hisruingbird
Coinaon flicker
Harsh hawk
Mallard
Cooper's hawk
Red-tailed hawk
SpecIes
Appendix
15
3u3.1
25.5
19.b
119.4
32.5
70.7
)ti.8
13.0
127.3
5.0
13.0
May
1,283.8
2.23
1,88
0.83
(1.78
1,021.9
11
84.0
308.4
10.0
/9.6
20.0
191.0
19.9
237.1
19.5
19.9
b.0
april
14.
21.5
59.9
19.1
216.5
179.0
19.9
42.7
25.5
318.3
79.6
127.3
139.3
218.8
39.8
50.9
139.3
19.9
106.1
39.8
159.2
141.4
99.5
119.4
1.4
39.8
106.1
70.7
July
83.7
59.7
318.3
June
99.5
39.8
19.9
59.7
12.7
12.7
59.7
232.1
25.5
3.2
8.8
5.9
5.9
79.6
August
75.3
14.6
212.2
/0.3
77.3
13.0
2.6
8.8
Septuaber
159.2
79.6
12.7
13.0
1.0
2/8.5
6.4
32.5
October
28.1
19.5
159.2
12.7
9.9
6.5
November
Month
238.1
143.2
106.1
119.4
19.9
2.6
6.5
9.5
December
391.9
119.4
31o.3
159.2
424.4
198.9
/9.6
59.7
79.6
35.4
Jdnuary
198.9
10.1
33.4
30.8
419.4
3.2
Eebruary
19.o
19.1
212.2
26.5
53.1
123.4
35.4
25.7
159.2
26.0
12.7
March
Species deneity (birds/lOU ha), June 1978-Hay 1979, West Sand Island Tree/Shrub, Cotuabia giver, Ore8ut,.
Cooper's hawk
Red-tailed hawk
Harsh hawk
Western gull
California gull
Rufous hummingbird
Common flicker
Downy woodpecker
Western flycatcher
Olive-sided flycatcher
Violet-green swallow
Tree swallow
Barn swallow
Steller's jay
Common crow
Black-capped chickadee
Winter wren
Rewick's wree
American robin
Varied thrush
Swainson's thrush
Golden-crowned kinglet
Kuby-ccowned kinglet
Cedar waxwtng
Starling
Rutton'
wire.)
Yellow-ruisped warbler
Yellow warbler
8lack-thrnted gray warbler
Wilson's warbler
Brown-headed cowbird
House finch
American goldfinch
Rufous-sided towhee
White-crowned sparrow
Golden-crowned sparrow
?ox sparrow
Song sparrow
Species
Appendix
1o4.2
10./
19.9
35.'.
79.6
57.3
35.4
4.9
10.1
44.2
26.5
05.0
305.6
2.2
50.9
391.9
19.9
ApriL
189.0
50.9
59.7
318.3
97.3
411.
31.8
139.3
8.8
8.8
58.5
0.5
195.0
20.4
1.4
1t.5
63.1
e.
79.0
May
Ln
ppendtx
14. (contInued)
Month
Spece
Number of lpecles
Deusity (btrd/ha)
Dlver8tty
Eveiinese
June
July
18
12
1,905.6 1,286.0
2.47
2.32
0.94
0.85
AuguL
14
665.0
2.06
0.18
Sepu*ber
October
8
8
49.0
587.9
1.31
0.66
1.65
0.19
Noveibec
Ueceiber January
b
8
1.25
654.7
1.54
0.74
29.5
0.70
10
1,812.4
2.04
0.89
'ebruary
MarcL
b
11
462.4
1.43
116.8
2.03
0.80
u.8S
ArII
nay
18
20
2.31
0.82
2.32
0.78
i550.0 1,862.)
15.
Golden-crowned kinglet
Ruby-crowned kinglet
Cedar wazwlng
Northetn shrike
StarlIng
Hutton's vireo
Orange-crowned warbler
Yellow warbler
Black-throated gray warbler
Swatnoa's thrush
Long-billed marsh wren
American robin
Varied thrush
HermIt thrush
Iiewick's wren
tree swallow
Barn swallow
Cliff swallow
Purple martin
Common crow
Stellar's jay
Black-capped chickadee
Winter wren
Violet-green wailow
Sharp-shinned hawk
Cooper's hawk
Red-tailed hawk
Marsh hawk
Peregrine falcon
Kilideer
Glaucous-winged gull
Western gull
California gull
Caspian tern
Short-eared owl
Rufous hummingbird
Common flIcker
Downy woodpecker
e.reen-..,Ineed teal
19.9
70.7
11.1
[.6
65.0
198.9
12.7
5.0
6.4
19.9
13.0
6.5
5.0
October
50.9
19.9
10.7
19.5
19.9
6.5
September
9.9
5.0
35.4
39.8
25.5
278.5
39.8
35.4
4.2
12.7
8.8
14.9
17.1
0.8
August
238.7
198.9
19.9
119.4
59.7
139.3
2.8
[9.6
25.5
2.8
198.9
45.5
305.9
318.3
318.3
35.4
2.6
1.4
11.8
July
2.6
June
15.5
318.3
12.1
389.0
59.7
6.5
Noveiber
Month
12.7
-
19.6
2.2
6.6
December
99.5
495.1
218.5
99.5
59.7
12.7
9.5
4.4
19.6
1.9
3.2
3.9
January
9.5
35.4
32.0
28.9
35.4
19.6
99.5
33.2
1ebrusry
25.5
19.5
17.7
1.6
154.0
14.2
159.2
o.5
5.0
Mwrch
SpecIes density (birds/IOU ha). June l918-t4y 1979, West Sand Island Transition, coluabia giver, Cun.
Common loon
Great blue heron
Species
Appendix
19.6
9.5
10.1
26.0
8.8
116.8
12.7
2.9
141.9
19.9
23s.1
3.2
1.9
April
35.4
22.3
19.9
45.
4.9
115.6
160.1
12.1
12.8
79.6
8.8
May
15. (continued)
Density (birds/ha)
Diversity
Evenness
Ruber of upecteS
Wilson's warbler
Northern oriole
Western meadowiark
brown-headed cowbird
Rouse finch
American goldfinch
Rufous-aided towhee
Savannah sparrow
Datk-eyed junco
White-crowned sparrow
Golden-crowned sparrow
ox sparrow
Song sparrow
Specie8
AppendiX
42.1
358.1
311.1
22.1
1/6.8
127.3
224.9
557.0
295.2
24.3
124.4
20
21
2,472.6
2,706.1
2.52
2.36
0.84
0.77
19.9
July
19.9
June
14
701.9
2.08
0.79
6
382.3
1.16
0.65
226.1
635.8
2.05
0.08
141
179.0
61.9
104.8
59./
17.7
19.9
October
238.7
Se1.tewber
19.8
37.6
79.6
17.7
Auguet
1.35
0.69
941.0
1
139.3
November
Month
0.18
1.87
984.9
11
19.6
103.4
39.8
35.4
37d.O
i.o
Decemauer
4,316.6
1.94
0.72
15
83.5
59.7
19.9
January
066.9
2.02
0.84
14
242.6
354
3b.4
FebLUSry
522.6
1.06
0.12
10
119.4
March
lb
139.3
19.9
321.1
id.2
Hay
2,081.6
1,690.8
1.vb
2.46
0.73
0.12
19
159.2
1s9.2
141.5
71.1
19.9
3.2
A4jttL
Appendix 16.
Mean percent covera of each habitat characteristic for
habitats comprising each island, Lower Columbia River,
19 78-79.
Percent cover
Habitat
complex
Moss!
lichen
Litter
Bare
ground
Forbs
Grass!
sedge
Miller Sands
MSM
MSSU
MSU
MSSB
MSB
MSTS
0.5
00b
43.3
00b
19.4
55C
585d
2.4
92.7
1.0
24.5
2.1
15.6
2.5
96.9
0.5
949
0.2
7.4
00b
12.1
60.8
2.2
0.0
2.5
95.3
5.5
84.5
14.8
58.2
2.8
36.8
22.7
2.3
79e
0.2
5.0
Jim Crow Sands
JCSB
JCSM
l.5f
csu
07g
01g
3.8
13.0
West Sand Island
WSIM
WSIDU
wsIu
WSIB
WSIT
WSITSI
25h
0.0
63h
0.0
21h
81h
56.2
t
346k
5.4
38.2
29.1
21.4
20.5
14.1
12.2
89.2
0.0
0.0
0.5
10.0
30.4
0.2
5.0
49.3'
90
Appendix 16.
(continued)
aData were transformed (arcsin IPT).
bpercent cover of moss/lichen was not considered for MSSU:MSS1,
NSSU:MSB, and NSSB:MSB comparison.
CMS
and MSB were similar.
dBare ground was eliminated from analysis because of high correlation
with moss/lichen (r
-0.81) and litter (r = -0.81).
epercent cover of grass/sedge for NSM was similar to MSU.
percent cover of moss/lichen for JCSM was similar to JCStJ.
cover of f orbs for JCSM was similar to JCSR.
hpercent cover of moss/lichen was similar for WSIU:WSITS and WSIM:WSIT
comparisons.
Percent cover of litter ws similar for WSIN:WSIU, WSIDU:WSIT, and
WSIT:WSITS comparisons.
3Percent cover of bare ground was similar for WSIM:WSIU comparison.
kpercent cover of grass/sedge was similar for WSfl4:WSIDIJ, WSIM:WSITS,
WSIM:WSIT, WSIDU:WSIT and WSIT:WSITS comparisons.
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