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USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 313
Marbled Murrelet Populations of Washington—Marine Habitat Preferences and Variability of Occurrence
Steven M. Speich 1 Terrence R. Wahl 2
1 Research Ecologist, Dames & Moore, Inc., 1790 E. River Road, SuiteE-300, Tucson, Arizona 85718
2 Naturalist, 3041 Eldridge, Bellingham, Washington 98225
Abstract: Marbled Murrelets occur in Puget Sound marine habitatsin relatively low numbers. The rates of occurrence of murrelets oncensuses within marine habitats is generally low. Further analysisis required to determine if low occurrence rates are a generalfunction of the movement of birds, or their consistent occurrenceon particular censuses and not on others. Qualitative data indicatethat Marbled Murrelet abundance in Puget Sound is now lower thanearlier this century. Such long-term information is unavailable forMarbled Murrelets along the Pacific Ocean coast of Washington.Census data from nearshore waters of the Pacific Ocean off GraysHarbor indicate that Marbled Murrelet abundance is reduced theresince 1989 and especially in 1993. This pattern is also reflected inseveral other more oceanic species suggesting basic and wide-spread changes in marine carrying capacity.
Early reports on the birds of Washington (Dawson andBowles 1909, Jewett and others 1953) consisted primarily ofinteresting general accounts, whereas several recently publishedreports have focused specifically on marine birds in Washington(Briggs and others 1992; Cody 1973; Grover and Olla 1983;Manuwal and Campbell 1979; Manuwal and others 1979;Paine and others 1990; Speich and Wahl 1989; Wahl 1984;Wahl and Speich 1984; Wahl and Speich, in press; Wahl andothers 1981; Wilson 1991). Despite these reported activitiesof researchers, including ourselves, little has been written ofthe habitat preferences of marine birds, including MarbledMurrelets, in Washington. Lately, a few general descriptionsand quantifications of the abundance of marine birds basedon marine habitats have appeared. While only the reports ofWahl and others (1981) and Long (1983) are pertinent toPuget Sound, and of Wahl (1984), Speich and others (1987),and Briggs and others (1992) to the Pacific Ocean coast,they only marginally included the Marbled Murrelet. Thesame paucity of information pertains to the foods of marinebirds in Washington marine areas, with the exception of afew species specific studies (e.g., Rhinoceros Auklet[Cerorhinca monocerata] [Wilson and Manuwal 1986]). Littlehas been written that specifically deals with Marbled Murreletsin Washington marine waters (Speich and others 1992;Thoresen 1989; Varoujean and Williams, this volume).
Marbled Murrelets are found throughout the Puget Soundregion, although their distribution varies spatially andtemporally (Speich and others 1992; Wahl and others 1981;Wahl and Speich 1983, 1984). Speich and others (1992)
attempted to establish the size of populations in each of themajor marine areas of Washington, to determine if a seasonalchange in numbers occurred, and to evaluate historical marineevidence for changes in the numbers of breeding birds. Theresults were a breeding population estimated at near 5,000Marbled Murrelets, evidence of an influx of birds into atleast Puget Sound during the winter, and an indirect conclusionthat the breeding population in Puget Sound had declinedfrom early periods, although the magnitude of the change isunknown and cannot be quantified. Our impression is thatmurrelets are variable in their occurrence, moving from onearea to another, often in short time periods, although birdsare often found in specific areas.
In this paper we quantify and discuss the seasonalgeographic and marine habitat distribution, abundance andvariability of Marbled Murrelets in Washington marine waters.Changes in abundance over the past 23 years of censusingcontinental shelf waters near Grays Harbor are explored.
MethodsThe inland marine areas of Washington, better known as
Puget Sound, are a complex of bays and passages, supportinga large variety of marine habitats and associated organisms(Long 1983, Simenstad and others 1979). They are connectedby larger deep water areas such as the Strait of Juan de Fuca,Admiralty Inlet, Haro Strait, Rosario Strait and Georgia Strait,all distinct habitats. During the 1978 and 1979 NationalOceanic and Atmospheric Administration (NOAA) MarineEcosystem Analysis Program (MESA), marine bird surveysof northern Puget Sound (Wahl and others 1981) censuseswere established and conducted. These quantified distributionand abundance of the marine birds found in all the majormarine habitat types and geographic areas of northern PugetSound. During the MESA program, northern Puget Soundwas divided into 11 major regions and 72 subregions, largelybased upon marine and terrestrial geography and water depth.Each subregion, and certainly each region, contained one ormore distinct marine habitat types, based on water depth,marine substrate type, slope and, in part, geography. Eachregion, subregion and habitat type was overlain with one ormore distinct marine census transects. These transects werefixed in location and were censused by one or morestandardized census methods. Census methods included smallaircraft, small boats, Washington State Ferries, and fixedlocations at points, about bays, and along beaches. Censusdata were lumped by location, marine habitat, subregion andregion, and time period, as appropriate. In the habitat analysisfor this paper, censuses were combined by habitat type and
Chapter 30
314 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
time period. Densities were calculated by dividing the totalnumbers of Marbled Murrelets observed by the total areacensused on all censuses within each habitat types for thespecific time period. Data from 1978 and 1979 were combined.
Surveys similar to the MESA bird program, partiallyfunded by NOAA and the Washington Department of Wildlife,were later established for southern Puget Sound (Wahl andSpeich 1983, 1984). These surveys were designed to allowthe quantification and appraisal of the general, seasonaldistribution and abundance of all marine waterbird speciesin southern Puget Sound. This program consisted of onecomplete summer survey of the area and extensive wintersurveys (as yet unpublished) over several years. All wintersurveys were from light aircraft and summer surveys werefrom small boats and light aircraft following the methods setforth in Wahl and others (1981, see above). Results areexpressed as birds per square kilometer and are summarizedby geographic area, inshore or offshore, and time period.
There are only limited data from the Pacific Ocean coastof Washington that quantify the occurrence of MarbledMurrelets in marine habitats. The best data are for the southernouter coast, the coast south of Point Grenville, including theGrays Harbor Channel and habitats in the shelf waters off themouth of Grays Harbor channel to the continental shelf break(Wahl 1984), and the onshore area in the vicinity of PointGrenville (Speich and others 1987, 1992). Along the northportion of the coast, the area north of Point Grenville, onlylimited data are available for the nearshore and offshorewaters of the continental shelf (Speich and others 1992).Censuses of birds over the continental shelf, to the shelfbreak, off of Grays Harbor, were made from chartered fishingboats, from 1971 to the present. Census frequency, especiallyduring the winter storm period, was often limited for accessto the ocean due to rough bar and sea conditions. All birdsobserved were counted and summarized for transects of varyinglength within specific water depth intervals, and results areexpressed as birds observed per linear kilometer, as describedin specific detail by Wahl (1984). Observations of birds ofspecified nearshore water areas from three land locationsnear Point Grenville were accomplished with the aid ofbinoculars and telescopes during the spring-summer periodsof 1984 and 1985 (Speich and others 1987). Observationswere expressed as Marbled Murrelets observed per squarekilometer for consecutive week periods. North of the PointGrenville study area, observations were made from boats(Zodiacs) while moving up and down the coast over nearshorewaters of the continental shelf (Speich and others 1992). Allbirds observed in moving zones about the vessel were recorded,with results expressed as birds per square kilometer.
ResultsAbundance and Occurrence by Habitat Types -Puget Sound
For northern Puget Sound, the seasonal densities andpercent occurrence of Marbled Murrelets were determined
for censuses within five broad habitat groups, each in turnsubdivided into several more specific habitat types (table 1,figs. 1-10).
Open Water Greater than 20 m DepthIn Sequim and Discovery bays, the large sheltered bays
at the eastern end of the Strait of Juan de Fuca, MarbledMurrelets reach peak abundance during the fall period (table1, fig. 1). No other habitat within this habitat group had ashigh a density, 2.5 birds/km2. The maximum density obtainedduring the winter period, 0.92 birds/km2, was also fromSequim and Discovery bays. Concentrations of MarbledMurrelets were also reported from this area on AudubonChristmas Counts, according to summary statements by Speichand others (1992).
Within this habitat group, the proportion of individualcensuses with Marbled Murrelets was generally near, andoften less than, 20 percent (0.2) (table 1, fig. 2). The exceptionwas the summer period for Sequim and Discovery bayswhere Marbled Murrelets were observed on 50 percent of allcensuses in the area, but the sample size (n = 2) is verysmall. Within this habitat group, the deep open waters withinthe San Juan Islands showed peak numbers (density) andoccurrence rate during the summer and fall periods (table 1,figs. 1 and 2).
Bays with Steep and Gradual SlopesHabitat types within the group, “bays—steep and gradual
slopes”, are described by location and type (table 1, figs. 3and 4). These habitat types generally are characterized bylow densities of Marbled Murrelets. High densities of fourand five murrelets per square kilometer, were found in habitatson steep slope and sand substrate within Whatcom and Skagitcounties (Chuckanut Bay) and within the San Juan Islands,during the winter period.
Although the densities of Marbled Murrelets in the habitattypes of this group are low, the rate of occurrence of censuseswith birds is nonetheless relative high. While the density ofbirds was relatively high during the winter in steep slopehabitats with sand substrate within the San Juan Islands,birds were only observed on about half of censuses (table 1,fig. 4). This suggests that relatively large numbers of birdswere present during the winter in this habitat type, but birdsmove about and were not always encountered on censuses.A similar pattern was observed in this habitat type in Whatcomand Skagit Counties, except in winter when birds were detectedon a greater portion of censuses, about 80 percent. Similarly,during the spring period, birds were detected on about 90percent of censuses, even though average density (1.42 birds/km2) (table 1) was lower than the winter period density(3.92 birds/km2).
Areas of Tidal ActivityThe occurrence of Marbled Murrelets in areas of tidal
mixing is not unexpected, as these are generally thought of asproductive areas where prey concentrate in nutrient and food-
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 315
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
con
tinu
es
Tab
le 1
—S
easo
nal d
ensi
ty a
nd o
ccur
renc
e of
Mar
bled
Mur
rele
ts in
nor
ther
n P
uget
Sou
nd m
arin
e ha
bita
t typ
es
Spr
ing
Sum
mer
Fal
lW
inte
r__
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
_
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
____
____
____
____
____
____
___
____
____
____
____
____
____
____
___
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
___
Bird
s/B
irds/
Bird
s/B
irds/
Hab
itat t
ypes
- L
ocat
ions
km2s.
d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
Ope
n W
ater
>20
m D
epth
Seq
uim
and
Dis
cove
ry B
ays
0.00
(0.0
0)17
00.
000.
33(0
.33)
21
0.50
2.54
(7.5
0)13
30.
230.
92(2
.37)
183
0.17
Can
adia
n G
ulf I
slan
ds0.
02(0
.08)
533
0.06
0.43
(0.7
9)19
50.
260.
24(0
.87)
7915
0.19
0.10
(0.3
0)94
110.
12
Str
ait o
f Jua
n de
Fuc
a0.
00(0
.00)
160
0.00
0.01
(0.0
1)9
10.
110.
06(0
.23)
284
0.14
0.02
(0.0
7)36
20.
06
Har
o S
trai
t0.
00(0
.00)
140
0.00
0.00
(0.0
0)6
00.
000.
00(0
.00)
230
0.00
0.02
(0.1
0)32
10.
03
Ros
ario
Str
ait
0.21
(0.8
2)48
70.
150.
01(0
.03)
131
0.08
0.41
(1.2
2)64
160.
250.
16(0
.65)
766
0.08
Geo
rgia
Str
ait
0.37
(0.9
1)44
80.
180.
00(0
.00)
90
0.00
0.16
(0.7
0)41
60.
150.
11(0
.50)
472
0.04
Wha
tcom
& S
kagi
t Cou
ntie
s0.
12(0
.44)
272
0.07
0.00
(0.0
0)5
00.
000.
32(1
.06)
263
0.12
0.20
(0.5
8)33
40.
12
San
Jua
n Is
land
s0.
17(0
.66)
100
90.
090.
32(0
.79)
307
0.23
0.80
(3.1
5)16
639
0.23
0.07
(0.3
2)19
111
0.04
Bay
s–S
teep
and
Gra
dual
Slo
pes
Ste
ep-S
and/
Str
ait o
f Jua
n de
Fuc
a0.
01(0
.02)
111
0.09
0.28
(0.2
8)2
10.
500.
00(0
.00)
90
0.00
0.01
(0.0
2)12
20.
06
Ste
ep-S
and/
Wha
tcom
& S
kagi
t Cou
ntie
s1.
43(1
.22)
87
0.88
0.11
(0.1
9)4
10.
250.
22(0
.46)
186
0.33
3.92
(3.1
7)15
120.
80
Ste
ep-S
and/
San
Jua
n Is
land
s0.
22(0
.82)
222
0.09
0.11
(0.2
8)12
20.
171.
15(3
.16)
265
0.19
5.08
(10.
37)
4826
0.54
Gra
dual
-San
d-E
elgr
ass/
San
Jua
n Is
land
s0.
01(0
.07)
231
0.04
0.00
(0.0
0)1
00.
000.
04(0
.12)
243
0.13
0.17
(0.4
5)34
100.
29
Gra
dual
-San
d-E
elgr
ass/
Wha
tcom
& S
kagi
t Cou
ntie
s0.
13(0
.27)
4315
0.35
0.31
(0.7
3)17
80.
470.
06(0
.24)
7216
0.22
0.06
(0.2
0)82
170.
21
Gra
dual
-San
d-M
udW
hato
cm &
Ska
git C
ount
ies
0.14
(0.2
7)13
50.
380.
40(0
.03)
22
1.00
0.01
(0.0
3)16
20.
130.
67(1
.57)
188
0.44
Gra
dual
-San
d-M
ud/
San
Jua
n Is
land
s0.
75(1
.40)
62
0.33
0.00
(0.0
0)4
00.
000.
30(0
.84)
91
0.11
0.00
(0.0
0)13
00.
00
Gra
dual
-Roc
k-C
obbl
e/as
sort
ed s
ites
0.34
(1.0
5)27
60.
221.
25(3
.54)
91
0.11
0.11
(0.4
3)46
60.
130.
26(0
.56)
5120
0.39
Are
as o
f Tid
al A
ctiv
ity
Var
ious
poi
nts
0.92
(1.4
8)29
110.
385.
62(1
0.10
)8
60.
751.
51(4
.35)
6131
0.51
0.37
(1.0
8)28
40.
14
Pas
sage
s/S
an J
uan
Isla
nds
0.13
(0.3
5)62
90.
150.
23(0
.53)
235
0.22
5.05
(32.
62)
8332
0.39
1.69
(2.9
7)11
260
0.54
Adm
iralty
Inle
t0.
03(0
.07)
91
0.11
0.12
(0.1
7)3
10.
330.
03(0
.11)
181
0.06
1.12
(2.1
9)19
100.
53
316 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
Tab
le 1
—co
ntin
ued
Spr
ing
Sum
mer
Fal
lW
inte
r__
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
_
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
Den
sity
Occ
urre
nce
____
____
____
____
____
____
___
____
____
____
____
____
____
____
___
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
___
Bird
s/B
irds/
Bird
s/B
irds/
Hab
itat t
ypes
- L
ocat
ions
km2s.
d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
km2
s.d.
No.
Pre
s.P
Sho
relin
es w
ith N
arro
w S
helf
Kel
p-C
obbl
e/S
trai
t of J
uan
de F
uca
1.00
(2.4
9)21
40.
194.
00(4
.24)
42
0.50
0.14
(0.6
8)23
10.
040.
00(0
.00)
240
0.00
Kel
p-C
obbl
e/W
hatc
om &
Ska
git C
ount
ies
0.54
(1.8
3)49
90.
181.
05(2
.13)
166
0.38
1.01
(1.8
6)65
280.
430.
10(0
.30)
547
0.13
Kel
p-C
obbl
e/W
hatc
om C
ount
y is
land
s0.
68(1
.60)
275
0.19
0.71
(1.7
5)7
10.
1419
.98
(41.
77)
3218
0.56
3.07
(8.2
6)34
130.
38
Kel
p-C
obbl
e/S
an J
uan
Isla
nds
0.04
(0.2
5)45
10.
020.
06(0
.23)
151
0.07
4.90
(17.
53)
6617
0.26
3.17
(8.3
6)64
180.
28
Cob
ble-
Roc
k/S
trai
t of J
uan
de F
uca 0
.68
(1.6
2)17
40.
240.
11(0
.23)
51
0.20
0.45
(1.2
3)30
80.
270.
13(0
.29)
325
0.16
Cob
ble-
Roc
k/S
an J
uan
Isla
nds
0.18
(0.7
3)25
20.
080.
20(0
.57)
91
0.11
0.28
(0.9
9)33
30.
090.
48(1
.09)
4810
0.21
San
d-M
ud/a
ssor
ted
site
s0.
29(0
.00)
11
1.00
0.04
(0.0
4)2
10.
500.
00(0
.00)
20
0.00
0.09
(0.1
6)4
10.
25
Sho
relin
e w
ith B
road
She
lf
Eel
gras
s-S
and/
Str
ait o
fJu
an d
e F
uca
0.00
(0.0
0)14
00.
000.
00(0
.00)
20
0.00
0.23
(0.7
4)20
20.
100.
00(0
.00)
270
0.00
Eel
gras
s-S
and/
Wha
tcom
& S
kagi
t Cou
ntie
s0.
66(1
.83)
123
0.25
4.44
(6.2
9)3
10.
330.
57(1
.40)
71
0.14
0.00
(0.0
0)8
00.
00
Kel
p-C
obbl
e/S
trai
t of
Juan
de
Fuc
a2.
03(5
.62)
4917
0.35
2.13
(6.0
4)15
50.
330.
01(0
.01)
811
0.01
0.43
(1.9
7)87
150.
17
Kel
p-C
obbl
e/S
an J
uan
Isla
nds
0.00
(0.0
0)4
00.
000.
28(0
.39)
31
0.33
0.12
(0.3
7)10
10.
100.
26(0
.64)
71
0.14
Cob
ble-
Roc
k/as
sort
ed s
ites
0.00
(0.0
0)9
00.
000.
00(0
.00)
20
0.00
0.60
(1.3
6)12
20.
171.
14(2
.16)
122
0.17
San
d-M
ud/S
trai
t of J
uan
de F
uca
3.33
(5.7
7)4
10.
250.
00(0
.00)
10
0.00
0.00
(0.0
0)5
00.
000.
00(0
.00)
50
0.00
San
d-M
ud/
Wha
tcom
& S
kagi
t Cou
ntie
s1.
08(4
.15)
192
0.11
0.24
(0.5
3)6
10.
170.
67(1
.78)
254
0.16
0.65
(1.8
0)23
30.
13
San
d-M
ud/S
an J
uan
Isla
nds
0.39
(0.5
5)3
10.
332.
35(0
.00)
11
1.00
4.02
(7.9
5)6
30.
501.
67(3
.47)
62
0.33
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
__
Lege
nd:
Den
sity
- s.d
. = s
tand
ard
devi
atio
n; O
ccur
renc
e -
No.
= n
umbe
r ce
nsus
es p
erfo
rmed
, Pre
s. =
num
ber
cens
uses
bird
s pr
esen
t; P
= p
ropo
rtio
n c
ensu
ses
bird
s pr
esen
t
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 317
rich upwelled or mixed waters. Within Puget Sound, suchareas are normally associated with narrow passages or pointswhere currents, and mixing, are intensified. The habitat typespresented here reflect those conditions (table 1, figs. 5 and 6).
Highest densities were calculated from observationsduring the summer period from “Various Points” (figs. 5 and6) where Marbled Murrelets were recorded on over 70 percentof all censuses. Densities of birds at these points were lowerduring the spring and fall, and the percent occurrence oncensuses was lower, but on average, Marbled Murrelets wereobserved on about half of all censuses during this period.Only during winter did both the density and occurrence ratedrop below values from other periods.
In Passages—San Juan Islands, the highest seasonaldensity was obtained for the fall period, 5.05 birds/km2 (table1, fig. 5), when birds were observed on about 40 percent ofall censuses (table 1, fig. 6). There, numbers observed andpercent occurrence of censuses were much lower during thespring and summer periods. During the winter period, thedensity (1.69 birds/km2) was considerably lower, while birdswere recorded on over 50 percent of all censuses.
Murrelets occurred at much lower rates on censuses inAdmiralty Inlet, (table 1, fig. 5), compared to other tidalareas. However, more birds were likely present as the areaof this tidal passage is larger than the other tidal habitatareas discussed. Similar to the other areas of tidal activity
Figure 1 —Seasonal abundance of Marbled Murrelets, open water greater than 20 m depth.
Figure 2 —Seasonal presence of Marbled Murrelets, open water greater than 20 m depth.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
318 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
in Puget Sound, occurrence on censuses is quite variable(table 1, fig. 6).
Shorelines with Narrow ShelfThis group is represented by three specific habitat types,
Kelp and Cobble, Cobble and Rock, and Sand and Mud.Within this general habitat group, there is considerable variationin densities (table 1, fig. 7) and in the proportion of censuseson which they are recorded (table 1, fig. 8). The highestdensity (19.98 birds/km2) determined for any habitat in PugetSound occurred in the fall in the Kelp and Cobble substrate inthe Whatcom County islands area, where birds were also
recorded on half of all surveys. A relatively high density(5.05 birds/km2) was also determined for the fall period forKelp and Cobble substrate in the San Juan Islands. Otherwise,with a couple of additional exceptions, densities of murreletsin this habitat group were generally low (table 1, fig. 7).
Shorelines with Broad ShelfThis group is represented by four habitat types, Eelgrass
and Sand, Kelp and Cobble, Cobble and Rock, and Sand andMud. Here again the variation in densities of MarbledMurrelets between and within habitat types is apparent (table1, fig. 9), although sample sizes in some cases are small
Figure 3 —Seasonal abundance of Marbled Murrelets, bays - steep and gradual slopes.
Figure 4 —Seasonal presence of Marbled Murrelets, bays - steep and gradual slopes.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 319
Abundance and Occurrence by Habitat Types -Pacific Coast
Grays Harbor Channel and Shelf WatersOver a 23-year period, Marbled Murrelets were recorded
in Grays Harbor channel in every month. The general patternof occurrence was one of high average densities during thespring, fall, and winter months (table 2, fig. 11), and higherdensities in habitats closer to shore. Overall, the highestdensities occurred in Grays Harbor Channel, followed byGrays Harbor Channel to 20 m depth, and 20 to 50 m depth.Only rarely were they recorded in deeper habitat areas. Thehighest densities occurred during the spring months and highestaverage density occurred in Grays Harbor Channel in March.
(table 1). The highest calculated densities occurred in Eelgrassand Sand substrate in Whatcom and Skagit Counties duringthe summer, Cobble and Rock substrate of “Assorted Areas”during the winter, and in the Sand and Mud substrate in theSan Juan Islands during the fall. The last habitat type hadoverall the highest determined seasonal densities.
Overall within this habitat group, the proportion ofcensuses with murrelets was below 40 percent (table 1, fig.10). The only exceptions were the spring and summer periodsof the Sand and Mud substrate in the San Juan Islands,however small sample sizes (table 1) suggest caution ininterpreting those values. As elsewhere, there is considerableseasonal within and between habitat variation.
Figure 5 —Seasonal abundance of Marbled Murrelets, areas of tidal activity.
Figure 6 —Seasonal presence of Marbled Murrelets, areas of tidal activity.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
320 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
The percent of censuses that Marbled Murrelets wererecorded in each of the habitat types, by month, followed thesame pattern as shown by densities according to Wahl’sunpublished observations. Overall, though, sample size issmall, with highest occurrence recorded during the winter,spring, and summer periods. Interestingly, over a period of23 years, murrelets were recorded on every census in GraysHarbor Channel in February, March, November, andDecember. They were also recorded in the habitat area fromGrays Harbor Channel to 20 m depth in February, March,and November. Even though the proportion of censuses onwhich murrelets were recorded was often high, densitieswere often low (e.g., February).
Point Grenville Inshore WatersDuring the spring and summer periods of 1984 and 1985
Marbled Murrelets were irregularly observed in the inshorewaters near Point Grenville (Speich and others 1987). Thesedata, previously reported in Speich and others (1992), showedthat they were absent from about 25 to 30 percent of allcensuses, mainly in April. In both years peak densities wererecorded in July, and in August 1985.
Continental Shelf WatersEarlier, Speich and others (1992) suggested that the
Marbled Murrelet population on the Pacific Ocean Coast ofWashington was largely found north of Point Grenville with
Figure 8 —Seasonal presence of Marbled Murrelets, shorelines with narrow shelf.
Figure 7 —Seasonal abundance of Marbled Murrelets, shorelines with narrow shelf.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 321
an uncertain number found in the waters off the southerncoast. However, the numbers there were thought to be low.This pattern was confirmed by the August 1993 aerial surveyof the coast (Varoujean and Williams, this volume).
Regional Distribution and Variability - Puget Sound
During the 1978 and 1979 survey of northern PugetSound (Wahl and others 1981), Marbled Murrelets werefound differentially distributed temporally and spatiallythrough the study area. Overall, the results of surveys duringthe spring-summer and winter periods showed on averagethat the obtained densities and the proportion of surveyswith murrelets were higher during the winter, compared
with the spring-summer period (see appendices 1 and 2, andfig. 3 in Speich and others [1992]). Indeed, the calculatedtotal for northern Puget Sound was higher in winter than thespring-summer period.
Not only were changes at the region level apparent, butchanges within each region, at the subregion level were alsofound (see appendices in Speich and others [1992]). Forexample, along the Strait of Juan de Fuca, at the north end ofthe Olympic Peninsula, the densities (and calculated totals)of murrelets decreased during the winter period in seven ofthe 20 subregions (35 percent), remained the same in nine ofthe 20 (45 percent) and increased in three (15 percent).There were no increases of densities in subregions west of
Figure 9 —Seasonal abundance of Marbled Murrelets, shorelines with broad shelf.
Figure 10 —Seasonal presence of Marbled Murrelets, shorelines with broad shelf.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
322 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
Fig
ure
11—
Mon
thly
abu
ndan
ce o
f Mar
bled
Mur
rele
ts in
Gra
ys H
arbo
r ha
bita
ts, 1
971
thro
ugh
1993
. See
tabl
e 2
for
sam
ple
para
met
ers.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in WashingtonT
able
2—
Mon
thly
rat
es o
f occ
urre
nce
of M
arbl
ed M
urre
lets
in n
ears
hore
hab
itats
off
Gra
ys H
arbo
r, W
ashi
ngto
n, r
ecor
ded
and
aver
age
d ov
er a
23-
year
per
iod
Mon
thG
rays
Har
bor
Cha
nnel
Gra
ys H
arbo
r C
hann
el to
20
m20
to 5
0 m
dep
th50
to 1
00 m
dep
th10
0 to
200
m d
epth
200
to 1
000
m d
epth
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
____
__
No.
No.
Bird
s/N
o.N
o.B
irds/
No.
No.
Bird
s/N
o.N
o.B
irds/
No.
No.
Bird
s/N
o.N
o.B
irds/
bird
skm
kmbi
rds
kmkm
bird
skm
kmbi
rds
kmkm
bird
skm
kmbi
rds
kmkm
Janu
ary
2221
.01.
0518
33.5
0.54
069
.30
012
9.7
00
201.
60
018
9.2
0
Feb
ruar
y1
6.0
0.17
410
.00.
402
10.5
0.19
038
.00
050
.50
023
.00
Mar
ch42
15.2
2.76
44.
40.
912
31.7
0.06
025
.60
646
.50.
130
22.5
0
Apr
il99
77.2
1.28
9166
.31.
370
120.
60
228
7.0
0.01
031
3.1
02
198.
60.
01
May
173
334.
10.
5212
128
9.4
0.42
1844
7.4
0.04
2079
8.6
0.03
212
49.2
00
696.
10
June
2847
.20.
5930
54.8
0.55
362
.20.
050
97.3
00
156.
50
083
.70
July
8130
5.2
0.27
3137
8.2
0.08
1050
2.4
0.02
079
2.2
00
1442
.00
073
5.4
0
Aug
ust
4260
8.9
0.07
1356
4.9
0.02
084
5.8
00
1659
.90
024
89.9
00
1472
.20
Sep
tem
ber
1551
3.8
0.03
552
4.2
0.01
288
0.0
00
1715
.90
029
21.0
00
1361
.40
Oct
ober
6523
8.6
0.27
5024
2.0
0.21
442
0.5
0.01
085
1.0
00
1200
.10
065
8.7
0
Nov
embe
r29
16.3
1.78
410
.50.
384
28.0
0.14
249
.90.
040
80.2
00
13.1
0
Dec
embe
r2
6.5
0.31
1513
.71.
090
17.6
00
25.6
00
45.8
00
15.1
0__
____
____
____
____
____
____
____
___
____
____
____
____
____
____
____
____
____
___
____
____
____
____
___
____
____
____
___
____
____
____
____
__
Tot
al59
921
9038
621
91.9
4534
36.0
2464
70.7
810
196.
42
5469
.0
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 323
Dungeness Spit: six showed decreases and six showed nochange. East of Dungeness Spit, four subregions showedincreases, one showed a decrease and four showed no change.The Dungeness Spit subregion showed no change in densityor projected total of birds.
A different pattern was observed when spring-summerdensities and calculated totals were compared with those ofwinter in more northern and eastern areas of northern PugetSound. Within the eastern bays and passages of Whatcomand Skagit counties, the area to the east of Rosario Strait,including Bellingham, Padilla, Samish, Fidalgo and Lummibays, murrelets were more plentiful during the winter periodin five of the seven (71 percent) subregions and there was nochange in the other two subregions (29 percent).
This pattern was also observed in the bays and passagesof the San Juan Islands. There, Marbled Murrelet densitiesand projected totals were greater in 11 (52 percent) of the 21subregions of this geographic area. Values decreased in onlyfive subregions (24 percent) and remained the same in fiveothers (24 percent).
Long-Term Trends in Abundance and Occurrence -Puget Sound
As reported earlier (Speich and others 1992), there areno data that will allow for the quantification of long termchanges in the abundance of Marbled Murrelets in Washingtonmarine waters. Qualitative statements in early accounts suggestthat the species was once more abundant than present inPuget Sound waters.
Long-Term Trends in Abundance and Occurrence -Pacific Ocean Coast
There are no data that we are aware of that allow a directappraisal of the long term stability, or instability, of populationsalong the Pacific Ocean coast of Washington, over the lastcentury, or even over the last several decades. The onlyquantified information available is the data set collected byWahl from offshore birding trips from Grays Harbor to overthe continental shelf break, covering the years 1971 through1993 (table 3, fig. 12). Therein, no long term trend in theabundance of Marbled Murrelets is evident, except that since1989, yearly abundance is visibly diminished. The lowestlevel of murrelet abundance over the entire study periodoccurred in 1993.
We are unable to directly predict if the pattern ofabundance observed in nearshore waters near Grays Harborare indicative of abundance patterns along the entire outercoast of Washington. However, some insight is obtainedby considering the yearly patterns of relative abundance ofother marine bird species occurring in nearshore andcontinental shelf waters off of Grays Harbor. Several marinespecies, consisting of four families of birds, and comprisinglocal breeders, migrants, and winter visitors, were lessabundant in recent years in the Grays Harbor study area.Wahl’s unpublished observations are summarized below.The lowest abundances since 1971 were recorded during
the past two years for eight species (Sooty Shearwater[Puffinus griseus], Red-necked Phalarope [Phalaropuslobatus], Sabine’s Gull [Xema sabini], Arctic Tern [Sternaparadisaea], Common Murre [Uria aalge], Cassin’s Auklet[Ptychoramphus aleuticus], Rhinoceros Auklet, TuftedPuffin [Fratercula cirrhata]),and during one of the lasttwo years for two species (Fork-tailed Storm-Petrel[Oceanodroma furcata], Red Phalarope [Phalaropusfulicaria]). Of interest, Common Murre abundance decreasednoticeably in 1989, and has remained low, after severalyears (1980-1988) of relative high abundance. Theabundance pattern of the last species corresponds to thatobserved for the Marbled Murrelet, relative low abundancesince 1989 (fig. 12).
DiscussionThe overall pattern of abundance (density) and occurrence
of Marbled Murrelets observed in the marine habitats ofPuget Sound is one of variability. Our impression of MarbledMurrelets in Puget Sound before this limited analysis was ofa species that moves about a great deal on several temporalscales: seasons, daily, and hourly. Indeed, we have oftenobserved Marbled Murrelets foraging in a particular areathen departing that area in a short period of time, and flyingout of our sight to another unknown location. Such movementsmay account for the generally low probability of encounteringmurrelets on censuses within habitat types. Thus, the observedoccurrence patterns are not surprising. However, there aresources of noise in the original census program that couldgive a partially false impression of this species’ patterns ofoccurrence. Specifically, not discounting our generalimpression of variability, we have noticed that they are oftenfound in specific areas, while other areas are less likely tocontain them. Thus, since any specific habitat type presentedis usually represented by several spatially distinct censusesand the mixing of censuses, some often with and withoutdetecting murrelets, leads to lowered occurrence rates. Wepredict that this will be the case, and that the habitat spatialscale we presented here for Puget Sound will turn out to betoo coarse.
Our field observations of Marbled Murrelets in PugetSound, during the course of formal censuses and otherwise,suggest that the foraging distribution is closely linked totidal patterns, in particular to specific locations when tidalflows are clearly evident. The locations of tidal activitycovered by data that are presented here (figs. 5 and 6) arewell known and particularly prominent places of tidal activity.However, tidal activity is observable throughout Puget Soundand is likely the single dominant and persistent physicalprocess there. We suggest that analysis of our data at thelevel of individual censuses may give insight into the relativeimportance of tidal activity in determining the movementsand foraging areas of Marbled Murrelets.
As recognized previously, there are seasonal regionalpatterns in the distribution and abundance of murrelets in
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
324 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
Table 3—Yearly rates of occurrence of Marbled Murrelets in nearshore habitats off Grays Harbor,Washington
Year Grays Harbor Channel Grays Harbor Channel to 20 m 20 to 50 m depth______________________________________ ___________________________________________________ __________________________________________
No. No. Birds/ No. No. Birds/ No. No. Birds/birds km km birds km km birds km km
1971 6 27.7 0.22 0 20.3 0 0 53.4 0
1972 51 98.6 0.52 0 66.3 0 0 108.0 0
1973 15 109.4 0.14 0 39.6 0 0 93.8 0
1974 7 40.0 0.18 0 51.2 0 0 88.7 0
1975 16 69.8 0.23 9 56.0 0.16 0 81.1 0
1976 11 36.8 0.30 7 88.6 0.08 0 117.1 0
1977 35 76.9 0.46 15 148.9 0.10 1 206.9 0
1978 10 90.2 0.11 10 81.8 0.12 0 150.7 0
1979 26 88.4 0.29 0 54.7 0 0 155.9 0
1980 7 75.6 0.09 51 77.7 0.66 0 100.1 0
1981 24 60.8 0.39 20 125.1 0.16 18 168.8 0.11
1982 88 111.1 0.79 16 128.2 0.12 2 202.5 0.01
1983 23 101.2 0.23 16 75.4 0.21 4 158.8 0.03
1984 24 128.4 0.19 7 100.8 0.07 11 213.4 0.05
1985 74 137.9 0.54 102 140.5 0.73 0 227.2 0
1986 24 81.8 0.29 7 73.5 0.10 0 136.1 0
1987 27 129.3 0.21 16 115.2 0.14 3 191.0 0.02
1988 69 112.6 0.61 50 104.6 0.48 2 179.5 0.01
1989 8 104.0 0.08 2 86.0 0.02 0 120.0 0
1990 11 111.5 0.10 13 140.0 0.09 2 173.0 0.01
1991 25 131.5 0.19 21 148.5 0.14 0 172.0 0
1992 13 141.5 0.09 22 126.5 0.17 2 165.5 0.01
1993 5 125.0 0.04 2 142.5 0.01 0 172.5 0
______ ____________ ______ ___________ ______ __________
Total 599 2190 386 2191.9 45 3436
Figure 12 —Yearly abundance of Marbled Murrelets in Grays Harbor habitats, 1971 through 1993. See table 3 for sample parameters.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
USDA Forest Service Gen. Tech. Rep. PSW-152. 1995. 325
Puget Sound. Of particular note, are the nearshore subregionsalong western portion of the Strait of Juan de Fuca wherethey are found less often during the winter period. This area,particular in Puget Sound, is exposed to winter storm activity,as it opens directly to the Pacific Ocean, while those Strait ofJuan de Fuca subregions to the east of Dungeness Spit arerelatively protected from western approaching winter storms.The densities in these subregions also suggest a shift of birdsfrom the Strait of Juan de Fuca during the spring and summerperiods, to areas in the San Juan Islands and the eastern baysduring the fall and winter periods. In addition, as demonstratedin Speich and others (1992), there is apparently an additionalinflux of Marbled Murrelets into the latter areas from thenorth, presumably British Columbia. A shift in MarbledMurrelets from the Strait of Juan de Fuca alone cannotaccount for the increases in numbers in these areas. Similarpatterns, though on a lesser scale, are likely present in otherareas of Puget Sound.
An estimate of the size of the Marbled Murrelet populationfound in the marine waters of Washington during the spring-summer breeding period has been made by Speich and others(1992), calculated at approximately 5,000 birds. Estimateswere made for each of four marine regions: southern outercoast, <500 birds; northern outer coast, 1,900; southern PugetSound, 480; and northern Puget Sound, 2,100. Except for thesouthern outer coast figure, in reality an educated guess,estimates of breeding populations were based on data fromcensuses that sampled specific areas, calculated projectednumbers for sub-regions (see Wahl and others 1981), thenadjusted for the proportions of non-breeding birds and thosenot present on the water during censuses but incubating andbrooding at inland nest sites (Speich and others 1992). Weconsider these only approximations of the numbers of breedingMarbled Murrelets, requiring a better defined baseline fordetecting and appraising potential changes in the future.
There may be several factors that could explain theobserved apparent decrease in murrelet abundance in theGrays Harbor study area in the nearshore continental shelfwaters. Some may suggest that the population has beenreduced by the accumulative removal of terrestrial nestinghabitat areas. Such action has the potential to affect MarbledMurrelet populations and may have in particular locations.However, such an evaluation is beyond this paper (see Ralph1994). In our paper, in the time period presented (1971–1993), there are recent indications of changes in the marinecarrying capacity of waters over the continental shelf andslope, off Grays Harbor and beyond. This is reflected in therecent record low abundances reported for several species ofmarine birds, birds representing several different foragingtechniques and positions in marine food changes, and ofvarious geographic affinity. On a slightly larger scale, thereduced abundance of Common Murres in the study areassince 1989 suggests that marine food chains have beenreduced, or have otherwise become unavailable to CommonMurres for several years, because breeding colonies as farsouth as Oregon have experienced depressed reproduction
rates (Varoujean, pers. comm.). The study area numbers inpart reflect the abundance of murres moving north fromOregon breeding colonies after the breeding period.Additionally, the Common Murre breeding population inWashington has declined over the past decade (Ainley andothers 1994; Wilson 1991). Although the Marbled Murreletis not as oceanic a species as other species reported on here,the documented declines in abundance and local breedingsuccess suggest that fundamental changes in marine systemshave occurred, likely expressed by the reduced availabilityof prey. Considering the temporal and geographic scale ofreported effects it is perhaps not surprising that MarbledMurrelet abundance in the Grays Harbor study area waslower the past two years in particular, or even for the pastseveral years.
The patterns of abundance and occurrence presentedherein are descriptive in nature, and represent the “what”stage of the continuing investigation of the Marbled Murreletin the marine waters of Washington, in this case, PugetSound. There is certainly the need to advance ourunderstanding of the marine biology of the Marbled Murreletbeyond the descriptive phases of investigation. We need toaddress, as we have started to do here, “why” MarbledMurrelets are found distributed as they are. However, suchpost priori explanations are limited by their nature in theirpotential to allow understanding of the causes of MarbledMurrelet distribution and abundance patterns. Clearly, thedevelopment and testing of a priori questions, hypotheses,and the development of models are appropriate and necessarysteps for meaningful advancement of our understanding ofMarbled Murrelet biology. However, as necessary anddesirable as these last steps are, a sound and basic knowledgeof the natural history of the species is necessary for theinterpretation and evaluation of study and test results and forour understanding of the factors influencing and controllingMarbled Murrelets (see interesting discussion in Oreskesand others [1994]).
Puget Sound is by its very nature a complex system ofbays, estuaries, channels, passages and straits, greatlyinfluenced by tidal and wind patterns, by the influx of freshwater and nutrients from several river systems, and by itsconnections to the Pacific Ocean. Perhaps a betterunderstanding of, and precision in predicting, the temporaland spatial variable distribution and abundance of the MarbledMurrelet in Puget Sound and along the Pacific Ocean coastwould result from an increased understanding of marinephysical and biological processes and how they link withMarbled Murrelets.
AcknowledgmentsThis paper is the result of combining the data from
several different projects. Consequently, the list of personsand organizations who contributed in many different ways islong. Specific acknowledgments are found in documentscited herein as are the many sources of support (see also
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington
326 USDA Forest Service Gen. Tech. Rep. PSW-152. 1995.
Speich and others 1992). Data from the Grays Harbor areawere made largely possible through the long interest andefforts of bird watchers and other naturalists that supportedvessel time. William Tweit and Dennis Paulson, by theirinterest and participation, contributed in no small way to thesuccess of the Grays Harbor effort. We especially appreciatethe professionalism of Deborah Kristan, as reflected in herreview, and her manipulating and editing of the Puget Sound
data set. The review and comments of C. John Ralph and S.Kim Nelson were particularly helpful, as was the technicalassistance of Linda Long.
The participation of S. Speich in the ConservationAssessment process was made possible by unconditionalsupport from the California Forestry Association, the NationalCouncil of the Paper Industry for Air and Stream Improvement,and the Pacific Lumber Company.
Speich and Wahl Chapter 30 Marine Habitat Preferences, Variability in Washington