Trophic importance of some marine gadids in northern Alaska and their body-otolith size relationships

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CALVIN M.
KAYA
Southwest Fisheries Center Honolulu Laboratory
National Marine Fisheries Service,
NOAA
lionolulu,Hawaii
Present
address: Department
of
Biology
Montana Ste.te University
Bozeman,
MT
59717
ANDREW
E.
DIZON
SHARON D. HENDRIX
Southwest Fisheries Center Honolulu Laboratory
National Marine Fisheries Service,
NOAA
lion.olulu,
HI
96812
TROPHIC
IMPORTANCE OF
SOME
MARINE
GADIDS
IN
NORTHERN ALASKA
AND
THEIR
BODY-OTOLITH SIZE RELATIONSHIPS
Natural
marine
ecosystems
are
beingsubjected
to
ever
increasing
human-induced
stresses, includ-
ing
expanding
commercial fisheries
and
activities
associated
with
the
exploration
and
development
of
offshore
petroleum
resources.
Numerous
studies
ofthe
food
habits
and
trophic
interactions
of
marine
vertebrate
consumers have been un-
dertaken
in
Alaska
during
the
last
5
yr
in
re-
sponse
to
increased
demand
for multispecies ap-
proaches
in
fishery
management
plans
and
the
legal
requirement
for
environmental
assessments
prior to
petroleum
development.
Through
these
and
other
studies
the
importance
of
three
species
-
walleye
pollock,
Theragra
chalco-
gramma,
saffroncod, Eleginus gracilis,
and
Arctic
cod, Boreogadus
saida-in
Arctic
and
subarctic
ecosystems
has
become
increasingly
apparent
(Klumov
1937;
Andriyashev
1954;
Lowry
and
Frost
in
press;
Pereyra
et
aU).
These
species
are
widespread
and
locally
abundant,
are
major sec-
ondary
consumers,
and
are
important
prey
of
other
species (Table
1).
Walleye pollock
are
found
throughout
the
North
Pacific
and
in
greatest
abundance
along
the
conti-
nental
shelf
break
of
the
Bering
Sea. Abundance
decreases
rapidly
north
of
St.
Matthew
Island,
and
they
are
caught
only
rarely
north
of
Bering
Strait
(Pereyra
et
al. footnote
1).
The
species supports a
commercial fishery
of
almost
1million tannually,
one of
the
largest
in
the
world. Walleye pollock
form amajor portion
of
the
diet
of
all
pinnipeds
in
the
southern
Bering
Sea, except
bearded
seals
and
walruses,
and
are
eaten
by
at
least
4species
of
cetaceans,
13
species ofseabirds,
and
10
species of
fishes
in
that
area.
Saffron cod occur
in
the
eastern
Bering
and
Chukchi
Seas
and
throughout
the
western
Arctic
Ocean (Andriyashev 1954).
They
are
also
present,
but
less
abundant,
in
the
Beaufort
Sea. Saffroncod
are
utilized for food by coastal Eskimos.
They
make
up
amajor
portion
of
the
diet
of
ringed
and
spotted
seals
and
white
whales
in
the
northern
Bering
and
southern
Chukchi
Seas.
They
are
also
1Pereyra,
W.
T.,
J.
E. Reeves,
and
R.
G.
Bakkala.
1976. De-
mersal fish
and
shellfish resources of
the
eastern
Bering Sea in
the
baseline year 1975. Processed rep., 619 p. Northwest
and
Alaska
Fisheries
Center, National
Marine
Fisheries
Service,
NOAA, 2725 Montlake Boulevard E.,
Seattle,
WA
98112.
FISHERY BULLETIN: VOL. 79,
NO.
1,1981. 187

TABLE
I.-Marine
mammals,
birds,
and
fishes
reported
to
eat
walleye
pollock,
saffron
cod,
and
Arctic
cod.
20
20
20,21
30
20,21
16,21
30
16,21
30
20
Walleye pollock Saffron cod Arctic cod
12
12
15
15
4
28
15
12,15
15,19
1
44
Species
Marine
mammals:
Northern
lur
seal, Callorhinus ursinus
Steller sea lion, Eumetopias jubatus
Pacific harbor seal, Phoca vitulina richardsi
Spotted seal,
P.
largha
Ribbon seal,
P.
fasciata
Ringed seal,
P.
hispida
Bearded seal, Erignathus barbatus
Fin Whale, Balaenoptera physalus
Minke whale, B. acutorostrata
Sei whale,
B.
borealis
Humpback whale, Megaptera novaengliae
White whale, Delphinapterus leucas
Harp seal, Phoca groenlandica
Narwhal, Monodon monocerus
Harbor porpoise, Phocoena phocoena
Polar bear, Ursus maritimus
Birds:
Glaucous gUll, Larus hyperboreus
Herring gUll,
L.
argentatus
Sabine's
gUll,
Xema sabini
Ross's gUll, Rhodostethia rosea
Ivory gull, Pagophila eburnea
Black-legged kittiwake, Rissa tridactyla
Red-legged kittiwake,
R.
brevirostris
Common murre, Uria aalge
Thick-biled murre,
U.
10m
via
Black gUilemot, Cepphus grylle
Pigeon
gu~lemot,
C.
columba
Walleye pollock
9
3,10,31
23,24,31
22,23,31
14,23,31
32
31,32
5,13
5,25
5
5
Saffron cod
32
14,23
11,23
8,23
5
5
5
5,25
25
Arctic
cod
22,23
22,23
2,6,11,23
2,11,32
1,5
1,5
5
1,5
1,2,25
1
1,2,18
12
12
15
17
17
1,12,30
7,21,30
7,28,30
1,12,28
12
Species
Tufted puffin, Lunda cirrhata
Horned puffin, Fratercula corniculata
Kittlitz's murrelet, Brachyramphus brevirostre
Parakeet auklet, CyclOrrhynchus psittaculus
Least auklet, Aethia pusilla
Arctic tern, Sterna paradisea
FUlmar, Fulmarus glacialis
Shearwaters, Puffinus spp.
Pelagic cormorant, Phalacrocorax pelagicus
Red-faced cormorant,
P.
urile
Red-throated loon, Gavia ste//ata
Jaegers, Stercorarius spp.
Fishes:
Mantic
cod, Gadus morhuB
Pacific cod,
G.
macrocephalus
Walleye pollock, Theragra chalcogramma
Sa~oncod,eegmusgracms
Pacific halibut, Hippoglossus stenolepis
Greenland halibut, Reinhardtius hippoglossoides
Sablefish, Anoploploma fimbria
Flathead sole, Hippoglossoides elassodon
American plaice,
H.
platessoides
Arrowtooth flounder, Atheresthes stomias
Snaillish, Uparis sp.
Eelpout, Lycodes spp.
Sculpins, Ice/us spiniger, Myoxocephalus spp.
Sheelish, Stenodus leuCichthys
Arctic char, Salvelinus alpinus
Atlantic salmon, Salmo sa/ar
21
21
21
21
21
21
4,26,32
26,32
29
26,32
26
29,32
26,29
32
32
32 32
32
32
27,28
1
10. Fiscus and Baines 1966
11. Johnson et
aI.
1966
12. Swartz 1966
13. Nemoto 1970
14. Fedoseev and Bukhtiyarov 1972
15. Watson
and
Divoky 1972
16. Ogi
and
Tsujita 1973
17. Divoky 1976
1. KJumov 1937
2. Vibe 1950
3. Wilke and Kenyon 1952
4. Andriyashev 1954
5. Tomilin 1957
6. McLaren 1958
7. Tuck 1960
8. KenlOn 1962
9. Fiscus et al. 1964
26. Pereyra et al. (text footnote 1).
27. Bendock,
T.
N.
1977. Beaulort Sea estuarine fishery study.
In
Environmental assessment
01
the Alaskan continental shel" annual
reports of principal investigators
for
the year ending March 1977.
Vol. VIII, p. 320-365. Environ. Res. Lab., Boulder, Colo.
28. Bain, H., and A. D. Sekerak. 1978. Aspects
01
the biology
of
arctic
cod, Boreogadus saida, in the central Canadian arctic. Report lor
Polar Gas Project by LGL Ltd., Toronto, Ontario, 104 p.
29. Smith,
R.
L.
1978. Food and leeding relationships
in
the benthic
and demersal fishes
01
the
Gull
01
Alaska and Bering Sea.
In
Environmental assessment of the Alaskan continental shelf, linal
18. Mansfield et
aI.
1975
19.
Bergman and Derksen 1977
20.
Divoky
in
press
21. Hunt et al. in press
22. Frost and Lowry 1980
23. Lowry
and
Frost
in
press
24. Pitcher 1980
25. Frost and Lowry
in
press
report of principal investigators. Vol.
I,
p. 33-107. Environ. Res.
Lab., Boulder, Colo.
30. Springer, A. M., and
D.
G.
Roseneau. 1978. Ecological studies
01
colonial seabirds at Cape Thompson and Cape Lisburne, Alaska.
In Environmental assessment of
the
Alaskan continental shelf,
annual reports
01
principal investigators lor the year ending March
1978.
Vol.
II, p. 839-960. Environ. Res. Lab., Boulder, Colo.
31. Lowry,
L.
F.,
K.
J.
Frost, and J. J. Burns. 1979. Potential resource
competition
in
the
southeastern Bering Sea: Fisheries and phocid
seals. Proc. 29th Alaska Sci. ConI., p. 287-296.
32. Frost and Lowry unpubl. data.

prey
of
other
cetaceans
and
numerous
birds
and
fishes.
Arctic cod
are
circumpolar
in
Arctic
waters
ex-
tending
south
to
at
least
lat.
60° Non
the
Alaska
coast, typically
in
association
with
sea ice (An-
driyashev 1954).
They
are
aspecies of
key
trophic
importance upon which
many
other
far
northern
marine
consumers depend
entirely
for amajor
portion
of
their
yearly
nutritional
requirements.
They
are
eaten
by
at
least
12
species
of
marine
mammals,
20 species
of
birds,
and
5species of
fishes. Arctic cod
are
especially
important
because
in
the
areas
and
at
the
times
when
they
are
abun-
dant
they
are
the
only forage fishes present.
Investigations of food
habits
of
marine
animals
almost
invariably
involve analysis
of
stomachcon-
tents. Morrow (1979) published
preliminary
keys
to otoliths of16 families offishes found
in
Alaskan
Waters
including
the
Gadidae,
whereby
fishes
eaten
by predators can be identified from otoliths
even
after
soft
parts
and
bones have beendigested.
In most
instances
the
size of
the
fish or
meal
can
also be
determined
from otoliths
through
back
cal-
culation offish
length
and/or
weight from various
measurements
of
otolith size (Morrow
1951;
Tem-
pleman
and
Squires
1956;
Southward
1962;
Gjosaeter 1973).
In
this
paper
we
present
relationships
ofotolith
length to fish
length
and
weight for pollock, saf-
fron cod,
and
Arctic cod
of
the
Bering, Chukchi,
and
Beaufort
Seas.
Methods
Samples offishes were obtained
by
otter
trawl-
ing
in
the
Bering, Chukchi,
and
Beaufort
Seas
(Table
2).
Soon
after
capture
all
fishes were iden-
tified, weighed to
the
nearest
0.1 g,
and
fork
length
llleasured to
the
nearest
millimeter.
The
sagittal
otoliths were removed
and
length
and
width
mea-
sured
to
the
nearest
0.1
mm
with
vernier
calipers.
When otolith lengths
and
widths were plotted
against
fish
lengths
as
scatter
diagrams,
the
rela-
tionship between otolith
length
and
fish
length
was found to be less
variable
than
that
of otolith
width
and
fish length. For
this
reason otolith
length
was
taken
as
the
criterion for otolith size
and
used
in
subsequent
calculations.
Casteel
(1976) discussed
in
detail
the
reasons for
using
length
as
the
best
measure
ofotolith size.
We
chose adouble regression method for
relat-
ing
otolith size to fish size (Fitch
and
Brownell
1968; Casteel 1976). For each species
the
relation-
ships
of
otolith
length
tofish
length
and
fish
length
to fish weight were calculated.
In
cases where two
equations were required to fit asingle relation-
ship,
the
inflection point was
determined
by itera-
tion.
The
::lpecified inflection
point
was
varied
by
increments
of 0.1
and
the
pair
of equations
which
minimized
the
combined
deviation
was
selected.
Results
and
Discussion
Regressions offish fork
length
on otolith
length
differed
markedly
among
the
three
species. Those
ofwalleye pollock
and
saffron cod formed two dis-
tinct
straight-line
sections each,
with
inflection
points
at
otolith lengths of
10
mm
in
walleye pol-
lock (fish
length
22 cm)
and
8.5
mm
in
saffron cod
(fish
length
15
cm) (Figures
1,
2).
The
regression
for Arctic
cod
was
rectilinear
over
the
range
of
samples (Figure 3).
Several sources
of
error
are
possible
when
es-
timating
the
size ofafish from
its
otoliths, among
which
are
normal
variability
in
the
ratio
of
fish
length
to otolith
length
and
differences
in
lengths
of
left
and
right
otoliths of
the
same
fish.
The
calculated regression coefficients show
that
such
variability
is
quite
small. Deviation between ac-
tual
measured
and
calculated fish
lengths
was
usually
<5%.
Since food
habits
studies
deal
with
TABLE
2.-Sources
ofAlaskan marine gadids measured to determine otolith length-fish size relationships. T=Theragra
chalcogramma; E=
Eleginus
gracilis; B=Boreogadus saida.
Vessel
a:ld
cruise
no.
Date Area Depth range (m) Trawls (no.) Species
NOAAtShip Surveyor (RP-4-SU· 76AI&II) Mar.-Apr. 1976 Bering 79-173 39 T
NOAA Ship Discoverer (RP-4-DI·76BIII) Aug. 1976 Bering/Chukchi 16-55
16
B.E
USCGC'Gmcwr(AWS7~
Aug. 1976 Beaufort
40·123
2B
NOAA Ship Miller Freeman (RD-4-MF-76BII) Oct. 1976 Bering 15-55
75
B.E
NOAA Ship Surveyor (RD-4·SU·77A
II
,
III)
Mar.-Apr. 1977 Bering
26·150
45
I.E
NOAA Ship Discoverer (RD·4-DI-77AVI) May-June 1977 Bering 30·150
36
B,
T
NOAA Ship Surveyor (RD·4-SU,77BII) June-July 1977 Bering/Chukchi 13·57
17
B,E
USCGC Glacier
(AWS77II1)
Aug.-Sept. 1977 Chukchi/Beaufort 31·400 33 B
ADF&G' skiff (Shishmaref 76)
Mar.
1976 Chukchi 5-10 5E
NOAA Ship Surveyor (RP-4-SU-78AV. VI) May-June 1978 Bering 17·210 78
T.
E
'National Oceanic
and
Atmospheric Administration. 'Alaska Department
of
Fish and Game.
'United States Coast Guard Cutter.
189

lSI
~r--------------------,
lSI
,.;r--------------------,
en
..
...
:
..
I:
.
Otoliths
~
8.5
mm
Y=
1.
74f3X-f3.
13913
N=
36
R=
13.932
..
Ot-olithe >8.5
mm
Y=
2.323X-4.839
N=
1113
R=
13.963
lSI
lSI
'"
..
'"
....
Ot-olit-hs
~
1~.
13
mm
Y=
2.
246X-f3.
5113
N=
158
R=
13.981
.
-"
Otoliths >
1~.~
mm
Y=
3.
175X-9.
77~
N=
98
R=
~.968
lSI
cO
lSI
..;
In
lSI
m
..
lSI
cO
.en
Q
I
t~
~~
...J
.c
~lSI
LL..;
'"
FIGURE
2.-Scatter
diagram
and
regression lines
and
equations
ofotolith length
against
fish fork length for Eleginus gracilis.
16.~
14.~
lSI
cO~--4---+----+----+--->----+----1
2.~
4.~
6.~ 8.~
l~.~ 12.~
lltolllh
L..,glh -
..
";~-4--+---+--+--4_-+-_-+-
_
_+_->___+---1
~.~
2.~ 4.~
6.~
8.~
1~.~ 12.~
14.~
16.~ 18.~
2~.~
22.~
lltolllh
L..,glh -
..
FIGURE
I.-Scatter
diagram
and
regression lines
and
equations
of
otolith
length
against
fish
fork
length
for
Theragra
chalcogramma.
mixed collections of otoliths,
the
cumulative
im-
portance
of
these
differences should
be
minimal.
The
relationships
between
fish
lengths
and
weights of
the
three
species were
best
fit by expo-
nential
equations
of
the
form:
weight
=a(length)b
(Table 3).
These
relationships
may
vary
somewhat
with
time
ofyear, geographic location, sex, repro-
ductive
status,
or fullness
of
stomach. Variation is
probably
most
pronounced
in
sexually
mature
in-
dividuals
with
mature
reproductive products, a
condition which
persists
for only afew
months
of
the
year. Since
small
(juvenile) fishes
are
eaten
by
most
marine
mammals
(Frost
and
Lowry 1980),
birds
(Hunt
et
al.
in
press),
and
other
fishes
(Frost
and
Lowry
unpubl.
data),
this
is probably a
small
source
of
error. Significant differences
in
weight-at-Iength
by sex
and
geographic
area
were
foundfor Arctic
and
saffroncods
by
Wolotira
et
al.2
but
they
justified
use
of
a
single
regression equa-
tion
since
the
differences were
small
(3-7%). Simi-
lar
differences have
been
noted
for walleye pollock
(Bakkala
and
Smith
3
).
Otoliths
are
valuable
indicators
of
the
diet
of
piscivorous
marine
consumers.
Published
keys
such
as
Morrow (1979) allow
determination
of
the
species
and
numbers
of
fishes
represented
by
otoliths
in
stomachs,
intestines,
or scats. By
using
the
relationships
between otolith size
and
body
TABLE
3.-Length-weight
relationships observed for walleye
pollock, saffron
cod,
and
Arctic
cod
in
the
Bering, Chukchi,
and
Beaufort
Seas
(weight =a(length)b).
Range in Regression
Number fork length coefficient
Species sampled (cm) ab(r)
Walieye pollock
109
6-57 0.0077 2.906 0.998
saffron cod 104 6-29 .0050 3.095 .991
Arctic cod 118 7-21 .0018 3.500 .987
190
'Wolotira,
R.
J.,
Jr. 1977. Demersal fish
and
shellfish re-
sources of Norton Sound,
the
southeastern Chukchi Sea
and
adjacent
waters
in
the
baseline year 1976. Processed rep., 292
p.
Northwest
and
Alaska Fisheries Center, National
Marine
Fisheries Service, NOAA, 2725 Montlake Boulevard E., Seattle,
WA
98112.
3Bakkala,
R.
G.,
and
G.
B.
Smith. 1978. Demersal fish re-
sources
of
the
eastern
Bering Sea: Spring 1976. Processed rep.,
233 p. Northwest
and
Alaska
Fisheries Center, National Marine
Fisheries Service, NOAA, 2725 Montlake Boulevard
E.,
Seattle,
WA
98112.

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size
it
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obtain
additional information
sUch
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of fishes
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by
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S. K.
1937.
Saika
(Boreogadus saida (Lepech.» i
ee
znachenie
dlja
nekotorik
zhiznennik
protzesov
arktiki
(Morue
polaire (Boreogadus saida)
et
son
importance
pour
cer-
Literature
Cited
9.0
8.0
......
.'
'1
".
• _
I.
.....
-.
'::,1::"
,
..
,.
Y=
2.
198X+1.588
N=
202
R=
0.981
lSI
..;
N
""
~
lSI
re
lSI
~
!lSI
I
~
llSl
i~
~-
lSI
~
lSI
~
lSI
as
..
lSI
cO
2.0 3.0
Many people assisted us
in
the
collection of
samples, especially
Larry
M.
Shults
who
spent
many
long
hours
sorting
through
trawls
and
measuring fish
with
us,
and
the
officers
and
crew
of
the
NOAA Ship Surveyor who gave
unstint-
inglyof
their
time
and
energy
to
make
our
project
asuccess. Lawrence
R.
Miller provided invaluable
assistance
in
the
computeranalysis ofour
data.
We
thank
J.
E.
Morrow
and
anonymous reviewers for
their careful review of
the
manuscript.
We
are
especially indebted to
John
Fitch
for his
many
helpful suggestions
and
the
moral support
he
lent
throughout
preparation
of
this
manuscript. Fi-
nancial support was provided by
the
U.S.
Bureau
of
Land
Management
Outer
Continental
Shelf
Environmental Assessment
Program
and
Federal
Aid
in
Wildlife Restoration Project W-17-9.
~0 ~0
&0
~0
Oto
11
th
Longth
-
OM
FIGURE
3.-Scatter
diagram
and
regression
lines
and
equations
ofotolith
length
against
fish fork
length
for Boreogadus saida.
191

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proces
vitaux
de
I'Arctique.)
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str.]
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K.
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KATHRYN
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FROST
LLOYD
F.
LOWRY
Alaska
Department
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Fish
and
Game
1300 College Road
Fairbanks,
AX
99701
CAROLINIAN RECORDS FOR AMERICAN
LOBSTER,
HOMARUS
AMERICANUS,
AND
TROPICAL SWIMMING CRAB,
CAILINECTES
BOCOURTI.
POSTULATED
MEANS
OF
DISPERSAL
Recent
reports
of
distributional
extension
for
decapod
crustaceans
occurring
along
the
east
coast
of
the
United
States
include
two poor-
ly
substantiated
records of
American
lobster,
Homarus
americanus
H.
Milne
Edwards,
and
none of
the
tropical swimming crab, Callinectes
bocourti
A.
Milne Edwards, from
the
Carolinas
south ofCape
Hatteras,
N.C. (Williams 1965, 1974
[Carolinas];
Cerame-Vivasand
Gray
1966 [Cape
Hatteras];
Williams
et
al.1968
[North Carolina];
Musick
and
McEachren 1972
[North
Carolina-
Virginia];
Milstein
et
al. 1977 [New
Jersey];
Bowen
et
al. 1979 [Middle Atlantic
area];
Herbst,
Weston,
and
Lorman
1979
[Cape
Hatteras];
Herbst,
Williams,
and
Boothe
1979
[Capes
Hatteras
and
Lookout];
Wenner
and
Boesch
1979 [Norfolk Canyon area]; Perschbacher
and
Schwartz 1979
[North
Carolina)). Occurrences
of both species
in
the
Carolinas
south
of Cape
Hatteras
are
documented
here
along
with
discus-
sion of
their
postulated
means
of dispersal.
Specimens
are
deposited
in
the
U.S. National
Museum of
Natural
History
(USNM), or
are
living
in
aquaria
at
the
North
Carolina
Marine
Re-
sources Center, Bogue
Banks
(NCMRC),
and
the
Hampton
Mariners
Museum, Beaufort (HMM).
Occurrence of Species
Homarus
americanus.-Distribution
of
the
American lobster has been given as,
"East
coast
of
America from
the
Strait
ofBelle Isle, Newfound-
land
(Canada) to Cape
Hatteras,
North Carolina
(U.S.A.);'
at
depths of 0-480 m,
usually
4-50 m
(Holthuis 1974). Reported occurrences of
this
spe-
cies
south
of Cape
Hatteras
are: one
caught
in
a
FISHERY BULLETIN:
VOL.
79, NO.1,
1981.