HomeMy WebLinkAboutAPA350THREESPINE STICKLEBACK -GASTE.ROSTEUS ACULEATUS
AlASKA DEPARTMENT OF FISH &GAME
HABITAT PROTECTION SECTION
RESOURCE ASSESSMENT BRANCH
FRESHWATER HABITAT
RELATIONSHIPS
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FRESHWATER HABnAT RELATIONSHIPS
THREESPINE STICKLEBACK (GASTEROSTEUS ACULEATUS)
By
Stephen S.Hale
Alaska Department of Fish and Game
Habitat Division
Resource Assessment Branch
570 West 53rd Street
Anchorage,Alaska 99502
May 1981
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ACKNOWLEDGEMENTS
Many people from the Alaska Department of Fish and Game and from the Auke
Bay Fisheries Laboratory of the National Marine Fisheries Service freely
gave their time and assistance when contacted about this project and it is
a pleasure to thank them and fishery biologists from other agencies,
especially those who provided unpublished data and observations from their
own work.The librarians of the Alaska Resources library and the U.S.Fish
and Wildlife Service were of great help.
This project was funded by the U.S.Fish and Wildlife Service,Western
Energy and Land Use Team,Habitat Evaluation Procedure Group,Fort Collins,
Colorado.Contract No.14-16-0009-79-119.
TABLE OF CONTENTS
THREESPINE STICKLEBACK
Page
I.INTRODUCTION 1
A.Purpose 1
B.Distribution 2
C.Life History Summary 2
I.Size and Age 4
2.Maturity and ~eproduct;on 6
3.Feeding and Competition 9
4.Distribution and Behavior 13
O.Ecological and Economic Importance 15
II.SPECIFIC HABITAT RELATIONSHIPS 16
A.Spawning 16
I.Temperature 16
2.Water Depth 16
3.Substrl!te 17
4.Aquatic Vegetation 17
5.light 1B
6.Size of Territory 19
B.Incubation 19
I.Temperature 19
2.Dissolved Oxygen 21
C.Juvenile Rearing 21
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O.Adults
1.Temperature
2.Water Depth
3.Current Velocity
4.Dissolved Oxygen
5.Chemical Parameters
6.Feeding
Ill.CONCEPTUAL SUITABILITY INDEX CURVES
IV.DEFICIENCIES IN DATA BASE AND RECOMMENDATIONS
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1.
2.
3.
4.
1.
LIST OF FIGURES
Oistribution of threespine st;c~leback in
Alaska (Morrow,1980)and main study sites.
Conceptual model of relationship between
threespine stickleback embryos and
temperature.
Conceptual model of relationship between
threespine stickleback adults and water
temperature,current velocity,and water
depth.
Conceptual model of relationship between
threaspine stickleback adults and dissolved
oxygen concentration and pH.
LIST OF TABLES
Data table Spawning
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II.Data table -Incubation of embryos
III.D,ta table -Adults
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A.Purpose
This report compiles existing information on the freshwater
habitat requirements.tolerances,and preferences of the
threespfne stickleback,Gasterosteus aculeatus (L.)and provides
a data bilse for habitat evaluation procedures.The threespine
stickleback 1s generally regarded as a hardy species,tolerant of
a wide range of habitat conditions.
I nfonnat;on on phys ;01 og1 ca 1 to 1erances and requ i rements have
been gathered from throughout the range of the species.However,
observations on actual conditions observed in oodies of water
where sticklebacks occur are generally restricted to Alaska.The
threespine has been extensively used as a laboratory fish for
behavioral and pollution studies.Much work has been done on the
physiology of the species in Europe~but relatively little in
North America.It is recognized that habitat requirements may
differ for stocks from different geographic areas,but present
data available from Alaska are insufficient to demonstrate this
within the State.
Within the Gasterosteus aculeatus complex,three forms are
generally recognized {McPhail and Lindsey,1970}.There is a
partially plated freshwater fonn (called leiurus),a heavily
plated marine form {called trachurus},and an intermediate form
(semiannatus).This report is restricted to the totally
freshwater form (leiurus),even though the anadromous forms may
use freshwater habitat during the br~eding season.
This report emphasizes habitat require,nents,primarily those of a
physical and chemical nature.Certain biological factors
affecting the well being of the population,such as feeding,
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predation,camoetition.parasites,and disease.are not
comprehensively treated.
B.Distribution
The threespine !iitickleback is widely dis~ributed in the no:"th'~rn
hemisphere in North America.Europe,and Asia.Except for
Europe,it is not found more than a few hundred kilometers fl"Om
the coast.On the Atlantic coast of North America,it rangl!S
from Baffin Island and the Hudson Bay area down to Chesapea'te
Bay.On the Pacific ~oast of North America,it occurs fron
Alaska to Baja California.On the e.r:stern coast of Asia.-t
ranges from the Bering Strait south to Japan and Korea (Scott and
Crossman,1973;Wootton,1976).
In Alaska,the threespine stickleback occurs in all coastal lreas
from Dixon Entrance to the Alaska Peninsula.the Aleutian Islands
and Bristol Bay (Morrow,1980).It also occurs in the westt!rn
tip of the Seward Peninsula and on St.Lawrence Island;howe\'er,
the freshwater leiurus form is not thought to be present in the
latter areas (McPhail and lindsey,1970;Wootton,1976).(Sl~e
Fi9ure I.1
The leiurus form occurs in both lakes and streams.
c.Life History Sunmary
An excellent synthesis of threespine stickleback 1 ife history has
been presented by Wootton (1976).Behavi ora 1 aspects of
threespine stickleback life history regarding reproduction
(spawning and incubation of eggs)have been extensively
documented and will not be dealt with in this report.Section
II.Specific Habitat Requirements,will examine data on
to1erances and preferences for phys i ca 1 and chemi ca 1 pa rameter;
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Pc •t.Dle.rlbutla"ot Thr •••plne
.lId .eln atudy aU ••.
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MAIN STUDY SITES
[i]Wood AI...Laka.
[&)IIlack Lak ••Clllgnlk Lak.
[!J Lall ••Kartull Lak.·
~L _LalI.
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atlcklabac.itt AI ••ka <Morrow,1980)
history aspects other th.an reproductive behavior for juveniles
and adults.emphasizing Alaskan studies.Much of this section
has been provided by Cannon (1981),with permission of the
author.
1.Size and Age
Cannon (1981)identified three discrete size classes of
threespine sticklebacks in lower Jean Lake on the Kenai
Peninsula.Standard lengths ranged from 22-71 mm.One size
class (Sl •32-33 mm)was captured in surface tows in June
and early July;weekly abundance for this size class in the
catch decreased during this time.A smaller size class (~
•28-29 mm)entered the catch in mid-June and rapidly
dominated the catch by mid-July.During July and August,
two size classes were distinguished in baited minnow traps
fished on the lake bottom.Stickleback (SL •3R-40 mm)were
taken in littoral areas;a larger size class (S-L •51-52 mm)
was captured at depths of 19-20 m.
In Lake Nerka (Wood River lakes)Burgner (1958)observed a
trimoda1ity of size distributions which suggested the
presence of three age groups in early summer.Stickleback
fry did not ap~ear in catches until August and were abundant
in September.The maximum age was reached at three years or
more and in adult fish.females were larger than males.The
life history of populations.inhabiting Karluk and Bare Lake
on Kodiak Island was studied by Greenbank and Nelson (1959).
Lifespan was determi ned to be 2+years,some i ndivi dual s
probably lived past a third winter.Young of the year were
first observed in collections made in early July;this size
cl ass was reported to be very abundant and soon dam;nated
the catch.Mature females attained a somewhat larger size
than did males.Narver (1968)described the general life
history of threespine sticklebacks in the Chignik Lakes on
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the Alaska Peninsula.Determination of age was accomplished
by exam;nat ion of 1ength-frequency.An i nterpretati on of
size distribution for a September sample plac~d Age 0 fishes
within the 20-30 nm fork length (F.L.)size range,Age 1+
within 40-45 mm range and Age 2+within 55-65 mm range.Fry
first appeared in tow net catches in early August;their
density increased rapidly.Engel (1971)also identified
three size classes in beach seine and minnow trap catches in
Johnson,Scout.and Bear Lakes on the Kenai Peninsula.
Length of life was estimated to be 2+years,mean lengths
for ripe males and females in Bear Lake were 50.5 mm (F.l.)
and 49.5 mm respectively.These lengths correspond to those
found by Greenbank and Nelson for Age 2 fish.The abundance
and size of threespine found in lake Aleknagik (Wood River
lakes)were examined by Rogers (1972).Age groups were
assigned on the basis of length frequency.Age 0 fish were
approximately 10-20 mm (SL)by the first week of August and
initially appeared in beach seine hauls in mid-July.Age 1
were 30-35 mm,Age 2 were 40-50 mm,and Age 3 were 55-65 mm.
The variations in age and size structure found in these
studies undoubtedly reflects differences in genetic and
environmental influences between the respective populations.
Direct comparisons are confounded by d~fferences in sampling
gear and measurement methods used.Rogers (1972)
demonstrated that in years of unfavorable growth,major
stickleback age groups showed distinctive length
frequencies;but in years of good growth,the older year
classes overlapped.
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2.Maturity anj Reproduction
Initial sexual maturity in the Lower Jean Lake stickleback
population probably occurs during the second summer of life
(Cannon.1981).Fi shes thou9ht to be Age 1 whi ch were
captured in tows and minnow pots were sexually mature;a
smaller size ~lass,Age 0,'was n-ot mature.Only one age
class at sexual maturity was reported by Greenbank and
Nelson (1959)and McPhail and Lindsey (1970).R0gers
(1968).8urgner (1958).and Narver (1968)suggested that
threespines mature at Age 2.Carl (1953)and Jones and
Hynes (1950)stated that breeding occurred during the first
year of life {Age 0).Variation in these estimates cannot
be explained with available information;however,
differences in length of season,food availability and
genetic character were probably influencing factors.
A decrease in Age 1 stickleback catch per unit effort for
surface tows in lower Jean lake during the course of the
summer and the presence of a similar ~ize class in shallow
shoreline areas suggested that there was a migration from
pelagic to littoral habitats (Cannon.1981).8ecause Age 1
fish were sexually mature and were strongly suspected of
spawning in littoral areas during this time,the movement
appeared to be associated with reproductive activity.
Tinbergen (1952)and Narver (1968)described a spawning
migration of schools of threespine stickleback into
shallows.Males then left schools to establish territories.
Saerends (1957)stated that when reproductive instinct in
threespine is act1vated,adult fish begin to migrate.These
migrations appeared to correspond with a reproductive
stimulus that is only satisfied when the fish reach shallow,
warm waters with abundant vegetation.
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Although characteristic behavior a~sociated with breeding
was not observed by Cannon (1981)in Lower Jean lake,the
majority of the 1ake population apparently spawned in the
lake.Spawning emigrations from the lake were not observed
during the summer.however.they may have occurred prior to
mi d-~ay.Sti ckl ebacks were not found in hi gh dens;ties
milling near the inlet and outlet creek weirs nor attempting
to pass through them.Greenbank and Nelson (1959)reported
that in Karluk Lake.threespine in breeding condition were
observed in the lake.but nesting was not witnessed.
Various authors have noted a variety of materflJ's used in
construction of nests.Tinbergen (1952)described nests
constructed of small twigs.grass and other debris;
Greenbank and Ne1 ..n (1959)and Vrat (1949)discovered nests
fonmerl from cemented sand gr~ins.
Male sticklebacks apparently require tho sight of aquatic
vegetation for the initfatfOn'of·nest building (Pelkwijk and
linbergen.1937.cited by Aron,on.1957).flarver (1968)
noted that the occurrence of stickleback.1n spawning
condition was greatest in areas where submergent flora was
plentiful.Hagen (1967)observed sticklebacks nesting in
still and standing backwaters near dense stands of aquatic
vegetation.Male sticklebacks eXhibiting breeding colors
an~females with distended abdomens were observed in ~r near
thick patches of Chara 1£.in Lower Jean Lake (Cannon.
1981).
Egg production in EuroDe3n populations has been estimat~d to
b.100-200 eggs per spawning (Assen.1967;~ootton.1973 b).
McPhail and Lindsey.1970)reported that in populations from
the Pacific Northwest.graVid females will often spawn
several times per season and l~y 50 to 200 eggs ~t a time.
The number of eggs per spawning and the number of spawnings
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per season have been shown to be positively correlated to
the size of the female (Wootton,1973 .).~e sU9gested that
pODul~tion estimates which ignored the high potential of egg
production and ~urvival provided by t~!f"AJ1ti-seasonal
spawni'lg capacity of large females and the protective
nestin~behavior of males could grossly underestilMte
stickleback production and would incorrectly assess the
importance of this species to the energetics of the
freshwater comounity.Potapova et al.(1966)found that the
quantity and vi.biTlty of egg production 15 related to
growth rate and lipid storage;large f ....les exhibit higher
fecundity.
The rigors of breeding evidently lead to increased
mortality.This view is supported bY detailed descriptions
of the complex nature of stickleback reproductive behavior
provided by Tinbergen (1952)and van lersel (1953,cited by
Wootton,1976).High mortalities reported by Rogers in Lake
Aleknagik may have resulted from a decreased resistance to
high temperature due to the stress of spawning.Narver
(1968)observed high death rates associated with spawnin9
stress in the Chignik Lakes.Alaska.These high mortalities
and the territorial spawning of sticklebacks were considered
possible density-dependent population regulators.Hagen
(l 967)descri bed post-reproductive mortality in the
threespine population inhabiting the Little Campbell River,
B.C.Many adults were found in extremely poor condition;
fungus and other parasitic eruptions were noted.
Sticklebacks in Karluk and 8are Lakes on Kodiak Island
apparently incurred a high mortality after spawning and
appeared nutritionally deprived (Greenbank and Nelson,
1959).Engel (1971)reported post-spawning mortality in
study lakes on the Kenai Peninsula.Alaska.
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3.Feeding and Competitio~
Feeding investigations of stickle~acks in North America have
determined that their diet is composed mainly of zooplankton
and ;nsects.The fi sh are opportuni sti c and food sources
3re related to seasonal and regional availability.Carl
(1953)found cladocera and cope pods in the majori ty of
stickleback stomachs collected in Cowichan Lake.B.C.;
insects,ostracods,anphipods and algae were also observed.
Greenbank and He 1son (1959)di scussed the importance of
chironomid larvae and pupae.cope pods and cladocera in the
feeding of sticklebacks in Karluk and Bare lakes.Other
i terns encountered aecasi Dna l1y were pea clams.as tracods 1
ratiters,snails,leacht:.J.planarians.fish eggs and water
mites.The importance of chironomids decreased over the
summer.Most larger sticklebacks fed on ostracods and fish
eggs;fewer of them fed on cladocera.Chironomid larvae and
entomostraca were found to be primary food types in
threespine sticklebacks in littoral areas of Lake Aleknagik
with Entomostraca being their major prey in limnet;c areas
(R0gers,1968).Sticklebacks inhabitin9 Black Lake,a
shallow lake.fed predominately on insect larvae;
threespines in Chignik Lake.which has only a small littoral
zone,fed totally on zooplankton (Parr,1972).
In Lower Jean Lake.feeding habits of threespines reflected
differences in their distribution (Cannon,1981)and
seasonal availability.Fish utilizing bottom waters preyed
on food species which were not caught in plankton hauls
because most of these species 1he on or near the bottom.
Threespines foraging in the limnetic surface zone preyed
largely on planktonic invertebrates.Age 1 sticklebacks
(St,30-32 nm)fed predominately on copepods in mid-June;
but for Age 0 sticklebacks (~.28-30 nm)rotifers were
dominate prey during J~ly.The availability of prey species
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in surface waters may have fluctuated during the summer,due
to variations in their abundance or distribution.Surface
and bottom feeding threespine sticklebacks in Lower Jean
Lake were ecologically separated (Cannon,1981).
Distributional differences were considered a mechanism by
which stickleback age classes minimized competitive stress
and maximized the utilization of lake resources.The
ability of threespines to utilize 1imnetic waters as well as
bottom environments demonstrated their adaptability to use a
wide rang.of conditions thus allowing exo10itation of
diverse aquatic niches.Separation of age classes between
1imnetic and bottom habitats were believed to be associated
with spawning migrations.Exposure to a wide variety of
habitat types was facilitated by these movements.
Anatomical and physiological differences between age groups
may have enforced separation.
Because spawning and early rearing occur in shallows,
stickleback life history is closely associated with littoral
habitats.The quantity of suitable littoral area
potentially serves as an important stickleback population
regulating factor (Narver,1968).Ecological expansion
would depend on the physical condition and the magnitude of
co-actions (competition and predation)found in adjacent
environs.For these reasons,the variety of potential
niches accessible to sticklebacks and consequently their
distributional patterns can be expp.cted to vary seasonally
and between systems.
Spatial differences exhibited between Age 0 and Age 2
sticklebacks and rearing sockeye salmon in Lr~er Jean Lake
provided a mechanism to reduce interspecific competition
(Cannon,1981).Rearing sockeye were not caught in surface
tows after mid-July;this decl ine occurred in conjunction
with a large recruitment of Age 0 sticklebacks.
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Feeding similarities between sticklebacks and rearing
sockeye have been reported .by Burgner (1958)in the Wood
River Lakes,Alaska,Narver (1968)in the Chignik Lakes.
Alaska,and Krokhin (1957)in the Kamchatka lakes,USSR.
Common prey preferences have suggested potential competition
for food.In nature,many similar species appear to coexist
while seemingly in competition;however,detailed
observations have revealed differences in habitat and
behavior that permit coexistence.
R0gers (1968)compared the food of sockeye salmon fry and
threespine sticklebacl<s in the Wood River Lakes.In
littoral areas.the benthos was used more by sticklebacks
than sockeye fry;surface insects were utilized more by
sockeye.In 1 imnetic areas.winged insects were an
important prey for sockeye,but were rarely ingested by
sticklebacks.Parr (1972),who studied the feeding of
j uvenil e sockeye and res i dent fi sh speci es i ncl udi ng
threespine in the Chignik lakes,indicated that a dis-
similarity existed between stickleback and sockeye feeding.
Sockeye more frequently fed on winged and pupal insects.In
lower Jean lake,a comparison of feeding between these
species (although limited to only a five week period)found
similar differences (Cannon,1981).Sticklebacks and
sockeye inhabiting surface waters during this period
utilized similar prey,but sockeye appeared to prefer winged
insect species more than sticklebacks.
Partitioning rations among sticklebacks and rearing sockeye
in Lower Jean Lake I1Ppeared to be accomp1 ished through
differences in spatial distribution (Cannon,1981).Age 2
threespine utilized bottom habitats.Age 1 sticklebacks and
juvenile sockeye exhibited similar distribution patterns and
feeding;however,spatial overlap and prey similarity are
only prerequisites to potential competition.A significant
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decrease in Age 1 threespines was observed during the summer
in Lower Jean Lake.This reduction was believed to be
associated with reproductive migrations which brought adult
fish bottom environments.The rapid recruitment of Age 0
sticklebacks into the near surface waters during July may
have influenced the apparent shift in vertical distribution
of juvenile sockeye into deeper water stratums and
restricted the return of Age 1 sticklebacks into 1imnetic
areas.The high preference of Age 0 sticklebacks for
rotifers was not shared with sockeye or adult threespine,a
small mouth size possibly was responsible for the apparent
differences in feeding preference,although Age 1 threespine
of approximately the same size fed predominately on cope pods
in June when rotifers were abundant in the net plankton.
Temporal variations in the vertical distributions of prey
species may have been involved.Rogers (1968)suggested
that temporal as well as spatial differences in
stickleback-sockeye feeding relationships could exist.
Narver (196B)developed a conceptual ""del of sockeye-
stickleback co-actions for populations in the Chignik Lakes.
He concl uded that threespi ne sti ck 1ebacks whi ch were
ecologically a littoral species would be displaced from
1imnetic areas by rearing sockeye if the young salmon
rema;ned abundant.Parr (1972)detenni ned that the
abundance of Age 0 sockeye i,the Chignik Lakes had an
adverse effect on resident fish populations including
threespine sticklebacks.Burgner (1958),Kerns (1965),and
Rogers (1972)have suggested a similar population regulation
of stickleback abundance by large Age 0 sockeye salmon
recruitments in the Wood River Lakes.Reduced spawning
escapement due to overharvest or environmental factors have
resulted in periods of reduced sockeye fry abundance in the
Wood River and Chignik lakes.During these years I
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5 tick1eback.s were able to rap;dly occupy vacant ni ches
(Burgner.1958;Naryer.196B).
The threespi ne st i ckl eback is the dam;nant fi sh spec;e$
inhabiting Lower Jean Lake (Cannon,1981).Its dominance is
favored by a prolonged spawning time.a substantial littoral
spawning habitat with abundant submergent vegetation.an
ability to utilize deep and shallow bottom hab1~ats as well
as pelagic areas for feeding,reduced intraspecific
competition between age classes via differences in their
spatial beh~v1or and diurnal migrations,and an apparently
low level of interspecific competition frG~rearing sockeye
salmon in limnetic areas.
4.Distribution and Behavior
An apparent diel migration of threespines in surface waters
of Lower Jean Lake was observed by Cannon (198l)during the
sumner.The abundance of sti ckl ebacks in surface tow net
samples collected in the Wood River Lakes by Burgner (195B)
remained high throughout the day and night.Diel
zooplankton migrations probably influenced stickleback
distribution in lower Jean lake.Stomach analysis of
threespine from different periods showed no significant
decrease in numbers of whole undigested zooplankton.If
feeding continued throughout the day and night.sticklebacks
conceivably would concentrate at depths where prey density
was high.Vertical migrations of limnetic zooplanlctens
conmonly occurring in Alaska lakes have been observed
(Rogers.1974).
A reducti on in i nterspeci fi c competi ti on between
sticklebacks and sockeyes could result from seasonal
variations in vertical distribution.Because threespine are
more resistant to high temperature and illumination and
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because they are equ i pped with a se 1f-conta i ne';:defense
mechanism,near-surface water residence would be less
ecologically intolerable for stickleback than for sockeye.
Outside of breeding season,stickle~ack feed in schools
(Tinbergen 1952).Generally.schoolfng fish are the same
size because fish of similar size swim at the same speed.
Size in sticklebacks varys between sexes and schools
comprised of all males or all females have been observed
(Narver.1968).Possible benefits attributed to schooling
behavior have included increased feeding efficiency,
predator detection and defense,increased ability to locate
a mate,enhanced learning ability,and efficiency of
movement (Eggers.1975).Schoo 11 ng was observed in
sticklebacks in surface waters (juveniles)and in shallow
littoral areas (adults)of lower Jean Lake (Cannon.1981).
MacMahon (1946)described the threespine stickleback as the
fiercest "reshwater fish in Britain for its size.Its
highly flexible fin motions are well suited to feeding in
the dense vegetati on and submerged debri s of the 11 ttora 1
environment.Hagen (1967),who conducted dispersion studies
of threespi ne.reported that sti ck1 ebacks (1 ei urus)were a
sedentary fish.Recaptures of marked fish were never made
beyond 200 m from the point of release.Sticklebacks in
Lower Jean Lake generally exhibited,lethargic swimming
activity (Cannon.1981).Adult sticklebacks observed in
schools feeding in algae beds and solitary fish which were
probably spawning moved slowly through the water.Even
pelagically feeding schools of Age 0 threespine swam
sluggishly near the surface.Rapid swinming motions
occurred during territorial defense and occasionally in
pursuit of prey,but only for short distances.
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O.Ecological and Economic lmoortance
The threespine stickleback is often abundant where it is found
and plays a significant ro~e as a predator,competitor,and prey
species in many lake ecosystems.There has been concern
regarding the stickleback a~a potential competitor with sockeye
salmon (Oncorhynchus nerka)fry but recent studies (Cannon,1981;
Manzer,1976;Rogers,1972;Wootton,1976)have indicated that
competition to the detriment of the sockeye fry does not often
occur.(See also discussion in Section I.e.,life History
Summary.)In artificial situations,such as the reclamation and
re-stocking of lakes with rainbow trout (Salmo gairdneri),the
presence of threespine sticklebacks may be detrimental to the
t~out population (Engel,1971).In many areas,sticklebacks are
an important prey species for predaceous fish such as trout,
sa lmon,and northern pi ke (McPha i1 and li ndsey,1970;Wootton
1976)and for fish-eatin9 birds (Scott and Crossman,1973).
The threespine stickleback has been harvested only to a minor
extent.They have been used for oil,meal,fertilizer,and
animal food,inc1udin9 sled d09 food (Wootton,1976).
Because the threespi ne sti ck 1eback is hardy,easy to keep,and
widely distributed,it has proven to be an important laboratory
fi sh.It has been extens he ly used for behavi ora 1 studi es and
for studies on the effect of water pollution (Wootton,1976).
-15-
II.SPECIFIC HABITAT REQUIREMENTS
A.Spawning
1.Temperature
The surface water temperature during the breeding season in
lower Jean Lake ranged from about 12 to l8°e (Cannon,1981).
In a st;-eam of southern British Columbia.the average
temperature during the breeding season was 16°C (Hagen.
1967)and a stream in England during the spawning season had
temperatures ranging from 16 to 19'C (Lindsey,1962).
Greenbank and Nelson (1959)stated that the water
temperature in two lakes on Kodia~Island may influence the
time of spawning.Threespine sticklebacks spawned earlier
in Bare lake (surface water temperatures from mid·May to end
of July were 4.4-22.B·C)than in the deeper Karluk Lake
(3.3-13.9°C).Ba9ge"nan (1957,eited by Wootton,1976)
found that increased temperatures (about 20°C).given a
sufficiently long daylength.accelerate the maturation
process.
2.Water Depth
Male threespine sticklebacks build their nests in streams or
in shallow areas of lake shores.Hagen (1967)found that
the average depth of nests in Little Campbell River,B.C.,
was 24 cm;some nests were built in water as shallow as 4
em.
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3.Substrate
The nests are constructed of small twigs.plant materia'.
and sand.Some nests may be constructed mostly of sand
grai ns (Greenbank and Ne han.1959;McPha i1 and Li ndsey.
1970).
Nests in the River Wear.England.were built on a -muddy·
bottom (Wootton,1976).The same substrate was used by the
leiurus form in Little Campbell River,B.C.(Hagen,1967).
Hagen gave ma 1e stf ckl ebacks a choi ce between liS J.nd"and
"lrIJd"substrates in the laboratory and found that they
demonstrated a strong preference to build their nests on the
mud.The leiurus form in Hayer lake.B.C.also occurred on
a soft mud bottom (Moodie,1972).Scott and Crossman (1973)
state that threesp1nes prefer sandy areas for nest building
(form not mentioned).
4.Aquatic Vegetation
Stickleback nests are usually found in or near aquatic
vegetation.
Hagen (1967)observed stiCKlebacK (leiurus)nests in the
Little Campbell River,B.C.near dense stands of aquatic
vegetation such as Oenanthe,Potomoseton,Nuphar,~,
Hyosotis,G1Yceria,1l2h!,lemma,and green algae.The fish
always nested among broad1eBved vegetation.
In the laboratory,Hagen (1967)presented males of the
leiurus form with a choice between Oenanthe (a plant found
in the headwaters)and Elodea (a lower river plant)and
found a strong preference for the fanner.In a survey of
streams on Vancouver Island,during breeding season.Hagen
found 1eiurus plentiful only in areas with dense aquatic
-17-
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vegetation.The leiurus fonn in Mayer Lake,B.C.•occurs-
only among the thick vegetation of inlet stream margins and
stream mouths and apparently does not occur in open water
(Moodie.1972).The littoral zone of Mayer Lake ;s densely
covered with Sphagnum and emergent grasses.
Na rver (1966)reported that spawn;"9 st;ck 1ebacks ;n the
Chignik.Lakes were most abundant where aquatic vegetation
was plentiful.However,Karluk and Bare lakes on Kodiak
Island.which have good populations of sticklebacks,have
only sparse aquatic vegetation (Greenbank and Nelson,1959).
5.Light
Gonad maturation of three~pine sticklebacks in the spring ;s
depend~nt on both an adequate light intensity and on an
adequate day1ength (Baggerman,1957,cited by McInerney and
Evans,1970).Baggerman found that high temperature (20·C)
by itself is not effective in inducing sexual maturation,
long photoperiods are also required.Baggerman also showed
that sticklebacks exposed to 269-323 lux (25-30 ft-cand1es)
matured slightly more rapidly than those exposed to 161 lux
(15 ft-cand1es).McInerney and Evans (1970)reported that
sticklebacks (presumably the trachurus form)exposed to an
energy level of 370 ergs/cm(sec in the laboratory had
maturation rates comparable to those of wild fish.This
energy level is equivalent to illuminance levels ranging
from 230 lux (at the green wavelength)to 5 lux (at the
purple wavelength).McInerney and Evans also tested the
effect of 1i ght qual i ty (wave 1ength)by expos;ng fi sh to
four segments of the visible spectrum ranging from 388-653
millimicrons (long ultraviolet to short red).They found no
major differences among the four in affecting the rate of
gonad maturation.
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6.Size of Territory
Wootton (1976)states that the maximum density of threespine
stick.leback nests is 4_5/m 2 and that the minimum distance
between nests 15 30-50 em.The average di stance between
nests in the River Wear.England,ranged from 143-237 em
(Wootton,1976).
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B.Incubation of 8mbryos
1.Temperature
The time to hatching is directly dependent on temperature.
Wooton (1976)plotted the data of several investigators and
found that the time to hatching varies from about 5 days at
25"C to about 15-43 days at B"C.At lB-19"C,hatchin9
occurs in about 8 days and the y01 k sac is absorbed in
another 4 days.Studies in Alask.a have reported hatching
times of 14 days at a water temperature ranging from 9_16°C
(Kodiak Island;Greenbank and Nelson,1959)and 5 days at a
water temperature varying from 21.1-22.8°C (Kenai Peninsula;
Engel,1971).Neither of these studies was in situ.
Heuts (1947),in laboratory studies in Belgium,found that
the best survival of eggs incubated in freshwater occurred
at water temperatures of 15-26°C.Heuts showed that the
eggs are adapted to a narrow range of temperatures.Lindsey
(1962)reared the freshwater fonn in the laboratory at
temperatures ranging from 10_28°C.No eggs were
successfully reared in freshwater at 10°,12°.14°,or 28°e,
and survival at 16°and lBoe was less than optimum.
However,there may have been factors present other than
temperature which lowered the survival rate.Lindsey also
found that fish reared at higher temperatures (22°e and
greater)had a higher proportion of females than fish reared
-19-
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at 20°C and below.Lindsey further showed that eggs from
females with few lateral plates have a higher optimum
development temperature than eggs from females with more
plates.The approximate optima were:26°C (2 maternal
plates),20-27"C (4 or 5 maternal plates),20-22"C (6
maternal plates),and 16"C (7 maternal plates).
Wootton (1976),citing the work of 5warup (1958.1959).
stated that abnormal development occurs when newly
fertilized e99s are exposed to very low (O"C)or very high
(33°C)temperatures for a duration as short as 1.5-3.0
hours.One of the abnormalities was the production of fish
with a triploid number of chromosones.These fish developed
and grew at the same rate as normal (diploid)fish.but were
misshapen.
Hagen (1967)measured water temperatures ranging from 16 to
23"C during the nesting of sticklebacks in Little Campbell
River,B.C.Some approximate surface water temperatures for
various lakes in Alaska measured at the approximate time
incubation occurs are:Bare Lake,61 -23°e,and Karluk
Lake,41 -15°e (Greenbank and Nelson.1959);lower Jean
lake,13 -18°e (Cannon,1981),Johnson and Scout Lakes on
the western Kenai Peninsula,12 -18°e,and Bear lake in the
Kenai Mountains.7 -15°C (Engel,1971);Wood River lakes,
10 -14"C (Rogers.1968);Lake Nerka,9 -lB"C (Burgner,
1958);Black Lake,12 -15"C.and Chignik Lake,9 -]J"C
(Parr,1972).
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c.
D.
2.Dissolved Oxygen
Given the fact that sticklebacks nest in shallow areas with
adequate light penetration and aquatic vegetation.dissolved
oxygen levels in the water near the nests are probably
rarely a limiting factor.However.the eggs are placed
inside covered nests,often on muddy bottoms containing much
dead organic matter.and often in areas of little or no
current,so circulation of water through the nest to replace
the oxygen used by the enmryos 1s necessary and is
accomplished by the fanning of the mole.In the absence of
fanning,van lerse1 (1953,ci~ed by Wooton,1976)noted that
eggs became moldy and died.The rate of fanning reaches a
peak shortly before hatching;presumably.this is when the
oxygen requirement of the _ryos is highest.When van
lersel ran water low in dissolved oxygen and high in carbon
di ox;de through a nest ;n the 1aboratory I the amount of
fanning by the male increased.
Juvenile Rearing
After leaving their nests,young sticklebacks form schools,which
may be an adaptation for cover,and eventually join the adults.
There is 1ittle information to suggest that young of the year
have habitat tolerances,preferences,or requirements different
from those of adults.
Adult Life
1.Temperature
The adult threespine stickleback is usually regarded as a
eurythermal fish (Wootton.1976).Jordan and Garside (1972)
studied the upper lethal temperature of threespine
sticklebacks (probably the trachurus form)from the harbor
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of Ha1ffax,Nova Scotia (where the sal inity ranges from
20·30 pptl which had been acclimated to various combinations
of temperature and sa1fnity.The highest upper lethal
temperature (28.8·C)was noted for fish acclimated to 20·C
and 30 ppt salinity,tested at 12 ppt,and the lowest upper
lethal temperature (21.6·C)was shown by fish acclimated to
10·C and a salinity of 0 ppt,tested at 30 ppt.Fish which
had been acclimated to freshwater at 20·C,and tested in
freshwater,had an upper lethal temperature of 27.2·C.
There were no significant differences in survival among the
different sized fi sh (tota 1 1engths ranged from 30-80 1lIII)
tested.Coad and Power (1973),citing 8ertin (1955)state
thot the threespine can tolerate temperatures around 25·C
and is also tolerant of temperature changes.Heuts (1947)
found with the leiurus form collected at O·C and placed into
water at 25 -28·C that fish with fewer lateral plates
survived longer than fish with more lateral plates.Hean
survival time was around 40 hours.Heuts also stated that
the freshwater form will not tolerate 11ow'temperatures.
The threespine stickleback is found in Alaskan lakes with
late spring to early fan temperature ranges of anywhere
from O·to 23·C (Burgner,1958;Cannon,1981;Engel,1971;
Greenbank and Nelson,1959;Parr,1972;Rogers,1968 and
1972).The stickleback may not be found at the extremes of
thi s range if they have a choi ce of more moderate
temperatures.Little information is available on
t~rlture preferences or on temperature distribution
during late fall,winter,and early spring.In the summer,
sticklebacks move into warmer,shallower,water but this is
probably more a function of reproductive and feeding
requirements than a demonstration of temperature
preferences.
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Temperature influences the rate of growth.Wootton (1976),
citing the work of Cole,reported that the mean growth
efficiency increases from 5.9%at 7.0°C to 11.3~at 20.0°C.
Beukema (1968,cited by ~ootton,1976)found that the
feeding rate also depends on temperature,the rf'te of
stomach evacuation ranging from about 16 hours at l"i-12°C to
"one night"at 18-20 0 e.
2.Water Depth
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3.
The depth distribution of threespine sticklebacks in lakes
results from temperature preferences,feeding migrations,
reproduction requirements,predation and competition.They
are generally a ~hallow water fish,particularly in the
suntner when they move into the shoals along the shore.
During the summer they range from the surface to the bottom
in lower Jean Lake (21.3m)(Cannon,1981)and Johnson (4.0
m),Scout (6.1 m),and Bear (18.3 m)lakes (Engel,1971)on
the Kenai Peninsula.In Karluk lake on Kodiak Island,they
have been caught from the surface down to 24.4 m,but none
were caught in attempts at 38.4 m or at 61.0 m (Greenbank
and Nelson,1959).Very few fish were caught below surface
waters in Lake Nerka during the sumner;the deepest
stickleback was caught at 7.3 m (Burgner,1958).
Current Velocity
Although threespine sticklebacks commonly occur in streams,
they prefer areas with little or no current (Hagen.1967).
Hagen conducted a thorough study in Little Campbell River,
B.C .•and determined that swift waters are an unfavorable
habitat.The average current velocity in a section of the
stream where the leiurus fonn was plentiful was 3 em/sec
(grad;ent about 1.5 m/km).A section with an average
velocity of 23 cmlsec had only a few sticklebacks of the
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leiurus form and a riffle section with an average velocity
of 74 cm/sec had none.When Hagen transplanted the fish
from the low current area into an area of fast current.they
migrated into areas of slower current.When Hagen gave
males a choice between standing water and moving water in
an aquarium.they demonstrated a strong preference for
nesting in the standing water.Hagen also reported that
some fish successfully passed through a 70 m long culvert
with a current velocity of 92 cm/sec,although this was not
a common occurrence.
4.Dissolved Oxygen
Krokhin (1957)calculated that the oxygen consumption of
threespine sticklebacks in Kamchatka lakes where the
temperature ranges from 2.0 to l4.3°C would range from 0.127
-0.365 m9 Oz/hr-g live weight.He found that fish in the
laboratory at a temperature range of 0.5 -19.5°C used
oxygen at a rate ranging from O.lZ -0.55 mg OZ/hr-g 1 ive
weight.Threespine sticklebacks from some English streams
had an oxygen consumption rate ranging from 1.0 microliters
OZ/hr-mg dry weight (1 microliter <1.43 micrograms)for 500
mg dry weight fish to about 2.5 microliters/hr-mg dry weight
for 70 mg dry weight fish (temperature not given;Lewis et
a1..197Z).
Jones (1948 and 195Z.cited by Wootton.1976)stated that
the minimum dissolved oxygen (00)level at which threespine
sticklebacks can exist is about 0.Z5 -0.50 mg/l.Jones
found that the avoidance response of sticklebacks is
triggered when the fish are exposed to water with a 00 level
of 0.3 mgll at low temperatures.At 20°C,the response
occurs at 2.0 mgll.indicating the fish have a lower
tolerance for low 00 conditions at higher temperatures.In
a survey of some English streams,Lewis et al.(1972)found
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that threespine sticklebacks were most abundant in waters
with 8 -12 mg O2/1.less abundant in waters with 6 - 8 mg
O2/1,and absent from waters with 2 - 5 mg O2/1.
In Lower Jean Lake,sticklebacks were caught near the lake
bottom during the S~~r where 00 levels ranged from 3.0 -
6.5 mg/1 (Cannon.1981).These levels apparently had no
effect on feeding activity.Dissolved oxygen levels above
the thermocline ranged from 10.1 -14.0 mg/l.Greenbank and
Nelson (1959)reported that the ,tick1eback population in
Karluk and Bare lakes are in waters where there is an
"abundance"of dissolved oxygen at all depths during the
sunmer.They also stated that the threespine stickleback
1115 known to survive over winter in shallow lakes in
northern temperate and subarctic zones,where ....dissolved
oxygen sinks to a trace ....".
5.Chemical Parameters
Threespine sticklebacks have been reported to occur 1n
waters with a pH range of:7.0 -8.7 in Karluk and Bare
Lakes (Greenbank and Nelson,1959);6.3 -7.0 in three Kenai
Peninsula lakes (Engel,1971);and 6.8 in Little Campbell
River,B.C.(Hagen.1967).Jones (1948.cited by Wootton,
1976)tes ted the tal erance of the fi sh to a wi de pH range
and reported that they avoided a pH of less than 5.6 or
greater than 11.4.
A survey of some English streams conducted by Lewis et a1.
(1972)showed that the threespine did not occur in waters
which smelled of hydrogen sulfide and rotting vegetation.
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6.Feeding
Krokhin (1957)calculated that threespine sticklebacks in
some Kamchatka lakes.where the annual temperature range is
2.0 -14.3"C.consume daily between 0.08 9/fish in winter and
0.23 g/fish in August.This represents 1.8 -5.1%of body
weight.The monthly food consumption ranged from 2.70 -
6.77 g/fish.for an annual total of 42.49 g.which is
equivalent to 8 or 9 times the average weight of an adult
(4.5 g).In Lake Oal 'neye.Krokhin (1970)calculated that.2
the monthly food consumption ranged from 0.80 gIg 1 ive
weight in January to 2.185 gIg live weight in August.
Manzer (1976)reported that the daily ration of threespine
stickleback in Great Central Lake on Vancouver Island,B.C.,
was 6.6%of the body weight in July and 7.8%in October.
Wootton (1976)states that.to support a population of
threespine sticklebacks.an area must produce suitable food
---on-the order of 10 -100 g/m 2-yr.
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III.CONCEPTUAL SUITABILITY INDEX CURVES
Conceptua 1 5uitabil ity ;ndex curves are presented for water
temperature.current velocity,water depth,dissolved oxygen
concentration,and pH in Figures 2 through 4.Data to support the
curves ;s included in Tables I.II.and III.These curves hould ,lot
be construed as a graphical presentation of actual data.Rather,they
are intended to be hypothetical models of the relationship between
threespine sticklebacks and certain environmental parameters.As with
any hypothesis,they must be tested and verified before being applied
to any particular situation.
The curves are based on publ ished and unpublished data and on
conversations with fishery biologists who hav~~orked with threespine
sticklebacks.Both experimental laboratory data and field
measurements and observations were used.The suitabil ity index for
each environmental parameter ranges from zero to one.An index of one
indicates an optimum or preferred level of that particular parameter
and an index of zero indicates a completely unsuitable level.
Data from laboratory physiological studies in European populations of
threespine sticklebaCKS indicates that the optimum temperature during
incubation of embryos is B or goC higher than the average surface
water temperature during the presumed incubation period in several
Alaskan lakes (Figure 2).Until the temperature tolerance of Alaskan
sticklebacks has been studied in physiological laboratory studies,it
must be assumed that the optimum temperature for European sticklebacks
(about 22°C)does not apply in Alaskan waters which rarely reach 22°C.
A problem encountered in constructing the curves is that much of the
data in the literature concerning environmental parameters of
threespine stickleback habitat does not relate various levels of the
parameters to some measure of habitat sUitability.Often,ranges of
the parameter are given based on measurements taken throughout the
area occupied by threespines but there is no indication that one point
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on th~range is any better or worse than any other point in terms of
habitat sUitability.
The curves are drawn using data from throughout the natural rar:ge of
the threespine stickleback.Although there probably are differences
in habitat preferences and tolerances for different geographical areiS
or even in different streams of the same geographical area,there i!-
not enough data to support drall'ling separate curves at this time.
However.one must be aware that any point on the curve,especially
toward either extreme.may bp.unsuitable for a particular stock.Hew
far the stock deviates from the curve fIlJst be determined by field
measurements and experimentation with that particular stock.
A second precaution regarding the curves concerns the interaction If
various parameters.A given level of one parameter can have a
different effect on the fish as the level of another parameter varitls.
For example,a dissolved oxygen concentration of 3 mg/l may be
suitable at a water temperature of 5°C.but unsuitable at a
temperature of 20°C.Ideally.given enough data.a separat..dissohed
oxygen curve should be drawn for each of several different
temperatures.The overall suitability of any particular habitat is a
summation of the interacting effects of many parameters.
A third precaution to consider is that the effect of less than optinJm
levels of any parameter on the fish depends strongly on the duration
of exposure.Stickleback embryos exposed to a water temperature of
33°C for 1.5 -3.0 hours experience an abnormal development (Swarup,
1958 and 1959,cited by Wotton,1976),but survive.Embryos exposeQ
to 33°~for a longer period of time would die.
A1so.di fferent ages with;n ali fe stage probab 1y have di fferen 1•
habitat requirements.For example.the oxygen requirement of emtryos
is highest just prior to hatching.However,because of insufficient
data.it is not possible to draw separate curves for different ages.
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Overall,the suitability index curves presented in this report provide
an indication of conditions which make a desirable threespine habitat
and conditions which make a less desirable habitat.Also.although
the curves are general,they can sho~:di fferences ; n habi tat needs
among threespi nes and other species.Further,the process of
constructing these curves is beneficial in defining areas where more
data is needed.Lastly.these curves can aid in the design of
experiments and sampl ;09 programs.As more data becomes available.
these hypothetical curves can be further refined.
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Table 1.THREESPINE STICKLEBACK
Adult -Spawning
Parameter Observed Values Remarks location Reference
Temperature.ea.12-18
'C
5.6 -22.8
3.3 -13.9
16
16 -19
Wa ter depth,4
em
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surface water temperature during Lower Jean lake
breeding season
surface water temperatures Bare Lake
during June and July Karlu~lake
average during breeding seasop.Little Campbell
River,B.C.
during spawning season Hobson's Brook.
England
shallowest nest Little Campbell
River.8.C.
average depth for nests
Cannon (1981)
Greenbank and
Nelson (1959)
Hagen (1967)
Lindsey (1962)
Hagen (1967)
-------------------Table II.THREESPINE STICKLEBACK
Incub3Lion of Embryos
Parameter Observed Values Remarks location Reference
Temperature,16 -26 best survival rate Belgium (laboratory)Heuts (1947)
·C
10 -14 no eggs successfully reared England (laboratory)Lindsey (1962)
16 -lB survival less than optimum
20 -26 best survival
2B no eggs successfully reared
>22 lower male:female ratio
than fish reared at ~O°C
16 -26 optimum development temperaturesi the,fish with fewer maternal platesw
~lldving higher optimum temperatures,
0,33 exposure to these temperatures England 7 Swarup (195B and
for 1.5 -3.0 hours causes (1 aboratory)1959,cHed b~
abnormal development Wootton,1976
B time to hatching is 15 -43 days various Wootton (1976)-5 time to hatch1ng 1s 5 days
25 time to hatching is 5 days
13 -lB approximate surfac~water temp.lower Jean lake Cannon (19B1)
during incubation ~erlod
12 -18 approximate surface water temp.Johnson lake Engel (1971)
during incubation period
Table II.Cont'd THREESPINE STICKLEBACK
Incubation of Embryos
Parameter Observed Values Remarks Location Reference
Temperature 12 -18 approximate surface water temp.Scout lake
'C during incubation period
7 -15 approximate surface water temp.Bear lake
during incubation period
67 -23 approximate surface water temp.Bare lake Greenbank and
during incubation period Nelson (1959)
47 -15 approximate surface water temp.Karluk lake
during incubation period
10 -14 approximate surface water temp.~ood River lakes R0gers (1968)
during incubation period
,
9 -18 approximate surface water temp.lake Nerka Burgner (1958),
during incubation period
12 -15 approximate surface water temp.Black Lake Parr (1972)
during incubation period
9 -13 approximate surface water temp.Chignik Lake
during incubation period
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-------------------Table Il I.THREESPINE STICKLEBACKS
Adults
Parameter Observed Values Remarks Location Reference
Temperature.27.2 Upper lethal temperature for the Halifax,Nova .Jordan and Garside
·C trachurus (1)form acclimated to Scotia (l972)ZOde 1n freshwater (laboratory)
25 tolerable general Bertin (1925,cited
by Coad and Power.
1973 )
25 -2B The le1urus form collected at 0°Belgium Heuts (l947 j
and placed into water with these (laboratory)
temperatures died after an average
of 40 hours;fish with fewer
lateral plates survived longer
than fish with more plates,
ww 7.0 mean growth efficiency of 5.9%7 Cole (unpublished,,
cited by Wootton.
1976 )
20.0 mean growth efficiency of 11.3%
11 -12 rate of stomach evacuation was ?Beukema (196R,
16 hours cited by Wootton,
1976)
18 -20 rate of stomach evacuation was
"one night"
Table III.Cont'd THREESPINE STICKLEBACKS
Adults
Loea ti on Reference
lower Jean lake Cannon (19B1)
Johnson and Scout Engel (1971)
lakes
Greenbank and
Nelson (1959)
R0gers (1968)
Bur9ner (1958)
Parr (1972)
Bare lake
Bear lake
Karluk Lake
Lake Nerka
Wood River lakes
Remarks
Jun -Sept 6 yrs;observed water temp.
fi sh present
early Jul -early Sep.;observed water temp Black lake
ft sh present
early June-mid-JulYi observed water temp.
fish present
Jun 1 -Sep.15;observed water temp.
ft sh present
Jun 1 -Sep.15;observed water temp.
fish present
mid May -mid Sept.;observed water temp.
fish present
mid Hay -mid Sep.;observed water temp.
fish present
Aug.,observed water temp.fish present
11.5 -15.0
Parameter Observed Values
Tempera ture •6.9-17.7
·C
3.9 -15.0
8.9 -18.3
4.4 -22.8
3.3 -15.0
6.7 -14.2
0.0 -20.0
,
w
f"
9.0 -13.0 late Jun -early Sep.;observed water temp Chignik lake
fi sh present
-- ----•- -- --- -- ----Table III.Cont'd THREE'IHE STICKLEBACKS
Adults
Parameter Observed Values Remarks location Reference
Water Depth I 0-21.3 (bottom)suntner lower Jean lake Cannon (19B1)
m o -4.0 (bottom)sunmer Johnson Lake Enge1 (1971)
0-6.1 (bottom)SUlTlT1er Scout lake
o -18.3 (bottom)sunmer Bear lake
o -24.4 sUlTITler Karluk Lake Greenbank and Nelson
(1959 )
38.4,61.0 none caught;sunmer
7.3 greatest depth at which fish lake Nerka 8urgner (1958),were caughti summerw
'('
o -10 Jun -Sept.Kamchatka Krokhln (1957)
Lakes,USSR
50 -60 Deco.Jan .•Feb.
Current 3 average in leiurus habitat Little Campbell Hagen (1967)
Velocity,River t 8.C.
em/sec
(stream 23 marginal habitat
population)
74 poor hab i ta t
92 fish were able to negotiate
a 70 m culvert with this velocity
Table Ill.Cont'd THREESPi~E STIC~LEBACKS
Adults
,
ll:,
Parameter
Dissolved
Oxygen
"gil
pH
Observed Values
0.25 -0.50
0.3
2.0
8 -12
6 - 8
2 - 5
3.0 -6.5
10.1 -14.0
<:5.6 or)1l.4
6.8
6.3 -7.0
7.0 -8.7
Remarks
minimum level tolerable
level at which avoidance response
is triggered at low temps.
level at which avoidance response
is triggered at 20'C
fish most abundant in reaches with
this level
fish less abundant in reaches with
this level
fish absent from reaches with this
level
healthy f1sh caught near lake bottom
at this level;summer
level above thermocline,summer
avoided by fish
measured value
r.reasured value
measured value
location
Grea t Brita 1n
(laboratory)
England
lower Jean lake
Great Britain
(laboratory)
Little Campbell
River,8.C.
kenai Peninsula
lakes
Karluk and Bare
lakes
Refp.rence
Jones (1948 and
1952,cited by
Wootton,1976)
lewis et a1 .•
(1972)
Cannon (1981)
Jones (1948,cited
by Wootton,1976)
Hagen (1967)
Engel (1971)
Greenbank
and Nelson (1959)
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THREESPINE
STiCKLEBACK
INCUBATION
Alaslaon
obs.rvatlons
i 10 !
•European•It:""'"physiological""c uud .•s,
~05
-"0--,
'",,
0
0 __10 __..-20 30
Woter temperoture,·C
Figure 2 Conceptual model of relatIonship between
Threesplne sttcldebock embryos and tdmperoture.
See text for Qualifications for use of this curve
(NOT RECO.....ENDED FOR APPl.ICATION TO SPECIFIC wATERSHEDS
WITHOUT FIELD VERIFICATION)
-37-
See text for qualifications
for use of these curves
t HOT RECO"'MENQ£D 'OA "PPUCATION
TO S"C.te WAT!RSH£DS W~THOUT
FIELD YEIlI ..IC.'TION)
ADULTS
0
0 10 20 30
Water temperature,'C
10....
I,
I
0.5l,
x IwO~::0.,.
I I
>-30 60 90...
~Current velocity,em/seciii
:!
5 1.0V>,
I ,.......Summ.,\,,
,rearing,dls~"but ton W,nt.,
I distribution,
I
0.5 '~Spawn,ng
:distribution,,,
0
20 40 60
Water depth,m
0.5
THREESPI"JE
STICKLEBACK 10
rlgure 3.Conceptual model of relatlonsnlp between ThreesPlne
stickleback adults and woter temperature.C\Jr,ent
",eloelty (streom pOPulations only).and woter depth
-3B-
Figure 4.Conceptual mod.1 of relatIonship between
Threespine stickl.back adults and dissolved
oxygen concentration and pH .
Dissolved nxygen,",gIl
12
1210
See text for Qualifications
for use of these curves
(NOT .-eCOMMENC!D FOR APP1..ICATION
TO ,"ePIC WATERSHEDS Wr1'HOUT
MLD VERI'ICATION)
8
86
pH
ADUL TS
4
42a
0'-+--::::::::....+----+----+a
0.5
1.0
x
wo
Z
>1.0...
~
c«...
=>
'"0.5
THREESPINE
STICKLEBACK
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•-39-
IV.DEFICIENCIES IN DATA BASE AND RECDHHENDATIONS
Although extensively used in behavioral and pollution studies
elsewhere,the interest in the threespine stickleback in Alaska has
centered mainly on its supposed role as a competitor with rearing
sockeye salmon and rainbow trout.These Alaskan studies,therefore,
have naturally concentrated on the distribution and feeding of
sticklebacks.
Physiological studies on Alaskan populations are needed to define the
tolerances to environmental parameters such as temperature and
dissolved oxygen concentration.This need ;s dramatically shown by
the difference between European physiological studies of temperature
tolerance and the known temperature regime existing in Alaskan lakes
where the sticklebacks are present (Fig.2).
Information is needed
within the State.To
on lake populations.
on stream populations of threespine sticklebacks
date.Statewide investigations have concentrated
More data from Southeast Alaska is needed to describe regional
differences within the State.The Southeast populations may be more
similar to the populations described 'in British Columbia than they are
to the Bristol Bay populations.Additionally.there is some
indication that sticklebacks in the Kenai Peninsula area are exposed
to and adapted to higher temperature regimes than sticklebacks in the
Bristol Bay area.
A study in Alaska of differences in habitat requirements among the
three forms of Gasterosteus aculeatus (leiuru1,trachurus,and
semiannatus)would be beneficial as strong differences have been
demonstrated elsewhere;for example,Hagen1s (1967)work in British
Columbia.Virtually nothing is known of the populations on the Seward
Peninsula and Saint Lawrence Island (presumably trachurus).Because
these populations are on the edge of the species range,studies of
-40-
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these fish may provide interesting information about the environmental
limits of the species.
Very little infonnation is available on the habitat needs of the
threespine sticKleback during the stage immediately after hatching )ut
prior to leavir-g the nests.During this period 9 they are still !In(l~r
the parental care of the male.There are also many questions ibollt
their life history between leaving the nest until they first begin to
show up ;n mi nnow traps and fyke nets as young of the yea r.In
Alaska,few observations have ever been made of actual nests
-41-
LITERATURE CITED
Aronson,l.R.1957.Reproductive and parental behavior Chapt.III.
Part 3,271 -3J4 in Margaret E.Brown (ed.).The physiology of
fishes.Vol.II.Behavior.Academic Press.New York.526 pp.
Assen.J.van den.1967.Territory in the threespined stickleback
Gasterosteu5 aculeatus L.An experimental study in intra-specific
competition.Behavior Suppl.16:1-164.
Baerends.G.P.1957.Ethological analysis of fish behavior.Pages
229 -269 in Mar9aret E.Brown.(ed.).The physiolo9Y of fishes.
Vol.I.Academic Press,New York.pp.
Burgner,Robert l.1958.A study of fluctuations in abundance.growth and
survival in early life stages of the red salmon (Oncorhynchyus nerka
Walbaum)of the Wood River lakes,Bristol Bay,Alaska.Ph.D.Thesis.
University of Washington~Seattle.200 pp.
Cannon,Richard.1981.Summer feeding and distributional behavior of
threespined stickleback,Gasterosteus aculeatus,in lower Jean lake,
Alaska.1974.M.S.Thesis,University of Alaska,Fairbanks.pp.
Carl,Clifford G.1953.limnobiology of Cowichan lake,British Columbia.
J.Fish.Res.Bd.Can.,9(9):417-449.
Coad,R.W.and G.Power.1973.Observations on the ecology and
phenotypic variation of the threespine stickleback,Gasterosteus
acu1eatus l.,1758,and the blackspotted stickleback,~.wheatlandi,
Putnam 1867,(Osteichthyes:Gasterosteidae)in Armory Cove,Quebec.
Canadian Field Naturalist 87:113-122.
Eggers,Douglas M.1975.A synthesis of the feeding behavior and growth
of juvenile sockeye salmon in the limnetic environment.Phd Thesis,
University of Washington,Seattle.217 pp.
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II
fl
Engel,Larry J.1971.Evaluation of sport fish stocking on the Kena;
Peninsula-Cook Inlet Areas.Alaska Department of Fish and Game.
Federal Aid in Fish Restoration,Annual Report of Progress,1970-
1971.Project F-9-3.Job No.G-ll-F.Vol.12:1-34.
Greenbank,John,and Philip R.Nelson.1959.life history of the
threespine stickleback Gasterosteus aculeatus Linnaeus in Karluk Lake
and Bare Lake,Kodiak Island,Alaska.U.S.Fish and Wildlife Ser.
Fish.Bull.153.Vol.59:537-559.
Hagen"M.J.1967.Isolating mechanisns in threesp;ne sticklebacks
(Gasterosteus).J.Fish.Res.Bd.Canada,24 (B):1637-1692.
Heuts.M.J.1947.Experimental studies on adaptive evolution in
Gasterosteus aculeatus L.Evolution.1:89-102.
Jones,J.W.,and H.B.N.Hynes.1950.The age and growth of Gasterosteus
aculeatus.Pygosteus pungitius.and Spinachia vulgaris.as shown by
their otoliths.Jour.An.Eco1.19:59-73.
Jordan,C.M.•and E.T.Garside.1972.Upper lethal temperatures of
threespine stickleback,Gasterosteus aculeatus (L.).in relation to
thermal and ostomotic acclimation,ambient salinity.and size.Can.
J.Zool.50:1405-1411.
Kerns.Orra E.1965.Abundance and size of juvenile red salmon and
major competitor species in 11 iamna lake and lake Clark,1962 and
1963.Univ.Washington Fish.Res.Inst.Circ.231.35 pp.
Krokhin.E.M.1957.Determination of the daily food ration of young
sockeye and three-spined stickleback by the respiration method.
(Transl.from Russian).Fish.Res.Bd.Can.Transl.Ser.No.209.
14 pp.
McPhail,J.D.,and C. C.Lindsey.
northwestern Canada and Alaska.
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Krokhin 7 Yeo M.1970.Estimation of the biomass and abundance of the
threespine stickleback (Gasterosteus aculeatus L.)in Lake Del1neye
based on the food consumption of plankton-feeding fishes.Problems of
Ichthyology.4:471-481.
Lewis,David B.,M.Walkey,and H.J.G.Dartna1l.1972.Some effects of
low oxygen tensions on the distribution of the three-spined
stickleback Gasteroseus acu1eatus L.and the nine-spined stickleback
Pungitius pungitius (L.).J.Fish.Biol.4:103-108.
Lindsey,C. C.1962.Experimental study of meristic variation in a
population of threespine sticklebacks.Gasterosteus aculeatus.Can.
J.Zool.40:271-312.
McInerney.J.E.•and O. O.Evans.1970.Action spectrum of the
photoperiod mechanism controlling sexual maturation in the threespine
stickleback.Gasterosteus aculeatus.J.Fish.Res.Bd.Canada 27:
749-763.
MacMahon,A.F.M.1946.Fish10re.London,Penguin Books.208 pp.
197D.Freshwater fishes of
Fish.Res.Bd.Can.Bu1.173.381 pp.
Manzer.J.I.1976.Distribution.food and feeding of the threespine
stickleback,Gasterosteus aculeatus.in Great Central Lake.Vancouver
Island,with comments on competition for food with juvenile sock~ye
salmon,Oncorhynchus nerka.Fish.Bull.74 (3):647-66B.
Moodie,G.E.E.1972.Morphology,life history,and ecology of an unusual
stickleback (Gasterosteus aculeatus)in the Queen Charlotte Islands.
Canada.Can.J.Zool.50:721-732.
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Salmon Studies:The study of red salmon in
Periodic Report No.4.U.S.Dept.of
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II
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II
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II
II
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n
Morrow.James E.1980.The freshwater fishes of Alaska.Alaska
Northwest Publishing Co .•Anchorage.248 pp.
Narver.David W.196B.Pelagial ecology and carrying capacity of sockeye
salmon in the Chignik Lakes,Alaska.Ph.D.Thesis,University of
Washington.Seattle.309 pp.
Parr,William H.t Jr.1912.Interactions between sockeye salmon and
lake resident fish in the Chignik Lakes.Alaska.H.S.Thesis.Univ.
Washington.Seattle.102 pp.
Potapova.T.L••T.V.Legedeva.and H.,.Shantunovkiy.1966.Dif-
ferences in the condition of females and eggs of the threespined
stickleback Gasterosteus aculeatus L.Prob.of Ichth.8:143-146.
Rogers,Donald F.1968.A comparison of the food of sockeye salmon fry
and threespine sticklebacks In the Wood River Lakes.Pages 1-43 in R.
L.Burgner (ed).Further studies of Alaska sockeye salmon.University
of Washington.Publications in Fisheries,Vol.III,Seattle.
Rogers.Donald E.1972.Abundance and size of juvenile sockeye salmo~,
Oncorhynchus nerka.and associated species in Lake Aleknagik,Alaska,
in relation to their environment.U.S.Fish and Wildlife Servo
Fishery Bulletin:71:(4).
Rpgers.D.E.1974.Alaska
the Nushagak District.
Comnerce._pp.
Scott,W.B.,and E.J.Crossman.1973.Freshwater fishes of Canada.
Fi'h.Res.Bd.Canada Bull.184.Ottaw •.966 pp.
Tinbergen,N.1952.The curious behavior of the stickleback.Scientific
American.121:2-6.
Vrat,Ved.1949.Reproductive behavior and development of the threespined
stickleback (Gasterosteus aculeatusl of California.Copeia.4:
252-260.
Wootton,R.J.1973a.The effect of size of food ration on egg production
in the female threesp1ned stickleback,Gasterosteus aculeatus l.J.
Fish Biol.5:B9-96.
Wootton,R.J.1973b.Fecundity of the three-spined sticklebacks,
Gasterosteus aculeatus (L.).J.Fish.Biol.5:683-688.
•Wootton,R.J.1976.The biology of the sticklebacks.Academic Press,
London.3B7 pp.
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