Environmental Biology of Fishes 63: 137–150, 2002.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Ontogenetic behavior and migration of Atlantic sturgeon,

Acipenser oxyrinchus oxyrinchus, and shortnose sturgeon,
A. brevirostrum, with notes on social behavior

Boyd Kynarda,b & Martin Horgana,c

a
U.S. Geological Survey, Biological Resources Division, S.O. Conte Anadromous Fish Research Center,
P.O. Box 796, Turners Falls, MA 01376, U.S.A. (e-mail: kynard@forwild.umass.edu)
b
Graduate Program in Organismic and Evolutionary Biology, University of Massachusetts, Amherst,
MA 01003, U.S.A.
c
Current address: Department of Zoology, Miami University, Oxford, OH 45056, U.S.A.

Received 4 July 2000 Accepted 25 June 2001

Key words: habitat preference, fish behavior, early life-history, adaptation

Synopsis

Ontogenetic behavior of Hudson River Atlantic sturgeon and Connecticut River shortnose sturgeon early life intervals
were similar during laboratory observations. After hatching, free embryos were photonegative and sought cover.
When embryos developed into larvae, fish left cover, were photopositive, and initiated downstream migration. Free
embryos may remain at the spawning site instead of migrating downstream because the risk of predation at spawning
sites is low. The two species are sympatric, but not closely related, so the similarities in innate behaviors suggest
common adaptations, not phylogenetic relationship. Atlantic sturgeon migrated downstream for 12 days (peak, first
6 days), shortnose sturgeon migrated for 3 days, and year-0 juveniles of both species did not resume downstream
migration. Short or long migrations of larvae may reflect different styles related to the total migratory distance from
spawning sites to juvenile rearing areas. Atlantic sturgeon need to move a short distance to reach rearing areas and
they had a long 1-step migration of 6–12 days. In contrast, shortnose sturgeon need to move a long distance to
reach all rearing areas. This may be accomplished by a 2-step migration, of which the brief migration of larvae is
only the first step. Early migrant Atlantic sturgeon were nocturnal, while late migrants were diurnal, and shortnose
sturgeon were diurnal. These diel differences may also be adaptations for long (Atlantic sturgeon) or short (shortnose
sturgeon) migrations. Cultured shortnose sturgeon, and possibly Atlantic sturgeon, have a dominance hierarchy with
large fish dominant when competing for limited foraging space. Social behavior may be more important in the life
history of wild sturgeons than is generally recognized.

Introduction populations of Atlantic sturgeon as near threatened

Atlantic sturgeon, Acipenser oxyrinchus oxyrinchus,
and shortnose sturgeon, A. brevirostrum, historically
coexisted in large Atlantic coast rivers of eastern North
America (Lee et al. 1980, Dadswell et al. 1984, Smith
1985, Smith & Clugston 1997). Due mainly to over-
fishing and river damming, one or both species has
been extirpated from many natal rivers. The World
Conservation Union Red List classifies Atlantic coast
and restoration options were recently reviewed by
Waldman & Wirgin (1998). Shortnose sturgeon is pro-
tected throughout its range (United States – federally
endangered, Canada – special concern; Kynard 1997).
Atlantic and shortnose sturgeon life histories dif-
fer significantly, particularly with regard to use of salt
water. Throughout their range, Atlantic sturgeon are
anadromous with juveniles rearing in their natal river
and estuary for several years before moving to sea
138
(Dovel & Berggren 1983, Smith 1985, Bain 1997).
Shortnose sturgeon in northern rivers of the Atlantic
coast, like the Hudson and Connecticut rivers, are
amphidromous, spending most of the year in fresh
water with brief visits during the foraging season to
salt water (Buckley & Kynard 1985, Kieffer & Kynard
1993). In the Connecticut and Hudson rivers, shortnose
sturgeon usually spawn in May (Bain 1997, Kynard
1997) and farther upriver than Atlantic sturgeon
(Galligan 1960, Bain 1997, Kynard 1997). The spawn-
ing area is the most upstream river reach used by either
species. Thus, sometime after hatching, the new gener-
ation of young sturgeon migrate downstream to river-
ine rearing areas and join older juveniles (Atlantic
sturgeon) or migrate downstream to permanent concen-
tration areas containing juveniles and adults (shortnose
sturgeon). Concentration areas of shortnose sturgeon
are occupied all year by mixed age individuals (Kynard
et al. 2000).
Life history of Atlantic sturgeon early life inter-
vals is poorly known (Smith 1985, Smith & Clugston
1997). Spawning occurs during June in the Hudson
River (Bain 1997). After hatching, free embryos (Balon
1999) and early larvae were captured by active bot-
tom netting within or just downstream of the esti-
mated spawning area (Bath et al. 1981), so early life
stages were benthic. Dovel & Berggren (1983) captured
year-0 and older juveniles in fresh water to within 5 km
of saline water (fresh water: salt water interface). Free
embryos, larvae, and year-0 juveniles of most sturgeon
species are not salt tolerant and even 5–10 ppt causes
mortality (white sturgeon, McEnroe & Cech 1985;
stellate sturgeon, Sbikin & Lapina 1981; and shortnose
sturgeon, Jenkins et al. 1993). This suggests another
disadvantage to early life intervals that migrate near
salt water. All evidence indicates that Atlantic sturgeon
migrants should stop before reaching the maximum
intrusion of salt water, which occurs at river km 98
(Dovel & Berggren 1983). Dettlaff et al. (1993) specu-
late that the poor tolerance of young sturgeon to salinity
may be a major factor dictating their migration style.
Recent laboratory studies revealed saltatory onto-
genetic behavioral changes occur when shortnose
sturgeon free embryos develop into feeding larvae. Free
embryos are photonegative and seek cover, whereas lar-
vae are photopositive and initiate downstream migra-
tion from a spawning area (Buckley & Kynard
1981, Richmond & Kynard 1995). Migration may be
brief because year-0 juveniles were captured 15 km
downstream from the spawning area in the Connecticut
River (Taubert 1980, Taubert & Dadswell 1980) and
25 km downstream from the suspected spawning area
in the Hudson River (Bath et al. 1981, Dovel1 ).
Social structure of wild sturgeons has not been
studied. For cultured sturgeon, the usual situation
is a great variability for body size (often, a two
to ten fold difference in size of large and small
individuals). For example: day-47 Siberian sturgeon,
A. baeri, weighed 0.3–3.2 g (Charlon & Bergot 1991);
day-42 Chinese sturgeon, A. sinensis, weighed 42–80 g
(P. Zhuang personal communication); month-4 amur
sturgeon, A. schrenckii, weighed 20–118 g (D. Li,
P. Zhuang, Z. Zhang, L. Zhang & T. Zhang unpublished
report); month-4 shortnose sturgeon weighed a mean of
5.5 g (range, 1.4–13.9 g; B. Kynard unpublished data);
and month-9 pallid sturgeon, Scaphirhynchus albus,
weighed a mean of 13.6 g (range, 7.7–18.9 g; B. Kynard
unpublished data). While much of this variability for
size is likely related to metabolic differences, could
social interactions also be involved? Metabolic differ-
ences in salmonids affect growth rate and body size,
and with agonistic behavior, establishes dominance of
large fish during foraging (Metcalf et al. 1995). Could
sturgeon, like salmon, of different body sizes differ
for their ability to obtain forage? This situation was
suggested by the behavior of year-0 cultured Russian
sturgeon, A. gueldenstaedti, where large individuals
were superior to small individuals competing for for-
aging space (Sibikin & Budayev 1991). We studied
cultured shortnose and Atlantic sturgeon juveniles for
the affect of body size on the ability of fish to obtain
an optimal foraging area.
We sought to confirm earlier observations on onto-
genetic behavior of shortnose sturgeon gathered by
Richmond & Kynard (1995), and additionally, we
investigated their swimming height above the bot-
tom, migration, and foraging related to body size. We
investigated Atlantic sturgeon for ontogenetic changes
in behavioral preference (illumination intensity, sub-
strate color, and cover) to compare with information
on shortnose sturgeon and we studied their migra-
tory and foraging behaviors. Free embryos (termed
embryos) and larvae were studied for ontogenetic
behavior and juveniles were studied for foraging.
All evidence indicates the ontogenetic behaviors are
innate, so the results give insight into behaviors of wild
sturgeon.
1
Dovel, W.L. 1981. The endangered shortnose sturgeon of the
Hudson Estuary: its life history and vulnerability to the activities
of man. Final Report, The Oceanic Society, Stamford. 139 pp.

Methods
Test fish and general procedures
On 5 July 1994, we transferred 500 Atlantic sturgeon
free embryos (5 days after hatching) of Hudson River
stock from the Northeast Fishery Center (U.S. Fish &
Wildlife Service) at Lamar, PA to the S.O. Conte
Anadromous Fish Research Center (Conte AFRC),
Turners Falls, MA, where studies were done. Most
Atlantic sturgeon were placed in a 120-l flow-through
circular tank. Fish were reared in Connecticut River
water at 21–24◦ C. Natural lighting and photoperiod for
the geographic location was used during all rearing.
Few embryos died and mortality of larvae was 5–10
per day, so that there were always several hundred fish
available for tests.
Fertilized eggs of Connecticut River shortnose
sturgeon were obtained in May 1995 (700 eggs from
two females fertilized by sperm from three males) and
May 1998 (1000 eggs from one female fertilized by
sperm from one male). Eggs were reared in 8-l hatch-
ing jars that overflowed and deposited swimming free
embryos into 18-l rearing tanks. All rearing and test-
ing of shortnose sturgeon was done in dechlorinated
city water of Montague, MA. City water was slightly
colder than ambient Connecticut River water in 1995,
but was heated to the temperature of ambient river water
in 1998. In 1995, 1000 embryos hatched during 2 days,
with 90% hatching on 18 May. Thus, 18 May was day 0
for most fish. In 1998, 19 May was day 0 for all test fish.
In 1994 and 1995, sturgeon were fed a diet of
brine shrimp, Artemia nauplii and commercial dry food
(BioKiowa), then after 60 days, a moist pellet (Biodiet,
Inc.). In 1998, fish were fed a sturgeon starter diet
(see Acknowledgements) and Artemia nauplii, then the
BioKiowa moist diet after 50 days. Commercial food
was dispensed by an automatic feeder at 2–4-h inter-
vals during larva and early juvenile rearing, and feed-
ing was reduced to 6-h intervals after 50 days. Artemia
nauplii were added five times per day for 3 weeks, then
gradually reduced to zero during a 7-day period.
In 1994 and 1995, we removed a few fish from the
rearing containers daily to visually determine the devel-
opmental stage. Using Richmond & Kynard (1995),
we identified the following developmental intervals
(1) free embryo – undeveloped eyes, mouth, and fin
folds, yolksac present, and (2) larva – yolksac absent,
fish feeding exogenously, fin rays and scutes present.
Ontogenetic observations ceased just prior to the juve-
nile interval, which begins when fish acquire all fin
139
rays (except caudal fin) at 47–58 mm standard length
(Snyder 1988). Throughout the paper, we refer to the
age of sturgeon by the day after hatching, i.e., when free
embryos became available for behavioral study. This
notation of age excludes the cleavage egg and embry-
onic development intervals (Balon 1999).
Illumination, substrate color, height
above bottom and cover
Preference of Atlantic sturgeon larvae for illumination
intensity and substrate color was tested in rectangular
19-l aquaria with 22-cm deep water using procedures of
Richmond & Kynard (1995). The illumination aquar-
ium was covered on four sides with black plastic and
a 40-wtt daylight fluorescent light was placed 0.5 m
above the aquarium. One-half the length of the top was
covered by a black plate creating 7–690 lx habitat on the
aquarium’s bottom. We divided the bottom area length-
wise into two equal-sized illumination areas: dark
(7–90 lx) and light (254–690 lx). For substrate tests, we
used an open top 19-l aquarium with the four vertical
sides covered by black plastic that created a uniform
690 lx illumination on the bottom. On the aquarium’s
underside, we placed two plastic squares (one black,
one white) each covering 50% of the bottom area.
Test procedures were similar for both illumina-
tion and substrate tests. Each day after fish hatched
(shortnose sturgeon) or developed into larvae on day 8
(Atlantic sturgeon), test fish were removed singly from
a rearing tank using an aspiration tube and introduced
at the water’s surface in the aquarium’s center. After
5 min, an individual was scored for location, then
removed with the aspiration tube, placed in a small con-
tainer, and after each replicate test was completed, all
fish were returned to the rearing tank. Each test day,
20 fish were observed in three replicates (daily total,
60 fish). A fish could be recaptured and retested on
another day, but they could not be retested the same day.
In both illumination and substrate tests, we reversed
the relative position of light and dark areas after each
replicate to avoid positional bias.
In 1998, shortnose sturgeon swimming height above
the bottom, swimming behavior, and preference for
bottom cover or open were examined in an artificial
stream tube that simulated a vertical section of river.
The tube was a clear plexiglass cylinder (15 cm inside
diameter × 153 m long with water 150 cm deep) that
was open at the top to introduce fish (Kynard et al.
2001a). A horizontal current (as would occur in
a stream) was created by a two-blade paddlewheel
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(total width, 6 cm) that turned clockwise in the cen- wide, with water 20 cm deep (Figure 1). A similar end-
ter of the tube. The paddlewheel blades extended from less channel was used to study migration of Atlantic
10 cm beneath the water’s surface to 8 cm above the salmon (Thorpe et al. 1988). Water inflow and out-
bottom. This enabled fish at the bottom or top to remain flow was about 1 l min−1 . A small submerged pump
there without disturbance from the turning of the pad- created a bottom current of 9 cm s−1 in the straight runs
dlewheel blades. A small electric motor mounted above and 5 cm s−1 in the curves (measured 3 cm off the bot-
the tube turned the paddlewheel creating a horizontal tom, in the center of the channel, in each of the 12
velocity of 2 cm s−1 . Depending on time of day, illu- sections). Five large rocks (diameter, 10–15 cm) were
mination (top to bottom) was 500–100 lx or 100–50 lx. placed across the channel at each curve to provide cover
A tan background placed outside the tube facilitated and to slow water velocity. Gravel substrate (cover for
seeing the small dark fish, which on hatching were only embryos) was removed after they left cover. The chan-
9–10 mm long. Tests using a black background, instead nel was surrounded by a black plastic curtain to prevent
of tan, showed that background color did not effect disturbance of fish.
vertical swimming behavior of fish (B. Kynard unpub- Fish moving in the oval channel were observed using
lished data). The tube’s length was marked at 5-cm an overhead video system. Infrared light to see fish at
intervals from bottom to water surface to plot move- night was provided by a 300-wtt light during Atlantic
ments of sturgeon. One-half of the bottom was covered sturgeon tests and by two infrared illuminators (Furman
with 5-cm diameter grey stones; the other one-half was Diversified, Inc.) during shortnose sturgeon tests. The
open (no stones). When fish were at the bottom of the bottom and side of the channel within camera view was
stream tube, we recorded the time spent in the open or
under rocks (in cover).
We tested eight shortnose sturgeon in the stream tube
each afternoon from days 0 to 15. Single sturgeon were
removed from a rearing container, introduced at the sur-
face by beaker, and observed for general behavior for
a 60-s acclimation period. Then, we recorded distance
above the bottom continuously for three 60-s periods
that began at 1, 5, and 10 min after introduction. Dis-
tance above the bottom was estimated to the nearest
1 cm and was recorded at every high and low point
as the sturgeon alternated swim-up bouts with drifting
pauses. During other types of swimming behavior, we
recorded fish location at 5-s intervals. We also recorded
the total time for fish to reach the bottom. The stream
tube was drained after each fish and refilled to provide
similar water temperature for all trials.
We also observed cover preference of both species
in an oval endless channel (described in Migration Sec-
tion). In 1994, 30 day-5 Atlantic sturgeon embryos
were placed over 30 cm2 of gravel-rubble in the oval
channel and observed several times daily for occur-
rence in cover or open. Similarly the following year,
we placed 50 day-15 shortnose sturgeon embryos over
gravel-rubble in the oval channel and observed them
daily for occurrence in cover or open.
Figure 1. Oval channel used to observe migration and compe-
Migration
tition for foraging sites of sturgeon. Twelve sections around the
channel’s periphery are numbered. The following notations des-
We observed migration (directed up- or downstream ignate channel features: flow direction = arrow, water inflow = I,
movements) in a green–blue, fiberglass, oval endless water outflow = O, submerged pump = P, feeder = F, and video
channel (Frigid Units, Inc.) that was 7.3 m long, 32 cm camera and infrared light = C.

coated with silver reflective tape to enhance observing
fish at night. Day-40 or younger sturgeon do not see
infrared light (Loew & Sillman 1993), so artificial illu-
mination should have no effect on the fish in tests.
In 1994, when the 30 Atlantic sturgeon emerged after
2 days from cover and began migration, we added 70
additional fish from rearing tanks to make a total of 100.
Migration was observed each day for 24 h (10 min per
hour) beginning on 7 July for 41 days (embryo-early
juvenile stages), and then biweekly until 29 October,
when water temperature had decreased to 10◦ C and
fish were becoming inactive. Dead larvae (0–5 daily)
were removed and replacement fish from the rearing
tanks were added to maintain 100 fish in the migration
channel.
In 1995, shortnose sturgeon migration was observed
beginning on 2 June when 50 day-15 free embryos
were placed into the oval tank and were observed using
video. Video observations were done daily to day 28,
and after this date, at about 2-week intervals until
27 October. Fish were observed for 10 min per hour
for 8 h on 3 June, 12 h on 4 June, and thereafter for
19–24 h daily. Observations in October were of a group
of 50 fish that were reared in the oval channel or rear-
ing tanks which were being tested in foraging com-
petition tests (see section on Foraging). For Atlantic
sturgeon after day 58, fish were visually observed daily,
and observed for 24 h with video on days 74, 90, 105,
118, 130, 137, 152,159, 164, and 170 until December.
For shortnose sturgeon after day 68, fish were similarly
observed on days 80, 85, 100, 115, 130, 145, and 162
until December.
Because the number of hours of video observations
was not the same for all days, we used the maxi-
mum number of fish passing downstream during any
1-h period to represent migration intensity for each
day. Diel activity was determined for each hour as the
number of fish moving downstream by the camera for
5 min during even hours (12:00, 14:00, 16:00 h, etc).
Orientation of fish (head up- or downstream) was noted
to indicate active swimming or drifting.
Foraging and body size
After observations on Atlantic sturgeon migration
ceased in October 994, large and small individuals were
observed in the oval channel for their location relative
to optimal feeding sites. A feeder dispensing food six
times per 24 h was located between sections 1 and 12
(Figure 1). Most food settled within 60 cm downstream
of the feeder; so we ranked section 1 as the most optimal
141
feeding area. Section 12, the nearest upstream section
to the feeder, was designated the second most optimal
area and sections 2–11 were designated less than opti-
mal sites. Two large and two small Atlantic sturgeon
were observed for 9 days (total, 25 time periods, 1–4
observations per day) from 2 November to 4 Decem-
ber 1994. Four times each day (morning, mid-day, and
early and late afternoon), we visually observed the sec-
tion occupied by each fish. We summed the totals for
large and small fish and plotted fish use by channel
section.
Foraging location of large and small shortnose
sturgeon was observed in the oval channel for 22 days
in October 1995. Fish in rearing containers and the
oval channel were measured, weighed, then some were
grouped into three non-overlapping size groups – small,
medium, and large to facilitate visual identification
of individual fish. Fish size in the three categories
were (category, n, total length (mean, range), and
weight (mean, range)): small – n = 17, 6.1 cm
(5.8–6.4 cm), 4.8 g (4.4–5.2 g); medium – n = 13,
8.3 cm (7.8–8.7 cm), 7.6 g (7.2–8.0 g); and large –
n = 22, 10.6 cm (9.9–11.1 cm), 9.6 g (8.9–10.1 g). We
placed 27 large and 23 small fish (50 total) in the oval
channel and visually counted the number of each group
in sections 1 and 12 and in sections 2–11 during the
morning, mid-day, and afternoon of each day.
We also conducted foraging tests with shortnose
sturgeon of large vs. small and medium vs. small indi-
viduals. We tested fish in two identical circular stream
channels that were 5.4 m long (outside wall) × 30 cm
wide with water 25 cm deep. Velocity in the center of
the 12 sections of the channel was 9–12 cm s−1 at 3 cm
above the bottom. Feeders that released food six times
per day were placed the same distance downstream
from the water inlets to create similar food distribu-
tion patterns. Most food settled to the bottom within
50 cm downstream of both feeders. Beginning at the
feeder, the outside channel wall was marked with large
numbers at 5-cm intervals to plot distance of each fish
downstream from the feeder.
In channel 1 we tested large and small fish and in
channel 2 we tested medium and small fish. After plac-
ing a pair of fish in each channel on day 0, thereafter,
each day for 22 days we added a fish to each channel
alternating between fish sizes (n = 23 fish per test). We
positioned a video camera directly over the center of
each circular channel to record the location of fish rel-
ative to the feeder. Sturgeon were observed for 10 min
per hour from 9:00 to 17:00 h for 21 days in November–
December 1995. Later, we reviewed the video tapes and
142
determined the distance (to the nearest 5 cm) from each
sturgeon’s snout to directly beneath the feeder.
Data analyses
We scaled sturgeon development to age and cumu-
lative temperature. Water temperature was recorded
daily and these data were used to calculate daily ther-
mal units and cumulative temperature units (CTU) in
degree-days after hatching. Cumulative degree-days
were calculated to mid-day of each day when behav-
ioral observations were made and reflected temperature
during the first 12 h of the current day and the sec-
ond 12 h of the previous day. For example, day-0 fish
accumulated 0 degree-days and day-1 fish accumulated
0.5×temperature ◦ C on day 0 plus 0.5×temperature ◦ C
of day 1.
We calculated the percentage of Atlantic sturgeon
using illuminated or white substrate each day and plot-
ted these as a time series. We also calculated binomial
95% confidence intervals for each day to see if they
included 50% (no preference). Linear regression was
used to determine if percent of fish preferring illumi-
nation or white substrate were related to sturgeon age.
For shortnose sturgeon in stream tube tests, we plot-
ted the time series of mean height above the bottom,
cover use, and time to reach bottom. Cover use was
only determined for fish that reached and stayed at
the bottom of the stream tube for at least one 60-s
period (usually three 60-s periods). Most fish spent
all of their bottom time under rocks. Fish that were
not exclusively using cover (out of rocks at least 10%
of their bottom time) were considered to be using the
open. Mean height above the bottom was only calcu-
lated for sturgeon after they reached the bottom of
the tube. We based this decision on previous work
(Richmond & Kynard 1995), which suggests that if the
shortnose sturgeon been introduced at the bottom of
the stream tube, they would have remained there while
in the embryo development interval. Therefore, height
information for sturgeon prior to reaching the bottom is
likely an artifact of the introduction method. For later
studies, the stream tube was modified to introduce fish
at the bottom (Kynard et al. 2001a).
Migration timing for both species was compared
using the number of fish passing during the peak hour of
each day considering all time periods. Using the daily
maxima moving downstream allowed direct compari-
son between the species even if they had diel differ-
ences in migratory behavior. We plotted the number of
fish passing as a time series. Within a species, diel dif-
ferences were tested using unpaired t-tests of day vs.
night observations for each day of sturgeon age.
For shortnose sturgeon foraging location in the oval
channel, we calculated the percentage of large and
small fish in each section each day. We combined
sections 2–11 and calculated 95% confidence intervals
for sections 1, 12, and the remainder during 22 days.
In the circular channel tests, the mean location of each
group of sturgeon was calculated for each day. A daily
statistic was calculated for each channel as the differ-
ence between mean location of small fish and mean
location of the larger size class fish. These differences
were plotted as a time series for both circular channels
and a paired t-test was used to compare differences for
the two channels.
Results
Illumination and substrate color
Atlantic sturgeon larvae to fish days 30–50 old pre-
ferred illumination and white substrate (Figure 2). Most
daily illumination confidence intervals were above
50%, while most white substrate binomial confidence
intervals included 50%. However, we feel that in both
cases the consistent results among days suggest that the
true values are above 50% and that a greater sample size
would have moved most binomial confidence intervals
above the 50% line. There was no major ontogenetic
shift in preference for illumination intensity or sub-
strate color as Atlantic sturgeon aged (linear regression
with age, both tests, >0.40).
Figure 2. Preference of Atlantic sturgeon larvae for illumination
intensity (illuminated, 254–690 lx or dark, 7–90 lx) and substrate
color (white or black). The horizontal line indicates 50% of fish
choosing the habitat.

Height above the bottom and cover preference
Day-0 free embryos of shortnose sturgeon in the
stream tube generally alternated swim-up with drift-
ing toward the bottom within about 2 min (Figure 3,
upper panel). Day-0 fish mostly swam into the cur-
rent (7 of 8 fish), whereas day-1 fish swam with the
current (6 of 6 fish; the other two fish showed no ten-
dency). This difference in number swimming with or
against the current was significant (chi-square, 0.001).
After day 1, there was no consistent trend among fish
for swimming with or against the current. Except for
day-1 fish, which took more than 7 min to reach the
bottom, fish to day 9 reached the bottom in 1–4 min.
No sturgeon that reached the bottom moved far above
the bottom until day 12 (perhaps, day 11 but data
are missing) when fish were larvae (Figure 3, upper
panel). After day 12, the mean distance that fish swam
above the bottom increased daily to about 1 m on
day 15.
Figure 3. Swimming height above the bottom and cover
preference of shortnose sturgeon free embryos and larvae in a
stream tube. Top panel shows the time to reach the bottom for fish
introduced at the water surface, and for fish that reach the bottom,
the subsequent mean swimming height (bars = maximum height)
above the bottom. Bottom panel shows the number of fish using
cover (rocks) or open. Days with less than eight fish indicate that
some fish did not reach the bottom. Data were not collected on
day 11.
143
Fish that reached the bottom used cover through
day 8 (Figure 3, lower panel). Some day-12 and older
fish used bottom cover, but each day more fish were
using open habitat and swimming above the bottom.
Cover and migration
Day-5 free embryos of Atlantic sturgeon placed over
gravel-rubble in the oval channel immediately sought
cover and remained there during days 6–7 (Figure 4).
Fish left cover and began migration on day 8 when
they emerged from the gravel and began swim-up and
drift behavior. The stomachs of several fish contained
brine shrimp, so foraging began with the habitat switch
when embryos developed into larvae. Migrant num-
bers were high until day 13, then declined 50% on
day 14, and about 10% each day to day 19 (12 days
total). Daily addition of five replacement fish did not
likely confound daily migrant counts because there
were 100 fish total. The CTU from days 0 to 8 (migra-
tion initiation) was 181 degree-days, 311 degree-days
to the last day when 100% were migrating (day 13),
and 465 degree-days to the last day of migration
(day 19).
All day-15 free embryos of shortnose sturgeon
placed over gravel in the oval channel sought cover
(Figure 4). When days-16–17 fish emerged from
cover, they were feeding larvae and used open habi-
tat. Although days-16–17 larvae emerged from cover,
visual and video observations indicated that none
Figure 4. Number of Atlantic sturgeon (1994) and shortnose
sturgeon (1995) moving downstream in the oval channel from
late free embryo to late-larva intervals. Daily migration intensity
was indicated by the highest (maximum) number of fish passing
each day during one 10-min period. Symbols are: triangle = visual
observation, circle = video observation. Lines connect consecu-
tive observations. Results of observations after day 68 are in text.
144

moved downstream on day 16 and a maximum of only

three fish passes were counted per hour on day 17.
Newly emerged larvae held position on the bottom, for-
aged, and some even moved short distances upstream.
Large numbers of days-18–19 larvae left the bottom
and actively swam headfirst downstream, alternating
swimming with bottom foraging. On day 20, most fish
ceased migrating, held position, and foraged. Shortnose
migration was short (2-day peak, 3-day total). The CTU
from day 0 (hatching) to day 18 (migration initiation)
was 248 degree-days and 279 degree-days to day 20
(migration end).
Shortnose and Atlantic sturgeon juveniles did not
resume downstream migration during observations
which continued until late October when water tem-
perature declined to 10◦ C. The increase in the num-
ber of day-28 and days-38–41 shortnose sturgeon
and days-20–25 Atlantic sturgeon moving downstream
(Figure 4) was not a second migration because peak
numbers of fish were not involved and many fish were
also moving upstream.
Initial swim-up and drift of Atlantic sturgeon larvae
was mostly passive, while downstream movement of
shortnose sturgeon larvae was entirely directed swim- Figure 5. Diel downstream movement of Atlantic and shortnose
ming. Based on the CTU values, Atlantic sturgeon sturgeon during migration of larvae. Symbols are: circle = mean
migrated at less CTU than shortnose sturgeon. Dur- number of fish, vertical bar = 95% confidence interval.
ing the first 1–2 days of Atlantic sturgeon migra-
tion, 60–75% of the larvae were facing upstream nocturnal (Figure 6). The hourly movement during
and drifting, but by the fourth day of migration the peak 2-day shortnose sturgeon migration show
(CTU = 243 degree-days), most were facing down- migrants were diurnal with day-18 fish most active in
stream, swimming actively like shortnose sturgeon the early morning (peak, 8:00 h) and day-19 fish active
with a similar degree-day level of development. Occa- later in the morning at 10:00 and 12:00 h.
sionally, migrants of both species swam upstream for
a short distance before continuing downstream. These
fish were always on or near the bottom and appeared to Foraging and fish size
be foraging. Post-migrants of both species were posi-
tively rheotactic and benthic. We observed large individuals of both species charging,
Atlantic sturgeon were nocturnal during the first pushing, and occasionally biting smaller fish. After a
one-half of the migration, then fish were active both day or so, it only took a charge or push, not overt aggres-
day and night (Figure 5). The number moving at sion, by large fish to illicit avoidance by smaller fish.
night was significantly higher on the first 6 days We did not quantify aggressive interactions between
(days 7–12; unpaired t-tests, <0.03); then, numbers sturgeon.
of night migrants were similar to day numbers on Large Atlantic sturgeon used all sections of the oval
day 13 or later (t-tests, >0.15). Shortnose sturgeon channel, except for section 5, more than small sturgeon,
migrants were most active in the day. Day move- but the use pattern of large and small fish was not sig-
ment was significantly higher on days 18–19 (t-tests, nificantly different (Figure 7). Section 5, where small
<0.005). After migration, night values were higher on sturgeon occurred at the highest frequency, was not an
days 20, 23, and 24 (t-tests, <0.02). optimal foraging site, but some food collected there at
Hourly movement during the peak 6 days of Atlantic the channel connection joint. The highest occurrence
sturgeon migration show fish were moving most of large sturgeon and the second highest occurrence
between 22:00–6:00 h with early migrants being most of small sturgeon was in section 1, the most optimal

Figure 6. Hourly downstream movement during peak migration
periods of Atlantic sturgeon (6 days) and shortnose sturgeon
(2 days).
Figure 7. Spatial distribution of two large and two small Atlantic
sturgeon juveniles in 12 sections of the oval channel during 9 days
(22 observations) in November–December 1994. Large fish occu-
pied significantly more sections (p = 0.005), but did not exclude
small fish from optimal feeding space (sections 1 and 12).
foraging section. Large fish occurred at high frequen-
cies in six additional sections, including the second
best feeding section, section 12. Large sturgeon used
11 channel sections more often than small sturgeon
145
Figure 8. Spacial distribution of 27 large and 23 small shortnose
sturgeon in the 12 oval channel sections. Data are based on 22 days
of observations in November 1995. Vertical bars are 95% confi-
dence intervals which show that a greater percentage of large fish
observations were in optimal feeding section 1, but no signifi-
cant difference between fish sizes in optimal feeding section 12.
A greater percentage of small fish observations were in sub-
optimal feeding sections 2–11.
(paired t-test, <0.005), so larger fish moved signifi-
cantly more than small fish.
Most large shortnose sturgeon in the oval channel
occurred in sections 1 and 12, the two sections near the
feeder (Figure 8). An examination of the 95% confi-
dence intervals in Figure 8 shows that a greater percent-
age of large fish occurred in section 1. The percentage
of small and large fish observed in section 12 was not
different, and a greater percentage of small fish than
large fish were recorded in sections 3–11.
In tests using the two circular channels, large
shortnose sturgeon were significantly closer to the
feeder than small fish, except on day 4 (Figure 9). Fish
on day 4 showed an aberrant pattern of unknown cause
in both channels. Neither medium nor small fish estab-
lished a consistent pattern of dominance near the feeder.
Mean difference in the occurrence between large and
small fish was significantly greater than the difference
between medium and small fish, whether the day-4 data
was included or not (19 or 20 days, <0.001).
Discussion
Atlantic sturgeon exhibited ontogenetic changes in
behavior, migration, and habitat preference similar
to shortnose sturgeon. Embryos sought cover (likely
a photonegative response) and photopositive larvae
dispersed downstream. This ontogenetic pattern was
exhibited by Connecticut River shortnose sturgeon
146
Figure 9. Distance from the feeder of small, medium, and large
shortnose sturgeon in two circular channels. Top panel shows
distance from the feeder of large and small fish, middle panel
shows distance from the feeder of medium and small fish, and the
lower panel shows the mean difference (cm) between large vs.
small and medium vs. small fish.
from three different sets of parents (Richmond &
Kynard 1995, present study) – a clear indication of the
innate nature of the behaviors.
Is the similarity of Atlantic and shortnose sturgeon
early behavior the result of close phylogenetic relation-
ship or common adaptations? Atlantic and shortnose
sturgeon are not closely related (Krieger et al. 1996,
Birstein et al. 1997, Birstein & DeSalle 1998, DiLauro
et al. 1999); however, early life intervals of the two
species are likely under similar selection pressures
because of their sympatry in coastal rivers. In inland
rivers, lake sturgeon, A. fulvescens, also migrate as lar-
vae (Kempinger 1988). Thus, migration of larvae is not
only an adaptation in coastal rivers, but has a broader
adaptive significance.
At least two downstream migratory patterns (embryo
and larva migrations) occur in North American
sturgeon during early life. Migration of free embryos
was suggested for white sturgeon, A. transmontanus
(E. Brannon et al.2 ) and was verified recently using
techniques in the present study (B. Kynard & E. Henyey
unpublished data). Additionally, both species of
the genus Scaphirhynchus begin migration as free
embryos, then continue as larvae (Kynard et al. 2001a).
The available data suggest that the adaptive signifi-
cance of migration initiation may be related to the pre-
dation risk of free embryos at a spawning site. If the
risk of predation is high, natural selection should favor
departure of embryos immediately after hatching; if
predation risk is low, selection should favor the later
departure of more developed larvae to enhance sur-
vival. If this conceptual model is correct, predation risk
of free embryos should be low for shortnose, Atlantic,
and lake sturgeon. Information that tests this hypothesis
is scarce, but preliminary data support the prediction of
low predation on shortnose sturgeon. In 1995, we used
trammel nets and gill nets to sample potential preda-
tors at the small Montague spawning area of Connecti-
cut River shortnose sturgeon. Predators were captured
during 132.5 net-h of effort (standardized to 50 m net
length) with an equal number of short 2–3 h net sets and
overnight net sets over a 19-day period. Only one preda-
tion event on sturgeon was documented: a 35 cm long
fallfish, Semotilus corporalis, had two sturgeon eggs
in its stomach along with other benthic prey. Over 40
individuals of other potential predators (native and non-
native species) were also captured and examined: rock
bass, Ambloplites rupestris, white sucker, Catostomus
commersoni, smallmouth bass, Micropterus dolomieu,
and channel catfish, Ictalurus punctatus. None of
these fish contained sturgeon progeny even though
food items in stomachs were overwhelmingly benthic
(e.g. mollusks, nymphs of Ephemeroptera, crustaceans,
and dipteran larvae). During drift net sampling (which
was concurrent with netting for predators), we caught
over 100 sturgeon eggs and embryos just down-
stream of the spawning site. This evidence sug-
gests that predation on shortnose sturgeon eggs and
embryos was low. Further, newly emerged shortnose
sturgeon larvae held position for a day or two prior to
migration, behavior that further suggests predation risk
is low.
2
Brannon, E., S. Brewer, A. Setter, M. Miller, F. Utter &
W. Hershberger. 1985. Columbia River white sturgeon (Acipenser
transmontanus) early life history and genetics study. Final Report
to Bonneville Power Administration on Project No. 83–316,
Portland. 68 pp.

Preference for illumination and white substrate by
larvae and early juveniles may be related to foraging.
Both preferences may enhance the ability of larvae
to visually distinguish moving prey from the back-
ground. Migrating larvae of shortnose and Atlantic
sturgeon alternated swimming in the water column
with bottom foraging where they held position or
even moved upstream, before resuming migration. This
movement pattern is typical of pause-travel preda-
tors (MacKenzie & Kiorboe 1995). Sturgeon vision
is not well developed, but fish have the ability to
differentiate moving objects that differ for illumina-
tion and contrast (Sbikin 1974, Loew & Silliman
1993, Silliman et al. 1999). Cultured juvenile sturgeon
locate food pellets or food scents using olfaction and
are not thought to use vision for foraging (Sbikin
1974, Kasumyan & Kazhlayev 1993, Kasumyan 1995).
However, sturgeon larvae in rivers forage on small,
mobile invertebrates like cladocerans and copepods
(Baranova & Miroshnichenko 1969, Miller et al.3 ).
During the day, an illuminated bright substrate may
enhance the visual ability of young sturgeon to see
moving prey; while at night, they likely use chemical
stimuli, electroreception, or both to locate prey. Fur-
ther, use of vision by larvae for feeding is suggested
by the sudden increase in the critical flicker frequency
response of sturgeon as they develop into larvae and
began to forage (Sbikin 1981). Thirty-day old shortnose
sturgeon juveniles observed chasing large zooplankton,
Daphnia spp., exhibited the entire predation sequence
of encounter, chase, attack, and capture when feeding
during the day in an aquarium (B. Kynard unpublished
data). Vision, not olfaction, should guide the swift pur-
suit of zooplankton that actively avoid capture.
Major changes in preference for illumination inten-
sity and substrate color did not occur with age of
Atlantic sturgeon to day 50 (Figure 2). Richmond &
Kynard (1995) also found no saltatory changes in pref-
erence for these factors with age of shortnose sturgeon
larvae and the preference for larger diameter substrate
by juveniles was likely related to changes in foraging.
3
Miller, A.I., P.J. Anders, M.J. Parsley, C.R. Spraque,
J.J. Warren & L.G. Beckman. 1991. Reproduction and early life
history characteristics of white sturgeons in the Columbia River
between Bonneville and McNary dams define habitat require-
ments for spawning and rearing of white sturgeon and quantify
the extent of habitat available in the Columbia River between
Bonneville and McNary dams. In: A.A. Nigro (ed.) Progress
Report to Bonneville Power Administration on Project No. 86-50,
Portland. 180 pp.
147
Micro- and macro-habitat selection of yearlings and
adults is not different (Kynard et al. 2000), so foraging
habitat preference may not change after the first year
of life.
The difference between the two species for diel
migration may be related to the length of migration,
foraging, and avoiding predation. Atlantic sturgeon lar-
vae are long-distance migrants and for the first one-half
of their migration, they moved mostly at night. This
behavior may reduce daytime predation risk while lar-
vae were still developing, yet still enable them to for-
age when there was daylight to aid visual detection of
prey. During the latter one-half of migration, the more
fully developed larvae migrated day and night (like
shortnose sturgeon). Long-distance migrant larvae of
stellate sturgeon, A. stellatus, in the Kuban River were
mostly captured at night (Pavlov et al. 1978), so noctur-
nal migration may be common. The daytime migration
of shortnose sturgeon suggests that the risk of predation
does not outweigh the advantages of migrating dur-
ing the day. The diurnal activity pattern of shortnose
sturgeon may be affected by prey activity patterns (par-
ticularly, vertical migration of zooplankton; Helfman
1986). When Connecticut River shortnose sturgeon lar-
vae migrate, daylight illumination extends to the river
bottom. During the day, zooplankton may be concen-
trated near the bottom with young sturgeon.
Shortnose sturgeon free embryos deprived of quick
access to cover by introduction at the water’s surface
of the stream tube sought bottom cover, as predicted
by the cover seeking behavior of embryos observed by
Richmond & Kynard (1995). Even embryos introduced
at the surface, an unnatural location for embryos in
the wild, proceeded downward and did not swim up
after reaching the bottom. However, days-13–14 fish
swam more in the water column and day-15 fish (larvae)
swam a maximum of 1 m above the bottom (Figure 3).
Based on the increased use of open habitat and the 170
degree day CTU of day-15 test fish, our observations
may have stopped just prior to the migration of larvae
(no concurrent information on migration is available).
However, wild migrant larvae may behave similarly to
the day-15 larvae because wild migrant larvae were
captured in 1-m diameter bottom-set nets, but not in
mid-water nets (B. Taubert unpublished data).
The short duration of shortnose sturgeon migra-
tion observed in the oval channel is supported by
field sampling of larvae in the Connecticut River
(B. Taubert unpublished data). Extensive netting for
embryos and larvae in 1977 and 1978 captured embryos
a short distance downstream from the spawning site,
148
i.e., maximum, 12 km for embryos and 15 km for larvae
(migration distance back-calculated after the spawning
site was located in 1993 by M. Kieffer & B. Kynard
(unpublished data). With a peak migration of 2 days,
most larvae in 1977–1978 moved 7.5 km per day. It
is interesting that some embryos moved almost as far
as larvae. Thus, if only field data on capture of embryos
and larvae were available, investigators could not know
if embryos, larvae, or both were the migratory interval.
The difference between shortnose and Atlantic
sturgeon for migration duration may reflect different
migration styles related to migration distance. Atlantic
sturgeon spawn at river km 136–182 (Van Eenennaam
et al. 1996), so larvae need only to migrate a max-
imum of 38–84 km to reach the rearing area just
upstream of saline water at river km 98. If the migra-
tion rate of Atlantic sturgeon is similar to the 7.5 km per
day rate of shortnose sturgeon, then Atlantic sturgeon
larvae can easily make this movement with a long
1-step migration of 6–12 days. However, Connecticut
River shortnose sturgeon spawn at river km 192–194
(Kynard 1997), and larvae would need to move
about 150 km to reach all downstream rearing areas
(Buckley & Kynard 1985, Kynard 1997). If migration
was 7.5 km per day, then the migration of larvae would
last 20 days. The disadvantages of a long migration of
larvae (high mortality from starvation and predation)
seem greater than the advantages of quickly reaching
the final juvenile rearing areas. No second migration
by year-0 fish of either species was detected in the
oval channel as of early winter and no second migra-
tion likely occurs during winter or early spring due to
energetic factors. Field evidence indicates the down-
stream distribution of young shortnose sturgeon is com-
pleted by yearlings (Hudson River, Dovel1 ; Southern
rivers, Kynard 1997) or in the case of the Connecticut
River, by yearlings and older life intervals (Kynard
1997). Experiments using cultured yearling shortnose
sturgeon that migrated downstream in summer and fall
support field observations for a 2-step migration, i.e.,
an initial migration of larvae and a second migration by
yearlings (B. Kynard & E. Henyey unpublished data).
During competition for foraging space, large
shortnose sturgeon dominated small individuals during
two separate experiments and tests with medium and
small sturgeon indicated absolute size (large), not rel-
ative size was most important. Large Atlantic sturgeon
were significantly different from small individuals for
activity, and large individuals occurred more often in
the optimal foraging area than small sturgeon, but
the difference was not significant. Because density
of Atlantic sturgeon in tests was low and food was
abundant, the conditions may not have been suffi-
ciently competitive to demonstrate dominance of large
fish. Low fish density and high food abundance affect
expression of agonistic behavior and expression of
dominance (Noakes & Grant 1986, Mikheyev 1995).
Salmonids with higher metabolic rates have a higher
energy level, grow faster, and also may have a higher
level of activity (Metcalf et al. 1995). Possibly, this
was responsible for the high level of activity of our
large Atlantic sturgeon.
Our experiments and those of Sbikin & Budayev
(1990) indicate that cultured sturgeon have a domi-
nance hierarchy when competing for limited foraging
space. Based on our preliminary observations of fish
interaction, the hierarchy seems mediated by agonistic
behavior. Does this information on cultured sturgeon
have any significance for understanding wild sturgeon?
Sbikin & Budayev (1991) concluded the dominance
behavior was related to the artificial conditions (partic-
ularly, fish density) and had no significance for wild
populations. However, this may not be the correct
interpretation. While the intensity of agonistic behav-
ior or expression of other innate behaviors may be
altered (increased or decreased) by artificial conditions
(Chiszar et al. 1975, Chebanov 1990, Mikhayev 1995),
we do not know of any example where agonistic behav-
ior or a dominance hierarchy was present in cultured
individuals of a species, but was absent or had no sig-
nificance for the wild population. Also, while data are
not extensive, young wild sturgeon vary for body size
like cultured sturgeon. Information on wild years 0–1
sturgeon of several species indicates there is consider-
able variability for body size, e.g., months 2–3 stellate
sturgeon, A. stellatus, weighed 0.2–0.7 g (Amirkhanov
1968); month-14 white sturgeon, A. transmontanus,
were 41–180 mm long (Miller et al.3 ), and day-114
lake sturgeon, A. fulvescens, were 160–240 mm long
(Kempinger4 ). Shortnose sturgeon live their entire lives
sharing several discrete foraging concentration areas,
a life history that could favor the evolution of forag-
ing competition that would limit population density
through emigration (Kynard 1997).
Studies with wild sturgeon are needed to exam-
ine dominance hierarchy in the absence of artificial
fish densities and conditions. Young wild sturgeon are
4
Kempinger, J.J. 1983. Early life history of lake sturgeon.
Progress Report – Study No. 227, Wisconsin Department of
Natural Resources, Madison. 33 pp.

difficult to observe, but there are small clear rivers
with large numbers of young sturgeon that could be
appropriate for study. Recent observations on several
species from two genera provide additional evidence
that sturgeon may be more social than commonly
believed, e.g., Kynard et al. (2001a) gathered evidence
that young Scaphirhynchus spp. use their black tails
as a visual signal for aggregation, and pre-spawning
males identified pre-spawning females using a chem-
ical pheromone (Kasumyan 1993, Kynard & Horgan
2001b). Sturgeon are one of the most primitive teleost
fishes (Bemis et al. 1997), but they may have a complex
of social behaviors usually attributed to less primitive
fishes.
Acknowledgements
Shortnose sturgeon were studied under National
Marine Fisheries Service Scientific Permit No. 726
and 944 to B. Kynard. We thank the Northeast Fishery
Technology Center (US Fish & Wildlife Service) for
providing Atlantic sturgeon embryos. Technical assis-
tance during data collection and fish maintenance
was provided by C. Cauthron, P. Kwiatrowski, and
J. Messier. E. Henyey scored larvae for diel activity.
The Conte AFRC provided funding.
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