Environmental Biology of Fishes 70: 145–154, 2004.

© 2004 Kluwer Academic Publishers. Printed in the Netherlands.

Effects of temperature on the early development, growth, and

survival of shortnose sturgeon, Acipenser brevirostrum, and
Atlantic sturgeon, Acipenser oxyrhynchus, yolk-sac larvae

Ryan S. Hardya & Matthew K. Litvakb,∗

a
Idaho Department of Fish and Game, 2750 Kathleen Avenue, Coeurd’Alene, ID 83815, U.S.A.
(e-mail: rhardy@idfg.state.id.us)
b
Department of Biology and Center for Coastal Studies and Aquaculture, University of
New Brunswick, P.O. Box 5050, Saint John, N.B., Canada, E2L 4L5
∗
Corresponding author (Tel.: 506-648-5565; Fax: 506-648-5811; e-mail: litvak@unbsj.ca)

Received 11 February 2002 Accepted 2 September 2003

Key words: escape response, early life history, yolk utilization efficiency, yolk utilization rate

Synopsis

We reared shortnose and Atlantic sturgeons at different temperatures after hatch and measured yolk utilization rate
and efficiency (YUE), maximum standard length, survival and development of escape response. Newly hatched
Atlantic sturgeon, were smaller in size, more efficient at utilizing yolk (incorporating yolk to body tissue) and
reached developmental stages sooner than shortnose sturgeon reared at the same temperatures (13–15◦ C). Within
each species, decreasing temperature delayed yolk absorption, escape initiation, time to reach maximum size, and
time to 100% mortality. However, YUEs and the size of the larvae at these ‘stages’ were independent of rearing
temperature for both species. These results suggest that even as temperature drives metabolic processes to speed up
development, these two species are still extremely efficient at transferring yolk energy to body tissues. The lower
efficiencies experienced by larval shortnose may reflect difference in yolk quality between the two species and/or the
Atlantic sturgeon’s higher conversion efficiency. The ability of these two sturgeon species to develop successfully
and efficiently under a wide range in temperatures may provide a competitive advantage over more stenothermic
species and explain their persistence through evolutionary time.

Introduction of environmental factors that affect the rate of yolk

Mortality in many fish species is very high during early
development due to their small size, limited swimming
ability, and sensitivity to variations in their surround-
ing environment (Rice et al. 1987, Miller et al. 1988).
During these vulnerable stages, fishes may be adversely
affected by environmental factors that influence sur-
vival and subsequent year-class strength (Houde 1994,
Claramunt & Wahl 2000).
Size of a fish at the time of the switch from
endogenous to exogenous feeding, referred to as the
‘critical period’ in development, may be a very impor-
tant factor affecting year-class strength of a species
(Hjort 1914, Cushing 1972). Therefore, examination
utilization, efficiency to which yolk energy is trans-
ferred to body tissues, and body size at each stage
of development is important to our understanding of
recruitment.
Water temperature is considered to be the most
important environmental factor influencing fish devel-
opment (Chambers & Leggett 1987). Limited informa-
tion exists on how water temperature affects the early
development of sturgeon. The few studies performed
on sturgeon (Wang et al. 1985, 1987, Gershanovich &
Taufik 1992) included larvae of different sizes at the
start of the experiment. Since eggs of many sturgeon
species are spawned within a narrow range of tem-
peratures and have a relatively short incubation period
146
(<10 days for many species), they may experience lit-
tle variation in temperature. However, after hatching,
larval sturgeon (with a stage duration of 40–60 days
in some species; Richmond & Kynard 1995) will
undoubtedly experience a wider range of temperature
variation than during the incubation period. Therefore,
it is of interest to determine the effect of temperature on
developing sturgeon larvae.
Two North American species, the shortnose stur-
geon, Acipenser brevirostrum, and Atlantic sturgeon,
Acipenser oxyrhynchus, have suffered recent declines
primarily due to anthropogenic factors such as con-
struction of dams, water pollution, and over-fishing
(Smith & Dingley 1984). Presently, no data are avail-
able on how environmental factors influence mortality
during the early life history (ELH) stages of these
two species (Kynard 1997, Smith & Dingley 1984).
Since mortality may be insignificant in the late juvenile
through adult stages, it is likely that survival through
the ELH stages of larval sturgeon is more important
in determining year-class strength (Gross et al. 2002).
The objective of this paper was to determine the effects
of temperature on growth and development, with spe-
cific attention to yolk utilization efficiency (YUE) of
these two species. We will discuss how these effects
may influence predation avoidance ability and larval
survival.
Materials and methods
Egg collection and incubation
We collected mature shortnose sturgeon between 1
and 15 May 1998, with short-set (<6 h) gill nets
in the Kennebecasis River, New Brunswick, Canada
(N45◦ 30 W66◦ 55 ; water temperature: 13–16◦ C). We
collected mature Atlantic sturgeon in a similar fashion
between 1 and 7 July 1998 in the Saint John River,
New Brunswick, Canada (N45◦ 33 W66◦ 02 ; water
temperature: 16–18◦ C). We injected one female and
two male shortnose sturgeon with hormone (LHRH-A)
on 17 May 1998. We fertilized the eggs with the
sperm from the two males on 18 May using procedures
specified by Doroshov et al. (1983). We followed this
procedure on 8 July 1998, to produce fertilized Atlantic
sturgeon eggs. We placed ∼500 ml of fertilized eggs
from each species into each of three MacDonald incu-
bation jars in a partial recirculation system. We set
flows to the jars at 3 l min−1 to allow adequate oxy-
genation of the fertilized eggs. Eggs were incubated
at a mean temperature of 17◦ C, which is similar to
or within the range of natural spawning temperatures
for both species (Huff 1975, Wooley & Crateau 1985,
for additional reviews see Kynard 1997).
Egg size considerations
It is possible that different sized females as well as
seasonal variations such as age, weight, and health can
all play a role in egg size, yolk quality, and subse-
quent larval sizes. In addition, it is true that egg size
can play a role in survival (Chambers et al. 1989).
To make certain that the eggs from the single female
used in this experiment were representative in size to
that of other females, egg sizes from four independent
shortnose sturgeon crosses and from two spawning sea-
sons (two crosses per season) were compared using
a two-way ANOVA. The resulting ANOVA showed
no significant difference in mean egg diameter within
or between years of shortnose crosses (P = 0.134;
3.5 mm ± 0.088 SE following hydration).
Experimental design
We set-up four treatment tanks (∼1 m × 3 m × 40 mm
water baths) which were maintained at 13◦ C, 15◦ C,
18◦ C, and 21◦ C. Temperatures were controlled by sub-
mersible aquarium heaters (18◦ C and 21◦ C), ambient
room temperatures (15◦ C), and with a water-chilling
unit (13◦ C). We chose temperature treatments based
on literature (Huff 1975, Wooley & Crateau 1985, for
additional review see Kynard 1997) and field data.
A fluorescent light over each tank provided a light
intensity of ≈700 lux at the water surface (Lutron
LX-101 lux metre; intensity was chosen based on
Richmond & Kynard 1995). We placed an opaque black
plastic sheet over the treatment tanks to ensure consis-
tent lighting. Photoperiod for the experimental tanks
was set at 15L : 9D, which is similar to natural spawning
conditions. Each of the four treatment tanks contained
three or four replicate trays for each species (1 000 ml
glass trays filled with 700 ml of de-chlorinated water).
Upon hatch, (shortnose: 192 h; Atlantic: 120 h), we
placed 40 larvae into each tray. Although there is much
debate over when a sturgeon is considered an embryo,
larvae, or juvenile, for the purposes of this study, we
considered hatched sturgeon embryos to be ‘larvae’ for
the duration of the experiment (until complete yolk uti-
lization). We replaced water in each of the replicate
trays (600 ml) twice daily with fresh de-chlorinated
water. We tested the replacement water daily to ensure

that dissolved oxygen, salinity, and pH levels were ade-
quate and similar to natural conditions (7.0–8.0 mg l−1 ,
0 ppm, 6.2, respectively).
Data collection and analysis
We sampled young fish (3–4) from each treatment
group, (one per replicate tray) every other day, starting
at hatch. Each fish was anesthetized in a 25 mg l−1 solu-
tion of tricaine methanesulfonate (MS-222), placed
under a microscope (Olympus SZ6045), and video-
taped (Sony DXC-1821 videocamera and S-VHS
Panasonic AG-5700 recorder) for later analysis with
an image analysis system (Optimas v5.2 BioScan Inc.,
Edmonds, Washington). After we collected morpho-
logical data, we preserved larvae in 10% phosphate-
buffered formalin. The data from each fish at each
temperature included: yolk-sac length (YSL), height
(YSH), and area (YSA), body area (BA), and stan-
dard length (SL). Elliptical yolk-sac volume (YSV)
(mm3 ) was calculated using the formula of a spheroid:
YSV = (π /6)LH2 where L is the length of the yolk-
sac and H is height (Blaxter & Hempel 1963). YSV
also included lipid volume due to difficulties in distin-
guishing between the yolk and lipid globule with image
analysis. We measured SL (mm) from the anterior-most
point of the developing rostrum to the posterior-most
point of the notochord. We measured BA (mm2 ) by dig-
itally drawing a line around the entire larva excluding
the yolk-sac and finfold areas.
In addition to morphological analysis, we examined
development of escape response, as well as percent
mortality over time for each temperature treatment.
Sturgeons, like many other fish species, exhibit a
‘C-type’ startle response (a rapid movement of the body
trunk in the shape of a ‘C’; Blaxter & Batty 1985) to
escape predation. We tapped each treatment tank daily
to stimulate this response through mechanical vibra-
tion. We recorded the date of the first occurrence of a
startle response. The time to 100% mortality was deter-
mined for each treatment replicate over time (excluding
loss through sampling).
We determined yolk utilization rate (YUR) for both
species by calculating slopes from a regression anal-
ysis (Proc REG. SAS Institute Inc. 1992) of YSV
for each temperature treatment over the duration of
the experiment. We determined YUE for each species
by comparing the rate of BA growth (slope of BA
against age regression: mBA) to the rate of yolk uti-
lized (absolute value of slope of YSA against age
regression: |mYSA|) until the complete absorption of
147
the yolk (YUE = mBA/|mYSA|). Growth rates to a
maximum SL on yolk reserves were determined by cal-
culating slopes from a regression analysis (Proc REG.
SAS Institute Inc. 1992) for each temperature treat-
ment to the point where maximum SL was attained.
We used two-way ANOVAs to determine the effect of
temperature treatment and species on time to yolk-sac
absorption, YUR, YUE, larval growth, size at escape
response, size at yolk-sac absorption, and larval sur-
vival (Proc GLM, general linear models; SAS Institute
1992). We separated the data into one-way ANOVAs
to reveal significance when an interaction (P < 0.2;
see Winer 1971 (page 379) for justification of a con-
servative type I error rate of 0.2) between species and
temperature was detected. If the model showed no inter-
action, the term was dropped and the model rerun.
Least-square means were used for a posteriori com-
parisons, and probabilities were adjusted for multiple
comparisons using Tukey’s correction (SAS Institute
Inc. 1992). Level of significance for main effects and a
posteriori comparisons were set at an α of 0.05.
We tested all data for normality (Proc Univariate;
SAS Institute 1992) and homogeneity of variances
(Fmax -test, Sokal & Rohlf 1981). Analyses were run
on log10 or arcsine-square-root transformed data in
cases of non-normality or heterogeneity of variance.
We used a non-parametric Wilcoxon rank test to exam-
ine the effect of temperature on the time to first escape
response due to lack of variance within treatments.
Lastly, measurements (YSV, YSA, SL, and BA) of the
larvae at hatch were also compared between tempera-
tures with one-way ANOVAs to confirm no significant
differences within each species at the start of the
experiment.
Results
Morphological comparisons at hatch
Diameter of shortnose sturgeon eggs (3.5 mm ± 0.088
standard error (SE)) after fertilization and hydra-
tion were significantly larger than Atlantic sturgeon
eggs (2.2 mm ± 0.115 SE) (df = 1, 4; P =
0.001). At hatch (the start of the experiment), the
YSA and YSV within each species did not differ
between temperature treatments. Shortnose sturgeon
larvae, however, possessed larger (all temperatures =
df = 1, 5; P < 0.005) YSA (mean 5.13 mm2 ±
0.159 SE) and YSV (mean 5.20 mm3 ± 0.279 SE) than
Atlantic sturgeon larvae (mean 4.06 mm2 ± 0.076 SE;
148
mean 4.17 mm3 ± 0.141 SE, respectively). Similarly,
the SL and BA within a species at hatch did not differ
between temperature treatments. Shortnose sturgeon
larvae were significantly longer (mean 10.73 mm ±
0.287 SE; all P < 0.002) and had greater BA
(mean 15.76 mm2 ± 0.191 SE) than Atlantic sturgeon
larvae (mean 9.81 mm ± 0.086 SE; mean 10.41 mm2 ±
0.206 SE, respectively). Since the measurements BA
and SL were affected in the same way throughout
the results of the experiment, only SL was used as
growth parameter and BA was only used in calculation
of YUE.
Time to yolk-sac absorption
The resulting two-way ANOVA on the time to complete
yolk-sac absorption for both shortnose sturgeon and
Atlantic sturgeon was significant (df = 7, 19; P =
0.0001) and exhibited an interaction between species
and rearing temperature (df = 3, 19; P = 0.0061).
With increasing temperature (13–21◦ C), resulting one-
way ANOVAs for both species showed a significant
reduction in the time to yolk-sac absorption (Atlantic:
df = 3, 8; shortnose: df = 3, 11; P = 0.0001; Figure 1).
Shortnose sturgeon took longer (df = 1, 5; all P >
0.05) to absorb the yolk-sac than Atlantic at 13◦ C, 15◦ C,
and 18◦ C. However, at 21◦ C there was no significant
difference between species.
Figure 1. Mean time (days) to yolk-sac absorption for shortnose
sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus,
at each rearing temperature. Values with different letters are sig-
nificantly different (lowercase = Atlantic sturgeon). Significance
was set at an α of 0.05. Horizontal bars with (∗ ) over treat-
ments indicate significant difference between species. Error bars
represent ±1 SE.
Standard length at yolk absorption
No effect of temperature was detected on the SL at
the time of yolk-sac absorption by species. However,
resulting one-way ANOVAs between species revealed
that shortnose sturgeon had a significantly greater SL
than Atlantic sturgeon larvae at each rearing tempera-
ture tested (Table 1).
Yolk utilization rate
ANOVAs were run on log10 transformed slopes to
meet assumption of homogeneity of variance. The
resulting two-way ANOVA model on the yolk-sac
utilization rate (YUR) (slopes of YSV) for both short-
nose sturgeon and Atlantic sturgeon was significant
(df = 7, 19; P = 0.0001) and showed an interaction
between species and rearing temperature (df = 3, 19;
P = 0.0010). One-way ANOVAs revealed that a reduc-
tion in temperature significantly decreased the rate of
yolk volume utilized over the experimental period for
both species (Table 2). Shortnose sturgeon larvae uti-
lized yolk volume at a faster rate than Atlantic sturgeon
larvae at lower rearing temperatures (13◦ C and 15◦ C;
Table 2).
Yolk utilization efficiency
The two-way ANOVA on YUE for both shortnose
and Atlantic sturgeon was significant (df = 7, 19;
P = 0.0020) and exhibited an interaction between
species and rearing temperature (df = 3, 19; P = 0.1841).
Subsequent one-way ANOVAs showed that within
each species YUE did not differ among tempera-
ture treatments. However, Atlantic sturgeon larvae
were more efficient than shortnose sturgeon at 13◦ C
(df = 1, 5; P = 0.0029) and 15◦ C (df = 1, 5; P = 0.0068)
(close at 18◦ C: P = 0.0770) in the amount of body tissue
incorporated for the amount of YSA utilized (Figure 2).
Escape response
Time to initial escape response, for both species (df =
7, 19; P = 0.0001), was significantly delayed at the
lowest rearing temperatures (13◦ C and 15◦ C; Figure 3).
Atlantic sturgeon larvae exhibited the initiation of the
C-type escape response significantly earlier than short-
nose sturgeon, at all temperature treatments (df = 1, 5;
P = 0.0001). SL of both species at initiation of escape
response did not differ among rearing temperatures.
 149

Table 1. Standard length at yolk absorption and initial escape response, and maximum SL attained for larval shortnose
sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus, at each temperature (13◦ C, 15◦ C, 18◦ C, and 21◦ C).

Stage measured Species Temp (◦ C) Mean SL (mm) ±1 SE Significance

∗
At yolk absorption Shortnose 21 17.75A 0.68 (0.04)
∗
18 17.82A 0.45 (0.02)
∗
15 17.46A 0.29 (0.00)
∗
13 18.55A 0.26 (0.00)
Atlantic 21 15.40a 0.45
18 15.15a 0.71
15 14.44a 0.17
13 14.46a 0.11
∗
At first escape response Shortnose 21 16.90A 0.13 (0.00)
∗
18 16.52A 0.13 (0.00)
∗
15 16.02A 0.30 (0.00)
∗
13 16.00A 0.52 (0.01)
Atlantic 21 13.54a 0.27
18 13.23a 0.29
15 13.60a 0.15
13 13.40a 0.23
∗
Maximum size attained Shortnose 21 18.56A 0.27 (0.00)
∗
during experiment 18 18.35A 0.03 (0.00)
∗
15 18.71A 0.42 (0.00)
∗
13 18.55A 0.26 (0.00)
Atlantic 21 15.58a 0.26
18 15.63a 0.43
15 15.01a 0.01
13 14.99a 0.42
Significance (P < 0.05) between temperatures (within each species) is indicated by different superscript letters (lowercase =
Atlantic; uppercase = shortnose). Significance between species is indicated by an (∗ ) and P value. SE = ±1 SE.

Table 2. Results from regression analysis on the effect of temperature (13◦ C, 15◦ C, 18◦ C, and 21◦ C) and species
(shortnose sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus) on YSV absorption rate over the
experimental period. Mean values represent replicate trays within each treatment (n = 3–4).

Species Temp Mean slope ±1 SE Mean r2 Model P’s Significance

(◦ C) (mm3 day−1 ) between spp.

Shortnose 21 −0.51A 0.02 0.91 <0.01 0.06

18 −0.31B 0.03 0.81 <0.00 0.16
15 −0.27C 0.01 0.83 =0.00 ∗
0.00
13 −0.18C 0.00 0.77 =0.00 ∗
0.00
Atlantic 21 −0.68a 0.04 0.91 <0.04
18 −0.41b 0.13 0.84 <0.01
15 −0.13bc 0.00 0.60 <0.00
13 −0.11c 0.02 0.59 <0.00
Significance between temperatures (within each species) is indicated by different superscript letters (lowercase =
Atlantic; uppercase = shortnose). The species, reared at the same temperature, which is significantly greater in
value is indicated by an (∗ ) and P value. Significance was set at an α of 0.05. SE = ±1 SE.

However, shortnose sturgeon larvae were significantly Maximum SL on yolk reserves

larger in SL at the initiation of escape response than
Atlantic sturgeon larvae at each rearing temperature The two-way ANOVAs on the time to reach a maximum
tested (Table 1). SL on yolk reserves, in both species, were significant
150
Figure 2. Mean YUE for shortnose sturgeon, A. brevirostrum,
and Atlantic sturgeon, A. oxyrhynchus, at each rearing tem-
perature. Values with different letters are significantly different
(lowercase = Atlantic sturgeon). Significance was set at an α of
0.05. Horizontal bars with (∗ ) over treatments indicate significant
difference between species. Error bars represent ±1 SE.
Figure 3. Mean time (days) to initiate first escape response
for shortnose sturgeon, A. brevirostrum, and Atlantic sturgeon,
A. oxyrhynchus, at each rearing temperature. Values with different
letters are significantly different (lowercase = Atlantic sturgeon).
Significance was set at an α of 0.05. Horizontal bars with (∗ ) over
treatments indicate significant difference between species.
(df = 7, 19; P = 0.0001) and showed an interaction
between species and rearing temperature (df = 3, 19;
P = 0.1979). Subsequent one-way ANOVAs revealed
that for both species, time to reach a maximum length
was significantly delayed with a reduction in tem-
perature (Atlantic: df = 3, 8; shortnose: df = 3, 11;
P = 0.0001; Figure 4). A species comparison revealed
that Atlantic sturgeon larvae reached a maximum SL
Figure 4. Mean time (days) to reach a maximum SL for shortnose
sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus,
at each rearing temperature. Values with different letters are sig-
nificantly different (lowercase = Atlantic sturgeon). Significance
was set at an α of 0.05. Horizontal bars with (∗ ) over treat-
ments indicate significant difference between species. Error bars
represent ±1 SE.
sooner than shortnose sturgeon larvae at all tempera-
tures (df = 1, 4; all P < 0.0457).
In both species, no effect of temperature was
detected on the maximum SL attained over the exper-
imental period. However, resulting one-way ANOVAs
revealed that shortnose sturgeon did attain a signif-
icantly greater SL than Atlantic sturgeon larvae at
each rearing temperature tested (Table 1). The two-
way ANOVA on growth rate to maximum SL (slopes
of SL) for both shortnose sturgeon and Atlantic stur-
geon larvae was significant (df = 7, 19; P = 0.0001)
and showed an interaction between species and rear-
ing temperature (df = 3, 19; P = 0.0929). Resulting
one-way ANOVAs revealed that a reduction in tem-
perature significantly decreased the rate of growth to
maximum SL on yolk reserves over the experimen-
tal period for both species (Table 3). Atlantic sturgeon
larvae grew at a faster rate than shortnose at 18◦ C.
Survival
The two-way ANOVA comparing the time to 100%
mortality between species was significant (df = 7, 19;
P = 0.0001) and showed an interaction between
species and temperature (df = 3, 19; P = 0.0128).
Subsequent one-way ANOVAs revealed that both
species of sturgeon survived longer at lower rearing
temperatures, with mortality increasing rapidly after
 151

Table 3. Results from regression analysis on the effect of temperature on growth rate to maximum SL for both
shortnose sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus. Mean values represent replicate
trays within each treatment (n = 3–4).

Species Temp Mean slope ±1 SE Mean r2 Model P’s Significance

(◦ C) (mm3 day−1 ) between spp.

Shortnose 21 0.57A 0.06 0.89 <0.00 0.19

∗
18 0.33B 0.02 0.85 <0.00 0.03
15 0.23C 0.00 0.84 0.00 0.20
13 0.19C 0.01 0.87 0.00 0.72
Atlantic 21 0.68a 0.02 0.97 <0.00
18 0.44b 0.03 0.93 <0.00
15 0.20c 0.02 0.88 0.00
13 0.20c 0.00 0.90 0.00
Significance between temperatures (within each species) is indicated by different superscript letters (lowercase =
Atlantic; uppercase = shortnose). The species, reared at the same temperature, which is significantly greater in
value is indicated by an (∗ ) and P value. Significance was set at an α of 0.05. SE = ±1 SE.

A21
A A18 A15 A13

100
Shortnose survival (%)

21C
80
18C
60 E21,18 E15 E13 15C

40 13C

20
0
0 5 10 15 20 25 30 35 40 45 50

A21 A18 A15 A13

B
100
Atlantic survival (%)

80
E21,18 E15
60
E13
40
20

0
0 5 10 15 20 25 30 35 40 45 50

Figure 5. Survival curves for larval shortnose sturgeon, A. brevirostrum, and Atlantic, A. oxyrhynchus, sturgeon (B) reared at each
temperature. The point of yolk-sac absorption (A-temp) and escape initiation (E-temp) are represented by arrows on the graph.

yolk-sac absorption (Atlantic: df = 3, 8; shortnose: would also be larger. Shortnose sturgeon spawn ear-
df = 3, 11; P = 0.0001; Figure 5A and B; Table 4). lier in the spring and at lower temperatures (Dadswell
1979, Hall et al. 1991, Kynard 1997) than Atlantic
Discussion sturgeon (Huff 1975, Wooley & Crateau 1985, Smith
1985). The prolonged developmental stages and larger
It has previously been reported that eggs of larger diam- size of larval shortnose sturgeon at hatch may be an
eter produce larger larvae (Blaxter & Hempel 1963, evolved mechanism necessary to survive through peri-
Chambers et al. 1989). Since shortnose sturgeon eggs ods in the spring that are relatively poor in primary
were larger in diameter than those collected from and secondary productivity caused by low temperatures
Atlantic sturgeon, it follows that morphological mea- (Seip & Reynolds 1990) and poor water clarity (Scholtz
sures (SL, BA, and YSA) of shortnose sturgeon at hatch et al. 1988).
152

Table 4. Results from two-way ANOVAs on the time to 100% mortality for each rearing temperature (13◦ C, 15◦ C,
18◦ C, and 21◦ C) between species (shortnose sturgeon, A. brevirostrum, and Atlantic sturgeon, A. oxyrhynchus).
Mean values represent replicate trays within each treatment (n = 3–4).

Variable Species Temp Mean ±1 SE Significance

(◦ C) days between spp.
∗
Time to 100% mortality (d) Shortnose 21 26.5A 0.95 0.00
∗
18 34.5B 1.25 0.00
∗
15 41.0C 2.00 0.03
13 40.7C 5.70 0.08
Atlantic 21 18.3a 0.94
18 22.0b 3.02
15 34.3c 0.94
13 37.3c 1.04
Significance between temperatures (within each species) is indicated by different superscript letters (lowercase =
Atlantic; uppercase = shortnose). The species, reared at the same temperature, which is significantly greater in
value is indicated by an (∗ ) and P value. Significance was set at an α of 0.05. SE = ±1 SE.

Yolk-sac utilization rates and time (post-hatch) to
complete yolk-sac absorption exhibited a significant
inverse relationship with temperature for both short-
nose and Atlantic sturgeon larvae. This has also
been reported for other sturgeon species (Wang et al.
1985, 1987, Gershanovich & Taufik 1992) and some
teleosts (May 1974, Howell 1980). In contrast, the
yolk-sac utilization efficiencies within each species
were not directly affected by rearing temperature. YUE
is directly dependent on the relationship of the rate
at which the yolk is utilized in metabolic processes
with the rate at which it is incorporated into body
tissue (Blaxter & Hempel 1966). The similar efficien-
cies within each species, along with a similar size at
absorption and maximum size attained, indicates their
ability to balance the rise in metabolic requirement at
higher temperatures with a reduction in the devel-
opmental time. A study of white sturgeon Acipenser
transmontanus found that the conversion of yolk
proteins to tissues was not affected between 11◦ C
and 17◦ C yet was lower at 20◦ C (lipid and ash
content varied between temperatures, Wang et al.
1987). Temperatures tested in our study are simi-
lar to that experienced under natural conditions for
these two species; however, to fully determine effi-
ciency bounds, a wider range of temperatures should be
examined.
Shortnose sturgeon larvae utilized their yolk at a
faster rate than Atlantic sturgeon at 13◦ C and 15◦ C
(close at 18◦ C), and were not as efficient in utiliz-
ing it for body tissue at these temperatures. Due to
their higher metabolic demands, larger larvae tend to
experience a decline in efficiency with development
(for reviews see Kamler 1992). In this case, shortnose
sturgeon (larger at all developmental stages) would be
expected to have lower efficiencies than Atlantic stur-
geon reared at the same temperatures. Additionally,
it may also be due to differences in yolk quality and
or species-specific growth rates. Since most marine
species retain more protein for growth than fresh-
water species (Kamler 1992), and mature Atlantic
sturgeon spend most of their time in a marine envi-
ronment (Smith 1985), the maternal influence on the
egg yolk quality may result in higher caloric value than
that of shortnose sturgeon. This may result in more
body tissue incorporated in Atlantic sturgeon larvae as
the yolk is utilized. Giberson and Litvak (2003) also
found that Atlantic sturgeon juveniles of similar size
reared at the same temperature as shortnose sturgeon
had significantly higher growth rates and conversion
efficiencies than shortnose sturgeon suggesting that
they are faster and more efficient at somatic growth
throughout ELH stages.
The development of escape response is essential in
allowing larvae to avoid and survive predatory attacks
(Fuiman 1989). Studies on teleosts have reported
escape response is directly linked to rearing tem-
perature (Shepherd et al. 2000). Few studies have
looked at how temperature affects the development or
mechanisms responsible for escape response in stur-
geon. With the use of scanning electron micrographs,
Richmond & Kynard (1995) found that shortnose
sturgeon larvae incubated at 13–15◦ C began form-
ing a lateral-line system by the 9th day post-hatch
in conjunction with the development of an active
swimming ability. However, the actual initiation of
a C-type escape was not tested. We found that the
C-type response started much later than expected based

on Richmond & Kynard’s (1995) results. Our study
revealed that the development of escape response in
larval shortnose and Atlantic sturgeon was prolonged
in lower rearing temperatures. Similar to growth rate
the systems responsible for anti-predator defenses were
delayed in their morphological development with a
reduction in temperature.
Both shortnose and Atlantic sturgeon larvae survived
considerably longer at lower rearing temperatures
(13◦ C and 15◦ C), with mortality increasing rapidly fol-
lowing the full absorption of the yolk. The acceleration
of mortality in yolk-sac larvae in higher temperatures is
often attributed to higher metabolic rates, which result
in faster absorption of endogenous energy sources
(Kamler 1992). Although not the focus of this study,
the apparent wide range of temperatures tolerated by
shortnose and Atlantic sturgeon is similar to what
was found with white sturgeon A. transmontanus and
lake sturgeon Acipenser fluvescens (Wang et al. 1985,
1987). Again, this suggests that to determine the toler-
ance of these species we need to increase the range of
temperatures tested.
Experiencing variations in river temperatures within
their range of tolerance, wild larval shortnose and
Atlantic sturgeon may experience both beneficial and
deleterious effects on survival. With a faster yolk uti-
lization and growth rate, larvae reared at high tempera-
tures may switch to exogenous feeding much sooner
than those reared at lower temperatures (Wang et al.
1987), thereby avoiding additional risks of predation
in the vulnerable yolk-sac stage (Howell 1980, Miller
et al. 1988). In contrast, the accelerated larval develop-
ment may result in a lack of synchronization with food
availability (Cushing 1972). This has been referred to
as the ‘match–mismatch’ hypothesis, which suggests
that larval survival and recruitment are conditioned by
the match of larvae with prey fields in time and space
(Cushing 1972).
In addition to timing of stage development (yolk
absorption, escape initiation, max. size), the size of
shortnose and Atlantic sturgeon larvae at the start of
these stages may determine their ability to effectively
swim and feed. At a larger size, a larva’s ability to sur-
vive, avoid predation, and effectively capture prey may
increase (Hunter 1972, Miller et al. 1988, Litvak &
Leggett 1992). Our findings indicate that at the critical
yolk-sac absorption stage, similar foraging and escape
opportunity may be afforded for both species of stur-
geon larvae developing in a wide range of temperatures.
Intraspecific comparisons revealed that both shortnose
and Atlantic sturgeon larvae exhibited similar sizes at
153
first escape response when raised at different tempera-
tures. This may allow relatively similar ability to avoid
predation when these mechanisms are fully developed.
It must be considered, however, that escape speeds may
vary at different temperatures (Shepherd et al. 2000),
and that a delay in development of the systems needed
to react to attacking predators may inevitably affect
survival.
Despite temperature, shortnose and Atlantic stur-
geon attained similar maximum sizes and were equally
efficient at doing so. Energy transfer is often lost
through metabolic processes and it stands to reason
that a larva developing in warmer temperatures would
not reach as great a size as those at lower temperatures.
This is not so with these two species, and is signif-
icant to the understanding of their ELH stages. This
balance in metabolic processes may give a competi-
tive advantage in survival and recruitment over other
species whose development is more closely tied to
environmental conditions.
Acknowledgements
Thanks to C.J. Hardy, J.D. Kieffer and B.A. MacDonald
for their editorial comments; lab technicians:
C.A. Doherty, A.V. Giberson and T.D. Shepherd for
all their work in maintaining the eggs until hatch;
A.V. Giberson for morphometric analysis; and P. Soucy
for all of his help in spawning and holding broodstock.
Funding for this work was provided by grants from
Sir James Dunn Research Center Fund, World Wildlife
Fund Canada and Canadian Wildlife Service, NB Trust
Fund, and NSERC to M.K.L.
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