Journal of Freshwater Ecology

ISSN: 0270-5060 (Print) 2156-6941 (Online) Journal homepage: http://www.tandfonline.com/loi/tjfe20

Assessment of pallid sturgeon relative condition in

the upper channelized Missouri River
Kirk D. Steffensen & Gerald E. Mestl
To cite this article: Kirk D. Steffensen & Gerald E. Mestl (2016): Assessment of pallid sturgeon
relative condition in the upper channelized Missouri River, Journal of Freshwater Ecology, DOI:
10.1080/02705060.2016.1196465
To link to this article: http://dx.doi.org/10.1080/02705060.2016.1196465

Published online: 05 Aug 2016.

Submit your article to this journal

Article views: 2

View related articles

View Crossmark data

Full Terms & Conditions of access and use can be found at

http://www.tandfonline.com/action/journalInformation?journalCode=tjfe20
Download by: [Southern Illinois University]

Date: 09 August 2016, At: 07:25

JOURNAL OF FRESHWATER ECOLOGY, 2016

http://dx.doi.org/10.1080/02705060.2016.1196465

Assessment of pallid sturgeon relative condition in the upper

channelized Missouri River
Kirk D. Steffensen and Gerald E. Mestl

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

Fisheries Division, Nebraska Game and Parks Commission, Lincoln, NE, USA

ABSTRACT

ARTICLE HISTORY

During the spring 2015 sampling efforts on the Missouri River, several
adult pallid sturgeon (Scaphirhynchus albus) were captured in a visually
emaciated condition. As pallid sturgeon recovery efforts continue
throughout the Missouri River, this may negatively impact the local
population and hamper recovery. Therefore, the objectives were to (1)
assess the annual variation in pallid sturgeon condition, (2) determine how
condition varies amongst gender and reproductive status and if condition
affects fecundity, (3) determine the annual proportion of reproductive
female pallid sturgeon, and (4) test several hypothesized factors which
may contribute to variations in pallid sturgeon condition. Overall, the
relative condition of pallid sturgeon has signicantly declined over the
past 12 years, except for the large adult-sized sh (> 840 mm fork length).
Pallid sturgeon condition began to decline in 2013 then rapidly declined
in 2014 and even more in 2015. The mean annual pallid sturgeon
condition reached record lows in 2015. Females had a higher overall Kn
compared to males and sexually reproductive sh had a signicantly
higher Kn for both genders. Overall, 28% of the females captured have
been in reproductive condition; however, that frequency appears to be
declining with condition. Also, relative condition was not correlated with
the number of eggs released by reproductively ready females. Correlation
between pallid sturgeon condition and hypothesized variables did not
result in any statistically signicant relationships that would explain the
variations in pallid sturgeon condition. As pallid sturgeon recovery
continues throughout the lower Missouri River, concern arises over
whether or not the current habitat available and river management can
support a viable pallid sturgeon population.

Received 4 May 2016

Accepted 22 May 2016
KEYWORDS

Endangered species; Missouri

River; recovery; relative
condition; Scaphirhynchus;
sturgeon

Introduction
Historically, the Missouri River was a large, free-owing dynamic river system that meandered
extensively back and forth across the oodplain, continuously creating new habitats (Galat & Lipkin
2000; Galat et al. 2005). In its unaltered state, the Missouri River had annual cycles of high and low
discharge that resulted in constantly changing depths and velocities. These conditions provided
access to a wide variety of habitats for sh and acted as spawning cues for many native species. The
Missouri River is now a highly modied system managed for ood control, navigation, irrigation,
hydropower, water supply, water quality, recreation, and sh and wildlife (US Army Corps of
Engineers 2001). Modications included construction of six mainstem dams, an extensive levee
system and a highly altered navigation channel from Sioux City, Iowa (river kilometer [rkm]

CONTACT Kirk D. Steffensen

kirk.steffensen@nebraska.gov

2016 Informa UK Limited, trading as Taylor & Francis Group

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

K. D. STEFFENSEN AND G. E. MESTL

1178.8) to the conuence with the Mississippi River (rkm 0.0). Overall, approximately 35% of the
river has been impounded by the mainstem dams, 32% has been channelized for navigation, and
33% remains relatively unaltered but is interspersed among reservoirs where temperature, ows,
and turbidity have been greatly altered (Keenlyne 1989; Hesse & Mestl 1993). The total water surface
of the channelized reach has been reduced by approximately 50% (Funk & Robinson 1974) and
annual sediment discharge has decreased by 100 150 million metric tons, more than is currently
transported annually by the entire Missouri Mississippi River systems to the Gulf of Mexico
(Meade & Moody 2010). The mainstem dams and current water management practices have altered
the natural hydrograph and sediment transport system that historically created the dynamic habitat
elements for the native fauna and ora (Funk & Robinson 1974; Hesse et al. 1993; Latka et al. 1993;
Pegg et al. 2003). Furthermore, these modications have eliminated the natural ow regime and created a high-energy channel that has promoted stream bed degradation and disconnected the Missouri River from much of its oodplain (Winston & Criss 2003).
Flows on the lower Missouri River are controlled by Gavins Point Dam (Yankton, SD; rkm
1305.2), which is the smallest and lowest dam on the mainstem Missouri River. Gavins Point Dams
purpose is to stabilize the irregular discharge from upstream hydro-peaking and to support navigation on the lower Missouri River, resulting in articially stable ows during the March to November
navigation season that do not mimic the natural hydrograph. Before dam construction, the hydrograph had an early peak in March, followed by a second peak in late June, which inundated the lowlying oodplain and islands, then continuously declining stages for the rest of the year. The current
channel conguration and water management prevents oodplain connection, except during
extreme water years. This lack of connection limits food availability reduces spawning areas and limits refugia area for age-0 shes (Junk et al. 1989). However, high discharges through the upper channelized reach of the Missouri River in 2010 and 2011 resulted in connection to the oodplain and
increased levels of macroinvertebrate and sh production (Steffensen, Eder et al. 2014); therefore,
increased food resources.
These aforementioned dam operations and physical modications were identied as contributing
to the population decline of the federally listed pallid sturgeon (Scaphirhynchus albus) on the
Missouri River (Dryer & Sandvol 1993; USFWS 2014). These same modications and water management practices are also implicated in population declines documented for several other native
shes in the Missouri River (Hesse et al. 1989, 1993; Steffensen, Shuman et al. 2014; Steffensen,
Shuman & Stukel 2014). To supplement the diminished wild pallid sturgeon population, conserve
unique genotypes, and to bolster the number of reproductive sh in the system, the Pallid Sturgeon
Conservation Augmentation Program was implemented in the lower Missouri River (i.e. Gavins
Point Dam [rkm 1305.2] to the conuence with the Mississippi River [rkm 0.0]). Stocking occurred
three times in the 1990s and annually since 2001 with more than 163,000 pallid sturgeon stocked
into the lower Missouri River (Huenemann 2015). The primary concern of the Pallid Sturgeon
Conservation Augmentation Program was to augment the suppressed pallid sturgeon population
while properly managing for genetic contributions; there was little discussion about the current
carrying capacity of the system and impacts to other benthic shes.
Adult pallid sturgeons are piscivorous and dependent on the native chubs and minnows as their
primary prey. Pallid sturgeon prey selectivity is unknown as the early diet studies only reported
small shes in their stomachs (Coker 1930; Cross 1967; Carlson et al. 1985), not species-specic
information. Recent food habit studies have been more descriptive of species consumed but were
conducted in modied reaches where the sh community has been highly altered. In the upper Missouri River (i.e. Fort Peck Dam [rkm 2850.9] to the headwaters of Lake Sakakawea [rkm 2523.5]),
Gerrity et al. (2006) found that sturgeon chub (Macrhybopsis gelida) and sicklen chub (M. meeki)
comprised 79% of all identiable sh recovered from juvenile pallid sturgeon stomachs. Channel
catsh (Ictalurus punctatus), athead chub (Platygobio gracilis), sand shiner (Notropis stramineus),
and shorthead redhorse (Moxostoma macrolepidotum) were also consumed. In the inter-reservoir
reach of the middle Missouri River bordering Nebraska and South Dakota, Wanner et al. (2007)

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

JOURNAL OF FRESHWATER ECOLOGY

found that Johnny darter (Etheostoma nigrum), channel catsh, silver chub (M. storeriana), and
emerald shiner (N. atherinoides) were the primary prey selected. In the channelized reach of the
lower Missouri River, just below our study area, Winders et al. (2014) found Ictaluridae and
Cyprinidae were the primary prey for pallid sturgeon. Specically, the Cyprinidae family was most
represented by Macrhybopsis but the level of digestion generally prohibited further identication.
Finally, Hoover et al. (2007) found that pallid sturgeon in the middle Mississippi River consumed
Cyprinidae, Sciaenidae, and Clupeidae.
Several of these species identied as prey for pallid sturgeon are below historic levels (Meek 1892;
Johnson 1942) and continue to decline locally (Hesse 1994; Steffensen, Shuman & Stukel 2014) and
basin-wide (USFWS 2001), especially the chub species. More specically, the range of sicklen chub
and sturgeon chub has been reduced in the mainstem Missouri River by more than 1609 km (i.e.
1287 km of impoundment and 322 km of inter-reservoir reaches), and sturgeon chub only occur in
11 of their 30 historic tributaries (USFWS 2001). Flathead chub populations have declined across
much of the mainstem Missouri River downstream from North Dakota to Missouri (Tibbs 1998).
Understanding the impacts of condition on the pallid sturgeon population may be critical for the
species recovery as condition affects survival, growth rates, and reproductive potential (Pope &
Kruse 2007). Blackwell et al. (2000) suggested that sh in good condition have more energy available
for gamete production thus increasing relative fecundity. The relationship between condition and
fecundity for pallid sturgeon from the lower Missouri River is unknown, but fork length and weight
displayed a positive correlation with the number of eggs released (Steffensen et al. 2013a). However,
this information needs to be assessed cautiously as the sample size was very limited (n D 11).
During the spring 2015 sampling efforts along the Nebraska reach of the lower Missouri River
(see Welker & Drobish 2012a, 2012b for sampling details), several adult pallid sturgeon were captured in visually poor condition. Furthermore, after all potential brood sh (i.e. those great than
800 mm FL) were assessed for reproductive readiness, the proportion of reproductive females was
below the previously observed levels. The majority of adult wild and hatchery-reared pallid sturgeon
in the lower Missouri River resides in the upper channelized (Sioux City, IA [rkm 1178.8] to Kansas
City, MO [rkm 591.4]) reach (K. Steffensen, NGPC, unpublished data). Negative changes in condition of these sh may have important implications for pallid sturgeon reproduction and ultimately
recovery of the species to a naturally, self-sustaining population. Therefore, the objectives of this
study were to (1) document the temporal variation in pallid sturgeon condition against time (i.e.
2003 2015) by pallid sturgeon proportional size distribution (PSD) categories; (2) assess variation
in condition amongst gender and reproductive status (i.e. reproductive or non-reproductive) then
determine if reproductive female condition was related to the number of eggs produced; (3) document the percentage of reproductive female pallid sturgeon compared to no non-reproductive sh;
and (4) correlate the overall annual pallid sturgeon condition against several hypothesized factors
which may contribute to the decline in pallid sturgeon condition.

Methods
The study area was a 423.4 rkm reach of the lower Missouri River along Nebraskas eastern border
from Lower Ponca Bend (rkm 1211.8) to the Nebraska Kansas state line (rkm 788.4). This reach
has been channelized with a series of dike structures on the inside bends and limestone rock armoring the outside bends. The Platte River (rkm 957.6) intersects this reach and seasonally impacts the
hydrograph of the Missouri River.
Pallid sturgeon capture data (i.e. 2003 2015) were obtained from the Nebraska Game and Parks
Commissions (NGPC) Pallid Sturgeon Population Assessment (PSPA) Program sampling efforts
from the upper channelized reach (rkm 1212 789) of the Missouri River (Welker & Drobish 2012a,
2012b) and NGPCs annual intensive effort directed at capturing reproductively ready pallid
sturgeon for brood stock for hatchery spawning (i.e. Conservation Augmentation Program). Pallid
sturgeons were classied as wild or hatchery-reared according to the presence of a hatchery mark

K. D. STEFFENSEN AND G. E. MESTL

and/or tag or by post hoc genetic determination (Schrey & Heist 2007; Schrey et al. 2007; DeHaan
et al. 2008). Gender and gonad maturation stage were determined using ultrasound and endoscopic
techniques described in Bryan et al. (2007) and Divers et al. (2009) and sh categorized as reproductive (stage IV, ready to spawn this season) or non-reproductive (stage II III; Moos 1978; Colombo
et al. 2007; Tripp et al. 2009). The number of eggs released was estimated using volumetric estimation by counting a subset of number of eggs/mL and extrapolating to the total volume extracted.
Complete or incomplete spawn was noted during a post-spawn biopsy and only females that
completely spawned were included in the fecundity analysis.
Relative condition (Kn) was used to assess change in pallid sturgeon condition with the pallid
sturgeon weight length relationship calculated based on the equation provided by Shuman et al.
(2011),

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

log10 .W/ D 6:2561 C 3:2932 log10 .FL/;

where FL and W are the observed fork length and weight for each individually captured pallid sturgeon. The Shuman et al. (2011) equation is based on a regional assessment (i.e. Missouri River), not
the entire species range (Murphy & Willis 1992); therefore, relative condition factor (Kn) was calculated as
Kn D W=W;
where W is the individuals observed weight and W is the predicted weight from the weight length
relationship as described by the Shuman et al. (2011) equation.
Seasonal (i.e. pre-spawn (March April) and post-spawn (May November)) and origin (wild vs.
hatchery-reared) effects were tested (ANOVA [analysis of variance] with a Tukey adjustment) to
determine if there were differences in Kn. If treatments were signicantly different, then further
assessment would be analyzed by season and origin. However, if treatments did not differ, then data
from each treatment were merged into a single cohort. To test the annual trends, the annual mean
Kn between years was calculated and regressed by the incremental PSD indices developed for pallid
sturgeon (Shuman et al. 2006; Guy et al. 2007). Pallid sturgeon length categories were stock to quality (S-Q; 330 629 mm fork length [FL]), quality to preferred (Q-P; 630 839 mm), preferred to
memorable (P-M; 840 1039 mm), and memorable to trophy (M-T; 1040 1269 mm). No trophysized sh (>1270 mm) were collected. Next, to determine if condition varied between gender and
reproductive status, annual mean and overall mean Kn by gender and reproductive condition were
compared (ANOVA with a Tukey adjustment). If annual differences were detected then a pairwise
comparison (LSMEANS within PROC GLM, SAS 9.4) tested for differences amongst years. In addition, the number of eggs produced was regressed against condition to determine if relative condition
affects fecundity and the percent of reproductive compared to non-reproductive females was calculated. Finally, overall pallid sturgeon condition was correlated against several hypothesized factors
that may contribute to the variation in pallid sturgeon condition. The hypothesized, independent
variables that were acquired and tested included (1) the mean annual discharge of the Missouri
River at the Omaha (USGS 06610000) and Nebraska City gages (USGS 06807000), (2) intraspecic
competition which used the Steffensen et al. (2013b) population viability model to estimate the current pallid sturgeon population in the lower Missouri River, (3) interspecic competition which
used the standardized, long-term electroshing catch per unit effort (CPUE) for athead catsh
Pylodictis olivaris (Porter et al. 2011 and M. Pegg, University of Nebraska-Lincoln, Pers. Comm.),
(4) food availability by calculating CPUE from the NGPCs PSPA Programs standardized sampling
efforts for (a) annual otter trawl catch rates for sturgeon and sicklen chubs which were described
as primary prey items by Gerrity et al. (2006), (b) annual otter trawl catch rates for a suite of species
documented by Winders et al. (2014), and (c) annual mini-fyke net catch rates for all small bodied
sh. Since the majority of pallid sturgeons are collected in March and April, before the independent

JOURNAL OF FRESHWATER ECOLOGY

variables were measured, independent variables were set forward one year to adjust to the observed
conditions. All statistical tests were performed using SAS 9.4 (SAS Institute, Cary, NC) and signicance was determined at a D 0.05.

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

Results
Relative condition did not differ between pre-spawn (March April) and post-spawn (May
November) season for any of the pallid sturgeon size categories (S-Q, t D 1.41, p D 0.159; Q-P,
t D 1.79, p D 0.074; P-M, t D 1.53, p D 0.128; M-T, t D 0.85, p D 0.398). Furthermore, there was no
difference in mean Kn between wild-origin sh and hatchery-reared pallid sturgeon within size
categories (S-Q, t D 1.06, p D 0.290; Q-P, t D 1.89, p D 0.060; P-M, t D 1.27, p D 0.204; M-T,
t D 0.27, p D 0.790); therefore, wild and hatchery-reared pallid sturgeon were grouped with no
season or origin effect and summarized annually.
From 2003 to 2015, a signicant downward trend in pallid sturgeon Kn has occurred in all size
categories, except for M-T sized sh (Table 1; Figure 1) but this exception is being inuenced by
only three M-T-sized sh captured from 2003 to 2007. However, from 2011 to 2015, all size categories have experienced a signicant decline and the annual pairwise comparison between years were
signicantly different for all PSD size classes, except for the M-T sized sh. Annual pairwise comparisons were signicantly different for S-Q sized sh with a signicant decrease in Kn from 2011 to
2012 compared 2013 2015. However, the past three years were similar. A similar trend occurred
with Q-P and P-M sized sh with a signicant annual decline in 2014 2015 compared to
2011 2013. Relative conditions for both size classes have reached an unprecedented low. Even
though there is no long-term difference for M-T sized sh, Kn for M-T declined to an annual mean
of 0.79 in 2015.
Pallid sturgeon appears to exhibit sexual dimorphism in Kn between males and females
(Figure 2). Female pallid sturgeon had a signicantly higher overall Kn compared to males (t D
2.44, p D 0.015; Table 2). Fish in reproductive condition had a signicantly higher Kn compared to
non-reproductive for both genders (Table 2). Furthermore, comparisons between annual mean Kn
for reproductive females did not vary annually; whereas, non-reproductive females and males and
reproductive males all varied annually (Figure 2). Overall, Kn greatly improved in 2012 for all potential reproductive sized sh (> 800 mm) followed by a rapid decline in Kn for all sh, excluding
reproductive females.
Forty reproductive female pallid sturgeons have been captured since 2008. Genetic concerns prevented 10 reproductive females from being spawned. Twenty sh successfully released eggs, but
three were determined to have had an incomplete spawn. Relative condition did not differ between
sh that successfully spawned (Kn D 0.97, SE D 0.02) compared to sh that did not release eggs
(Kn D 0.93, SE D 0.01; t D 0.93, p D 0.360). Overall, 28% of the females captured have been in
reproductive condition; however, from 2008 to 2013, 34% (n D 32) were in reproductive condition
compared to only 17% (n D 8) the past two years (Figure 3). For females that successfully spawned,
Table 1. Regression statistics (over study duration [2003 2015] and short-term [2011 2015]), ANOVA model, and annual differences in relative condition (Kn) by incremental proportional size distribution (PSD) size categories for all pallid sturgeon captured
from 2003 to 2015 in the upper channelized Missouri River. Letters denote statistical comparison within each size category.
PSD size categories are: S-Q D 330 629 mm fork length (FL), Q-P D 630 839 mm), P-M D 840 1039 mm, and M-T D
1040 1269 mm).
Regression (2003 2015) Regression (2011 2015)
ANOVA model
Mean annual Kn
p
r2
p
F
p
2011 2012 2013 2014 2015
Size category
r2
S-Q
0.14
< 0.0001
0.11
< 0.0001
22.57 < 0.0001 0.90a 0.91a 0.84b 0.84b 0.84b
Q-P
0.16
< 0.0001
0.28
< 0.0001
15.04 < 0.0001 0.94a 0.94a 0.87b 0.82c 0.81c
P-M
0.28
< 0.0001
0.33
< 0.0001
15.84 < 0.0001 0.97a 0.95a 0.89b 0.85c 0.83c
M-T
0.28
0.199
0.35
0.029
1.44
0.199 0.96 1.02 0.89 0.89 0.79


No pairwise comparisons made between years as the model was not signicant.

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

K. D. STEFFENSEN AND G. E. MESTL

Figure 1. Annual mean (SE) relative condition factor (Kn) by incremental proportional size distribution (PSD) size classes for all
pallid sturgeon captured from 2003 to 2015 in the upper channelized Missouri River. Solid regression line indicates the overall
change in Kn, while the dashed regression line displays the change over the past ve years.

the number of eggs was highly variable and ranged from 12,220 to 54,705 (mean D 25,087) or 4721
to 13,128 (mean D 7443) per kg of body weight. Relative condition for the reproductively ready pallid sturgeon that successfully spawned was not correlated with the number of eggs released
(Figure 4).
Correlations between pallid sturgeon condition and hypothesized independent variables did not
result in any signicant relationships (Table 3). The annual catch rate of potential prey sh
described by Winders et al. (2014) with a one year offset had the strongest correlation followed by
athead catsh catch rates. Whereas, the pallid sturgeon population estimate and discharge variable
showed the weakest correlation.

Discussion
The majority of the pallid sturgeon, especially the wild pallid sturgeon, resides in the upper channelized reach of the lower Missouri River. The observed decrease in condition is probably related to the
increased bioenergetic demands and reduced food resources because of habitat loss caused by channelization; however, no statistical differences were determined. Karr (1991) noted that higher trophic level sh (i.e. piscivorous pallid sturgeon) are more vulnerable to habitat loss and degradation,
which likely affects shes condition. This low condition has decreased maturation rates with reduced
fecundity and is likely interrupting reproductive cycling. The fact that the annual condition of pallid
sturgeon in Nebraska varies so much from year to year may indicate that condition may be very
closely linked to the amount of prey in the system. Many native chubs appear to be as imperiled as

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

JOURNAL OF FRESHWATER ECOLOGY

Figure 2. Annual mean (SE) relative condition factor (Kn) by gender and reproductive assessment for adult (> 800 mm) pallid
sturgeon transported to local hatcheries as potential brood sh.

pallid sturgeon (Hesse 1994; Galat et al. 2005; Steffensen, Shuman & Stukel 2014) but have not
received the attention through federal listing as a threatened or endangered species. When the Conservation Augmentation Program was initiated, biologists assumed that the carrying capacity of the
lower Missouri River was adequate to support the recovery goal to create a self-sustaining population (Dryer & Sandvol 1993; USFWS 2014); however, the current rivers carrying capacity remains
unknown. Under the current river management it is questionable if the upper reach of the lower
Missouri River, in its current condition, can support a self-sustaining population of pallid sturgeon.
Keenlyne and Evenson (1993) developed the original relative weight equation for pallid sturgeon
and concluded that sh from the channelized Missouri River were underweight compared to
sh from the upper Missouri River (i.e. Fort Peck Dam [rkm 2850.9] to the headwaters of Lake
Sakakawea [rkm 2523.5]). Shuman et al. (2011) detected differences in Kn by incremental PSD
Table 2. Mean relative condition (Kn) and pairwise comparisons for pallid sturgeon by gender and reproductive assessment.
Pairwise comparison
ANOVA model
Gender
Overall Kn
Reproductive assessment
Mean Kn
t
p
F
p
Reproductive
0.97
0.18
0.9869
Female
0.91
4.78
< 0.0001
Non-reproductive
0.89
11.18
0.001
Reproductive
0.90
3.74
0.001
Male
2.43
0.0158
0.88
Non-reproductive
0.86
7.58
< 0.0001

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

K. D. STEFFENSEN AND G. E. MESTL

Figure 3. Percentage of reproductive females (gray bar) compared to non-reproductive females (black bar) for female pallid
sturgeon collected in the upper channelized reach of the lower Missouri River.

Figure 4. Number of eggs released by relative condition factor (Kn) for 17 reproductively ready female pallid sturgeon collected in
the upper channelized reach of the lower Missouri River and successfully spawned at a hatchery facility.

Table 3. Correlation statistics for independent variables hypothesized to be affecting pallid sturgeon Kn in the upper channelized
reach of the Missouri River.
Annual comparison
1-yr offset
Variable
r
p
r
p
Omaha discharge
0.12
0.6900
0.13
0.6628
Nebraska City discharge
0.15
0.6264
0.22
0.4604
Pallid sturgeon population estimate
0.05
0.8688
0.22
0.4722
Flathead catch rates
0.44
0.1548
0.31
0.3204
0.11
0.7309
0.14
0.6566
Chubs catch ratesa
b
0.12
0.6991
0.46
0.1133
Potential prey sh catch rates
Mini-fyke net catch rates
0.24
0.4217
0.35
0.2434
a

Sturgeon chubs and sicklen chubs only.

Suite of small bodied sh described in Winders et al. (2014).

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

JOURNAL OF FRESHWATER ECOLOGY

between the Pallid Sturgeon Recovery Priority Management Areas on the Missouri River but did not
provide any temporal differences. Pallid sturgeon in the lower Missouri River (i.e. Gavins Point Dam
[rkm 1305.2] to the conuence with the Mississippi River [rkm 0.0]) were always characterized as
being a smaller, slender sh compared to the remnant adults in the upper Missouri River. This likely
reects the demands of living in a highly engineered channelized environment; however, pallid sturgeon relative condition in the channelized Missouri River was always assumed adequate.
Condition indices may not always provide the best assessment for the change in sturgeon condition. Beamish et al. (1996) concluded that when food resources are limited, lake sturgeon (Acipenser
fulvescens) increase water uptake. We acknowledge this potential bias, but Kn for pallid sturgeon
along Nebraskas eastern border of the Missouri River are declining and sh are, on visual examination alone, emaciated. If pallid sturgeon can alter their body weight through water retention, then
conditions in the upper channelized Missouri River may be worse than our analysis is predicting. As
Kn declines, protein and lipid reserves are likely depleted and mortality may occur. Lipids tend to
be the primary energy source for gonadal development (Beamish et al. 1996). Without sufcient
lipid reserves to produce viable gametes, natural reproduction is unlikely. Until conditions can be
improved, recovery of the species is unlikely and species persistence will remain dependent on
hatchery supplementation. Concerns exist that the current levels of hatchery supplementation may
exceed the current carrying capacity for this reach of the Missouri River but the results of this study
do not support those concerns.
The mean condition factor for pallid sturgeon from the lower Missouri River along Nebraskas
eastern border since 2003 was around 0.90 but highly variable. However, it is unknown what the
mean condition factor should be for pallid sturgeon. Quist et al. (1998) suggested relative weight target values between 80 and 90 for the sympatric shovelnose sturgeon (Scaphirhynchus platorynchus)
excluding the large adults found in Montana. Although fairly similar in body shape, this recommendation may not be valid for the piscivorous pallid sturgeon, especially for the larger PSD size classes
(i.e. P-M and M-T). The diets of these two species have low similarity niche (Gerrity et al. 2006).
For the most part, only reproductively ready pallid sturgeon are achieving a Kn greater than 0.90,
which is the effect of gonadal development, while the current mean population Kn is approaching
the lower end of that recommendation.
Survival of hatchery-reared pallid sturgeon stocked in the lower Missouri River is similar to other
sturgeon species (Steffensen et al. 2010). Hatchery-reared sh in the lower Missouri River were growing (Shuman et al. 2011), exhibiting migratory behavior and reaching sexual maturity (DeLonay et al.
2010; DeLonay et al. 2012; Steffensen et al. 2013a). The numbers of reproductive sh in the lower
Missouri River was increasing prior to 2013; however, the proportion of reproductive sh collected
annually continues to decrease. Furthermore, the number of reproductively ready female pallid
sturgeon captured steadily increased from 2008 to 2013 but has decreased the past two years. A
similar trend was observed with capture-recaptured, telemetry-tagged pallid sturgeon. Several females
were predicted to be in reproductive condition in 2015 based on past reproductive histories but did
not achieve reproductive readiness. There is also some evidence that sh may not be maturing at the
same approximate size as they were prior to 2011. Prior to 2011, Steffensen et al. (2013a) determined
that the minimum length-of-maturity was approximately 800 mm, but over the past two years the
minimum length of reproductive females sampled has been 895 mm. Several known hatchery-reared
pallid sturgeon from 2001 and 2002 year classes were captured this past year and were determined to
be still immature and yet to reach their rst maturation cycle (J. Haas, NGPC, Pers. Comm.). Despite
all of this, a few pallid sturgeon in the upper reach of the lower Missouri River are still maintaining a
higher condition factor, becoming reproductively ready, and producing viable gametes.
Relative fecundity for females that reach reproductive readiness captured in the upper reach
(above Kansas City, MO [rkm 591.4]) of the lower Missouri River was considerably lower than sh
from the lower reach (below Kansas City, MO). George et al. (2012) estimated total fecundity of two
pallid sturgeon (mean weight 3.2 kg) captured in the lower Missouri River at 57,192 and 58,913 eggs
or approximately 18,140 eggs per kg of body weight. The reproductive females captured during our

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

10

K. D. STEFFENSEN AND G. E. MESTL

study were slightly larger (average weight D 3.7 kg) but only produced an average of 7400 eggs per
kg body weight. Additionally, Keenlyne et al. (1992) reported approximately 9900 eggs per kg body
weight for an old (age-41), 30.8 kg pallid sturgeon taken from North Dakota.
Even though a local difference was observed, relative fecundity for pallid sturgeon in the upper
reach of the lower Missouri River was similar to other large-sized sturgeon species. Beluga sturgeon
Huso huso (Linnaeus) was reported to have a relative fecundity between 2750 to 10,500 eggs per kg
body weight (Raspopov 1987), while white sturgeon ranged from 3192 to 8582 with a mean of 5648
eggs per kg body weight (Chapman et al. 1996). Smaller bodied sturgeon species have been reported
to produce more eggs per kg body weight with shortnose sturgeon (A. brevirostrum) having an average relative fecundity of 11,568 (Dadswell 1979), while shovelnose sturgeon, in the local proximity
of our study, had 18,355 eggs per kg body weight in the Platte River (Hamel et al. 2015).
Comparatively, pallid sturgeon captured in the Platte River, a major tributary to the Missouri River,
which bisects our study area, had a mean Kn ranging from 0.95 to 0.99 from 2009 to 2012 (Hamel
et al. 2014). If our data-set is truncated to the same time period, the reported Kn values for the Platte
River are still higher compared to Kn observed on the Missouri River (Kn D 0.93, SE D 0.003), but
the difference is lessened. Beamesderfer (1993) reported that white sturgeon (Acipenser transmontanus) relative weight (Wr) varied from 77 to 117 with the lowest Wr from areas with limited food
availability and declining populations. This suggests that sh in the Platte River are likely nding additional food resources to enhance their condition as the population between the Platte and Missouri
Rivers are sympatric. Furthermore, the available habitat and ow regime in the Platte River are more
favorable which allow pallid sturgeon at least to maintain or improve their conditions.
We documented that pallid sturgeon in the upper channelized Missouri River can rapidly
increase body weight and improve Kn when food resources become available, although not statistically correlated small bodied shes catch rates. There was a rapid increase in body weight and Kn
after the 2010 high water and 2011 major ood events when additional food and habitat resources
were readily available. There was increased reproduction of many native species during the major
ood in 2011 (Steffensen, Eder et al. 2014) and likely provided abundant food resources for pallid
sturgeon. Pallid sturgeon responded and reached a mean Kn greater than 0.95 after these two years.
However, the benets of 2010 and 2011 oods were short lived and were followed by a rapid decline
of pallid sturgeon condition over the past three years when food resources possibly decreased. Also
during the 2011 ood, adult pallid sturgeon were utilizing the slower inundated oodplain habitats
which likely reduced their biogenetics demands compared to residing in the swift currents of the
main channel (Haas et al. 2014). Therefore, current river management needs to focus on the lower
trophic levels and pallid sturgeon condition will have a positive response to increased food
availability.

Acknowledgments
The authors acknowledge the Nebraska Game and Parks Commission crews from the Pallid Sturgeon Population
Assessment Program, Habitat Assessment and Monitoring Program, and Pallid Sturgeon Research Program for their
hard work over the past 12 years and the hundreds of volunteers who have assisted them during their annual pallid
sturgeon brood stock collection efforts since 2008. Finally, the authors also thank Rick Holland and the Journal of
Freshwater Ecologys reviewers for their comments on earlier drafts of this manuscript.

Disclosure statement
No potential conict of interest was reported by the authors.

Notes on contributors
Gerald Mestl received his masters of science degree in zoology from Oklahoma State University. Gerald serves as the
Missouri River Program Manager for the Fisheries Division of the Nebraska Game and Parks Commission.

JOURNAL OF FRESHWATER ECOLOGY

11

Kirk Steffensen gained his MS degree in sheries science from the University of Nebraska-Lincoln. Kirk is a sheries
biologist II for the Nebraska Game and Parks Commission where he serves of the project leader of the Missouri River
Pallid Sturgeon Population Assessment Project.

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

References
Beamesderfer RC. 1993. A standard weight (Ws) equation for white sturgeon. Calif Fish Game. 79:63 39.
Beamish FWH, Jebbink JA, Rossiter A, Noakes DLG. 1996. Growth strategy of juvenile lake sturgeon (Acipenser
fulvescens) in a northern river. Can J Fish Aquatic Sci. 53:481 489.
Blackwell BG, Brown ML, Willis DW. 2000. Relative weight (Wr) status and current use in sheries assessment and
management. Rev Fish Sci. 8:1 44.
Bryan JL, Wildhaber ML, Papoulias DM, DeLonay AJ, Tillitt DE, Annis ML. 2007. Estimation of gonad volume,
fecundity and reproductive state of shovelnose sturgeon using sonography and endoscopy with application to the
endangered pallid sturgeon. J Appl Ichthyol. 23:411 419.
Carlson DM, Pieger WL, Trial L, Haverland PS. 1985. Distribution, biology and hybridization of Scaphirhynchus
albus and S. platorynchus in the Missouri and Mississippi rivers. Env Bio Fish. 14:51 59.
Chapman FA, Van Eenennaam JP, Doroshov SI. 1996. The reproductive condition of white sturgeon, Acipenser transmontanus, in San Francisco Bay, California. Fish Bull. 94:628 634.
Coker FB. 1930. Studies of common sh of the Mississippi River at Keokuk. US Bur Fish Bull. 45:141 225.
Colombo RE, Garvey JE, Wills PS. 2007. Gonadal development and sex-specic demographics of the shovelnose sturgeon in the middle Mississippi River. J Appl Ichthyol. 23:420 427.
Cross FB, 1967. Handbook of shes of Kansas. Lawrence (KS): University of Kansas Museum of Natural History
(Miscellaneous Publication 45).
Dadswell MJ. 1979. Biology and population characteristics of the shortnose sturgeon, Acipenser brevirostrum LeSueur
1818 (Osteichthyes: Acipenseridae) in the Saint John Estuary, New Brunswick, Canada. Can J Zoo. 57:2186 2210.
DeHaan PW, Jordan GR, Arden WR. 2008. Use of genetic tags to identify captive-bred pallid sturgeon (Scaphirhynchus albus) in the wild: improving abundance estimates for an endangered species. Conserv Genet. 9:691 697.
DeLonay AJ, Jacobson RB, Papoulias DM, Wildhaber ML, Chojnacki KA, Pherigo EK, Bergthold CL, Mestl GE. 2010.
Ecological requirements for pallid sturgeon reproduction and recruitment in the lower Missouri River: annual
report 2009. Columbia (MO): US Geological Survey.
DeLonay AJ, Jacobson RB, Papoulias DM, Wildhaber ML, Chojnacki KA, Pherigo EK, Haas JD, Mestl GE. 2012.
Ecological requirements for pallid sturgeon reproduction and recruitment in the lower Missouri River: annual
report 2010. Columbia (MO): US Geological Survey.
Divers SJ, Boone SS, Hoover JJ, Boysen KA, Kilgore KJ, Murphy CE, George SG, Camus AC. 2009. Field endoscopy for
identifying gender, reproductive stage and gonadal anomalies in free-ranging sturgeon (Scaphirhynchus) from the
lower Mississippi River. J Appl Ichthyol. 25:68 74.
Dryer MP, Sandvol AJ. 1993. Pallid sturgeon recovery plan. Bismarck (ND): US Fish and Wildlife Service.
Funk JL, Robinson JW. 1974. Changes in the channel of the lower Missouri River and effects on the sh and wildlife.
Jefferson City (MO): Missouri Department of Conservation (Aquatic Series 11).
Galat DL, Lipkin R. 2000. Restoring ecological integrity of great river: historical hydrographs aid in dening reference
conditions for the Missouri River. Hydrobiol. 422/423:29 48.
Galat DL, Berry CR, Gardner WM, Hendrickson JC, Mestl GE, Power GJ, Stone C, Winston MR. 2005. Spatiotemporal
patterns and changes in Missouri River shes. In: Rinne JN, Hughes RM, Calamuso B, editors. Historical changes
in large river sh assemblages of the Americans. Bethesda (MD): American Fisheries Society, Symposium 45;
p. 249 291.
George SG, Slack WT, Hoover JJ. 2012. A note on the fecundity of pallid sturgeon. J Appl Ichthyol. 28:512 515.
Gerrity PC, Guy CS, Gardner WM. 2006. Juvenile pallid sturgeon are piscivorous: a call for conserving native cyprinids. Trans Am Fish Soc. 135:604 609.
Guy CS, Neumann RM, Wilis DW, Anderson RO. 2007. Proportional size distribution (PSD): a further renement of
population size structure index terminology. Fish. 32:348.
Haas JD, Eder BL, Archer MW, Adams JD, Ruskamp RL. 2014. Documenting presence of pallid sturgeon on lower
Missouri River oodplains. Prairie Nat. 46:97 99.
Hamel MJ, Pegg MA, Hammen JJ, Rugg ML. 2014. Population characteristics of pallid sturgeon, Scaphirhynchus albus
(Forbes and Richardson, 1905), in the lower Platte River, Nebraska. J Appl Ichthyol. 30:1362 1370.
Hamel MJ, Rugg ML, Pegg MA, Patino R, Hammen JJ. 2015. Reproductive traits of shovelnose sturgeon Scaphirhynchus platorynchus (Ranesque, 1820) in the lower Platte River, Nebraska. J Appl Ichthyol. 31:592 602.
Hesse LW. 1994. The status of Nebraska sh in the Missouri River, selected chubs and minnows (Cyprinidae): sicklen chub (Macrhybopsis meeki), sturgeon chub (M. gelida), silver chub (M. storeriana), speckled chub (M.
aestivalis), athead chub (Platygobio gracilis), plains minnow (Hybognathus placitus) and western silvery minnow
(H. argyritis). Trans Neb Acad Sci. 21:99 108.

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

12

K. D. STEFFENSEN AND G. E. MESTL

Hesse LW, Mestl GE. 1993. An alternative hydrograph for the Missouri River based on the precontrol condition. N
Am J Fish Man. 13:360 366.
Hesse LW, Schmulback JC, Carr JM, Keenlyne KD, Unkenholz DG, Robinson JW, Mestl GE. 1989. Missouri River
shery resources in relation to past, present and future stresses. In: Dodge DP, editor. Proceedings of the International Large River Symposium; Toronto: Canadian Special Publication of Fisheries and Aquatic Sciences 106.
Hesse LW, Mestl GE, Robinson JW. 1993. Status of selected shes in the Missouri River in Nebraska with recommendations for their recovery. In: Heese LW, Stalnaker CB, Benson NG, Zuboy JR, editors. Restoration planning for
the river of the Mississippi River ecosystem. Washington (DC): National Biological Survey, Biological Report 19;
p. 327 340.
Hoover JJ, George SG, Kilgore KJ. 2007. Diet of shovelnose sturgeon and pallid sturgeon in the free-owing Mississippi River. J Appl Ichthyol. 23:494 499.
Huenemann TW. 2015. Central lowlands and interior highlands pallid sturgeon spawning and stocking summary.
Lincoln (NE): Nebraska Game and Parks Commission.
Johnson RE. 1942. The distributions of Nebraska shes [MS thesis]. Ann Arbor (MI): University of Michigan.
Junk WJ, Bayley PB, Sparks RE. 1989. The ood pulse concept in river-oodplain systems. In: Dodge DP, editor.
Proceedings of the International large River Symposium. Ottawa: Canadian Special Publications of Fisheries and
Aquatic Sciences.
Karr JR. 1991. Biological integrity: a long-neglected aspect of water resource management. Eco App. 1:66 84.
Keenlyne KD. 1989. A report of the pallid sturgeon. Pierre (SD): US Fish and Wildlife Service.
Keenlyne KD, Evenson PD. 1993. Standard and relative weight for the pallid sturgeon (Scaphirhynchus albus). Proc S
Dak Ac Sci. 2:41 49.
Keenlyne KD, Grossman EM, Jenkins LG. 1992. Fecundity of pallid sturgeon. Trans Am Fish Soc. 121:139 140.
Latka DC, Nestler J, Hesse LW. 1993. Restoring physical habitat in the Missouri River: a historical perspective. In:
Heese LW, Stalnaker CB, Benson NG, Zuboy JR, editors. Restoration planning for the river of the Mississippi River
ecosystem. Washington (DC): National Biological Survey, Biological Report 19; p. 350 359.
Meade RH, Moody JA. 2010. Causes for the decline of suspended-sediment discharge in the Mississippi River system,
1940 2007. Hydrol Proc. 24:35 49.
Meek SE. 1892. Notes of the shes of western Iowa and eastern Nebraska. Bul US Fish Comm. 10:217 248.
Moos RE. 1978. Movement and reproduction of shovelnose sturgeon Scaphirhynchus platorynchus in the Missouri
River, South Dakota [PhD thesis]. Vermillion (SD): University of South Dakota.
Murphy BR, Willis DW. 1992. Proper distinction between relative weight and relative condition factor. N Am J Fish
Man. 12:665 666.
Pegg MA, Pierce CL, Roy A. 2003. Hydrological alteration along the Missouri River basin: a time series approach.
J Aquatic Sci. 65:63 72.
Pope KL, Kruse CG. 2007. Condition. In: Guy CS, Brown ML, editors. Analysis and interpretation of freshwater sheries data. Bethesda (MD): American Fisheries Society, p. 423 472.
Porter TK, Mestl GE, Porath MT. 2011. Population Characeristics of athead catsh in channelized and unchannelized reaches of the middle river from 1997 2008. In: Michaletz PH, Travnichek VH, editors. Conservation, ecology, and management of catsh: the second international symposium. Bethesda (MD): American Fisheries
Society, p. 105 118.
Quist MC, Guy CS, Braaten PJ. 1998. Standard weight (Ws) equation and length categories for shovelnose sturgeon.
N Am J Fish Man. 18:992 997.
Raspopov VM. 1987. Fecundity of beluga sturgeon, Huso huso, of the Caspian Sea. J Appl Ichthyol. 27:140 141.
Schrey AW., Heist EJ. 2007. Stock structure of pallid sturgeon analyzed with microsatellite loci. J Appl Ichthyol.
23:297 303.
Schrey AW, Sloss BL, Sheehan RJ, Heidinger RC, Heist EJ. 2007. Genetic discrimination of middle Mississippi River
Scaphirhynchus sturgeon into pallid, shovelnose and putative hybrids with multiple microsatellite loci. Conserv
Genet. 8:683 693.
Shuman DA, Willis DW, Krentz SC. 2006. Application of a length-categorization system for pallid sturgeon (Scaphirhynchus albus). J Freshw Eco. 21:71 76.
Shuman DA, Klumb RA, Wilson RH, Jaeger ME, Haddix T, Gardner WM, Doyle WJ, Horner PT, Ruggles M,
Steffensen KD, et al. 2011. Pallid sturgeon size structure, condition and growth in the Missouri River basin. J Appl
Ichthyol. 27:269 281.
Steffensen KD, Powell LA, Koch JD. 2010. Assessment of hatchery-reared pallid sturgeon survival in the lower
Missouri River. N Am J Fish Man. 30:671 678.
Steffensen KD, Pegg MA, Mestl GE. 2013a. Population characteristics of pallid sturgeon in the lower Missouri River.
J Appl Ichthyol. 29:687 695.
Steffensen KD, Pegg MA, Mestl GE. 2013b. Population prediction and viability model for pallid sturgeon (Scaphirhynchus albus, Forbes and Richardson, 1905) in the lower Missouri River J Appl Ichthyol. 29:984 989.
Steffensen KD, Eder BL, Pegg MA. 2014. Fish community response to oodplain inundation in a regulated river.
J Freshw Eco. 29:413 427.

Downloaded by [Southern Illinois University] at 07:25 09 August 2016

JOURNAL OF FRESHWATER ECOLOGY

13

Steffensen KD, Shuman DA, Klumb RA, Stukel S. 2014. The status of shes in the Missouri River, Nebraska: pallid
sturgeon (Scaphirhynchus albus). Trans Neb Acad Sci. 34:3 15.
Steffensen KD, Shuman DA, Stukel S. 2014. The status of shes in the Missouri River, Nebraska: shoal chub (Macrhybopsis hyostoma), sturgeon chub (M. gelida), sicklen chub (M. meeki), silver chub (M. storeriana), athead chub
(Platygobio gracilis), plains minnow (Hybognathus placitus), western silvery minnow (H. argyritis) and brassy minnow (H. hankinsoni). Trans Neb Acad Sci. 34:49 67.
Tibbs J.E. 1998. Flathead chub (Platygobio gracilis) rangewide status assessment. Columbia (MO): University of
Missouri.
Tripp SJ, Phelps QE, Colombo RE, Garvey JE, Burr BM, Herzog DP, Hrabik RA. 2009. Maturation and reproduction
of shovelnose sturgeon in the middle Mississippi River. N Am J Fish Man. 29:730 738.
US Army Corps of Engineers. 2001. Missouri River mainstem reservoir system master water control manual, Missouri
River Basin. Omaha (NE): US Army Corps of Engineers.
US Fish and Wildlife Service. 2001. Updated status review of Sicklen and sturgeon chub in the United States. Denver
(CO): US Fish and Wildlife Service.
US Fish and Wildlife Service. 2014. Revised recovery plan for the pallid sturgeon Scaphirhynchus albus. Billings (MT):
US Fish and Wildlife Service.
Wanner GA, Shuman DA, Willis DW. 2007. Food habits of juvenile pallid sturgeon and adult shovelnose sturgeon in
the Missouri River downstream of Fort Randall Dam, South Dakota. J Freshw Eco. 22:81 92.
Welker TL, Drobish MR. 2012a. Pallid sturgeon population assessment project, Volume 1.6. Yankton (SD): US Army
Corps of Engineers, Omaha District.
Welker TL, Drobish MR. 2012b. Missouri River standard operating procedures for sh sampling and data collection,
Volume 1.6. Yankton (SD): US Army Corps of Engineers, Omaha District.
Winders KR, Dattilo JE, Huffmon TR, Travnicehek VH. 2014. Season diet composition of juvenile and adult pallid
sturgeon Scaphirhynchus albus (Forbes and Richardson, 1905) in the channelized lower Missouri River. J Appl
Ichthyol. 30:1133 1140.
Winston WE, Criss RE. 2003. Oxygen isotope and geochemical variation in the Missouri River. Environ Geo.
43:546 556.