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Mississippi River Basin Paddlefish Population

Dynamics: Implications for the Management of
a Highly Migratory Species

Article in Fisheries · October 2016

DOI: 10.1080/03632415.2016.1221701

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FEATURE

Mississippi River Basin

Paddlefish Population
Dynamics:
Implications for the Management
of a Highly Migratory Species
Ryan N. Hupfeld*
Southeast Missouri State University, 1 University Plaza, Cape Girardeau, MO 63701. E-mail: ryan.hupfeld@dnr.iowa.gov
Quinton E. Phelps, Sara J. Tripp, and David P. Herzog
Missouri Department of Conservation, Big Rivers and Wetlands Field Station, Jackson, MO

*Present address for Ryan N. Hupfeld: Iowa Department of Natural Resources, 21914 Park Loop, Onawa, IA 51040.

600 Fisheries | Vol. 41 • No. 10 • October 2016

Fisheries | www.fisheries.org 601
Management of migratory fishes is a difficult task for fishery managers because fish cross jurisdictional boundaries and
encounter differing regulations. States within the Mississippi River basin have differing commercial and recreational regula-
tions for riverine Paddlefish Polyodon spathula populations, and the current population dynamics have not been evaluated
on a system-wide basis. The objective of this study was to simulate the current reproductive potential of the population
using various harvest scenarios. At the current exploitation rate of all locations pooled (u = 14.8%), the population may
experience recruitment overfishing at a 610 mm minimum length limit and has the potential to experience recruitment
overfishing at a 710 mm length limit (u = ~20%). Any added stressors (e.g., Asian carp) may cause further mortality; thus,
adopting a precautionary approach is necessary to maintain sustainable Paddlefish fisheries. At an 810 mm length limit,
the population would not realistically experience recruitment overfishing. Implementing a basin-wide minimum length limit
of at least 810 mm on riverine Paddlefish fisheries is suggested. Given the migratory nature of Paddlefish, developing a
form of governance to ensure management strategies are complementary, and a sustainable source of funding is available
is imperative to ensure sustainability. To accomplish this, developing a Mississippi River Basin Interstate Fishery Commis-
sion may be beneficial.

Dinámica poblacional del pez espátula del Mississippi: implicaciones para el manejo de una
especie altamente migratoria
El manejo de peces migratorios es un reto importante para los administradores de recursos ya que los peces cruzan fron-
teras jurisdiccionales y se exponen a diferentes regímenes de regulación. Los estados que comparten la cuenca del Río
Mississippi, poseen diferentes esquemas de regulaciones recreativas y comerciales para el manejo de las poblaciones del
pez espátula, Polyodon spathula, y aun no se ha evaluado la dinámica poblacional vigente a escala de cuenca. El objetivo
de este estudio fue simular el potencial reproductivo actual de la población bajo distintos escenarios de captura. Apli-
cando la tasa de explotación vigente a todas las localidades muestreadas (u = 14.8%), la población pudiera experimentar
una sobrepesca en el reclutamiento, usando una talla mínima de captura de 610 mm, y existe el potencial de que ocurra
sobrepesca en el reclutamiento incluso a una talla mínima de captura de 710 mm (u = ~20%). Cualquier factor adicional
que genere estrés (e.g. la carpa asiática) puede causar una mayor mortalidad; por consiguiente es necesario adoptar un
enfoque precautorio con el fin de mantener sana la pesquería del pez espátula. En términos reales, una talla límite de 810
mm sería suficiente como para evitar una sobrepesca en el reclutamiento. De aquí se recomienda implementar en la pes-
quería del pez espátula, una talla mínima de captura de por lo menos 810 mm en toda la cuenca. En virtud de la naturaleza
migratoria del pez espátula, es necesario desarrollar un plan de manejo a nivel cuenca para asegurar la sustentabilidad del
recurso. Para lograr esto, pudiera ser útil desarrollar un Comisión de Pesca Interestatal de la cuenca del Río Mississippi.

Dynamiques de population du paddlefish dans le bassin du Mississippi: implications pour la

gestion des espèces hautement migratoires
La gestion des poissons migratoires constitue une tâche complexe pour les gestionnaires de pêcheries, car le poisson
traverse les frontières juridictionnelles et est donc soumis à différentes règlementations. Les états qui bordent le bassin du
fleuve Mississippi appliquent différentes règlementations en matière de pêche commerciale et récréative pour les popula-
tions de poissons spatulaire (Polyodon spathula), et les dynamiques de populations actuelles n’ont pas été évaluées sur
tout le système. L’objectif de cette étude consistait à simuler le potentiel de reproduction actuel des populations en ap-
pliquant différents scénarios de pêches. Au rythme d’exploitation actuel de tous les sites sondés (u = 14.8 %), la population
peut s’attendre à une surpêche des recrues à une limite de longueur de 610 mm minimum et a le potentiel pour subir une
surpêche des recrues à une limite de longueur de 710 mm (u = ~20 %). Tout facteur de pression supplémentaire (comme la
carpe asiatique) peut entraîner une mortalité supérieure ; il est donc nécessaire d’adopter une approche précautionneuse
pour maintenir des pêcheries de paddlefish (poisson spatule) durables. A une longueur limite de 810 mm, la population
pourrait ne pas subir de surpêche des recrues. L’application d’une limite de longueur d’au moins 810 mm sur l’ensemble du
bassin sur les pêcheries de paddlefish fluvial est suggérée. En raison de la nature migratoire du paddlefish, il est impératif
de mettre en œuvre un plan de gestion sur tout le bassin, et d’unifier les stratégies de gestion de l’espèce, pour en assurer
la durabilité. Pour cela, il est conseillé de mettre en place une Commission inter-état du bassin du fleuve Mississippi.

INTRODUCTION for many species due to their ability to produce high-quality roe
to make caviar (Pikitch et al. 2005). For example, the Caspian
Many fish stocks and species have experienced excessive
Sea sturgeon harvest decreased from 28,500 metric tons in 1985
fishing pressure, which has subsequently initiated population
to 1,345 metric tons in 2005, largely attributed to overfishing
declines and, in some cases, complete collapse (Hutchings
and bycatch of juveniles and immature fish (Pourkazemi
and Reynolds 2004; Allan et al. 2005; Cooke and Cowx
2006). Furthermore, Pikitch et al. (2005) identified numerous
2006). For example, a recreational Walleye Sander vitreus
major sturgeon fisheries with harvest reports of less than 15%
fishery in Alberta, Canada, experienced declines due to lack
of historic catches, along with 35% of new sturgeon fisheries
of productivity and high fishing pressure (Sullivan 2003).
collapsing within 7–20 years.
Additionally, Hutchings and Myers (1994) attributed the
Another example of a typical K-selected species that
collapse of the Atlantic Cod Gadus morhua population near
has demonstrated declines is Paddlefish Polyodon spathula.
Newfoundland and Labrador solely to overexploitation.
Paddlefish were once abundant throughout the Mississippi
Overexploitation can be especially prominent for K-selected
River basin but have experienced population declines and
species that exhibit late age at maturation and spawning
range reduction due to habitat degradation and overexploitation
periodicity, such as the order Acipenseriformes (sturgeons and
(Combs 1982; Gengerke 1986; Graham 1997; Scholten 2009).
paddlefishes). The order Acipenseriformes is considered one of
Paddlefish is a highly sought-after fish both recreationally (i.e.,
the most endangered groups of fishes in the world (Pikitch et al.
flesh) and commercially (i.e., black caviar) and has experienced
2005). Despite declines, commercial fishing pressure still exists

602 Fisheries | Vol. 41 • No. 10 • October 2016

declines due to exploitation. For example, Scholten and Bettoli and recaptured in the Osage River, Missouri, after crossing at
(2005) determined that the Paddlefish population in Kentucky least five political boundaries (S. Tripp, Missouri Department of
Lake was experiencing growth overfishing and had the potential Conservation, personal communication). Additionally, Pracheil
to experience severe recruitment overfishing, largely due to lack et al. (2012) estimated that 39% of wild Paddlefish movements
of population monitoring and sufficient regulation. Scholten occurred across different state boundaries. S. Tripp (unpublished
and Bettoli (2005:1286) stated, “Under certain conditions, manuscript, Missouri Department of Conservation) determined
Paddlefish can be easily overfished in a commercial gill- that 45% of Paddlefish moved out of the upper Mississippi
net fishery” and “are especially vulnerable to entanglement River with an average distance traveled of approximately 387
gear when they congregate in spawning areas below dams.” km, encompassing numerous rivers (e.g., middle Mississippi
Additionally, bycatch mortality of males and juvenile females River, lower Mississippi River, Ohio River, Illinois River, Des
can be high when water temperatures are warm (>15°C; Bettoli Moines River, Missouri River, and Meramec River) and political
et al. 2007). Pierce et al. (2015) found that less than 15% boundaries. Among the political boundaries mentioned, all have
exploitation substantially increased mortality rates in Lake differing management strategies, minimum length limits, and
Francis Case, South Dakota. Similarly, Donabauer et al. (2009) seasons (Tables 1 and 2). Because fishes cross jurisdictional
indicated that while commercial fishing was occurring, the total boundaries, and thus differing harvest regulations, management
annual mortality rate of Paddlefish in Ozark Lake, Arkansas, frequently becomes a more complicated issue.
was substantially higher than when commercial harvest was There are numerous examples of successful management
absent. Tripp et al. (2012) determined that the lower basin and rehabilitation of migratory fish populations across the
of the Mississippi River Paddlefish population was at risk of world, despite the difficulties associated with interjurisdictional
overfishing at the current minimum length limits. Furthermore, management. For example, the Atlantic States Marine Fisheries
Sharov et al. (2014) considered all Paddlefish stocks overfished Commission, along with state agencies, developed a recovery
based on estimates of fishing mortality. Illegal harvest of plan for the commercially and recreationally overfished stocks
Paddlefish has occurred in recent years, which could lead to of Striped Bass Morone saxatilis (ASMFC 1981; Field 1997;
further overexploitation occurring as well. In March 2013 in Richards and Rago 1999). Multiple parties working together
Missouri, eight individuals were indicted for federal crimes of developed an interstate fisheries management plan that was
illegal trafficking of Paddlefish and their eggs for use as caviar very successful in recovering the Striped Bass population until
following a multiyear investigation by more than 140 state and it was declared fully recovered 14 years after implementation
federal agents (Jerek 2013). The caviar market is a very lucrative
business and has likely increased pressure on Paddlefish within Table 1. Commercial minimum length limits and seasons for Pad-
the Mississippi River basin due to the decline of Eurasion dlefish within the Mississippi River basin.
sturgeon populations and increasing prices of caviar in recent Length
years (Bettoli et al. 2007; Colvin et al. 2013). The closure of the limit
Shovelnose Sturgeon Scaphirhynchus platorynchus commercial River (EFL) Season
fishery on the middle Mississippi River may also cause an Arkansas
increase in harvest pressure for Paddlefish. With an increase in
Mississippi River 890 mm 11/20–4/10
harvest pressure and concerns that Paddlefish stocks are being
overfished in certain areas, it is imperative that Paddlefish Illinois
populations are monitored and sufficient regulations are in place Mississippi River (downstream of Lock
710 mm 10/1–5/31
to prevent further overharvest (Quinn 2009; Colvin et al. 2013). and Dam 19)
Management of highly sought-after fisheries is complicated Ohio River 810 mm 11/1–4/30
and comes with a lot of pressure from many constituents Indiana
(e.g., commercial and recreational fishers, Convention
Ohio River (excluding restricted zones
on International Trade in Endangered Species [CITES]). around five lock and dams)
810 mm 11/1–4/30
Paddlefish regulation has largely been through recreational
Iowa
and commercial minimum length limits and seasons; however,
it is implemented on a state-by-state basis and is inconsistent Mississippi River Closed No season
across the Mississippi River basin. Differences also exist in Missouri River Closed No season
net attendance and reporting requirements, which can lead to Kentucky
incomplete harvest information from state to state. Differing
All waters open to commercial fishing
regulations can make management a more complicated issue, (excluding Kentucky and Barkley lakes)
810 mm 11/1–5/31
especially when dealing with a highly migratory fish species
Mississippi
such as Paddlefish. Conover and Grady (2000), Henley et al.
(2001), and Tripp et al. (2014) determined that Paddlefish Mississippi River 890 mm 11/20–3/31
within the Mississippi River basin crossed multiple state Missouri
boundaries while making long-distance movements. Further, Year-
Stancill et al. (2002) documented a Paddlefish that traveled Mississippi River 610 mm
round
~1,900 km and passed through five dams while crossing at Missouri River Closed No season
least seven political boundaries. A Paddlefish tagged in Moon
Ohio
Lake, Mississippi, traveled ~2,092 km, crossing at least seven
political boundaries before being captured in the Missouri River Ohio River Closed No season
near Levasy, Missouri (G. Lucas, Mississippi Department of Tennessee
Wildlife, Fisheries, and Parks, personal communication), and a
Mississippi River 864 mm 11/15–4/15
Paddlefish was tagged below Gavin’s Point Dam, South Dakota,

Fisheries | www.fisheries.org 603

Table 2. Recreational length limits and seasons for Paddlefish fishes to state agencies (Hooker et al. 2009). This act was put
within the Mississippi River basin. in place to encourage state agencies to support and promote the
Length limit management and research of interjurisdictional fishery resources
River (EFL) Season throughout their range (Hooker et al. 2009).
Arkansas Despite the success of several coastal marine fishery
No length
commissions and the binational Great Lakes Fishery
Mississippi River No season Commission, there are no inland fishery commissions in the
limit
Iowa
United States for interjurisdictional river fisheries. Specifically,
the Mississippi River basin is the largest drainage system in
Maximum: 838
Mississippi River
mm
3/1–4/15 North America and drains more than 40% of the continental
United States (Turner and Rabalais 1991). Within this
Protected slot:
Missouri River
890–1,143 mm
3/1–4/15 extensive system, there are 31 states with fishery management
authority. Furthermore, 91 fish species have been identified by
Illinois
the Mississippi Interstate Cooperative Resource Association
Mississippi River (between Maximum: 838
3/1–4/15 (MICRA) as interjurisdictional species, yet no fishery
Iowa and Illinois) mm
commission has been developed to help manage these highly
Mississippi River (between Minimum: 610 9/15–12/15, 3/15– migratory species. Without an inland fishery commission and
Missouri and Illinois) mm 5/15
thus a unified basin-wide management plan, management of
Indiana interjurisdictional species, such as Paddlefish, on a state-by-
Ohio River Closed Closed state basis differs. There have been efforts to address these
Kentucky
differing management strategies through MICRA. MICRA is a
cooperative agreement of 28 state natural resource departments.
No length
Mississippi River
limit
2/1–5/10 MICRA was organized in 1991 as a partnership to improve
management of interjurisdictional fish in the Mississippi River
Mississippi
basin. MICRA’s mission is “to improve the conservation,
Mississippi River
Maximum: 762
Year-round management, development, and utilization of interjurisdictional
mm fishery resources (both recreational and commercial) in the
Missouri Mississippi River basin through improved coordination and
Minimum: 610 communication among the responsible management entities”
Mississippi River 3/15–4/30
mm (Mississippi Interstate Cooperative Resource Association n.d.).
Minimum: 610 In 1992, MICRA developed a Paddlefish/Sturgeon
Missouri River 3/15–4/30
mm Committee, with one of the major goals being to coordinate and
Ohio develop complementary management strategies and regulations
for Paddlefish and sturgeon in states throughout the Mississippi
Ohio River Closed No season
River basin (Grady et al. 2005; Scholten 2009). From 1995 to
Tennessee 2004, MICRA initiated a basin-wide Paddlefish stock assessment
Mississippi River
No length
4/24–5/31
project (Grady et al. 2005). This study included multiple
limit state agencies (22 states) to assess habitat use, distribution,
movements, and extent of harvest and exploitation of Paddlefish
(Richards and Rago 1999). Lake Sturgeon Acipenser fulvescens (Grady et al. 2005). This was the largest multiagency study that
populations in Wisconsin have been managed since 1993 under has been attempted within the Mississippi River basin. Although
a rehabilitation plan, developed with tribal, state, and federal collaboration with other state agencies was highly successful,
representatives, called the Menominee Reservation Lake there are still many needs that should be addressed that were
Sturgeon Enhancement Committee (Runstrom et al. 2002). In outside the scope of the project (Grady et al. 2005). These efforts
2001, the first successful Lake Sturgeon natural reproduction conducted by MICRA have been a major step in improving
was documented in 50 years and was an important step in management of Paddlefish populations within the Mississippi
reaching one of the goals of the Menominee Reservation River basin. However, the current population dynamics (e.g.,
Sturgeon Restoration Plan in establishing self-sustaining recruitment, growth, and mortality) and harvest regulations for
Lake Sturgeon populations (Runstrom et al. 2002). As the Paddlefish have not been evaluated on a system-wide basis.
aforementioned studies suggest, large-scale, interjurisdictional These dynamic rate functions enable fisheries managers to detect
management is imperative in restoring and providing long-term changes at the population level due to fishing mortality and
sustainable fisheries for any fish species that cross jurisdictional are especially important in determining changes in Paddlefish
boundaries. Numerous fisheries commissions have been populations to prevent overharvest. As the aforementioned
put in place to help conserve and manage highly migratory studies suggest, Paddlefish have great potential to experience
fish stocks across the world (e.g., the National Oceanic and overexploitation due to their life history characteristics and
Atmospheric Administration’s Fisheries West Coast Regions have been overharvested in the past (Scholten and Bettoli 2005;
Highly Migratory Species Program, International Commission Bettoli et al. 2007; Tripp et al. 2012; Sharov et al. 2014). Due
for the Conservation of Atlantic Tunas, Pacific States Fisheries to international trade of Paddlefish caviar and the listing of the
Marine Commission, and Great Lakes Fishery Commission) species as a CITES Appendix II species, Paddlefish populations
and have been highly successful in rebuilding and maintaining are now under scrutiny. In order to provide the U.S. Fish and
fish stocks. Furthermore, the Anadromous Fish Conservation Wildlife Service with the data necessary to make nondetriment
Act of 1965 and the Interjurisdictional Fisheries Act of 1986 findings required for CITES Paddlefish export permits, intensive
provide financial assistance for research of interjurisdictional management to understand the Paddlefish population status

604 Fisheries | Vol. 41 • No. 10 • October 2016

and harvest regulations effectiveness is needed. As such, the
objective of this study was to assess the population dynamics for
Paddlefish populations in the upper Mississippi River, middle
Mississippi River, lower Mississippi River, Missouri River,
and Ohio River and simulate the reproductive potential of the
populations under various harvest scenarios. Ultimately, this
study will provide the information necessary for management
agencies to detect changes in Paddlefish populations and adopt
necessary changes to prevent overexploitation throughout the
Mississippi River basin.

METHODS
Information on Paddlefish populations were obtained from
the upper Mississippi River (river kilometer [rkm] 587–322),
middle Mississippi River (rkm 313–0), lower Mississippi
River (rkm 1,535–1,153), Ohio River (rkm 1,160–1,579), and
Missouri River near the Osage River confluence (rkm 233–0)
during the spring of 2013 (Figure 1). Sampling locations were Figure 1. Map of study area where Paddlefish were collected dur-
ing the spring of 2013. Locations included in the study are the up-
stratified by location (river reach) and points were randomly per Mississippi River (rkm 587–322), middle Mississippi River (rkm
selected. Paddlefish were collected using 127-mm bar-mesh 313–0), lower Mississippi River (rkm 1,535–1,153), Ohio River (rkm
floating monofilament gill nets. Gill nets were hobbled, 1,160–1,579), and Missouri River near the Osage River confluence
(rkm 233–0).
reducing net height from 9.1 to 6.1 m. Gill nets were monitored
continuously, and fish were removed immediately. All Paddlefish
potential ratio was simulated for various exploitation rates
were weighed (g), eye-to-fork length (EFL) was measured to
(i.e., 0%–95%) for fish of varying minimum length limits (e.g.,
the nearest millimeter, and maturation stage was recorded (Fv,
610, 710, 810, 864, and 890 mm). Exploitation was derived
F1, F2, F3, F4, F5, F6 and Mv, M1, M2,, M3; Colombo et al. 2007;
for each location by using calculated parameters: S (annual
Tripp et al. 2009). Maturation stage Fv was determined by small,
survival), M (instantaneous mortality rate), and Z (slope of
well-ordered ovarian folds with small amounts of fat (virgin
weighted catch curve regression). A 610-mm length limit was
female), F1 by the presence of ovarian folds with large amounts
simulated because that is the current minimum length limit in
of fat, F2 by the presence of white to yellow small oocytes, F3
Missouri. Length limits of 710, 810, 864, and 890 mm were
by the presence of oocytes that are light green and yellow in
simulated because these are the current minimum length limits
color, F4 by the presence of large oocytes that are black in color,
that other state agencies within the Mississippi River basin have
F5 is an actively spawning female, F6 by having translucent
employed. Spawning potential ratio is used to determine the
ovaries (spent female), Mv by the presence of pink ribbon-
critical number of adults needed to maintain recruitment in the
like testes with small amounts of fat (virgin male), M1 by the
population and to provide an idea of the reproductive potential
presence of yellow testes with large amounts of testicular fat,
of the population (Goodyear 1993). Spawning potential ratio
M2 by the presence of large pink testes, and M3 by the presence
values can range between 0 and 1 and decrease as higher total
of compressed pink testes (spent male; Colombo et al. 2007;
annual mortality rates reduce reproductive output. An SPR
Tripp et al. 2009). Paddlefish at maturation stages M2 and F4
level of 0.4 (i.e., 40%) was used because it has been considered
were considered reproductively viable (i.e., fish that have the
a conservative target threshold for recruitment overfishing
potential to spawn within the sampling year; Hupfeld and Phelps
in less resilient fish populations (i.e., sturgeon species and
2014). The entire female gonad was removed and weighed to
Paddlefish; Goodyear 1993; Hupfeld et al. 2014; Sharov et al.
the nearest gram, and a 50-g subsample of eggs was weighed
2014). The model requires input parameters of recruitment,
and counted to estimate absolute fecundity (Reed et al. 1992).
growth, mortality, maximum age, weight-to-length relation,
A 2-cm section of the lower left dentary bone was removed,
fecundity, and maturation schedule (Table 3; Slipke et al. 2002).
cleaned, and dried. Multiple thin sections were obtained using
Conditional natural mortality was calculated using the Hoenig
a Buehler Isomet low-speed saw using methods explained by
method (Hoenig 1983). Recruitment was held constant. Growth
Scarnecchia et al. (1996). Dentary sections were examined
was assessed using a von Bertalanffy model, using age data
independently by two readers under a dissecting microscope,
coupled with length at capture to estimate overall mean length
and age was estimated by counting annuli along the mesial arm
at age. Differences in growth among locations were analyzed
of the section. If differences in age estimate existed between
using the residual sum of squares (Ratkowsky 1983; Chen et
readers, the structure was viewed again by both readers to
al. 1992). Sexes were lumped for all simulations conducted
resolve discrepancies. If a consensus could not be ascertained,
because the fishery is not managed for one sex alone (e.g., males
the sample was discarded. Age determination using the lower
are harvested for flesh and females are harvested for eggs and
dentary bone is well established; however, the underestimation
flesh). Additionally, males are captured as bycatch and have the
of older individuals is possible (Scarnecchia et al. 2006; Pierce
potential to sustain postrelease delayed mortality due to warm
et al. 2011; Sharov et al. 2014).
water temperatures, being checked for eggs by incision, and
Using the yield per recruit modeling option in Fisheries
injuries in fishing gear. However, a conservative management
Analysis and Modeling Simulator, the spawning potential ratio
approach may be needed due to suggested minimum length
(SPR) was modeled for each Paddlefish sampling location
limits potentially being low for a commercial roe fishery on
to evaluate the effectiveness of minimum length limits on
females. Total annual mortality rates were assessed using catch
the population status (Slipke and Maceina 2010). Spawning
curve analyses. Weighted regressions were used to reduce bias

Fisheries | www.fisheries.org 605

Table 3. Life history parameters of Paddlefish populations used for spawning potential ratio modeling from the upper Mississippi River
(UMR), middle Mississippi River (MMR), lower Mississippi River (LMR), Ohio River (OHR), Missouri River (MOR), and pooled locations.

Parameter Description UMR MMR LMR OHR MOR Pooled

B0 Intercept a
−4.10 −4.83 −5.19 −5.57 −7.34 −4.70
B1 Slopea 2.74 2.97 3.11 3.25 3.87 2.94
L∞ Theoretical maximum length at time = ∞ (mm)b 1,382.61 1,301.56 1,143.15 1,051.90 1,156.40 1,099.60
t0 Theoretical time when length = 0 (years) b
−0.165 −0.132 0.374 0.155 −0.890 −0.018
K Growth coefficientb 0.070 0.088 0.117 0.127 0.084 0.109
Agemax Maximum age observed 22 18 22 18 23 23
Maturation Age at sexual maturation 11 10 9 10 10 10
Cm Conditional natural mortalityc 0.17 0.21 0.17 0.21 0.17 0.17
% Females Percentage of fish that are females 56 50 50 47 49 50
Fecundity : EFL Interceptd −0.776 −13.398 −4.262 −0.995 −1.909 −0.903
Fecundity : EFL Sloped 2.054 6.291 3.223 2.103 2.458 2.083
Annual ­mortality Total annual mortality (%) 31.4 26.0 33.7 31.3 29.8 34.5
a
Parameters of the log10-transformed length : weight regression.
b
Parameters of the von Bertalanffy growth curve.
c
Calculated by the Hoenig method.
d
Parameters of the log10-transformed fecundity : EFL regression.

associated with weak or missing year classes in

the population (Ricker 1975; Slipke and Maceina
2000).

Figure 2. Length–frequency distributions of Paddlefish sampled in the upper
Mississippi River, middle Mississippi River, lower Mississippi River, Ohio River,
and Missouri River in 2013, with 127-mm bar-mesh floating gill nets. Length dis-
tribution indices include number of fish (N), range of lengths, and mean lengths
(SE).
RESULTS
A total of 719 Paddlefish were collected
throughout the study. Lengths ranged from 226
to 1,105 mm EFL and ages ranged from 2 to 23
years (Figures 2 and 3). Across all locations,
approximately 33% of Paddlefish collected were
reproductively viable females, similar to Tripp et
al. (2012). Paddlefish EFL (log10 transformed) was
an adequate predictor of Paddlefish weight at all
locations (P < 0.0001, r2 > 0.80). Mean length at
age for all aged fish was sufficiently described by
von Bertalanffy growth functions at all locations
(P < 0.0001, r2 > 0.98). Growth rates were
similar among locations (P > 0.005). Total annual
mortality rates between locations appeared similar.
Specifically, the upper Mississippi River had an
annual mortality rate of 31.4%, middle Mississippi
River 26%, lower Mississippi River 33.7%,
Missouri River 29.8%, and Ohio River 31.3%.
Exploitation (u) rates among locations appeared
similar, with the upper Mississippi River having
an exploitation rate of 15.8%, middle Mississippi
River 20.1%, lower Mississippi River 15.6%, Ohio
River 19.3%, and Missouri River 15.6%.
In the upper Mississippi River (e.g., u =
15.8%), SPR estimates indicated that at the
current minimum length limit (MLL) in Missouri
(610 mm), the Paddlefish population is likely
experiencing recruitment overfishing; however,
at the current MLL in Illinois (710 mm), the
population would not experience recruitment
overfishing until approximately 18% exploitation
occurred (see Figure 4 for SPR simulations for
each location). At a conservative MLL of 810
mm, the upper Mississippi River population

606 Fisheries | Vol. 41 • No. 10 • October 2016

would not realistically experience recruitment
overfishing. In the middle Mississippi River (e.g.,
u = 20.1%), the Paddlefish population is likely
experiencing recruitment overfishing at 610
and 710 mm MLLs. At an MLL of 810 mm, the
population would likely not experience recruitment
overfishing until approximately 31% exploitation.
In the lower Mississippi River (e.g., u = 15.6%),
the population may currently be experiencing
recruitment overfishing at the 610 mm MLL;
however, at an MLL of 710 mm the population
would not experience recruitment overfishing until
approximately 18% exploitation occurred. At a more
conservative MLL of 810 mm, the lower Mississippi
River population would not realistically experience
recruitment overfishing. In the Ohio River (e.g.,
u = 19.3%), the population would not experience
recruitment overfishing until approximately 22%
exploitation occurred at the current MLL of 710
mm in Illinois. At the current MLL of 810 mm in
Kentucky, the population would not realistically
experience recruitment overfishing. In the Missouri
River (e.g., u = 15.6%), the population may currently
be experiencing recruitment overfishing at the
current 610 mm MLL. At a 710 mm MLL, the
Missouri River population would not experience
recruitment overfishing until approximately 26%
exploitation and would not realistically experience
recruitment overfishing at an 810 mm MLL. At
the current exploitation rate of all locations u =
14.8%, the population may experience recruitment
overfishing at a 610 mm MLL (Figure 5). At a 710
mm MLL, the population would not experience
recruitment overfishing until an approximately 20% Figure 3. Age–frequency distributions of Paddlefish sampled in the upper Mis-
sissippi River, middle Mississippi River, lower Mississippi River, Ohio River, and
exploitation rate. At an 810 mm, MLL the population Missouri River in 2013, with 127-mm bar-mesh floating gill nets. Age distribu-
would not realistically experience recruitment tion indices include number of fish (N), catch–curve regression (r2), signifi-
overfishing at current exploitation rates. cance of regression (P), and total annual mortality (A).

DISCUSSION in the lower Tennessee River may have been experiencing

This study provides the first evaluation of the dynamic
rate functions and potential length limit guidance on a large
scale to prevent overexploitation on the riverine Paddlefish
populations within the Mississippi River basin. Annual mortality
rates in this evaluation appeared similar to those found in the
lower Mississippi River (34% and 38%; Tripp et al. 2012)
and Wisconsin River (26.7%; Runstrom et al. 2001) but were
substantially lower than the intensively fished lower Tennessee
River (68%; Scholten and Bettoli 2005). Growth rates appeared
similar to those found in the lower Mississippi River (Tripp et al.
2012) and lower Tennessee River (Scholten and Bettoli 2005).
Numerous studies have demonstrated that Paddlefish are
susceptible to overharvest and, in some cases, may deplete the
population completely (Stockard 1907; Coker 1923; Larimore
1950). There are concerns that Paddlefish populations within
the Mississippi River basin may be near or are currently
experiencing growth and recruitment overfishing. For example,
Sharov et al. (2014) determined that all Paddlefish stocks may
currently be enduring overfishing. In addition, Tripp et al. (2012)
determined that the minimum length limits within the lower
Mississippi River may not prevent recruitment overfishing at the
current minimum length limit of 610 mm in Missouri. Scholten
and Bettoli (2005) also suggested that the Paddlefish population
growth and recruitment overfishing at the minimum length
limit of 864 mm. Similarly, based on SPR simulations, riverine
Paddlefish populations within the study area may be near, or are
currently experiencing, recruitment overfishing in most locations
at the current length limits.
Paddlefish populations have experienced population
declines due to numerous other stressors such as habitat
degradation, pollution, sedimentation, and fragmentation (e.g.,
construction of dams; Combs 1982; Gengerke 1986; Graham
1997; Scholten 2009). Though Paddlefish have withstood
many of these stressors, the uncertainties of how Silver Carp
Hypophthalmichthys molitrix and Bighead Carp H. nobilis could
affect Paddlefish populations should be of concern due to their
ability to alter the zooplankton community (Sass et al. 2014).
For example, Irons et al. (2007) determined that Bighead and
Silver carp may be negatively influencing body condition of
native planktivores (e.g., Bigmouth Buffalo Ictiobus cyprinellus
and Gizzard Shad Dorosoma cepedianum) in the Illinois River.
Additionally, Schrank et al. (2003) determined that Paddlefish
growth could be negatively influenced by Bighead Carp when
resources are limited. Due to the difficulties and costs with
managing the aforementioned stressors and controlling Silver
and Bighead carp, fishery managers need to monitor Paddlefish

Fisheries | www.fisheries.org 607


Figure 4. Spawning potential ratio models for the upper Mississippi River,
middle Mississippi River, lower Mississippi River, Ohio River, and Missouri River
under five simulated minimum length limits (610, 710, 810, 864, and 890 mm).
Spawning potential ratio of 40% (dashed line) indicates a conservative target
threshold for recruitment overfishing in less resilient fish populations (i.e.,
sturgeon species and Paddlefish). Vertical solid line indicates current exploi-
tation rate at each location.
populations within the Mississippi River basin and have
appropriate regulations to ensure sustainability.
Currently, commercial and recreational regulations across
state boundaries within the Mississippi River basin differ. There
are numerous instances where contiguous states within the
Mississippi River basin have differing management regulations.
For example, Missouri and Illinois, which share boundaries
along the Mississippi River, have differing regulations despite
managing the same Paddlefish population. In areas where
conservation agencies have minimum length limits in place
that may precede maturation, it may have deleterious effects
on Paddlefish population basin-wide. Because of this, the
population dynamics of Paddlefish within the Mississippi
River basin may be affected through cumulative harvest. State
conservation agencies with low minimum length limits may be
benefiting to the detriment of other states with higher minimum
length limits (e.g., larger Paddlefish crossing state boundaries
with a high minimum length limit to a state with a lower
minimum length limit). Because minimum length limits of 610
and 710 mm may jeopardize the interconnected basin-wide
population at current exploitation rates, implementing a basin-
wide commercial and recreational minimum length limit of at
least 810 mm to prevent recruitment overfishing from occurring
608 Fisheries | Vol. 41 • No. 10 • October 2016
within the Mississippi River basin is suggested, while
still allowing state agencies to have more conservative
but not less restrictive regulations (e.g., Mississippi
and Arkansas currently have an 890 mm commercial
fishing length limit). This would also ensure that state
agencies that have greater protection of Paddlefish
do not lose the benefit of their regulations to states
with less-restrictive length limits. Given the growing
demand of caviar, increases in illegal harvest,
recently established competition with invasive species
(e.g., Bighead and Silver carp), and the increasing
popularity of recreational fishing, it is essential that
regulations are in place to maintain sustainable harvest
for future generations.
Fishery commissions have adopted an approach
where multiple entities collaborate to develop a
unified management strategy to set regulations and
make recommended habitat improvements and
have been quite successful. In the case of our study,
despite knowing that there is an issue with managing
interjurisdictional fish species on a state-by-state basis,
state agencies are hesitant to give up authority on
Paddlefish fisheries. This could be due to the potential
loss of economic benefits (e.g., license sales), as well
as concerns about losing input on the management of
the fishery. Pracheil et al. (2012) suggested a similar
approach to manage Paddlefish through swimways
made up of different river basins. These data were
based on mark–recapture data, and management
zones were set up based on these movements. Though
these data are very useful, the interbasin movement
patterns were likely underestimated due to tagging
and sampling efforts being unevenly distributed
among states. Pracheil et al. (2012) have further
explained Paddlefish movements on a larger scale
than most literature available. However, Paddlefish
movement data are still lacking at the population level
and at all life stages. There are no data examining
Paddlefish movement patterns from larval and juvenile
individuals. Knowing that movement data are lacking, fishery
managers should be hesitant to set up boundaries based on
Pracheil et al.’s (2012) suggestions. Managing Paddlefish based
on boundaries will still have individuals crossing jurisdictional
boundaries, and management will likely be faced with similar
issues as today (e.g., state-by-state management). Though there
are likely areas where management boundaries for Paddlefish
would need to be developed (e.g., Lock and Dam 19, Keokuk,
Iowa; Bagnell Dam, Bagnell, Missouri), further understanding
Paddlefish movement and river usage at all life stages is
imperative before developing boundaries and implementing
appropriate management actions. Thus, developing a basin-
wide fishery commission could be very beneficial. A fishery
commission would develop a unified management approach
across the entire range of Paddlefish. A commission would
allow fishery managers to develop research projects to evaluate
movement patterns in order to develop management boundaries,
set appropriate complementary regulations, have a centralized
database, and make appropriate habitat modifications/
rehabilitations to overall better manage Paddlefish. Sharov et
al. (2014) called Paddlefish fisheries data poor. In past years,
federal funding has been very limited to promote collaborative
Mississippi River basin projects; this could be an opportunity
to develop a funding source to better understand Paddlefish
populations and implement recommendations and suggestions
from Sharov et al. (2014). Managing Paddlefish on a population-
relevant scale and continuing to develop interjurisdictional
management may not only benefit the Paddlefish populations,
and thus the commercial and recreational fisheries, but could
also help to relieve responsibilities associated with CITES on
the management of Paddlefish. Although interjurisdictional
management can be a daunting task, managing highly
migratory fishes on a large scale is imperative to maintaining
sustainable fisheries. As previous studies throughout this article
demonstrate, successful management of fishes on a population
relevant scale, with multiple governing bodies, has been
accomplished in recent years across the world (ASMFC 1981;
Field 1997; Richards and Rago 1999; Runstrom et al. 2002;
Grady et al. 2005).
This study provides a basis for states to select initial metrics
and targets for sustainable management of Paddlefish. This is
especially important when fishing mortality can have a large
influence on the reproductive potential of Paddlefish populations
(Boreman 1997). Thus, harvest regulations and management
strategies need to be complementary among adjacent state
agencies that share this resource. Given the results uncovered
in this study, and to ensure sustainability, it is suggested that
Paddlefish management in these large, interconnected rivers be
transformed from state agencies only working cooperatively
with each other to considering the development of a formal
arrangement (governance) across the entire Mississippi River
basin where Paddlefish occur. Currently, states do not have the
financial resources needed to adequately manage commercially
and recreationally exploited populations. A form of governance,
or fishery commission, may be an avenue to provide sustainable
funding in the future to state agencies.
Future research should focus on movement patterns of
Paddlefish on a population level at all life stages to better
understand where management zones could potentially be
implemented and to confirm exploitation rates in these large
river environments. Additionally, population dynamics of
Paddlefish populations in the northern and southern portions
of the Mississippi River basin need to be further evaluated to
determine whether regulations are sufficient in those areas as
well. To accomplish this, developing a Mississippi River Basin
Interstate Fishery Commission, similar to the Great Lakes
Fishery Commission, could be beneficial. A fishery commission
could help with funding, developing formal inter-state
agreements on management objectives and evaluation, sharing
data, and establishing complementary regulations/enforcement.
ACKNOWLEDGMENTS
We thank Greg Conover and John Scheibe for comments that
greatly improved this article. We also thank Tom Stefanavage
and Les Franklin for assistance in data collection.
FUNDING
This study was funded in part by the U.S. Army Corps of
Engineers’ Upper Mississippi River Restoration–Environmental
Management Program’s Long Term Resource Monitoring
element implemented by the U.S. Geological Survey, Upper
Midwest Environmental Sciences Center, and by the Missouri
Department of Conservation through a Cooperative Agreement
with Southeast Missouri State University.
Figure 5. Spawning potential ratio model of all locations pooled
(i.e., the upper Mississippi River, middle Mississippi River, lower
Mississippi River, Ohio River, and Missouri River) under five
simulated minimum length limits (610, 710, 810, 864, and 890
mm). Spawning potential ratio of 40% (dashed line) indicates a
conservative target threshold for recruitment overfishing in less
resilient fish populations (i.e., sturgeon species and Paddlefish).
Vertical solid line indicates current exploitation rate.
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