203
American Fisheries Society Symposium 66:203–221, 2009
© 2009 by the American Fisheries Society
Harvest of Paddlefi sh in North America
JEFFREY W. QUINN*
Arkansas Game and Fish Commission
2 Natural Resources Drive, Little Rock, Arkansas 72205, USA
Abstract.—Paddlefi sh Polyodon spathula have been intensively harvested
in both sport and commercial fi sheries. Recent harvests (2000–2006) were
surveyed from state agencies and compared to historical harvests (1965–
1975). Seven major sport fi sheries had recent annual harvests greater than
1,000 fi sh, and most large sport fi sheries appeared to have sustainable har-
vests due to intensive management. Recent commercial harvest was greater
than sport harvest across the species’ range. Most of the commercial harvest
was from Arkansas, Kentucky, and Tennessee. Annual commercial harvest
from the Ohio River increased from 6,000 to 196,000 kg from 1965–1975 to
2000–2006. Annual harvest remained substantial from the Arkansas River
(37,000 kg), the lower Tennessee River (121,000 kg), and the Mississippi River
(103,000 kg). Harvests of paddlefi sh (sport and commercial) compiled from
the literature were highly variable and ranged between 0.01 and 5.06 fi sh/
ha and 0.04–43.43 kg/ha (median = 0.12 fi sh/ha, 1.73 kg/ha). Stock depres-
sion has been associated with a fi rst-year harvest as low as 1.46 kg/ha, and
harvests greater than 5 kg/ha were usually associated with overfi shing
or opening a previously closed fi shery. Case histories from the Tennessee
and Ohio River systems documented that paddlefi sh were susceptible to
overharvest in lentic waters and river reservoirs, but the threat posed by
commercial harvest from large rivers will remain unresolved until more
fi sheries-independent data becomes available. Anthropogenic alterations
to habitat, overreliance on harvest data, and lack of fi sheries-independent
data limit our historical understanding of the degree of threat that harvest
is to paddlefi sh populations.
* Corresponding author: [email protected].us
Introduction
Harvest is a threat to paddlefi sh Polyodon
spathula populations throughout the spe-
cies’ range because they are harvested for
recreation or commercially for the roe or
fl esh. Overfi shing is a global problem for
marine (Sissenwine and Rosenberg 1993)
and freshwater fi sheries (Allen et al. 2005).
Early paddlefi sh researchers documented
high susceptibility of paddlefi sh to commer-
cial seines in oxbow lakes, and these fi sher-
ies were often depleted by overharvest (e.g.,
Stockard 1907; Coker 1923; Larimore 1950).
Multiple commercial fi sheries have been
overexploited to the extent that they were
no longer commercially viable (e.g., Norris
Lake, Lake Cumberland, and Old Hickory
Reservoir; Pasch and Alexander 1986).
Global harvest of paddlefi sh and sturgeon
has been reported to have declined since
1960 due to the collapse of Caspian Sea stur-
geon fi sheries (Pikitch et al. 2005), and sev-
eral closely related sturgeon species are now
rare or endangered (Birstein et al. 1997).
204 QUINN
The goal of this chapter is to review
harvest of paddlefi sh in North America.
The specifi c objectives were to (1) survey
recent (2000–2006) commercial and sport
harvests and compare the results with past
surveys (1965–1975; Carlson and Bonis-
lawsky 1981), (2) compare harvests with
density and biomass estimates to better
understand the limits of paddlefi sh har-
vest, (3) and review select commercial fi sh-
ing case-history studies. My intent was to
improve understanding of what may con-
stitute a sustainable harvest while limiting
the amount of information repeated from
previous reviews (e.g., Carlson and Bonis-
lawsky 1981; Gengerke 1986; Pasch and Al-
exander 1986; Graham 1997; Mosher 1998;
Todd 1998; CITIES 2000; Jennings and
Zigler 2000; Williamson 2003; McDougal
2005; Pikitch et al. 2005).
Commercial Fisheries
Modern commercial paddlefi sh fi sher-
ies used large-mesh gill and trammel nets
(127–152 mm bar mesh) to selectively tar-
get gravid female paddlefi sh for the roe
due to its high value. As of 2006, only six
states allowed commercial harvest of pad-
dlefi sh (Arkansas, Illinois, Indiana, Ken-
tucky, Missouri, and Tennessee). Missis-
sippi recently allowed commercial harvest
with restrictions designed to prevent roe
harvest (i.e., season was open from May 1
to October 31), but 2006 was the last year
Mississippi allowed harvest during the
summer months. Sport and commercial
harvest data were unavailable from Missis-
sippi (D. Riecke, Mississippi Department
of Wildlife, Fisheries, and Parks, personal
communication).
From 1894 to 1975, paddlefi sh com-
mercial harvest declined in the Missis-
sippi, Missouri, Ohio, and Red rivers and
increased in the impounded Tennessee,
Arkansas, and Cumberland rivers (Carlson
and Bonislawsky 1981). Peak harvest from
the Mississippi River was estimated at
853,000 kg during 1899 (Carlson and Bon-
islawsky 1981), which refl ects the massive
seining operations for fl esh harvest that
depleted numerous oxbow lakes (Stockard
1907; Hussakof 1911; Coker 1923; Larimore
1950). Sedimentation of aquatic habitats
was likely related to declining commercial
catches of paddlefi sh for the Missouri Riv-
er (Funk and Robinson 1974).
Commercial Harvest Survey
Methods
Commercial harvest data were surveyed
from appropriate state agencies from 2000
to 2006. Data reporting standards were not
consistent among states with commercial
harvests (Scholten 2009, this volume). Ma-
jor reporting issues included (1) reporting
period was not standard among states, (2)
some states did not collect either number
(e.g., Illinois) or weight of fi sh landed (e.g.,
Arkansas), and (3) weight of fi sh and roe
harvested was reported as processed (e.g.,
dressed fl esh, screened roe) or unpro-
cessed weights (whole fi sh and egg sack
weights).
The effect of nonstandardized report-
ing period was minimized by organizing
data so that large spring harvests were
synchronized for each year. Missouri and
Illinois reported harvest by calendar year
(January 1 to December 31), Kentucky by
license year (March 1 to February 28), and
Arkansas, Indiana, and Tennessee report-
ed by harvest season. Arkansas considered
harvest from the 2000–2001 roe season
(November to April) to be the 2001 har-
vest year, whereas Tennessee considered
it the 2000 harvest year. For this paper, the
harvest year was standardized as July 1 to
June 31, and the year designation followed
the Arkansas convention of the year being
assigned the later date of the harvest year.
Thus, harvest from July 1, 2005 to June 30,
2006 was designated the 2006 harvest year
and had spring harvests that were com-
205HARVEST OF PADDLEFISH
parable with
those from the 2006 calendar
year. Monthly harvest data were obtained
from Kentucky, and harvest was compiled
so that harvest year was reported in this
paper instead of license year.
Mean weight was used to calculate ei-
ther harvest by weight (Arkansas) or by
number (Illinois, Kentucky, and Missouri)
because of inconsistent reporting stan-
dards. Appropriate mean weights were
obtained from state agency biologists.
Mean weight for the Arkansas River was
13.6 kg (Quinn et al. 2009, this volume).
Mean weight of paddlefi sh harvested from
the Ohio River in Indiana (7.3 kg; T. Stefa-
navage, Indiana Department of Natural
Resources, personal communication) was
used to calculate fi sh harvested from Ken-
tucky because most of Kentucky’s harvest
came from the Ohio River. Mean weight of
paddlefi sh harvested from Illinois (11.4 kg)
was applied to Missouri data because both
states have harvests that were largely from
the Mississippi River. Harvest by number
(and calculated weight) for Arkansas was
likely underestimated but did refl ect all the
gravid females harvested.
States with commercial harvest gen-
erally reported whole fl esh weights (e.g.,
Illinois, Indiana, Kentucky, and Missouri)
of fi sh landed, but Tennessee collected
dressed weight of fi sh harvested. Esti-
mates of whole fl esh weights landed from
Tennessee were obtained by summing
the estimated weight of each harvested
fi sh (G. Scholten, Tennessee Wildlife Re-
sources Agency, personal communication).
Weight was estimated from the reported
eye-to-fork length (EFL) using an observed
length–weight relationship for Kentucky
Lake (Scholten and Bettoli 2005). Most
states reported processed roe weights (e.g.,
Arkansas, Illinois, and Tennessee), but Ken-
tucky and Indiana reported unprocessed
roe weights. Conversion data were not
available for the Ohio River, so the conver-
sion for the Yellowstone/Sakakawea stock
was used (processed roe weight was 62%
of the unprocessed roe weight; V. Riggs,
Montana Department of Fish, Wildlife, and
Parks, unpublished data).
Commercial Harvest Results
Mean annual commercial fl esh harvest of
paddlefi sh in North America was 487,600
kg from 2000 to 2006 (Table 1), and the
annual mean number of paddlefi sh com-
mercially harvested was 47,400 fi sh (Table
2). Estimated mean annual processed roe
harvest from North America was 25,380 kg
from 2000 to 2006 (Table 3).
Mean annual commercial harvest from
2000 to 2006 was much higher than reports
from 1965 to 1975 (260,000 kg; Carlson
and Bonislawsky 1981) and 1992 (223,000–
269,000 kg; Graham 1997). Although the re-
cent (2000–2006) average was only 44% of
peak fl esh harvest from 1899 (1,105,000 kg;
Carlson and Bonislawsky 1981), recent har-
vest appeared substantial because (1) only
six states were open and reported harvest
(compared to 11 states in 1984; Gengerke
1986), (2) some waters were closed in states
open to commercial fi shing (e.g., Red River,
Arkansas), (3) several states had roe fi sher-
ies skewed towards gravid females (e.g.,
Arkansas, Tennessee), (4) and large com-
mercial seining operations were no longer
a commonly used harvest method. Recent
(2000–2006) mean commercial harvest was
104% of the 1894 harvest, 77% of the 1922
harvest, and 113% of the 1931 and 1975
harvests from Carlson and Bonislawsky
(1981).
The majority of the commercial paddle-
fi sh harvest came from the states of Arkan-
sas, Kentucky, and Tennessee (Tables 1–3).
Arkansas and Tennessee led the nation in
annual processed roe harvested at slightly
greater than 8,000 kg. Tennessee’s harvest
came primarily from Kentucky Lake (Ten-
nessee River; 80%) and the Mississippi Riv-
er (17%). Eighty-three percent of the harvest
206 QUINN
Table 1. Commercial fl esh harvests (kg whole weight) of paddlefi sh in six states with an
open season from 2000 to 2006. Year was based on (1) calendar year for Illinois and Mis-
souri, and (2) harvest season for other states (e.g., July 1, 2000 to June 30, 2001 is de-
fi ned as the 2001 year). Harvests were unavailable for Mississippi.
State 2000 2001 2002 2003 2004 2005 2006 Mean
Arkansas
a
– – 62,500 60,800 106,950 111,600 118,300 99,400
Illinois 16,700 19,600 29,900 25,300 31,600 30,000 – 25,500
Indiana 13,900 8,900 28,700 25,700 23,500 26,500 31,100 22,600
Kentucky 118,600 165,300 279,100 199,900 166,100 240,100 219,900 198,400
Missouri 6,100 8,500 9,500 6,900 7,400 4,500 – 7,200
Tennessee 148,700 181,500 157,400 118,700 92,100 104,200 139,000 134,500
Total 487,600
a
Estimated using a 13.6-kg mean weight of fi sh harvested; data from 2002 from an incomplete
season.
Table 2. Estimated number of paddlefi sh harvested in six states with an open commercial
fi shing season from 2000 to 2006. Year was based on (1) calendar year for Illinois and
Missouri, and (2) harvest season for other states (e.g., July 1, 2000 to June 30, 2001 is
defi ned as the 2001 year). Harvest data were unavailable for Mississippi.
State 2000 2001 2002 2003 2004 2005 2006 Mean
Arkansas – – 4,600 4,500 7,800 8,200 8,700 7,300
Illinois
a
1,500 1,700 2,600 2,200 2,800 2,600 – 2,600
Indiana 2,400 1,500 4,300 2,700 2,400 3,100 3,500 2,900
Kentucky
b
14,500 20,200 34,100 24,400 20,300 29,300 26,900 24,300
Missouri
a
500 800 800 600 700 400 – 600
Tennessee 11,000 13,500 11,800 8,300 6,800 7,400 8,900 9,700
Total 47,400
a
Estimated using a 11.4-kg mean weight of fi sh harvested.
b
Estimated using a 7.3-kg mean weight of fi sh harvested.
Table 3. Estimated commercial harvest of processed paddlefi sh roe (kg) in six states from
2000 to 2006. Year was based on (1) calendar year for Illinois and Missouri, and (2) har-
vest season for other states (e.g., July 1, 2000 to June 30, 2001 is defi ned as the 2001
year). Harvest data were unavailable from Mississippi, but they were closed for the major-
ity of the roe season.
State 2000 2001 2002 2003 2004 2005 2006 Mean
Arkansas – – 6,120 6,760 6,640 8,330 11,380 8,280
Illinois 1,040 280 2,550 1,460 1,820 3,110 1,900 1,740
Indiana 1,580 960 2,590 1,830 1,250 2,020 2,870 1,650
Kentucky 4,760 4,330 6,930 4,500 4,000 6,950 6,930 4,810
Missouri – – – 790 810 610 – 740
Tennessee – – – 6,330 5,220 8,150 12,900 8,160
Total 23,380
207HARVEST OF PADDLEFISH
in Arkansas harvest came from the Arkan-
sas (40%), Mississippi (22%), and White
(21%) rivers. Kentucky led the nation in
total commercial paddlefi sh fl esh harvest,
with the majority of the fl esh harvest com-
ing from the Ohio River (87%). The fl esh
harvest from Kentucky was higher than
any other state because year-round fi shing
was legal without size limits. Missouri had
the least commercial harvest because the
only open water body was the Mississippi
River and use of unattended entanglement
gear was illegal. Illinois and Indiana had
intermediate harvests.
Annual commercial harvest from the
Ohio River increased from 6,000 to 196,000
kg from 1965–1975 to 2000–2006 (Table 4).
Thomas et al. (2005) documented increased
paddlefi sh abundance in lock chambers of
the Ohio River from 1957 to 2001, which
coincided with improved water quality
(increased dissolved oxygen and pH). Har-
vest declined for the Missouri and Red riv-
ers because they were closed to commercial
paddlefi sh harvest. Annual fl esh harvest
from the Mississippi River remained sub-
stantial (1965–1975 average: 41,000 kg;
2000–2006 average: 103,000) despite com-
mercial closures in Iowa and Louisiana
(Graham 1997) and a partial closure in Illi-
nois (Quist et al. 2009, this volume). Recent
mean annual harvest of 19,000 kg from the
White River was similar to historical Ar-
kansas Game and Fish Commission mean
annual harvest estimates (SD) of 38,000
61,000 kg (1950–1989; N = 26 years).
Commercial harvest data from Mis-
souri were obtained to evaluate harvest
trends from 1945 to 2006 because they have
a complete and continuous harvest record.
Missouri’s harvest regulations remained
relatively stable over time. The Missouri
and St. Francis rivers were closed to har-
vest during 1989. A 610-mm-EFL minimum
length limit was enacted during 1991, but
it was unlikely that this length limit highly
infl uenced harvest because fi sh less than
610 mm were not recruited to large mesh
gill nets (e.g., Paukert and Fisher 1999).
Harvest time-series data from Missouri
were highly variable (Figure 1), and mean
annual fl esh harvest (SD) was 6,800
4,200 kg. Similar to Quist et al. (2009, this
volume), harvest appeared low during
the early to mid-1960s and appeared to be
sustained at relatively high levels for most
Table 4. Estimated commercial fl esh harvest (metric tons) of paddlefi sh in the major rivers
of North America from 2000 to 2006. Year was based on (1) calendar year for Illinois and
Missouri, and (2) harvest season for other states (e.g., July 1, 2000 to June 30, 2001 is
defi ned as the 2001 year). Harvest data were unavailable from Mississippi. One metric ton
is equivalent to 1,000 kg.
River 2000 2001 2002 2003 2004 2005 2006 Mean
Ohio 112 164 286 185 160 247 218 196
Tennessee
a
149 168 126 114 84 88 119 121
Mississippi
a
– – 75 79 149 106 – 103
Arkansas
b
– – – 25 31 38 57 37
White
a
– – 18 14 16 45 4 19
Cumberland
c
12 10 23 15 12 14 30 17
Ouachita
a
– – – 3 3 4 6 4
Illinois 2 1 7 6 1 1 – 3
Black
a
– – – 1 2 3 3 2
a
Estimated using a 11.4-kg average weight.
b
Estimated using a 13.6-kg average weight.
c
Estimates from Tennessee were based on calculating a mean weight from each fi sh harvested
from the Kentucky Lake length–weight relationship.
208 QUINN
0
5,000
10,000
15,000
20,000
25,000
30,000
1945
1950
1955
1960
1965
1970
1975
1980
1985
1990
1995
2000
2005
Year
Harvest (kg)
Figure 1. Harvest of paddlefi sh from Missouri from 1945 to 2006. Data were obtained from
Vince Travnichek, Missouri Department of Conservation. The Missouri River and St. Fran-
cis River were closed to harvest during 1989, and during 1991, a 610-mm-EFL minimum
length limit was enacted.
of the 1970s. Two major harvest peaks oc-
curred during 1980 (16,800 kg) and 1989
(26,000 kg; Figure 1). During 1980, large
harvests were also observed for the Ten-
nessee and Cumberland rivers (340,000
kg dressed weight; Pasch and Alexander
1986). Missouri commercial harvest was
comparatively low from 1992 to 1997 (Fig-
ure 1), and Todd (1998) and Williamson
(2003) suggested that harvest declined
from 1990 to 1997 for Illinois, Missouri, and
Tennessee. From 1993 to 1998, national har-
vest was likely infl uenced by the February
16 through April 15 closed season regula-
tion in Tennessee (Williamson 2003).
Sport Fisheries
Blind snagging was the primary method
sport fi shers use to capture paddlefi sh,
and
archery was a lesser-used harvest method
(Mestl et al. 2005; Scarnecchia et al. 2007).
Recreational fi shing with snagging had
low-size selectivity (Gengerke 1978; Con-
over and Grady 2000; Scarnecchia et al.
2007), and harvest of juvenile paddlefi sh
may occur where permitted. The distribu-
tion of paddlefi sh sport fi sheries were re-
viewed by Carlson and Bonislawsky (1981),
Gengerke (1986), and Graham (1997). Ma-
jor recreational sport fi sheries tend to ex-
ist where paddlefi sh make spawning runs,
especially in tailwaters below dams. The
Lake Sakakawea/Yellowstone River snag
fi shery was unique because donated sport-
harvested roe was processed and sold and
the proceeds were applied toward paddle-
fi sh management in North Dakota and
Montana (Scarnecchia et al. 2007).
Carlson and Bonislawsky (1981) sug-
209HARVEST OF PADDLEFISH
gested that sport harvest was greater than
commercial harvest in the Mississippi Riv-
er basin, and they reported harvest for 11
major sport fi sheries. They reported that
maximum sport harvest was at Kentucky
Dam, Kentucky, where anglers snagged
102,800 fi sh during 1978 with average size
of only 3 kg. However, this was an ex-
treme harvest outlier. Sport harvest from
the Kentucky Dam tailwater was 15,050,
1,000, and 1,790 fi sh in 1979, 1994, and 2006
(Carlson and Bonislawsky 1981; Timmons
and Hughbanks 2000; P. Rister, Kentucky
Department of Fish and Wildlife Resourc-
es, personal communication). Carlson and
Bonislawsky (1981) reported that the aver-
age weight of paddlefi sh harvested in snag
fi sheries varied from 3 to 28 kg.
Sport harvest data were surveyed from
appropriate state agency and university per-
sonnel. The number of fi sh harvested was
reported for all sport fi sheries, but mean
weight data were generally unavailable.
Mean annual sport harvest in North
America was estimated as 12,700 fi sh from
2000 to 2006 (Table 5). Sport harvest data
were not available for the major Lake of
the Ozarks, Truman Lake, and Osage River
fi sheries in Missouri, which were expected
to add roughly 2,000–5,000 fi sh to the total
harvest (Graham 1997). Sport harvest data
for most of the Arkansas, Ohio, and upper
Mississippi River tailwater fi sheries were
not available (Graham 1997; Williamson
2003), but cumulative harvest from these
waters may be important. For example, the
Markland tailwater, Ohio River, had a har-
vest of 500 paddlefi sh during 1997 (Henley
et al. 2001a), and 1,320 paddlefi sh were har-
vested from the Mississippi River at Lock
and Dam 12 in Iowa during 1976 (Carlson
and Bonislawsky 1981). Thus, the estimate
of 12,700 fi sh was very conservative, and a
more accurate range-wide sport harvest es-
timate may reasonably be expected to be be-
tween 15,000 and 20,000 fi sh. Overall, seven
major sport fi sheries had harvests greater
than 1,000 fi sh annually, including the Lake
of the Ozarks and Truman Lake fi sheries.
Comparing Harvest to
Population Estimates
Mean paddlefi sh population density among
12 select waters was 2.7 fi sh/ha (95% confi -
dence interval; 0.1–5.3; median = 0.9; Table
Table 5. Number of paddlefi sh harvested from major paddlefi sh sport fi sheries from 2000
to 2006. Data were unavailable for the Lake of the Ozarks and Truman Lake fi sheries in
Missouri.
Fishery State 2000 2001 2002 2003 2004 2005 2006 Mean
Grand Lake/
Neosho River OK – – – 2,930 2,700 – – 2,810
Yellowstone/Missouri/
Sakakawea ND, MT 2,870 1,930 2,570 2,100 1,300 2,420 2,250 2,210
Kentucky Dam tailwater KY – – – – – – 1,790 1,790
Missouri River/
Gavins Point NE, SD 940 1,030 730 1,200 2,090 1,950 1,940 1,410
Table Rock Lake MO – 1,260 1,370 – – – – 1,310
Missouri River/
Fort Peck MT – – 540 870 790 1,030 1,290 900
Barkley Dam tailwater KY 810 – – – – – – 810
Ozark Lake/
Pool 13 tailwater AR – – – – 600 – – 600
Neosho River,
Chetopa Dam KS 100 – 480 1,590 510 380 440 510
Fort Gibson/Grand River OK – – – – – 320 – 320
210 QUINN
6), and maximum reported density was 13.1
paddlefi sh/ha for Sibley Lake, an oxbow of
the Alabama River (Hoxmeier and DeVries
1997). Biomass estimates for paddlefi sh in
natural waters typically ranged between 11
and 22 kg/ha (Onders et al. 2001), but max-
imum biomass exceeded 40 kg/ha (Jenkins
et al. 1982). A rotenone evaluation of Bar-
kley Lake, Kentucky estimated biomass at
6.8 kg/ha (Aggus et al. 1979), and biomass
ranged from 0 to 10.5 kg/ha among seven
Louisiana oxbow lakes (Lambou 1959).
Mean biomass estimates for paddlefi sh at
four long-term (>25 years; SD) Arkansas
River rotenone sites were 0, 0, 21.7 10.3,
and 25.6 11.2 kg/ha (Quinn and Limbird
2008).
Harvests of paddlefi sh compiled from
the literature were highly variable and
ranged between less than 0.01 and 5.06
fi sh/ha and less than 1 and 43 kg/ha (me-
dian = 0.12 fi sh/ha, 1.73 kg/ha; Appendix
1). Maximum reported harvest for a large
water body (>2,000 acres) was 42 kg/ha
from Wilson Reservoir, Tennessee River,
Alabama (Tarzwell and Bryan 1945). This
harvest is realistic because biomass esti-
mates of paddlefi sh greater than 914 mm
EFL ranged from 32 to 87 kg/ha for Ozark
Lake, Arkansas River (Quinn et al. 2006).
Also, Jenkins et al. (1982) reported harvests
of 32.6 and 43.4 kg/ha for Energy and
Bards Lakes, which are 150- and 129-ha
subimpoundments of Lake Barkley.
The highest sustainable harvest rates
found in the literature were 1.9–3.5 kg/ha
for the Neosho/Grand Lake sport fi shery,
with an associated exploitation rate of 15–
17% and population density of 1.02–1.38
fi sh/ha (Combs 1982). Based on the liter-
ature, harvests less than 1 kg/ha did not
appear to be problematic. However, stock
depression for Norris Reservoir occurred
at a relatively modest 1.5 kg/ha (0.06 fi sh/
ha), and after the fi shery recovered, this
reservoir had a biomass of 11.2 kg/ha (0.64
fi sh/ha; Alexander and Peterson 1982). Us-
ing the suggested 15% maximum exploi-
tation rate (Combs 1982) and an expected
biomass range of 11–22 kg/ha (Onders et
al. 2001), harvests greater than 3.3 kg/ha
should raise management concerns for pro-
ductive waters. Harvests greater than 5 kg/
ha reported in the literature were usually
related to opening a previously closed fi sh-
ery or overharvest (Appendix 1). Compar-
ing harvests to population density and bio-
mass estimates is useful for understanding
sustainability of harvests, but it should not
replace understanding of site-specifi c fi sh-
eries characteristics and demographic rates
(e.g., mortality, growth, recruitment).
Table 6. Paddlefi sh density estimates (fi sh/ha) for select waters in North America. CI =
confi dence interval.
Area Density
Water (ha) fi sh/ha (95% CI) Citation
Barkley Lake, KY 23,400 1.2 Aggus et al. 1979
Grand Lake, OK 22,000 1.14 (0.86–1.45) Combs 1982
Keystone Reservoir, OK 10,600 0.81 (0.37–1.25) Paukert and Fisher 2001
Lake Cumberland, KY 20,336 0.64 Hageman et al. 1986
Lake Francis Case, SD 33,994 0.79 Unkenholtz 1979
Lake Sakakawea, MT 156,000 0.21 Ryckman 2003
Norris Reservoir, TN 13,695 0.64 (0.34–1.35) Alexander and Peterson 1982
Ozark Lake, AR 4,249 3.53 (0.87–6.19) Quinn et al. 2004
Mississippi River, IA 10,918 0.99 (0.40–2.47) Gengerke 1978
Sibley Lake, AL 32 13.10 Hoxmeier and DeVries 1997
South Cross Creek R., KY 192 8.79 (8.12–9.47) Boone and Timmons 1995
Watts Bar, TN 15,783 0.22 (0.14–0.42) Alexander et al. 1985
211HARVEST OF PADDLEFISH
Case Histories for Commercial
Fisheries
Kentucky Lake
Kentucky Lake was the best studied com-
mercial paddlefi sh fi shery and it provided
a valuable case history for understanding
the effects of harvest (Bronte and John-
son 1983, 1984, 1985; Hoffnagle and Tim-
mons 1989; Timmons and Hughbanks
2000; Scholten and Bettoli 2005; Bettoli et
al. 2007). This large main-stem Tennes-
see River reservoir was 296 km long and
64,800 ha and had a residence time of 14 d
(Voigtlander and Poppe 1989). The riverine
portion fl ows through Tennessee, and the
Kentucky portion was primarily reservoir
habitat. Paddlefi sh segregated in this river
reservoir, with juvenile paddlefi sh remain-
ing in the downstream, lacustrine portion
of the lake (Scholten and Bettoli 2005).
Kentucky Lake was reopened to com-
mercial entanglement gear during 1979
after being closed for 3 years (Bronte and
Johnson 1983). Based on fi sh observed from
commercial landings, Bronte and Johnson
(1985) estimated that total annual mortal-
ity for fi sh greater than age 6 was 44% and
maximum age was 14 during 1980–1981.
Gravid females were greater than age 9
and ranged from 930 to 1,002 mm EFL and
16.8–22.7 kg. Age at recruitment into the
fi shery was 6–7 years and length at recruit-
ment was 700 mm EFL. Bronte and John-
son (1985) concluded that fi sh growth was
rapid to large sizes, and fi sh recruited into
the fi shery before contributing to repro-
duction.
Hoffnagle and Timmons (1989) indi-
cated that Kentucky Lake was overfi shed
during 1985–1986 after a period of heavy
exploitation from 1984 to 1985. They ob-
served 702 paddlefi sh captured by com-
mercial fi shers. The sex ratio was 1:1, but
only 2% were gravid females. Total annual
mortality from catch curve was 69%, and
maximum age was 16 years. Mean length
and weight of the catch was only 691 mm
EFL and 5.7 kg.
From 1991 to 1994, Timmons and
Hughbanks (2000) found 14% exploitation
and 22% total annual mortality for ages
9–15, and they concluded that the paddle-
fi sh stock of Kentucky Lake was recover-
ing. Sex ratio was 0.64:1 males:females.
Mean length of paddlefi sh was 857 mm
EFL (range: 690–1,220 mm EFL), where-
as mean length was 691 mm EFL during
1985–1986 (Hoffnagle and Timmons 1989).
Mean length, weight, and maximum age in-
creased by 166 mm EFL, 5.8 kg, and 5 years
(16–21) since 1985–1986. Todd (1998) noted
that commercial harvest of paddlefi sh de-
clined in Tennessee waters from 221,000 to
29,000 kg between 1990 and 1996.
Scholten and Bettoli (2005) found
very high total annual mortality (68%) for
paddlefi sh in Kentucky Lake during 2003.
They found that harvest was positively re-
lated to the number of fi shable days when
mean discharge was less than 850 m
3
/s
because fi shers could not deploy nets dur-
ing high fl ows. They found evidence that
no mature females were protected by the
864-mm-EFL minimum length limit, and
recruitment overfi shing likely occurred
during low-fl ow years. They suggested a
1,016-mm-EFL minimum length limit was
needed to maintain spawning potential ra-
tios greater than 20% at high exploitation
rates (62%).
Tennessee River, Alabama
The Tennessee River commercial paddle-
fi sh fi shery in Alabama provides a classic
paddlefi sh stock-depression, overfi shing
example. Pasch and Alexander (1986) re-
viewed harvests for the Tennessee River
snag fi sheries (i.e., Guntersville, Wheeler,
Wilson, and Pickwick lakes; total area =
56,000 ha) and found major declines in
commercial harvest from 1942 to 1952
(293,000–43,500 kg; 5.23–0.77 kg/ha). This
fi shery apparently recovered during the
212 QUINN
late 1950s and 1960s (Tarzwell and Bryan
1945; Bryan and White 1958). From 1948 to
1973, harvest ranged from 21,800 kg (1956)
to 281,700 kg (1964), with a secondary har-
vest peak of 175,900 kg in 1973 (Pasch et
al. 1978). Paddlefi sh harvest increased in
the late 1970s and reached 150,000 kg (2.68
kg/ha) by 1980 due to increased roe prices
(Gengerke 1986). Hoxmeier and DeVries
(1996) failed to fi nd paddlefi sh in the Ala-
bama portion of the Tennessee River dur-
ing 1993–1994 after the fi shery was closed
during 1988, although their efforts were
not extensive. Fisheries biologists from
Alabama, based upon fi eld sampling and
creel reports, concluded that the paddle-
fi sh population in the Alabama portion of
the Tennessee River has not recovered as
of 2006 (S. Cook, Alabama Wildlife and
Freshwater Fisheries Division, personal
communication). Additional evidence that
Guntersville Lake has not recovered is
that opening of the Tennessee portion be-
low Nickajack Lake produced a very small
commercial harvest (Appendix 1). The
Tennessee River and Kentucky Lake case
histories provided evidence (e.g., Pasch
and Alexander 1986) that river reservoirs
are susceptible to overfi shing.
Ohio River
The Ohio River commercial fi shery (Ken-
tucky, Indiana, and Illinois) has been inten-
sively fi shed from 1999 to 2006 and shows
signs of heavy exploitation (Henley et al.
2001a). Estimated average size and age at
harvest was only 5.45 kg, 710 mm EFL,
and 8.5 years (Henley et al. 2001a). Dur-
ing 1995–2001, total annual mortality esti-
mates ranged from 47% to 68% (Henley et
al. 2001b), and length-frequency distribu-
tions were truncated at 850 mm EFL. The
Ohio River management team tracked the
percentage of fi sh age 10 and older and
found that it peaked in 127-mm bar mesh
gill nets at 71% during 1997 and declined
to a low of 46% during 2001 after a period
of intensive fi shing. The Ohio River case
history indicated that intensive commer-
cial harvest resulted in a small size distri-
bution and a relatively young population
with high annual mortality.
Discussion
Anthropogenic alterations to habitat, over-
reliance on harvest data, and lack of fi sh-
eries-independent data limit our historical
understanding of the degree of threat that
harvest is to paddlefi sh populations. Hu-
man-alteration of large North American
rivers (e.g., Barnickol and Starrett 1951;
Sheehan and Rasmussen 1999) occurred
at the same time that commercial harvests
of paddlefi sh reportedly declined (Carl-
son and Bonislawsky 1981; Gengerke 1986;
Pikitch et al. 2005). Factors that increased
uncertainty with the quality of harvest data
included lack of data collection standards,
lack of quality control in several states, and
inconsistent reporting by fi shers. Draw-
ing inferences about historical popula-
tion trends from commercial harvest data
should be done with caution because (1)
fi shing effort, habitat, and harvest regu-
lations varied over time, and some states
closed commercial fi shing (Graham 1997);
(2) data collection formats were often not
thoroughly documented; (3) many fi sher-
ies shifted from fl esh to roe harvest; and (4)
harvest data only represent minimal catch
estimates (Carlson and Bonislawsky 1981).
Thankfully, fi sheries-independent stock
surveys have increased in recent decades
since the development of the Mississippi
Interstate Cooperative Resource Associa-
tion (Grady et al. 2005).
The magnitude of recent commercial
harvest estimates (Tables 1 and 2) indicates
that previous comments and analyses sug-
gesting range-wide population declines
and depletion of paddlefi sh stocks may
have been overstated (e.g., Gengerke 1986;
Graham 1997; Pikitch et al. 2005). Pikitch
213HARVEST OF PADDLEFISH
et al. (2005) described a steeply declining
paddlefi sh harvest trend over time, and
their results refl ected (1) that states such
as Arkansas stopped collecting paddlefi sh
harvest data after National Marine Fisher-
ies Service eliminated funding in the late
1980s, and (2) several states closed paddle-
fi sh harvest (e.g., Iowa, Louisiana). Graham
(1997) stated that paddlefi sh have declined
dramatically in most areas during the past
100 years; however, state-by-state status re-
views suggested that paddlefi sh were com-
mon throughout much of the species range
where habitats were not severely altered
(Gengerke 1986; Graham 1997).
Market conditions and fi shing effort
were primary factors infl uencing harvest
of paddlefi sh (Gengerke 1986; Quist et al.
2009; Scholten 2009). Increased demand
and effort occurred when Caspian Sea
sturgeon caviar imports were banned or
reduced. Major peaks in effort occurred
during the 1940s associated with World
War II (Bryan and White 1958), during the
embargo on Iranian caviar in the late 1970s
through the 1980s (Alexander and Peterson
1982), and from 2005 to present associated
with limited Caspian Sea quotas (Colombo
et al. 2007).
A growing body of evidence suggested
commercial harvest was a threat to paddle-
fi sh in lentic waters, including impounded
rivers such as the Arkansas, Ohio, and Ten-
nessee rivers (Leone et al. 2005; Scholten
and Bettoli 2005). Stockard (1907) noted
depletion of paddlefi sh from oxbow lakes
from seining operations. Paddlefi sh typi-
cally grow faster in reservoirs (Paukert
and Fisher 2001) and become susceptible to
commercial harvest before reaching sexual
maturity (Pasch and Alexander 1986; Schol-
ten and Bettoli 2005). Main-stem reservoirs
on large rivers, with abundant lentic habi-
tats, such as those on the Tennessee River
and Arkansas River, had high commer-
cial harvests and appeared susceptible to
overharvest without adequate regulation
(Pasch and Alexander 1986; Scholten and
Bettoli 2005). The degree of threat posed
by harvest of paddlefi sh from large open
rivers such as the Mississippi and lower
White River, Arkansas, will remain unre-
solved until fi sheries independent data be-
comes available.
Commercial fi shing has the potential
to quickly reduce the abundance of large,
old fi sh in a population (e.g., Hoffnagle
and Timmons 1989; Henley et al. 2001b),
and maximum age usually declined with
increasing harvest. Maximum age for Ken-
tucky Lake was 7.2, 14, and 21 years when
total annual mortality was 68%, 44%, and
14%, respectively (Bronte and Johnson 1984;
Timmons and Hughbanks 2000; Scholten
and Bettoli 2005). Three Arkansas River
pools appeared to follow the pattern of de-
clining maximum age with increased har-
vest, and maximum ages were 16, 15, and
13 years for pools with 51%, 56%, and 66%
total annual mortality (F. Leone, Arkansas
Game and Fish Commission, unpublished
data). Francis et al. (2007) suggested that
maintaining old fi sh in fi sh populations
was one of the key tenants of ecosystem-
based fi sheries science because old fi sh
often had higher spawning success. Scar-
necchia et al. (2007) suggested that harvest
regulations should be developed to protect
a range of ages of mature fi sh, including
middle-aged and old spawning females.
Published population modeling stud-
ies indicated that paddlefi sh are suscep-
tible to recruitment overfi shing (Boreman
1997; Scholten and Bettoli 2005), and fi eld
studies confi rmed that recruitment was
depressed for some paddlefi sh popula-
tions by the harvesting of adult spawners
(Alexander and McDonough 1983). Alex-
ander and McDonough (1983) noted a se-
vere decline in age-0 paddlefi sh at the Gal-
latin Steam Electric Plant on Old Hickory
Lake, Tennessee after heavy exploitation
during the late 1970s. Hilborn and Walters
(1992) noted that studies often fail to relate
214 QUINN
recruitment to stock size, but when fi sher-
ies are fi shed extensively, recruitment will
eventually decline.
The functions used to describe stock–
recruitment relationships can provide
greatly different results in population
models (e.g., Rieman and Beamesderfer
1990). Population modeling indicated that
some gill-net fi sheries may not experience
recruitment or growth overfi shing with ad-
equate minimum length limits if bycatch
mortality of undersized fi sh is low (Schol-
ten and Bettoli 2005; Leone et al. 2005). By-
catch mortality of undersized fi sh may be
a threat for maintaining sustainable com-
mercial paddlefi sh fi sheries (Barnickol and
Starrett 1951; Bettoli and Scholten 2006).
Increased management by state agen-
cies has reduced the threat of harvest to
paddlefi sh over the species range. Rec-
reational harvests were often intensively
managed through complex harvest regu-
lations, and harvest quotas were used to
create sustainable fi sheries (e.g., Ryck-
man 2003; Mestl et al. 2005). Commercial
harvest was closed for much of the range
of paddlefi sh (e.g., Alabama, Iowa, Loui-
siana, Minnesota, Nebraska, New York,
North Dakota, Oklahoma, Pennsylvania,
South Dakota, Ohio, Texas, West Virginia,
and Wisconsin). Management of commer-
cial fi sheries in Arkansas and Tennessee
has focused on protecting sexually imma-
ture paddlefi sh to commercial exploitation
through high minimum length limits, and
harvest seasons were set to limit bycatch
mortality (Scholten 2009). Several states
open to commercial fi shing for paddlefi sh
had some rivers or river sections that were
closed to provide a refuge (e.g., Arkansas,
Illinois). However, management for sus-
tainable paddlefi sh fi sheries has faced nu-
merous political challenges (Bettoli et al.
2007).
A major challenge for fi sheries manag-
ers is characterizing what is a sustainable
harvest. Recent freshwater commercial
fi sheries management has focused on
maintaining spawning potential ratios
(SPR; Goodyear 2003) greater than 20–30%
to prevent recruitment overfi shing (e.g.,
Slipke et al. 2002; Goodyear 2003; Schol-
ten and Bettoli 2005; Leone et al. 2005;
Maceina and Pereira 2007). Slipke et al.
(2002) reviewed recruitment overfi shing
in freshwater fi sheries and demonstrated
the utility of SPR for freshwater fi sher-
ies management. Traditional commercial
fi sheries stock assessments and biological
reference points (e.g., FMSY, F0.1) used
to assess growth overfi shing for marine
fi sheries (Hilborn and Walters 1992) have
not been applied to paddlefi sh roe fi sher-
ies. During the next decade, paddlefi sh
managers will likely fi eld test the utility
of SPR targets for preventing recruitment
overfi shing.
Acknowledgments
I would like to thank all the biologists who
provided information and harvest data that
contributed to this review, including Doug
Henley (Kentucky Department of Fish and
Wildlife Resources), Rob Maher (Illinois
Department of Natural Resources), Gerald
Mestl, Tom Mosher (Kansas Wildlife and
Parks), Bill Posey (Arkansas Game and
Fish Commission), Vic Riggs (Montana
Department of Fish, Wildlife, and Parks),
Paul Rister (Kentucky Department of Fish
and Wildlife Resources), Fred Ryckman
(North Dakota Game and Fish Depart-
ment), George Scholten (Tennessee Wildlife
Resources Agency), Tom Stefanavage (In-
diana Department of Natural Resources),
Vince Travnichek (Missouri Department of
Conservation), and Trish Yasger (Missouri
Department of Conservation). This manu-
script was improved by discussions I have
had with Arkansas commercial fi shers, and
constructive reviews by Jason Olive, Frank
Leone, George Scholten, Craig Paukert,
and two anonymous reviewers.
215HARVEST OF PADDLEFISH
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219HARVEST OF PADDLEFISH
Appendix 1. Harvest, exploitation (u), and roe harvest of paddlefi sh from select sport and commercial fi sheries in North American
waterbodies. Mean harvest is provided where multiple years are listed. Type of fi shery included commercial (C) and sport (S).
Mean Roe
Drainage and Area Harvest Harvest weight Harvest Harvest harvest
Waterbody or state Years(s) Type (ha) (fi sh) (kg) (kg) (fi sh/ha) (kg/ha) u (kg)
Arkansas River
Ozark Lake
1
* 2004 C 4,291 1,075 15,910 14.8 0.25 3.71 19% 1,473
Ozark Lake (5 d)
1
* 2006 C 4,291 1,065 16,934 15.9 0.25 3.95 25% 2,048
Ozark Lake (150 d)
1
* 2006 C 4,291 2,260 35,934 15.9 0.53 8.37 4,326
Pool 13
1
* 2003 C 2,308 1,588 23,026 14.5 0.69 9.98 714
Pool 13
1
* 2006 C 2,308 2,453 35,569 14.5 1.06 15.40 35% 2,030
Lake Dardanelle
2
2003–2006 C 13,887 1,041 15,615** 0.07 1.12 3,365
Neosho River
3
1979–1980 S 22,000 3,008 57,011 19.0 0.14 2.71 17%
Fort Gibson Lake
4
1957–1960 C 8,900 370 0.04
Clinch River
Norris Reservoir
5
*** 1959 C 13,840 781 20,256 25.9 0.06 1.46
Norris Reservoir
5
1960 C 13,840 233 5,684 24.0 0.02 0.41
Norris Reservoir
5
1961 C 13,840 63 1,038 22.2 0.01 0.08
Norris Reservoir
5
1959–1961 C 13,840 359 8,993 22.2 0.03 0.65
Cumberland River
Bards Lake
6
*** 1980 C 129 653 5,602 8.5 5.06 43.43 0
Old Hickory Reservoir
7
1977 C 9,100 630 2,835 4.5 0.07 0.31
Old Hickory Reservoir
8
1999–2006 C 9,100 66 0.01 8
Cheatham Reservoir
8
1999–2006 C 3,015 174 0.06
Energy Lake
6
*** 1980 C 150 495 4,883 9.9 4.07 32.55 0
Lake Cumberland
9
1983–84 C 20,336 3,566 37,764 10.5 0.18 1.86 ~12%
Barkley Lake, KY
10
1979 C 18,960 7,015 62,000 8.8 0.37 3.27
Barkley Lake, KY
10
1980 C 18,960 3,685 21,500 5.8 0.19 1.13
Barkley Lake KY, TN
11
1991–1994 B 23,400 3,597** 37,764 10.5 0.18 1.61 14%
Barkley Lake, TN
8
1999–2006 C 7,407 150 0.02
Barkley Dam
tailwater
12
1979 S 2,276 6,828 3.0
Mississippi River
AR
2
2003–2006 C 40,740 2,143 24,430** 0.05 0.60 1,656
IL
13
1999–01 C 1,914** 21,828 503
KY
13
1999–01 C 270** 3,080 779
MO
13
1999–01 C 543** 6,195
MO, open river
14
2003–2005 C ~55,000 465 5,305 0.01 0.10 647
Pool 13, IA
12
1976 S 10,918 1,320 7,920 6 0.12 0.73
220 QUINN
Appendix 1. Continued.
Mean Roe
Drainage and Area Harvest Harvest weight Harvest Harvest harvest
Waterbody or state Year(s) Type (ha) (fi sh) (kg) (kg) (fi sh/ha) (kg/ha) u (kg)
Pool 13, IA
15
1975–1977 C 10,918 760** 8,812 0.81 4–12%
Pool 24, MO
14
2003–2005 C 4,472 102 1,163** 0.02 0.26 3
Pool 26, MO
14
2003–2005 C 6,490 849 9,679** 0.13 1.49 85
TN
8
1999–2006 C 40,150 1,504 0.04 2,948
Reelfoot Lake
8
2002–2006 C 4,220 56 0.01
Missouri River
Unchannelized
reach
16, 12
1972–1974 B 3,294 19,764** 6 7%
Yellowstone/Missouri
17
1973–1974 S 156,000 4,452 42,000 18 0.03 0.27 8%
Ohio River
KY, IL, IN
18, 19
1999 C 67,178 11,711 63,827 5.5 0.174 0.95 2,733
KY, IL, IN
18, 19
2000 C 67,178 29,194 159,109 5.5 0.582 2.37 10,071
Osage River
Lake of the Ozarks
20
1958–61 S 22,275 5,822 79,569 14.0 0.26 3.57 19%
Lake of the Ozarks
21
1978–1988 S 22,275 2,809 48,315 17.2 0.13 2.17
Harry S. Truman
Lake
22
1992 S 22,000 4,041 0.18
Ouachita River, AR
2
2003–06 C 3,225 325 3,705** 0.10 1.15 486
Tennessee River
Wilson Reservoir
23
*** 1942 C 6,475 324,000 42.03
AL
23
*** 1942–52 C 80,664 267,302 3.31
Cherokee
8
1999–2002 C 12,262 98 0.01
Chickamauga
8
2000–2006 C 13,962 356 0.03 300
Guntersville
8
2003–2006 C 468 8 0.02
Kentucky Lake
8
1999–2006 C 43,784 7,460 0.17 4,593
Kentucky Dam
tailwater
12
1978 S ~1,450 102,837 308,511 3.0 70.9 212.8
Kentucky Dam
tailwater
12
1979 S ~1,450 15,050 45,150 3.0 10.4 10.4
Kentucky Dam
tailwater
8
2006 S ~1,450 1,788 1.2
Pickwick Lake
8
1999–2006 C 2,492 35 0.02 11
Watts Bar
8
2001–2006 C 15,783 63 <0.01 46
White River
221HARVEST OF PADDLEFISH
Appendix 1. Continued.
Mean Roe
Drainage and Area Harvest Harvest weight Harvest Harvest harvest
Waterbody or state Years(s) Type (ha) (fi sh) (kg) (kg) (fi sh/ha) (kg/ha) u (kg)
Black River, AR
2
2003–06 C 162 234
Table Rock Lake
21
1990 S 17,400 350 0.02
Lower White River
24
1987–1988 C ~8,148 108,818 13.3 19–31% Lower
White River
2
2003–06 C ~8,148 1,743 19,870 0.21 2.4 1,642
1
Quinn et al. 2009;
2
Posey 2006;
3
Combs 1982;
4
Houser 1965 ;
5
Carrol et al. 1963;
6
Jenkins et al. 1982;
7
Pasch et al. 1980;
8
Tennes-
see Wildlife Resources Agency, unpublished data;
9
Hageman et al. 1986;
10
Alexander and Peterson 1982;
11
Timmons and Hughbanks
2000;
12
Carlson and Bonislawsky 1981;
13
Williamson 2003;
14
Missouri Department of Conservation, unpublished data;
15
Gengerke 1978, Pool
12 tailwater;
16
Rosen et al. 1982, reach below Gavins Point Dam;
17
Rehwinkle 1978, Lake Sakakawea stock;
18
Henley et al. 2001b;
19
Henley
et al. 2001a;
20
Purkett 1963;
21
Graham 1988;
22
Graham 1997;
23
Guntersville, Pickwick, and Wheeler reservoirs, Pasch and Alexander 1986;
24
Robison and Buchanan 1988, Filipek 1990; *First year harvest after the fi shery was closed; **Estimated using an assumed mean weight;
***Documented overharvest.
