Articles

Overfishing of Inland Waters

J. DAVID ALLAN, ROBIN ABELL, ZEB HOGAN, CARMEN REVENGA, BRAD W. TAYLOR, ROBIN L. WELCOMME,
AND KIRK WINEMILLER

Keywords: overfishing, fishing down, freshwater biodiversity, ecosystem function, fish harvest

verexploitation of the worlds fisheries is the

subject of much recent concern (FAO 2002, Pauly et al.
2002, Hilborn et al. 2003). Although the global production of
fish and fishery products continues to grow, the harvest from
capture fisheries has stagnated over the last decade. Today numerous fish stocks and species have declined since their historical peaks, and some have even collapsed, leading to urgent
calls for more stringent management and the establishment
of protected areas (Roberts et al. 2003). However, the discussion of the current fisheries crisis has focused nearly exclusively on marine resources, and to some extent on
associated threats to marine biodiversity, particularly those affecting charismatic animals such as seabirds, marine turtles,
dolphins, and whales. The fisheries of inland waters have received only slight consideration within global analyses (FAO
1999, Hilborn et al. 2003, Kura et al. 2004). Here we summarize
and evaluate the evidence that overfishing in inland waters is
occurring and is a contributing factor to the decline of freshwater biodiversity. We define inland fisheries as the capture
of wild stocks of primarily freshwater fish, including migratory species that move between fresh water and the oceans.
Although aquaculture significantly augments the supply of certain species and contributes a substantial fraction to the overall harvest, we focus on capture fisheries of natural stocks.
Fishing and the activities surrounding itprocessing,
packing, transport, and retailingare important at every
scale, from the village level to national and international
economies. Fishing is a crucial source of livelihoods in developing nations, particularly for low-income families in
rural areas where job options are limited. Small-scale commercial and subsistence fishing often provides the employment
of last resort when more lucrative labor opportunities cannot be found (Kura et al. 2004). This is particularly true for

inland fisheries. Although there are no global estimates of the

number of people engaged in inland fisheries, in China alone,
more than 80% of the 12 million reported fishers are engaged
in inland capture fishing and aquaculture (Kura et al. 2004).
The contribution of fisheries to the global food supply is
also significant. In 2000, fish and fishery products constituted
15.3% of the total animal protein consumed by people
(FAO 2003). About 1 billion peoplelargely in developing
countriesrely on fish as their primary animal protein source
(calculation based on Laurenti 2002), and this is especially true
for poor rural communities. For example, within the lower
Mekong basin, the average consumption of fish and other
aquatic animals is estimated at 56 kilograms (kg) per capita
per year (Hortle and Bush 2003), and may reach 71 kg per
capita per year in high-yielding fishing areas such as the
floodplains around Tonle Sap Lake in Cambodia (Ahmed et
al. 1998). In comparison, the global average is 16 kg per
capita per year (FAO 2002).

J. David Allan (e-mail: [email protected]) works at the School of Natural Resources and Environment, University of Michigan, Ann Arbor, MI 48109.
Robin Abell is with the Conservation Science Program, World Wildlife Fund,
Washington, DC 20037. Zeb Hogan works at the Center for Limnology, University of Wisconsin, Madison, WI 53706. Carmen Revenga is with the Global
Priorities Group, The Nature Conservancy, Arlington, VA 22203. Brad W.
Taylor is affiliated with the Department of Zoology and Physiology at the
University of Wyoming in Laramie and the Department of Biological Sciences,
Dartmouth College, Hanover, NH 03755. Robin L. Welcomme works at Long
Barn, Stoke by Clare, Suffolk CO10 8HJ, United Kingdom. Kirk Winemiller
is with the Interdisciplinary Program in Ecology and Evolutionary Biology and
the Department of Wildlife and Fisheries Sciences, Texas A&M University,
College Station, TX 77843. 2005 American Institute of Biological Sciences.

December 2005 / Vol. 55 No. 12 BioScience 1041

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Inland waters have received only slight consideration in recent discussions of the global fisheries crisis, even though inland fisheries provide
much-needed protein, jobs, and income, especially in poor rural communities of developing countries. Systematic overfishing of fresh waters is largely
unrecognized because of weak reporting and because fishery declines take place within a complex of other pressures. Moreover, the ecosystem
consequences of changes to the species, size, and trophic composition of fish assemblages are poorly understood. These complexities underlie the
paradox that overexploitation of a fishery may not be marked by declines in total yield, even when individual species and long-term sustainability
are highly threatened. Indeed, one of the symptoms of intense fishing in inland waters is the collapse of particular stocks even as overall fish
production risesa biodiversity crisis more than a fisheries crisis.

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framework, and subsidies to the fishery

through hatchery stocking (Post et al.
2002). In relatively remote areas of the
Orinoco, we have observed rapid declines
of large peacock bass (Cichla spp.)
subsequent to the fisherys discovery by affluent anglers (K. W., personal
observation).
The reporting of global fisheries statistics to the Food and Agriculture Organization of the United Nations (FAO)
relies on data provided by member countries, with the potential for distortions.
Watson and Pauly (2001) argued that increases in Chinas reported marine catch,
beginning in the 1980s, were consistent
neither with a careful examination of
other catch statistics nor with a model
based on oceanographic conditions, suggesting instead that catches had been exaggerated. In 1998, in response to
domestic and foreign criticism, China
Figure 1. Fisheries landings from inland waters, 19502002: (a) world, (b) Africa,
declared a zero-growth policy in which re(c) Asia, and (d) China as a percentage of Asia. Data are from the Food and Agriported catches were frozen at 1998 levels.
culture Organization of the United Nations (FAO 2004). Abbreviation: MMT, milPerhaps reflecting similarly questionable
lion metric tons.
reporting for inland waters, Chinas reported
inland catch leveled off after 1998,
The status of inland fisheries
and
its
share
of
Asias
total
catch,
which had increased strongly
The total catch from inland waters, exclusive of aquaculture
since
the
1960s,
fell
(figure
1).
harvest and recreational fishing, was 8.7 million metric tons
The use of catch statistics to assess stocks, a common pracin 2002, the last year for which statistics are available (FAO
tice
with marine species, is difficult with inland species because
2004). Of this total, Asia accounted for 65% and Africa for
much
of the inland catch includes artisanal, recreational,
24%, with the remainder landed in South America (4%),
and
illegal
fisheries; moreover, landings are dispersed, and
Europe and the former USSR (4%), North America (2%), and
are
underreported
by a factor of three or four, according to
Oceania (less than 1%). Fisheries landings from inland wathe
FAO
(1999).
FAO
records date back only to 1950, and
ters have experienced a more than fourfold increase, roughly
separate
accounting
of
inland fisheries was introduced only
3% annually, since data were first compiled in 1950 (figure 1).
in
the
1990s;
thus,
historical
declines of important fish stocks
China accounts for about one-quarter of the worlds inland
are
poorly
documented.
Finally,
catch data are rarely reported
catch. Other countries with significant inland catches are Inat
the
species
level,
making
stock
assessments even more
dia, Bangladesh, Cambodia, Indonesia, Egypt, Tanzania,
challenging.
Myanmar, Uganda, and Thailandall developing or transitional economies where inland fish harvests have rapidly increased over the last 10 to 15 years.
Overfishing and freshwater biodiversity
Developed regions of the world have seen the opposite
The status of inland waters and their species should be of
trend. North America, Europe, and the former Soviet Union
broad concern, yet threats to freshwater fisheries and assoall show declining trends in inland capture fisheries, as many
ciated biodiversity have received scant attention from coninland commercial fisheries have been abandoned and reservation groups and the media. This imbalance seems
placed by recreational fisheries, which may add substantially
particularly dangerous considering evidence that freshwater
to the total fisheries harvest but are not always reported
ecosystems and the species they support are, on average,
(Cooke and Cowx 2004). The global recreational harvest is
more threatened than marine ecosystems (Ricciardi and
poorly documented, but may be on the order of 2 million metRasmussen 1999).
ric tons (FAO 1999). Even when fish are released, recreational
Inland waters and their species experience myriad direct
fishing can result in substantial postrelease mortality
and indirect stresses in addition to overfishing, including al(Muoneke and Childress 1994) and reduced growth and
tered flows and habitat fragmentation due to dams and other
fitness (Cooke et al. 2002). The extent of decline due to recreinfrastructure, pollution, habitat degradation, nonnative
ational fishing is often unappreciated, even in well-managed
species introductions, and detrimental interactions with
regions, because of inadequate records, lack of a historical
hatchery-reared fish (Allan and Flecker 1993). Although the

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Characteristics of inland fisheries

Inland fisheries are complex in their multigear and multispecies aspects, in their interannual variability as driven by abiotic factors, and in their social and economic context. Many
inland fisheries, particularly those of large tropical river
basins, occur within species-rich, ecologically diverse assemblages where population dynamics are difficult to observe and
interpret. As in most ecological communities, a few species are
highly abundant, more are moderately abundant, and many
are rare (Winemiller 1996); this distribution of abundances
also applies to the numerical distribution of fish species
caught by individual gears (Welcomme 1999). Another
widespread trend is for small individuals and species to
greatly outnumber large individuals and species (McDowall
1994), a consequence of the low ecological efficiency of
food chains, and of the coupling of high reproductive output and high juvenile mortality that characterizes most fish
species. About 50% of species present in any system do not
grow larger than 15 centimeters (cm) standard length, and
90% of species never grow larger than about 50 cm (figure
2; Welcomme 1999). Wherever larger fish are targeted, their
relatively small numbers and lower population growth
rates, relative to small species, make them more prone to depletion. In addition, a greater proportion of large species are
piscivorous, as illustrated by the trophic composition of
the fish species present in the various inland waters of West
Africa (figure 3). Although many exceptions can be cited
including the detritus-eating Mekong giant catfish (Pangasianodon gigas), the omnivorous major carp of India (e.g.,
Catla catla), and the herbivorous tambaqui of South America (Colossoma macropomum)in many instances removal
of the largest fish translates into the removal of apex predators, with the potential for substantial top-down effects on food
web dynamics.

To fully exploit the wide diversity of inland fish species, each

with its own habits and size range, mankind has developed
an equally extensive armory of fishing gears, broadly classified as active and passive (von Brandt 1984). These include
gill, cast, seine, lift, and hand nets; traps; lines; poisons; and
dynamite. In developing countries, high reliance on fishing,
together with the depletion of large high-value stocks, results
in the exploitation of a great variety of species and habitats
by diverse fishing methods and large concentrations of fishers (figure 4; Welcomme 2001). In affluent societies, in contrast, only a handful of fishing methods (gill nets, hoop nets,
longlines) are employed to capture those large species with
the highest economic value.

Fishing down in inland waters

As harvest intensity increases over a period of years, the classic model for a fishery predicts a rise in catch with increasing effort until reaching a relatively high level where the
maximum sustainable yield may be harvested over the long
term. Further increases in effort beyond this maximum are
expected to lead ultimately to declining catches and possible
collapse. The reality is far more complex. Especially when fishers are able to shift their effort to other members of the fish
assemblage, overfishing leads to numerous changes in both
the target species and the assemblage, and may not immediately cause declines in total catch.
As in marine systems, intensive, multispecies fishing in
inland waters can lead to what is known as fishing down the
food webthe successive removal of the larger elements of
a multispecies fish assemblage and their replacement by

Figure 2. Relationship between the maximum length attained by

a species (in centimeters) and the number of species attaining
that length, averaged for a number of river systems. Total number
of species is expressed as a cumulative percentage, with dotted
lines showing approximate maximum size attained by roughly
50% and 90% of the species pool. Data include the Niger, the
Mekong, the Magdalena, the So Francisco, North American
rivers, and several Indian rivers. Data are from Welcomme
(1999).
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importance of other anthropogenic stressors in relation to fishing may appear greater for inland waters than for the seas, the
contributions of pollution, species introductions, and other
human impacts in the collapse of coastal ecosystems should
not be underestimated (Boesch et al. 2001). As Jackson and
colleagues (2001) document in the context of marine fisheries,
overfishing commonly was the first disturbance in the historical progression, followed by other factors including pollution and eutrophication, mechanical habitat destruction,
introduced species, and climate change. Fish stocks are unable to recover from historical overfishing because of a host
of current pressures, and in their altered state may be more
vulnerable to disturbances, including species invasions and
outbreaks of disease. A recent assessment of inland fisheries
(FAO 1999) concluded that most inland capture fisheries
that rely on natural stock reproduction were overfished or being fished at their biological limit, and that the principal factors threatening inland capture fisheries were habitat loss
and environmental degradation. Overfishing, then, may not
always be the sole or even the primary threat, but in conjunction with other stresses it can be a serious one.

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smaller elements of the assemblage, which typically are at lower

trophic levels (Regier and Loftus 1972, Pauly et al. 1998).
Usually this means successive elimination of larger individuals and species, although some fish populations respond to
heavy fishing pressure with reductions in mean body size
and size at maturation. A synthesis of evidence from inland
waters supports the following general model (figure 5; Welcomme 2001). Fishing down initially leads to an increase in
the weight of total catch as the number of harvested species
and individuals increases, followed by a plateau or slight decline in total catch. As overfishing reduces the mean size of individuals and species in the assemblage, fishers reduce the
mesh size of gear they use. Where mesh sizes are universally
small, in the 2-cm size range, the assemblage may experience
intense selection on size and age at maturity. Even in the absence of data on mean fish length, consistent declines in net
mesh sizes may be an indicator of the state of the fishery,
as small-meshed nets are expensive and time-consuming
to make and usually will be adopted by fishers only out of
necessity.
Fisheries based on gill nets will successively concentrate on
smaller species and length classes as mesh sizes are reduced,
resulting in an increase in the number of species forming the
catch (figure 5c). In contrast, multigear fisheries are generally
designed to catch all species and ages of the target assemblage,
so the total number of species in the catch is high initially and
declines as large species are fished out. As the fishing-down
process causes larger species with slow life cycles to be replaced
by smaller species with more rapid life cycles, remaining
populations are expected to exhibit higher growth and mortality rates, higher ratios of productivity to biomass, and possibly population cycles with greater amplitude (Welcomme
2001).
In floodplain river fisheries, catch responds to the intensity of flooding in previous years, and the lag in response
1044 BioScience December 2005 / Vol. 55 No. 12

Figure 4. Fishing in the Mekong River basin, Cambodia.

(a, b) Cambodian fishermen crowd the river, using gill
nets and purse seines. One boat can catch in excess of 500
kilograms of fish over a 1-week period. (c) An aerial view
of bag nets, cone-shaped nets 25 meters in diameter and
125 meters long, which can catch up to 10 metric tons of
fish per day during the peak season. (d) Close view of a
bag net. (e) On the Tonle Sap Lake, local people live in
floating villages; they harvest fish using (f) arrow and
(g) fence traps. Bamboo fences often block migrating fish,
directing fish into traps. Small cyprinids, known collectively as trey riel, make up almost half of the total catch
from the Tonle Sap River and are so important that the
Cambodian currency (the riel) derives its name from the
fish. Photographs: Zeb Hogan.
indicates the time taken for a year class to enter the fishery.
In the 1950s, some river catches were correlated with floods
as much as four to five years earlier, indicating that the fishery targeted older fish and exhibited a low level of exploitation relative to potential. More recent catches often are
correlated with floods of the same year, indicating that the fish-

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Figure 3. The percentages of fish species in West African

rivers that are piscivorous in a given maximum length
class (standard length in centimeters). Data are from
Lvque and colleagues (1990, 1992).

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ery is based mainly on very young fish, and that it is at risk
(Welcomme 1979).

Evidence of overfishing

High-value individual targets

The declines of the Murray cod of the Murray-Darling river
system in Australia, some sturgeon stocks of Eurasia, the
tilapiine species Oreochromis esculentus and Oreochromis
variabilis of Lake Victoria, and perhaps the Pacific salmon of
the Columbia River are examples of the historical influence
of overharvest. The Mekong giant catfish and the Nile perch
of Lake Victoria provide contemporary cases.
The Murray cod. Native to the vast Murray-Darling river system of southern Australia, the Murray cod (Maccullochella
peelii) is a large (up to 114 kg), long-lived (estimated at 50 to
100 years), and relatively slow-growing species (Rowland
1989). A large fishery had developed by the 1860s, and government concern for the fishery was expressed as early as 1880.
Railroad consignments and sales in urban markets indicate
that catches increased to a peak in 1918 and then gradually
declined, and fishing became unprofitable for large operators
by the 1930s. The Depression and World War II may have contributed to a decline in fishing pressure, and cod populations
had recovered somewhat by the early 1950s.
The Murray cod then underwent a second, precipitous
decline, which most likely had several causes, illustrating the
wider panoply of threats that began to develop during the 20th

Figure 5. Characteristics of fishing down, including

trends in various parameters of a multispecies fish assemblage in response to increasing effort: (a) total catch;
(b) mean maximum length of assemblage and catch, and
mesh size of nets; (c) number of species accessible to netbased and multigear fisheries.
century. Catch statistics show an increase from 1940 to 1955,
and then a steep and rapid decrease to much lower levels by
the early 1960s. River regulation and species introductions
were probably the primary causes, although pollution and the
growth of recreational fishing may also have contributed
(Rowland 1989). The major impoundments on the MurrayDarling were constructed mainly after 1936, and their cumulative effect on river regulation was substantial by the
1950s1960s. Although reproduction in the cod occurs annually regardless of flooding, year-class strength is linked to
the occurrence of floods, which now are highly restricted. In
addition, Eurasian perch (Perca fluviatilis), an exotic, became
abundant by the 1940s, resulting in increased competition and
predation on young Murray cod.
The history of the Murray cod illustrates several points that
may have general application: A large, highly valued fish, selectively targeted during the latter half of the 19th century before diverse human influences were manifest, is reliably
known to have been fished beyond the level of commercial
profitability. Some recovery appears to have occurred when
fishing pressure was relaxed, but then river regulation, exotic
species, and pollution became severe. Reduced larval recruitment rather than capture of mature fish has become the
primary reason for low abundances, although the increasing
recreational fishery after the 1950s may also have contributed,
and hatchery stocking has played a substantial role in its recent recovery (Rowland 1989).
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Despite the challenge of evaluating the effects of fishing owing to complex system responses and the presence of other
pressures, there is ample evidence that overfishing is a significant factor in the decline of numerous species and fisheries, and is of global importance as a threat to inland water
biodiversity. We identify two main types of overfishing and
illustrate each type with case studies. In the first type of overfishing, intensive fishing of a targeted species leads to marked
declines in catch per unit effort and size of individuals captured. Such overfishing most likely was primarily responsible for the decline of a number of fish species prior to the
contributions of dams, habitat alteration, and pollution. In
the second type, known as assemblage or ecosystem overfishing (Murawski 2000), overfishing of an assemblage is
demonstrated when catches proceed beyond the asymptotic
maximum of a plateau-type curve or decline below the asymptotic or plateau level. Symptoms include sequential declines of species and depletion of individuals and species of
large size, especially piscivores; declines in the mean trophic
level of the assemblage; and changes in the responsiveness of
populations to environmental fluctuations (such as shorter
time lags). The associated decrease in size of the fish caught
is unacceptable in some parts of the world, such as Latin
America, but the preference for small fish in many African
cuisines and the use of fish pastes and sauces in Asia encourage harvest of extremely small species, thereby allowing
the entire assemblage to become depleted.

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Assemblage overfishing
Overfishing of an entire assemblage may be most common
in tropical regions, where fish diversity and the reliance of
local people on fish harvests both are high. However, it clearly
occurs in temperate latitudes as well, including in the Laurentian Great Lakes, where a number of valued species were
overharvested in succession.
Laurentian Great Lakes. The fish assemblage of the Laurentian Great Lakes has undergone continual change since the
earliest records, brought about by fishing, nutrient enrichment,
and myriad invasive species. However, major changes after the
late 1800s and continuing into the first half of the 20th century reflect an intensive and selective fishery targeting a succession of species and resulting in a succession of collapses
(Smith 1968). The lake sturgeon (Acipenser fulvescens) was intentionally overfished because this large fish frequently damaged gear intended for more valuable species, and by the late
1920s it was so reduced that restrictions were imposed. The
cisco (Coregonus artedi) experienced a collapse in the mid1920s attributed to overfishing, and while occasional strong
year classes occurred subsequently, their extreme rarity since
the mid-20th century is most likely due to the additional
pressure of increasingly unfavorable environmental conditions.
A trap net introduced in 1928 was a very effective new gear
for lake whitefish (Coregonus clupeaformis), and its use
expanded rapidly through the US (but not the Canadian)
waters of the Great Lakes. Although specific areas were rapidly
depleted, overall catch increased until the 1930s, when declines
in catch led to gear restrictions. Thus, several Great Lakes stocks
1046 BioScience December 2005 / Vol. 55 No. 12

were successively overfished in the early 20th century, when

additional threats emerged, including the establishment of the
invasive sea lamprey (Petromyzon marinus) after about 1940,
the expanding influence of pollution, and a further sequence
of invading species. Moreover, despite the loss of sturgeon and
the collapse of certain stocks in parts of the Great Lakes, the
total production of the fishery was relatively stable until the
1940s, although somewhat reduced from the high values
seen before 1920.
Oueme River fisheries. The Oueme River, Republic of Benin,
experienced intensive fishing by the 1950s, with the number
of fishermen estimated to be 25,000 in 1957 and 29,800 in 1968
(Welcomme 1971). The surrounding environment was still
largely undeveloped at that time, although it was used for intensive drawdown agriculture and cattle grazing. Assemblage
overfishing was evidenced by changes in the composition
and mean length of the fish caught. Large species reaching
maximum lengths of around 60 cm, such as Lates niloticus
(Nile perch), Heterotis niloticus, and Distichodus, were a significant fraction of the catch in the 1950s, but had disappeared
from the lower Oueme or reduced their size at first maturation by the 1970s. They were replaced by smaller species such
as Labeo, Clarias, Heterobranchus, Schilbe, Synodontis, and
larger mormyrids of maximum lengths of about 40 cm. As
fishing down continued into the 1990s, the fishery became
dominated by numerous small species of cichlids, mormyrids,
clariid catfishes, and the bagrid catfish Chrysichthys auratus,
attaining maximum lengths of 10 to 30 cm.
Tonle Sap fisheries. Overfishing along the Mekong River
threatens not only large species but the overall catch as well.
The number of fishers in the Tonle Sap River basin has increased from 360,000 in the 1940s to an estimated 1.2 million
in 1995 (Hortle et al. 2004). During the same period, catch per
fisher has decreased by 50%, but overall catch has nearly
doubled. Although large and medium-sized fish dominated
the 1940s catch, by 1996 the catch was heavily dominated by
small fish, largely because of increased fishing pressure and
assemblage overfishing. Small cyprinids now make up more
than 40% of the total catch of the Tonle Sap system, and
populations of large migratory catfish and carps have declined.
Fishers report that catches of river catfish have dropped by
90% in some fishing lots of the Tonle Sap Lake, for example,
from about 100 metric tons 20 years ago to just 5 metric
tons, or even 1 metric ton, today (Z. H., personal observation).
Nonetheless, the shift from large to smaller species is very difficult to demonstrate because of the lack of long-term data.
What is clear is that small, low-value cyprinids now dominate
the fishery, whereas anecdotal evidence indicates that migratory and larger speciesonce much more abundanthave
declined as a result of fishing pressure.

Ecosystem consequences of overfishing

Loss of apex predators results in a relaxation of top-down control of prey populations and stronger top-down control at the

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Large fish of the lower Mekong. The Mekong, the largest

river in Southeast Asia, is home to over 1200 species of fish
the highest level of fish diversity in any river system except the
Amazon and the Congoincluding some of the largest
species of freshwater fish in the world (figure 6). Several
species of Mekong fish are now endangered, and the wild population of at least one species, the Mekong giant catfish, is close
to extinction (Hogan et al. 2004). This species, a Mekong
endemic, reaches a maximum length of 3 meters and a total
weight of up to 300 kg. A large fishery had developed for the
Mekong giant catfish in Luang Prabang (Laos) as early as 1890,
and declines were observed in northeastern Thailand around
1940. Recent data reveal a fishery characterized by a pattern
of increasing catch and increasing effort, followed by a declining catch with sustained high fishing effort, typical of an
overexploited population (figure 7).
The same trend appears to be taking place with other large
Mekong species. Anecdotal and published records (Mattson
et al. 2002) point to the steep decline of the catch of large
species such as the Sutchi catfish (Pangasianodon hypophthalmus), the giant barb (Catlocarpio siamensis), and the
freshwater whipray (Himantura chaophraya; figure 6).
Indeed, most of the worlds largest freshwater fish are at risk
according to the IUCN Red List (table 1), and overexploitation contributes in a number of these cases.

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Table 1. Conservation status of the worlds largest freshwater fish species.
Common
name

Scientific
name

Maximum
size

Distribution

IUCN Red List

category

Major
threats

Mekong giant catfish

Pangasianodon gigas

300 cm, 300 kg

Mekong River basin

Critically endangered

Harvest, habitat
loss

Giant barb
Isok barb
Freshwater whipray

Catlocarpio siamensis
Probarbus jullieni
Himantura chaophraya

300 cm, 300 kg

180 cm, 100 kg
500 cm, 600 kg

Mekong River basin

Mekong River basin
Mekong River basin

Not evaluated
Endangered
Vulnerable

Giant pangasius
Sutchi catfish

300 cm, 300 kg

250 cm

Mekong River basin

Mekong River basin

Data deficient
Not evaluated

Goonch
Largetooth sawfish

Pangasius sanitwongsei
Pangasianodon
hypophthalmus
Bagarius yarrelli
Pristis microdon

200 cm
650 cm

Mekong River basin

Mekong River basin

Not evaluated
Endangered

Piraba or valentn
(giant catfish)
Pirarucu (bonytongue)
Huchen

Brachyplatystoma
filamentosum
Arapaima gigas
Hucho hucho

360 cm, 200 kg

Amazon and Orinoco

River basins
Amazon River basin
Danube River basin

Not evaluated

Taimen

Hucho taimen

200 cm, 100 kg

Not evaluated

Chinese paddlefish

Psephurus gladius

300 cm, 300 kg

Selenge River basin

(Lake Baikal)
Yangtze River basin

Yangtze sturgeon

Acipenser dabryanus

250 cm

Yangtze River basin

Critically endangered

Murray cod

Maccullochella peelii

200 cm, 113 kg

Critically endangered

Nile perch

Lates niloticus

200 cm, 200 kg

Wels catfish

Silurus glanis

500 cm, 306 kg

Colorado pikeminnow
Alligator gar
Lake sturgeon

Ptychocheilus lucius
Atractosteus spatula
Acipenser fulvescens

200 cm
305 cm, 137 kg
274 cm, 125 kg

Tigris River salmon

Barbus esocinus

230 cm, 136 kg

Murray River basin

(Australia)
Congo, Niger, and Nile
River basins
Widespread in Europe
and Asia
Colorado River basin
Mississippi River basin
Saint Lawrence, Great
Lakes
Tigris River basin

Data deficient
Endangered

Critically endangered

Harvest, habitat
loss

Harvest
Harvest, habitat
loss

Harvest, habitat
loss
Harvest, habitat
loss
Harvest, habitat
loss

Not evaluated
Not evaluated
Vulnerable
Not evaluated
Vulnerable

Habitat loss
Harvest, habitat
loss

Not evaluated

Source: IUCN Red List (www.redlist.org/info/categories_criteria2001.html).

next trophic level below. Because species richness tends to be

lower at higher trophic levels, the likelihood that another
species will fill the role of an extirpated apex predator may be
diminished (Raffaelli 2004). Although the influence of predator composition and diversity on ecosystem stability remains
poorly understood, a synthesis of experimental studies found
significant destabilizing effects of predator removals on herbivore biomass (Halpern et al. 2005). Exclusion of top predators also has been found to release smaller predators that
control dominant herbivores, thereby altering algal assemblages and their associated invertebrates. Because herbivorous
fishes can regulate primary producers through direct grazing
on benthic algae (Flecker 1996) and macrophytes (Bain
1993), changes in piscivore density can have a marked influence on primary production and abundance of basal resources via indirect pathways in trophic cascades.
Freshwater fishes also influence nutrient dynamics in freshwater ecosystems, both directly via excretion and indirectly
via grazing and bioturbation of algae and detritus. Detritivorous gizzard shad (Dorosoma cepedianum) have been shown
to affect inorganic nutrient ratios and planktonic primary pro-

duction in a North American lake (Vanni 1996). Sediment

grazing by fishes altered abundances of attached algae and the
response of algae to limiting nutrients in an Andean piedmont
river in Venezuela (Flecker et al. 2002).
One of the most spectacular and certainly best documented influences of fish on the supply of nutrients is due to
the death and decay of great spawning runs of Pacific salmon
after entering fresh waters. As salmon populations have decreased because of overfishing and other causes, declines
have also occurred in lake productivity and juvenile salmon
recruitment, leading to a number of fertilization experiments. Virtually without exception, the addition of nutrients
has resulted in greater algal production and increased biomass
of zooplankton and salmon smolts, and a few studies report
greater survival of smolts and adults as well (Hyatt et al.
2004).
Although artificial fertilization can replicate the nutrient
subsidy of formerly large spawning runs, a reduction in the
seasonal flux of anadromous fishes into inland waters has
serious implications for a diverse assemblage of terrestrial and
semiaquatic animals, and terrestrial plant assemblages. In
December 2005 / Vol. 55 No. 12 BioScience 1047

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450 cm, 200 kg

200 cm

Harvest
Harvest, habitat
loss
Harvest

Articles

North America, spawning salmon and their fry have been

shown to be important to a wide variety of mammals, from
mink to North American brown bears, as well as to piscivorous birds; furthermore, salmon influence the densities of insectivorous passerine birds in the riparian areas of salmon
streams as a result of the indirect effects of salmon on insect
prey (Willson and Halupka 1995). Nutrients derived from decaying fish that are transported into terrestrial habitats by subsurface flow, flooding, and consumers fertilize terrestrial
ecosystems and enhance the growth and diversity of plants and
soil microbes (Gende et al. 2002). Thus, declines in spawning runs of anadromous fishes result in the loss of an important flux of resources into terrestrial systems.
Mass migrations of fishes within fluvial basins are responsible for active transport of nutrients and energy at the
landscape scale. Prochilodontid fishes, the major species in
commercial fisheries in the large river basins of South America, are renowned for seasonal migrations of hundreds of
kilometers, connecting food webs of distant, and sometimes
divergent, ecosystems. During the falling-water period, young
prochilodontids migrate en masse from productive whitewater
floodplains into unproductive blackwater rivers, thus subsidizing apex predators in a species-rich food web (Winemiller
and Jepsen 2004). Large pimelodid catfishes of the Amazon
and Orinoco Rivers migrate many hundreds of kilometers between nursery areas in the lower river reaches and adult feed1048 BioScience December 2005 / Vol. 55 No. 12

ing and spawning areas in the upper reaches of the basins.

These large piscivores may induce top-down effects on local

Figure 7. Yield as a function of effort for the Mekong

giant catfish fishery in Chiang Khong, Thailand. The
initial increase in catch as effort increases, followed by a
steep decline in catch but sustained high effort, may indicate a fisheries crash. In the past, increases in fishing
effort have resulted in increased catch. This is no longer
the case today. Source: Giant Catfish Fishermens Club
of Chiang Khong.

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Figure 6. Large fish species of the Mekong River: (a) Mekong giant catfish, Pangasianodon gigas;
(b) Sutchi catfish, Pangasianodon hypophthalmus; (c) freshwater whipray, Himantura chaophraya;
(d) giant barb, Catlocarpio siamensis. Photographs: Zeb Hogan.

Articles

Future directions
Overfishing threatens both the biodiversity of inland waters
and the ecosystem goods and services on which people rely.
However, its importance as a threat is underappreciated, because intensive fishing frequently acts synergistically with
other pressures, and its consequences for inland fisheries
and ecosystems are poorly understood and documented.
The development community is beginning to appreciate the
need to promote sustainable catches instead of increased
production (World Bank 2003), but individual countries,
municipalities, villages, and even some fisheries managers may
not be as farsighted.
Overfishing is being driven by overcapacity and excess effort, which in turn are due to the generally open access
regimes of many inland fisheries and to the effective use of
fisheries as an occupation of last resort in developing
economies. Managing fisheries today is not limited just to satisfying the commercial fishing industry, but must accommodate the wide array of economic and social benefits that
people derive from freshwater ecosystems, including food
security and economic growth. The practical effect of this is
a widening of the group of stakeholders that have legitimate
interests in how fisheries are managed. Setting up appropriate institutional structures and legal frameworks that will

allow wider stakeholder participation in resource management

is essential for the successful implementation of better fishery management strategies. Proposals have been made on
many rivers for co-management systems that will substitute
for the previously centralized approaches (Hartmann et al.
2004). Such decentralized systems will by definition involve
local people to a greater degree in decisionmaking, resulting
in more flexible management systems, with greater likelihood of formulating and enforcing regulations that correspond best to the needs of the fishery at the local level.
Fishery science and management is shifting its focus from
single species to ecosystem-based fishery management (Pikitch et al. 2004). Assessing the effects of fishing on communitywide interactions and on ecosystem structure and function
is a new challenge, and extending this assessment beyond fish
populations (e.g., to benthos or producers) will be especially
difficult. Harvest reserves and no-take zones, strategies with
similar potential for achieving benefits beyond the fishery itself, have attracted impressive attention from the marine
conservation and management communities (Hilborn et al.
2004) and also merit greater attention in inland waters (Hoggarth et al. 1999). Owing to the high degree of impact that
freshwater systems experience from upstream and upland environmental threats, reserves in inland waters should be designed within a framework of integrated river basin
management.
Until knowledge of ecosystem processes and their response
to human actions is better developed, we suggest that four core
principles should guide the management of inland fisheries
for the long-term benefit of the widest range of stakeholders,
including the environment: sustainability of yields, maintenance of biodiversity, protection from other anthropogenic
stressors (habitat degradation, invasive species, pollution,
etc.), and provision of socioeconomic benefits to a broad
spectrum of resource consumers. Fishery management should
be guided by a precautionary approach in setting management
targets and limits (Pikitch et al. 2004). Managing inland fisheries, in most cases, will require developing plans that consider pressures occurring across multiple scales, through a
process that includes both fishers and other stakeholders
who have an interest in how a basins resources are used.
Biodiversity conservation and fishery management should be
undertaken together, as the two constituencies ultimately
have many shared goals.

Acknowledgments
We thank those who provided us with case studies and documentation, including Stuart Rowland, Jean Jackson, Alistair
Richardson, and Ronald Bjorkland. Helpful comments on earlier drafts of the manuscript were provided by Brian Allan,
Jim Diana, Peter Esselman, Ben Halpern, Ed Houde, Michael
Nussman, Margaret Palmer, and three anonymous reviewers.
J. D. A. thanks the David H. Smith Program of The Nature
Conservancy for support during the initial development of
this manuscript.
December 2005 / Vol. 55 No. 12 BioScience 1049

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fish populations, particularly when they enter rivers that

drain unproductive watersheds (Winemiller and Jepsen 1998).
Commercial fisheries have caused declines in at least one of
these migratory catfish (Brachyplatystoma vaillantii) in the
lower reaches of both major South American rivers (Barthem
and Goulding 1997).
Finally, overfishing of inland waters has the potential for
severe and unexpected impacts on the health of human populations, particularly in developing countries. For example,
the trematode parasite responsible for schistosomiasis is vectored by snails of several genera that are consumed by numerous species of cichlid fishes. Substantial increases in
recent years in the human incidence of schistosomiasis in
freshwater systems, such as Lake Malawi, have been attributed
to increases in vector snail populations released from predatory control by cichlid fishes, a likely consequence of overfishing (Stauffer et al. 1997). Investigations of the effects of
overfishing on waterborne diseases such as schistosomiasis and
cholera are currently in their infancy, but the consequences
for human populations of depleting fish assemblages in these
systems may be profound.
In sum, the functional or actual elimination of species
from the ecosystems of inland waters is likely to have numerous consequences of varying severity. Progressive reductions in assemblage diversity mean that fewer species are
available to perform critical functions, and consequences
will be greatest when species with disproportionately strong
influences on nutrient, habitat, or assemblage dynamics are
lost. The ecological correlates of overfishing, although unevenly documented, almost certainly are profound and widespread.

Articles
References cited

1050 BioScience December 2005 / Vol. 55 No. 12

Downloaded from http://bioscience.oxfordjournals.org/ by guest on October 22, 2014

Ahmed M, Navy H, Vuthy L, Tiongco M. 1998. Socio-economic Assessment of Freshwater Capture Fisheries of Cambodia: Report on a Household Survey. Phnom Penh (Cambodia): Mekong River Commission.
Allan JD, Flecker AS. 1993. Biodiversity conservation in running waters. BioScience 43: 3243.
Bain MB. 1993. Assessing impacts of introduced aquatic species: Grass carp
in large systems. Environmental Management 17: 211224.
Barthem R, Goulding M. 1997. The Catfish Connection: Ecology, Migration,
and Conservation of Amazonian Predators. New York: Columbia
University Press.
Boesch D, Burreson E, Dennison W, Houde E, Kemp M, Kennedy V, Newell
R, Paynter K, Orth R, Ulanowicz R. 2001. Factors in the decline of
coastal ecosystems. Science 292: 15891591.
Cooke SJ, Cowx IG. 2004. The role of recreational fishing in global fish
crises. BioScience 54: 857859.
Cooke SJ, Schreer JF, Dunmall KM, Phillip DP. 2002. Strategies for quantifying sub-lethal effects of marine catch-and-release anglinginsights
from novel freshwater applications. American Fisheries Society Symposium 30: 121134.
[FAO] Food and Agriculture Organization of the United Nations. 1999.
Review of the State of World Fishery Resources: Inland Fisheries. Rome:
FAO Fisheries Department. FAO Fisheries Circular no. 942.
. 2002. The State of World Fisheries and Aquaculture 2002. Rome:
FAO Fisheries Department.
. 2003. Food Balance Sheet. FAOSTAT database. Rome: FAO. (20 October 2005; http://faostat.fao.org/)
. 2004. FISHSTAT Plus: Universal software for fishery statistical time
series. Version 2.30. (8 October 2005; www.fao.org/fi/statist/FISOFT/
FISHPLUS.asp)
Flecker AS. 1996. Ecosystem engineering by a dominant detritivore in a
diverse tropical stream. Ecology 77: 18451854.
Flecker AS, Taylor BW, Bernhardt ES, Hood JM, Cornwell WK, Cassatt SR,
Vanni MJ, Altman NS. 2002. Interactions between herbivorous fishes and
limiting nutrients in a tropical stream ecosystem. Ecology 83: 18311844.
Gende SM, Edwards RT, Willson MF, Wipfli MS. 2002. Pacific salmon in
aquatic and terrestrial ecosystems. BioScience 52: 917928.
Halpern BS, Borer ET, Seabloom EW, Shurin JB. 2005. Predator effects on
herbivore and plant stability. Ecology Letters 8: 189194.
Hartmann W, Dugan P, Funge-Smith S, Hortle K, Kuemlanagen B, Lorenzen K, Marmulla G, Mattson N, Welcomme R. 2004. People and fisheries
management. Pages 6192 in Welcomme RL, Petr T, eds. Proceedings of
the Second International Symposium on the Management of Large
Rivers for Fisheries, vol. 1. Bangkok (Thailand): Food and Agricultural
Organization of the United Nations, Regional Office for Asia and the
Pacific. RAP Publication 2004/16.
Hilborn R, Branch TA, Ernst B, Magnusson A, Minte-Vera CV, Scheuerell MD,
Valero JL. 2003. State of the worlds fisheries. Annual Review of Environment and Resources 28: 359399.
Hilborn R, et al. 2004. When can marine reserves improve fisheries management? Ocean and Coastal Management 47: 197205.
Hogan ZS, Moyle PB, May B, Vander Zanden MJ, Baird IG. 2004. The
imperiled giants of the Mekong. American Scientist 92: 228237.
Hoggarth DD, Cowan VJ, Halls AS, Aeron-Thomas M, McGregor JA,
Garaway CA, Payne AI, Welcomme RL. 1999. Management Guidelines
for Asian Floodplain River Fisheries, part 1: A Spatial, Hierarchical and
Integrated Strategy for Adaptive Co-management. Rome: Food and
Agricultural Organization. FAO Fisheries Technical Paper no. 384/1.
Hortle KG, Bush SR. 2003. Consumption in the Lower Mekong Basin as a
measure of fish yield. Pages 7682 in Clayton T, ed. New Approaches for
the Improvement of Inland Capture Fishery Statistics in the Mekong
Basin. Bangkok (Thailand): Food and Agriculture Organization of the
United Nations, Mekong River Commission.
Hortle KG, Lieng S, Valbo-Jorgensen J. 2004. An Introduction to Cambodias
Inland Fisheries. Phnom Penh (Cambodia): Mekong River Commission.
Mekong Development Series no. 4.

Hyatt KD, McQueen DJ, Shortreed KS, Rankin DP. 2004. Sockeye salmon
(Oncorhynchus nerka) nursery lake fertilization: Review and summary
of results. Environmental Reviews 12: 133162.
Jackson JBC, et al. 2001. Historical overfishing and the recent collapse of coastal
ecosystems. Science 293: 15891591.
Kura Y, Revenga C, Hoshino E, Mock G. 2004. Fishing for Answers: Making
Sense of the Global Fish Crisis. Washington (DC): World Resources
Institute.
Laurenti G. 2002. Fish and Fishery Products: World Apparent Consumption
Statistics Based on Food Balance Sheets. Rome: FAO Fisheries Department. FAO Fisheries Circular no. 821, rev. 6.
Lvque C, Paugy D, Teugels GG, eds. 1990. The Fresh and Brackish Water
Fishes of West Africa, vol. 1. Paris: Editions de lORSTOM.
. 1992. The Fresh and Brackish Water Fishes of West Africa, vol. 2.
Paris: Editions de lORSTOM.
Mattson N, Buakhamvongsa K, Sukumasavin N, Tuan N, Vibol O. 2002.
Cambodian Giant Fish Species: On Their Management and Biology.
Phnom Penh (Cambodia): Mekong River Commission. MRC Technical
Paper no. 3.
McDowall RM. 1994. On size and growth in freshwater fish. Ecology of
Freshwater Fish 3: 6779.
Muoneke MI, Childress WM. 1994. Hooking mortality: A review for recreational fisheries. Reviews in Fisheries Science 2: 123156.
Murawski S. 2000. Definitions of overfishing from an ecosystems perspective. ICES Journal of Marine Science 57: 649658.
Pauly D, Christensen V, Dalsgaarrd, Froese JR, Tores F. 1998. Fishing down
marine food webs. Science 278: 860863.
Pauly D, Christensen V, Gunette S, Pitcher TJ, Sumaila UR, Walters CJ,
Watson R, Zeller D. 2002. Towards sustainability in world fisheries.
Nature 418: 689695.
Pikitch EK, et al. 2004. Ecosystem-based fishery management. Science 305:
346347.
Post JR, Sullivan M, Cox S, Lester NP, Walters CJ, Parkinson EA, Paul AJ,
Jackson L, Shuter BJ. 2002. Canadas recreational fisheries: The invisible
collapse? Fisheries 27: 617.
Raffaelli D. 2004. How extinction patterns affect ecosystems. Science 306:
11411142.
Regier HA, Loftus KH. 1972. Effects of fisheries exploitation on salmonid communities in oligotrophic lakes. Journal of the Fisheries Research Board
of Canada 29: 959968.
Ricciardi A, Rasmussen JB. 1999. Extinction rates of North American freshwater fauna. Conservation Biology 13: 12201222.
Roberts CM, et al. 2003. Application of ecological criteria in selecting
marine reserves and developing reserve networks. Ecological Applications
13: S215S228.
Rowland SJ. 1989. Aspects of the history and fishery of the Murray cod,
Maccullochella peeli (Mitchell) (Percichthyidae). Proceedings of the
Linnean Society of New South Wales 111: 201213.
Smith SH. 1968. Species succession and fishery exploitation in the Great Lakes.
Journal of the Fisheries Research Board of Canada 25: 667693.
Stauffer JR, Arnegard ME, Cetron M, Sullivan JJ, Chitsulo LA, Turner GF,
Chiotha S, McKaye KR. 1997. Controlling vectors and hosts of parasitic
diseases using fishes. BioScience 47: 4149.
Vanni MJ. 1996. Nutrient transport and recycling by consumers in lake
food webs: Implications for algal communities. Pages 8195 in Polis GA,
Winemiller KO, eds. Food Webs: Integration of Patterns and Dynamics.
New York: Chapman and Hall.
von Brandt A. 1984. Fish Catching Methods of the World. Surrey (United
Kingdom): Fishing News Books.
Watson R, Pauly D. 2001. Systematic distortions in world fisheries catch.
Nature 414: 534536.
Welcomme RL. 1971. Evolution de la pche intrieure, son tat actuel et ses
possibilits. Report to the Government of the Republic of Dahomey.
Rome: Food and Agriculture Organization of the United Nations/United
Nations Development Programme.
. 1979. Fisheries Ecology of Floodplain Rivers. London: Longman.

Articles
. 1999. A review of a model for qualitative evaluation of exploitation
levels in multi-species fisheries. Fisheries Management and Ecology
6: 119.
, comp. 2001. Inland Fisheries: Ecology and Management. Oxford
(United Kingdom): Fishing News Books.
Willson MF, Halupka KC. 1995. Anadromous fish as keystone species in
vertebrate communities. Conservation Biology 9: 489497.
Winemiller KO. 1996. Dynamic diversity: Fish communities of tropical
rivers. Pages 99134 in Cody ML, Smallwood JA, eds. Long-term
Studies of Vertebrate Communities. Orlando (FL): Academic Press.

Winemiller KO, Jepsen DB. 1998. Effects of seasonality and fish movement
on tropical river food webs. Journal of Fish Biology 53 (suppl. A):
267296.
. 2004. Migratory neotropical fish subsidize food webs of oligotrophic blackwater rivers. Pages 115132 in Polis GA, Power ME, Huxel
GR, eds. Food Webs at the Landscape Level. Chicago: University of
Chicago Press.
World Bank. 2003. Reaching the rural poor: A renewed strategy for rural
development. Washington (DC): World Bank.

Downloaded from http://bioscience.oxfordjournals.org/ by guest on October 22, 2014

December 2005 / Vol. 55 No. 12 BioScience 1051