Zebrafish female top, male bottom. Photo by |
Zebrafish is member of the minnow family
(Cyprinidae); it is native to the southeastern Himalayan region. Zebrafish readily adapt to life in aquaria
and they have a fast development time and short generation time; for these
reasons they have become a model fish for laboratory investigations, typically
development, gene function, and toxicology.
Atlantic Menhaden is a member of the herring family (Clupeidae); it lives in estuaries and coastal waters from Nova Scotia to northern Florida. They are filter feeders that swim with their mouth open and gill openings spread. They travel in large schools and are preyed upon by many fish, marine mammals, and birds, Atlantic Menhaden mature in two years at a size of 18-22 cm; females produce between 100,000 to 600,000 eggs, depending on size. Common names include the pogy, mossbunker, bunker, fat-back, and bug-mouth. Native Americans called them 'munnawhatteaug' (=fertilizer).
Atlantic Menhaden |
Red
Snapper is a long-lived, early-maturing fish in the family Lutjanidae. They are broadly distributed in the Gulf of
Mexico and are an important sport and commercial fish. Red Snapper have been overfished for several
decades and are one of the most controversial fisheries with high landings from both recreational and commercial fishers.
Red Snapper caught in January 2013 was 38.25 inches and was released due to closed season. |
This
gradual decline in the average size of fishes caught in intensely harvested
populations has been documented in numerous fisheries (Silliman 1975; Ricker
1981; Sharpe and Hendry 2009; Arlinghaus et al. 2010). Cautious managers warned that intensive size
selective fishing could lower fisheries productivity over time (Hutchings 2009;
Rijnsdorp et al. 2010; Heino et al. 2013). The effects of size selective fishing are not
only plausible, but potentially widespread.
The
mathematics of optimum size based harvesting (Reed 1980) is based on the
assumptions of stationarity and identical fish in each age group. The population is further assumed to interact
with no others (as predator or prey or competitor). The maximization of yields based on Reed’s mathematical
model demonstrated that under intense levels of exploitation, harvesting
restricted to two age groups maximizes yields. The mathematical theorem does not support the
application of minimum size limits as an optimal strategy. Rather, the optimal size to harvest depends on
exploitation rate. And old, more fecund
age groups, are protected from harvest.
Uusi-Heikilla and others (2015) recently
explored the effects of intensive harvest on Zebrafish populations to address
the controversy of whether harvesting causes genetic as opposed to mere
phenotypic changes. The question is
critically important to fisheries manager as genetic changes are more slowly
reversible. The use of Zebrafish allowed
the investigators to follow populations over 5 generations of intense size
selective pressure and examine functional genomic markers related to life
history traits. The parental stock
began with 1500 wild-collected Zebrafish to ensure maximum genetic variation.
The design employed three experimental treatments, with two replicates per
treatment, and 450 zebrafish per replicate tank. Harvest rate was 75% per generation. Treatments were designed to mimic random size
selection, minimum size limit (size selective), and maximum size limit (large
size selection). In addition to life
history traits, the investigators examined 371 single nucleotide polymorphisms
to see if allelic frequencies differed among treatments.
As
hypothesized, the minimum length limits had surprising undesired genetic effects.
Twenty-two loci showed genetic differentiation and eight of these were
statistically significant. After just
five generations of harvesting, adult body size of the Zebrafish shrunk by 7%,
which also affected the egg production of the surviving fish. The now-smaller
Zebrafish produced fewer and smaller eggs!
Furthermore, the large-selected
Zebrafish were significantly more explorative and bolder. This experiment
supports a cause-and-effect relationship between size-selective harvest
practices and changes in fish life history traits and productivity. The implications for recovery of overfished
populations are important.
There
are at least three options to restore overfished fisheries: (1) moratorium on
all harvest; (2) daily creel limits or restrictive size limits, or (3) protect
large fecund individuals via a maximum size limit or protected area. A harvest moratorium rebuilt populations and
restored age structure in Atlantic striped bass (Morone saxatilis) (Richards and Rago 1999; Secor, 2000). A
transition from minimum length limit to a protected slot limit transformed an
extremely truncated size distribution in a smallmouth bass fishery to a trophy
fishery (Copeland et al. 2006). The designation
of marine protected reserves is a tool for protecting species from fishing
(Bohnsack et al. 2004). Reserves permit
recovery of overexploited fish populations if larval export seeds exploited
habitats (Harrison et al. 2012).
Many
commercially or recreationally important fish species have high fecundity and
display a weak response between parents and recruits. Big yearclasses can still be produced by low spawning stocks. This leaves most fish managers with doubt
about strong effects of size selective harvest.
Further, not all fish have the same life history and role. Small fish
species should be exploited at levels well below those producing a maximum
sustainable yield. This will provide
some forage fish to survive to feed higher trophic levels (Pikitch et al 2014). Fishing forage fish intensely increases
likelihood of collapse (Essington et al. 2015).
The
Atlantic menhaden is exploited with a reduction fishery (i.e, it reduces the catch, into fishmeal and fish oil) and a bait fishery. The
fishery expanded from New England after the Civil War to the rest of the
coastal states and peaked by 1950, when over 20 reduction factories processed
the harvest. Since the 1960s the
menhaden fluctuated and reduction factories closed and reopened, until odor
abatement regulations caused most of them to close. Today a single reduction factory in
Reedville, Virginia, processes the landings; Virginia is allocated 85% of the
total allowable catch. Menhaden
constitute the largest landings, by volume, along the east coast, and rank
second in the U.S behind the Alaskan Pollock. The Atlantic Menhaden were once
overfished and sport and commercial fisheries dependent on the Atlantic Menhaden
(e.g., Menhaden Defenders) have
lobbied hard to change the way this fishery is managed. Not too many forage fish have their own lobbying
group. Yet, the total allowable catch
in recent years is still managed based on single species concepts, as if the
Atlantic Menhaden was not eaten by a myriad of fish, mammals, and birds. Talk about your inappropriate spherical
cow!
Based
on the latest stock assessment, the Atlantic menhaden “stock status is not
overfished and overfishing is not occurring” (SEDAR 2015). But nothing in this 643-page assessment
addresses the influence of abundance of Atlantic Menhaden on striped bass,
bluefish, weakfish, tuna, or other coastal fishes. Atlantic Menhaden holds the distinction of being the only fish species controlled by the General Assembly and not the Virginia
Marine Resources Commission. The politicians, not the professionals, make
decisions.
The
Red Snapper fishery in the Gulf of Mexico collapsed in the late 1980s. Because the fish and fishery occur in all
Gulf states and federal waters, the fishery is managed by the Gulf Fishery Management
Council under the authority of the Magnuson-Stevens Fisheries Management Act. The council was slow to enact protections and, therefore, the Red
Snapper remained overfished for several decades. Even in an overfished state, the economic
value of the fisheries is $80 million. The
Red Snapper story illustrates why the simple theory was difficult to implement (Cowan
et al. 2010). First, bycatch of juvenile
Red Snapper in shrimp trawls was not regulated and caused high mortality among
immature Red Snapper. Second, management
plans required the allocation of allowable catch to recreational and commercial
fisheries. Third, regulations such as
minimum size limits resulted in regulatory discard mortality, that is fish that
are required by law to be released due to size, season, or bag
requirements. Undersized fish had to be
released, much to the distress of anglers who assumed that many of the fish,
caught from deeper waters and exhibiting protruding swimbladders, would die
anyway. Finally, artificial reefs and
oil and gas platforms were deployed in the 1970s and 1980s and used as a red
herring, as if the artificial structures would increase Red Snapper abundance
(Galloway et al. 2009).
The
Red Snapper is a long-lived species that can live over 50 years, but some will
mature at age 3 or 4. The large old fish are highly fecund. Red Snapper have been called “bet hedgers” as
an evolutionary strategy – females produce millions of very small eggs over her
lifetime, with an infinitesimally small chance of surviving to be an adult Red
Snapper.
Concept Map depicting cause and effect relationships of big old fat fecund female fish (BOFFF; Hixon et al. 2013) |
Intense exploitation in the Gulf fishery
truncates the age distribution such that few fish over 5 years old
survive. Fish are gaining weight at a
very high rate at this age range from 5 to 20 years. Also the fecundity is much higher for these
big old fat fecund female fish (BOFFFF; Hixon et al. 2013). Older red snapper also spawn more frequently and a 32-inch female produces 24 times as many eggs as a 16-inch female (Porch et al. 2013).
One
major factor affecting recruitment of Red Snapper is periodic occurrence of
hypoxia, which caused the loss of all red snapper and most of the invertebrates
on and around the reefs in 2001 (Workman and Foster 2002). Since
this periodic recruitment failure is an expected occurrence, it makes sense to
maintain a larger spawning stock biomass so that large year classes of Red
Snapper may be produced in good years.
Texas State record Red Snapper (40 pounds) caught June 1, 2014. Source |
These
three fishes teach us a lot about exploited populations of fish. Any assessment of the status of an exploited
fish population and evaluation of alternative management strategies requires
certain assumptions. However, sometimes
I feel like choking on the spherical cow.
As we move forward we need to
apply lessons learned from other fisheries and follow common sense (KISS) principles.
· Little fish feed the
world. Total allowable catch should be adjusted based on needs of predators or ecosystem services provided.
· Protect the BOFFFF --
they survive unfavorable recruitment times and facilitate resilience.
· Size-selective
harvesting causes changes in key life-history traits, leading to low maximum
body size and poor reproductive output.
· Intense fishing selects
for shy behaviors.
· Effects of fishing
proceed at a faster pace than the scientific approach to evaluating actions. The best available science often adopts simplifying assumptions.
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