The
Atlantic menhaden population is subject to a high natural
mortality rate. There is a somewhat reduced probability
of death from natural causes when the population is
being harvested. Natural mortality is also higher during
the first two years of life than during subsequent years.
Scientific studies indicate an annual instantaneous
natural mortality rate (M) of 0.45 in the absence of
fishing; this rate is equivalent to an annual reduction
in population numbers of 36%. This rate is quite high
compared to other pelagic marine species. Atlantic herring,
for example, is characterized by an 18% annual natural
mortality rate. During the 1955-1987 period, under exploitation,
the annual natural mortality rate for age-1 Atlantic
menhaden was 30% and, for ages 2 and older, it was 20%.
Natural mortality removes an estimated 30% of the exploited
population at age 1 and 20% each year thereafter.
Menhaden
natural mortality is probably due primarily to predation,
since the fish are so abundant in coastal waters during
the warmer months of the year. All large piscivorous
sea mammals, birds, and fish are potential
predators on Atlantic menhaden. Menhaden are preyed
upon by species such as bluefish, striped bass, king
mackerel, spanish mackerel, pollock, cod, weakfish,
silver hake, tunas, swordfish, bonito, tarpon, and a
variety of sharks .
Coastal
pollution and habitat degradation threaten marine fish
species, such as Atlantic menhaden, which spend their
first year of life in estuarine waters and the rest
of their life in both ocean and estuarine waters.
Other
poorly understood sources of natural mortality for Atlantic
menhaden are diseases and parasites. There is no information
available concerning the extent of parasitism or its
possible effect on survival. Studies described the incidence
of ulcerative mycosis (UM), a fungal infestation which
was observed in menhaden over much of their range in
1984 and 1985 and in a more restricted area in 1986.
A large fish kill in Pamlico Sound, North Carolina in
November, 1984 was associated with UM, but its primary
effect may be to weaken fish, making them more susceptible
to other causes of mortality, such as predation, parasites,
other diseases, and low dissolved oxygen concentrations.
The overall impact of UM on the 1984 and 1985 year classes
could not be assessed, but it was not believed to be
significant. However, other scientists believed that
the mortality effects of a disease or other event must
be "truly catastrophic" to be detectable.
Another
source of natural mortality for Atlantic menhaden (and
many other species) may be "red tide." The term refers
to the color of water caused by the rapid multiplication
("bloom") of single-celled planktonic organisms called
dinoflagellates, which produce a toxic compound. The
toxin accumulates in the tissues of filter-feeding animals
which ingest the dinoflagellate. An outbreak of red
tide occurred along the coast of the Carolinas during
November, 1987 - April, 1988 when Gulf Stream water
containing the dinoflagellates was transported into
coastal waters. Menhaden recruitment in Beaufort Inlet
during this period was severely reduced.
A
new species of toxic dinoflagellate was identified as
the causative agent in a major menhaden kill in the
Pamlico River, North Carolina, in May, 1991. Problems
with toxic phytoplankton organisms may increase in the
future since their appearance has been correlated with
increasing nutrient enrichment in estuarine and coastal
waters which are subject to increasing organic pollution.
An
additional source of mortality are fish "kills" which
occur when schools of menhaden enter enclosed inshore
bodies of water in such large numbers that they consume
all available oxygen and suffocate. The mean lethal
dissolved oxygen concentration for menhaden has been
reported to be 0.4 mg/l. Bluefish are known to follow
(or even chase) schools of menhaden inshore, feeding
on them, and may contribute to their mortality by preventing
them from leaving an area before the oxygen supply is
depleted. Oxygen depletion is accelerated by high water
temperatures which increase the metabolic rate of the
fish; at the same time, oxygen is less soluble in warm
water. Menhaden which die from low oxygen stress can
immediately be recognized by the red coloration on their
heads caused by bursting blood capillaries. Just before
death, the fish can be seen swimming very slowly in
a disoriented manner just below the surface of the water.
This is a common phenomenon which has been observed
throughout the range of the species. Menhaden spotter
pilots have reported menhaden "boiling up" from the
middle of dense schools, and washing up on the beach,
apparently from oxygen depletion within the school.
This phenomenon was observed during December, 1979 in
the ocean off Atlantic Beach, North Carolina. Other
scientists reported a similar event off Core Banks,
North Carolina, in December 1997. Other species are
not nearly as susceptible simply because they do not
enter enclosed inshore waters in such large numbers.
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