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Exotic
Invaders in Elkhorn Slough:
Who Are They and Where Did They Come From?
Large bays with international ports are
known to harbor diverse populations of exotic species. For example,
about 150 non-indigenous invertebrates have been documented in
San Francisco Bay, 100 in Pearl Harbor, and fifty in Puget Sound.
These invaders arrived by various means, but recently the most
significant mechanism of introduction has probably been ballast
water, which is pumped into vessels at one port and discharged
at another. As a result, the fauna of many large bays has become
homogenized, losing much of its regional character as native species
are replaced or marginalized.
While biological invasions of large bays have received much attention
lately, very little is known about the extent of this problem
in smaller embayments. Are estuaries without international shipping
safe from exotic invaders?
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1: There was no decline in the cumulative number of exotic
species collected at Elkhorn Slough over time (each sampling
period represents only about ten hours of search effort),
suggesting many more invaders remain to be found. |
In a nutshell, our study at Elkhorn Slough revealed that the
answer is a resounding "no." We have recently documented
fifty-five exotic invertebrate invaders in the Slough. Of these,
twenty-four had not previously been known to occur there. We carried
out a rather modest search for invasive invertebrates and have
calculated that we collected one new exotic species for every
1.5 hours of search effort (Figure 1). Therefore, more hours spent
searching will doubtless turn up more exotic invertebrate species.
In addition, there are likely to be many other exotic species
in other groups that we did not examine (dinoflagellates, fishes,
bacteria, etc.).
Overall, exotic species account for only about 10 percent of
the Slough's rich invertebrate fauna. However, due to the
incredible abundance of some of these species, we suspect that
they account for a far more significant proportion of the total
invertebrate
biomass. For instance, the Japanese mud snail, Batillaria attramentaria,
is found in virtually every high intertidal area of the Slough,
often in great densities (see next article). Another conspicuous
invader of the Slough is Hymeniacidon sinapium, a bright
orange sponge that forms massive aggregations on Slough mudflats
and channels (Figure 2). The filtering activities of this sponge
may be influencing planktonic communities at the Slough. European
green crabs (Carcinus maenas) also occur in the Slough.
Voracious predators, these crabs have been shown to alter clam
and worm communities at Bodega Bay dramatically. As a final example,
the burrows of a small isopod resembling a pill bug, Spheroma
quoyanum, riddle almost every bank around the Slough and likely
increase the already high erosion rates in the Slough. Exotic
invaders at the Slough therefore have the potential to influence
native communities and physical habitat structure negatively.
Some exotic species found in Elkhorn Slough were introduced with
oysters, which were cultured there from the early 1900s to the
1970s. Japanese and Atlantic oysters were imported, and other
creatures inadvertently hitched a ride with them. However, new
introductions have continued unabated in the fifty years since
oyster-culturing dwindled and ceased.
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| Figure
2: Massive aggregations of an invasive sponge, Hymeniacidon
sinapium, in the Elkhorn Sough Reserve. |
Due to the similarity in exotic species composition between the
Slough and San Francisco Bay (about 90 percent of the invasive
species found in the Slough are also found in the Bay), we suspect
that most of the recent introductions into the Slough occurred
indirectly via the Bay, rather than directly from distant waters.
Many exotic species are first introduced to this coast at major
harbors via international shipping. After becoming established
and abundant, they may then spread up and down the coast, for
instance on the hulls of regional boats or as larvae travelling
by natural currents.
The effect of international shipping, including ballast water
dumping, is thus not limited to areas with major harbors but rather
reverberates up and down the coast to smaller and more isolated
estuaries. Appropriate management to limit future invasions involves
not only minimizing direct transport vectors that bring exotic
species from distant shores to major ports -- for instance
by treating ballast water -- but also requires measures such
as frequent cleaning of small boats to reduce intraregional transport
from port areas to other smaller embayments.
Kerstin Wasson1, Chela Zabin2, Laura Bedinger3, Cristina
Diaz3, and John Pearse3
1Elkhorn Slough National Estuarine Research
Reserve
2Department of Zoology, University of Hawaii
3Institute of Marine Sciences, University of
California Santa Cruz
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A
Battle between Snails Elucidates Details of Biological Invasion
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A non-native Japanese mudsnail, Batillaria
attramentaria, was introduced to Elkhorn Slough and other
sites along the West Coast of North America fifty to seventy years
ago with imports of Japanese oysters. Batillaria has been
displacing a highly similar native mudsnail, Cerithidea californica
(sometimes called the "California horn snail"), in several
salt marshes along our coast, such as Tomales Bay and Bolinas
Lagoon. Cerithidea however no longer exists in Elkhorn
Slough; its absence there is suspected to be due to competitive
exclusion by Batillaria.
Batillaria achieves superiority over Cerithidea
through a combination of three major advantages. First, Batillaria
is a better competitor for the diatoms that each species grazes
off the surface of the marsh mud. Notably, Batillaria processes
its food more efficiently and can thus convert the shared, limited
food resource into snail tissue at a faster rate than Cerithidea.
Second, both snails are susceptible to infection by trematode
parasites, but Batillaria is typically infected at a lower
rate. Because these parasites disable a snail's reproductive
system, this lower infection in Batillaria equates to a
major advantage in the number of eggs that a Batillaria
individual can produce in its lifetime compared to a Cerithidea.
(Interestingly, one of the parasites infecting Batillaria
is itself a non-indigenous species, thus adding to the growing
list of non-indigenous species in Elkhorn Slough.) Finally, Batillaria
has a lower background mortality rate; that is, Batillaria
simply has a lower probability of death than Cerithidea of the
same age.
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| sIn
certain pockets of Elkhorn Slough Batillaria achieves its
densest populations (up to 10,000 snails/m2 a) in its entire
non-native distribution along the North American Pacific coast. |
In order to test the relative importance of Batillaria's
three advantages over Cerithidea, we defined the biology
and interactions of the snail species as connected mathematical
expressions. Output of this mathematical model of the snails'
interactions identified which advantage was most responsible for
the exclusion of Cerithidea by Batillaria, and thus
suggests means of human intervention that may more successfully
control or delay the impact of the exotic species.
The model indicated that displacement and ultimate local exclusion
of Cerithidea by Batillaria takes between fifty-five
to seventy years. Historical and museum records and analyses of
long-term population patterns corroborate this result, indicating
that the model accurately captures the biology of the species.
Importantly, the long time to Cerithidea exclusion highlights
that, over short time scales, harmful invaders may appear quite
innocuous. Additionally, the model indicated that competition
and susceptibility to parasites are relatively weak mechanisms
in driving the overall success of Batillaria. Rather, we
learned that Batillaria's lower background mortality
rate plays the key role governing its displacement of Cerithidea.
Therefore, creative management techniques can focus attention
on equalizing species-specific differences in mortality to neutralize
Batillaria's invasion most effectively.
Despite the impending local extinction of Cerithidea and
an immediate increase in the invader's population, all aspects
of Cerithidea's biology in the model were disconcertingly
slow to exhibit signs of impact from the invasion. Most of Cerithidea's
biological parameters (such as density, population biomass, egg
production, mean size, proportion parasitized, and individual
growth rate) took at least twenty to twenty-five years from Batillaria's
first introduction to show significant declines. With such a pronounced
delay between an invader's introduction and the manifestation
of its harmful effects, biological monitoring programs currently
used to prioritize intervention for imperiled native species appear
inadequate. Rather, the studies of Batillaria-Cerithidea
interactions indicate that mechanistic field experiments and mathematical
modeling of native-invader interactions may provide a better,
earlier warning of impending invader impact on native biota.
Jeb Byers
University of California Santa Barbara
University of Washington
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