Senior
Seminar 2002
Introduced Species in Hawaii
Preface Introduction Impacts Vectors Prevention/Management Action
Samoan Crab Philippine Mantis Shrimp White-Spotted Jellyfish
You will notice that this page in the web site is structured much differently than the other sites. This is because marine bioinvasions are much different than terrestrial invasions. The other pages within this site focus on a number of species that have invaded Hawaii. This page includes more general information about marine exotic species, because they are different in many ways from terrestrial introduced species. To be completely honest, I was unaware that introduced species were even a problem in marine environments. But throughout my research for this project I have become increasingly aware of the dangers to our oceans due to introduced species. If I was unaware that marine bioinvasions were a problem, I am sure that there are many others that are also in the dark. I have learned a great deal about the problems that invasive species have caused and continue to cause in Hawaii and all over the world, and along with this knowledge comes the responsibility to share this information with others. I hope that this website will be more than just part of a senior seminar that is required for graduation. I hope that this website will succeed in educating people (even if just a few!) about both marine and terrestrial introduced species in Hawaii and how their impacts on the Hawaiian environment are applicable all over the world.
The
introduction of alien species is most evident in a terrestrial environment.
This is because humans are obviously terrestrial and as a species
we tend to only notice things that directly affect us. The amount
of research and attention given to terrestrial introduced species is enormous
in comparison to that given to marine alien species. This lack of
attention and concern has allowed introduced species to become an increasing
threat to marine resources all over the world. Everyday numerous species
are transplanted to foreign waters and an unknown number of these species
become established. After habitat destruction, introduced species
are the greatest cause of the loss of biological diversity (Vitousek et
al. 1997). These introductions noticeably impact subsistence
fishing, fishing industries, human welfare, and the stability of the ecosystem
(Carlton 1989).
Hawaii is a very isolated oceanic archipelago
and greatly depends on planes and ships to bring in supplies that the
islands cannot produce themselves. The Hawaiian Islands are 1390
kilometers from the nearest island, Johnston Island, and are 3890 kilometers
away from the nearest continental landmass, North America. The Hawaiian
marine fauna is clearly of Indo-Pacific origin, most likely due to ocean
currents coming from tropical Asia (Brock 1960). Oceanic islands,
such as Hawaii, are extremely vulnerable to biological invasions because
they have been isolated from the selective forces that have shaped the
evolution of continental species (Loope et al. 1988). While this
concept concerns terrestrial exotic species, it is also applicable to
the marine environment. The waters surrounding the Hawaiian archipelago
contain various types of habitats and are teeming with a diverse array
of species. This ecosystem is very unstable because of its evolutionary
isolation and any sort of disturbance can cause irreversible damage. The
marine ecosystem is very important to the Hawaiian people, as it is responsible
for providing much of their income. The surrounding seas are culturally
significant for the native people, the ocean provides food, jobs, and
is responsible for bringing millions of tourists and their money to the
islands every year. But everyday as countless numbers of exotic
marine species are brought to the waters surrounding the Hawaiian Islands,
this important ecosystem is put in ever-increasing amount of danger.
As previously mentioned, introduced marine species impact an environment in which many people depend on for their income (Carlton 1989). Introduced species can cause alterations in species interactions, change nutrient cycling, change energy flow, and spread disease, all of which result in unpredictable and most likely damaging effects on the marine ecosystem (Carlton 2001). Many serious diseases have occurred as a direct result of transferring live aquatic animals to new locations (Humphries 1995). Introduced marine species can become so dominant that the native species become extremely rare, as in the case of the introduced Philippine mantis shrimp in Hawaii (Kinzie 1968 & 1989). Introduced marine species have been known to foul jetties, marinas, boats, and buoys. Alien species are even capable of stressing or even destroying fisheries. There is no way to determine the actual economic impact that marine introduced species have; but the amount of money lost from the destruction of fisheries, the removal of fouling organisms, and the money spent on research and education, is an enormous sum (Carlton 2001).
A
vector is the physical means for transporting a species into a foreign
environment. To become an introduced species the species must survive
transport on a vector, survive its initial introduction into a new environment,
reproduce, and become naturalized. There are many factors that affect
the success of an introduced species, including climate, food resources,
the biology of the species, and the occurrence of competition, predators,
parasites, and disease.
One frequently asked question is why all
of a sudden are marine bioinvasions becoming a problem, when marine organisms
have been moved on ship hulls for many years? There are many answers
to this question. One reason may be that introductions are not successful
every time. A particular organism can be transported many times
but not become naturalized, but it only takes one successful introduction
to establish a population. Another reason may be the increased frequency
in which we use vectors such as boats and planes (Carlton 2001).
As the Hawaiian population has grown, there has been increased demand
for products from places outside the islands. This means that the
frequency of cargo ships and planes arriving to the islands increases,
resulting in more foreign species being transported. Increased tourism
in Hawaii can also be attributed to increased frequency of these vectors.
We have also increased the distances we travel and trade; therefore
we introduce a new host of species from all over the world. As we
evolve so do the vectors in which exotic species are transported.
New vectors, such as drilling platforms, allow for a different set of
species to be introduced. As we alter the marine environment by
creating marinas, docks, jetties, beaches, we create new habitats for
introduced species to settle. As we over fish we eliminate potential
competitors and predators that could inhibit successful introductions
(Carlton 2001). Basically the short answer to the question is that
humans create new vectors, increase the frequency of vectors, and destroy
the ecosystem, making the environment more vulnerable to invasion by foreign
species.
Common Vectors of Marine Bioinvaders
Floating
Marine Debris
Photo Courtesy of M.
Posch
*Species can be transported on pieces of garbage floating on the ocean surface (Carlton 2001).
Recreational
Equipment
Photo Courtesy of
Buckeye
Marine Photo
Courtesy of Chris
Mac
Photo Courtesy of Divers
Supply
*Small recreational vehicles such as speedboats, yachts, and jet skis have become very popular and are sometimes transported large distances. Many marine-fouling organisms can be transported on these vehicles. It is possible that snorkeling and SCUBA gear could also serve as a vector (Carlton 2001).
Seaplanes
Photo
Courtesy of
Martin
Hadley
*Fouling organisms
can attach to the planes rudders and be easily transported over long distances
from one location to the next. Organisms are also transported via
the pontoon water (Carlton 2001).
Canals
Photo of the Atlantic entrance to the Panama Canal Courtesy
of MACR.com
*It is possible for
organisms to be moved through sea level, lock, or irrigation canals. Although
canals probably contribute to the introduction of freshwater organisms
more than marine organisms, there is speculation that some introductions
do occur through the Panama Canal by water movement and by ship (Carlton
2001).
Public and Private
Aquaria
This is a typical marine aquarium with live rock, fish, and invertebrates
Courtesy of Atlantis
Aquatics
*Public aquaria can accidentally or intentionally release display organisms. The water that these organisms are transported in can also be a source of introductions if not disposed of properly. The aquarium industry is also a source of marine introductions. Invertebrates, fish, live rock, and seaweeds are shipped all over the world. The ultimate fate of these organisms lies in the hands of their buyers, and often times these organisms are intentionally or accidentally released (Carlton 2001). During the summer of 2000, Caulerpa taxifolia of the Mediterranean was released by a home aquarist in southern California and it quickly became established, taking over many of the native species (Anderson and Keppner 2001 in Carlton 2001). Species belonging genus, Caulerpa are commonly found in home aquariums, this macroalgae can spread very rapidly and dominate reefs and it would be a real threat to the reefs if released in Hawaii.
Research and Restoration
Photo courtesy of Amanda
Robinson
Arbicia punctulata, used in
developemental
research
*Organisms used for
research or organisms associated with the research organisms can be intentionally
or accidentally introduced. In an effort to restore locally devastated
populations of native species, people bring in these native species from
other areas. Sometimes other organisms associated with these species
are accidentally transported as well (Carlton 2001).
Drilling Platforms
and Dry Docks
Courtesy of the
Center for Southeast Asia Studies
Off-shore drilling
platform
*Worldwide trade and exploration has become increasingly important, therefore the need to move drilling platforms, dry docks, and navigation buoys has also grown. Dry docks are large, floating structures used to repair ships and drilling platforms are used for resource detection and extraction. Both of these structures have large surfaces for fouling organisms to attach and they also have ballast water systems. These structures are often moved over long distances allowing for species to be introduced into foreign waters (Carlton 2001).
Fisheries and
Mariculture
Both photos courtesy of Douglas
Jager
Commercial fishing boat Commercial
fishing net
*The fisheries and
mariculture industries are responsible for introducing many marine species.
These introductions occur both intentionally and accidentally. Many
times organisms are held in containers in marine environments for aquaculture
purposes. It is sometimes possible for these organisms to escape
and any other organisms that are associated with the farmed species may
be released also. This is the case for many shellfish, such as clams,
crabs, oysters lobsters, etc. Another way in which introductions
happen is when species are released by the government in hopes of initiating
a new fishery or enhancing an active fishery. This type of introduction
often occurs in the form of an illegal release by private citizens in
hopes of jump-starting an industry (Carlton 2001). Hawaii is very
guilty of this practice. The Hawaiian islands have relatively few
native forms of shellfish capable of supporting a strong shellfish industry.
Private citizens of Hawaii have recognized this insufficiency, and
have attempted to introduce shellfish since at least the late 1800's.
Around 1920 the Hawaiian government also began to release exotic shellfish
in hopes of adding to the economy (Edmondson and Wilson 1940). The
government and private citizens of Hawaii were successful in many cases,
such as in Scylla serrata (you can find out
more information about this crab further down the page). Live seafood
intended for consumption, is sometimes released out into the wild.
The waste materials associated with live, fresh, or frozen seafood often
contains other organisms (sometimes encysted) that can be introduced if
not disposed of properly. Live bait can also be released back into
the wild by fishermen/women. Organisms may also be introduced via
the movement or drifting of fishing gear like nets, traps, trawls, etc.
Organisms intentionally or accidentally transported in "live well"
(a compartment where bait or caught fishes are kept alive) water can also
be responsible for marine introductions (Carlton 2001). There are
many other ways in which the fisheries and mariculture industries can
introduce species, but these are the most common vectors.
Ships
All photos courtesy
of SERC
Marine Invasion Reasearch Laboratory and Watershed Radio
*Diagram showing how ballast-mediated *Researchers
sampling a ship's *A
ship exchanging coastal ballast water
species introductions occur ballast
water with
open ocean water to reduce the
risk
of introducing exotic species
*Ships are the primary
vectors that transport marine organisms to foreign locations. Hundreds
of species are known to be transported both on and in ships. There
are two ways in which marine organisms can be introduced by a ship, through
fouling or ballast water. Ballast water is pumped into a ship to
balance for a lack of cargo, and most ships carry some ballast water even
when the ship is full of cargo. A ship can take in or discharge
water in port or at sea and the water is held in ballast tanks. Most
ships will take coastal water into their ballast and then discharge the
water upon reaching its destination. Because coastal environments
are relatively similar, there is a greater chance of an introduced organism
surviving in the coastal environment than in the open ocean. Open
ocean ballast water exchange is diffucult to do in rough seas and it is
also more time consuming, therefore many ships prefer to exchange ballast
water in port. Until around the late 1800's "dry ballasts"
were used, which involved heavy objects such as rock, sediment, or sand.
The "dry ballasts" were also a vector for the introduction
of many organisms.
*Most of the time ballast water is pumped into a ship while it is near
the coast. This coastal water is usually packed with many organisms.
The water can contain drifting organisms known as plankton, free-swimming
organisms known as nekton, fouling organisms that can attach to the walls
of ballast tanks, and benthic organisms known as benthos. Many of
the organisms transported through ballast water are in their larval stages
and become benthic as adults. Sea anemones, barnacles, crustaceans,
mollusks, polychaetes, urchins, seaweeds, and many other organisms are
introduced in their larval stages. There are other organisms such
as copepods, jellyfish, dinoflagellates, and other organisms that are
transported in their adult stages (Carlton 2001). Organisms that
become introduced species are often overlooked because they are inconspicuous
and difficult to identify (Brock 1960). There have been many studies
done on the role of ballast water in species introductions. One
such study that sampled ballast water at the beginning of the voyage right
after the water was taken in and then sampled the ballast water at the
end of the voyage before the water was exchanged. This study suggested
that the transit stage serves as a significant selective force for ballast
water organisms. There was change in both the number and the taxonomic
diversity of organisms living in the water from the beginning to the end
of the transit. The study suggests that certain taxonomic groups
are more likely to survive transit, therefore they have a better probability
of becoming an introduced species (Smithe et al. 2000).
A study done by Wonham et al. (2000) has shown that fish in their
adult stage have also been transported in ballast water and the role of
ballast water in fish introductions has been greatly underestimated.
*The National Invasive Species Act of 1996 directed the U.S. Coast Guard
and the Smithsonian Environmental Research Center to create the National
Ballast Water Information Clearinghouse. The Clearinghouse is responsible
for monitoring patterns of ballast water exchange in U.S. commercial shipping
ports. The Clearinghouse began the National Ballast Survey (NABS)
to examine these patterns. The NABS suggests that all ships entering
U.S. ports from outside the Exclusive Economic Zone undertake a voluntary
open-ocean exchange, in which they exchange as much of their water as
possible in the open ocean. The NABS then requires that all ships
entering U.S. ports report their ballast water activities. If after
three years the NABS feels that the voluntary program is not successful
the Secretary of Transportation will make this program mandatory (Miller
et al. 2000). This program began in July 1999, but after
the first 12 months only 12,170 out of 58,000 vessels fulfilled the mandatory
reporting requirement. The results of the NABS have not yet been
reported (Carlton 2001). One big problem with this voluntary exchange
program, besides the fact that it is voluntary, is that it does not regulate
intracoastal ballast water exchange. It has been suggested by Lavoie
et al. (1999) that intracoastal transfer of ballast water on
ships can be an important vector for marine introductions. They
suggest that future management plans concerning ballast water exchange
include domestic ships.
*Hawaii marine ecosystems have fallen victim to introductions by ballast
water transport. In a study done in Pearl Harbor 22% of the species are
considered to be introduced (Coles et al. 1997). Many of
these species were introduced by ballast water transfer that occurs in
the harbor. Most of the marine introduced species in Hawaii were
introduced by ballast water or fouling. This is very evident when
reading A Guidebook of Introduced Marine Species in Hawaii (Defelice et
al. 2001).
*The most important
part of introduced species management is prevention. Once an introduction
occurs it is almost impossible to eliminate the organism and eradication
efforts are far more costly than the costs of preventing the introduction.
There are very few laws that concern the introduction of species
into foreign environments and laws currently in place are usually not
known by the public and not strictly enforced. The U.S. National
Invasive Species Act of 1996 (previously discussed) is a step in the right
direction for the United States and hopefully other countries will follow
suit (Carlton 2001).
*Once an introduction occurs there are no laws in place to specify how
eradication efforts should proceed. There are many options for post-introduction
management, but there are negative side effects to many of these methods.
Many times eradication efforts lead to further environmental damage.
The mechanical removal of organisms is the most common method of post-introduction
control. The organisms can be removed by hand or by machine. It
is sometime possible to turn the removal of an invasive species into an
industry, like using the removed organisms for fertilizer or selling the
organisms at the market as food products (Carlton 2001). The release
of chemicals is also used to reduce invasive populations. This method
can cause a problem if the chemicals kill organisms not intended as a
target.
Photo
courtesy of Diane
Drigot
Civilian and military
volunteers remove alien red mangrove (Rhizophora mangle) and pickleweed
(Batis maritima), major invasive plants in the Hawaiian wetlands.
*There are other methods of post-invasion control that have not been used yet, but are strong possibilities. Genetic engineering of introduced species could be used to alter the tolerances, reproduction, or other important processes (Carlton 2001). Biological control is also a method that has been strongly considered. Biological control species can be parasites, parasitoids, pathogens, predators, or competitors. A study done on biological control of marine pests concluded that biological control is a strong possibility in marine environments, but due to spatial and safety concerns much more study must be done before such introductions happen (Lafferty and Kuris 1996).
*It is very obvious
that something needs to be done to keep our oceans from being devastated
by introduced species. The difficulty with making laws to protect
our oceans is that the ocean is continuous. What happens in Canada
can affect waters in the United States and visa versa. There is
a great need for all countries to cooperate in making and enforcing laws
that concern exotic marine species.
*In the United States there are surprising few laws concerning the introduction
of species and there is a great need for more strict laws. I think
that it should be mandatory for all ships to do open ocean ballast water
exchange, unless there is extremely bad weather. I also think that
there should be more money given to marine bioinvasions research by the
government. We need more research so that we can deal with these
invasions more effectively. And last but not least, there needs
to be more public education about introduced species. Many people
are unaware that their actions are putting the oceans at risk and if they
are educated they will be less likely to be involved in marine introductions.
*Check out links below to find out more about what
Hawaii is doing about marine introduced species.
Scylla serrata
Phylum Arthropoda
Subphylum Crustacea
Class Malacostraca
Order Decapoda
Infraorder Brachyura
Family Portunidae
Common names: Mangrove Crab, Mud Crab, Samoan Crab
Photo Courtesy of J.
Poupin
Description
This crab is the largest and most aggressive
portuid crab in Hawaii, about 18cm in width. The carapace is broad,
smooth, and long. On the front of the carapace are four blunt teeth
of equal size and the anterolateral border has nine sharp teeth. The
chelipeds of the males are very large and powerful. These crabs
inhabit muddy bottoms in brackish water, in mangrove areas, and river
mouths (Edmondson 1954, DeFelice et al. 2001). These crabs
are gonochoristic and physiological maturity in males is reached when
the carapace reaches between 90-110mm. The male and female begin
the mating process when a female is in premolting condition. The
crabs remain paired for 3 to 4 days until the female molts, and then they
copulate (Knuckey 1996). The female then migrates offshore with
the fertilized eggs, where they hatch in a couple of weeks (Hill 1996).
Geographic
Range
The native range of the mangrove crab is in the
Indo-Pacific. From South Africa to Tahiti, north to Okinawa, Japan,
and south to Port Hacking, Australia and the Bay of Islands, New Zealand.
The crab now inhabits most of the Indo-Pacific including Japan,
China, Philippines, Australia, Indonesia, East and South Africa, and the
Red Sea. In Hawaii the mangrove crab can be found in the waters
surrounding all of the islands (DeFelice et al. 2001). Scylla
Serrata is not considered invasive in any of these areas, due to
fishing pressures on the crab. If for some reason Scylla Serrata
was not exploited as a food resource in Hawaii or any other area
of it's non-native range, the crab would most likely become invasive because
there would be no control on the population.
Vectors
The Samoan crab was first introduced into Kaneohe Bay
in 1926 to start a commercial crab fishery. A total of 98 crabs were released
on Oahu, Hawaii, and Molokai between 1926 and 1935 (Brock 1960). A paper
by Edmondson and Wilson written in 1940 reported that the mangrove crab
was already an established species. A study done in 1981 in Kahana Estuary,
Oahu, Hawaii found species of crabs in the estuary, and Scylla serrata
was the most abundant even though they were highly fished (Maciolek and
Timbol 1981).
Why
is it successful
This species is a large and very aggressive carnivorous species and it
most likely feeds on native invertebrates. Brock (1960) believes
that this crab may be successful because the estuaries in which these
crabs were released have low rates of tidal flushing, which may facilitate
rapid population group in the estuaries.
Impacts
This species has been intoduced in Hawaii and through
the Indo-Pacific to start commercial crab fisheries. This crab is
an important source of income for many people. The ecological impacts
of this crab have not been studied in Hawaii, but this crab most likely
feeds on native invertebrates.
Management
There is nothing being done to eradicate the mangrove
crab from Hawaii. The population of mangrove crabs stays in control
due to the commercial fishery surrounding this species. I think
that as long as this crab continues to be a part of the fisheries industry
there is not much that needs to be done to control this species. While
this crab is not considered to be an invasive species in Hawaii or in
any of it's non-native areas, this species does have the potential to
become a pest if its population is not kept under control by the crab
fishing industry. If the time comes that this species is no longer
fished, there will have to be another way to control the populations to
prevent the population from becoming invasive.
Gonodactylaceus falcatus
Phylum
Arthropoda
Subphylum Crustacea
Class Malacostraca
Order Stomatopoda
Family Gonodactylidae
Common name: Philippine Mantis Shrimp
photo courtesy
of J.Hoover
Description
This species grows to about 6cm. The females
are reddish brown and the males are dark green. The sixth abdominal
segment has six inflated lobes. The telson also has these lobes,
along with three pairs of marginal teeth and one pair of accessory teeth.
These mantis shrimp inhabit dead branching coral heads, coralline
algae clumps, and holes in the substrate or rock. These shrimp are
carnivorous and use their raptorial claws to grab live prey. They
have seperate sexes and the female carries the fertilized eggs until hatching.
This species reproduces twice a year.
Geographic
Range
The native range of the Philippine mantis shrimp
is of the Indo-Pacific. It can now be found in shallow reef surrounding
Oahu. This species is not invasive in Oahu and it has not been found
anywhere else outside of it's native range.
Vectors
The mechanisms by which this shrimp was introduced was
most likely through ship hull fouling (Carlton 2001). Kinzie (1968)
suggested that it was introduced in Oahu waters by the concrete barges
towed back from South China and the Philippines after World War II, because
this species was not found in Hawaii until after the war.
Why
is it successful
This is a very aggressive species and has been observed
to physically drive out a similar native species (Pseudosquilla ciliata)
from their crevices. It is able to outcompete the native species
and therefore fill their niche
(Kinzie 1968, 1984).
Impacts
This species has been able to displace the native mantis
shrimp, Pseudosquilla ciliata, from Kaneohe
Bay, Oahu (Kinzie 1968, 1984). Other impacts on the environment
have not been studied, but these impacts are probably minimal since the
individuals of this introduced species fill an already existing niche.
While this species is not considered to be invasive and it's impact appears
to be minimal, I still think that it is important to include. It
is important because researchers have no clue how many native species
have been replaced by non-native species and many times introductions
go unnoticed. It is also important to realize that we may not predict
there to be any adverse effects caused by this species, but that doesn't
mean that there won't be.
Management
Nothing has been done about this species and there is
probably very little that could be done except for to minimize the spread
of this species to other islands.
Phyllorhiza
punctata
Phylum Cnidaria
Class Scyphozoa
Order Rhizostomeae
Family Magistiidae
common name: White-spotted Jellyfish
Photo courtesy of
Dock Watch
Description
The maximum size for this jellyfish is 50cm in
diameter. It is usually bluish-brown in color and has many opaque
white spots. The white-spotted jellyfish has eight oral arms which
terminate with large brown bundles of stinging cells, called nematocysts.
The nematocysts are used for protection and capturing plankton. The
adult jellyfish swim near the surface in estuaries, harbors, and bay areas,
but the habitat of the larval stages is unknown. The white-spotted
jellyfish have basiic cnidarian reproduction (see diagram below). These
jellyfish have been observed to be more abundant in the winter months
in Hawaii.
Jelly fish life cycle courtesy of ptpleasant
Geographic
Range
This jellyfish is native to Australia. It's
present distribution includes Australia, the Hawaiian Islands, the Caribbean,
and the Gulf of Mexico. So far in Hawaii this jelly fish has only
been observed in the waters surrounding Oahu, specifically Pearl Harbor,
Honolulu Harbor, Ala Wai Canal, Yacht Harbor, and Kaneohe Bay. This jellyfish
is invasive in the Gulf of Mexico and while it has not become invasive
in Hawaiian waters yet, there is a strong possibility that is soon could
become invasive.
Vectors
This jellyfish was most likely introduced into Hawaii
as a scyphistomae by ship fouling or as a medusa, ephyrae, or planktonic
planula in ballast water.
Why
is it successful
This jellyfish is most likely successful because of its'
reproductive cycle. I t can be transported as a medusa, ephyrae,
planula, or scyphistomae via ballast water or fouling, therefore it has
a higher probability of being introduced. If the jellyfish is transported
in the planula or scyphistomae stage it has the ability to break off into
many young medusa ephyrae. This means that if introduced the population
can grow very rapidly. If transported in the ephyra or medusa stage
there must be another individual of the opposite sex available for fertilization
to take place. These jellyfish have very few natural predators in
Hawaii and there is plenty of planton for them to feed on, therefore they
are able to be very successful.
Impacts
The ecological impact of these jellyfish has not been
studied in Hawaii, however these organisms are known to eat planktonic
crustaceans, fish eggs, and fish larvae in other locations. There
was a population explosion of the white-spotted jellyfish in the Gulf
of Mexico, where it threatened commercially important organisms such as
fish, shrimp, and crabs. This sort of population fluctuation has
not been observed in Hawaiian water, but it is possible that it could
occur.
Management
Nothing has been done to control this species in Hawaii
and there is little that can be done once it has been introduced, because
of its' alternation of generations life cycle (Carlton 2001). Probably
the best management strategy that Hawaii can have concerning this introduced
species is to try to prevent the spread of this species to other islands.
Anderson, L. W. and S. Keppner. 2001. Caulerpa taxifolia: marine algal invader provokes quick response in U.S. waters. ANS Digest 4: 21-23.
Brock, V. E. 1960. The introduction of aquatic animals into Hawaiian water. Internationale Revue der gesamten Hydrobiologie 45: 463-480.
Carlton, J. T. 1989. Man's role in changing the face of the ocean: biological invasions and implications fro conservation of near-shore environments. Conservation Biology 3: 265-273.
Carlton, J. T. 2001. Introduced species in U.S. coastal waters: environmental impacts and management priorities. Pew Oceans Commission, Arlington, Virginia.
Coles, S. L., R. C. DeFelice, L. G. Eldredge, J. T. Carlton, R. L. Pyle, and A. Suzumoto. 1997. Biodiversity of marine communities in Pearl Harbor, Oahu, Hawaii with observations on introduced exotic species. Bishop Museum Technical Report 10: 1-3.
DeFelice, R. C., L. G. Eldredge, and J. T. Carlton. 2001. Nonindigenous invertebrates. Eldredge, L. G. & C. Smith, coordinators, Guidebook to the Introduced Marine Species in Hawaiian Waters. Bishop Museum Technical Report 21: 1- 60.
Edmondson, C. H. 1954. Hawaiian Portunidae. Occasional Papers of Bernice P. Bishop Museum 21: 217-274.
Edmondson, C. H. and I. H. Wilson. 1940. The shellfish resources of Hawaii. Proceedings of the Sixth Pacific Science Congress, University of California Press, Berkeley. 241-243.
Hill, B. J. 1996. Offshore spawning by the portunid crab Scylla serrata (Crustacea: Decapoda). Marine Biology 120: 379-384.
Humphries, J. D. 1995. Introductions of aquatic animals to the Pacific Islands: disease threats and guidelines for quarantine. Perspectives in Aquatic Exotic Species Management In the Pacific Islands 2: 1-47.
Kinzie, R. A. 1968. The ecology of the replacement of Pseudosquilla ciliata by Gonodactylus falcatus (Crustacea: Stomatopoda) recently introduced into the Hawaiian Islands. Pacific Science 22: 465-475.
Kinzie, R. A. 1984.
Aloha also means goodbye: a cryptogenic stomatopod in Hawaii. Pacific
Science 38: 298-311.
Knuckey, I. A. 1996. Maturity in male mud crabs, Scylla serrata, and the use of mating scars as a functional indicator. Journal of Crustacean Biology 16: 487-495.
Lafferty, K. D. and A. M. Kuris. 1996. Biological control of marine pests. Ecology 77: 1989-2000.
Lavoie, D. M., L. D. Smitch, and G. M. Ruiz. 1999. The potential for intracoastal transfer of non-indigenous species in the ballast water of ships. Estuarine, Coastal and Shelf Science 48: 551-564.
Loope, L. L, O. Hamann, and C. P. Stone. 1988. Comparative Conservation Biology of Oceanic Archipelagoes. BioScience 38: 272-282.
Maciolek, J. A. and A. S. Timbol. 1981. Environmental features and macrofauna of Kahana Estuary, Oaho, Hawaii. Bulletin of Marine Science 31: 712-722.
Miller, A. W., G. M. Ruiz, L. Takata, B. Steves, and A. H. Hines. 2000. Measuring ballast water delivery and management patterns in the United States: The national ballast water information clearinghouse and national ballast water survey. In Marine Bioinvasions: Proceeding of the First National Conference, J. Pederson, ed. Massachusetts Institute of Technology, MIT Sea Grant College Program, MIT SG 00-2, Cambridge, Massachusetts.
Smith, L. D., D. M. Lavoie, G. M. Ruiz, and B. S. Galil. 2000. Changes in ballast water biota during intracoastal and transoceanic voyages. In Marine Bioinvasions: Proceeding of the First National Conference, J. Pederson, ed. Massachusetts Institute of Technology, MIT Sea Grant College Program, MIT SG 00-02, Cambridge, Massachusetts.
Vitousek, P. M., H. A. Mooney, J. Lubchenco, and J. M. Melillo. 1997. Human domination of Earth's ecosystems. Science 277: 494-499.
Wonham, M. J., J. T. Carlton, G. M. Ruiz, and L. D. Smith. 2000. Fish and ships: relating dispersal frequency to success in biological invasions. Marine Biology 136: 1111-1121.
National Marine Invasions Center - this site provide information about the National Ballast Water Survey, marine invasive species, provides good links, and has a lot of other good information.
Introduced Marine Species of Hawaii - you can download A Guidebook of Introduced Marine Species in Hawaii, it is a very helpful resource.
USGS - this site provides information about aquatic introduced species, it is easy to use and allows you to view information about the specific taxa that you choose.
Hawaii Coral Reef Initiative Plan - find out what is being done to protect Hawaii's coral reefs from destruction by introduced species.
Center For Research On Introduced Marine Pests - provides some of the most most current research that the center is working on, as well as some other useful information and links.
Designed by: Jenny Jamison Contact: jamisje@earlham.edu
Last Revised: 12/9/02