Senior
Seminar 2002
Introduced Species in Hawaii
Pig Goat Sheep Deer Rodents Mongoose Cat Wallaby
Pronghorn Humans Literature Cited
Introduction
In Hawaii there is only
two native species of mammals, the Hawaiian Hoary Bat (Lasirus cinereus
semotus),
and the Hawaiian Monk Seal (Monachus schauinslandi). The
lack of native mammals leaves Hawaiian ecosystems very vunerable, native
species have not evolved defenses to the mammalian predators and herbivores
that have been introduced in the last 300-1000 years. Therefore making
native species very vulnerable to attack. Add on the warm tropical climate
in Hawaii, lack of competitiors and predators, and this archipelago provides
an ideal habitat for nearly all introduced mammals to become established.
In the following page 15 species will be profiled, examining their impacts
and management that must be conducted to save this archipelago from losing
it's native flora and fauna.
Sus
scrofa
Common Name: Feral Pig (below)
Picture courtesy of: usgs.gov/s+t/SNT/noframe/pi179f19.htm
Geographic
Range
Original: Europe (Atkinson and Atkinson 2000)
Hawaiian: Pigs are found on all but two of the Islands (Kahoolawe and
Lanai where they were formerly present), the island of Hawaii supports
the largest and densest population of the entire archipelago. (Department
of Land and Natural Resources 2002)
Vectors
The Polynesian race of pigs
was brought with the first human inhabitants around 400 A.D. (Spatz and
Mueller-Dombois 1975).
The European race was brought to the islands
by Captain James Cook in 1778 (Tomich, 1986).
Why
is it successful
As mentioned in the introduction
the lack of native mammals leaves the Hawaiian ecosystems very vulnerable.
Plants have not evolved defenses against native browsing animals, especially
one as voracious as the pig. Entire forests and have been destroyed and
leveled by this species. Also lack of native predators allows this species
to flourish with no population regulators. Loope et al. (1988) suggests
that pig invasions are governed by favorable combinations of climate,
genetics, population dynamics and forest use by people. The Polynesian
stock of the feral pig are thought not to have been a significant role
in destruction of the Hawaiian ecosystems. There are two factors being
hypothesized to explain the feralization of pigs and the destruction of
habitats. Loope et al. (1988) suggests that a lack of proteinaceous foods
were available in early native forests, not allowing the pigs to exist
outside human settlements. The introduction of the european race of pig
(common farm pig found in America and Europe) could have also played a
major role, as this larger much more aggressive stock could have been
much harder to handle then the smaller and more docile Polynesian type
(typical pot-bellied pig). This newly introduced stock completely replaced
the older stock. Both factors were likely to have influences on feralization
of the as introduction of earthworms, and fleshy-fruited plants by Europeans
would provide the necessary proteins for a sustainable feral population
to exist.
Impacts
Feral Pigs are a huge problem
nearly everywhere they are introduced, Texas, Tennessee, Florida, Australia,
and the Galapagos Islands in Ecuador. They are found to impact nearly
every native plant community in the Hawaiian islands (Aplet et al.
1991). They dig deep holes in the ground searching for soil invertebrates
and the starchy centers of native plants. The
wallows they dig fill with water allowing mosquitoes carrying avian malaria
to breed (Loope et al. 1988). They
have been found to selectively seek out native species such as ferns,
tree-ferns, lobeliads, and mints for food (Loope et al. 1988). With destruction
of native vegetation comes the destruction of native bird and insect populations
as well. The pig facilitates the invasion of other introduced species.
Behaviors of uprooting vegetation and digging wallows leaves areas open
allowing early successional invasive plants to become established (Spatz
and Mueller-Dombois 1975, Aplet et al. 1991). Pigs eat and carry
invasive seeds which are deposited through excrement or phoresy (Atkinson
and Atkinson 2000). Such
imapcts may not all come from the european race of pigs though as Kirch
(1982) cites a number of endemic terrestrial mollusks that were found
to be driven to extinction during these prehistoric times. Although it
is hypothesized that destruction of forests by man had led to the extinction
of these species (Kirch 1982), it may be possible that feral populations
of Polynesian pigs may have existed and ate native mollusks as a source
of protein.
Management
Control measures that are
being taken against the pig are few but very effective. Hunting is probably
the number one control method. Unfortunately, the state of Hawaii has
set hunting seasons for pigs as well as bag limits. In order to completely
eradicate the pig from the archipelago it would seem advantageous to have
a year-round season with no bag limits. Other,
methods such as snaring have proven to be quite effective as well (Anderson
and Stone 1993). Although there is opposition against the eradication
of Feral Pigs in Hawaii, Native polynesian groups argue that by erasing
the pig from the islands they are erasing a integral part their culture,
and certain hunting outfits also oppose this idea, as there will be nothing
left to hunt if the pigs are removed (Atkinson and Atkinson 2000). Passive
measures are also being taken to keep pigs out of national parks and other
nature reserves. Fencing has been a popular and effective strategy as
of late (Loope et al. 1988) (see pictures below). Fencing has
played a major role of the recovery of previously pig-damaged areas, however
fencing only keeps the pigs out, it does not control the populations.
Even if the pigs are removed other measures must be utilized to remove
the species that were brought in by the pig.
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Left: Native bog without pigs. Middle:
Three years after pigs found the bog. Right : Seven years after the bog
was fenced.
Pictures courtesy
of: www.rarehawaii.org/pigpage/pigs.htm
Geographic
Range Vectors Why
is it successful Impacts Ovis
aries Geographic
Range Vectors Why
is it successful Impacts Management Axis
axis Geographic
Range Vectors Why
is it successful Impacts Management Mus
musculus Geographic
Range Vectors Why
is it successful Impacts Management Herpestes
auropunctatus Geographic
Range Vectors Why
is it successful Impacts Management Felis
catus Geographic
Range Vectors Why
is it successful Impacts Management Petrogale
penicillata Geographic
Range Vectors Why
is it successful Impacts Management Antilocapra
americana Geographic
Range Vectors Why
is it successful Management Homo
sapiens Although
you may not technically be able to call humans an invasive or introduced
species, because we migrated to the islands on rafts of floating vegetation,
much like many of the ancestors to todays modern endemic species that
are found on the island, it must be realized that we have caused more
trouble on these islands than all of the above species combined. Humans
are responsible for bringing all of the species on this entire web site
to the Hawaiian Islands. Although at the time some of the reasons for
these introductions may have seem justified, we now realize that they
have only hurt the islands more. Many native species have felt direct
impact from humans as soon as we arrived on these islands. Kirch (1982)
and Atkinson (1977) cite that prehistoric man hunted native birds for
their feathers, and there is no evidence that this hunting was decreased
as bird populations began to diminish. It seems that every time fossil
deposits are discovered containing early human populations we find a whole
range of other species that are no longer with us today (Atkinson and
Atkinson 2000, Kirch 1982). Man has affected these islands in many other
ways as well, burning and logging of native forestsand the widespread
use of substances that pollute the air, ground and water. However one
advantage of Man is that we can stop what we are doing and we can try
to change things. We can take back what we introduced into this ecosystem,
and we can prevent more species from entering. This should be the most
important strategy that man has, prevention and post-introduction control
and management. Literature
cited: Designed by: Ryan Welsh Contact:
welshry@yahoo.com
Common Name:
Feral Goat (below)
Picture
courtesy of:
www.sls.lcs.mit.edu/~klivescu/photos/hawaii/
Original: Asia
Hawaiian: Goats are found on all but two of the islands (Lanai and Nihau
where they were formerly present) (Atkinson and Atkinson 2000).
They
were brought to Hawaii by Captain James Cook in 1778 (Tomich 1986).
Goats were introduced to nearly all islands in the Pacific
Ocean by European sailors, they were brought to islands as a food reserve
for shipwrecked sailors (Atkinson and Atkinson 2000).
Goats are generalist, they
will feed on nearly any type of vegetation and are able to survive in
nearly all habitats. In Hawaii goats are present from low to high altitudes,
where they are able to exists and affect both wet and dry habitats (Stone
1984 in Atkinson and Atkinson 2000).
Goats
are extremely destructive herbivores that will eat nearly any type of
vegetation available. They will eat young trees and young shoots of plants,
before any establishment can be made. On the Galapagos Islands, they have
been witnessed to completely change a landscape from a low wooded area,
to a grassland in less than ten years (Loope et al. 1988). Large
population of goats on hawaii are able to decimate forests and eat native
vegetation. It is also possible that disturbances caused by goats allow
early successional invasive plant species to become established, as they
may be carried through excrement and phoresy.
Management
Management
strategies of controlling goats are very similar to those of the pig expand
hunting seasons and increase bag limits. However hunting alone will not
affect the populations. From 1920 to 1970 approximately 70,000 feral goats
were removed from Hawaii Volcanoes National Park, with no noticeable effect
(Loope et al. 1988). Atkinson
and Atkinson (2000) report that New Zealand uses radio-collared goats
to locate herds and after a herd is located, hunters are dispatched to
the area. Fencing has become
a popular strategy as well, as Mueller-Dombois and Spatz (1975) report
that two years after a goat exclosure fence was put up, a previously undescribed
leguminous plant species (Canavalia kauensis) appeared.
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Common Name: Feral Sheep
Ovis ammon
Common Name: Mouflon Sheep (below)
Picture courtesy of: www.ansi.okstate.edu/breeds/sheep/mouflon/
Feral Sheep:
Original: Europe
Hawaiian: Hawaii (formerly existing on Kahoolawe) (Atkinson and Atkinson
2000)
Mouflon Sheep:
Original: Europe and Asia
Hawaiian: Hawaii and Lanai (Atkinson and Atkinson 2000)
Feral Sheep were brought
by Captain James Cook in 1778 (Loope 1999).
The exact date when Mouflon was brought
to the islands is unknown, but they were most likely brought over for
food and sport at a later date then the Feral Sheep.
Both species are herbivores
that feed on a varied types of vegetation.
Very similar situation to
the goat, both species of sheep are extensive and destructive herbivores.
They have been found to decrease populations of the Mamane (Sophora
chrysophylla) a endemic leguminous tree, by stripping the bark off
thus facilitating damage from insects and and other disease causing organisms
(Scowcraft and Sakai 1983). These species have also been found to be a
significant factor in the decline of the Mauna Kea Silversword (Argyroxiphium
sandwichense), which due to browsing pressures the wild population
has been limited to 42 plants all confined on cliffs and rock faces that
are inaccessible to the sheep (Robichaux et al. 2000).
Management strategies are
very similar to the those used with the pig and the goat. Expand hunting
seasons, increase bag limits and use fencing to keep the animals out.
There is a big push to keep sheep in some of the reserves, especially
the Mouflon as it is considered to be a very popular game animal.
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Common Name: Axis Deer (below right)
Odecoileus hemionus
Common Name: Mule Deer (below left)
Pictures courtesy of www.botany.hawaii.edu/bot350/
& www.ranchbrokers.com
Axis Deer:
Original: Indian subcontinent
Hawaiian: Maui, Lanai, Molokai, and Oahu (Atkinson and Atkinson 2000)
Mule Deer:
Original: North America
Hawaiian: Kauai (Atkinson and Atkinson 2000)
Axis Deer
were first introduced to the islands in 1867, and subsequent introductions
continued on the islands until 1959 (Waring 1996). They were introduced
by order of the state legislature, to promote as a game species (Rarehawaii
2002).
Mule Deer were brought
to Kauai in 1961 from Oregon by the State of Hawaii Fish and Game Division,
these were brought to be promoted as a game species as well (Rarehawaii
2002).
Axis Deer: Warm
Hawaiian climate allows for aseasonal breeding (Graf and Nichols 1966),
and extreme palatability of native grasses allows for greater foraging
thus increase in fawn output (Waring 1996).
Mule Deer: Varied diet
allows the deer to feed on a number of different plants. Feeding largely
on introduced species such as strawberry guava, thimbleberry, passion
flower, and blackberry. It feeds on some native fruits such as maile,
a'ali'i, 'uki'uki, pilo, and koa as well (Rarehawaii
2002).
Axis
Deer are primarily a grazing species, and prefers to eat newly sprouting
grasses (Elliot and Barrett 1985). This can destroy populations of many
native grasses before they are able to be established.
Since Mule Deer are known to eat a variety
of introduced species, it is very likely that they serve as a vector to
carry the seeds of these species allowing them to spread.
Currently there are no known
management strategies for control of either species of deer, although
there are hunting seasons available for both species. Culling of both
species and fencing could be useful strategies as well. Waring (1996)
suggests that fencing would be an ideal control measure for Axis Deer
as they do not readily jump obstacles.
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Common Name: House
Mouse
Rattus rattus
Common Name: Black Rat
Rattus norvegicus
Common Name: Norway Rat
Rattus exulans
Common Name: Polynesian Rat (below)
Pictures courtesy of: biology.usgs.gov/pierc/
House Mouse:
Original: Africa or Asia
Hawaiian: All islands (Atkinson and Atkinson 2000)
Black Rat:
Original: India
Hawaiian: All islands (Atkinson and Atkinson 2000)
Norway Rat:
Original: Mongolia
Hawaiian: All islands except Kahoolawe (Atkinson and Atkinson 2000)
Polynesian Rat:
Original: Indo-Malaysian region
Hawaiian: All islands (Atkinson and Atkinson 2000)
All four species of rodents
were brought to the islands as stowaways on ships (Atkinson and Atkinson
2000). Atkinson (1977) has gathered evidence to suggest that the Black
Rat arrived on the islands sometime in the 1870's and also dates the Norway
Rat at first arrival around 1816 and the House Mouse 1825. The Polynesian
Rat was the only species presumed to have been brought to the islands
by the Polynesians in the 5th century (Kirch 1982).
All four species are generalists,
that will feed on anything ranging from seeds and berries to vertebrate
flesh (Wirtz 1972). Many species of insects and birds have not evolved
defense mechanisms against predators therefore making them very vulnerable
to these introduced species.
The affects of these rodents
on native species are vast. Being generalists they are able to feed on
a variety of food types. Polynesian Rats are known to prey upon the nestlings
and eggs of the Laysan Albatross (Diomedea immutabilis), Black-footed
Albatross (Diomedea nigripes), Wedge-tailed Shearwater (Puffinus
pacificus), Bonin Island Petrel (Pterodroma hypoleuca),
Red-tailed Tropicbird (Phaethon rubricauda), Masked Booby (Sula
dactylatra), Brown Booby (Sula leucogaster), Red-footed
Booby (Sula sula), Sooty Tern (Sterna fuscata), and
Common Noddy (Anous stolidus) (Wirtz 1972). Atkinson (1977) has
attributed the Black Rat as the main factor for the decline of many of
the native land bird species as well. Though the Norway Rat and the House
Mouse are not as agile climbers they are thought to feed extensively on
native insects (Lindsey et al. 1999) and the seeds to native
plants decreasing those poulations as well (Atkinson and Atkinson 2000).
Poisoning or baiting appears
to be the only strategy that is currently being used to control these
species. Research is currently being conducted to find a bait that will
affect all four species (Atkinson and Atkinson 2000, Dunlevy et al.
2000). Effective compounds currently being used are anticoagulant
poisons such as brodifacoum and bromodialone (Atkinson and Atkinson 2000).
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Common
Name: Small Indian Mongoose (below)
Pictures courtesy of: www.hawaii-forest.com
Original: Asia
Hawaiian: Hawaii, Maui, Molokai and Oahu
The mongoose was brought
to the islands in 1883 as a bio-control agent for rats in sugarcane fields
(Baldwin et al. 1952).
The mongoose is an agile
predator with a wide range of prey and a very high reproductive rate (breeds
2-3 times a year producing litters of 3 young) as well as a young age
of first reproduction (10 weeks) (Baldwin et al. 1952).
Although mongoose were introduced
to the islands to prey on rats they have preyed upon many other native
species as well. They are known to be the number one predator against
the endangered Nene (Nesochen
sandvicensis) eating both eggs and incubating females (Stone
et al. 1983). They will eat the eggs and adults of nearly any
forest species of bird to enter the understory. They have been known to
eat the young of the endangered Hawaiian Crow (Corvus hawaiiensis),
and they have eliminated populations of the Hawaiian Dark-rumped Petrel
(Pterodroma phaeopygia) on all islnds the mongoose is present
(Baldwin et al. 1952). Mongoose have also been known
to eat and carry the seeds of invasive plants such as the Strawberry Guava
(Baldwin et al. 1952).
Currently there appears
to be no attempts to eradicate the mongoose from any of the Hawaiian islands.
Coblentz and Coblentz (1985) have not found an effective strategy in eradicating
the mongoose. It is mentioned however that they may be controlled by using
anticoagulant poisons, but it should be tested further to measure its
impact on other species (Stone et al. 1983). Other control measure
include shooting and trapping, especially in breeding areas of rare birds
(Atkinson and Atkinson 2000).
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Common Name: Feral Cat (below)
Pictures courtesy of: biology.usgs.gov/pierc
Original: Africa
Hawaiian: All islands (Atkinson and Atkinson 2000)
Cats were most likely introduced
to the islands in the late 18th century by European sailors, who often
carried them on ships as a control for rodent infestation (Atkinson and
Atkinson 2000).
With an abundance of easily
catchable prey, and the ability to adapt nearly any habitat Hawaii has
cats have become very successful thoughout Hawaii.
After the Black Rat, the
cat is thought to be a major reason for the decline of many endemic forest
birds (Atkinson 1977). Although cats have been found to prefer small mammals
as prey, they will switch to native birds as well as a host of invertebrate
species many of which are endemic (Smucker et al. 2000). For
instance, in 56 cat scats collected the remains of 44 birds were found,
40 of them were endemic species. The remains of the endemic Hawaiian Hoary
Bat were found as well (Smucker et al. 2000).
Management strategies for
cats include live-trapping, shooting, poisoning, snaring, gin traps and
the introduction of the viral disease feline panleucopaenia (Bloomer and
Bester 1992, Innes et al. 1999). Not surprisingly there is a
civilian effort to end to killing of cats on the islands, as it is seen
as cruelty to animals. There are attempts to create cat colonies, where
they put food out, and spay and neuter each cat as it visits the area.
For homeowners however the best control of feral cats is to keep them
inside and have them spayed or neutered. Therefore effectively stopping
them from harming native species and breeding with other feral or demestic
cats.
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Common Name: Brush-tailed Rock Wallaby (below)
Pictures courtesy of:
www.royle.abel.co.uk/slwap/animals/rockwall.htm
Original: Australia
Hawaiian: Oahu (Lazell 1981)
Three wallabies (A male,
female and juvenile) were brought to the island in 1916 to be part of
a private zoo, the breding pair was known to have escaped from the zoo
when a dog attacked the tent in which they were being held (Tomich 1986).
Since the population of
these wallabies hover at around 100 individuals, they can not be considered
to be that successful (Lazell 1981). However they are known to live in
only one valley (Ewa Kalihi), which has a series of sheer cliffs and narrow
rocky ledges (Lazell et al. 1984). This probably protects this
species from most introduced predators as there are only a few accounts
of predations by dogs (Lazell et al. 1984).
Currently there is no suggested
impact that this species has on the native hawaiian ecosystem. However
they have been found to eat some introduced species of plants such as
the Christmasberry, which they may be spread through excretion (Lazell
et al. 1984).
As of right now there are
no management strategies for the wallabies and they are actually protected
by the Hawaiian government (Lazell 1981). The population on Hawaii was
thought to be a population of wallaby that had gone extinct on the Australian
mainland and it was not until very recently that this notion was dismissed
through molecular genetic analysis (Eldridge and Browning 2002).
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Common Name: Pronghorn (below)
Pictures courtesy of: www.amazilia.net
Original: North America
Hawaiian: Lanai
Pronghorn was brought to
the island in 1959 by the state Department of Agriculture and Conservation,
to be maintained as a hunting species.
Fortunately for the islands
of Hawaii, this introduction was not very successful. Fifty-six animals
were set to be introduced in 1959, their process of introduction is as
follows: (Taken from Environment
Hawaii 1999)
"When they arrived in Honolulu on
December 9, 1959, there were 44 antelope remaining, the rest having died
en route of injury or pneumonia due to cold and rainy weather on the way.
These were then sprayed with Malathion as a preventative to the introduction
of ectoparasites into the islands, and then trucked to Hickam Air Force
Base, where they remained overnight. Four antelope were given to the Honolulu
Zoo; the remaining 40 were flown to Lana`i for release. Two more died
between Honolulu and Lana`i. The 38 animals reaching Lana`i were released
from their crates into a small holding pen which was located on a large,
grassy flat on the northeast side of the plateau at about 1,700 feet elevation.
When they were all out of the crates and grouped in a single herd, the
gate was opened and they were eased into the open and freedom.
There
are no natural water holes or streams on Lana`i, so artificial water units
were established for the antelope. However, one factor was overlooked
in the release, and was to cause serious trouble almost immediately: the
antelope had never seen salt water, and were used to large fresh water
lakes in which to quench their thirst. They took one look at the large,
blue "lake" several miles down slope from the release area,
and headed directly for the Pacific Ocean! Unfortunately, they bypassed
the water units placed in the release area. Sometime during the first
night following their release, they found their way through the heavy
algaroba forest lining the coast and came out on the narrow sand beach
between the forest and the sea.
The next day, they were found wandering
disconsolately up and down the narrow beach, searching vainly for drinking
water, and unable to return to the cooler, open country above because
of the solid forest which they refused to enter. A crew of volunteers
was immediately rounded up and the antelope, now suffering from lack of
water, were herded up the beach to an open ridge that led to the higher
rangeland and water units. During the drive, some of the animals became
confused and took to the water, swimming out over the reef towards the
open sea; however, the surf turned them, and they returned to shore with
no losses. A few became lost in the trees and did not make it up the ridge
with the main herd. Most of these were subsequently chased -- or captured
and carried -- up to the open range of the release area, but at least
two died on the beach, probably from the effects of drinking salt water.
The majority regained the release area and
found the water units. They remained there for a number of days before
commencing to wander. By January 20th, the known number of survivors had
been reduced to 18. A number had died from the effects of having their
eyeballs punctured by the thorns on the algaroba trees while they were
at the beach, and others had wandered from the main herd and could not
be found. Most of the 18 were suffering from scours, probably brought
on by the severe change in diet, and it is possible that some losses were
caused by this."
Impacts
The effects of the pronghorn
are very similar to the problems that deer, sheep and goats have caused
through the overgrazing on native species and spread of invasive species
through excrement and phorsey.
Nothing is currently being
done to manage the pronghorn, Tomich (1986) says that the last reports
of pronghorn were in the mid 1980's and that the species has probably
died off. However, Atkinson and Atkinson (2000) still list the species
as being present on the island, although no further information is provided.
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Common Name: Human
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