Senior
Seminar 2002
Introduced Species in Hawaii
Photos courtesy of:
http://www.cursorius.com/haw_introduced.html
Japanese White-eye Kalij Pheasant House Finch
Introduction
In the Hawaiian Islands, more species of birds have been
introduced than anywhere else on earth (Moulton and Pimm 1986, Shehata
et al. 2001). At least 142 species of birds, from six continents,
have been introduced since 1850 (Moulton et al. 2001). Around
54 of these introduced species have established breeding populations on
the islands (Loope et al. 2001).
Vectors
Most of the introduced birds in Hawaii were introduced
intentionally during the late 1800s and early 1900s. The majority of these
species were from the orders Galliforme, Columbiforme, and Passeriforme
(Moulton et al. 2001). Intentional introductions were common
until the 1950’s when they were restricted by the government (Pyle
1995). No additional species from the above three orders have been introduced
since 1981, however multiple species of parrots have been sighted since
1990 (Moulton 2001).
A society called the Hui Manu was formed strictly for the purpose of introducing birds to Hawaii (Moulton and Pimm 1983). The Hui Manu may have been motivated to bring new species because they wanted to replace the native species that were declining (van Riper 2000). State and local government organizations and private citizens also made efforts to establish populations of exotic birds (Moulton et al. 2001). Unlike accidental introductions, most intentional introductions involved large numbers of individuals, and often, efforts were made to enhance survival after their release. Galliformes is an example of a group of species that were intentionally released and are still being maintained. Apparently, the motivation for introducing birds to Hawaii was the perceived need to enrich the native avifauna of Hawaii by adding exotic birds. Galliformes and most Columbiformes were probably introduced for recreational hunting, while Passeriformes were introduced for reasons such as aesthetics and biological control of other introduced species (Moulton et al. 2001). The cattle egret (Bubulcus ibis), barn owl (Tyto alba), and Guam swiftlet (Aerodramus bartschi) are examples of birds that were introduced by governmental agencies to biologically control other invasive species (Loope et al. 2001).
Why
are some introduced birds successful?
Human activities created habitats that are favorable for
many of the species that were intentionally introduced. Most introductions
occurred after habitats below 600 m had been drastically changed through
deforestation, grazing by introduced mammals, and agriculture (Moulton
and Pimm 1983).
Human effort is responsible for the success of many species of Galliformes. The success of many Galliformes released at the Puu Waawaa Ranch, which was established for the purpose of introducing game birds, is probably at least partially attributable to the efforts made by the ranch owners. By the mid 1960s, 43 fenced sanctuaries had been constructed and were supplied with food and water (Lewin 1971). In 1971, the ranch was providing 212,715 lbs of bird food, costing around $10,600 US, to the sanctuaries annually (Lewin 1971). In these areas, predator control was practiced intensively (Lewin 1971).
Many of the species that have been introduced to Hawaii have adapted to live in unnatural habitats created by humans such as parks, houses, and zoos. Since most native birds are not found in highly disturbed areas, introduced birds did not experience much competition with native species (Moulton and Pimm 1983). In some urban areas, only introduced species of birds are found. For example, in Waikiki, five species of introduced birds made up 94.0% of all of the birds observed during one study (Williams and Fleischer 1989). Although many introduced species are found associated with human settlements, there are also species such as the Japanese white-eye that have colonized both disturbed and undisturbed habitats.
The introduction success of birds in Hawaii has been studied extensively to attempt to determine the role of various factors in predicting establishment success. Abiotic conditions, behavioral flexibility, and migratory behavior have been found to impact establishment success. The number of individuals introduced also has been found to relate to the likelihood of establishment success, with higher introduction numbers being more likely to become established (Veltman et al. 1996).
Blackburn and Duncan (2001) studied the determinants of establishment success using data from all of the known bird introductions in the world. Their study provided evidence that establishment success is determined by the suitability of the abiotic conditions at the introduction site more than by species richness or climate. They did not find significant differences between highland and lowland or island and mainland locations, but instead found that birds are more successful at invading sites that are similar to their origin and are within their biogeographic range. This is probably because such sites share climatic and habitat features more frequently.
Species with large native geographic ranges are more likely to be successfully introduced to new areas because there is a greater chance that they will encounter habitats and resources they are capable of utilizing, and they are usually more tolerant of variances in abiotic conditions such as climate (Blackburn and Duncan 2001). Moulton et al. (2001) found that birds that were successfully introduced to Hawaii had a larger native range size on average than unsuccessful species. Generalists are probably more successful than specialists for similar reasons.
Behavioral flexibility can also be used to predict establishment success. Sol and Lefebvre (2000) used brain size and foraging innovations in the area of origin as measures of behavioral flexibility and found that birds with higher behavioral flexibility were more successful at invading novel environments. This trend is logical because behavioral flexibility allows birds to exploit new food sources and habitats without a long evolutionary process.
Impacts
Competition with Native Avifauna
Much of the endemic Hawaiian avifauna is endangered or threatened. Competition
with introduced birds is possible cause of decline in endemic bird populations
(van Riper et al. 1986). Interspecific competition in native
forests was found to be greater between native and introduced species
than among native species or among established introduced species in areas
where introduced and native species are found together (Mountainspring
and Scott 1985). Loss of habitat is also a major cause of the declining
native populations because many native species have close habitat associations
and are specialists (Moulton and Pimm 1986).
Dispersion
of Non-native Plants
Introduced birds play a destructive role in Hawaii by distributing invasive
plants. The germination of some seeds may be enhanced rather than harmed
by being digested by a bird. Introduced species of birds disperse both
native and introduced plants. According to Cuddihy and Stone (1990), the
spotted dove it thought to be responsible for the rapid spread of invasive
Lantana camara, and the Japanese white-eye, house finch, and other introduced
birds are blamed for dispersing an invasive plant called Myrica faya.
Three species of parrots, the rose-ringed parakeet (Psittacula krameri),
red-crowned amazon (Amazona viridigenalis) and mitred conure (Aratinga
mitrata) have established breeding populations on some islands in Hawaii,
and they are a threat because they can potentially damage native plants,
spread invasive seeds, and are agricultural pests (Loope et
al.
2001). Introduced parrots have been very destructive in other areas where
they have been introduced (Loope et
al.
2001).
Reservoir
for Avian Parasites and Diseases
Introduced birds function as vectors and reservoirs for avian diseases
that are spread to native species of birds. Avian diseases, especially
avian malaria and avian pox, are one of the greatest factors leading to
the decline of native Hawaiian birds (van Riper and van Riper 1986, Banko
et
al.
2001, van Riper and Scott 2001). According to Jacobi and Atkinson (1995),
a major avian malaria and avian pox epidemic in midelevation forest birds
was documented during a National Biological Service study in 1992. Avian
malaria is probably the most important factor preventing endemic bird
populations from recovering in low elevations (Jarvi et
al.
2001). Many introduced birds develop nonfatal infections that can be transmitted
to other birds by introduced mosquitoes.
Mosquito-borne diseases are a relatively new problem for Hawaiian birds. Culex quinquefasciatus, the mosquito that serves as a vector of avian malaria and avian pox, was not present in Hawaii until it was unwittingly introduced into Maui in 1826 when sailors dumped water containing larvae from Mexico into a stream while refilling their water dregs (Warner 1968). Before this event, mosquito transmitted diseases were not a threat to the Hawaiian avifauna even though migrating birds may have harbored malaria infections. The Culex mosquito has spread to all of the islands, but it is mainly in low to midelevation areas because this subspecies is typically found below around 600 m above sea level (Warner 1968). Mosquitoes can inhabit areas as high as 1800 m above sea level in some years when conditions are favorable (VanderWerf 2001).
Many native Hawaiian birds are extremely susceptible to malaria and pox because they did not evolve in an environment with strong selective pressure for resistance to these diseases. The iiwi is so susceptible to malaria that in a study where juveniles were infected with a dose equivalent to the bite from a single infected mosquito, 90% died (Jacobi and Atkinson 1995). Introduced birds are found in all forests, but most native birds have been found to be restricted to forests with few or no mosquitoes (Shehata et al. 2001).
Some native species have developed immunogenic and behavioral responses that decrease their vulnerability to diseases (van Riper and van Riper 1986). Shehata et al. (2001) found that a population of the endemic amakihi (Hemignathus virens) living in a lowland forest may have evolved resistance to malaria. In this study, the amakihi individuals were not infected with malaria, but several introduced birds in the same area harbored infections. The amakihi may have been able to evolve a resistance more quickly than many other native birds because they reach sexual maturity in only six months (Shehata et al. 2001).
Even though some native birds have evolved resistance to avian malaria, they are still succeptible by avian pox. Avian pox causes lesions on the legs, feet, faces and internal membranes of birds, and can result in the loss of toes, blindness, emaciation and death (VanderWerf 2001). The pox virus can be spread through arthropod bites or by direct contact between a contaminated surface and broken skin or mucous membrane (VanderWerf 2001). This disease was widespread among endemic Hawaiian birds in the late 1800s (van Riper and Scott 2001). Pox was probably responsible for some of the extinctions that have occurred since then, and it continues to affect birds in Hawaii today (van Riper and Scott 2001). VanderWerf (2001) found that in areas where the prevalence of pox in elepaio was high, the density of elepaio and other bird species was low, which is evidence that pox is limiting the success of native birds in these areas and increasing the risk of local extinction. In this study, the site with the highest prevalence of pox and also malaria was at a lower elevation (1,550 m) where more mosquitoes were found, and the at the highest elevation sites (1,800-1,900 m), pox lesions and mosquitoes were very rare.
Avian pox is not always fatal. VanderWerf (2001) found that elepaio individuals that survived the initial infection of pox did not experience decreased future survival and reproductive success, and they probably became at least somewhat immune to subsequent infections. However, the mortality rate of elepaio with active lesions may be as high as 40% on Oahu (VanderWerf 2001), so the disease is still very detrimental to this native species. New diseases such as encephalitis, which is also transmitted by the Culex mosquito, are a huge threat to native Hawaiian avifauna (Warner 1968), and precautions should be taken to prevent additional diseases from entering the islands.
Other Impacts
Several introduced birds have detrimental impacts on specific native birds.
The barn owl (Tyto alba) preys upon the Newell’s shearwater (Puffinus
auricularis newelli) and other native birds (Ainley et
al.
2001). The mallard duck (Anas platyrhynchos) hybridizes with the Hawaiian
duck (Anas wyvilliana), an endemic species, which contributes to its decline
(Rhymer 2001). Insectivorous introduced birds impact Hawaiian plants by
preying upon insects that are pollinators for native plants (Cuddihy and
Stone 1990).
Management
Introduced birds are not targeted by current efforts to
manage invasive species (Ikuma et al. 2002). Other types of introduced
organisms have the potential to cause more damage than birds and have
been given higher priority. It would be very difficult to exterminate
or even control introduced birds. Chemical control might be a possibility
in lower elevation areas where there are few or no native birds, but developing
a safe program would be expensive and the potential hazards might exceed
the benefits. Mechanical removal is also unfeasible because birds are
difficult to locate and capture because they are highly mobile. Mobility
allows birds to escape from areas while they are being targeted and then
to rapidly repopulate them. Biological control also is not promising because
it is unlikely that another organism would target only invasive birds.
Some measures can be taken to avoid promoting introduced birds. Bird feeders should not be used because only introduced birds in residential areas are likely to benefit from them. Native forests should be protected and restored because the majority of the introduced birds are most successful in disturbed habitats.
Zosterops
japonicus
Common Name: Japanese White-eye
Photo courtesy
of: http://www.stanford.edu/~petelat1/whiteeye.html
Description
The Japanese white-eye is the most abundant bird
in Hawaii (van Riper 2000). It is found in a wide range of habitats at
both high and low elevations (Mountainspring and Scott 1985). It is most
common along forest edges at low elevations (van Riper 2000). Sakai (1988)
found that the Japanese white-eye was the first bird to become reestablished
in an area of Hawaii where the native rainforest was cleared, which demonstrated
this species' ability to colonize new habitats quickly. The principal
food of this species is nectar, but it also feeds on fleshy fruits and
arthropods (Mountainspring and Scott 1985).
Geographic
Range
The Japanese white-eye is native to Japan, eastern
and southern China, Taiwan, Vietnam, Hainan Island, Ryukyu Island, Volcano
Island, and the northern Philippine Islands (van Riper 2000).
Vectors
This species was introduced in 1929 (Mountainspring and
Scott 1985). It was intentionally released onto Oahu from Japan by the
Territorial Board of Agriculture and Forestry (van Riper 2000). This species
was introduced to the island of Hawaii in 1937 in an attempt to eradicate
insects (van Riper 2000). It currently inhabits all of the Hawaiian Islands
(van Riper 2000).
Why
is it successful?
The Japanese white-eye is the most successful introduced
bird in Hawaii. The white-eye has a broad native range. In Japan, this
species is common in many areas, from evergreen and deciduous forests
to urban areas, and is found at various elevations (Yamashina 1961). The
broad native range and generalist characteristics of the white-eye are
at least partially responsible for its success in Hawaii because these
factors allow it to utilize a wider variety of resources. The white-eye
has demonstrated behavioral flexability, which is another factor that
has been shown to increace introduction success, by utilizing a wide variety
of food sources in various habitats in Hawaii (van
Riper 2000).
Impacts
The Japanese
white-eye competes with endemic species more than most introduced birds
do because, unlike most of the introduced birds in Hawaii, it has become
established in native montane forests (Mountainspring and Scott 1985).The
presence of endemic species such as the iiwi (Vestiaria coccinea),
apapane (Himatione sanguinea) and elepaio (Chasiempis sandwichensis)
was found to be negatively correlated with the presence of the Japanese
white-eye even when habitat suitability was accounted for (Mountainspring
and Scott 1985). The negative correlation for these species was confined
to certain geographic areas for some species: wet forests for the elepaio,
and high elevation forests for the iiwi (Mountainspring and Scott 1985).
The omnivorous feeding behavior of the Japanese white-eye enables it to compete with several native specialists in different forests, and it is particularly successful because, unlike native specialists, it can switch food sources when local supplies are depleted. The elepaio specializes on foliage insects at lower heights in the foliage column, and the Japanese white-eye also exploits insects in this microhabitat (Mountainspring and Scott 1985). The iiwi specializes on nectar resources (Jacobi and Atkinson 1995), which are also exploited by the Japanese white-eye. Mountainspring and Scott (1985) suggest that since the average Japanese white-eye density is 3-16 times higher than the elepaio density and 2-5 times higher than the iiwi, the Japanese white-eye is probably depressing these native populations. In this study, the negative correlation for the amakihi and the Japanese white-eye was weak, but competition was still suspected in the areas where the average amakihi density exceeded the average white-eye density. Like the Japanese white-eye, the amakihi feeds on insects, flowers, and fruit rather than just on one of these types of foods (Mountainspring and Scott 1986), so the amakihi may not be as heavily impacted by the white-eye because it can also utilize other food resources. This study also demonstrated that the Japanese white-eye is probably partially responsible for the decline of the Hawaii creeper, an endangered endemic species.
Invasive plants such as Miconia calvescens, Lantana camara, and Myrica faya are distributed by the Japanese white-eye (Medeiros and Loope 1997, van Riper 2000). Because of its ability to invade and distribute invasive plants into undisturbed forests, the Japanese white-eye has the potential to be a particularly damaging invasive plant vector.
Lophura
leucomelanos
Common name: Kalij Pheasant
Photo courtesy
of: http://www.ejphoto.com/kalij_pheasant_page.htm
Description
The kalij pheasant is one of the most successful introduced Galliformes.
Around 78 species of game birds have been introduced to Hawaii (Cuddihy
and Stone 1990). The first bird introduced to Hawaii by Polynesians was
the galliform, Gallus gallus, which is more commonly known as
the chicken or red jungle fowl (Moulton et al. 2001). Unlike
most of the introduced Galliformes, the pheasant has been able to expand
its range and does not rely upon constant care by humans.
Geographic
Range
The kalij pheasant is native to the forests of
Pakistan, India, Nepal, Sikkim, Bhutan, Burma, and Thailand, and typically
found from 600-3400 m above sea level (Lewin and Lewin 1984).
Vectors
A total of 67 individuals of this species from Michigan
and Texas game farms were intentionally introduced to the Puu Waawaa Ranch
on the island of Hawaii in 1962 (Lewin 1971, Lewin and Lewin 1984, Loope
et al. 2001). Humans have been particularly interested in introducing
and promoting Galliformes such as the kalij pheasant in Hawaii for recreational
hunting. By 1977, the kalij pheasant was abundant enough to be declared
a legal game species (Lewin and Lewin 1984). It dispersed throughout many
parts of the island of Hawaii within 14 years without human assistance,
and humans transported several individuals to seemingly suitable forests
that had not been invaded (Lewin and Lewin 1984). Attempts to introduce
the kalij pheasant have also been made in Tennessee, Virginia, Oregon,
Washington, and British Columbia, but all of these efforts failed to establish
breeding populations (Lewin and Lewin 1984).
Why
is it successful?
This pheasant is able to exploit a wide variety of food
resources including insects, earthworms, bird eggs, slugs, snails, seeds,
leaves, flower buds, and rotting fruits (Lewin and Lewin 1984). Recently
disturbed areas are favorable for the kalij pheasant because they are
able to forage on the disrupted invertebrates and feed on starches from
cut trees (Lewin and Lewin 1984). Without the introduced species that
this pheasant feeds upon, it probably would not be as successful (Lewin
and Lewin 1984). The
kalij pheasant has not been found to utilize sanctuaries, so their success
is probably more related to the suitability of biotic and abiotic factors
in the disturbed habitats of Hawaii than to human efforts (Lewin 1971).
The island of Hawaii is probably more similar in climate to the native
range of the kalij pheasant than the locations where introduction efforts
failed in North America.
Impacts
The kalij pheasant is a threat to the native flora and
fauna of Hawaii because it has been found to consume and disperse the
seeds of invasive plants. The main food of the kalij pheasant is the invasive
banana poka (Passiflora mollissima: Cuddihy and Stone 1990). In 82% of
the kalij pheasant samples collected during one study, banana polka seeds
were found (Lewin and Lewin 1984). Banana poka is considered to be one
of the most threatening invasive plants (Lewin and Lewin 1984). According
to Lewin and Lewin (1984), other introduced plants dispersed by this pheasant
include thimbleberry (Rubus rosaefolius).
Some Galliformes, including the kalij pheasant, ring-necked pheasant (Phasianus calchinus) and chukar (Alectaris chukar), are known to disperse the seeds of native shrubs (Cuddihy and Stone 1990). The kalij pheasant has been found to disperse the Uncinea uncinata, an endemic sedge (Lewin and Lewin 1984). On Maui, introduced Galliformes are thought to play an important role in dispersing native species of plants where native birds are extinct or endangered (Cole et al. 1995). They may also have a negative effect on endangered endemic birds such as the nene (Branta sandvicensis) through competition since their niches overlap (Cole et al. 1995).
The kalij pheasant also is a threat because it feeds on rare endemic land snails and may serve as a disease reservoir (Lewin and Lewin 1984). Galliformes have been found to carry nematode and cestode parasites (Lewin 1971). Further study is necessary to understand the extent of this threat to native birds.
Carpodacus
mexicanus
Common Name: House Finch
Photo courtesy
of: http://www.enature.com/fieldguide/showSpeciesRECNUM.asp?recnum=BD0050
Description
The house finch is established on all of the
main Hawaiian islands (Moulton and Pimm 1983). The diet of this species
consists mainly of seeds and other vegetable matter (Hill 1993), but it
also consumes insects (Gough). Papaya fruit and the seeds of ironwood
(Dasuarina equisetefolia), Formosan koa (Acacia confuse), pink tecoma
(Tabebuia pentaphyla), broad-leafed plantain (Plantago major), and beggar
tick (Bidens pilosa) make up most of its diet in Hawaii (Hill 1993). In
North America, it is most abundant in settled areas with buildings, lawns,
and some trees (Hill 1993). The habitat preference in Hawaii is similar,
but it is also found in fragmented native forests (Hill 1993).
Geographic
Range
The house finch is native to western North America.
As human activities altered the landscape, the range of the western population
expanded its range eastward (Hill 1993). This species was also introduced
to New York, where it became established and has expanded westward, reaching
the Mississippi River by 1980 (Kammermeier 1999).
Vectors
The house finch
was introduced to Hawaii before 1870 (Englund and Preston 2000), and was
established on all of the islands by 1901 (Hill 1993). The individuals
introduced probably originated in the San Francisco Bay area, CA (Hill
1993).
Why
is it successful?
The house finch is probably successful because it has
adapted to coexisting with humans in altered habitats, and this type of
habitat is available in Hawaii. It has been found to benefit from human
alterations because they have provided sources of water, food, and cover
(Hill 1993). The large native range of this species, and the similarities
between disturbed habitats in Hawaii and in North America also make this
species well suited for establishment in Hawaii.
Impacts
The house finch serves as a reservoir for avian pox and malaria because
it is susceptible to these diseases but is somewhat resistant to their
debilitating effects. This resistance is evident because birds have been
found with mild infections (Warner 1986). This partial immunity allows
the disease to be carried and spread without killing the host individual.
If this species did not have some degree of immunity, they would not be
a reservoir because many infected individuals would die and therefore
cease to be carriers. In North America, the house sparrow is vulnerable
to mycoplasmal conjunctivitis, a bacterial eye disease (Kammermeier 1999).
The house sparrow is the main species affected by this disease in North
America, but it is very possible that native species of Hawaiian birds
may also be vulnerable. If mycoplasmal conjunctivitis reaches Hawaii,
the house finch may harbor and spead this disease.
The house finch is a problem in Hawaii because it causes extensive damage to grain crops. In one study on a farm in Oahu, this species consumed or destroyed 30-50% of the sorghum crop (Hill 1993). This species also competes with native species and serves as vectors for invasive plants.
Links
(see Literature Cited for additional references)
Avian
Pox: Chapter 19 from Field Manual of Wildlife Diseases: Birds.
This document provided detailed information about avian pox in the USA.
http://www.nwhc.usgs.gov/pub_metadata/field_manual/chapter_19.pdf
Hawaii’s
Forest Birds: An Inventory of Research Conducted, 1992-1998.
This site summarizes avian disease research in Hawaii and provides contact
information for many of the researchers.
http://www.hawaii.edu/scb/lib_researchFB7pr3.htm
Hawaii’s
Forest Birds Sing the Blues
This site describes the threats native forest birds in Hawaii are facing,
which include problems caused by introduced species.
http://www.fonz.org/zoogoer/zg1995/honeycreepers.htm
Our
Living Resources: Hawaii
This is an excellent document that describes some of the research that
has been undertaken by United States governmental organizations about
birds in Hawaii and in the rest of the country.
http://biology.usgs.gov/s+t/pdf/Hawaii.pdf
Seasonal
Prevalence and Transmission of Avian Pox and Malaria in Hawaiian Forest
Birds
This site provides useful information and pictures of birds with pox and
malaria.
http://biology.usgs.gov/pierc/PMPoxMalaria.htm
Literature
Cited
Ainley,
D. G., R. Podolsky, L. Deforest, G. Spencer and N. Nur. 2001. The status
and population trends of the Newell’s Shearwater on Kauai: insights
from modeling. Studies in Avian Biology 22: 108-123.
Banko, P. C., R. E. David, J. D. Jacobi and W. E. Banko. 2001. Conservation status and recovery strategies for endemic Hawaiian birds. Studies in Avian Biology 22: 359-376.
Blackburn, T. M. and R. P. Duncan. 2001. Determinants of establishment success in introduced birds. Nature 414: 195-196.
Cole, R. F., L. L. Loope, A. C. Medeiros, J. A. Raikes, and C. S. Wood. 1993. Conservation implications of introduced game birds in high-elevation Hawaiian shrubland. Conservation Biology 9: 306-313.
Cuddihy, Linda W. and Charles P. Stone. 1990. Alteration of Native Hawaiian Vegetation: Effects of Humans, Their Activities, and Introductions. Honolulu: University of Hawaii Cooperative National Park Resources Studies Unit.
Englund, R. A. and D. J. Preston. 16 March 2000. Avian species assessment for the Kuhio Highway, Manoa stream ford crossing project. Bishop Museum. Hawaii Biological Survey: Contribution No. 2000-005. URL http://hbs.bishopmuseum.org/pdf/hbs2000-005.pdf (11 November 2002)
Gough, G. House finch Carpodacus mexicanus. United States Geological Survey. URL http://www.mbr-pwrc.usgs.gov/id/framlst/i5190id.html (24 November 2002)
Hill, G. E. 1993. House Finch (Carpodacus mexicanus). Report No. 46 in A. Poole and F. Gill (editors). The Birds of North America. The Academy of Natural Sciences, Philadelphia, PA, and American Ornithologists’ Union, Washington, DC.
Ikuma, E. K., D. Sugano and J. K. Mardfin. 2002. Filling the gaps in the fight against invasive species. Honolulu, HI: Legislative Reference Bureau Report No. 1, 2002. URL http://www.state.hi.us/lrb/rpts02/gaps.pdf (22 November 2002)
Jacobi, J. D. and C. T. Atkinson. 1995. Hawaii’s Endemic Birds. Pages 376-381 in LaRoe, E. T., G. S. Farris, C. E. Puckett, P. D. Doran, and M. J. Mac, eds. 1995. Our living resources: a report to the nation on the distribution, abundance, and health of U.S. plants, animals, and ecosystems. U.S. Department of the Interior, National Biological Service, Washington, DC. 530pp URL http://biology.usgs.gov/s%2Bt/pdf/Hawaii.pdf (11 November 2002)
Jarvi, S. I., C. T. Atkinson and R. C. Fleisher. 2001. Immunogenetics and resistance to avian malaria in Hawaiian honeycreepers (Drepanidinae). Studies in Avian Biology 22: 254-263.
Kammermeier, L. 1999.
Population Dynamics of the House Finch. Birdscope, Volume 13, Number 2:
15.
URL http://birds.cornell.edu/publications/birdscope/Spring1999/hofipop99132.html
(15 November 2002)
Lewin, V. 1971. Exotic game birds of the Puu Waawaa Ranch, Hawaii. Journal of Wildlife Management 35: 141-154.
Lewin, V. and G. Lewin. 1984. The Kalij Pheasant, a newly established game bird on the island of Hawaii. Wilson Bulletion 96: 634-646.
Loope, L. L., F. G. Howarth, F. Kraus and T. K. Pratt. 2001. Newly emergent and future threats of alien species to pacific birds and ecosystems. Studies in Avian Biology 22: 291-304.
Medeiros, A. C. and L. L. Loope. 1997. Status, ecology, and management of the invasive plant, Miconia calvescens DC (Melastomaceae) in the Hawaiian Islands. Bishop Museum Occasional Papers 48: 23-36. URL http://hbs.bishopmuseum.org/pdf/melastome97.pdf (22 November 2002)
Moulton, M. P., K. E. Miller and E. A. Tillman. 2001. Patterns of success among introduced birds in the Hawaiian Islands. Studies in Avian Biology 22: 31-46.
Moulton, M. P. and S. L. Pimm. 1983. The introduced Hawaiian avifauma: biogeographic evidence for competition. American Naturalist 121: 669-690.
Moulton, M. P. and S. L. Pimm. 1986. The extent of competition in shaping an introduced avifauna. Pages 80-97 in J. Diamond and T. J. Case, editors. Community Ecology. New York: Harper and Row Publishers.
Moulton, M. P., J.
G. Sanderson and R. F. Labisky. 2001. Patterns of success in game bird
(Aves: Galliformes) introductions to the Hawaiian islands and
New Zealand. Evolutionary Ecology Research 3: 507-519.
Mountainspring, S. and J. M. Scott. 1985. Interspecific competition among Hawaiian forest birds. Ecological Monographs 55: 219-239.
Pyle, R. L. 1995. Birds of Hawaii. Pages 372-375 in LaRoe, E. T., G. S. Farris, C. E. Puckett, P. D. Doran, and M. J. Mac, eds. 1995. Our living resources: a report to the nation on the distribution, abundance, and health of U.S. plants, animals, and ecosystems. U.S. Department of the Interior, National Biological Service, Washington, DC. 530pp URL http://biology.usgs.gov/s+t/pdf/Hawaii.pdf (11 November 2002)
Rhymer, J. M. 2001. Evolutionary relationships and conservation of the Hawaiian Anatids. Studies in Avian Biology 22: 61-67.
Sakai, H. F. 1988. Avian response to mechanical clearing of a native rainforest in Hawaii. Condor 90: 339-348.
Shehata, C., L. Freed and R. L. Cann. 2001. Changes in native and introduced bird populations on Oahu: infectious disease and species replacement. Studies in Avian Biology 22: 264-273.
Sol, D. and L. Lefebvre. 2000. Behavioral flexibility predicts invasion success in birds introduced to New Zealand. OIKOS 90: 599-605.
VanderWerf, E. A. 2001. Distribution and potential impacts of avian pox lesions in Elepaio at Hakalau Forest National Wildlife Refuge. Studies in Avian Biology 22: 247-253.
van Riper, C. and S. G. van Riper. 1986. The epizootiology and ecological significance of malaria in Hawaiian land birds. Ecological Monographs 56: 327-344.
van Riper, C. and J. M. Scott. 2001. Limiting factors affecting Hawaiian native birds. Studies in Avian Biology 22: 221-233.
van Riper, S. G. 2000. Japanese White-eye (Zosterops japonicus). Report No. 487 in A. Poole and F. Gill (editors). The Birds of North America. The Academy of Natural Sciences, Philadelphia, PA, and American Ornithologists’ Union, Washington, DC.
Veltman, C. J., S. Nee and M. J. Crawley. 1996. Correlates of introduction success in exotic New Zealand birds. American Naturalist 147: 542-557.
Warner, R. E. 1968. The role of introduced diseases in the extinction of the endemic Hawaiian avifauna. Condor 70: 101-120.
Williams, R. N. and R. C. Fleischer. 1989. Distributions and Habitat Associations of Birds in Waikiki, Hawaii. Pacific Science 43:152-160.
Yamashina, Y. 1961. Birds in Japan. Tokyo: Tokyo News Service, LTD.
Designed by:
Christy Hand
Contact: handch@earlham.edu
Last
Revised: 5 December 2002