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Birds of Hawaii
by
Robert L. Pyle
Bishop Museum, Hawaii
The wild birds inhabiting Hawaii are unique and known worldwide.
Native breeding birds rank among the world's highest in endemism,
endangerment, and extinction, and Hawaii's total bird life contains
a higher proportion of non-native species than perhaps any other
area of comparable size. Interest in Hawaii's birds centers on the
status and trends of its populations, understanding their ecological
requirements, and developing measures to protect and conserve
their remaining populations, which are dwindling at an alarming rate.
The unique nature of Hawaii's bird life results primarily from isolation.
The Hawaiian Islands, a linear archipelago extending some 2,650 km
(1,646 mi) from Kure to Hawaii, is 4,000 km (2,484 mi) from the nearest
point in North America and 3,400 km (2,111 mi) from Asia. The wild
colonizers, individual birds or small groups out over the ocean, were
the first to stumble on Hawaii, where they remained to live and breed.
This process has been going on for millions of years, with two species
repeating the same process within the past 15 years. Then came
evolution of new species in situ, as many of these original colonizers
changed through adaptation and filled unused ecological niches in these
young islands. During the past 2,000 years, humans began inhabiting the
islands, bringing with them some birds that otherwise would never have
reached Hawaii on their own. In addition, some strong-flying species that
regularly migrate long distances have found Hawaii and developed annual
migration patterns that bring them to the islands for part of each year during
the nonbreeding season.
Current Status
It is convenient to categorize the wild birds of Hawaii into residents and visitors.
Resident species remain permanently in Hawaii; visitors regularly come to
Hawaii for only part of each year. Each group can be further divided. Residents
are either native species that arrived or evolved here naturally or alien species
brought in by humans. Visiting birds either come to Hawaii to breed or breed
elsewhere and come during nonbreeding season. True pelagic species, which
spend all their time at sea except when breeding, are considered to have visited
Hawaii if they have occurred in Hawaii's offshore waters within the 200-nautical
mile zone.
Native Residents
Native resident species may be either indigenous, meaning that others of the
same species or subspecies reside elsewhere in the world, or endemic, meaning
that they are found nowhere else. The latter may be endemic at subspecies level,
at species level, or at genus or higher level. For example, endemic at subspecies
level means that others of its species are found elsewhere, but the subspecies
occurs only in Hawaii.
Alien Residents
Polynesians first settled in Hawaii roughly 2,000 years ago (Kirch 1982). Only one bird species brought by the early Polynesians still survives in Hawaii as an established alien species, the red junglefowl (Gallus gallus), ancestor of the domestic chickens. The Hawaiians called it moa, not to be confused with the huge, extinct, flightless birds in New Zealand of the same name. How many other bird species may have been brought by the Polynesians and failed to become established is unknown.
Origins of the 54 alien species established in Hawaii.
Since Captain Cook first visited Hawaii in 1778, alien bird species have been brought to the islands in a steady stream. Only a few have been successful in establishing a continuing breeding population. Of the 54 alien species now considered to have established populations, fully half have origins in Asia (Fig. 1). Fewer are from North America and Africa; a few have come from Australia and South America. Among the continents, only Antarctica is not represented by an established alien species. Penguins have indeed been brought to Hawaii, and one thriving population is in captivity today. But were they to escape, they would not find sufficient krill and ice to maintain a wild population in the islands.
Breeding Visitor Species
Dispersal of the 13 breeding visitor species when not breeding.
Visiting species that come to Hawaii to breed are basically pelagic, that is, living on the open ocean. They come to land to breed, but depart again as soon as parental duties end. Many go to the food-rich boundaries of ocean currents just north of the equator, but some species range throughout the North Pacific (Fig. 2). None appears at any other land during nonbreeding season. First-year birds of most species remain at sea for 3 years or more before returning to breed. Breeding visitors are the albatrosses, shearwaters, petrels, terns, and some tropicbirds. Other seabird species, including boobies, frigatebirds, and noddies, are classed as residents since they remain based at their breeding areas throughout the year, going to sea usually for only a few days at a time.
Nonbreeding Visitor Species
Origins of the 155 visitor species that do not breed in Hawaii.
A great many birds that breed elsewhere depart their nesting grounds after chick rearing is finished, some wandering freely and others following traditional migration routes. Some species, notably the familiar Pacific golden-plover (Pluvialis fulva) and some other shorebirds and ducks, have developed migration patterns that bring large numbers to Hawaii regularly each year, with some individuals even coming to the same plot of ground each winter (Johnson et al. 1981). For other species, just a few individuals turn up each year. For still others, an individual or two may be reported in only a few years out of ten. A number of species have been recorded in Hawaii fewer than a dozen times, perhaps only once or twice. All regular visitors and most others are strong flyers, accustomed to making long migration flights annually, or are larger birds able to store enough energy to reach Hawaii on their own. Almost all are waterbirds.
Only nine species of passerine landbirds are known to have straggled to Hawaii, and most of these have been recorded only one or two times each. Note that absolutely no species of small landbird migrates regularly to Hawaii, either for breeding or in nonbreeding season. Virtually all nonbreeding visitor species nest in the northern hemisphere, most of them in the far north (Fig. 3). A few shearwaters and petrels, a skua, and the great crested tern (Sterna bergii) are the only exclusively southern hemisphere nesters that have straggled to Hawaii.
All Species
Of the 272 species resident now or recorded as visitors (Pyle 1992), roughly 40% are permanent residents, about equally divided between native and alien species (Fig. 4). The breeding visitors, all seabirds, are relatively few. The remaining 55% of the species are nonbreeding visitors. This large percentage for nonbreeding visitor species is not surprising, since these include many species that have wandered to Hawaii as individual stragglers. But in terms of total individuals, the picture is reversed. The nonbreeding visitor species account for probably the fewest individuals, while the breeding visitor seabirds have much larger populations in their huge nesting colonies in the unpopulated Northwest Hawaiian Islands. But the largest of all in total population are the alien residents, which include the ubiquitous Japanese white-eye (Zosterops japonicus), zebra dove (Geopelia striata), and other residents found almost everywhere in the main populated islands.
Bird species currently resident in or visiting Hawaii.
Birds known to have been in Hawaii in the past, but which are no longer there, can be summarized as follows: 16 species (resident-native) have become extinct since Captain Cook's visit; 35 or more species (subfossils, probably native residents) were extinct before Captain Cook's visit; and about 150 species are alien introductions not established. Adding these to the 272 species here now constitutes about 475 species of birds known to have occurred in Hawaii.
Trends
Native Landbirds
Meaningful estimates of total populations of landbirds in Hawaii are difficult to derive. Native species have been confined, at least since Captain Cook's visit, to thickly vegetated and wet higher elevation forests on steep slopes or occupied by deep muddy bogs. Not surprisingly, naturalists over the years could make no real estimates of landbird populations for the island group or even for an individual island, despite the relatively small total areas that were occupied by many of these endemic species.
It was not until the Hawaii Forest Bird Survey in the late 1970's to early 1980's that thoroughly planned fieldwork was conducted, leading to the first comprehensive population estimates for native Hawaiian landbirds. Pioneering techniques for field surveys in such terrain and for statistical analysis were used to obtain population estimates for the native landbird species on all forested islands except O'ahu and Ni'ihau (Scott et al. 1986, 1988). For O'ahu Island, Shallenberger's surveys during the latter 1970's in the Koolau Mountains (Shallenberger and Vaughn 1978) and in the Waianae Mountains (unpublished) have been the most comprehensive.
More recently, Ellis et al. (1993) estimated populations for each native forestbird on each Hawaiian island, based on information available at the end of 1992. These are not directly comparable with the earlier estimates derived from field surveys. However, these estimates and numerous other less comprehensive surveys over the years involving some species on some islands do reinforce a general consensus that Hawaiian forestbirds have declined steadily both in the long term during the past century and in the short term in the past decade.
Resident Waterbirds and Visitors
The Hawaii Division of Forestry and Wildlife has conducted statewide counts of wetland birds semiannually during recent decades. These have included resident wetland species (not the seabirds) and nonbreeding visitor species. Variations in these population counts over the years reflect changes in available wetland habitat, thoroughness of coverage, and possibly some irregular interisland movements. Engilis and Pratt (1993) analyzed these statewide counts for the resident species during 1978-87. Data from earlier surveys covering only certain islands and using less rigorous counting techniques are not readily comparable. Longer-term historical trends in populations of four endangered wetland species are being examined for the Hawaii Wetland Bird Recovery Plan now in preparation by the Recovery Team for the U.S. Fish and Wildlife Service (A. Engilis, personal communication).
The breeding visitor seabirds gather to nest in large colonies in the Northwest Hawaiian Islands. Gross estimates of population numbers for these species made in the 1960's and again in the late 1970's are not comparable for trends analysis because of varying techniques used in attempting to arrive at meaningful numbers in these huge colonies totaling in the millions. Harrison (1990) discussed the difficulties involved in making representative counts and finds no evidence of long-term trends in species numbers, although some wide fluctuations occurred earlier this century. One notable feature has been the return of the Laysan albatross (Diomedea immutabilis) as a breeding visitor to Kaua'i and O'ahu in the main Hawaiian Islands. Since 1977, a steady increase in numbers now measured in the hundreds has local interest but has had a rather small effect on the total statewide population of millions.
Extinction
The rate of extinction within Hawaii's endemic birds is by far the highest in the United States and is approached worldwide only within a few other isolated island groups. At the time of Captain Cook's visit in 1778 some 93 species and subspecies of native birds were breeding in Hawaii, as determined by subsequent discovery and scientific description. In the ensuing two centuries, at least 23 of these have gone extinct (A.O.U. 1983) and another 13 are imperiled. Recent discoveries of the bones of prehistorically vanished species now reveal a vast array of former birds that became extinct long before Captain Cook arrived. Thirty-five of these have already been scientifically described (Olson and James 1991) and must represent only a small fraction of the forms of birds that existed prehistorically in Hawaii.
Extinctions over the past 200 years (Fig. 5) show a disproportionate number of bird species vanishing during the 1890's, a decade concluding a period of intense discovery and collecting of Hawaii's birds. A similar large decline in the 1980's represents nine forms not reported since then. Ralph and van Riper (1985) discussed the factors that have contributed to the decline in Hawaiian bird populations since the arrival of the Polynesians.
Extinction of native breeding birds since 1778. Steps mark the decade of the last record for each form considered extinct (A.O.U. 1983). The 70 forms shown as currently existing include 13 in peril, with steps marking the decades of their last known records. Yellow represents prehistoric forms.
Aliens
An early listing of the alien species in Hawaii was that of Caum (1933), who identified 92 species as alien introductions. These may be categorized as established, not established, or uncertain. Most (75%) of the alien species established in 1933 are still present (Pyle 1992). Few of those deemed uncertain or not established in 1933 have persisted until today. Introductions continued during the 1940's and 1950's but thereafter were severely curtailed by stronger governmental restrictions on importation of wild birds. Of the 54 alien species considered established in Hawaii today, 31 (57%) had been introduced more than 60 years ago, and 23 (43%) have been introduced and have become established since 1933.
Conclusion
Hawaii's birds comprise four groups: native and alien resident species, and breeding and nonbreeding visitor species. Factors affecting population levels differ markedly among the groups. Although current status of species within all groups is fairly well understood, assessing meaningful trends for species is difficult for lack of comparable quantitative data on statewide populations over time.
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