Chapter 7. In the Air
Native Birds of the Hawaiian Islands

[Illustration or Photograph: Native Birds]

Birds, birds, birds! Birds by the hundreds, thousands, millions! Seabirds, landbirds, wetland birds, migrating birds. Hawks and owls and crows.
Albatross and native ducks. Stilts perched on tall legs in the taro fields. A goose adapted to walking on barren lava flows. In the dense higher forests are birds with every manner of beak for probing bark, cracking open seeds, or extracting the nectar of native flowers.
These are the native birds of the Hawaiian Islands.

Wetland Birds of the Hawaiian Islands

Hawaiian Black-Necked Stilt Ae‘o

Hawaiian Coot ‘Alae Ke‘oke‘o

Koloa the Hawaiian Duck Koloa Maoli

Hawaiian Moorhen ‘Alae ‘Ula

Black-Crowned Night-Heron ‘Auku‘u

Seabirds of the Hawaiian Islands
Noddy Terns Noio Kōhā
The Great Frigatebird ‘Iwa
Laysan Albatross Mōlī
Pacific Golden Plover Kōlea
Newell’s and Wedge-Tailed Shearwaters ‘A‘o; ‘Ua‘u Kani
Red-Tailed and White-Tailed Tropicbirds Koa‘e ‘Ula; Koa‘e Kea

Forest and Mountain Birds of the Hawaiian Islands
Hawaiian Owl Pueo
Hawaiian Hawk ‘Io
Hawaiian Crow ‘Alalā
Hawaiian Goose Nēnē
Honeycreepers and Honeyeaters
(and other small forest birds)
Original Colonizing Species
Populations and Extinctions
Hawaiian Featherwork
Hawaiian Honeycreepers;
Evolutionary Processes / Adaptive Radiation
‘Akialoa and ‘I‘iwi;
The ‘Elepaio
The Puaiohi—Captive Breeding

Threats to Endemic Forest Birds




Wetland Birds of the Hawaiian Islands

[Illustration: Koloa Duck or Black-Necked Stilt]

Hawaiian Black-Necked Stilt Ae‘o Himantopus mexicanus knudseni

Hawaiian Coot ‘Alae Ke‘oke‘o Fulica alai

Hawaiian Duck Koloa Maoli Anas wyvilliana

Hawaiian Moorhen ‘Alae ‘Ula Gallinula chloropus sandvicensis

Black-Crowned Night-Heron ‘Auku‘u Nycticorax nycticorax hoactli

Threats to Wetland Bird Species of the Hawaiian Islands




Hawaiian Black-Necked Stilt (Himantopus mexicanus knudseni)
Hawaiian Name: Ae‘o
Endemic subspecies.
Status: Endangered Species. Found on Hawai‘i Island, Maui, Lāna‘i, Moloka‘i, O‘ahu, Kaua‘i and Ni‘ihau.
[Illustration: Hawaiian Black-Necked Stilt]

Perfectly suited to a wetland environment, the Hawaiian Black-necked stilt is a wading bird, easily identified by its long, skinny pink legs, black and white forehead, and white breast. Black-necked stilts grow to about 16 inches (41 cm) tall. The birds’ long, jointed legs bend in the opposite direction of the human leg.

He kukuluāe‘o.
A stilt.
A thin, long-legged person.
(Pukui: 709-79)

Hawaiian stilts are known as ae‘o, which means “one standing tall,” or kukuluae‘o, which is also the Hawaiian term for wooden stilts that were used for amusement by Hawaiian children in ancient times. The wooden stilts were most commonly made from the wood of ‘ohe (Reynoldsia sandwicensis), a tree also known by the Hawaiian term ‘ohe kukuluae‘o.
Hawaiian Black-necked stilts are wading birds that often gather in groups. They make a chirping sound similar to “kip kip” or “keek, keek” and use their long beaks to probe the shallow water mud flats for food, including worms, aquatic insects, crabs, fish and mollusks.
The stilt breeding and nesting season is from December to August (predominantly from March to August). The stilt builds a nest in a shallow depression in a small mound, often on the banks of lo‘i kalo (taro patches) or in low-lying vegetation areas near the water. The stilt lines its nest with rocks and twigs.
The female stilt usually lays four eggs. The eggs are well camouflaged, and the stilt aggressively defends the nest. The eggs incubate for 24 to 26 days.
The stilt chick is covered with a downy, tannish-brown coat speckled with black, and leaves the nest soon after hatching. The chicks hide under cover until ready to fly. Parents don’t feed the chick but instead help them find suitable food sources. Sometimes the parent will fake a broken wing, feigning injury in order to draw predators away from the nest and hatchlings.
Many hatchling chicks don’t make it through their first months. At least 44 stilt chicks were hatched on Kaua‘i’s Hanalei National Wildlife Refuge in 1994, but only one was observed to have fledged. The precise causes for this low survival rate are unknown, but all of the following may play a role: diseases, parasites, poor food supply and/or food quality, and predation by bullfrogs, cats, dogs, pigs, owls, and possibly also cattle egrets and black-crowned night-heron. Large introduced bullfrogs (Rana catesbeiana) also have been seen preying on stilt chicks.
Historically, stilt populations have also suffered due to loss of wetland habitat as well as the proliferation of invasive, non-native plant species. Many stilts have died due to a soil bacteria that has caused botulism in the stilts’ food sources. This has caused sporadic but large die-offs of affected birds.
Black-necked stilts (ae‘o) in the Hawaiian Islands constitute a subspecies of the Black-necked stilts found in North and South America, which have less black on their neck and face, and a shorter bill and tail. The North American stilts are found as far south as Brazil and also in the Galapagos Islands.
Hawaiian Black-necked stilts are known to occasionally fly between Hawaiian Islands, particularly between Ni‘ihau and Kaua‘i. The birds are found mostly in lowland areas, but are also seen at an elevation of 2,300 feet (700 m) on the island of Hawai‘i at the Pu‘u Wa‘awa‘a reservoir.

Hawaiian Black-Necked Stilt Population
Stilts were once hunted as game birds in the Hawaiian Islands. The U.S. Fish and Wildlife Service listed the Hawaiian subspecies as endangered on October 13, 1970. In 1982, the population of stilts was estimated to be less than 1,000, and found mostly on Maui and O‘ahu.
Stilts are found on O‘ahu, Hawai‘i Island, Maui, Lāna‘i, Moloka‘i, Kaua‘i and Ni‘ihau. There are currently about 1,400 to 1,800 Black-necked stilts in the Hawaiian Islands, with the biggest populations of the birds on Maui, Kaua‘i and O‘ahu. The stilt is also found on the island of Hawai‘i’s Kona Coast, where the stilt population grew to more than 220 in 2001 from an estimated 105 in 1998.
The Lāna‘i stilt population was first documented in 1989, and seems to be gradually increasing. Moloka‘i’s stilt population has also grown steadily since 1989 (an estimated 20 stilts) with at least 148 stilts on Moloka‘i in 2001. A key factor in the growing populations appears to be the creation of new habitat suitable for the stilts. On Lāna‘i a wastewater treatment plant created the habitat, while on Moloka‘i the birds benefited from the restoration of loko i‘a (fishponds).
[Photograph: Hawaiian Black-necked stilt]

Hawaiian Coot (Fulica alai)
Hawaiian Name: ‘Alae Ke‘oke‘o or ‘Alae Kea
Endemic subspecies.
Status: Endangered Species. Found on Hawai‘i Island, Maui, Lāna‘i, Moloka‘i, O‘ahu, Kaua‘i.
‘Alae ke‘oke‘o, the Hawaiian coot, was formerly considered a subspecies of the American coot (Fulica americana), which is found from Panama to Canada and occasionally seen in the Hawaiian Islands. The Hawaiian coot, now classified as a unique species (Fulica alai), is generally darker, and has a larger frontal shield and a more slender bill than its North American relatives. The Hawaiian coot also displays different patterns of white on its head than the American coot.
While the Hawaiian moorhen (‘alae ‘ula) has a red bill (‘ula means “red”), the Hawaiian coot (‘alae ke‘oke or ‘alae kea) has a white bill (ke‘oke‘o and kea both mean “white”).
[Illustration: Hawaiian Coot]

The Hawaiian coot is about 14 to 15 inches (35 to 38 cm) long, and mostly dark gray to black on top, with white undertail feathers. The bill is ivory white, as is the bulbous frontal shield (or frontal knob), though rarely a bird will have a brown or red frontal shield.
Coots feed on insects, fish, and tadpoles as well as the leaves and seeds of aquatic plants. Coots don’t fly much but are able to sustain flight low to the water.
Coots may nest throughout the year, particularly between March and September, building a floating nest on wetland vegetation (e.g., in lo‘i kalo, or taro patches) using sedges, weeds, taro stems, grass, and other aquatic plants. The nest may rise and fall with water levels.
After building its nest, the coot lays usually from three to ten eggs (averaging four to six eggs). The creamy to tan-colored eggs are speckled with black and incubate for three to four weeks.
The coot chick is downy black with a reddish-orange neck, head, and bill and has a baldish appearance due to the absence of down on the crown and forehead. The chick’s reddish-orange bill has a black tip. Soon after hatching, the chicks are able to swim.

Coot Population
Historical records do not indicate there ever were a large number of coots, yet the bird was significant in Hawaiian mythology and some early accounts reported large populations on Kaua‘i. A 1982 count documented 700 coots, mostly on Kaua‘i, O‘ahu and Maui.
The Hawaiian coot is federally listed as an endangered species. There are currently an estimated 2,000 to 4,000 coots living throughout the main Hawaiian Islands, with likely few to none of the birds on Kaho‘olawe and Lāna‘i.
The largest coot populations are found on Kaua‘i, O‘ahu and Maui. Coots are currently listed as endangered on Hawai‘i Island, Maui, Lāna‘i, Moloka‘i, Oahu, and Kaua‘i. Coots are known to fly between the Hawaiian Islands.
[Photograph: Hawaiian Coot]

Hawaiian Duck(Anas wyvilliana)
Hawaiian Name: Koloa Maoli (“Native Duck”)
Endemic
Status: Endangered Species. Found on Hawai‘i Island, O‘ahu, Kaua‘i; possibly extinct on Maui.
Male koloa are about 20 inches (51 cm) long while female koloa are about 17 inches (43 cm) long. Koloa are mottled golden brown in color, with a greenish head and olive-colored bill. Male koloa have a darker head than females, and some females have a bill with an orange tip.
The koloa’s secondary wing feathers (speculum) are greenish-blue to metallic purple in color, with white borders. The duck’s feet and legs are orange.
[Illustration: Hawaiian Duck]

Koloa eat insects, mollusks, and aquatic vegetation. Koloa breed and nest year round, particularly between December and May, beginning at about one year of age. The birds often nest near mountain streams and taro patches as well as near irrigation ditches, low elevation wetlands, and river mouth areas.
A well-hidden nest is built using feathers and down, and usually from two to ten white to tan-colored eggs are laid (the average clutch size is eight eggs). The koloa eggs incubate for about 28 days and then hatch. After about nine weeks the birds learn to fly.

Koloa Population
Koloa are widely distributed on Kaua‘i, where more than 80% of all koloa are found. Once abundant on all of the main Hawaiian Islands except Lāna‘i and Kaho‘olawe, by 1915 koloa became rare on all of the Hawaiian Islands except Kaua‘i.
By 1947, an estimated 30 or fewer koloa were on O‘ahu and these birds were gone by 1960. In 1962 the World Wildlife Fund and the Federal Aid to Wildlife Restoration Act provided money for a bird restoration program. By 1979, more than 350 koloa had been released on O‘ahu and 300 more on the island of Hawai‘i.
The koloa’s population is now estimated at less than 2,500. More than 80% of all koloa are found on Kaua‘i. Some koloa have been relocated to O‘ahu and Hawai‘i Island. The koloa was listed as a federally endangered species in 1967. The koloa is currently listed as endangered on Hawai‘i Island, O‘ahu, Kaua‘i, and listed as endangered and possibly extinct on Maui.
The koloa evolved from the common mallard (Anas platyrhynchos) into a unique Hawaiian species (not a subspecies). The common mallard is found in North America, Europe and Asia. The endemic Hawaiian species closely resembles the female North American mallard. One threat to the koloa comes from interbreeding with domestic mallards, resulting in hybridization of the endemic Hawaiian species.
Two of the largest koloa populations in the Hawaiian Islands are found on the island of Kaua‘i, at the Hanalei Valley National Wildlife Refuge and the Alekoko (Menehune) Fishpond. These populations are also thought to be the least hybridized (and thus the most genetically pure). In addition to many other threats to this wetland species, koloa chicks may be eaten by dogs, cats, pigs, muskrats, rats, bass, and bullfrogs.
[Photograph: Koloa maoli]



Hawaiian Moorhen(Gallinula chloropus sandvicensis)
Also called: Hawaiian Gallinule or Mudhen
Hawaiian Name: ‘Alae ‘Ula (red forehead)
Endemic subspecies.
Status: Endangered Species. Found on O‘ahu and Kaua‘i; possibly extinct on Hawai‘i Island, Maui and Moloka‘i.
The moorhen is easily identified by its bright red frontal shield. Because the moorhen sometimes feeds out on the mud flats, it is also known as the mudhen. Hawaiian legend tells of how the ‘alae ‘ula brought fire from the gods to the Hawaiian people. During the journey, the moorhen was scorched by the flames, giving the bird its red forehead, or frontal shield. The demigod Māui tried to catch the ‘alae ‘ula, “...in order to learn the secret of making fire...He caught them before they could hide and forced them to yield the secret of fire.”[1]
[Illustration: Hawaiian Moorhen]

The endemic Hawaiian moorhen (‘alae ‘ula) is a subspecies of North America’s common moorhen, also found in Eurasia. A non-migratory bird, the adult Hawaiian moorhen is about 13 inches (33 cm) long, with a black head and neck, and a slate-gray to bluish-black back, which may be somewhat iridescent. The flanks and undertail feathers are white. The moorhen’s red bill has a yellow to light green tip. Aside from its red shield, the moorhen’s dark gray plumage looks very similar to the ‘alae ke‘oke‘o (Hawaiian coot, Fulica alai).
The moorhen’s feet are not webbed, and the bird’s yellowish-green legs and feet may show red near the top. Moorhen are adept at walking across floating wetland vegetation. An ancient Hawaiian proverb, “Hanopilo ka leo o ka ‘alae,” translates to “Hoarse is the voice of the mudhen,” and was said of someone “...who talks himself hoarse.”[2]
Generally a shy and secretive bird, the moorhen likes to hide in dense cover, usually only coming out into the open in the morning and evening to feed. They inhabit wetlands, including freshwater marshes and lo‘i kalo (taro patches). Moorhen feed on aquatic plants (e.g., taro tops and young shoots), as well as animals, including mollusks and insects.
March through August is the main breeding period for moorhen, but they may breed year round. The moorhen’s courtship behaviors include bowing and arching as well as nibbling. During breeding season the bird’s frontal shield may become enlarged and is also a deeper red color. When it comes time to nest, the moorhen will build a well-hidden nest, often on folded reeds, using plants and mud.
Moorhen lay from five to nine cream-colored eggs (which are spotted with grey, black and brown). The eggs incubate for about 22 days. The downy moorhen chick has a bright red bill and pale yellow to brown body, and is able to swim soon after hatching. Immature (juvenile) moorhen are olive-brown to grayish-brown in color.
The moorhen, also called the Hawaiian gallinule or mudhen, is thought to have been quite common throughout the Hawaiian Islands in the 1800s until the population declined rapidly in the early and mid-1900s. The Hawaiian moorhen was federally listed as endangered in 1967 and remains an endangered species today. The moorhen is listed as endangered on O‘ahu and Kaua‘i, and endangered and possibly extinct on Maui, Hawai‘i Island, and Moloka‘i.
A 1982 count by the State Division of Forestry and Wildlife documented 194 moorhen on O‘ahu and Kaua‘i with perhaps a few scattered birds on Maui and Moloka‘i. In the 1970s and 1980s, aquaculture and taro cultivation created more suitable moorhen habitat, increasing the bird’s survival rate.
The moorhen’s secretive nature leads to uncertainty in current population estimates, which range from 150 to 900 birds left in the wild. Moorhen are found only on O‘ahu and Kaua‘i, with few to none on Maui and Moloka‘i and probably none on Hawai‘i Island. Efforts to reintroduce moorhen on Hawai‘i Island and Maui have not been successful.
[Photograph: Hawaiian Moorhen]

Black-Crowned Night-Heron (Nycticorax nycticorax hoactli)
Hawaiian Name: ‘Auku‘u
Indigenous: American continent.
[Illustration: Black-Crowned Night-Heron]

The Black-crowned night-heron is about 25 inches (63 cm) long with a wingspan of almost 4 feet (1.2 m). The night-heron has yellow legs, and a black head, back and bill, but otherwise is mostly gray. The birds are often seen standing motionless at the edges of lagoons, marshes, canefield ditches, lo‘i kalo (taro patches), ponds and exposed reef areas. The night-heron may be heard making a fairly loud “kwok” sound while flying.
When in breeding plumage, the male night-heron develops four or five long white head plumes, while the female may have just two or three. These white nuptial plumes grow out from the back (occipital) region of the bird’s head. Immature night-heron are streaked with white and brownish-rust colors that turn mostly gray as the bird matures.
An ancient Hawaiian proverb, “Kohā ka leo o ka ‘auku‘u,” translates to “The voice of the ‘auku‘u is heard to croak,” and referred to “...a snooping gossip. The ‘auku‘u bird lives in the upland and goes to the lowland for fish, often snatching them from people’s ponds.”[3]
The Black-crowned night-heron feeds primarily on crustaceans and fish, but also eats mice, frogs, aquatic insects, and chicks of other bird species (e.g., Hawaiian Black-necked stilts, Brown noddies, Sooty terns). The night-heron is a solitary wading bird. Looking for prey beneath the water, it is most active at dawn and dusk, and may feed during the night as well as during the day.
Black-crowned night-heron breeding occurs around May. Using large sticks and twigs, the night-heron constructs a nest in a tree and lays usually two to four bluish-green eggs. The Hawaiian Black-crowned night-heron is indistinguishable from the North American Black-crowned night-heron.
A Hawaiian chant to the ancient chief Kuali‘i mentions the ‘auku‘u, a bird that was once found on all the main Hawaiian Islands but today is much less common. Kuali‘i was a warrior chief known for his strength and bravery.
[Photograph: Black-Crowned Night-Heron]


Threats to Wetland Bird Species of the Hawaiian Islands
A primary threat to native wetland birds is the destruction of their habitat due to urbanization, diversion of water, and development. Other threats to wetland birds include disease, parasites, deterioration of food quality and supply, and introduced species such as cats, dogs, pigs, rats, barn owls, mongoose, cattle egrets, and even bullfrogs (which may prey on small chicks).
Water contaminants, such as insecticides, heavy metals and petroleum products are also destructive to the survival of native wetland species. Introduced plants such as the mangrove and the water hyacinth also make habitat unsuitable for native birds.
The cattle egret, an introduced species, may prey on young native birds as well as compete for food resources. Introduced fish, such as tilapia, may also consume valuable food resources, affecting native bird populations.
For a more in-depth discussion of threats to native Hawaiian species, see Overview of Native and Polynesian-Introduced Species of the Hawaiian Islands, Chapter 5.
[Photograph: Wetland bird in habitat]


Seabirds of the Hawaiian Islands

[Illustration: Black or Brown Noddie]




Noddy Terns Noio Kōhā
The Great Frigatebird ‘Iwa
Laysan Albatross Mōlī
Pacific Golden Plover Kōlea
Newell’s and Wedge-Tailed Shearwaters ‘A‘o; ‘Ua‘u kani
Red-Tailed and White-Tailed Tropicbirds Koa‘e ‘Ula; Koa‘e Kea




Overview of Hawaiian Seabirds

About 9,700 species of birds inhabit Earth, but only about 3% of these species are marine birds. The Hawaiian Islands have at least 22 species of native seabirds, which combine to form a total population of more than 12 million seabirds in the Hawaiian Islands. Most of the Hawaiian Islands’ seabirds live on the Northwestern Hawaiian Islands, which are hundreds of miles northwest of the eight main Hawaiian Islands.

[Photograph: Seabird over the ocean]

‘Au i ke kai me he manu ala.
Cross the sea as a bird.
To sail across the sea. Also applied to a hill that juts out
into the sea or is seen from far out at sea.
(Pukui: 237-28)




Hawaiian Noddy Tern (Anous tenuirostris melanogenys)
Hawaiian Name: Noio Kōhā
Status:
[Illustration: Noddy tern]

More than 60% of seabirds in the Hawaiian Islands are noddies and terns, and there are three native species of each. Noddies are actually a type of tern, but are distinguished from other terns by their distinctive nodding and bowing behavior during mating ceremonies.
Noddies are seabirds, but they lack substantial oil glands at the base of their tails, so they cannot alight on the water for more than a few minutes. Noddies and terns present a fish to their mate during mating ceremonies. A male noddy will nod and bow when he feeds his mate before she lays her egg.

Ua ho‘i ka noio ‘au kai i uka, ke ‘ino nei ka moana.
The seafaring noio bird returns to land, for a storm rages at sea.
A weather sign.
(Pukui: 2787-307)

Black noddies have yellow-orange legs and feet, and vary in color from dark gray-brown to sooty black. They live throughout the year along the coasts of the main Hawaiian Islands as well as throughout the Northwestern Hawaiian Islands. Black noddies build substantial nests on rocky ledges, in caves, or in trees. They may live to 25 years of age.

Brown noddies nest on the ground, usually laying a single egg near ground vegetation. Brown noddies are found in the Northwestern Hawaiian Islands as well as on the offshore islets of the main Hawaiian Islands.

[Photograph: Noddies bowing]




The Great Frigatebird(Fregata minor palmerstoni)
Hawaiian Name: ‘Iwa
Indigenous: Tropical Pacific.
To look up and see a frigatebird soaring overhead is like stepping back into the age of dinosaurs. With a forked tail and bent wings, the frigatebird looks like an ancient pterodactyl in flight.
Frigatebirds are also called “man-o’-war birds,” a name given to them by sailors who thought they resembled the favorite pirate vessels known as frigates.
There are five kinds of frigatebirds in the world, but the only type of frigatebird in the Hawaiian Islands is ‘iwa, also known as the great frigatebird. A wide-ranging species, frigatebirds banded in the Hawaiian Islands have later been seen in the Philippines.
[Illustration: Frigatebird (wing profile)]

He ‘iwa ho‘ohaehae nāulu.
An ‘iwa that teases the rain clouds.
A beautiful maiden or handsome youth who rouses jealous envy in others.
(Pukui: 645-73)

Anatomy
The great frigatebird’s body may average 43 inches (109 cm) long with a wingspan that may exceed 90 inches (229 cm), yet the bird usually weighs less than 3 pounds (1.4 kg). The female frigatebird is bigger than the male. Females are black with some white feathers on the upper breast and throat, while males have all black feathers.
Male frigatebirds have an inflatable red pouch of skin under their throat, and they often blow up this throat (gular) pouch like a balloon, particularly when they are near a colony of birds and want to get a female’s attention. Frigatebirds can glide for several hours with very little effort, often soaring at heights above 500 feet (152 m), higher than any of the other seabirds in the Hawaiian Islands. This is helpful since they have very little webbing between their toes, and don’t land on the water if they can avoid it. Lacking oil glands to waterproof their feathers, frigatebirds don’t dive beneath the ocean’s surface for fish like some other seabirds.

Feeding
Frigatebirds can dive very fast and make sharp spiraling turns, and are probably the most acrobatic of all of Hawai‘i’s seabirds. Frigatebirds may chase and dive down upon other seabirds, such as red-footed boobies and shearwaters, in order to force them to drop or disgorge their food. The frigatebird then uses its agility and speed to swoop down and catch the food before it hits the water. This piracy explains how the frigatebird got its Hawaiian name, ‘iwa, which means “thief.”
In addition to stealing food from other seabirds, frigatebirds fly low over the water and use their long, hooked bills to grab floating food, including squid, fish, newly hatched sea turtles, and even mālolo (flying fish).

Frigatebird Nesting
Frigatebirds are migratory, traveling between their nesting areas and places where food is more plentiful. Though frigatebirds are commonly seen flying over the Kaua‘i shoreline, they do not nest there. Most of Hawaiian frigatebirds nest on the Northwestern Hawaiian Islands, which begin hundreds of miles northwest of Kaua‘i.
Frigatebirds are seen near in the Northwestern Hawaiian Islands during winter months, and are seen on the main Hawaiian Islands more often in summer. Frigatebirds are also known to nest on Ka‘ula Island, which is about 22 miles (35 km) southwest of Ni‘ihau and also on Mokumanu (“Bird Island”[iv]), off of Mōkapu Point on the east side of O‘ahu.
Frigatebirds are biennial breeders, nesting every other year beginning at about five years of age. In breeding season the males gather together and show off their puffed up red throat pouches. The birds gather nesting material (branches and twigs) to build relatively flat nests in bushes or other vegetation.
Nesting material is gathered primarily by the female frigatebirds, but it’s primarily the male that builds the nest. The female then lays one white egg around March or April. Male and female frigatebirds take turns incubating the egg, which basically means that they sit on it. After about 1½ months, the chick hatches.
The baby chick has no feathers at first, and so the parents must protect the baby from the sun. Soon the hatchling is covered with white, downy feathers. The baby gets fed by the adults about every 18 hours and stays in the nest about 4½ months before it grows adult feathers and fledges (learns to fly), which usually occurs in October. Frigatebirds have been documented to have reached 34 years of age.
Some ancient mariners kept frigatebirds on their ships. The sailors knew that when a frigatebird was released it would fly straight toward land, helping the sailors know which way to go. Frigatebirds feathers were also utilized in ancient Hawaiian featherwork.
[Photograph: Frigatebird, red throat pouch]


Laysan Albatross (Diomedea immutabilis)
Hawaiian Name: Mōlī
Indigenous: Central and North Pacific.
Along the oceanside cliffs of the Hawaiian Islands, adult albatross perform elaborate courtship dances with each other. Then they breed, nest, lay their eggs, and nurture their young. Quite a site to see, fat and fluffy baby albatross are sometimes bigger even than their parents.
Eventually the albatross parents leave, and the young ones must learn to fly on their own. Finally the fledglings head out to sea, not to return until years later when it is their turn to nest.
[Illustrations: Albatross mating dance; baby albatross.]

Physiology, Migration, Feeding and Nesting
The Laysan albatross may be nearly 3 feet (91 cm) long with a wingspan up to 80 inches (2 m). Adult Laysan albatross weigh from 5 to 7 pounds (2 to 3 kg), making them the largest seabirds in the entire Pacific region. Laysan albatross are white, with black on their upper wings and around their eyes. This black coloration around the eyes is a common trait in seabirds, and helps to reduce glare.
Albatross may live for more than 40 years, and may stay at sea up to five or more years before returning to land to nest. The birds often spend the summer months about 1,000 miles (1,600 km) or more from the Hawaiian Islands, on and over the waters of the North Pacific Ocean. Laysan albatross do not dive beneath the water, but instead fish from a sitting position on the surface. This makes them particularly vulnerable to shark attacks, which occur quite frequently in the birds’ Northwestern Hawaiian Islands habitat.
Laysan albatross usually arrive in the Hawaiian Islands by late October or early November, and stay at their nesting areas until June or July. Even during the nesting season they may fly thousands of miles in search of food, then return directly to their island nests to feed their young. Laysan albatross feed mostly on large squid as well as the eggs of the mālolo (flying fish).
As graceful as Laysan albatross are in flight, they are fairly awkward on land. Laysan albatross sometimes have difficulty taking off for flight unless they can run down a slope or use some other suitable launching area. Once in the air, however, Laysan albatross can soar for hours without flapping their wings, and may even sleep while airborne.
Laysan albatross prefer to nest in the same area where they were born, often returning to the exact same location. They also tend to mate with the same partner for life. The female lays one egg in a nest depression, usually in November or December. The parents alternate tending the egg, which incubates for about 64 days, and then the chick may take several days to break free from the shell. After about 5½ months the chicks fledge (learn to fly).
During the nesting season in the Hawaiian Islands, albatross often perform bizarre courtship dances that include prancing around with their mate and thrusting their beaks skyward along with bill snapping and vocalizations. These courtship displays, as well as the difficulty the chicks have learning to fly, are likely reasons for the birds’ nickname: “gooney bird.”

Endangered Status
Worldwide, many types of albatross are endangered due to hunting. Early explorers utilized their meat, oil and feathers. In just one 17-year period, more than five million albatross were killed for their downy body feathers, which were used as stuffing for quilts and mattresses sold in European markets. Also highly valued were the albatross’ long white tail feathers and wing feathers, which were prized for pen plumes.
Laysan albatross nest on the main Hawaiian Islands, particularly on O‘ahu, Ni‘ihau and Kaua‘i, though many of their traditional nesting areas no longer provide good habitat because they have been developed. Still the birds persist in some areas near where homes and other structures have been built. The albatross remain particularly vulnerable to domestic animals (e.g., cats and dogs) that pose a continuing threat to the birds (especially to baby albatross).
On Kaua‘i, Laysan albatross have been making a comeback, particularly at Kīlauea Lighthouse National Wildlife Refuge on the island’s north shore, and at the Pacific Missile Range Facility on Kaua‘i’s southwest shore. Most albatross in the Hawaiian archipelago are found on the Northwestern Hawaiian Islands, a group of reefs, islets and atolls located hundreds of miles northwest of Kaua‘i. At the far northwest of the Northwestern Hawaiian Islands are the French Frigate Shoals, Midway Atoll and Kure Atoll, which are home to the largest albatross populations of the Hawaiian Islands.
Midway Atoll has the largest Laysan albatross population anywhere, with nearly 400,000 nesting pairs. Midway also has nearly 20,000 black-footed albatross (Diomeda nigripes) and there are occasional sightings of the rarely seen short-tailed albatross (Diomedea albatrus), which is federally listed as an endangered species. The short-tailed albatross is also known as the golden gooney because of its yellow head and neck feathers and its pink bill.
As of January 2001, there were an estimated 1,150 short-tailed albatross remaining worldwide, making them one of the world’s most endangered seabirds. The species main breeding area is Torishima Island off Japan. One short-tailed albatross banded at Torishima in 1993 was spotted years later at Midway.

Satellite Tagging of Laysan Albatross in the Hawaiian Islands
In 1999, ornithologists working in conjunction with Kīlauea Point National Wildlife Refuge and the U.S. Fish and Wildlife Service attached transmitters to Laysan albatross nesting near Kīlauea Lighthouse on Kaua‘i’s north shore. When the birds left Kaua‘i to find food for their young, the transmitters sent signals to a satellite that relayed the birds’ latitude and longitude data back to Earth.
Results from a previous study based on Tern Island found that one albatross flew 24,843 miles (40,000 km) in just 90 days. The bird traveled all the way across the North Pacific and back, a distance equal to flying all the way around the world. Another bird made repeated trips east to the San Francisco Bay area and then back to the Hawaiian Islands to feed its young. Continuing research will reveal more about the large, graceful albatross birds, including their unique courtship habits and their amazing journeys in search of food.
[Photograph: Albatross]



Pacific Golden Plover (Pluvialis fulva)
Hawaiian name: Kōlea
Indigenous
Pacific golden plovers migrate to the Hawaiian Islands during fall to feed, then return north to Alaska again for the summer months to lay their eggs. This is similar to the migration habits of koholā (Megaptera novaeangliae, humpback whales), which also come from Alaska to spend their winters in the Hawaiian Islands. The humpback whales, however, come to the Hawaiian Islands in winter to mate and give birth, and go north in the summer to feed.
An ancient Hawaiian proverb, “O Kā‘elo ka malama, kāpule ke kōlea,” (“Kā‘elo is the month when the breasts of the plovers darken,”) refers to “...the month when the plovers are fat and ready to fly on their migration to the north.”[v]
[Illustration: Pacific golden plover]

The Pacific golden plover’s non-stop flight between Alaska and the Hawaiian Islands may cover more than 3,500 miles (5,633 km) but take less than three days. After spending May, June, and July on the Alaskan tundra, the plovers head south in August (some arrive in July). The plovers often arrive in flocks that disperse once they reach the Hawaiian Islands. While plover adults usually begin arriving in the Hawaiian Islands in late August, juveniles arrive in late September, staying until late April, or May, when they again head north.

Anatomy and Physiology
Pacific golden plovers are about 11 inches (28 cm) long, weigh about ½ pound (.23 kg), and live to about eight years of age. Plovers are dark brown, and spotted with gold on top, but paler on the underside. Plovers have a large, dark eye region, and a fairly large head relative to their body.
Before leaving the Hawaiian Islands in April, the female plover molts into a beautiful beige-gold breeding plumage. The male plover gets a “tuxedo” look, with a pure white stripe along the sides of the head and down the neck. The abdomen, breast, cheek and throat are black.

Feeding
Pacific golden plovers are known to return to the same general area each year to feed, often returning to the same specific patch of grass, where they will remain throughout the season. They prefer open grassy areas, including pastures, lawns, and golf courses. In the Hawaiian Islands the plover is very territorial, and will defend its feeding area against other plovers, though it will often ignore other bird species. Some plovers move around a bit more, and are not as territorial when it comes to their feeding grounds.
During its winter stay in the Hawaiian Islands, the plover feeds on insects and other invertebrates, as well as certain flowers and leaves. Plovers often run in short bursts, intermittently stopping to look for insects. Feeding for about four months in the Hawaiian Islands, plovers fatten up for the long journey north.

Migration
Beginning in late April, the plovers head north again to the Alaskan and Siberian tundra, a journey that may span over 3,500 miles (5,633 km). Plovers are often very solitary birds during their stay in the Hawaiian Islands, but before they begin their migration north they gather together in large flocks and then head north.
The departure dates may be quite regular among certain Hawaiian Island plover populations. For example, the O‘ahu birds are said to always leave within days of April 25, but this may vary.
Once the plovers begin their journey north, they may sustain speeds of 60 to 70 miles per hour (97 to 113 km/hour), making them among the world’s fastest birds when it comes to sustained level flight, which excludes speeds attained by diving birds such as raptors.
During their migrations, plovers fly at elevations up to 20,000 feet (6,100 m). With a tailwind, the birds may reach speeds of over 100 miles per hour (161 km/hour), and the flight to the northern breeding grounds may be completed in 50 to 60 hours.
Some plovers may remain in the Hawaiian Islands during the summer months. New efforts to track plover migrations, estimate population size, and learn more about plovers are being coordinated through the website: www.hawaii.edu/bird. The Hawaiian Pacific golden plover population is currently estimated at about 2,500 birds. Plovers have also been seen in South Pacific Islands, and these plovers may be migrating much farther than the birds that winter in the Hawaiian Islands.

Nesting
Once plovers reach their arctic nesting grounds, the males build nests and share in the duties of caring for the chick. Most but not all males change partners from year to year. Plovers typically lay four, greenish-brown eggs (one every other day). Together the eggs total about two-thirds of the female’s body weight, and may be more than 50% yolk, which is a higher percentage of yolk than the eggs of most other birds. The plover’s nest is lined with lichens and leaves. The eggs are incubated by the female at night and the male during the day.
Less than a month after hatching, the downy goslings and ducklings begin to fledge (learn to fly). In early August the parents begin to leave, but the chicks will not follow until one or two months later. This next generation of plovers must then find their way to the remote Hawaiian archipelago thousands of miles away. The journey south will be the beginning of a cycle of migration that will then be repeated each year.
[Photograph: Plover]


Shearwaters—Newell’s and Wedge-Tailed
[Illustration: Newell’s and wedge-tailed shearwater]

Newell’s (Townsend’s) Shearwater (Puffinus auricularis newelli)
Hawaiian Name: ‘A‘o
Endemic subspecies
Status: Threatened subspecies. Found on Kaua‘i, O‘ahu, and Hawai‘i Island.
The Newell’s shearwater, a local variety of the Townsend’s shearwater, has a black back, a white breast, and is about 13 inches (33 cm) long, with a wingspan that may exceed 3 feet (91 cm).
Newell’s shearwaters breed on the Hawaiian Islands from April to November, flying to their nesting colonies each day at dark and leaving again before dawn. The sound they make is a repeated “ah-oh,” which explains their Hawaiian name, ‘a‘o.

He ‘a‘o ka manu noho i ka lua, ‘a‘ole e loa‘a i ka lima ke nao aku. It is an ‘a‘o, a bird
that lives in a burrow and cannot be caught even when the arm is thrust into the hole.
Said of a person who is too smart to be caught.
(Pukui: 545-64)

Newell’s Shearwaters spend about six months over the eastern tropical Pacific, and then in April they return to their mountain nesting sites, which are frequently in areas with dense uluhe ferns (Dicranopteris linearis). There they dig new burrows (or fix up their old ones), and then nesting occurs. The sounds of the birds above the colonies at night have been described as similar to the sounds of crying babies, mules, or even ghoulish laughter.
Shearwaters are seabirds that skim close the surface of the ocean and then plunge into the water to catch fish or squid spotted from the air. The birds have what is called a tube nose at the base of their beak. The tube nose is connected to a gland that removes extra salt from the bird’s food and water. Shearwaters sometimes stay at sea for years, and their webbed feet allow them to kick off from the water’s surface.
The fish and squid that shearwaters feed on are often pushed toward the ocean’s surface by schools of tuna. Fishermen have long known that seeing a gathering of shearwaters may be a sign of the valuable tuna species the fishermen seek.
[Photograph: Newell’s shearwater]


Wedge-Tailed Shearwaters (Puffinus pacificus chlororhynchus)
Hawaiian name: ‘Ua‘u Kani
Indigenous (migratory).
[Illustration: Wedge-tailed shearwater]

Wedge-tailed shearwaters are the most commonly seen Hawaiian seabird offshore of the main Hawaiian Islands. Wedge-tailed shearwaters are about 18 inches (46 cm) long, with a wingspan of about 3 feet (91 cm) and a wedge-shaped tail. They are brownish-gray, have a pointed beak and webbed feet. Birds in the “light phase” (most Hawaiian shearwaters) are dark on top but whitish-colored underneath.
Wedge-tailed shearwaters nest in burrows on the ground. The burrows are small little caves about 2 to 3 feet (60 to 90 cm) long. The shearwaters begin preparing their nests in April and lay a single white egg in June. Each parent sits on the egg for about ten days at a time as the other parent feeds at sea. After about 52 days the chick hatches (usually in August), and both parents feed the nestling partly digested food (regurgitated squid and fish). About two weeks before the chick learns to fly, the parents leave the nesting island. The chicks must then survive on their own stored fat. The young bird soon learns to fly and then is able to get its own food.
Shearwaters are often called “the moaning birds” because they make strange wailing or crying sounds when they are settled in their colonies, particularly at dawn and dusk. The literal translation of the wedge-tailed shearwater’s name, ‘ua‘u kani, is “calling ‘ua‘u,” which describes the sound made by the bird. The shearwaters’ habit of gliding so close to the surface of the water that they appeared to slice or cut it caused English sailors to name the bird “wedge-tailed.”
Wedge-tailed shearwaters arrive at the Hawaiian Islands around March. Like the Laysan albatross, the wedge-tailed shearwaters return to the same location where they were born to lay their eggs. The chicks hatch in July or August, and after about 3½ months the chicks learn to fly.
By November the shearwaters migrate along the equatorial countercurrent to the coast of Central America, reaching such disparate locations as Panama and Japan and sometimes staying at sea for up to four years. Many of the birds migrate back to the Hawaiian Islands in March, repeating their migration path each year.

Shearwater Populations
Archaeological evidence shows that ancient Hawaiians utilized shearwaters and petrels as a food source. Abundant remains of shearwaters have been found in ancient Hawaiian settlements. Pigs and dogs brought by the early Polynesian settlers of the Hawaiian Islands also had an effect on the populations of the ground nesting shearwaters.
After Westerners arrived, cats and mongoose further decimated the shearwater population. By 1931 the birds were thought to be extinct in the Hawaiian Islands, but they were “rediscovered” in 1954.

Saving the Shearwaters
Every autumn many of the little fledgling Newell’s shearwater birds leaving their colonies and head for the sea where they will feed on their own for the first time. During this maiden journey, the birds often become disoriented by bright lights near roadways and other areas, causing them to become disoriented and land. This leaves the birds vulnerable to cars as well cats, dogs and other predators. These incidents are known as strandings.
On Kaua‘i, residents have been saving these stranded birds for more than 20 years. Most shearwater strandings occur in October and November. The peak period of the month for strandings is during the new moon when the sky is dark. Also vulnerable to strandings due to nighttime light are other night-flying seabirds, including the ‘ua‘u (Pterodroma phaeopygia sandwichensis, Hawaiian dark-rumped petrel) an endangered species.
The Kaua‘i Island Utility Cooperative has recently completed work on all of its 3,049 light poles on Kaua‘i, installing a housing that blocks the light from escaping upward. This was recommended by island’s Save Our Shearwater program. Other night-flying seabirds are also vulnerable to

Shearwater Aid Stations
On Kaua‘i, the public is encouraged to rescue stranded shearwaters and leave them in small, protected cages that are provided at special aid stations. These aid stations are located at Kaua‘i County Fire Departments as well as other locations.
The birds placed in the cages at the aid stations are picked up each morning by state and federal wildlife biologists. The birds are checked for injuries, banded, and then released along the shoreline so they may fly out to sea. Up to 2,000 Newell’s shearwaters (an estimated 90% of all strandings), are returned to safety each year, with more than 25,000 birds rescued to date. Hundreds of fledgling wedge-tailed shearwaters are also rescued.

Rescuing Shearwaters
Be careful when attempting to rescue stranded shearwaters, as the birds may occasionally bite. Often the disoriented birds will be found sitting on the ground, and it will not be too difficult of a task to relocate them to an aid station. Stay clear of the bird’s head as you grasp its folded wings and tail. Then put the bird in a ventilated box for transport.
The Department of Land and Natural Resources (DLNR) advises the public that stranded birds may have unseen injuries and should not be tossed up into the air in an attempt to allow them to fly. Local residents may assist in helping the native bird species survive by reducing unnecessary outdoor lights, especially floodlamps, which may cause the birds to land due to temporary blindness.
[Photograph: Newell’s shearwater in burrow]


Tropicbirds

White-Tailed Tropicbird(Phaethon lepturus dorotheae)
Hawaiian Name: Koa‘e Kea
Indigenous: Tropical Pacific.

Red-Tailed Tropicbird(Phaethon rubricauda rothschildi)
Hawaiian Name: Koa‘e ‘Ula
Indigenous: Tropical Pacific.
[Illustration: Red-tailed and White-tailed Tropicbirds: Comparison]

You will see them high in the sky, their long, thin bodies effortlessly soaring above the ocean or over mountain valleys. The red-tailed tropicbird and the white-tailed tropicbird are both almost all white in color, except the red-tailed tropicbird has a red bill and red tail feathers, while the white-tailed tropicbird has a yellow bill and white tail feathers.
At 36 to 42 inches (7.6 to 107 cm) long, the red-tailed tropicbird is a bit larger than the white-tailed tropicbird, which is 23 to 32 inches (58 to 81 cm) long. Both types of tropicbirds plunge dive into the water to catch fish and squid, and both birds may give a loud scream-like sound while in flight.
[Illustration: Tropicbirds soaring.]

Ke koa‘e iho ia, he manu lele no ka pali kahakō.
That is the tropic bird, one that flies at the sheer cliffs.
Said of a person who is hard to catch.
(Pukui: 1749-188)

The red-tailed tropicbird has a 44-inch (112-cm) wingspan, compared to the 36-inch (91-cm) wingspan of the white-tailed tropicbird. Red-tailed tropicbirds also have two long, red tail feathers as well as black feathers around their eyes. White-tailed tropicbirds have black eye stripes black bars across their wings and back, and long, white tail feathers. The distinctive long tail feathers of the tropicbirds, also called streamers, are actually highly specialized inner tail feathers.
Extremely graceful in flight, red-tailed tropicbirds are awkward on land, and barely able to walk without falling. This is due to the their fully webbed feet. Tropicbirds plunge dive into the ocean to catch fish and squid, sometimes going as deep as 10 feet (3 m).
Feathers from both the red-tailed and white-tailed tropicbirds were utilized in ancient Hawaiian featherwork. The feathers were utilized to make royal feather standards called kāhili, consisting of feather clusters attached to long poles. Kāhili were symbols of chiefly rank. Especially valued for kāhili were the tail feathers of the red-tailed tropicbird.

Breeding
Tropicbird breeding extends from March to October. Red-tailed tropicbirds are known for their elaborate displays of courtship during flight. The birds sometimes repeatedly circling each other in an upward and backwards flight motion. A bird engaged in this backward flight appears a bit like it is rowing a boat, and may be nearly stationary in wind during this elaborate courtship dance, which usually lasts less than ten seconds.
The red-tailed tropicbird lays its eggs on the ground, usually under a shrub, beach vegetation or a rock overhang. The white-tailed tropicbird may nest inland and along the coastline, and lays its egg on a crater wall or on a ledge on a steep cliff face. The chicks of both birds are fully feathered after about six weeks and by about two months of age they learn to fly.
Tropicbirds are commonly seen near Kīlauea National Wildlife Refuge on Kaua‘i’s north shore, and circling high above Kaua‘i’s Nāpali Coast and Waimea Canyon. The birds are relatively rare, however, on the other Hawaiian Islands. On the island of Hawai‘i, tropicbirds are seen at Hawai‘i Volcanoes National Park in Kīlauea Crater. On O‘ahu the birds are sometimes seen on the windward side of the Pali.
[Photograph: Side by side comparison photos of red-tailed tropicbird and white-tailed tropicbird]


Forest and Mountain Birds of the Hawaiian Islands

[Illustration: ‘I‘iwi]

He ali‘i ka manu.
A bird is a chief.
A bird flies and perches higher than any human.
(Pukui: 534-63)


Hawaiian Owl Pueo
Hawaiian Hawk ‘Io
Hawaiian Crow ‘Alalā
Hawaiian Goose Nēnē
Honeycreepers and Honeyeaters
(and other small forest birds)
Original Colonizing Species
Populations and Extinctions
Hawaiian Featherwork
Hawaiian Honeycreepers;
Evolutionary Processes / Adaptive Radiation
‘Akialoa and ‘I‘iwi;
The ‘Elepaio
The Puaiohi—Captive Breeding

Threats to Endemic Forest Birds





The Hawaiian Owl(Asio flammeus sandwichensis)
Hawaiian Name: Pueo
Also called: Short-Eared owl.
Endemic
Status: Species of Concern (federal listing: actually a subspecies). Endangered on O‘ahu (state listing).
Considered by Hawaiians to be an ‘aumakua, or guardian spirit, the pueo is about 15 inches (38 cm) long with yellow eyes, a round facial disc, a black bill, and brown and white feathers. The pueo is an endemic Hawaiian subspecies of the short-eared owl. The pueo’s call is a repeated "tchak" sound, though the bird is usually silent.
The pueo’s habitat includes lowland pastures and fields as well as higher forest areas at elevations up to 8,000 feet (2,438 m). Pueo often hunt over grassland areas, mostly at dawn and dusk but also during daylight hours. They eat insects and rodents (e.g., mice) as well as other birds including both native and introduced species. The pueo may also prey upon the native ‘ōpe‘ape‘a (Lasiurus cinereus semotus, Hawaiian hoary bat).
Pueo often hover briefly over their prey before diving down for the attack. Unlike most other owls, the pueo often soars high in the sky, and is diurnal (active during the day).
[Illustration: Pueo]

Malu ke kula, ‘a‘ohe ke‘u pueo.
The plain is quiet; not even the hoot of an owl is heard.
All is at peace.
(Pukui: 2130-232)

There is very little research and documentation of pueo nesting behavior. Pueos are thought to nest throughout the year in the Hawaiian Islands, and the peak breeding months are March through June, the same as most other Hawaiian forest bird species.
During courtship, the pueo gives a series of low hoots and claps its wings. The owl lays usually from three to six white eggs in a nest built on the ground in a grassy area. Nests are built on the ground, but rarely seen. One researcher discovered a pueo nest of six eggs in a depression in tall grass on the downhill side of a dead snag of māmane (Sophora chrysophylla).

Pueo Population
No pueo fossils have been found dating to the time before the first Polynesian settlers arrived in the Hawaiian Islands. Pueo’s need open grassland habitats, and researchers theorize that the pueo may not have become established on the Hawaiian Islands until after the Polynesian settlers arrived and altered the landscape, increasing the amount of grassland and creating other suitable habitat that then made pueo colonization attempts successful.
The pueo is found on all of the main Hawaiian Islands, with the largest populations found on Maui, Kaua‘i and the island of Hawai‘i. The O‘ahu pueo population is listed as endangered by the State of Hawai‘i. The pueo’s roundish facial area helps to distinguish it from the non-native barn owl, which is also found in the Hawaiian Islands.
The legend of Pu‘upueo (“Owl hill”) in Mānoa, Honolulu tells of a beautiful woman named Kahala-o-Puna. All three times that Kahala-o-Puna was killed by her jealous husband she was resuscitated by the owl that lived on the hill, which is now known as Pu‘upueo.
[Photograph: Pueo]


The Hawaiian Hawk (Buteo solitarius)
Hawaiian Name: ‘Io
Endemic
Status: Endangered Species. Found on island of Hawai‘i Island only.
A sacred hawk in Hawaiian mythology, the ‘io is about 17 inches (43 cm) long, with the female on average a little bigger than the male. “Dark-phase” birds are dark brown above and below, while “light-phase” birds are dark on top but lighter colored below with dark streaks. The immature ‘io has a light-colored head that darkens as the bird becomes an adult. The legs also go from a green to a more yellowish color.
[Illustration: ‘Io]

He ‘io au, he manu i ka lewa lani.
I am an ‘io, the bird that soars in the heavenly space.
A boast. The highest chiefs were often called ‘io (hawk),
king of the Hawaiian birds.
(Pukui: 639-72)

In air, the ‘io soars gracefully, searching for insects, rodents, and other birds (including young game birds). The ‘io is also a natural predator of the native ‘alalā (Asio flammeus sandwichensis, the native Hawaiian crow) and the native ‘ōpe‘ape‘a (Lasiurus cinereus semotus, Hawaiian hoary bat). The ‘io is often are seen soaring high in the air in large circles.
A vocal hawk the ‘io makes the most sound during the breeding season. The ‘io’s vocalization is a high-pitched “kee-oh” sound. Little is known about breeding and nesting behavior, though most nesting is through to occur from May to October.
The hawk builds a nest in a tree using twigs and leaves, and then lays usually from one to three light blue eggs. Both parents participate in the feeding and aggressive protection of the fledglings.

‘Io Population
The ‘io was listed as endangered in 1967, and in 1982 the ‘io population was estimated to be between 1,400 and 2,500 hawks. The ‘io population likely remains near that number today. Fortunately, the ‘io population is spread through a wide range of habitats and elevations, making it less vulnerable to sudden declines in population. On Hawai‘i Island the hawk may be seen over the slopes of Mauna Kea and Kohala Mountains as well as Mauna Loa.

The ‘Io in Ancient Hawai‘i
‘Iolani was a sacred hawk of Hawaiian mythology, and also one of the names of King Kamehameha IV (Alexander Liholiho ‘Iolani). ‘Iolani means “Hawk of heaven,” or “Royal hawk,” and the flight of the ‘io was believed to be a sign of royalty.
In December of 1879, the cornerstone was laid for ‘Iolani Palace in midtown Honolulu. A project of King Kalākaua, ‘Iolani Palace was built on the site of the earlier royal palace, called Hale Ali‘i. Hale Ali‘i was built in 1845 for King Kamehameha III (Kauikeaouli), and named ‘Iolani in 1863 at the request of King Kamehameha V (Lot Kapuāiwa Kamehameha) who wanted a name chosen to honor his deceased brother, the former King Kamehameha IV (Alexander Liholiho ‘Iolani). ‘Iolani Palace was the seat of the Hawaiian monarchy for King Kalākaua and later for Queen Lili‘uokalani. (See O‘ahu, Chapter 2; Chapter 11, Timeline: 1895, Jan. 6; 1879, December 31.)
[Photograph: ‘Io]


The Hawaiian Crow(Corvus hawaiiensis)
Hawaiian Name: ‘Alalā
Endemic
Status: Endangered Species. Found on Hawai‘i Island only, and in propagation.
The ‘alalā is about 19 inches (48 cm) long, with a black beak and legs, and dark brown and black feathers. The Hawaiian crow’s color is a duller black than its North American relatives. The crow also has tinges of brown on its wings, and hair-like throat feathers that are stiff and grayish.
The ‘alalā often makes a “cawak” sound as well as other loud vocalizations, especially during the breeding season. The word ‘alalā comes from ala (to rise up) and lā (the sun), and refers to the crow’s habit of filling the forest with its raucous sounds early in the morning.
[Illustration: ‘Alalā]

He ‘alalā, he manu leo nui.
It is the crow, a loud-voiced bird.
Said of a person who talks too loud.
(Pukui: 527-62)

The ‘alalā is an omnivorous bird that feeds on fruit and nectar as well as insects and carrion. Its diet has changed due to changes in its habitat, but the original populations of ‘alalā fed primarily on native berries, such as ‘ōhelo (Vaccinium reticulatum) as well as the fruit and flower of the ‘ie‘ie vine (Freycinetia arborea). ‘Alalā may also eat mice.
In spring (from April to June) the ‘alalā builds a nest using twigs and grasses, and breeding occurs from March to July. The ‘alalā lays up to five greenish-blue eggs, though it’s likely only two will survive. Fledglings may remain with their parents for up to one year.

‘Alalā Population
The ‘io (Buteo solitarius,Hawaiian hawk) is a natural predator of the ‘alalā, but this was not a significant factor in the precipitous decline of the species over the last century, though it has been a factor more recently (see below). The primary causes of the ‘alalā’s historic population decline include the intentional killing of the birds by humans as well as loss of habitat.
Farmers began killing the ‘alalā in the 1890s because the birds would get into the poultry yards and feed pens. The ‘alalā population began seriously declining in the mid-1900s due to avian diseases (e.g., avian malaria, avian pox), predation by feral mammals, shootings by farmers, and cattle grazing on land that had previously been ‘alalā habitat. Another avian disease affecting ‘alalā is taxoplasma, which is transported to the birds through the droppings of feral cats.
In 1967, the ‘alalā was listed as a federally endangered species. In 1976, a captive rearing program at Pōhakuloa on the island of Hawai‘i attempted to raise ‘alalā, but the effort was unsuccessful. By 1978 less than 100 ‘alalā were left.
In 1981, when the wild ‘alalā population was estimated at somewhere between 30 and 150 birds, unsuccessful attempts were made to breed ‘alalā at Pōhakuloa on Hawai‘i Island. Then three ‘alalā eggs from captive ‘alalā nests were artificially incubated and hatched at the Honolulu Zoo.
In 1987, the ‘alalā was on the edge of extinction with just one bird in North Kona and several more in central Kona. A 1991 National Audubon Society lawsuit against the United States Fish & Wildlife Service and McCandless Ranch initiated the capture of some of the last wild ‘alalā as part of a recovery plan.
In 1992, the wild population of ‘alalā was believed to be just 12 birds as work to save the bird species from extinction was undertaken by the Peregrine Fund, under contract by the U. S. Fish & Wildlife Service. Eggs were taken from Kealia Ranch, Kai Malino Ranch, and McCandless Ranch, and ‘alalā chicks were raised at captive propagation facilities at Olinda on Maui (Maui Bird Conservation Center) and at Volcano on Hawai‘i Island (Keauhou Bird Conservation Center). Both facilities are operated by the Zoological Society of San Diego.
In 1993, seven ‘alalā were born, five of which were released into the wild in their native habitat on the island of Hawai‘i. In 1995, seven more chicks were released on Hawai‘i Island. By 1997, four more ‘alalā had been released into the wild, and the number of ‘alalā in captivity reached 40.
In 1997, when the wild ‘alalā population was estimated to be just 15 birds, the species suffered a severe setback when five captive-released ‘alalā were killed by ‘io (Buteo solitarius, Hawaiian hawks). It is possible that some of the ‘alalā preyed upon by ‘io had been weakened by taxoplasma.
In response to the multiple ‘alalā deaths, the U. S. Fish & Wildlife Service began trapping the ‘io (also an endangered species), and relocating the hawks to Volcano, but they quickly returned to the McCandless Ranch in South Kona. The Wildlife Service then caught ten ‘alalā and shipped them to zoos on the United States Mainland as part of a captive breeding program. Five of these ‘alalā later died.
By 1999, releases of captive-bred ‘alalā were halted due to the high death rate of released birds. As of the year 2000, there were 33 ‘alalā, with 30 in captivity.
By the beginning of 2004, there were 40 ‘alalā in captivity. No ‘alalā had been seen in the wild since June of 2002, when one bird was sited at Kealia Ranch in South Kona, though several persons claimed they had heard ‘alalā. If the species is re-established in the wild, it is likely that some “cultural” knowledge among the species will be lost, as the young birds will have not have been raised by their parents in the wild, when much of the generational information is typically learned.
The primary remaining habitat for ‘alalā is found in south Kona on the island of Hawai‘i. This mixed forest of koa (Acacia koa) and ‘ōhi‘a lehua (Metrosideros species) is on the 5,300-acre (2,145-ha) Kai Malino Ranch and another 1,258 acres (509 ha) nearby. This unique leeward rainforest also supports other endangered bird species, including the ‘akia pōlā‘au (Hemignathus munroi), the Hawai‘i creeper (Oreomystis mana), and the ‘io(Buteo solitarius, Hawaiian hawk). Another native species seen in the region is the ‘ōpe‘ape‘a (Lasiurus cinereus semotus, Hawaiian hoary bat).
Continued threats to the ‘alalā include loss of habitat due to the logging of koa, which is highly valued for fine furniture and other items. (See Koa in Native Plants and Ferns of the Hawaiian Islands, Chapter 8.) Another species of crow once lived on O‘ahu, and a separate species lived on Maui, but both are now extinct. Hope remains that the extremely endangered ‘alalā will not also suffer that fate.
[Photograph: ‘Alalā]


Nēnē—The Hawaiian Goose(Branta sandvicensis)
Endemic
Status: Endangered Species. Found on Maui, Kaua‘i, and Hawai‘i Island. Recent releases on Moloka‘i (see below).
Many thousands of years ago, some Canadian geese were blown off course or perhaps caught in a storm, and ended up in the Hawaiian Islands. Today their descendents have evolved into a unique Hawaiian bird known as the nēnē. This endemic goose species is now Hawai‘i’s official state bird.
[Illustration: Nēnē]

“Unele! Unele!” wahi a ka nēnē.
“Honk! Honk!” says the goose.
A play on nele (a lack, poverty), this saying implies a going without,
a lack of success, chagrin, and so forth.
(Pukui: 2878-314)

The nēnē is about 2 feet (61 cm) long, which is a typical size for a goose. The nēnē’s head, face, and the back of its neck (the nape) are black, and the cheeks and the sides of the neck are a light tan color, with a buffy striped pattern (distinct horizontal bands). The nēnē’s lower body has this same light brown color and is striped, but the top of the body is a darker gray or brown.
The nēnē’s bill is black, as are its legs and feet. The webbing between the toes on the nēnē’s feet is much reduced compared to the fuller webbing on the feet of its ancestor, the Canadian goose. This adaptation is better for walking on high, dry lava flows and other habitats frequented by the nēnē.
When nēnē fly they make a “ney ney” sound, but on the ground nēnē sometimes make a different noise, slightly similar to a cow’s moo. Nēnē aren’t very shy, and sometimes even approach humans.

Feeding and Habitat
Nēnē eat grasses, seeds, buds, flowers, berries and leaves, and are especially fond of native plants such as berries of naupaka (Scaevola species), ‘ōhelo (Vaccinium species), kūkaenēnē (Coprosma ernodeoides), pūkiawe (Styphelia tameiameiae) and ‘ulei (Osteomeles anthyllidifolia, Hawaiian hawthorn). Nēnē prefer high, dry areas like old lava flows as well as wetlands and forest uplands.

Nesting
Researchers previously believed that nēnē mate for life, though recently a nēnē at Maui’s Haleakalā National Park was seen to have three different mates in just a several month period. The first had been the nēnē’s mate for about ten years when another male chased it away. A few months later another male displaced that mate. In ancient Hawaiian lore, the promiscuity of the nēnē is implied in several native Hawaiian proverbs or sayings that link the birds’ amorous behavior to the behavior of certain men.
The nēnē builds its nest on the ground and lines it with feathers. This ground nesting is one reason nēnē became so endangered (ground predators include mongooses, pigs, rats, and domestic animals).
By the age of two, nēnē begin laying eggs. They nest between October and March, and one nēnē may lay from two to five (usually four or five) creamy, white eggs. They sit on the eggs for about 30 days while the eggs incubate. Sometimes the mother leaves the nest during this time and when she does, she covers the eggs with the downy feathers that are part of the nest lining.
During nesting, the adults go through a four to six week process called molting, at which time the adult birds cannot fly. The infant chick is able to run around just as soon as the chick’s downy feathers dry. The parents provide food for the baby until the hatchling is about 10 to 12 weeks old, when the gosling learns to fly.

Nēnē Population
Genetic research has revealed that there was a marked decline in the population of nēnē (and other bird species) between about A.D. 1100 A.D. and A.D. 1750. These population declines (and in some case extinctions) in native Hawaiian bird species in the centuries before Western contact with the Hawaiian Islands is generally attributed to Hawaiians hunting the birds for food.
Fossil records show that nēnē were once common at lower elevations on Kaua‘i, but then became extinct by the time Captain Cook arrived in 1778. Apparently the early Polynesian settlers had eliminated the nēnē on Kaua‘i, but the birds survived on other Hawaiian Islands. There were an estimated 25,000 nēnē living on the island of Hawai‘i in 1778. Predatory animals, habitat destruction, hunting, and egg collecting eventually decimated the nēnē populations of all the Hawaiian Islands.
The Maui nēnē population became extinct around 1890. By 1951, there were only about 30 nēnē left in the wild as well as some captive nēnē in European and American zoos. Nēnē were federally listed as endangered in 1967.
Beginning in 1949, scientists began raising nēnē in captivity, including more than 1,600 birds raised at Pōhakuloa on Hawai‘i Island. Almost all of these birds were released in Maui’s Haleakalā Crater and on Hawai‘i Island with the goal of re-establishing natural nēnē populations.
Beginning in 1951, nēnē were also raised at England’s Slimbridge Wildfowl Trust, and 200 of these nēnē were released on Maui in the 1960s (at least two were still alive in 1990). In all, more than 2,000 nēnē were raised in captivity between 1960 and 1990 and released on three of the Hawaiian Islands (Kaua‘i, Maui, and Hawai‘i Island). More than 80% of these nēnē were raised at the Endangered Species Facility in Olinda, Maui.
Nēnē restoration efforts have been a great success. The wild nēnē population increased to more than 1,300 birds statewide in 2004, though nēnē are still considered endangered. It’s estimated that the population atop Maui’s Haleakalā Volcano is about 250 nēnē, with about 60 more of the birds living in West Maui.
In addition to the wild nēnē population in the Hawaiian Islands, there are also up to 2,000 nēnē in captivity in U.S. and European collections and zoos.
The island of Kaua‘i is now considered the ideal place for the nēnē because there are no mongooses on Kaua‘i, as there are on the other main Hawaiian Islands. On Kaua‘i the population of nēnē has rapidly grown, and they are seen in various low elevation habitats (e.g., lower Hanalei Valley) as well as higher elevations (e.g., Kōke‘e State Park). On other Hawaiian Islands, however, nēnē populations are generally restricted to higher elevations.
Nēnē were first re-introduced to Moloka‘i in the early 1960s, with numerous bird releases occurring until 1978. In December of 2001, ten nēnē were released at Pu‘u O Hōkū Ranch in eastern Moloka‘i in the Cape Hālawa region. The birds were reared as part of a captive flock at the state’s Maui Bird Conservation Center at Olinda, Maui, which is operated by the Zoological Society of San Diego using funds provided by the federal government and the State of Hawai‘i. In 1995, a nēnē reintroduction program began at Hana‘ula in West Maui.
At least another 55 nēnē have been released on Moloka‘i, including eight nēnē released at Pu‘u O Hōkū Ranch in June of 2003. The nēnē releases at Pu‘u O Hōkū occurred as part of the State of Hawai‘i’s first “safe harbor” agreement in which the private landowner, in turn for providing habitat for the endangered birds, is given assurances that no further restrictions will be placed upon the land due to the endangered species living there. In the last decade nēnē have also been released in the West Maui Mountains and on Kaua‘i.

The Nēnē Count
For more than a decade an annual nēnē count has been conducted at Haleakalā National Park, where about 250 of the birds now exist. The survey is held in August, the time of year when single nēnē adults and family groups of nēnē flock before breeding season. Various agencies participate in the nēnē count, including the U.S. Fish and Wildlife Service, the Nature Conservancy, the University of Hawai‘i and others.
[Photograph: Nēnē]


Honeycreepers and Honeyeaters
and other small forest birds
[Illustration: Beaks of different honeycreepers]

The songs of small forest birds of the Hawaiian Islands echo through the high tropical forests. Some of these birds have long, thin beaks while others have short stout beaks. Some have beaks designed for probing bark and grasping insects, while others are better suited for eating fruit or sipping nectar. With colors are as varied as their beaks, the honeycreepers and honeyeaters evolved into many different species that eat just about every food the forest has to offer, from seeds to insects to fruit.
The Hawaiian Islands have an extremely high rate of endemic avifauna. In other words, many of Hawai‘i’s native birds are unique to the Hawaiian Islands, and are found nowhere else on Earth. The main reasons that Hawai‘i has so many endemic (unique) bird species are Hawai‘i’s geographic isolation as well as the abundance of ideal bird habitat throughout the Hawaiian Islands.

Original Colonizing Species
Millions of years ago, several families of birds made the long journey to the Hawaiian Islands. Perhaps they were blown off course in a storm, or otherwise lost their way and ended up on the remote archipelago. Those first colonizing bird populations (consisting of probably very few birds) took advantage of the lack of predators in the Hawaiian Islands, and thrived in their new habitats. Eventually the birds multiplied, and over many generations established significant populations throughout the Hawaiian Islands.
Groups of the growing bird populations utilized different food sources and ecological opportunities (niches). Eventually the populations became genetically isolated from one another and evolved into many distinct species. When one species evolves into many species it is known as adaptive radiation, or evolutionary divergence. In all, about 15 to 20 original colonizing bird populations (from at least 11 different bird families) evolved into well over 100 different bird species in the Hawaiian Islands.

Populations and Extinctions
When the first Polynesians settled in the Hawaiian Islands they immediately began to have an impact on native bird populations. For example, while nēnē (Branta sandvicensis) had once thrived on Kaua‘i, it is believed the bird was extinct on Kaua‘i by the time Captain Cook arrived in 1778, though nēnē populations on other islands, particularly Hawai‘i Island, were still significant.
When Captain Cook arrived in the Hawaiian Islands in 1778, there were about 80 species of endemic Hawaiian landbirds and at least 24 species and subspecies of native Hawaiian seabirds (4 endemic) and at least 34 species and two subspecies of waterbirds (29 endemic). Before Cook arrived, more than 35 endemic Hawaiian landbirds had already gone extinct.
Bird species that went extinct prior to Cook’s arrival met their demise from a variety of causes, including humans hunting the birds for food, feathers, and other reasons (e.g., collectors capturing the birds). Pigs and dogs brought by the Polynesian settlers of the Hawaiian Islands also had an effect on island bird populations, as did the rats that hitched a ride on the Polynesians’ voyaging canoes. These particular Polynesian-introduced species were detrimental to ground-nesting birds, which were also the most likely birds to be hunted by the early settlers for food.
About 23 of the bird species that became extinct prior to Western contact were ground-nesting birds, now known only from subfossil discoveries. (For a more in-depth discussion of extinct Hawaiian birds, see Overview of Native and Polynesian-Introduced Species of the Hawaiian Islands, Chapter 5.)

Hawaiian Featherwork
Another factor contributing to bird extinctions in the Hawaiian Islands before European contact was the elaborate featherwork that Hawaiian ali‘i (royalty) fashioned from the feathers of various birds, particularly small endemic forest birds such as the ‘ō‘ō (Moho species). The featherwork produced by the Hawaiians is considered among the finest in all of Polynesia. This is a result of the Hawaiians’ skillful craftsmanship as well as a reflection of the array of colorful bird species found on the Hawaiian Islands.
Hawaiians crafted various items using feathers, including ‘ahu ‘ula, the long royal capes and cloaks worn by ali‘i. Also fashioned from feathers were mahiole, the ceremonial warrior helmets bearing a resemblance to Homeric Greek military helmets. The coarser seabird feathers (e.g., tail feathers of tropicbirds) were used to make kāhili, the royal feather standards that were symbols of chiefly rank. Feathers were also used to make lei hulu (feather lei), which were worn around the head or neck, or untied around the shoulders.
Tropicbirds (koa‘e), frigatebirds (‘iwa), and various other species provided feathers for Hawaiian featherwork, but the most valued plumage was that of the small endemic forest birds. The scarlet feathers of the ‘i‘iwi (Vestiaria coccinea)and the green feathers of the ‘ō‘ū (Psittirostra psittacea) were often used. Above all other feathers, plumage hunters valued the yellow feathers of the ‘ō‘ō (Moho species) and mamo (Drepanis pacifica), two predominantly black birds with tufts of yellow feathers near their wings and tail.
The mamo, which went extinct in the early 1900s, was a mostly black bird with yellow feathers above and below its tail. One royal yellow cape made for King Kamehameha I was said to have required the feathers of about 80,000 birds (mostly feathers of the mamo).
The prized yellow thigh feathers of the ‘ō‘ō (Moho species) were a major factor in that species extinction, island by island. At one point, ‘ō‘ō feathers were even considered more valuable than the lei niho palaoa (whale tooth pendant) made with the teeth of the palaoa (Physeter macrocephalus, sperm whale) and highly prized by ali‘i (royalty). (See Lei Niho Palaoa in Island Flowers and Lei section, Chapter 3.)
A unique variety of ‘ō‘ō on Kaua‘i was referred to as the ‘ō‘ō ‘ā‘ā (Moho braccatus). Of the four species of ‘ō‘ō in the Hawaiian Islands, the Kaua‘i species likely survived the longest because it lacked the bright yellow wing feathers of the other ‘ō‘ō species and these yellow feathers were desired for use in featherwork. The last sighting of the ‘ō‘ō‘ā‘ā occurred in Kaua‘i’s Alaka‘i Swamp in 1987. With a variable and echoing song, the bird was known as one of the finest singers of all the Hawaiian birds.
In ancient times, birds captured for Hawaiian featherwork were caught by a variety of methods, including using fine mesh nets made from olonā (Touchardia latifolia). The birds were also caught by smearing the gummy sap from certain plants onto a stick, and catching the birds when they alighted upon the sticky substance. Used for this purpose was the sap of ‘ulu (Artocarpus altilis, breadfruit), as well as pāpala kēpau (Pisonia species), ‘āla‘a (Pouteria sandwicensis) and other trees.
An ancient Hawaiian proverb states, “E hō‘ike mai ana ka lā‘au a ke kia manu,” (“The stick of the birdcatcher will tell”), meaning “...we will know how successful one is by what he produces. One knew whether a birdcatcher was successful by counting the birds on his gummed stick.”[vi]

Hawaiian Honeycreepers
The Hawaiian Islands are considered by scientists to be one of the world’s most fascinating natural laboratories of genetics and biology. A prominent example: about 50 unique species or subspecies of Hawaiian honeycreeper birds evolved from a single colonizing population of finches. Many of these bird species are now extinct. As of 1995, only 21 honeycreeper species or subspecies remained, and 14 of those were listed as endangered.
Researchers theorize that the original colonizing finches first took hold on the Hawaiian archipelago sometime between 15 and 30 million years ago. At that time, the eight main Hawaiian Islands had not yet come into existence. Kaua‘i is only about five million years old, and the other main Hawaiian Islands are even younger.
The basaltic rocks of Midway Atoll, 1,580 miles (2,543 km) northwest of the island of Hawai‘i and 1,309 miles (2,107 km) northwest of Honolulu, have been dated to nearly 28 million years old.
The original colonizing finch birds inhabited the islands that were the predecessors to today’s Hawaiian Islands. These ancient islands have now been carried northwest by the Pacific Plate and eroded away.
Once a bird species took hold on the emerging island chain, it then populated each successive island over time. This trans-migration between geologically changing habitat was occurring even as the birds themselves continued to evolve into a multitude of new and different species.

Evolutionary Processes / Adaptive Radiation
Honeycreepers are one of the most studied examples of adaptive radiation in the field of evolutionary biology. After the original colonizing population of finches arrived, they began to increase in numbers and spread throughout the Hawaiian Islands. They inhabited different areas and utilized different food sources, each sub-population taking advantage of different ecological opportunities (niches).
A great diversity of beak shape, tongue morphology, and feather color evolved in the different bird populations. The birds also developed a wide variety of songs and behaviors.
Natural selection favors physical traits that are beneficial to survival in the particular environment in which the animal lives. When different sub-groups of the original finch population became reproductively isolated from each other, they eventually began to possess very different physical traits from one another.
Some bird species developed powerful, seed-eating bills that were thick, short and efficient for cracking seeds. Other species evolved more delicate, pointed beaks that were well-adapted for catching insects or sipping nectar. Still others evolved crossed beaks good for probing bark, pecking at woody stems and twigs or opening leafbuds.
The honeycreeper family is a remarkable and world-renowned example of the evolutionary process of adaptive radiation whereby one species evolves into many distinct species. In the case of the honeycreepers, an ancestral population of birds became isolated in various Hawaiian habitats, and then over thousands of generations, evolved into about 50 species and subspecies of unique birds.
One of the most unusual Hawaiian honeycreepers to evolve was the little olive and yellow-colored bird called ‘akia pōlā‘au (Hemignathus munroi), which is federally listed as an endangered species. The ‘akia pōlā‘au developed a small, stout bill on the bottom for pecking, but a long curved, hooked bill on top, which it uses to pry open bark as it explores the trunks of trees for larvae.
[Illustration or Photograph: ‘Akia pōlā‘au]

‘Akialoa and ‘I‘iwi
Some bird species evolved beaks ideal for extracting the nectar from flowers. Two birds with specialized bills for sipping nectar are the ‘akialoa (Hemignathus species) and ‘the ‘i‘iwi (Vestiaria coccinea).
The ‘akialoa is a small olive or yellow-green bird with a long, curved bill. Different subspecies of the ‘akialoa once inhabited different Hawaiian Islands, but today the only one that is not extinct is the Kaua‘i ‘akialoa (Hemignathus procerus), which also has the longest, curved beak. Federally listed as an endangered species, the ‘akialoa uses its long beak to sip honey from the base of leaves of the hala pepe (Pleomele species) and ‘ie‘ie (Freycinetia arborea).
The ‘i‘iwi is a small, vermilion (red) bird with black wings and tail, and orange legs. The ‘i‘iwi has a pronounced salmon-colored bill that is unusually long and curves downward, and is among the most spectacular of all the honeycreeper bills. Researchers believe that the ‘i‘iwi evolved a long, down-curving bill because it fit perfectly into the deep corolla (the petal array) of certain native lobelias (Lobelioideae) with long, tubular-shaped flowers. (See Introduction to Native Plants and Ferns of the Hawaiian Islands, Chapter 8.)
These lobelia flowers likely evolved their long, curved corolla at the same time that the ‘i‘iwi evolved a long and curved bill to fit into the flower. Both species benefited, with the flower providing food for the bird, and the bird pollinating the flower (the stamen delivers pollen on the bird’s forehead). Eventually only the ‘i‘iwi had the proper beak to gain access to the nectar of this particular long-throated flower, aiding the survival of the flower species as well as the bird.
When two species (in this case a bird and a flower) evolve together to the benefit of each, it is known as co-evolution. The two species are also said to have a symbiotic relationship, with both species benefiting from the other.
[Photograph: ‘Akialoa]

Today many of the native lobelioid flower species are very rare or extinct, and the i’iwi has adapted to this change in food supply by feeding on the blossoms of the ‘ōhi‘a lehua tree (Metrosideros species). The ‘ōhi‘a blossom has no petals or corolla, so from an evolutionary standpoint the ‘i‘iwi’s long beak is no longer necessary for feeding.
Researchers have noted that, as evolutionary theory would predict, the ‘i‘iwi’s beak now seems to be evolving shorter again over time. A comparison of current ‘i‘iwi beaks with museum species from 1902 shows that ‘i‘iwi beaks have indeed become 3% shorter. The bird is apparently undergoing a sort of reverse evolution, in which a trait (a long, decurved beak) that formerly evolved because it was beneficial is now disappearing because it has no particular benefit.
[Illustration or Photograph: ‘I‘iwi feeding on lobelia.]

One of the most commonly seen of the honeycreepers is the ‘apapane (Himatione sanguinea), which survives on all of the main Hawaiian Islands. Its deep crimson color is very similar to the color of the ‘ōhi‘a lehua (Metrosideros species) blossom upon which the bird feeds.
[Photograph: ‘Apapane feeding on ‘ōhi‘a lehua blossom.]

Many Hawaiian species exist in just one location, making them extremely vulnerable to extinction and dependent on the last remote pockets of native habitat. For example, the endangered crested honeycreeper (Palmeria dolei) lives only on the upper eastern slopes of Maui’s Haleakalā Volcano in forests of ‘ōhi‘a lehua (Metrosideros species) above 4,500 feet (1,372 m).

The ‘Elepaio
The ‘elepaio (Chasiempis sandwichensis) is a small bird with a gray back, blackish tail, and white rump,was also able to adapt to the changing environment of the Hawaiian Islands. Not a honeycreeper, the ‘elepaio is a member of the flycatcher family (Muscicapidae). The bird’s name comes from its song, which sounds like e-le-pa-i-o, and is also described as a squeaky "wolf-whistle.”
[Photograph: ‘Elepaio]

‘Elepaio feed on insects and weave small nests of roots and grass. They lay usually two or three eggs, and the hatchlings fledge (learn to fly) in about two weeks. Variety has evolved in the ‘elepaio’s different populations, which look different on each Hawaiian Island.
Canoe makers of ancient Hawai‘i considered the ‘elepaio as their friendly guardian spirit. The bird was said to guide canoe builders to the best koa trees (Acacia koa) for use as hulls for their voyaging canoes.
If an ‘elepaio landed in a tree and spent some time pecking at it, then the tree was considered unsuitable for canoe building. However, if the bird landed on the tree and then began singing that was a good omen, and meant the tree would be a good one to use for building a canoe. An ancient proverb states, “Ua ‘elepaio ‘ia ka wa‘a.” (“The ‘elepaio has [marked] the canoe [log]”), meaning “...there is an indication of failure. Canoe makers of old watched the movements of the ‘elepaio bird whenever a koa tree was hewed down to be made into a canoe. Should the bird peck at the wood, it was useless to work on that log, for it would not prove seaworthy.”[vii]
One of the most recent additions to the federal endangered species list is the O‘ahu ‘elepaio (Chasiempis sandwichensis ibidis). The bird inhabits less than 5% of its former range, which has declined nearly 90% since 1940. Sightings have dropped precipitously since 1960. The O‘ahu ‘elepaio is now found in just seven small populations near the Wai‘anae and Ko‘olau Mountains, with an estimated total population of between 200 and 500 birds.

The Puaiohi—Captive Breeding
Many of native Hawaiian bird species are endemic to just one island. For example, in the Alaka‘i Swamp lives the puaiohi (Myadestes palmeri), a little pink-legged bird found nowhere else on Earth. Also called the small Kaua‘i thrush, the puaiohi is found only in the uplands of the Garden Island.
The puaiohi has an olive-brown back and a gray belly, and likes to nest in holes on cliffs. Puaiohi feed on native fruits, such as ‘ōlapa (Cheirodendron species) and pilo (Capparis sandwichiana), as well as insects.
[Photograph or Illustration: Puaiohi]

An extremely endangered species, the puaiohi’s population was at one point believed to be less than 30 birds. Then in 1995 scientists learned more about their habitat and discovered more of these rare birds. The population is now estimated to be about 250.
Some puaiohi are being raised in captivity at the Keauhou Bird Conservation Center on the island of Hawai‘i, a project sponsored by the Peregrine Fund, and the Zoological Society of San Diego with the approval of the state Board of Land and Natural Resources. At least 47 captive-raised birds have been released in the wild and have produced many new nests and fledglings.
The San Diego group also manages the endangered bird facility at Olinda, Maui. The Maui facility is engaged in captive propagation of nēnē (Branta sandvicensis, Hawaiian goose), ‘alalā (Corvus hawaiiensis, Hawaiian crow), and other species. Researchers plan on using the propagation techniques that were successful with the puaiohi to breed the endangered Maui parrotbill (Pseudonestor xanthophrys) and the Big Island palila (Loxioides bailleui).
Released birds are fitted with transmitters to gain more information that helps in the effort to prevent the extinction of the species. For example, a puaiohi nest observed in 1999 near a streamside cliff on a fallen log was determined to be the nest of two captive-bred puaiohi that had earlier been released in the wild.

Threats to Endemic Forest Birds
Avian malaria and avian pox are two of the primary causes of the endangerment and extinctions of small endemic Hawaiian forest birds. Other major factors affecting forest bird populations include habitat destruction resulting from agriculture, ranching, and urban development, as well as predation by mongoose and domestic animals (cats and dogs). Cattle, deer, sheep, goats, and pigs have destroyed large amounts of native Hawaiian habitat. Introduced bird species also compete with native bird species for habitat and food sources.
In the past, human hunting has also played a major role in reducing native bird populations, including the extinctions of several flightless bird species that were apparently captured by Hawaiians who utilized the ground-nesting birds for food and feathers before Captain Cook arrived in the Hawaiian Islands in 1778 establishing Western contact. Archaeological evidence has revealed that ancient Hawaiians also utilized various other bird species for food and/or feathers, including shearwaters and petrels. (See Overview of Native and Polynesian-Introduced Species of the Hawaiian Islands, Chapter 5.)



Chapter 7 Source Notes

All Hawaiian words in this text, the Hawaiian Encyclopedia, include proper diacritical marks, including the ‘okina (hamzah, or glottal stop) and kahakō (macron), except when the word is cited directly as part of a title or in a direct quote, in which case the words are spelled as originally published.
See Appendix 1: Note On Sources for more information about diacritical marks; objectivity; Hawaiian word spellings and meanings; scientific, cultural, and historic information; new research and discoveries; verifying dates and details of historic events; and Hawaiian sovereignty and other native issues.

[Note: This list is still in progress and being updated.]
A case of bill overkill. Discover Magazine, May, 1996.
Aikin, Ross R. Kilauea Point Lighthouse: The Landfall Beacon on the Orient Run. Kilauea Point Natural History Association. Lihue: Island Printers, 1988.
All About Hawaii (1948-1949): Thrum’s Hawaiian Annual and Standard Guide. Publications Division: Honolulu Star-Bulletin, Honolulu, Hawaii, 1949.
Ariyoshi, Rita. The National Geographic Traveler: Hawaii. Washington, D.C.: National Geographic Society, 2000.
Atlas of Hawai‘i: Second Edition. Honolulu: University of Hawai‘i Press, 1983.
Atlas of Hawai‘i: Third Edition. Honolulu: University of Hawai‘i Press, 1998.
Beletsky, Les. Hawaii: The Ecotravellers’ Wildlife Guide. London: Academic Press, 2000.
Berger, Andrew J. Bird Life in Hawaii. Honolulu: Island Heritage Limited, 1983.
Berger, Andrew J. Hawaiian Birdlife: Second Edition. Honolulu: The University Press of Hawaii, 1981.
Bishop Museum: Hawaii’s Extinct Species. Internet site: http://hbs.bishopmuseum.org/endangered/extinct.html, 7/28/2002.
Bishop Museum: Hawai‘i’s Extinct Species—Birds. Internet site: http://hbs.bishopmuseum.org/endangered/ext-birds.html, 7/28/2002.
Bishop Museum: The How Many Species Are there in Hawaii Web Page. Includes: How many species are there in Hawaii?, from the “Records of the Hawaii Biological Survey for 1994” as published in the Bishop Museum Occasional Paper volume 41: 3-18; and Number of Hawaiian Species: Supplement 1, originally published in Bishop Museum Occasional Papers 45: 8-17 (1996). Internet site: http://hbs.bishopmuseum.org/hispp.html, 10/17/2002.
Bishop Museum—The role of the Hawaii Biological Survey. Internet site: http://hbs.bishopmuseum.org/hbs3.html.
Boynton, David. Students taking part in Kolea Watch: Native bird ready for annual migration north. The Garden Island, 4/15/2002.
Bushnell, O.A., and Daws, Gaven; Illustrated by Feher, Joseph. The Illustrated Atlas of Hawaii. Aiea, Hawaii: Island Heritage Publishing, 1970.
Chang, Lester. It’s raining Newell shearwaters again: Annual falling—and rescuing—of the birds resumes. The Garden Island, 10/07/2000.
Clark, Hugh. Land deal creates habitat for endangered birds. The Honolulu Advertiser, 12/27/2001.
Conrow, Joan. Puaiohi thrives in the Alaka‘i Swamp. The Garden Island, 6/2/2000.
Creamer, Beverly. Plans set for rare birds to meet, mate: Biologists ready for high-tech effort on Maui. The Honolulu Advertiser, 3/15/2002.
Culliney, John L. Islands in a Far Sea. San Francisco: Sierra Club Books, 1988.
Daws, Gavan. Shoal of Time: A History of the Hawaiian Islands. Honolulu: University of Hawaii Press, 1968.
Dayton, Kevin. 2 endangered Hawaiian hawks returning to Islands. The Honolulu Advertiser, 12/14/2003.
Denny, Jim. The Birds of Kaua‘i. Honolulu: University of Hawai‘i Press, 1999.
Endangered animals in the Hawaiian Islands: Kaua‘i ‘O‘o. Internet site produced by the US Fish and Wildlife Service: http://www.r1.fws.gov/pacific/wesa/ookauai.html.
Eyre, David L. By Wind, By Wave. Honolulu: The Bess Press, 2000.
Federal Register, 2002, Vol. 67, U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS), Listings by State and Territory as of 7/20/2002. Internet site: http://ecos.fws.gov/servlet/TESSWebpageUsaLists?state=HI, 7/28/2002.
Friary, Ned, and Bendure, Glenda. Hawaii: 5th Edition. Victoria, Australia: Lonely Planet Publications Pty Ltd., 3/2000.
Gonser, James. Endangered bird may get extra federal protection. The Honolulu Advertiser, 12/18/2000.
Handy, Emory, Bryan, Buck, Wise and Others. Ancient Hawaiian Civilization: A Series of Lectures Delivered at the Kamehameha Schools. Mutual Publishing: Honolulu, Hawaii, 1999.
Hawaiian and Pacific Islands National Wildlife Refuges. United States Department of the Interior Fish and Wildlife Service, September, 1985.
Hawaiian Coot: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/coothi.html, 6/17/2000.
Hawaiian Crow: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/alala.html, 6/17/2000.
Hawaiian Duck: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/duckkoloa.html, 6/17/2000.
Hawaiian Goose: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/nene.html, 6/17/2000.
Hawaiian Hawk: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/io.html, 6/17/2000.
Hawaiian Moorhen: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/moorehenhi.html, 6/17/2000.
Hawaiian Stilt: Threatened and Endangered Species; Endangered Animals in the Hawaiian Islands. U.S. Fish and Wildlife Service. Internet site: http://www.r1.fws.gov/pacific/wesa/aeo.html, 6/17/2000.
Hawai‘i Data Book: A Statistical Reference to Hawai‘i’s Social, Economic and Political Trends. Edited by Robert C. Schmitt. Honolulu: Mutual Publishing, 2002.
Hawaii’s Birds. Honolulu: Hawaiian Audubon Society, 1993.
Hayworth, Philip. Hawai‘i’s Marine Zoos: The Waikīkī Aquarium, Sea Life Park and Maui Ocean Center showcase the incredible diversity of marine life flourishing in Hawaiian waters. Spirit of Aloha Magazine, May/June, 2002.
Hurley, Timothy. ‘Alala recovery left to captive methods. The Honolulu Advertiser, 12/19/2003.
Hurley, Timothy. Bird boom worries Kona airport: Stilt habitat raises concerns of flight hazards. The Honolulu Advertiser, 6/09/2002.
Hurley, Timothy. Nēnē mother may have been maliciously killed. The Honolulu Advertiser, 5/23/2002.
Hurley, Timothy. Society to manage Maui endangered bird facility. The Honolulu Advertiser, 1/1/2000.
Hurley, Timothy. State bird-nēnē-gets day of tribute: Hawaiian goose numbers 1,300. The Honolulu Advertiser, 9/25/2003.
Hurley, Timothy. State to release nēnē on Moloka‘i: Flock to be first on island in 100 years. The Honolulu Advertiser, 12/21/2001.
Hurley, Timothy. Tragic ending to a mother goose tale: Celebrity nēnē run over by car. The Honolulu Advertiser, 1/18/2002.
Ii, John Papa. Fragments of Hawaiian History. Translated by Mary Kawena Pukui; Edited by Dorothy B. Barrère. Honolulu: Bishop Museum Press, 1959.
Joesting, Edward. Kauai: The Separate Kingdom. Hawaii: University of Hawaii Press, 1984.
Kamakau, Samuel Manaiakalani. Ka Po‘e Kahiko: The People of Old. Translated from the Newspaper Ke Au ‘Oko‘a by Mary Kawena Pukui. Arranged and edited by Dorothy B. Barrère. Bishop Museum Special Publication 51. Honolulu: Bishop Museum Press, 1991.
Kāne, Herb Kawainui. Ancient Hawai‘i. Captain Cook, Hawai‘i: The Kawainui Press, 1997.
Kawaharada, Dennis. The Settlement of Polynesia, Part 1. Polynesian Voyaging Society. Internet Site: http://leahi.kcc.hawaii.edu/org/pvs/mirgrationspart1.html.
Kent, Harold Winfield. Treasury of Hawaiian Words in One Hundred and One Categories. Masonic Public Library of Hawai‘i. Honolulu: University of Hawaii Press, 1986.
Kilauea Point Natural History Association. Hawaii’s Wetlands. Kauai: Hawai‘i Nature Focus: Nature Studies for Children, # 11.
Kilauea Point Natural History Association. The Great Frigatebird: Iwa. Kauai: Hawai‘i Nature Focus: Nature Studies for Children, #7.
Kilauea Point Natural History Association. The Nene—The Hawaiian Goose. Kauai: Hawai’i Nature Focus, #14, 1993.
Kilauea Point Natural History Association. Wedge-tailed Shearwater. Kauai: Hawai‘i Nature Focus #2.
Kirch, Patrick Vinton. Feathered Gods and Fishhooks. Honolulu: University of Hawai‘i Press, 1985.
Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume One: Historical Ethnography (Marshall Sahlins, with the Assistance of Dorothy B. Barrère). Chicago and London: The University of Chicago Press, 1992.
Kirch, Patrick V., and Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawaii. Volume Two: The Archaeology of History (Patrick V. Kirch). Chicago and London: The University of Chicago Press, 1992.
KIUC finishes work to shield lights from endangered seabirds. The Garden Island, 6/13/2003.
Krauss, Bob, and Gleasner, Bill. Kauai. Aiea, Hawaii: Island Heritage Publishing, 1978.
Life in Early Hawai‘i: The Ahupua‘a—Third Edition. Honolulu: Kamehameha Schools Press, 1994.
Malo, David. Hawaiian Antiquities: Mo‘olelo Hawai‘i. Translated from the Hawaiian by Dr. Nathaniel B. Emerson, 1898. Bernice P. Bishop Museum Special Publication 2: Second Edition. Honolulu: Bishop Museum, 1951.
Manguchei, Kendyce. Poor man’s time machine uncovers Kaua‘i’s past at Maha‘ulepu. The Garden Island, 6/23/2002.
Maui ranch plans nēnē habitat: Deal with state to establish ‘safe harbor’ in Upcountry. The Honolulu Advertiser, 12/15/2003.
Merriam Webster’s Collegiate Dictionary: Tenth Edition. Springfield, Massachusetts: Merriam-Webster, Incorporated, 1996.
Mitchell, Donald D. Kilolani. Resource Units in Hawaiian Culture: Revised Edition. Honolulu: Kamehameha Schools Press, 1992.
Mullins, Joseph G. Hawaiian Journey. Honolulu: Mutual Publishing, 1978.
Nene goslings released in Na Pali Coast valley. The Kauai Times, 4/23/1995.
Oliver, Anthony Michael. Hawaii Fact and Reference Book. Mutual Publishing, 1995.
Oliver, Douglas. Polynesia in Early Historic Times. Honolulu, Hawai‘i: The Bess Press, 2002.
Peregrine helps our puaiohi. The Garden Island, 4/29/1996.
Pratt, H. Douglas. A Pocket Guide to Hawai‘i’s Birds. Honolulu: Mutual Publishing, 1996.
Pratt, H. Douglas. Enjoying Birds in Hawaii: A Birdfinding Guide to the Fiftieth State. Honolulu: Mutual Publishing, 1993.
Pukui, Mary Kawena, and Elbert, Samuel H. Hawaiian Dictionary: Revised and Enlarged Edition. Honolulu: University of Hawaii Press, 1986.
Pukui, Mary Kawena, and Elbert, Samuel H, with Mookini, Esther T. and Nishizawa, Yu Mapuana. New Pocket Hawaiian Dictionary. Honolulu: University of Hawaii Press, 1992.
Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983.
Pukui, Mary Kawena, and Elbert, Samuel H. & Mookini, Esther T. Place Names of Hawaii: Revised & Expanded Edition. Honolulu: University of Hawaii Press, 1974.
Rogers, Michael. Little Birds Lost. National Wildlife, October/November, 1993.
Ronck, Ronn. Hawaii Almanac. Honolulu: University of Hawaii Press, 1984.
Roth, Susan. Hawai‘i expert testifies for House bill on invasive species: Administration says effort not a priority. The Honolulu Advertiser, 3/15/2002.
Royte, Elizabeth. On the brink: Hawaii’s vanishing species. National Geographic Magazine, September, 1995.
Sahlins, Marshall. Anahulu: The Anthropology of History in the Kingdom of Hawai‘i. Chicago: The University of Chicago Press, 1992.
Scott, Susan. Nature Watch: Noddies. Hawaii Magazine, June 1996.
Scott, Susan. Plants and Animals of Hawai‘i. Honolulu: Bess Press, 1991.
Seven ‘alala chicks released into wild. ‘Elepaio: Journal of the Hawaiian Audubon Society, Volume 55, #1, February, 1995.
Shearwater safety goal of KIUC street light program. The Garden Island, 5/05/2003.
Snetsinger, Thomas J. Observations of Pueo nests on the slopes of Mauna Kea. 'Elepaio: Journal of the Hawaiian Audubon Society, February, 1995.
Soar with Kauai's albatross. Pacific Islands Ecoregion External Affairs Office, 1/12/1999.
Sprague, Brandon. In pursuit of Kauai’s dwindling native bird population. The Garden Island. (no date)
Status of O’ahu ‘Elepaio requires further study. ‘Elepaio, (Source: U.S. Fish and Wildlife Service): Journal of the Hawaiian Audubon Society, Volume 55, Number 6, August/September, 1995.
Taxonomy: Species Moorhen, Common, Hawaiian. Internet site: http://fwie.fw.vt.edu/www/esis/lists/e101014.htm, 3/13/1996.
TenBruggencate, Jan. Albatross back in islands. The Honolulu Advertiser, (no date).
TenBruggencate, Jan. Amazing kōlea faithfully return. The Honolulu Advertiser, 8/26/2002.
TenBruggencate, Jan. Dark evenings of autumn a risky time for fledglings. The Honolulu Advertiser, (no date).
TenBruggencate, Jan. Echoes of Iniki: Some victims still staggering. The Honolulu Advertiser, 9/5/1993.
TenBruggencate, Jan. Group sues for forest bird habitat. The Honolulu Advertiser, 1/4/2000.
TenBruggencate, Jan. Help keep watch on our kōlea. The Honolulu Advertiser, 3/25/2002.
TenBruggencate, Jan. Identify ‘elepaio habitat, court says. Honolulu Advertiser, 6/27/2000.
TenBruggencate, Jan. Kaua‘i limestone 315,000 years old. The Honolulu Advertiser, 2/19/2002.
TenBruggencate, Jan. Kauai thrush may be coming back. The Honolulu Advertiser, (no date).
TenBruggencate, Jan. Last habitat for crow endangered. The Honolulu Advertiser, 2/15/1996.
TenBruggencate, Jan. Loss of variability left nēnē vulnerable. The Honolulu Advertiser, 2002.
TenBruggencate, Jan. Muted lights brighten sea birds’ future. The Honolulu Advertiser, 6/05/2003.
TenBruggencate, Jan. Nēnē finds home on Moloka‘i. The Honolulu Advertiser, 1/07/2002.
TenBruggencate, Jan. Nene flourish on Kauai. The Honolulu Advertiser, 7/19/1995.
TenBruggencate, Jan. Rare birds socializing again at Midway. The Honolulu Advertiser, 1/15/2001.
TenBruggencate, Jan. Sinkhole yields treasure trove of bird fossils. The Honolulu Advertiser, 2/25/2002.
TenBruggencate, Jan. Songbird showing promise in wild: Efforts tested on Kaua‘i thrush. The Honolulu Advertiser, 2/22/2002.
TenBruggencate, Jan. Three rare puaiohi hatch in captivity. The Honolulu Advertiser, 5/1996, (uncertain date).
The Kumulipo: A Hawaiian Creation Chant. Translated and edited with commentary by Martha Warren Beckwith. Honolulu: University of Hawaii Press, Facsimile reproduction of first edition, University of Chicago Press, 1951. Including Foreword by Katharine Luomala, The University Press of Hawaii, 1972.
United States Fish & Wildlife Service: Pacific Islands—Endangered Species. Internet site: http://pacificislands.fws.gov/wesa/endspindex.html, 7/15/2002.
U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): Listings by State and Territory, as of 7/28/2002. Internet site: http://ecosfws.gov/webpage/webpage_usa_lists.html?state=all&module=undefined, 7/28/2002.
U.S. Fish & Wildlife Service, Threatened and Endangered Species System (TESS): U.S. Listed Vertebrate Animal Species Report by Taxonomic Group as of 7/28/2002. Internet site: http://ecos.fws.gov/webpage/webpage_vip_listed.html?module=undefined&code=V&listings=0, 7/28/2002.
Varez, Dietrich. Nene. Hawaii Magazine, April, 1996.
Viernes, Kathleen J.F. Bullfrog predation on an Endangered Common Moorhen chick at Hanalei National Wildlife Refuge, Kaua‘i, ‘Elepaio, Volume 55, Number 6, August/September, 1995.
Walther, Michael. Fading songs of the forest: Scientists work to save endangered birds. Hawaii Magazine, August, 1996.
Wetland Wildlife. Hawaii Magazine, December, 1996.
Where nēnē can wander again. The Honolulu Advertiser, 6/06/2003.
Wichman, Fredrick B. Kaua‘i: Ancient Place-Names and Their Stories. Honolulu: University of Hawai‘i Press, 1998.
Wildlife service launches plan to recover population of alala. The Garden Island, 12/19/2003.
Wilson, Christie. Haleakalā launches annual count of nēnē. The Honolulu Advertiser, 8/09/2001.
Ziegler, Marjorie. Ae‘o: Hawaiian Stilt. Hilo, Hawai‘i: Environment Hawai‘i, Volume 13, Number 2, August, 2002.


[1] p. 310, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 2830.
[2] p. 57, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 472.
[3] p. 194, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 1812.
[iv] Pukui, Mary Kawena, Elbert, Samuel H., and Mookini, Esther T. Mookini. Place Names of Hawaii: Revised & Expanded Edition. Honolulu: University of Hawaii Press, 1974.
[v] p. 262, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 2399.
[vi] p. 36, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 287.
[vii] p. 306, Pukui, Mary Kawena. ‘Ōlelo No‘eau: Hawaiian Proverbs & Poetical Sayings. Honolulu: Bishop Museum Press, 1983. Proverb 2777.