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Home , Black mamo, Ciridops, Common gallinule, Drepanis, Hawaiian rail, Hawaiian stilt

The Present Status of the of ' ANDREW J. BERGER2

THEGREAT EXPANSES of open ocean that sep­ waii, and in 1962 birds were first released in arate the from the major the Paliku Cabin area of Haleakala Crater on continental land masses of North America and . There is still debate as to whether or Asia resulted in the of a number of not the Nene originally inhabited Maui. unique landbirds. Unfortunately, a higher per­ The Nene was on the verge of in centage of of birds have become extinct the 1940s, and the species is still included in in Hawaii than in any other region of the the list of endangered species. In 1949 a Nene world. Approximately 40 percent of the en­ Restoration Program was begun by using a pair demic Hawaiian birds are believed to be ex­ of captive birds obtained from Herbert Ship­ tinct, and 25 of the 60 birds in the 1968 list man of Hawaii. This has been a very success­ of "Rare and Endangered Birds of the United ful program, and Nene have been raised in States" are Hawaiian CRare and Endangered captivity both at the Severn Wildfowl Trust Fish and Wildlife of the United States, 1968 at Slimbridge, England, and at the State of edition," Bureau of Sport Fisheries and Wild­ Hawaii Fish and Game rearing station at life, Washington, D. C.). Most of the native Pohakuloa on the Saddle Road of Hawaii birds of have long been extinct, and few (Elder, 1958) . native landbirds are to be found on any of the The program at Pohakuloa has been increas­ main islands below 3,000 feet elevation. ingly effective throughout the years, primarily . Three general groups of birds are found in through the dedicated efforts of Mr. Ah Fat Hawaii today: endemic, indigenous, and intro­ Lee. Over 500 Nene have been raised at Po­ duced. hakuloa during the period of 1949 through ENDEMIC HAWAIIAN BIRDS 1968. Most of these pen-reared birds have been released at several known of wild Ten families of birds are recognized as hav­ Nene on the slopes of Maun a Loa; a smaller ing endemic genera, species, or subspecies in number have been released in Haleakala Hawaii (although taxonomic dispute still ex­ Crater. ists regarding the relationship of some Ha­ The Nene is a highly specialized , waiian forms to closely related North American adapted for living in a rugged of lava forms); in addition, one entire famil y of birds flows far from any standing or running water (Drepanididae) is endemic to the Hawaiian (Miller, 1937). Among the more noticeable chain of islands (Amadon, 1942; Mayr, 1943). anatomical specializations for this terrestrial An "endemic" form is one that occurs in one life is a reduction in the webbing between the region only and is not found in any other toes. The birds spend much of the time on part of the world. Ornithologists believe that sparsely vegetated lava flows on Mauna Loa the ancestors of these birds reached Hawaii and Hualalai, at elevations between approxi­ from the areas indicated in Figure 1. The mately 5,000 and 8,000 feet. Here the birds endemic Hawaiian birds, listed according to often build their nests on the lava although these 11 families, are discussed briefly. typically well concealed in clumps of vegeta­ 1. ( s, geese, and swans) tion. The nests are lined with the birds ' own down ; the clutch consists of from 2 The NENE or HAWAIIAN GOOSE (B rante to 5 . sandvicensis) is endemic to the island of Ha- The KOLOA or HAWAIIAN DUCK (Anas wy­ 1 Manuscript received June 13, 1969. villiana) originally was found on all of the 2 D epartm ent of Zoology, University of H awaii, . Supported by NSF Grant GB-5 612. main Hawaiian Islands except and Ka- 29 I.» o

HauiaJlan Th rush. Nene, Owl. Crow. Hawk. Ga l/lnule. Coot.

Stilt._Black i ·crowned Night Heron

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RAROTONGA, - .~ ~ ., ~

N PITCAIRN ."'" '-< ~ § ~l ~ -< ..... F IG. 1. Map of the Pacific Basin to show regions from which the ancestors of endemic Hawaiian birds are presumed to have originated, The broken \0 -....J arrow indicates the annual migratory flights of the Pacific Golden Plover between Alaska and the Hawaiian Islands, o Birds of Hawaii-BERGER 31 hoolawe. A decline in numbers of Koloa on on , and, after these operations ceased, most of the islands was noted by several writ­ Japanese hunters also used large num­ ers after the turn of the present century, and bers of the for food. Theodore Roose­ in recent years this duck has been found only velt established the Hawaiian Islands National on . A propagation program is now un­ Wildlife Refuge in 1909, but Alfred M. Bai­ derway at Pohakuloa, Hawaii. ley believed there were only seven ducks left Man probably was the most serious predator by 1912. Alexander Wetmore (1925) counted on the Koloa, and the birds could be hunted 20 birds during the Tanager Expedition of legally during the early 1920s, when the bag 1923. Since that time, when the last of the rab­ limit was 25 ducks per day. Duck hunting bits were believed to have been killed, both was prohibited for a period of two years be­ the vegetation and the have ginning in 1939, and hunting was further made a remarkable recovery. Because of the closed during World War II. Fortunately, dense vegetation of the areas inhabited by the duck hunting (both for the Koloa and the ducks, it is virtually impossible to make an wintering migratory ducks) has been prohib­ accurate count of the birds, but the population ited since that time. The decline in taro and is now thought to fluctuate between 100 and rice acreages, however, has reduced suitable 600. The downward fluctuations in population habitat for the birds. .that do occur are thought to result in part Wild dogs are known to be serious preda­ from severe winter storms, but there may be tors on both ducklings and the adults in their other, as yet unknown, reasons. The present flightless stage during the annual postnuptial habitat is thought to be adequate for about molt. A number of other (e.g., large­ 600 ducks. mouth bass, bullfrog) have been known to It is imperative for the future welfare of the kill small ducklings on Kauai. The role of the Laysan Duck (as well as for the surviving in the great reduction or extinction honeycreeper and the tens of thousands of of the Koloa throughout most of its former nesting ) that predators (such as , range is unknown, but it may be significant , dogs) and pest and plants (which that Kauai, the last stronghold of the Koloa, . would alter the ecology of the island) be pre­ is the only main island on which the mon­ vented from gaining access to Laysan Island. goose has not been introduced. Wild cats, rats, Nests of the Laysan Duck are built on the and pigs also destroy nests. ground and are well concealed among the vege­ The main breeding season on Kauai appears tation. Little is known about the breeding biol­ to be from December through May, although ogy in the wild, however. There are more than the species seems to breed throughout the year, 150 birds in zoos and private aviaries; breeding inasmuch as nests or downy young have been pairs also are held at Pohakuloa, Hawaii. found in all months except August (Swed­ 2 . Accipitridae (hawks, kites, and eagles): berg, 1967). The well-concealed nests are built on the ground. Clutch size is reported to The HAWAIIAN HAWK or 10 (Buteo soli­ be from 2 to 10 eggs, with a mean of 8.3 tarim), for entirely unknown reasons, has al­ eggs for wild birds. The Koloa is tolerant of ways inhabited Hawaii only. The is now varying climatic and ecological conditions. On uncommon and has been placed on the list of Kauai the birds nest from level to 3,500 rare and endangered species. The chief reason feet elevation, and in areas of annual rainfall for the decline in numbers of this interesting varying from 35 to 125 inches. bird is believed to be shooting by uninformed people who consider all hawks to be " The LAYSAN DUCK (Anas laysanemis) was hawks." All available evidence, however, sug­ in danger of extinction during the early part gests that rodents form the main food of the of this century (Rothschild, 1893-1900). 10. Although much rarer than the Hawaiian Again man was the agent of destruction. The Owl, the hawk sometimes can be seen soaring birds were hunted for sport and for food by high in the air on the slopes of both Mauna the personnel of the guano mining company Loa and Mauna Kea. 32 PACIFIC SCIENCE, Vol. 24, January 1970

Few nests of this species have been found Several birds survived the long sea voyage to and very little is known of its breeding habits. England in the 1890s. The birds build nests of twigs and sticks in Like many rails on oceanic islands, the Lay­ trees, and one author has described a lining san had evolved into a flightless condition . made of the stems and fronds of ferns . Two The birds ate many kinds of insects, the flesh or three eggs are thought to form a clutch. from the carcasses of other birds, and the eggs of the smaller seabirds, such as and 3. Rallidae (rails, gallinules, and coots): petrels. Although the rails apparently some­ times broke open the eggs, they are said usu­ The (P orzanula palm eri) had a historical life of 116 years. The species was ally to have waited until Laysan discovered on Laysan Island in 1828, and it (Psittirostra cantans) broke through the shells with their more powerful bills and then chased probably became extinct in 1944. the finches away to eat the contents. The devastation wrought by the rabbits re­ The breeding season apparently extended leased on Laysan Island in 1903 is well de­ from late March through July on Laysan, but picted by Alexander Wetmore (1925) after downy young were seen on Midway in March . he visited Laysan in 1923 as a member of the The nests were built on the ground or in grass Tanager Expedition: "On every hand extended tussocks and were constructed of dried stems a barren waste of sand. Two coconut palms, and of juncus and other plants. Unlike a stunted hau tree and an ironwood or two, Mainland rails, which lay large clutches of planted by former inhabitants, were the only eggs, the Laysan Rail laid 2 to 4 eggs in a bits of green that greeted the eye. Other vege­ clutch (Baldwin, 1947) . tation had vanished. The desolateness of the scene was so depressing that unconsciously we The H AWAIIAN RAIL ( Pe11J11Ila sandwichen ­ talked in und ertones. From all appearances, sis) was last collected on the island of Hawa ii Laysan might have been some desert, with the (near Olaa ) about 1864, and was last seen gleaming lake below merely a mirage." about 1884 (Greenway, 1958, p. 235). Munro It was estimated that there were about 2,000 (19 44, p. 51) believed that this rail "fre­ rails on Laysan as late as 1915, but Wetmore quented most of the larger islands" and that .and his party found only two birds, and the "it certainly was on ," although there species is thought to have been extirpated there is no good evidence for this belief. The nest, before 1936. eggs, and newly hatched young were never A pair of Laysan Rails was introduced to described. Midway Island in 1891. A large population The causes of extinction of this small (about had built up by the turn of the century and 5Yz inches in total length ) , flightless rail are still existed in 1939. The extermination of the unknown, but it seems certain that rats, dogs, Laysan Rail on Midway, however, was very and cats played a large role in the extermina­ rapid after the onset of W orId War II when tion of this unique species. Hawaiian chiefs the U.S. Navy took over, and rats gained ac­ are said to have hunted the rail with bows and cess to both Sand and Eastern islands. The last arrows. rails were seen on Eastern Island in June of The GALLINULE ( Gallimtla cbloropus sand ­ 1944. vicel1sis ) is considered conspecific with the The Laysan Rail could easily have been saved Common Gallinule of North America and from extinction if Government officials had Eurasia, although the Hawaiian birds are non­ heeded the pleas of ornithologists, but they migratory and have been inhabitants for an were unable to obtain the necessary transpor­ unknown length of time. Their distinctness is tation to restock Laysan Island (or other is­ indicated by their subspecific name. These are lands) from the Midway population after the birds of fresh-water ponds and marshes, and, vegetation began to recover on Laysan. Al­ because of the continuing disappearance of though small and flightless, the Laysan Rail such habitats in Hawaii, the birds are consid­ was a hardy bird , easily reared in captivity. ered endangered on all islands they still in- Birds of Hawaii-BERGER 33

habit. They formerly inhabited all of the main linule chicks, the young are able to move about islands except and Lanai. Attempts to shortly after hatching, when the down has reestablish the birds on Hawaii and Maui ap­ dried . pear to have been unsuccessful. Essential habi­ tat is being destroyed to make way for housing 4. Recuruirostridae (avocets and ) developments, and , rats, dogs, and The BLACK-NECKED or HAWAIIAN cats are serious predators on the birds. The (Himantopus himantopus knttdseni) is a large prospects for survival of this species are con­ (16 inches), striking, black-and-white bird sidered by personnel of the Bureau of Sport with very long reddish legs. This species is Fisheries and Wildlife as "not good." endemic to the islands of Niihau, Kauai, Oahu, The gallinule builds its nest of reeds and Molokai, Maui, and Hawaii, but is now greatly other aquatic vegetation. Like its continental reduced in numbers in most of its former relatives, the bird is thought to lay large range. clutches of eggs (6 to 13). The newly hatched The stilt was considered a game bird until young have red bills and are covered with 1941, and still is sometimes shot illegally. The black down feathers ; they are precocial and birds also are subject to predation by the mon­ are able to run about and swim within a few goose and by feral dogs and cats. A major hours after hatching. reason for the decline of this species, however, has been the continual draining of marshes and The (Falic« americana alai) other wetland areas. also is considered conspecific with the North It is estimated that the total population of American members of this widely distributed the stilt now numbers about 1,500 birds, which species. Like the gallinule, however, the Ha­ are found chiefly on Oahu and Maui. One of waiian birds are nonmigratory and have been the major nesting and feeding habitats is at inhabitants of the Hawaiian Islands for a long Kanaha Pond on Maui. This marsh area is in period of time. They are given subspecific constant danger of being filled in because of designation. pressures to enlarge the runways at .the Ka­ Coots occupy the same general type of fresh­ hului Airport. Another important breeding water ponds as gallinules, but they prefer more area is found among the ponds on the Kaneohe open water. The coot is found on all the main Marine Air Station on Oahu; efforts have been islands, and is especially common on Kauai made with military personnel to have some of and at Kanaha Pond on Maui. Munro (1944, these ponds set aside as a sanctuary for the p. 54) reported seeing "from 500 to 600 on stilt. a lagoon near Lihue, Kauai," in 1891, but no­ The nest of the stilt is a simple "scrape" where are they so abundant now. One or more made on the ground by the birds themselves; birds sometimes can be observed on the reser­ small stones, bits of wood, and other debris voir along the Old Pali Road in Nuuanu often are added to the scrape. The normal Valley, Oahu. Coots were on the game bird clutch is 4 eggs. The newly hatched, precocial list until 1939. They are now classified as an young are covered with a coat of variegated endangered species, with an estimated total brownish down, which makes them very diffi­ population of 1,500 birds. cult to find after they leave the nest. The young Coots typically build relatively large floating are brooded for some time after hatching, but nests of aquatic vegetation. Little is known of they run from the nest and hide in the sur­ the clutch size of the Hawaiian Coot, but it rounding vegetation when disturbed . presumably lays fewer than the 8 to 12 eggs of the Mainland birds. The newly hatched chicks 5. Stridgidae (owls) are covered with black down except on the The or SHORT-EARED OWL (Asio head, neck, and throat where the down is red­ flammelts sandwichensis) differs from most dish-orange. The down is short or absent on species of continental owls in that it is diurnal the forehead and crown of the head, giving in habits. It is found in open (such the bird a bald-headed appearance. Like gal- as along the western part of the Saddle Road 34 PACIFI C SCIENCE, Vol. 24, January 1970 of Hawaii), over lava flows, and in forested native birds are to be expected to survive; areas (both ohia and mamane-naio . for est ~ ) , population levels of goats and pigs also must and often it is seen near towns. This speCies be controlled. appears to be tolerant of wide climatic ex­ N othing is known about the breeding habits tremes-from relatively dry areas (a bout 20 of this thrush. inches of annual rainfall ) to the extremely The HAWAIIAN THRUSH (Pbaeornis obsmflls) wet Kokee area of Kauai. The Pueo is resident developed races on all of the main islands ex­ on all of the main islands of the chain, and cept Maui. The races found on Oahu and it was prominent in Hawaiian mythology. Lanai are presumed to be extinct; reports of The Pueo builds its nest on the ground. The survival of the Molokai race need to be con­ females are said to lay from 3 to 6 eggs in a firmed (Richardson, 1949) . clutch. Th e Kauai race, or the Large Kauai Thrush

6. Coroidae (CI'01llS, jays, and magpies) (P. o, myadestina) appears to be even rarer than the Small Kauai Thrush, whereas the The HAWAIIAN CROW (Coruus tl'opim s) is Large Kauai Thrush was said by early writers endemic to the island of Hawaii only, being to be the most common forest bird on Kauai found in the Kona and Kau distri cts. The bird in 1891. is now rare, and it is estimated that the total The Hawaii race (P. o, obsmms) still is population may be no more than 30 birds. fairl y common in suitable habitat ( Berg~r, In former times, they were much more com­ 1969a) . The birds inhabit the ohia forests 111 mon and were found at elevations from 1,000 regions of high annual rainfall, in general to 8,000 feet. Shooting is probably resp ~nsibl e above 3,000 feet elevation. The best areas are for the decline of this sole representative of on the Saddle Road, Stainback Highway, and the crow family to have reached the Hawaiian in the more undisturbed, wet forests of Ha­ Islands. Th e effect of the great alteration of waiian Volcanoes N ational Park, but this the environment on the decline of the species thrush also is found in ohia forests at higher is unknown. elevations on the Kona coast. Very little is known about the fee~i ng habits A nest of this species and was first or breeding biology of the Haw al1~n C~ow . found near the Saddl e Road on May 11 , 1968, The birds build nests of twigs and sticks, lined by Andrew J. Berger ( 1969). The nest, built with finer plant materials. The eggs have been on the trunk of a tree fern less than 5 feet described as having a greenish background above the ground, contained a single egg with brown markings around the larger end which was heavily covered by small, irregu­ of the egg. In April 1964, Dr. P. Quentin larly shaped, reddish-brown markings. Tomich ( 1967) found a nest containing five eggs in an ohia tree. 8. Sylviidae ( Old W orid uiarbler family) 7. Turdidae (thm sh family) Th e LAYSANMILL ERBIRD (Acl'ocephallts fa­ miliaris familial'is) was one of three species Th e SMALL KAUAI THRU SH ( Pbaeornis pal­ of endemic birds to become extinct on Laysan mel'i) is now known to inhabit .only the o hi~ prior to 1923 because of the destruction of forests in the Alakai Swamp regIOn of Kauai, the habitat by the rabbits. How the ancestors The size of the remaining pop ulation is un­ of this small bird (about 5 inches in total known but because of its restricted distribu­ Vz length ) managed to reach Laysan and tion, this s ~ e cics is thought to be rare and it is included in the list of rare and endangered is, of course, unknown. Because of their Old W orld affinities, however, it is assumed that species, as are most of the endemic Haw a ~~an they came from Asia and "island-hopped" to birds. Both this thru sh and the Hawaiian reach Laysan and Nihoa islands. Thrush (P. obsm ms) appear to tolerate very little change in environment. Hence the fur­ The N IH OA MILLERBIRD (AcrocephalttS fa­ ther spread of exotic plants into the dep~hs of miliaris kingi) has one of the most limited the Alakai Swamp must be prevented If the distributions of any bird species: Nihoa con- Birds of Hawaii-BERGER 35

tains 156 acres. Personnel of the Bureau of and 3 eggs in one nest. The incubation period Sport Fisheries and Wildlife estimated a total is 14 days, and the nestling period, 16 days. population between 500 and 600 in 1967. The Frings found the breeding season in species is endangered because of its limited Valley to extend from mid-January to mid­ distribution, and it is imperative that rats, cats, June. The season differs on the other islands and dogs be prevented from gaining access to for as yet unanalyzed reasons. Nihoa and the other islands in the Hawaiian Islands National Wildlife Refuge. 10. Meliphagidae ( family) The birds are secretive in habits, usually This is a large Old World family, contain­ staying in the dense cover afforded by Cheno­ ing 160 species of birds. Two genera and five podium sandtoicbeum (goosefoot) and Sida species were found in Hawaii, but all except fal/ax (ilima). Several nests have been found one species are now thought to be extinct. in this vegetation, but little else is known about either the breeding biology or the feeding The four species of the genus had habits of the Nihoa Millerbird. patches of bright yellow feathers, prized by the early Hawaiians who used them for their 9. M uscicapidae (Old W orld flycatcher family) feather capes and headdresses. The role that The EL EPAIO (Chasiempis sandwichensis) , the Hawaiians played in causing the extinction important in Hawaiian folklore, has a puzzling of the several species of the 00 is unknown, distribution in that races have developed on but it may be significant that the sole known Kauai, Oahu, and Hawaii, but there is no evi­ surviving species (on Kauai) has fewer yellow dence that the species was ever found on the feathers than any of the other species. The other main islands in the chain (Wilson and evidence also suggests that the Oos, like most Evans, 1890-1899) . of the other endemic Hawaiian landbirds, are Although not as common as reported during intolerant to any extensive changes in their the early 1900s, the Elepaio has been able to environment. Also unknown is the role played adapt to man-made changes in the environ­ by the three species of rats in the islands as ment as no other endemic landbird has been predators on the eggs and young of tree-nest­ able to do. The Oahu race (c. s.gayi) is still ing birds; some species of rats are agile climb­ fairly common in the mixed forests of the ers and have been seen in tall trees and in island, and a small population is resident in tree ferns in the ohia forests. the lowland introduced forest near the head of Manoa Valley. The KAUAl 00 (Moho braccatm) , formerly The Kauai race (C . s. sclateri) is common thought to be extinct, was rediscovered by in the Kokee State Park area as well as in the Dr. Frank Richardson in 1960 in the depths Alakai Swamp. The Hawaii race (C. s. sand­ of the Alakai Swamp region (Richardson and wichensis) is found both in the wet ohia for­ Bowles, 1964). The bird is very rare and ests and in the dry mamane-naio forest on nothing is known of its breeding habits. Mauna Kea. The OAHU 00 (Moho apicalis) is thought to Frings (1968) found that the Oahu Elepaio have become extinct within a short period after defended a territory of 4.9 acres. The nest site 1837 (Greenway, 1958, p. 423) . is selected by the female, but both sexes take The MOLOKAl 00 (Moho bishoPi) was last part in nest-building activities. The average reported in 1904 and is now presumed to be height above ground of 32 nests was 25 feet. extinct. The small cup-shaped nests are very neat and The HAWAII 00 (Moho nobilis) has not compact, and contain large quantities of spi­ been reliably reported since 1934 and is listed der web, which aids in holding the plant as "probably extinct." It is certain that the materials together. The eggs have a white birds no longer inhabit the forests where they background covered with reddish-brown spots, were collected in the 1890s, but there are vast which are concentrated at the larger end of forest areas on Hawaii which have not been the egg. The clutch size was 2 eggs in 15 nests visited by ornithologists, and this species may 36 PACIFIC SCIENCE, Vol. 24, January 1970

still exist in remote and relatively undisturbed Drepanididae (formerly, also, Drepaniidae) . areas. No suitable evidence has been found since that Th e (Chaetoptila angllStipll/1na) was time to refute this interpretation, although the a large bird, about 13 inches in total length. possibility exists that we are not dealing, in Its color pattern was unlike that of any other fact, with a true monophyletic family. N ever­ Hawaiian bird: a black face mask, greenish­ theless, I choose to follow the classification of brown wings and tail, and a heavily streaked the Hawaiian Honeycreepers proposed by Ama­ pattern of brown and white feathers on the don (1950) .There is a more recent system of head, upper back, and und erparts. The type classification (Greenway, 1968), but that au­ specimen was collected by the Pickering and thor had no more information on or Peale expedition in 1840 on the island of breeding biology than was available to Ama­ Hawaii, the only known range of the species. don. According to Munro (1944, p. 88) several Unfortunately, all of the highly specialized additional specimens were collected by Mills honeycreepers have become extinct on Oahu, about 1859. The species apparently has not Molokai, and Lanai, as have most of those on been seen since that time. Hawaii. Kauai, the only island on which the mongoose was not introduced, is the only island 11. Drepanididae (H awaiian honeycreepers) which still has all of the endemic birds known This endemic Hawaiian family exhibits to have occurred there. Most of these are con­ among its numerous species the most striking fined to the Alakai Swamp region, and many example of adaptive radiation from an assumed are now rare. The east and northeast slopes of single ancestral species of any bird family in Haleakala also have proven a haven for the the world . It demonstrates admirably, there­ survival of unique Hawaiian birds. fore, the results of evolutionary processes on One should note that the largest number of oceanic islands. That this family of birds has species of honeycreepers are now found in two not been studied more intensively in the past relatively undisturbed wilderness areas: the can be attributed, in part, to the fact that Alakai Swamp of Kau ai and the outer, wind­ Charles Darwin visited the Galapagos Islands ward slope of Haleakala. These are areas which and not the Hawaiian Islands. have not been much disturbed by wild cattle, The members of this family reached all of nor have they been desecrated by State for ­ the main H awaiian Islands, and two species esters and ranchers. were found on certain of the Leeward Islands Despite the remarkable bill adap tations in historic times. The tragic remnant popula­ foun d in the Hawaiian Honeycreepers, very tion of three individuals of the Laysan Honey­ little is known about this family of unique creeper (also incorrectly called the Laysan birds. The nests, eggs, and newly hatched "H oneyeater"}, a race of the Apapane, became young were never described for any of the extinct in 1923. Th e -billed extinct species (except for the Laysan H oney­ and still inhabit those respective creeper) . A nest with eggs of the Palila was islands. foun d for the first time by Andrew J.Berger It might be noted here that the name "finch" in 1968 (Berger, 1969b); the first nests of the was given to these birds by taxonomists in the Akepa and the Creeper to be foun d were re­ 1890s, because they thought that these large­ ported by C. Robert Eddinger in 1969. The billed birds (as well as some on the main incubation periods for this family also were islands) belonged to the finch family (F ringil­ first determined by Eddinger in 1969. lidae) . This interpretation was based almost exclusively on the superficial resemblance in A . SlIbfamily Psittirostrinae bill shape and size. Later students of Hawaiian AMAKIHI ( virellS) : This , the second birds (particularly Perkins, 1901), concluded most common living honeycreeper, is found on that a large number of the endemic Hawaiian all of the main islands. The fou r subspecies birds had evolved from a single ancestral spe­ are distributed as follows: L. v. stejnegeri, cies; all of these were included in the family Kauai; L. v. cbloris, Oahu; L. v. u/ilsoni, Maui, Birds of Hawaii-BERGER 37

Molokai, and Lanai; L. v. uirens, Hawaii. The region of Kauai. It appears to be rare on Maui Amakihi is a characteristic bird of the wet and Hawaii, and is presumed to be extinct on ohia forests on the windward slopes of the Oahu. The males of the Hawaii and Maui races islands, but it is also a common permanent have reddish-orange , whereas the resident of the dry mamane-naio forest on Kauai male has a yellow crown and underparts Mauna Kea. This broad climatic distribution and olive-green back and wings. suggests that the Amakihi may be the most KAUAI AKIALOA (Hemignathus procerus) : adaptable of the surviving species of honey­ This highly specialized honeycreeper with its creepers. long (over 2 inches) and strongly decurved ANIANIAU (Lo xops parva) : This species is bill was long feared to be extinct, but it was endemic to Kauai, and now is limited in dis­ rediscovered in the Alakai Swamp region in tribution to the Kokee and Alakai Swamp re­ 1960 by Richardson and Bowles (1964) . The gions of the island, where the bird is fairly bird must be very rare, probably close to extinc­ common. The nest and eggs of this species tion, and has been found by very few observers. were first described in 1969 (Berger, Eddin ­ AKIALOA (Hemignathus obscurusy: Subspe­ ger, and Frings, 1969). cies of a second closely related species of GREATER AMAKIHI (Lo xops sagittirostris) : Akialoa formerly inhabited Oahu, Lanai, and This bird, which has been called also (inappro­ Hawaii. Those on Oahu and Lanai are certainly priately) the Green Solitaire, had a very short extinct, and the Hawaii race is presumed to be known history. The species was first collected extinct. near the Wailuku River on Hawaii in 1892. It NUKUPUU (Hemignathus Ittcidus): The was rediscovered by Perkins in 1895, but has strongly decurved bill of the Nukupuu is not been observed since early in the present unique among birds in that the lower mandible century. The early collecting sites probably is only about half as long as the upper man­ were near the upper limits of the present sugar dible. Subspecies formerly were distributed as cane fields. However, there are extensive cloud follows: H . I. banepepe, Kauai; H. I. lucidus, forests along the Hamakua Coast of Hawaii Oahu; H . I. affinis, Maui. The Oahu race is where this species might still survive. extinct; the Maui race was rediscovered in 1967 ; and the Kauai race is very rare, inhabit­ CREEPER (Loxops maculata) : The six sub­ ing the depths of the Alakai Swamp. species of this small bird with a relatively short bill are : L. m. bairdi, Kauai; L. m, maculata, AKIAPOLAAU (Hemignatbus wilsoni): The Oahu ; L. m, flamm ea, Molokai ; L. m. montana, upper mandible of this closely related species Lanai; L. m. neuuoni, Maui; L. m. mana, also is long and strongly decurved but the Hawaii. The creeper is a relatively common lower mandible is straight, robust, and only bird in the Alakai Swamp region of Kauai and about half as long as the upper mandible . The on the windward slope of Haleakala Crater, Akiapolaau has woodpecker-like habits in that Maui. It is uncommon on Hawaii, rare on it pounds its lower mandible into dead branches Oahu and Molokai, and presumed to be extinct and tree trunks, searching for grubs and insects; on Lanai. An unusual feature of the Molokai the birds often forage on branches close to the race is that the males are reddish-brown, ground. The Akiapolaau is endemic to the whereas the males of the other races have island of Hawaii . The bird is very rare, and, in yellowish-green or brownish feathers, especially recent years, has been sighted only in the on the dorsal surface. mamane-naio forest on Mauna Kea, but for­ merly, at least, the species was found in Vol­ AKEPA (Loxops coccinea): This species dif­ canoes National Park. ferentiated into subspecies on Kauai (L. c. caeruleirostris), Oahu (L. c. ruta) , Maui (L. c. MAUl PARROTBILL (Pseudonestor xantbo­ ochracea), and Hawaii (L. c. coccinea) . The phrys): This remarkable stub-tailed bird with Akepa is fairly common in the Alakai Swamp a large parrot-like bill is known to have oc- 38 PACIFIC SCIEN CE, Vol. 24, January 1970 curred only at higher elevations on the very would be the last endemic H awaiian bird to wet, windward slopes of Haleakala, Maui. Vir­ be discovered. In 1923, however, Alexander tually nothing is known about this rare bird. We tmore visited Nihoa and discovered the It was observed in the upper reaches of Ni hoa Millerbird. Kipahulu Valley in August 1967. The Nihoa Finch is a successful species, with an estimated population in 1967 of be­ au (Psittirostra psittacea): The Ou is a tween 4,800 and 5,000 birds on Nihoa's 156 large-billed, yellow-headed bird with a green­ acres. The survival of the species, however, ish back. The species once inhabited Kauai, depends upon maintenance of the native vege­ Oahu, Molokai, Lanai, Maui, and Hawaii. It is tation and prevention of the introduction on extinct on Oahu, Molokai, and Lanai; Maui is the island of rats and other mammalian preda­ not even listed as part of the forme r range in tors. 'most books, and the species has not been seen there for many years. The Ou apparently was PAULA (Psittirostra bailleui): The Palila is last reported seen on Hawaii in 1955. The bird superficially similar to the Ou, being a large­ is rare on Kauai but can be found in the Alakai billed and large-headed bird with a yellow Swamp. head and throat but with a gray back. Th e bill of the Palila is dark in color, rather than light, LAYSAN FINCH (Psittir ostra cantans cantans ): and is differently shaped than t he bill of the This species was first described by S. B. Wilson Ou. The Palila is found only on Hawaii. It in 1890. It was undoubtedly because of its had a wider distribution on that island in the omnivorous feeding habits that the Laysan past but is now known to occur only in the Finch managed to survive the destruction by mamane-naio forests on the slopes of Mauna rabbits of the vegetation on Laysan Island. Kea, in general at elevations above 6,500 feet. During the extended breeding season of the The survival of the Palila, therefore, is entirely seabirds, the Laysan Finch breaks the eggs of depen dent on the recovery and continued main­ the several species of nesting terns, especially, tenance of this forest. The present tree line is and eats their contents. Wetmore found sev­ located at approximately 9,300 feet and is eral dozen Laysan Finches on the island in gradually receding because of overgrazing by 1923. The population has increased steadily the feral sheep on the State-owned game man­ since that time, and there were between 8,000 agement areas on Mauna Kea. A superabun­ and 10,000 Laysan Finches on the island in dance of wild pigs also inhabits this relatively 1967.Personnel of the U.S.Bureau of Sport dry forest region. Fisheries and Wildlife have since released birds on Southeast Island of Pearl and H ermes GREATE R KOA F INCH (Psittirostra palm eri) : Reef ; the population is now thought to num­ Th is and the next two species of Koa Finch ber between 75 and 100 birds. provide ornithologists with a real puzzle. Th e The Laysan Finch also was introduced on three species were first discovered and described Midway Island in 1891 and again in about during the period between 1888 and 1892, all 1905. Bailey (1956, p. 124) wrote, however, on the Kona slope of Mauna Loa. Th ere are that "the disappearance of the finches and the no reliable records of anyone of these species [Laysan] rails was very rapid when rats over­ having been seen since 1896 . ran the islands during the war years, and few if any existed on Midway after 1944." LE SSER KOA FINCH ( Psittirostra flaviceps) About two dozen Laysan Finches were pre­ GROSB EAK FINCH ( Psittirostra kona) sented to the in 1966, and a similar number were sent to the Un iversity of B. Subfamily D repaniinae Michigan. APAPANE ( Himatione sangl/inea ) : This is NIHOA FINCH ( Psittirostra cantans ultima ) : the most common of the surviving species of This honeycreeper was named "ultima" in honeycreepers, and it has undergone no im­ 1917 by W . A. Bryan because he thought it portant geographic variation. One race (H. s, Birds of Hawaii-BERGER 39 sanguinea) inhabits all of the six main Ha­ accompanying Captain Cook first collected this waiian islands . A second race (R. s. freethii) striking yellow and black bird with its very inhabited Laysan Island until becoming extinct long, decurved, black bill, which was endemic in 1923. only to the island of Hawaii in historic times. The Apapane is the most conspicuous of The head and body feathers were black but the native birds in the wet ohia forests on all the rump, upper and lower tail coverts, thighs, major islands , in general now above 3,000 feet bend of wing, and part of the under wing elevation. It also is found along the Mauna coverts were bright yellow. Munro (1944, p. Loa Strip Road in Volcanoes National Park, 91) reported that the birds were still' being among both the scattered ohia trees and the collected for their yellow feathers in 1880, groves of koa (Baldwin, 1953) . The species and that one man shot as many as 12 in one rarely moves through the rnamane -naio forest day with a shotgun. The species apparently on Mauna Kea, and is not known to nest there. was last seen in 1899. The places to observe this species most easily BLACK ( ftmerea): This jet are at Kokee State Park on Kauai, Hosmer's black bird with white on the wing feathers Grove on Maui, and Volcanoes National Park had an even larger, decurved bill than the on Hawaii. Mamo. Endemic to Molokai, the species was CRESTED HONEYCREEP ER (P alm eria d olei): discovered in 1893. The last specimens appar­ This remarkably plumaged honeycreeper (to­ ently were collected in 1907, and the species tally unlike any other species) once inhabited is thought to be extinct. both Molokai and Maui. The species is extinct on Molokai, and almost nothing is known INDIGE NOUS HAWAI/AN BIRDS about the birds on Maui. There they are found in the cloud forest on the northeast slope of Indigenous birds are those native to Hawaii Haleakala, presumably at elevations above but whose normal range of distribution in­ 5,000 feet. cludes a much wider geographical area. In­ cluded among these indigenous Hawaiian birds ULA-AI-HAWANE (Ciridops anna ): Accord ­ are many seabirds, the Black-crowned Night ing to Munro ( 1944, p. 99), the common Heron, and a number of migratory species that name means "the red bird that feeds on the spend the nonbreeding season in the Hawaiian hawane" ( the native Hawaiian palm, Pritchardia Islands. Most of these birds are illustrated in spp .) . The color pattern of this species, too, the books by Ord ( 1967) and Peterson was unlike that of any other honeycreeper. (1 961) . The species apparently was first collected about 1859 but it was not described until 1879. BLACK-CROWNED NIGHT HERON (Nycticorax Palmer and Munro obtained one specimen nycticorax hoactli) : This heron is considered from natives in the early 1890s, the last of indigenous rather than endemic (see gallinule this species ever reported. and coot) because the Hawaiian birds have not been recognized as subspecifically distinct I/WI (Vestiaria coccinea): This is the most in plumage characters from the American con­ striking in appearance of the common honey­ tinental birds. This subspecies has a very large creepers. The head and body feathers are a breeding range, extending from Washington brilliant vermillion; there is a white patch in and Oregon south to northern Chile and south­ the otherwise black wings ; and the long, de­ central . curved bill is salmon colored . This species once The Black-crowned Night Heron, found on inhabited all of the main islands . It is extinct all the main islands, inhabits marshes ponds on Lanai and probably Molokai, and it is rare and lagoons, where it feeds on aquati~ insects: and presumably on the verge of extinction on fish, frogs, and mice. The birds roost and nest Oahu. On Kauai , Maui, and Hawaii, however, in trees. The future of this species in Hawaii, it is still fairly common . like that of the gallinule and coot, is dependent MAMO (Drepanis pacifica): The naturalists on the preservation of suitable wetland habitat. 40 PACIFIC SCIENCE, Vol. 24, January 1970

The Seabirds Brown (Sttla lettcogaster) These birds belong to several families of Red-footed Booby (Stlla sttla) oceanic birds (as indicated below) . They breed Fregatidae () by the tens of thousands on the Leeward Is­ Great (Fr egata minor) lands, and certain species nest on the offshore Laridae (, terns, and noddys) islands of Kauai and Oahu (especially Moku Sooty (S terna ftlScata) Manu and Manana Island (Fisher, 1948a, Gray-backed Tern (Sterna llmata) 1966 ) . The offshore islands are State wildlife Blue-gray Noddy ( Procelsterna cemlea ) refuges, and permission is required before Brown No ddy (AnotlS stolidtlS) visiting these islands. White-capped or Hawaiian Noddy (Anous A few species nest on the main islands. Red­ minttttlS) footed (Sul« sltla) have established Fairy Tern ( Gygis alba) breeding colonies near the Kilauea Lighthouse M igratory species on Kauai and on Ulupau Head, Oahu. N ewell's Manx (Pitffimts r«ffimts newel/i ) Th e most conspicuous of these is the Pacific is known to breed on almost inaccessible cliffs Golden Plover (Plttvialis do minica) , which on Kauai. The Hawaii an Dark-rumped Petrel spends the nonbreeding season in Hawaii, in­ ( Pterodroma phaeopygia sand wichensis) nests habiting grassy areas in the cities as well as in on the walls of Hal eakala Crater, Maui, and, the mountains. Most of the birds molt into the in smaller numbers, on Hawaii. Both species full breeding plumage before leaving for are thought to have nested formerly on all of Alaska, usually in April. the high islands, and both are listed as endan­ A number of species of ducks and shore­ gered species. birds also are winter residents in the Hawaiian Families containing species that nest in the Islands (see Ord , 1967; Bryan, 1958; and Hawaiian Islands and the species are these: Clapp and Woodward, 1968).

INTRODUCED OR EXOTIC BIRDS Diomedeidae ( or gooney birds) Black-footed (Diomedea nigripes) Virtually all of the landbirds that one sees (Diomedea im mtltabilis) in Honolulu, as well as in lowland areas on Procellariidae (, petrels, fulm ars) all islands, are introduced species: for exam­ Wedge-tailed Shearwater ( P«ffintlS pacifiCtts) ple, doves, mynahs, white-eyes, cardinals, Christmas Island Shearwater (P«ffintlS nativi­ mockingbirds, linnets (F isher, 1948b; Edd in­ tattlS) ger, 1967a, 1967b; Walker, 1967; Warner, Newell's Manx Shearwater ( Pttffi111tS pttf­ 1968) . To see endemic birds, one must get fimts newel/i ) into the mountains and the native forests' and Dark-rumped Petrel (Pterodroma phaeo­ few native birds remain on the island of Oahu . pygia sandwichensis ) In addition to the 76 species of game birds (Pterodroma hyp olettca) known to have been introduced in the main Bulwer's Petrel (B uli oeria bttlwerii ) Hawaiian Islands as of 1967, at least 60 spe­ Hydrobatidae (storm petrels) cies of non-game birds have been released. Harcourt's Storm Petrel ( Oceanodroma cas­ Th ese cover the gamut from the Chinese Fish­ tro) ing Cormorant (Phalacrocorax carbo) and the Sooty Storm Petrel ( Oceanodroma mark­ Guam Edible-nest Swiftl et ( Col/ocalia inex­ hami ) pectata) to a wide variety of birds. Phaethontidae (tropicbirds) Fortunately, most of the introductions have White-tailed Tropicbird (Ph aethon leptlt1'tts) been unsuccessful. Red-tailed Tropicbird (Ph aethon mbricaltda) Th e exact numb er of birds which have been Sulidae (boobies and gann ets) introduced is uncertain because an unknown Blue-faced or Masked Booby (Sttla dac­ number of cage birds have been released ille­ tylatra) gally by presumably well-meaning, but igno- Birds of Hawaii-BERGER 41

rant, citizens. A surprising number of weaver­ --- 1969b. The eggs and young of the finches (family Ploceidae, to which the House Palila. Condor, vol. 71. In press. Sparrow, Passer domesticus, belongs) have BERGER, A. J., C. R. EDDINGER, and S. C. been released intentionally by citizens on the FRINGS. 1969. The nest and eggs of the slopes of Diamond Head in recent years, and Anianiau. Auk, vol. 86, pp. 183-187. a number of these seem to be established as BRYAN, E. H., JR. 1958. Check list and sum­ breeding birds. mary of H awaiian birds. Books about Ha­ The success of an exotic bird introduced waii, Honolulu. into a foreign environment is best exemplified, CLAPP, R.B., and P. W . WOODWARD. 1968. perhaps, by the Mejiro or Japanese White-eye New records of birds from the Hawaiian ( Zosterops palpebrostls japonicus; family Zos­ Leeward Islands. Proceedings of the U.S. teropidae) . According to Bryan (195 8, p. 21) , National Museum, vol. 124, no. 3640, 39 the White-eye was introduced to Oahu from pp. Japan in 1929. This species has spread (appar­ EDDINGER, C. R. 1967a. A study of the breed­ ently unaide d by man) to all of the main ing behavior of the mynah (Acridotheres islands ; it is found both in the very dry and tristis 1.) . Elepaio, vol. 28, pp. 1-5, 11-15. the very wet habitats , and from sea level to - -- 1967b. Feeding helpers among imma­ treeline on the mountains of Hawaii and Maui. ture White-eyes. Condor, vol. 69, pp. 530­ Another successful exotic is the Red-billed 531. Leiothrix (Leiothrix It/tea; family Timaliidae) , which was released in 1918 and again in 1928­ ELDER, W . H. 1958.Biology and manage­ 1929. This species prefers the wetter areas ment of the Hawaiian Goose. Transactions (both native and introduced vegetation) , but of the 23rd N orth American Wildlife Con­ is now widely distributed on the main islands. ference, Washington, D. C. It is a common bird in the native forests, FISHER, H. I. 1948a. Laysan Albatross nesting where it, as well as the White-eye, may be on Moku Manu Islet, off Oahu, T. H. competing seriously with the endemic birds. Pacific Science, vol. 2, p. 66. --- 1948b. The question of avian intro ­ LITERATURE CITED ductions in Hawaii. Pacific Science, vol. 2, pp . 59-64. AMADON, DEAN. 1942.Relationships of the --- 1966 . Airplane-albatross collisions on Hawaiian avifauna. Condor, vol. 44, pp . . Condor, vol. 68, pp. 229­ 280- 281. 242. --- 1950. The Hawaiian honeycreepers FRINGS, S. C. 1968. The breeding biology of (Aves, Drepaniidae). Bulletin of the Amer­ the Oahu Elepaio, Chasiempis sandioicbensis ican Museum of N atural History, vol. 95, gayi. Unpublished thesis, University of Ha­ article 4. wall. BAILEY, A. M. 1956. Birds of Midway and GREENWAY, J. c., JR. 1958. Extinct and van­ Laysan Islands. Denver Museum of Natural ishing birds of the world. American Com­ H istory, Museum Pictorial No . 12. mission for International W ild Life Protec­ BALDWIN, P. H. 1947. The life history of the tion, Special Publication 13, New York. Laysan Rail. Condor, vol. 49, pp . 14-21. --- 1953. Annual cycle, environment and --- 1968. Drepanididae, Hawaii an honey­ evolution in the H awaiian honeycreepers creepers. In: Check- of the (Aves, Drepaniidae) . Universi ty of Califor­ world . Vol. XI V. Museum of Comparative nia Publications in Zoology, vol. 52, pp . Zoology, Cambridge, Massachusetts. 285-398. MAYR, ERNST. 1943. The zoogeographic posi­ BERGER, A. J. 1969a. Discovery of the nest of tion of the Hawaiian Islands. Condor, vol. the Hawaiian Thrush. The Living Bird, 45, pp. 45-48. Eighth Annual, Laboratory of , MILLER, A. H. 1937. Structural modifications Cornell University. in the Hawaiian Goose (N esochen sand- 42 PACIFIC SCIENCE, Vol. 24, January 1970

vicemis) , a study in adaptive evolution. ment of Land and Natural Resources, Hono­ University of California Publi cations in lulu . Zoology, vol. 38, pp. 11-242. TOMICH, P. Q. 1967. Arthropoda associated MUNRO, GEORGE. 1944. Birds of Hawaii. with a nest of the Hawaiian Crow.Proceed­ Tongg Publishing Co., Honolulu. ings of the Hawaiian Entomological Society, ORD, W . M. 1967. Hawaii's birds. Hawaii vol. 19, pp . 431-432. Audubon Society, Honolulu. WALKER, R.1. 1967. A brief history of exotic PERKINS, R. C. 1. 1901. An introduction to game bird and mammal introductions into the study of the Drepanididae. Ibis, 1901, Hawaii, with a look to the future. Confer­ pp . 562-585. ence of Western Association of State Game PETERSON, R. T. 1961. A field guide to west­ and Fish Commissioners, Honolulu, July 19, ern birds. 2nd ed. H oughton Mifflin Co., 1967. Boston. WARNER, R. E. 1961. Hawaii's birds- birth RICHARDSON, FRANK. 1949. The status of na­ and death of an island biota. Pacific Dis­ tive land birds on Molokai, Hawaiian Is­ covery, vol. 14, pp. 6- 13. lands. Pacific Science, vol. 3, pp. 226-230. --- 1968. Th e role of introduced diseases RICHARDSON, FRANK, and JOHN BOWLES. in the extinction of the endemic Hawaiian 1964. A survey of the birds of Kauai, Ha­ avifauna. Condor, vol. 70, pp. 101- 120. waii. Bernice P. Bulletin WETMORE, ALEXANDER. 1925. Bird life 227. among lava rock and coral sand, the chron­ ROTHSCHILD, WALTER. 1893-1900. The avi­ icle of'.a scientific expedition to little known fauna of Laysan and the H awaiian posses­ islands of Hawaii. National Geographic, sions. R. H . Porter, London. vol. 48, pp . 77-108. SWEDBERG, G. E. 1967. Th e Koloa. State of WILSON, S. B., and A. H . EVANS. 1890-1899. H awaii Division of Fish and Game, Depart- Aves Hawaiienses. R. H. Porter, London .