(DRAFT) - Taxonomy
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



TAXONOMY

NAME - PALILA OTHER COMMON NAMES - PALILA; 'O'U-PO'OPAPALE;'O'U PO'OPAPALE ELEMENT CODE - CATEGORY - Birds PHYLUM AND SUBPHYLUM - CHORDATA, CLASS AND SUBCLASS - AVES, ORDER AND SUBORDER - PASSERIFORMES, FAMILY AND SUBFAMILY - FRINGILLIDAE, GENUS AND SUBGENUS - LOXIOIDES, SPECIES AND SSP - BAILLEUI, SCIENTIFIC NAME - LOXIOIDES BAILLEUI AUTHORITY - TAXONOMY REFERENCES - COMMENTS ON TAXONOMY - Palila Loxioides bailleui (Oustalet, 1877) KINGDOM: Animal GROUP: Bird PHYLUM: Chordata CLASS: Aves ORDER: Passeriformes FAMILY: Fringillidae The Palila is a large-billed, large-headed bird about 15.25 cm long with a golden-yellow head and throat but with a gray back; the abdomen is grayish white; the lores are blackish; the bill is dark in color and is not hooked or parrot-like (01). The female's throat and breast are duller than in the male; the crown is only slightly yellowish (02). The Palila is currently placed in the monotypic genus Loxioides, but in the past it was included with several other allied species in Psittirostra (02,03). The taxonomic author is Oustalet, who described the species in 1877 (04). The Palila is also known by the Hawaiian name 'o'u-po'opapale (or o'u' po'opapale). The holotype is located in the Museum National d'Histoire Naturelle in Paris, France (05). In the United States collections may be found in the [number of specimens are given in brackets]: Museum of Natural History in New York, NY [21]; Bernice Pauahi Bishop Museum Taxonomy - 1 (DRAFT) - Taxonomy Species PALILA Species Id ESIS101017 Date 14 MAR 96 in Honolulu, HI [18]; Museum of Vertebrate Zoology at the University of California, Berkeley, CA. [10]; Museum of Comparative Zoology at Harvard University, Cambridge, MA [5]; National Museum of Natural History in Washington, D.C. [5]; Carnegie Museum in Pittsburgh, PA [2]; California Academy of Science collection in Los Angeles, CA [2]; Academy of Natural Sciences collection at Philadelphia, PA [1]; Los Angeles County Museum, Los Angeles, CA [1]; and the Peabody Museum at Yale University, New Haven, CT [1] (05). Illustrations of the species may be found in "Hawaiian birdlife" by Berger (01), "The Hawaiian honeycreepers (Aves, Drepaniidae)" by Amadon (02), "Hawaii's birds" by Shallenberger (06), "A field guide to western birds" by Peterson (07), "Birds of Hawaii" by Munro (08), "The avifauna of Laysan and the neighbouring islands" by Baron Rothschild (09), "Aves Hawaiienses: the birds of the Sandwich Islands" by Wilson and Evans (10), "Forest bird communities of the Hawaiian Islands: Their dynamics, ecology, and conservation" by Scott, et al. (11). Taxonomy - 2
                                  (DRAFT) - Status
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



STATUS

Coded Status E: Federal Endangered Non-consumptive recreational COMMENTS ON STATUS - U.S. STATUSES AND LAWS: The Palila (Loxioides bailleui) has been designated an Endangered species pursuant to the Endangered Species Act of 1973 (50 CFR 17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540), as amended. The species has this status wherever found including the State of Hawaii. Critical Habitat has been designated on the Island of Hawai'i (50 CFR 17.95 (b)) in portions of the Mauna Kea Forest Reserve, U.S. Army Pohakuloa Training Area, Ka'ohe Game Management Area, and other areas. This species is protected by the Lacey Act (P.L. 97-79, as amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import, export, transport, sell, receive, acquire, or purchase any wild animal (alive or dead including parts, products, eggs, or offspring): (1) in interstate or foreign commerce if taken, possessed, transported or sold in violation of any State law or regulation; or (2) if taken or possessed in violation of any U.S. law, treaty, or regulation or in violation of Indian tribal law. It is also unlawful to possess any wild animal (alive or dead including parts, products, eggs, and offspring) within the U.S. territorial or special maritime jurisdiction (as defined in 18 U.S.C. 7) that is taken, possessed, transported, or sold in violation of any State law or regulation, foreign law, or Indian tribal law. RESPONSIBLE FEDERAL AGENCIES: USFWS -Responsible for the management/recovery, listing, and law enforcement/protection of this species. DOD -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public land under their control. Also responsible for management/recovery on Department of Defense lands. All Federal agencies have responsibility to ensure that any action authorized, funded, or carried out by that agency is not likely to jeopardize the continued existence of the species or result in the destruction or adverse modification of Critical Habitat (50 CFR 402), and to utilize their authorities to carry out programs for the conservation of the species. Status - 1 (DRAFT) - Status Species PALILA Species Id ESIS101017 Date 14 MAR 96 STATE STATUSES AND LAWS: STATE: Hawai'i DESIGNATED STATUS: Endangered ADMINISTRATIVE AGENCY: Department of Land and Natural Resources, Division of Forestry and Wildlife. STATE STATUTE: Hawai'i Revised Statutes, Chapter 195D. INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS: The species is not listed by CITES. It is listed as an endangered species in the IUCN Red Data Book, 1977; and is also listed by the Convention on Nature Protection and Wildlife Preservation in the Western Hemisphere, 1970. ECONOMIC STATUSES: The Palila is one of the rare birds sought on Hawai'i by bird watchers. 67/03/11:32 FR 04001/ - Original listing 69/03/08:34 FR 05034/05035 - Republication of list 70/08/25:35 FR 13519/13520 - Proposed relisting 70/10/13:35 FR 16047/16048 - Final rule on previous list 76/12/22:41 FR 55729/55732 - Proposed ruling on Critical Habitat 77/03/11:42 FR 13569/ - Correction of error in proposed rule 77/08/11:42 FR 40685/40690 - Final rule, Critical Habitat 85/07/22:50 FR 29901/29909 - Five year review Status - 2
     

HABITAT ASSOCIATIONS

HABITAT - TERRESTRIAL TERRESTRIAL LAND USE - Cropland and Pasture Mixed Rangeland Evergreen Forest Land COMMENTS ON HABITAT ASSOCIATIONS - The present habitat of the Palila is the subalpine mamane-naio (Sophora chrysophylla-Myoporum sandwicense) woodland on Mauna Kea (13). Palila are found from 2,000 to 2,850 m elevation in woodlands with at least 5% canopy cover and of 2-10 m height (13). Palila are thermally stressed at 37 degrees C (17). The Palila shows habitat preferences because densities of birds can be as much as 100 times greater in some habitat types than in others. The species prefers areas with greater crown cover, taller trees, and higher proportion of native plants in the understory (13). Possibly some of the areas meeting the minimal habitat requirements would be unable to sustain a population; the birds occurring in these areas at present may merely represent population overflow or dispersal (11). An area of possible occurrence includes the sparse woodlands and open pastures with scattered trees on the northern and northwestern slopes of Mauna Kea, in the Humu'ula Saddle, and on the eastern slopes of Mauna Kea. Nest trees average 6.9 m in height and 83.4 cm in circumference at breast height. Both of these measurements are near the maximum for trees in the area (19). Nest placement averages 5.2 m above ground, and are placed on horizontal branches, some on terminal or lateral forks (19). Habitat Associations - 1
                                (DRAFT) - Food Habits
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



FOOD HABITS

TROPHIC LEVEL - OMNIVORE LIFESTAGE FOOD FOOD PART General Coniferae General Arthropods General Evergreen Trees-Flowers/Fruit/Seed Food Habits - 1
                         (DRAFT) - Environment Associations
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



ENVIRONMENTAL ASSOCIATIONS

G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS G Coastal Features: Vegetated offshore islands Environment Associations - 1
                               (DRAFT) - Life History
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



LIFE HISTORY

FOOD HABITS: The staple food item of Palila is the green, fully-inflated pods of the mamane tree (Sophora chrysophylla); other food items contributing significantly to the diet include mamane flowers, mamane leaf buds, mamane flower buds, naio (Myoporum sandwicense) flowers, naio berries, and insects (01,06,08,13,18,19,20,21). Little is known about seasonal variation in the food parts consumed. Juvenile and adult diets are similar (19). HOME RANGE/TERRITORY: Palila are territorial during the breeding season (19). Early in the breeding cycle, the territory is a "mate-defended area", with the male excluding all conspecifics from the vicinity of the female, usually by song and direct chase (19). As incubation begins, defense centers more on the nest, with females infrequently chasing other birds (19). Two measured territories were 0.39 ha and 0.06 ha (19). Territories are abandoned when the young fledged (19). The birds then join small feeding flocks (19). Some individuals return to the same area to nest the following year (13). PERIODICITY: The Palila is a diurnal bird (22). Breeding activity occurs from March through September and moult from May through December (19). Birds have a wider distribution in the non-breeding season (21). MIGRATION PATTERNS: The Palila is non-migratory (22). COVER/SHELTER REQUIREMENTS: The Palila requires mamane-naio woodland vegetation for cover and shelter. These woodlands have at least 5% canopy cover and are 2-10 m in height. Preferred habitat contains greater crown cover, taller trees, and a higher proportion of native plants in the understory (13). REPRODUCTIVE SITE REQUIREMENTS: Requirements, as opposed to observed patterns, are unknown. Average tree height of 26 nest trees was 6.9 m, circumference at breast height 83.4 cm; both values were near the maxima for trees in the area (19). Nest placement averaged 5.2 m above ground; 15 nests were on horizontal branches, 4 on terminal forks, and 7 on lateral forks (19). Mean distance of 26 nests from the trunk was 2.7 m and from the end of the branch 1.05 m (19). Nest base usually consists of large dead twigs broken off and collected from the nest tree (19). The bowl is composed of interwoven grasses and sometimes sheep wool, and lined with lichens or fine grasses and rootlets (19). Width of 26 nests averaged 14.7 cm, total height 7.7 cm, bowl depth 3.9 cm, and bowl width 7.4 cm (19). REPRODUCTIVE CHARACTERISTICS: Breeding occurs from March through September, with most intensive Life History - 1 (DRAFT) - Life History Species PALILA Species Id ESIS101017 Date 14 MAR 96 breeding from June to August (19). Low advertising flights are given by males during courtship, and sexual chasing occurs in the early breeding season (19). There is but one nesting per year (19). Pair bond appears to be the norm for mating relationships (19). Nest sites are selected by females (19). Clutch size is usually 2 (19). Incubation period is 15-16 days (19). PARENTAL CARE: Length of development from hatching to fledging is 21-27 days; parents may accompany young for over 30 days after fledging (19). Only the females broods, and females feed the young more than males (19). Development is altricial (19). Family bond duration is unknown. POPULATION BIOLOGY: The population is currently limited by (a) levels of food availability; (b) adverse habitat modification (c) possibly avian malaria and pox in Pohakuloa Flats; (d) drought and storms; (e) thermal stress; and (f) human disturbance (11,13). Survival rate, mortality rate, rate of increase, sex ratio, turnover rate, and longevity are unknown. The draft revised Recovery Plan suggests a population density of 25 birds/square km covering 200 square km or more throughout the mamane belt on Mauna Kea. For the Palila population to increase, the mamane forest must be expanded first. Reproductive success (number of eggs laid that fledged young) was 52 pct. during 1972-1975 (19). Population varies as much as 50% between years from a high of 6,410 birds in 1981 and low of 1,317 birds in 1985 (13,23). SPECIES INTERRELATIONSHIPS: The Palila's life cycle is closely tied with that of the mamane tree (01,02,06,07,08,09,10,11,13,16,18,19,20,21). Occurrence of avian malaria (Plasmodium relictum) and avian pox (Poxvirus avium) would probably preclude the presence of Palila in an area because of susceptibility and low resistance to these diseases (11,13,24,25,26, 27). The obligate vector for malaria and facultative vector for pox in these areas is the mosquito, Culex quinquefasciatus (28). OTHER LIFE HISTORY DESCRIPTORS: None. Life History - 2
                           (DRAFT) - Management Practices
                                   Species PALILA
                                Species Id ESIS101017
                                   Date 14 MAR 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Beneficial Suppressing wildfire Beneficial Restricting/regulating human use of habitats Beneficial Land Acquisition Beneficial Controlling/Restricting Agricultural Practice Beneficial Reforestation Beneficial Controlling/Removing Nonnative Vegetation Beneficial Disease Control Measures Beneficial Controlling/Removing Exotic Vertebrates Beneficial Controlling/Removing Feral Animals Beneficial Controlling/Removing Domestic Animals Adverse Artillery/Explosions Existing Artillery/Explosions Adverse Existing Adverse Food Supply Reduction Existing Food Supply Reduction Adverse Disease Existing Disease Adverse Parasites Existing Parasites Adverse Predation Existing Predation Adverse Exotic/Feral/Introducted Species Existing Exotic/Feral/Introducted Species Adverse Erosion Existing Erosion Adverse Vegetation Composition Changes Existing Vegetation Composition Changes Adverse Fire Existing Fire Adverse Forest Alteration Existing Forest Alteration Adverse Harvesting Existing Harvesting COMMENTS ON MANAGEMENT PRACTICES - Habitat destruction of low elevation dry forests and woodlands by Hawaiians probably significantly reduced the range of the Palila during the period from 400 A.D. to perhaps 1000 A.D. (12). Palila were extirpated from Kona between 1896 and 1936, probably as a response to the spread of avian diseases, particularly malaria and pox (13). Feral ungulates - first cattle, then sheep and to a lesser extent goats, and now (1985) mouflon - have had a severe impact on the mamane-naio ecosystem on Mauna Kea (13,14). The degradation and fragmentation of the subalpine woodland by feral ungulates followed in some areas by erosion, has contributed greatly to the reduction in range and population size of the Palila (13). Fire affected several hundred hectares in 1979 (13). Predators [black rats (Rattus rattus, Management Practices - 1 (DRAFT) - Management Practices Species PALILA Species Id ESIS101017 Date 14 MAR 96 cats (Felis catus)] doubtlessly have some impact on the population, but this has not been quantitatively studied (15). Climate affects the population through droughts and storms; production of the chief food item is related to rainfall (13,16). Palila are absent or very rare in apparently good habitat in the Pohakuloa Flats area; probable causes are excessive disturbance from military activity (explosions, off-road vehicles, etc.), disease, high temperatures, food supply shortages at certain seasons, and site tenacity (13). Future threats are difficult to envision, but might possibly include wildfire, excessive feral ungulate damage, excessive disturbance from development activities at Halepohaku, excessive disturbance from increased military activity, and imported disease vectors or parasites (11). APPROVED PLAN: U.S. Fish and Wildlife Service. 1986. Revised Palila Recovery Plan. U.S. Fish and Wildlife Service, Portland, OR. 49 pp. With its life history inextricably tied to a primary food source - the ripening seeds of the mamane tree - the Palila provides a relatively straight forward, albeit controversial, case of endangered species management. Forests of mamane, and with them the Palila, once enjoyed a wider distribution than they do today. Now that extensive stands of the tree can only be found on Mauna Kea, the range of the bird is also very restricted. The revised Palila Recovery Plan considers that before the species may be delisted it must, for a period of 5 years, exceed population densities of 25 birds/square km covering 200 square km or more throughout the mamane belt on Mauna Kea. For the Palila population to increase, the mamane forest must be expanded first. The Recovery Plan recommends the following actions to improve Palila habitat or in other ways benefit the species: 1) Eliminate mouflon, feral sheep, and feral goats that are increasing again in the mamane forest and detrimentally impacting it. 2) Improve stock proof fences protecting these forests. 3) Increase the Mauna Kea forest reserves by incorporating additional state lands, utilizing leases to limit agricultural practices, and easements, and land acquisition. 4) Control incursion of alien plants. 5) Prevent fires. 6) Continue studies of limiting factors, especially to learn why Palila are absent from the Pohakuloa Flats. 7) Determine the effect of avian disease upon the Palila needs to be determined and appropriate management actions taken. 8) As understanding of the mamane forest increases, assess and implement approaches for habitat rehabilitation. 9) Initiate an information and education program to encourage public support of these conservation efforts, which have often been criticized by hunters demanding that Mauna Kea be managed for sheep, mouflon, and goats. Management Practices - 2 (DRAFT) - Management Practices Species PALILA Species Id ESIS101017 Date 14 MAR 96 Recovery actions ongoing or completed include: 1) Studies of how feral sheep and mouflon impact mamane forest, completed. 2) Agency conducted hunts to eradicate feral sheep and goats, completed, a federal court order initiated this action. 3) Fence maintenance, ongoing. 4) Determining Critical Habitat, completed. 5) Surveys and research on Palila biology, ongoing with much accomplished. 6) Incorporation of other State lands with the Mauna Kea forest reserves, ongoing, initial contacts have been made by the U.S. Fish and Wildlife Service (USFWS). Management Practices - 3
                                   (DRAFT) - References
                                      Species PALILA
                                  Species Id ESIS101017
                                      Date 14 MAR 96



     

References

***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE ***** 01 Berger, A.J. 1981. Hawaiian birdlife, 2nd ed. Univ. Press of Hawaii, Honolulu. 02 Amadon, D. 1950. The Hawaiian honeycreepers (Aves:Drepaniidae). Bull. Amer. Mus. Nat. Hist. 95:151-262. 03 American Ornithologists' Union. 1983. Check-list of North American birds, 6th ed. Allen Press, Lawrence, KS. 04 Oustalet, M. 1877. Bull. Sci. Soc. Philom. Paris, Ser. 7,1. P. 100. 05 Banko, W.E. 1979. History of endemic Hawaiian bird specimens in museum collections. Avian History Report 2, Coop. Nat. Park. Res. Studies Unit, Univ. Hawaii, Honolulu. 06 Shallenberger, R. 1984. Hawaii's birds, 3rd rev. ed. Hawaii Audubon Soc., Honolulu. 07 Peterson, R.T. 1961. A field guide to western birds, 2nd rev. ed. Houghton Mifflin, Boston. 08 Munro, G.C. 1944. Birds of Hawaii. Tongg Publ. Co., Honolulu. 09 Rothschild, W. 1893-1900. The avifauna of Laysan and the neighboring islands. 3 Vols. R.H. Porter, London. 10 Wilson, S.B. and A.H. Evans. 1890-1899. Aves Hawaiienses: the birds of the Sandwich Islands. R.H. Porter, London. 11 Scott, J.M., S. Mountainspring, F.L. Ramsey, and C.B. Kepler. 1986. Forest bird communities of the Hawaiian Islands: Their dynamics, ecology, and conservation. Studies in Avian Biology 9. 12 Olson, S.L. and H.F. James. 1982. Prodromus of the fossil avifauna of the Hawaiian Islands. Smithsonian Contr. Zool. 365. 13 Scott, J.M., S. Mountainspring, C. van Riper III, C.B. Kepler, J.D. Jacobi, T.A. Burr, and J.G. Griffin. 1984. Annual variation in the distribution, abundance, and habitat response of the palila (Loxioides bailleui). Auk 101:647-664. 14 Tomich, P.Q. 1969. Mammals in Hawaii: A synopsis and notational bibliography. Spec. Publ. 57, B.P. Bishop Mus, Honolulu. 15 Atkinson, I.A.E. 1977. A reassessment of factors, particularly Rattus rattus L., that influenced the decline of endemic forest birds in the Hawaiian Islands. Pacific Sci. 31:109-133. 16 van Riper, C., III. 1980. The phenology of the dryland forest of Mauna Kea, Hawaii, and the impact of recent environmental perturbations. Biotropica 12:282-291. 17 Weathers, W.W. and C. van Riper III. 1982. Temperature regulation in two endangered Hawaiian honeycreepers: The palila (Psittirostra bailleui) and the Laysan finch (Psittirostra cantans). Auk 99:667-674. 18 van Riper, C., III. 1978. The breeding biology of the amakihi (Loxops virens) and palila (Psittirostra bailleui) on Mauna Kea, Hawaii. Ph.D. diss., Univ. Hawaii, Honolulu. 19 van Riper, C., III. 1980. Observations on the breeding of the palila, Psittirostra bailleui of Hawaii. Ibis 122:462-475. 20 Perkins, R.C.L. 1903. Vertebrata. Pages 365-466. IN: Fauna Hawaiiensis, Vol. 1, Part IV. D. Sharp, ed. The University Press, Cambridge, England. References - 1 (DRAFT) - References Species PALILA Species Id ESIS101017 Date 14 MAR 96 21 van Riper, C., III, J.M. Scott, and D.M. Woodside. 1978. Distribution and abundance patterns of the Palila on Mauna Kea, Hawaii. Auk 95:518-527. 22 Mountainspring, S. 1985. Personal observations. U.S. Fish and Wildlife Service, Mauna Loa Field Station. 23 Anon. [n.d.] File data. Mauna Loa Field Station, U.S. Fish and Wildlife Service. 24 van Riper, C., III. Unpublished data. Cooperative National Park Resources Studies Unit, University of California, Davis, CA. 25 Warner, R.E. 1968. The role of introduced diseases in the extinction of the endemic Hawaiian avifauna. Condor 70:101-120. 26 van Riper, C., III, S.G. van Riper, M.L. Goff, and M. Laird. 1982. The impact of malaria on birds in Hawaii Volcanoes National Park. Tech. Rept. 47, Coop. Nat. Res. Stud. Unit Univ. Hawaii, Honolulu. 27 van Riper, S.G. and C. van Riper, III. 1985. A summary of known parasites and diseases recorded from the avifauna of the Hawaiian Islands. Pages 298-371. IN: Hawaii's terrestrial ecosystems: Preservation and management. C.P. Stone and J.M. Scott, eds. Univ. Press Hawaii, Honolulu. 28 Laird, M. and C. van Riper, III. 1981. Questionable reports of Plasmodium from birds in Hawaii, with recognition of P. relictum ssp. capistranoae (Russel, 1932) as the avian parasite there. Pages 159-165. IN: Parasitological topics. E.V. Canning, ed. Soc. Protozool., Spec. Publ. 1. ***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY ***** 01 Berger, A.J. 1981. Hawaiian birdlife, 2nd ed. Univ. Press of Hawaii, Honolulu. 02 Olson, S.L. and H.F. James. 1982. Prodromus of the fossil avifauna of the Hawaiian Islands. Smithsonian Contrib. Zool. 365. 03 Scott, J.M., S. Mountainspring, F.L. Ramsey, and C.B. Kepler. 1986. Forest bird communities of the Hawaiian Islands: Their dynamics, ecology, and conservation. Studies in Avian Biology 9. 04 Scott, J.M., S. Mountainspring, C. van Riper III, C.B. Kepler, J.D. Jacobi, T.A. Burr, and J.G. Griffin. 1984. Annual variation in the distribution, abundance, and habitat responses of the palila (Loxioides bailleui). Auk 101: 647-664. 05 Anon. [n.d.] Field data files. Mauna Loa Field Station, U.S. Fish and Wildlife Service. 06 van Riper, C., III, J.M. Scott, and D.M. Woodside. 1978. Distribution and abundance patterns of the Palila on Mauna Kea, Hawaii. Auk 95:518-527. 07 Perkins, R.C.L. 1903. Vertebrata. Pages 365-466. IN: Fauna Hawaiiensis, Vol. 1, Part IV. D. Sharp, ed. The University Press, Cambridge, England. 08 Rothschild, W. 1893-1900. The avifauna of Laysan and the neighbouring islands. R.H. Porter, London. References - 2