Palila
Encyclopedia
The Palila is a critically endangered
Critically endangered
Version 2010.3 of the IUCN Red List of Threatened Species identified 3744 Critically Endangered species, subspecies and varieties, stocks and subpopulations.Critically Endangered by kingdom:*1993 Animalia*2 Fungi*1745 Plantae*4 Protista-References:...

 finch-billed species of Hawaiian honeycreeper
Hawaiian honeycreeper
Hawaiian honeycreepers are small, passerine birds endemic to Hawaii. Some authorities still categorize this group as a family Drepanididae, but in recent years, most authorities consider them a subfamily, Drepanidinae, of Fringillidae, the finch family...

. It has a golden-yellow head and breast, with a light belly, gray back, and greenish wings and tail. The bird has a close ecological
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

 relationship with the māmane
Mamane
Sophora chrysophylla, known as Māmane in Hawaiian, is a species of flowering plant in the pea and bean family, Fabaceae, that is endemic to Hawaii. It is highly polymorphic, growing as a shrub or tree, and able to reach a height of in tree form. Yellow flowers are produced in winter and...

 tree (Sophora chrysophylla), and became endangered due to destruction of the trees and accompanying dry forests
Hawaiian tropical dry forests
The Hawaiian tropical dry forests are a tropical dry broadleaf forest ecoregion in the Hawaiian Islands. They cover an area of on the leeward side of the main islands and the summits of Niihau and Kahoolawe. These forests are either seasonal or sclerophyllous. Annual rainfall is less than and...

.

Physical description

The palila has a yellow head and breast, with white to light gray plumage
Plumage
Plumage refers both to the layer of feathers that cover a bird and the pattern, colour, and arrangement of those feathers. The pattern and colours of plumage vary between species and subspecies and can also vary between different age classes, sexes, and season. Within species there can also be a...

 ventrally, medium gray plumage dorsal
Dorsum (biology)
In anatomy, the dorsum is the upper side of animals that typically run, fly, or swim in a horizontal position, and the back side of animals that walk upright. In vertebrates the dorsum contains the backbone. The term dorsal refers to anatomical structures that are either situated toward or grow...

ly, and olive
Olive
The olive , Olea europaea), is a species of a small tree in the family Oleaceae, native to the coastal areas of the eastern Mediterranean Basin as well as northern Iran at the south end of the Caspian Sea.Its fruit, also called the olive, is of major agricultural importance in the...

-green wings and tail. The wing
Wing
A wing is an appendage with a surface that produces lift for flight or propulsion through the atmosphere, or through another gaseous or liquid fluid...

s and the tails are greenish. The bird also has a heavy dark bill
Beak
The beak, bill or rostrum is an external anatomical structure of birds which is used for eating and for grooming, manipulating objects, killing prey, fighting, probing for food, courtship and feeding young...

 with swollen sides, a brown iris
Iris (anatomy)
The iris is a thin, circular structure in the eye, responsible for controlling the diameter and size of the pupils and thus the amount of light reaching the retina. "Eye color" is the color of the iris, which can be green, blue, or brown. In some cases it can be hazel , grey, violet, or even pink...

, and dark feet with yellowish sole
Sole
Sole may refer to:* Sole , the bottom of the foot* Sole , the bottom supporting member of the shoe-Business:* Sole proprietorship, a business which legally has no separate existence from its owner...

s. The palila is one of the largest living Hawaiian honeycreeper
Honeycreeper
The typical honeycreepers are small birds in the tanager family. They are found in the tropical New World from Mexico south to Brazil.They occur in the forest canopy, and, as the name implies, they are specialist nectar feeders with long curved bills....

s, measuring around 6–7.5 in (15.2–19.1 cm).

There is some sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...

. Males tend to have brighter colors overall, as well as clear-cut black lore
Lore
Lore may refer to:* Loré , a city and subdistrict in Lautém District* Lore , the region on each side of a birds face between eye and bill* Lore , a fictional android* Lore Sjöberg, an internet humourist...

s. The corresponding area contrasts less with the dirty-yellow heads in the marginally smaller females.

The bird's song is inconspicuous, containing whistling, warbling and trill
Trill
Trill may refer to:* Trill , a type of musical ornament* Trill consonant, a type of sound used in some languages*Trill, a type of bird food-Fiction:* Trill , two symbiotic races of aliens in the fictional Star Trek universe...

ing notes. The call is characteristic, however, being a clear, bell-like whistle
Whistle
A whistle or call is a simple aerophone, an instrument which produces sound from a stream of forced air. It may be mouth-operated, or powered by air pressure, steam, or other means...

, chee-clee-o or te-cleet. This is loudly communicated between birds advertising food during the morning and evening, and according to native informants, it is given most frequently during the day as rain approaches (Rothschild
Rothschild
Rothschild is a common German surname. It is a habitational name from a house distinguished with a red shield , the earliest recorded example dating from the 13th century...

 1900).

Systematics and nomenclature

The Hawaiian honeycreepers (Drepanididae) are sometimes included in the true finch
Finch
The true finches are passerine birds in the family Fringillidae. They are predominantly seed-eating songbirds. Most are native to the Northern Hemisphere, but one subfamily is endemic to the Neotropics, one to the Hawaiian Islands, and one subfamily – monotypic at genus level – is found...

 family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 (Fringillidae). Oustalet
Émile Oustalet
Jean-Frédéric Émile Oustalet was a French zoologist.Oustalet was born at Montbéliard, in the department of Doubs. He studied at the Ecole des Hautes-Etudes and his first scientific work was on the respiratory organs of dragonfly larvae...

 scientifically described the palila in 1877. Named Loxioides bailleui by him, it was for some times united with several other "parrot
Parrot
Parrots, also known as psittacines , are birds of the roughly 372 species in 86 genera that make up the order Psittaciformes, found in most tropical and subtropical regions. The order is subdivided into three families: the Psittacidae , the Cacatuidae and the Strigopidae...

-billed" Hawaiian honeycreeper species in [[ʻŌʻū|Psittirostra]]. Currently, the palila has again been moved to genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Loxioides
Loxioides
Loxioides is a genus of finches in the Hawaiian honeycreeper subfamily, Drepanidinae. It contains the following species:* Loxioides bailleui Oustalet, 1877 - Palila...

, which was long considered monotypic. The native
Hawaiian language
The Hawaiian language is a Polynesian language that takes its name from Hawaii, the largest island in the tropical North Pacific archipelago where it developed. Hawaiian, along with English, is an official language of the state of Hawaii...

 name ōū poopapale ("capped [[ʻŌʻū|ōū]]") probably refers to this species too (Rothschild 1900, FWIE 1996). Despite its bill and habits being somewhat similar to the ōū, its color pattern betrays a very close relationship with the genus Telespiza
Telespiza
Telespiza is a genus of finches in the Hawaiian honeycreeper subfamily, Drepanididae. All species in it are or were endemic to the Hawaiian Islands.-Species:* Telespiza cantans Wilson, 1890 - Laysan Finch...

.

Distribution and status

Currently, the palila can be found only on the upper slopes of Mauna Kea
Mauna Kea
Mauna Kea is a volcano on the island of Hawaii. Standing above sea level, its peak is the highest point in the state of Hawaii. However, much of the mountain is under water; when measured from its oceanic base, Mauna Kea is over tall—significantly taller than Mount Everest...

 on the island of Hawaii
Hawaii (island)
The Island of Hawaii, also called the Big Island or Hawaii Island , is a volcanic island in the North Pacific Ocean...

. Palila live from about 6500 to 9500 ft (1,981.2 to 2,895.6 m) AMSL. The population density
Population density
Population density is a measurement of population per unit area or unit volume. It is frequently applied to living organisms, and particularly to humans...

 of the bird increases in areas where māmane
Mamane
Sophora chrysophylla, known as Māmane in Hawaiian, is a species of flowering plant in the pea and bean family, Fabaceae, that is endemic to Hawaii. It is highly polymorphic, growing as a shrub or tree, and able to reach a height of in tree form. Yellow flowers are produced in winter and...

 (Sophora chrysophylla) grows more plentiful, and the birds do not appear to venture far from māmane stands. Essentially, this means that the species is confined - and may always have been so - to the area above the moist forest belt at around 3000–4500 ft (914.4–1,371.6 m).

Palila are today found in less than 10 percent of their historical range; they were found at elevations down to 4000 feet (1,219.2 m) as late as the 19th century. Loxioides bailleui was abundant throughout Hawaii until the beginning of the 20th century. It lived on the upper slopes of Mauna Kea, the northwest slopes of Mauna Loa
Mauna Loa
Mauna Loa is one of five volcanoes that form the Island of Hawaii in the U.S. state of Hawaii in the Pacific Ocean, and the largest on Earth in terms of volume and area covered. It is an active shield volcano, with a volume estimated at approximately , although its peak is about lower than that...

, and the eastern slopes of Hualālai
Hualalai
Hualālai is a dormant shield volcano on the island of Hawaii in the Hawaiian Islands. It is the third-youngest and the third most active of the five volcanoes that form the island of Hawaii, following Kīlauea and the much larger Mauna Loa, and also the westernmost. Its peak is above sea...

. Then, as early as 1944, scientists believed the bird almost extinct.

On March 11, 1967, the palila were listed as an endangered species
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

 under the ESA
Endangered Species Act
The Endangered Species Act of 1973 is one of the dozens of United States environmental laws passed in the 1970s. Signed into law by President Richard Nixon on December 28, 1973, it was designed to protect critically imperiled species from extinction as a "consequence of economic growth and...

. In 1975, it was estimated that only 1,614 palila existed. In 1978, the 9th Circuit Court of Appeals
United States Court of Appeals for the Ninth Circuit
The United States Court of Appeals for the Ninth Circuit is a U.S. federal court with appellate jurisdiction over the district courts in the following districts:* District of Alaska* District of Arizona...

 ruled that feral
Feral
A feral organism is one that has changed from being domesticated to being wild or untamed. In the case of plants it is a movement from cultivated to uncultivated or controlled to volunteer. The introduction of feral animals or plants to their non-native regions, like any introduced species, may...

 sheep
Domestic sheep
Sheep are quadrupedal, ruminant mammals typically kept as livestock. Like all ruminants, sheep are members of the order Artiodactyla, the even-toed ungulates. Although the name "sheep" applies to many species in the genus Ovis, in everyday usage it almost always refers to Ovis aries...

 and goat
Goat
The domestic goat is a subspecies of goat domesticated from the wild goat of southwest Asia and Eastern Europe. The goat is a member of the Bovidae family and is closely related to the sheep as both are in the goat-antelope subfamily Caprinae. There are over three hundred distinct breeds of...

s had to be removed from critical habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 of the bird. From annual counts between 1980 and 1996, variable estimates of population ranged from 1,584 to 5,685 mature birds, though there are no consistent trends. In 1997, the west slope of Mauna Kea contained 72% of the population. The entire population, an estimated 4,396 birds, occupied an estimated 78 square kilometres (19,274.2 acre) (BLI 2004).

Ecology and behavior

The palila favors māmane
Mamane
Sophora chrysophylla, known as Māmane in Hawaiian, is a species of flowering plant in the pea and bean family, Fabaceae, that is endemic to Hawaii. It is highly polymorphic, growing as a shrub or tree, and able to reach a height of in tree form. Yellow flowers are produced in winter and...

 and māmane-naio (Myoporum sandwicense
Myoporum sandwicense
Myoporum sandwicense is a species of flowering tree in the figwort family, Scrophulariaceae. Common names include Naio, bastard sandalwood or false sandalwood. It is native to Hawaii and Mangaia in the Cook Islands.-Description:...

) dry forests
Hawaiian tropical dry forests
The Hawaiian tropical dry forests are a tropical dry broadleaf forest ecoregion in the Hawaiian Islands. They cover an area of on the leeward side of the main islands and the summits of Niihau and Kahoolawe. These forests are either seasonal or sclerophyllous. Annual rainfall is less than and...

. A habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

 mix containing, apart from said forest, patches of grassland
Grassland
Grasslands are areas where the vegetation is dominated by grasses and other herbaceous plants . However, sedge and rush families can also be found. Grasslands occur naturally on all continents except Antarctica...

s, pūkiawe (Styphelia
Styphelia
Styphelia is a genus of shrubs in the heather family, Ericaceae. The genus is endemic to Australia and the Pacific Islands.Species include:*Styphelia adscendens R.Br. – Golden Heath...

 tameiameiae
Styphelia tameiameiae
Styphelia tameiameiae, known as Pūkiawe in the Hawaiian language, is a species of flowering plant in the heather family, Ericaceae, that is native to the Hawaiian and Marquesas Islands. The specific epithet honors King Kamehameha I, who formed the Kingdom of Hawaii. It grows as a tree up to tall...

) shrubland on lava fields
Lava plain
A lava plain, also called a lava field or lava bed, is a large expanse of nearly flat-lying lava flows. Such features are generally composed of highly-fluid basalt lava, and can extend for tens or even hundreds of miles across the underlying terrain...

, and other types of native understory
Understory
Understory is the term for the area of a forest which grows at the lowest height level below the forest canopy. Plants in the understory consist of a mixture of seedlings and saplings of canopy trees together with understory shrubs and herbs...

 vegetation is optimal for their survival.

The diet of the palila is almost exclusively the immature seeds of māmane when these are available. These contain much vile-tasting phenol
Natural phenol
Natural phenols, bioavailable phenols, plant phenolics, low molecular weight phenols or phenoloids are a class of natural products. They are small molecules containing one or more phenolic group. These molecules are smaller in size than polyphenols, containing less than 12 phenolic groups...

ic compounds in the seed coat and a lethal amount of quinolizidine
Quinolizidine
Quinolizidine is a nitrogen-containing heterocyclic compound. Some alkaloids are derivatives of quinolizidine....

 alkaloid
Alkaloid
Alkaloids are a group of naturally occurring chemical compounds that contain mostly basic nitrogen atoms. This group also includes some related compounds with neutral and even weakly acidic properties. Also some synthetic compounds of similar structure are attributed to alkaloids...

s in the embryos themselves. By some undetermined means, adult palila are able to cope with a dose of these toxin
Toxin
A toxin is a poisonous substance produced within living cells or organisms; man-made substances created by artificial processes are thus excluded...

s that would kill other small animals in mere minutes. The amount of toxin in māmane varies, and the palila can be seen to avoid certain trees. It is possible that these contain the highest amounts of poison, but how the birds would be able to recognize this is not known (Banko et al. 2002).

The bitter taste of the seed coats probably does not affect the birds (see below). Nonetheless, the seed coats are not very nutritious, and are thus discarded. Palila bills are adapted to open Fabales
Fabales
Fabales is an order of flowering plants. It is included in the rosid group of the eudicots in the Angiosperm Phylogeny Group II classification system...

 pods. The birds hold the pod with one foot and pry it open with the bill to expose the seeds. They then tear away the visible portion of the seed coat and extract the embryo, leaving the remaining coat in the pod. Seeds that drop out of the pod intact during opening are picked up and positioned longitudinally
Anatomical terms of location
Standard anatomical terms of location are designations employed in science that deal with the anatomy of animals to avoid ambiguities that might otherwise arise. They are not language-specific, and thus require no translation...

 in the bill. The seed coat is then neatly cut open by the bill's edge and the embryo nudged out with the bird's tongue. The seed coat, still remaining in one piece, is then dropped.(Banko et al. 2002)

Palila also eat naio berries and other fruit (such as the introduced Cape gooseberry: Rothschild 1900), and māmane flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

s, buds, and young leaves. Additionally, they feed on caterpillar
Caterpillar
Caterpillars are the larval form of members of the order Lepidoptera . They are mostly herbivorous in food habit, although some species are insectivorous. Caterpillars are voracious feeders and many of them are considered to be pests in agriculture...

s, particularly those of Cydia species (māmane codling moths) and more rarely on those of Uresiphita polygonalis virescens
Uresiphita polygonalis
Uresiphita polygonalis is a moth of the Crambidae family. It is found in the Pacific, including Hawaii and New Zealand.The wingspan is about 27 mm. Adults range in color from green to red....

 (māmane snout moth
Pyraustinae
Pyraustinae is a large subfamily of the lepidopteran family Crambidae, the crambid snout moths. It currently includes over 1,400 species, the majority of them tropical but some found in temperate regions including both North America and Europe....

). These caterpillars as well as other insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s, along with the very nutritious māmane seeds, provide the palila's main source of protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

. Nestlings, apparently not yet able to cope with the amount of poison contained in the seeds, are fed to a large extent on Cydia caterpillars. These destroy or discard the māmanes toxins they take up with their food, so that the caterpillars themselves are non-toxic. They do contain high amounts of phenolic compounds they probably sequester from their food and quite likely taste as bad. Palila do not seem to mind the adverse taste or are physically unable to perceive it, given that they go to great lengths to obtain this food during breeding season. (Banko et al. 2002)

The abundance of māmane seeds affects reproduction
Biological reproduction
Reproduction is the biological process by which new "offspring" individual organisms are produced from their "parents". Reproduction is a fundamental feature of all known life; each individual organism exists as the result of reproduction...

 rates and adult survival. Palila start to eat the seeds at higher elevations and then gradually move downslope. During drought
Drought
A drought is an extended period of months or years when a region notes a deficiency in its water supply. Generally, this occurs when a region receives consistently below average precipitation. It can have a substantial impact on the ecosystem and agriculture of the affected region...

s, when māmane seeds are scarce, most birds do not attempt to breed
Breed
A breed is a group of domestic animals or plants with a homogeneous appearance, behavior, and other characteristics that distinguish it from other animals or plants of the same species. Despite the centrality of the idea of "breeds" to animal husbandry, there is no scientifically accepted...

.

The birds normally breed from February to September. The female constructs a loose, cup-shaped nest
Nest
A nest is a place of refuge to hold an animal's eggs or provide a place to live or raise offspring. They are usually made of some organic material such as twigs, grass, and leaves; or may simply be a depression in the ground, or a hole in a tree, rock or building...

 around 4 inches (10.2 cm) in diameter high up in a māmane or naio tree. For this it uses grass
Grass
Grasses, or more technically graminoids, are monocotyledonous, usually herbaceous plants with narrow leaves growing from the base. They include the "true grasses", of the Poaceae family, as well as the sedges and the rushes . The true grasses include cereals, bamboo and the grasses of lawns ...

es, stem
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...

s, root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...

s, lichen
Lichen
Lichens are composite organisms consisting of a symbiotic organism composed of a fungus with a photosynthetic partner , usually either a green alga or cyanobacterium...

, and branch bark
Bark
Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines and shrubs. Bark refers to all the tissues outside of the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner...

 from the māmane trees provide the building material. Lichen and small leaves
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 layer the inside of the nest. Usually the palila clutch size is two eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

. Both parents regurgitate
Regurgitation (digestion)
Regurgitation is the expulsion of material from the mouth, pharynx, or esophagus, usually characterized by the presence of undigested food or blood.Regurgitation is used by a number of species to feed their young...

 food to feed their young. The juveniles
Juvenile (organism)
A juvenile is an individual organism that has not yet reached its adult form, sexual maturity or size. Juveniles sometimes look very different from the adult form, particularly in terms of their colour...

 remain in the nest for up to 31 days before fledging
Fledge
Fledge is the stage in a young bird's life when the feathers and wing muscles are sufficiently developed for flight. It also describes the act of a chick's parents raising it to a fully grown state...

.

Legal background

The palila was the first animal to have a 9th circuit
United States Court of Appeals for the Ninth Circuit
The United States Court of Appeals for the Ninth Circuit is a U.S. federal court with appellate jurisdiction over the district courts in the following districts:* District of Alaska* District of Arizona...

 federal case cited in its own name. Prior to Palila v. Hawaii Department of Land and Natural Resources
Palila v. Hawaii Department of Land and Natural Resources
Palila v. Hawaii Department of Land and Natural Resources was an ecological court case pertaining to the Palila and the Māmane-Naio ecosystem of Mauna Kea. The case stems from the introduction of goats and sheep onto Hawaii island in the late 18th century, which became feral and damaged the local...

 852 F.2d 1106 (U.S. 1988), cases were cited under the represented party e.g. Lujan v. Defenders of Wildlife
Lujan v. Defenders of Wildlife
Lujan v. Defenders of Wildlife, , was a United States Supreme Court case decided on June 12, 1992, in which the court held that a group of American wildlife conservation and other environmental organizations lacked standing to challenge regulations jointly issued by the U.S...

 504 U.S. 555 (U.S. 1992), this opened a door for environmental protection agencies who in prior years had lost on issue of standing, e.g. Lujan.

Counsel: For PALILA, Plaintiff - Appellee: Michael R. Sherwood, Esq., Earthjustice
Earthjustice
Earthjustice is a non-profit public interest law firm based in the United States that specializes in cases protecting natural resources, safeguarding public health, and promoting clean energy...

 Legal Defense Fund, San Francisco, California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

.

Judges: Before: Schroeder, Noonan, and O'Scannlain, Circuit Judges.

External links

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