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''Sophora chrysophylla'', known as ''māmane'' in Hawaiian, is a species of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
in the
pea Pea (''pisum'' in Latin) is a pulse or fodder crop, but the word often refers to the seed or sometimes the pod of this flowering plant species. Peas are eaten as a vegetable. Carl Linnaeus gave the species the scientific name ''Pisum sativum' ...
and
bean A bean is the seed of some plants in the legume family (Fabaceae) used as a vegetable for human consumption or animal feed. The seeds are often preserved through drying (a ''pulse''), but fresh beans are also sold. Dried beans are traditi ...
family,
Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...
, that is
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to
Hawaii Hawaii ( ; ) is an island U.S. state, state of the United States, in the Pacific Ocean about southwest of the U.S. mainland. One of the two Non-contiguous United States, non-contiguous U.S. states (along with Alaska), it is the only sta ...
. It is highly polymorphic, growing as a
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
or
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only ...
, and able to reach a height of in tree form. Yellow
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s are produced in winter and spring.


Biology

''S. chrysophylla'' has ridged golden brown branches. The tree has pinnately compound
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
with 6 to 10 pairs of leaflets. Each leaflet is long and wide. Leaves are smooth, or with gray or yellow
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and ...
s on the underside. The specific name is derived from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
words χρυσός (''chrysós''), meaning "
gold Gold is a chemical element; it has chemical symbol Au (from Latin ) and atomic number 79. In its pure form, it is a brightness, bright, slightly orange-yellow, dense, soft, malleable, and ductile metal. Chemically, gold is a transition metal ...
," and φυλλον (''phyllos''), meaning "leaf." Flowers are found at the bases of leaves or the ends of branches in
clusters may refer to: Science and technology Astronomy * Cluster (spacecraft), constellation of four European Space Agency spacecraft * Cluster II (spacecraft), a European Space Agency mission to study the magnetosphere * Asteroid cluster, a small ...
– that is, they occur in axillary or terminal
raceme A raceme () or racemoid is an unbranched, indeterminate growth, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are ...
s. The corolla is yellow. The
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
size ranges from long, and wide. The tree blooms in winter and spring. The height of the flowering season is in mid-spring. ''Māmane'' wood is dense, hard and durable. Seedpods are persistent, and remain on the tree for most of the year. They are twisted, brown to brownish-gray, have four wings and are long and usually wide. Seedpods are tightly constricted around the yellow-orange or brown to grayish-black
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, which are long. Untreated, the seeds have
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, ...
rates of less than 5%. The tree is
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
and highly polymorphic.


Habitat

''Māmane'' is an endemic species of Hawaii, and can be found on all main islands except
Niihau Niihau (Hawaiian language, Hawaiian: ), anglicized as Niihau ( ), is the seventh largest island in Hawaii and the westernmost of the main islands. It is southwest of Kauai, Kauai across the Channels of the Hawaiian Islands#Kaulakahi Channel, Ka ...
and Kahoolawe. It inhabits low shrublands, high shrublands,
dry forests The tropical and subtropical dry broadleaf forest is a habitat type defined by the World Wide Fund for Nature and is located at tropical and subtropical latitudes. Though these forests occur in climates that are warm year-round, and may receive ...
, mixed mesic forests, and, rarely, wet forests. It can grow at elevations of , being limited by the
tree line The tree line is the edge of a habitat at which trees are capable of growing and beyond which they are not. It is found at high elevations and high latitudes. Beyond the tree line, trees cannot tolerate the environmental conditions (usually low ...
. ''Māmane'' is most common and grows the tallest in montane dry forests at elevations of . ''Māmane'' and ''naio'' (''
Myoporum sandwicense ''Myoporum sandwicense'', commonly known as ''naio'', bastard sandalwood or false sandalwood is a species of flowering plant in the figwort family, Scrophulariaceae. It is a tree or shrub highly variable in its form, the size and shape of its le ...
'') define a dry woodland
ecotone An ecotone is a transitional area between two plant communities, where these meet and integrate. Examples include areas between grassland and forest, estuaries and lagoon, freshwater and sea water etc. An ecotone may be narrow or wide, and it ma ...
on the
subalpine Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
areas of East
Maui Maui (; Hawaiian language, Hawaiian: ) is the second largest island in the Hawaiian archipelago, at 727.2 square miles (1,883 km2). It is the List of islands of the United States by area, 17th-largest in the United States. Maui is one of ...
(
Haleakalā Haleakalā (; Hawaiian: ), or the East Maui Volcano, is a massive, active shield volcano that forms more than 75% of the Hawaiian Islands, Hawaiian Island of Maui. The western 25% of the island is formed by another volcano, Mauna Kahalawai, als ...
) and the
Island of Hawaii Hawaii is the List of islands of the United States by area, largest island in the United States, located in the Hawaii, state of Hawaii, the southernmost state in the union. It is the southeasternmost of the Hawaiian Islands, a chain of volcani ...
.


Uses

The hard, durable
wood Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...
of ''māmane'' was used by the
Native Hawaiians Native Hawaiians (also known as Indigenous Hawaiians, Kānaka Maoli, Aboriginal Hawaiians, or simply Hawaiians; , , , and ) are the Indigenous Polynesian people of the Hawaiian Islands. Hawaiʻi was settled at least 800 years ago by Polynesian ...
for ''pou'' (house posts) and ''kaola'' (beams) up to in diameter, ''ōō'' ( digging sticks),
spear A spear is a polearm consisting of a shaft, usually of wood, with a pointed head. The head may be simply the sharpened end of the shaft itself, as is the case with Fire hardening, fire hardened spears, or it may be made of a more durable materia ...
s, ''kope'' (
spade A spade is a tool primarily for digging consisting of a long handle and blade, typically with the blade narrower and flatter than the common shovel. Early spades were made of riven wood or of animal bones (often shoulder blades). After the a ...
s), '' papa hōlua'' (
sled A sled, skid, sledge, or sleigh is a land vehicle that slides across a surface, usually of ice or snow. It is built with either a smooth underside or a separate body supported by two or more smooth, relatively narrow, longitudinal runners ...
) runners, ''papa olonā'' ('' Touchardia latifolia'' scrapers), ''au koi'' (
adze An adze () or adz is an ancient and versatile cutting tool similar to an axe but with the cutting edge perpendicular to the handle rather than parallel. Adzes have been used since the Stone Age. They are used for smoothing or carving wood in ha ...
handles), and ''wahie'' (
firewood Firewood is any wooden material that is gathered and used for fuel. Generally, firewood is not heavily processed, and is in some sort of firelog, recognizable log or branch form, compared to other forms of wood fuel like pellet fuel, pellets. ...
).
Cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
ranch A ranch (from /Mexican Spanish) is an area of landscape, land, including various structures, given primarily to ranching, the practice of raising grazing livestock such as cattle and sheep. It is a subtype of farm. These terms are most often ap ...
ers used it as
fence A fence is a structure that encloses an area, typically outdoors, and is usually constructed from posts that are connected by boards, wire, rails or net (textile), netting. A fence differs from a wall in not having a solid foundation along its ...
posts. In
herbal medicine Herbal medicine (also called herbalism, phytomedicine or phytotherapy) is the study of pharmacognosy and the use of medicinal plants, which are a basis of traditional medicine. Scientific evidence for the effectiveness of many herbal treatments ...
, the
flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s are used as an
astringent An astringent (sometimes called adstringent) is a chemical that shrinks or constricts body tissues. The word derives from the Latin '' adstringere'', which means "to bind fast". Astringency, the dry, puckering or numbing mouthfeel caused by t ...
. The wood was also used in
religious Religion is a range of social- cultural systems, including designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relate humanity to supernatural ...
ritual A ritual is a repeated, structured sequence of actions or behaviors that alters the internal or external state of an individual, group, or environment, regardless of conscious understanding, emotional context, or symbolic meaning. Traditionally ...
s to ward off evil. A '' kahuna nui'' (high priest) would wrap a piece of ''māmane'' wood in a dark '' kapa'' cloth and hold it up to symbolize authority.


Ecology and conservation

''Māmane'' is essential for the endangered '' palila'' (''Loxioides bailleui''), which feeds almost exclusively on the plants' immature
seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s when these are in season. It also nests in the ''māmane'' branches.
Caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
s of ''
Cydia Cydia is a graphical user interface of APT for iOS. It enables a user to find and install software unauthorized by Apple on jailbroken iPhones, iPads and iPod Touch devices. It also refers to the digital distribution platform for software on ...
'' moths also eat the ''māmanes seeds, and in turn are being eaten by the ''palila''. Both the bird and the larvae utilize the
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
only, leaving the seed coat untouched. To other animals, ''māmane'' seeds are highly poisonous. House finches die within minutes after eating the seeds. The ''māmane'' employs a two-layered biochemical defence system: The seed coats contain some 4%
phenol Phenol (also known as carbolic acid, phenolic acid, or benzenol) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile and can catch fire. The molecule consists of a phenyl group () ...
ic compounds, which give them a vile taste. They are also somewhat
toxic Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subst ...
and have a high
fibre Fiber (spelled fibre in British English; from ) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorp ...
content. Seed predators trying to eat the seeds will probably not be killed but at least have a nasty experience. Should a predator be able to penetrate the seed coat, the embryo contains deadly quantities (>4% of
dry weight Vehicle weight is a measurement of wheeled motor vehicles; either an actual measured weight of the vehicle under defined conditions or a gross weight rating for its weight carrying capacity. Curb or kerb weight Curb weight (American English) or k ...
) of quinolizidine alkaloids. For example cytisine, anagyrine, 5,6-dehydrolupanine, lupanine, N-methylcytisine, and baptifoline (Banko ''et al.'' 2002). The ''palila'' and the moths, however, have
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
the ability to deal with the poisonous compounds. The ''palila'', for example, can deal with dozens of times the dose of cytisine that would kill a
laboratory mouse The laboratory mouse or lab mouse is a small mammal of the order Rodentia which is bred and used for scientific research or feeders for certain pets. Laboratory animal sources for these mice are usually of the species ''Mus musculus''. They ...
. Both seed predators seem to be able to recognize and avoid the most poisonous trees, which would be
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
in action. ''Cydia'' caterpillars are able to break up the toxic compounds. They do not sequester the alkaloids for their own use, but are found to contain about as much phenolic compounds as the seed coats do. This seems not to be sufficient to deter predators however as they are also cryptically colored. The ''palila'' is apparently impervious to the phenolic aroma (as they eat ''Cydia'' caterpillars which would smell and taste like the plant), discarding the seed coats due to their low nutritional value. How the ''palila'' deals with the toxins is not known.(Banko ''et al.'' 2002)
Feral goat The feral goat is the domestic goat (''Capra hircus'') when it has become established in the wild. Feral goats occur in many parts of the world. Species Feral goats consist of many breeds of domestic goats, all of which stem from the wild go ...
s and
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
voraciously eat the seedpods of the plant, negatively impacting the tree's
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
. Toxicity of the leaves is unknown; it is notable that '' Uresiphita polygonalis virescens'' caterpillars which feed on the leaves have
aposematic Aposematism is the Advertising in biology, advertising by an animal, whether terrestrial or marine, to potential predation, predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the pr ...
coloration. Clearly, this issue requires further study. Ranch
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
can kill trees through stomping on the
root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...
s.
Wildfire A wildfire, forest fire, or a bushfire is an unplanned and uncontrolled fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identified as a ...
has also destroyed some of the trees, though it is generally resistant to fire, and can grow quickly after one occurs. As regards
diseases A disease is a particular abnormal condition that adversely affects the structure or function of all or part of an organism and is not immediately due to any external injury. Diseases are often known to be medical conditions that are asso ...
, the
canker A plant canker is a small area of dead tissue, which grows slowly, often over years. Some cankers are of only minor consequence, but others are ultimately lethal and therefore can have major economic implications for agriculture and horticultur ...
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
'' Botryosphaeria mamane'' causes witch's broom disease, which kills heavily-infected trees (Gardner 1997). ''Māmane'' grows well in areas where there are no sheep or goats, and cattle populations are limited. The plant is being reforested in order to increase the population of both the tree itself and the ''palila'', and revive the indigenous ecosystem.


References

* Banko, P.; Cipollini, M.L.; Breton, G.; Paulk, E.; Wink, M. & Izhaki, I. (2002): Seed chemistry of ''Sophora chrysophylla'' (Mamane) in relation to the diet of the specialist seed predator ''Loxioides bailleui'' (Palila) in Hawaii. ''Journal of Chemical Ecology'' 28(7): 1393–1410. PDF fulltext
* Gardner, D.E. (1997): ''Botryosphaeria mamane'' sp. nov. associated with witches'-brooms on the endemic forest tree ''Sophora chrysophylla'' in Hawaii. ''
Mycologia ''Mycologia'' is a peer-reviewed scientific journal that publishes papers on all aspects of the fungi, including lichens. It first appeared as a bimonthly journal in January 1909, published by the New York Botanical Garden under the editorship of W ...
'' 89(2): 298–303. (HTML abstract and first page image)


Footnotes


External links


Plant Profile

Hawaiian Plants – Mamane
{{Taxonbar, from=Q2064969 chrysophylla Endemic flora of Hawaii Trees of Hawaii Biota of Hawaii (island) Biota of Kauai Biota of Lanai Biota of Maui Biota of Molokai Biota of Oahu Plants described in 1865 Taxa named by Berthold Carl Seemann