Sophora Chrysophylla
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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 (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
in the pea and bean family, Fabaceae, that is
endemic Endemism is the state of a species being found 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 found else ...
to
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only state ...
. It is highly polymorphic, growing as a shrub 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, including only woody plants with secondary growth, plants that are ...
, and able to reach a height of in tree form. Yellow
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are produced in winter and spring.


Biology

''S. chrysophylla'' has ridged golden brown branches. The tree has pinnately compound leaves with 6 to 10 pairs of leaflets. Each leaflet is long and wide. Leaves are smooth, or with gray or yellow hairs on the underside. The specific name is derived from the
Greek Greek may refer to: Greece 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 ...
words χρυσός (''chrysós''), meaning "
gold Gold is a chemical element with the symbol Au (from la, aurum) and atomic number 79. This makes it one of the higher atomic number elements that occur naturally. It is a bright, slightly orange-yellow, dense, soft, malleable, and ductile me ...
," and φυλλον (''phyllos''), meaning "leaf." Flowers are found at the bases of leaves or the ends of branches in clusters – that is, they occur in axillary or terminal racemes. The corolla is yellow. The petal 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 A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s, which are long. Untreated, the seeds have germination rates of less than 5%. The tree is
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...
and highly polymorphic.


Habitat

''Māmane'' is an endemic species of Hawaii, and can be found on all main islands except
Niihau Niihau ( Hawaiian: ), anglicized as Niihau ( ), is the westernmost main and seventh largest inhabited island in Hawaii. It is southwest of Kauaʻi across the Kaulakahi Channel. Its area is . Several intermittent playa lakes provide wetland ha ...
and
Kahoolawe Kahoolawe (Hawaiian: ), anglicized as Kahoolawe (), is the smallest of the eight main volcanic islands in the Hawaiian Islands. Kahoolawe is located about southwest of Maui and also southeast of Lānaʻi, and it is long by wide, with a total l ...
. 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 the habitat at which trees are capable of growing. It is found at high elevations and high latitudes. Beyond the tree line, trees cannot tolerate the environmental conditions (usually cold temperatures, extreme snow ...
. ''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 transition area between two biological communities, where two communities meet and integrate. It may be narrow or wide, and it may be local (the zone between a field and forest) or regional (the transition between forest and gras ...
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 (
Haleakalā Haleakalā (; Hawaiian: ), or the East Maui Volcano, is a massive shield volcano that forms more than 75% of the Hawaiian Island of Maui. The western 25% of the island is formed by another volcano, Mauna Kahalawai, also referred to as the West ...
) and the
Island of Hawaii Hawaii ( ; haw, Hawaii ) is the largest island in the United States, located in the state of Hawaii. It is the southeasternmost of the Hawaiian Islands, a chain of volcanic islands in the North Pacific Ocean. With an area of , it has 63% of th ...
.


Uses

The hard, durable
wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...
of ''māmane'' was used by the Native Hawaiians for ''pou'' (house posts) and ''kaola'' (beams) up to in diameter, ''ōō'' (
digging stick A digging stick, sometimes called a yam stick, is a wooden implement used primarily by subsistence-based cultures to dig out underground food such as roots and tubers, tilling the soil, or burrowing animals and anthills. It is a term used in a ...
s),
spear A spear is a pole weapon 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 hardened spears, or it may be made of a more durable material fasten ...
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 (; alternative spelling: 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 ...
handles), and ''wahie'' ( firewood).
Cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
ranch A ranch (from es, rancho/Mexican Spanish) is an area of land, including various structures, given primarily to ranching, the practice of raising grazing livestock such as cattle and sheep. It is a subtype of a farm. These terms are most ofte ...
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 netting. A fence differs from a wall in not having a solid foundation along its whole length. ...
posts. In herbal medicine, the
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
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". Calamine lotion, witch hazel, and yerba mansa, a Californian pla ...
. The wood was also used in
religious Religion is usually defined as a social- cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatur ...
ritual A ritual is a sequence of activities involving gestures, words, actions, or objects, performed according to a set sequence. Rituals may be prescribed by the traditions of a community, including a religious community. Rituals are characterized ...
s to ward off evil. A '' kahuna nui'' (high priest) would wrap a piece of ''māmane'' wood in a dark ''
kapa Kapa is a fabric made by native Hawaiians from the bast fibres of certain species of trees and shrubs in the orders Rosales and Malvales. Description and uses It is similar to ''tapa'' found elsewhere in Polynesia (the Hawaiian phoneme corr ...
'' cloth and hold it up to symbolize authority.


Ecology and conservation

''Māmane'' is essential for the endangered ''
palila The palila (''Loxioides bailleui'') is a critically endangered finch-billed species of Hawaiian honeycreeper. 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 rel ...
'' (''Loxioides bailleui''), which feeds almost exclusively on the plants' immature
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s when these are in season. It also nests in the ''māmane'' branches. Caterpillars of ''
Cydia Cydia is a graphical user interface of APT (Advanced Package Tool) for iOS. It enables a user to find and install software not authorized by Apple on jailbroken iPhones, iPads and iPod Touch devices. It also refers to digital distribution pl ...
'' moths also eat the ''māmanes seeds, and in turn are being eaten by the ''palila''. Both the bird and the larvae utilize the embryo only, leaving the seed coat untouched. To other animals, ''māmane'' seeds are highly poisonous.
House finch The house finch (''Haemorhous mexicanus'') is a bird in the finch family Fringillidae. It is native to western North America and has been introduced to the eastern half of the continent and Hawaii. This species and the other two American rosefi ...
es 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 called carbolic acid) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile. The molecule consists of a phenyl group () bonded to a hydroxy group (). Mildly acidic, it ...
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 subs ...
and have a high fibre content.
Seed predator Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source,Hulme, P.E. and Benkman, C.W. (2002) "Granivory", pp. 13 ...
s 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 (U.S. English) or kerb ...
) of
quinolizidine alkaloids Quinolizidine alkaloids are natural products that have a quinolizidine structure; this includes the lupine alkaloids. Occurrence Quinolizidine alkaloids can be found in the plant family of legumes, especially in papilionaceous plants. While t ...
. For example
cytisine Cytisine, also known as baptitoxine, cytisinicline, or sophorine, is an alkaloid that occurs naturally in several plant genera, such as ''Laburnum'' and ''Cytisus'' of the family Fabaceae. It has been used medically to help with smoking cessat ...
, anagyrine, 5,6-dehydrolupanine, lupanine, N-methylcytisine, and baptifoline (Banko ''et al.'' 2002).
The ''palila'' and the moths, however, have
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
the ability to deal with the poisonous compounds. The ''palila'', for example, can deal with dozens of times the dose of
cytisine Cytisine, also known as baptitoxine, cytisinicline, or sophorine, is an alkaloid that occurs naturally in several plant genera, such as ''Laburnum'' and ''Cytisus'' of the family Fabaceae. It has been used medically to help with smoking cessat ...
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 mice are usually of the species '' Mus musculus''. They are the most commonly ...
. 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 heritable traits characteristic of a population over generations. Cha ...
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 aegagrus 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 goats, all of which stem from the wild goat ...
s and
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals 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 domesticated ...
voraciously eat the seedpods of the plant, negatively impacting the tree's
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
. 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 by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste o ...
coloration. Clearly, this issue requires further study. Ranch
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
can kill trees through stomping on the
root In vascular plants, the roots are the 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 below the su ...
s.
Wildfire A wildfire, forest fire, bushfire, wildland fire or rural fire is an unplanned, uncontrolled and unpredictable fire in an area of combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identi ...
has also destroyed some of the trees, though it is generally resistant to fire, and can grow quickly after one occurs. As regards diseases, the canker ascomycete ''
Botryosphaeria ''Botryosphaeria'' is a genus of pathogenic fungi in the family Botryosphaeriaceae. There are 193 species, many of which are important disease-causing agents of various important agricultural crops. Species *'' Botryosphaeria abietina'' *'' Bot ...
mamane'' causes
witch's broom Witch's broom or witches' broom is a deformity in a woody plant, typically a tree, where the natural structure of the plant is changed. A dense mass of shoots grows from a single point, with the resulting structure resembling a broom or a ...
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 ...
'' 89(2): 298–303. (HTML abstract and first page image)


Footnotes


External links


Plant ProfileHawaiian 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