''Amaranthus brownii'' was an

annual Annual may refer to: * Annual publication, periodical publications appearing regularly once per year **Yearbook ** Literary annual * Annual plant * Annual report * Annual giving * Annual, Morocco, a settlement in northeastern Morocco * Annuals (b ...

herb In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal ...

in the family

Amaranthaceae Amaranthaceae is a family of flowering plants commonly known as the amaranth family, in reference to its type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making it ...

. The plant was found only on the small island of

Nihoa Nihoa (; haw, Nīhoa ), also known as Bird Island or Moku Manu, is the tallest of ten islands and atolls in the uninhabited Northwestern Hawaiian Islands (NWHI). The island is located at the southern end of the NWHI chain, southeast of N ...

in the

Northwestern Hawaiian Islands The Northwestern Hawaiian Islands or Leeward Hawaiian Islands are a series of islands and atolls in the Hawaiian island chain located northwest (in some cases, far to the northwest) of the islands of Kauai and Niihau. Politically, they are all p ...

, growing on rocky

outcrop An outcrop or rocky outcrop is a visible exposure of bedrock or ancient superficial deposits on the surface of the Earth. Features Outcrops do not cover the majority of the Earth's land surface because in most places the bedrock or superficial ...

s at altitudes of . It was one of nine species of ''

Amaranthus ''Amaranthus'' is a cosmopolitan genus of annual or short-lived perennial plants collectively known as amaranths. Some amaranth species are cultivated as leaf vegetables, pseudocereals, and ornamental plants. Catkin-like cymes of densely pac ...

'' in the

Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...

, as well as the only

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 elsew ...

Hawaiian species of the genus. It is now considered

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

. ''A. brownii'' was first discovered during the

Tanager Expedition The ''Tanager'' Expedition was a series of five biological surveys of the Northwestern Hawaiian Islands conducted in partnership between the Bureau of Biological Survey and the Bishop Museum, with the assistance of the United States Navy. Four ex ...

in 1923 by botanist Edward Leonard Caum. It differed from other Hawaiian species of ''Amaranthus'' with its spineless leaf axils, linear leaves, and

indehiscent Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that op ...

fruits. It was one of 26 vascular plants on Nihoa, 17 of which are indigenous, six alien, and three endemic only to Nihoa; these latter three included ''A. brownii'', the Nihoa fan palm or ''loulu'', and the Nihoa carnation. ''A. brownii'' was considered the rarest plant on Nihoa and was not directly observed on the island after 1983. Past expeditions collected plant samples and seeds, but no specimens managed to survive

ex-situ conservation Svalbard GLOBAL SEED BANK, an ''ex situ'' conservation. ''Ex situ'' conservation literally means, "off-site conservation". It is the process of protecting an endangered species, variety or breed, of plant or animal outside its natural habitat; ...

efforts outside of its native habitat. Consequently, there are no known plants or seeds from ''A. brownii'' in any botanical garden. Conservation and recovery plans for ''A. brownii'' were proposed by the

United States Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...

(FWS), which administers the island of Nihoa as part of the

Hawaiian Islands National Wildlife Refuge Hawaiian may refer to: * Native Hawaiians, the current term for the indigenous people of the Hawaiian Islands or their descendants * Hawaii state residents, regardless of ancestry (only used outside of Hawaii) * Hawaiian language Historic uses * ...

in the

Papahānaumokuākea Marine National Monument The Papahānaumokuākea Marine National Monument (PMNM) (roughly ) is a World Heritage Site, World Heritage listed National Monument (United States), U.S. National Monument encompassing of ocean waters, including ten islands and atolls of th ...

. In 1996, the plant was listed by the FWS as an endangered species. In 2003, the FWS designated the island of Nihoa as a

critical habitat Critical habitat is a habitat area essential to the conservation of a listed species, though the area need not actually be occupied by the species at the time it is designated. This is a specific term and designation within the U.S. Endangered Spec ...

for the plant and it was classified as critically endangered on the IUCN Red List. Following a lack of sightings for over 35 years despite intensive surveys, the species was classified as extinct on the IUCN Red List in 2018.

Taxonomy

The species was first collected during a ten-day visit to the island of Nihoa by the

. Botanist Edward Leonard Caum collected the first specimen on June 17, 1923, and a second was collected by cartographer Charles S. Judd on June 20, 1923. Forest B. H. Brown, botanist of the Bayard Dominick Expedition to the

Marquesas Islands The Marquesas Islands (; french: Îles Marquises or ' or '; Marquesan: ' ( North Marquesan) and ' ( South Marquesan), both meaning "the land of men") are a group of volcanic islands in French Polynesia, an overseas collectivity of France in th ...

(1921–1922), helped provide descriptions and comments for some of the species described by

Erling Christophersen Erling Christophersen (April 17, 1898 – November 9, 1994) was a Norwegian botanist, geographer and diplomat. He participated in and led several notable scientific expeditions in the 20th century, including the fifth Tanager Expedition (1924) to ...

and Caum. They named ''A. brownii'' after Brown in 1931 with the publication of their paper "Vascular plants of the Leeward Islands, Hawaii". In the paper they originally described ''A. brownii'' as one of 20 vascular plant species on the island of Nihoa. The FWS does not recognize a common name.

Description

''Amaranthus brownii'' was the only endemic species of Hawaiian ''Amaranthus'' in the Hawaiian Islands. It was an herbaceous

annual plant An annual plant is a plant that completes its life cycle, from germination to the production of seeds, within one growing season, and then dies. The length of growing seasons and period in which they take place vary according to geographical ...

that grows to a height of and had narrow, linear leaves, small green flowers, and fruit that held a single, dark red seed. ''A. brownii'' was

monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...

; that is, the male and female flowers were found together on the same plant.United States Fish and Wildlife Service (2003).
Endangered and Threatened Wildlife and Plants; Designation of Critical Habitat for Five Plant Species From the Northwestern Hawaiian Islands, Hawaii
(PDF). ''Federal Register'' (United States Government Printing Office) 68 (99): 28053–28075. Retrieved 2011-05-12; See also: "Listing actions". ''Endangered Species Update'' (University of Michigan, School of Natural Resources and Environment). April–June 2004. It differed from other Hawaiian species of ''Amaranthus'' with its spineless leaf axils, linear leaves, and

fruits (fruit which does not open to release seeds when ripe). The fruits were ovoid and between 0.8–1 mm long and 0.6–0.8 mm wide. The plant is thought to have been

anemophilous Anemophily or wind pollination is a form of pollination whereby pollen is distributed by wind. Almost all gymnosperms are anemophilous, as are many plants in the order Poales, including grasses, sedges, and rushes. Other common anemophilous plan ...

(pollinated by wind).

Distribution and habitat

''Amaranthus brownii'' had a very limited

range Range may refer to: Geography * Range (geographic), a chain of hills or mountains; a somewhat linear, complex mountainous or hilly area (cordillera, sierra) ** Mountain range, a group of mountains bordered by lowlands * Range, a term used to i ...

; it was found only on the island of Nihoa, located northwest of

Kauai Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the 21st largest island ...

. It is thought that this endemic plant had probably always been rare and restricted to Nihoa. Its former habitat is managed by the

and protected as part of the

in the

. ''A. brownii'' was one of three endemic and endangered species only found on Nihoa, along with the Nihoa fan palm (''

Pritchardia remota ''Pritchardia remota'', the Nihoa pritchardia, Nihoa fan palm, or Loulu, is a species of palm endemic on the island of Nihoa, Hawaii, and later transplanted to the island of Laysan. It is a smaller tree than most other species of '' Pritchardia' ...

'') and the Nihoa carnation (''

Schiedea verticillata ''Schiedea verticillata'', known as the Devils Slide schiedea or Nihoa carnation, is an endangered species of plant in the family Caryophyllaceae, endemic to the island of Nihoa in the Northwestern Hawaiian Islands, where it was discovered in ...

'').Pacific Islands Fish and Wildlife Office (2008).
Recovery Plan for Three Plant Species on Nihoa Island
. Endangered Species in the Pacific Islands. United States Fish and Wildlife Service. Retrieved 2008-09-05; See also: U.S. Fish and Wildlife Service 1998. At least nine other native plant species can be found in its habitat, including Hawaiian goosefoot (''

Chenopodium oahuense ''Chenopodium oahuense'' is a species of flowering plant in the amaranth family known by the common names ''aweoweo'', ''alaweo'', ''alaweo huna'', ''aheahea'', ''ahea'', ''ahewahewa'', and ''kahaihai''.Eragrostis variablis''), ''koali awa'' (''

Ipomoea indica ''Ipomoea indica'' is a species of flowering plant in the family Convolvulaceae, known by several common names, including blue morning glory, oceanblue morning glory, ''koali awa'', and blue dawn flower. It bears heart-shaped or 3-lobed leaves an ...

''), goat's foot (''

Ipomoea pes-caprae ''Ipomoea pes-caprae'', also known as bayhops, bay-hops, beach morning glory or goat's foot, is a common pantropical creeping vine belonging to the family Convolvulaceae. It grows on the upper parts of beaches and endures salted air. It is one ...

'' subsp. ''brasiliensis''), ''

Panicum torridum ''Panicum'' (panicgrass) is a large genus of about 450 species of Poaceae, grasses native throughout the tropical regions of the world, with a few species extending into the northern temperate zone. They are often large, Annual plant, annual or P ...

'', ''naupaka'' (''

Scaevola sericea ''Scaevola taccada'', also known as beach cabbage, sea lettuce, or beach naupaka, is a flowering plant in the family Goodeniaceae found in coastal locations in the tropical areas of the Indo-Pacific. It is a common beach shrub throughout the Ara ...

''), ''

Sicyos pachycarpus ''Sicyos'' is a flowering plant genus of the family Cucurbitaceae. Members of the genus are commonly known as burr cucumbers. Selected species Out of 130 species which have been attributed to the genus, just 37 are currently accepted by botani ...

'', ''ilima'' (''

Sida fallax ''Sida fallax'', known as yellow ilima or golden mallow, is a species of herbaceous flowering plant in the ''Hibiscus'' family, Malvaceae, indigenous to the Hawaiian Archipelago and other Pacific Islands. Plants may be erect or prostrate and are ...

''), and Nelson's horsenettle (''

Solanum nelsonii ''Solanum nelsonii'' (often misspelled ''Solanum nelsoni''), common names pōpolo and Nelson's horsenettle, is a partially woody sprawling shrub-like perennial plant in the family Solanaceae, part of the ''Solanum'' or nightshade genus. This pois ...

'').

The plant grew during the moist, winter season from December through July in Nihoa's coastal dry shrubland habitat in shallow soil on rocky

s in exposed areas between . At the time of its discovery in 1923, botanists first observed the plant growing in great quantity on the

ridge A ridge or a mountain ridge is a geographical feature consisting of a chain of mountains or hills that form a continuous elevated crest for an extended distance. The sides of the ridge slope away from the narrow top on either side. The line ...

s towards Miller Peak and on eastern ridges of the island. Expeditions in the early and mid-1960s failed to identify any specimens, but in 1969,

ethnobotanist Ethnobotany is the study of a region's plants and their practical uses through the traditional knowledge of a local culture and people. An ethnobotanist thus strives to document the local customs involving the practical uses of local flora for m ...

Douglas E. Yen of the

Bishop Museum The Bernice Pauahi Bishop Museum, designated the Hawaii State Museum of Natural and Cultural History, is a museum of history and science in the historic Kalihi district of Honolulu on the Hawaiian island of Oʻahu. Founded in 1889, it is the lar ...

collected specimens near Miller Peak. Derral R. Herbst and Wayne Takeuchi of the FWS collected the last known specimen on July 27, 1980. Carl C. Christensen also visited Nihoa in 1980 to reevaluate endemic species last observed on the Tanager Expedition. Sheila Conant and Mark S. Collins visited Nihoa in 1980 as well; Conant returned twice in 1981, first with Mark J. Rauzon and later with Audrey L. Newman. In 1983, Conant visited the island with Wayne C. Gagné. Conant found ''A. brownii'' growing on the island in 1981 and 1983, by which time only two populations of 35 plants were thought to exist: 23 plants were found near Miller Peak and 12 plants in Middle Valley. The two plant populations were separated by a distance of approximately . Prehistoric Polynesian habitation on Nihoa may have initially led to a decrease in the plant population of ''A. brownii''. Major threats to the plant included

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

, fire, and

hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...

with other ''Amaranthus'' species. Inbreeding was a serious threat, as the small plant population must reproduce within its own circle resulting in genetic defects. ''A. brownii wa''s also forced to compete with the non-native weed (''

Portulaca oleracea ''Portulaca oleracea'' (common purslane, also known as little hogweed, or pursley) is an annual (actually tropical perennial in USDA growing zones 10–11) succulent in the family Portulacaceae. Description The plant may reach in height. It ...

''), the plant's main alien species threat. In 2002 and 2004, the invasive gray bird grasshopper (''

Schistocerca nitens ''Schistocerca nitens'' is a species of grasshopper known by several names, including vagrant grasshopper and gray bird grasshopper. It is a close relative of the desert locust, which is in the same genus. This grasshopper is native to southern N ...

'') presented an even larger threat to ''A. brownii''. First recorded on the island in 1977, the increasing population density of gray bird grasshoppers led to massive defoliation on the island, leaving ''A. brownii'' at greater risk of

predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...

. In 2004 alone, an estimated 400,000 gray bird grasshoppers destroyed almost 90% of the vegetation on Nihoa.Lockwood, Jeffrey A.; Alexandre V. Latchininsky (2008).
Confessions of a Hit Man
. ''Conservation Magazine'' (Seattle, WA: Society for Conservation Biology) 9 (3). Retrieved 2011-05-12. All these threats may have contributed to its ultimate extinction.

Conservation

According to zoologist and conservationist Sheila Conant, ''A. brownii'' was important due to its uniqueness in the Northwestern Hawaiian Islands as "the only Hawaiian endemic in this large genus which contains many economically and nutritionally important species." However, in more than a decade of field surveys on Nihoa, no living plants were identified. Wildlife refuge staff visited the island during the dry season at least 21 times between 1983 and 1996. Originally, botanists thought the absence of the plant in field surveys might have been explained by the time of visits During the summer months, the stems of ''A. brownii'' dried up and could not be distinguished from other

herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...

s. However, a seven-day visit to the island in April 2006 still did not find any specimens. Additional winter surveys were performed, but none were successful.

Ex-situ conservation Svalbard GLOBAL SEED BANK, an ''ex situ'' conservation. ''Ex situ'' conservation literally means, "off-site conservation". It is the process of protecting an endangered species, variety or breed, of plant or animal outside its natural habitat; ...

efforts to propagate ''A. brownii'' by seed in

botanical garden A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...

s were also unsuccessful. During the 1981 expedition, ''A. bronwii'' seeds were collected by Sheila Conant and presented to the Waimea Arboretum on the Hawaiian island of

Oahu Oahu () (Hawaiian language, Hawaiian: ''Oʻahu'' ()), also known as "The Gathering place#Island of Oʻahu as The Gathering Place, Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over t ...

and the

Kew Gardens Kew Gardens is a botanical garden, botanic garden in southwest London that houses the "largest and most diverse botany, botanical and mycology, mycological collections in the world". Founded in 1840, from the exotic garden at Kew Park, its li ...

London, England London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a major s ...

. Although the seeds at the Waimea Arboretum

germinated 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, fer ...

and grew for a while, no plants survived beyond the stage of

seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embry ...

development. Information about the outcome of the seeds sent to Kew Gardens is unavailable. A proposal for listing ''A. brownii'' under the U.S.

Endangered Species Act The Endangered Species Act of 1973 (ESA or "The Act"; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of ec ...

was originally submitted on June 16, 1976, but was withdrawn on December 10, 1979, as out of date and incomplete. It was proposed again on March 24, 1993, and was federally listed as an

endangered species An endangered species is a species that is very likely to become extinct in the near future, either worldwide or in a particular political jurisdiction. Endangered species may be at risk due to factors such as habitat loss, poaching and inv ...

on August 21, 1996. On May 22, 2003, the FWS designated on the island of Nihoa as a

for ''A. brownii'', as well as ''Pritchardia remota'', ''Schiedea verticillata'', and two other species also found on Nihoa and other Hawaiian islands, ''ohai'' (''

Sesbania tomentosa ''Sesbania tomentosa'', commonly known as Oahu riverhemp and ''ōhai'', is an endangered species of plant in the pea family, Fabaceae, that is endemic to the main Hawaiian Islands as well as Nihoa and Necker Island. It inhabits low shrublands ...

'') and '' Mariscus pennatiformis''. Also in 2003, ''A. brownii'' was internationally classified as critically endangered on the

IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...

. As of 2010, ''A. brownii'' was one of 51 Hawaiian plant species listed as endangered or threatened under the Endangered Species Act.

Notes

a. Wagner and Herbst list five naturalized species of ''

'' in Hawaii in addition to the endemic ''A. brownii''. The authors note that the information may be both inaccurate and incomplete due to errors caused by a lost collection and lack of data. '' A. graecizans'', '' A. retroflexus'', and a third unknown species (possibly extinct or reclassified) have been proposed as additional naturalized candidates. b. Nihoa: fl, fr, June 20, 1923, C. S. Judd No. 2 c. Herbst & Takeuchi 6545; BISH. Also see th
database record
at the U.S. National Herbarium: Herbst, D.R.; Takeuchi, W. No. 6545; Collection Date: 27 Jul 1980; Hawaiian Islands, Nihoa, Middle Valley. Alt. 91 m.; Barcode: 00453038 USNM No.: 02921853. d. According to Mark J. Rauzon, anthropologist

Kenneth Emory Kenneth Pike Emory (November 23, 1897 – January 2, 1992) was an American anthropologist who played a key role in shaping modern anthropology in Oceania. In the tradition of A. L. Kroeber and other pioneering anthropologists who trained him, ...

, a member of the Tanager Expedition, identified 66 archaeological sites on the island of Nihoa, and to date 86 sites have been found. Emory estimated that 7.7 percent of the island (12 out of 156 acres) was used for terraced, dry-land crop production, and along with fish and seafowl, Emory believed 100 people (or more) could have survived on a long-term basis. However, questions about good potable water sources and the fact that only six skeletons have been found cast doubt on this figure. e. Compare the destruction of vegetation on Nihoa in 2004 to that of

Laysan Laysan (; haw, italics=no, Kauō ), located northwest of Honolulu at , is one of the Northwestern Hawaiian Islands. It comprises one land mass of , about in size. It is an atoll of sorts, although the land completely surrounds a shallow c ...

island. In 1894, Max Schlemmer introduced rabbits to Laysan. Eventually, the rabbit problem and bird poaching led U.S. President

Theodore Roosevelt Theodore Roosevelt Jr. ( ; October 27, 1858 – January 6, 1919), often referred to as Teddy or by his initials, T. R., was an American politician, statesman, soldier, conservationist, naturalist, historian, and writer who served as the 26t ...

to declare the Northwestern Hawaiian Islands a bird sanctuary in 1909. By 1918, 26 plant species had disappeared from the island and the

Laysan millerbird The Laysan millerbird (''Acrocephalus familiaris familiaris'') was a subspecies of the millerbird, similar in appearance to the remaining subspecies, the Nihoa millerbird. Its dorsal side was brown, and its belly was grayish. Its name derives fr ...

had become extinct. The Tanager Expedition arrived in 1923 and exterminated the last of the surviving rabbits.

References

External links

* *
Amaranthus brownii Christoph. & Caum
'. ''Hawaii Biological Survey''.

. Retrieved May 20, 2011. {{Taxonbar, from=Q4740157 brownii Endemic flora of Nihoa Extinct flora of Hawaii Plants described in 1931 Tanager Expedition Plant extinctions since 1500

Taxonomy

Description

Distribution and habitat

Conservation

Notes

References

Further reading

External links