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{{short description|Extinct species of flowering plant}}
{{Taxobox
{{good article}}
| name = ''Amaranthus brownii''
{{speciesbox
| image = Amaranthusbrownii low res.jpg
|image = Amaranthusbrownii low res.jpg
| image_caption =
|status = EX
| image_width = 250px
|status_system = IUCN3.1
| status = CR
|status_ref = <ref name="iucn status 19 November 2021">{{cite iucn |author=Bruegmann, M. |author2=Caraway, V.L. |author3=Keir, M. |date=2018 |title=''Amaranthus brownii'' |volume=2018 |page=e.T44077A83794603 |doi=10.2305/IUCN.UK.2018-2.RLTS.T44077A83794603.en |access-date=19 November 2021}}</ref>
| status_system = iucn2.3
|genus = Amaranthus
| regnum = [[Plant]]ae
|species = brownii
| divisio = [[Flowering plant|Magnoliophyta]]
|authority = [[Erling Christophersen|Christoph]]. & [[Edward Leonard Caum|Caum]]
| classis = [[Magnoliopsida]]
| ordo = [[Caryophyllales]]
| familia = [[Amaranthaceae]]
| genus = ''[[Amaranthus]]''
| species = '''''A. brownii'''''
| binomial = ''Amaranthus brownii''
| binomial_authority = [[Erling Christophersen|Christoph]]. & [[Edward Leonard Caum|Caum]]
}}
}}
'''''Amaranthus brownii''''', also known as '''Brown's amaranth''', or more commonly '''pigweed''', is an [[Annual plant|annual]] [[herbaceous plant|herb]] in the [[Amaranthaceae]] family. It is one of nine species of ''Amaranthus'' in [[Hawaii]],{{Ref_label|A|a|none}} but the only [[endemic]] Hawian species of the genus. Botanists from the [[Tanager Expedition]] first collected the plant in 1923 while exploring [[Nihoa]], a small island in the [[Northwestern Hawaiian Islands]]. This unique plant is endemic to the island of Nihoa and is found nowhere else. The plant shares the island with 12 other endemic plants, two of which&#8212;''[[Pritchardia remota]]'' and ''[[Schiedea verticillata]]''&#8212;can only be found on Nihoa like itself. ''Amaranthus brownii'' is considered the rarest plant on Nihoa, and has not been observed ''[[in situ]]'' since 1983.<ref name="FWS003"/>


'''''Amaranthus brownii''''' was an [[Annual plant|annual]] [[herbaceous plant|herb]] in the family [[Amaranthaceae]]. The plant was found only on the small island of [[Nihoa]] in the [[Northwestern Hawaiian Islands]], growing on rocky [[outcrop]]s at altitudes of {{convert|120|–|215|m|ft|abbr=on}}. It was one of nine species of ''[[Amaranthus]]'' in the [[Hawaiian Islands]], as well as the only [[endemism|endemic]] Hawaiian species of the genus. It is now considered [[extinction|extinct]].
==Morphology==
''A. brownii'' is an herbaceous [[annual plant]] that grows from 30-90 cm (1-3 ft) in height and has narrow, linear leaves, small, green flowers, and fruit that holds a single, dark red seed. ''A. brownii'' is [[monoecious]] and the male and female flowers are found together on the same plant.<ref name="FWS003"/> There are nine recorded species of Hawaiian Amaranthus, but ''A. brownii'' is the only native species in the islands.<ref name=Evenhuis_Eldredge/> ''A. brownii'' differs from other Hawaiian species of ''Amaranthus'' with its spineless leaf axils, linear leaves, and [[indehiscent]] fruits (fruit which does not open to release seeds when ripe)<ref name="FWS003"/> that are ovoid and between 0.8-1 mm long and 0.6-0.8 mm wide.<ref name="Wagner-1999"/>


''A. brownii'' was first discovered during the [[Tanager Expedition]] in [[1923 in science|1923]] by botanist [[Edward Leonard Caum]]. It differed from other Hawaiian species of ''Amaranthus'' with its [[Thorns, spines, and prickles|spineless]] leaf axils, [[Leaf shape|linear leaves]], and [[Dehiscence (botany)|indehiscent]] fruits. It was one of [[Flora of Nihoa|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|''ex situ'' conservation]] efforts outside of its native habitat. Consequently, there are no known plants or seeds from ''A. brownii'' in any botanical garden.
==Distribution and habitat==
The plant has a very limited [[Range (biology)|range]], as it is endemic to and only found on Nihoa. Nihoa is part of the [[Hawaiian Islands National Wildlife Refuge]] in the [[Papahānaumokuākea Marine National Monument]] and is managed by the [[United States Fish and Wildlife Service]].<ref name="DNLR"/>


[[Island restoration|Conservation and recovery plan]]s for ''A. brownii'' were proposed by the [[United States Fish and Wildlife Service]] (FWS), which administers the island of Nihoa as part of the [[Hawaiian Islands National Wildlife Refuge]] in the [[Papahānaumokuākea Marine National Monument]]. In 1996, the plant was listed by the FWS as an endangered species. In 2003, the FWS designated the island of Nihoa as a [[Endangered Species Act#Critical habitat|critical habitat]] for the plant and it was classified as [[Critically endangered species|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.<ref name="iucn status 19 November 2021" />
[[Image:NWHI Reserve.jpg|thumb|left|Papahānaumokuākea Marine National Monument]]
''A. brownii'' was first collected on Nihoa by botanist [[Edward Leonard Caum]]<ref name="BruegmannFWS"/> during a ten-day visit by the [[Tanager Expedition]] beginning on June 10, 1923. Scientists conducting research on the island observed the plant growing in great quantity on the [[ridge]]s towards Miller Peak and on eastern ridges of the island. Expeditions in the early and mid-1960s failed to identify any specimens, however, in 1969, [[Ethnobotany|ethnobotanist]] Douglas E. Yen of the Bishop Museum Project collected specimens near Miller Peak. Sheila 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:<ref name="Conant">{{cite journal
| last = Conant
| first = Sheila
| title = Recent observations on the plants of Nihoa Island, Northwestern Hawaiian Islands
| journal = Pacific Science
| volume = 39
| issue = 2
| pages = 135–149
| publisher = [[University of Hawaii Press]]
| date = 1985
| url = https://scholarspace.manoa.hawaii.edu/bitstream/10125/921/1/v39n2-135-149.pdf
| format = PDF
| accessdate = 2008-07-22
}}</ref> 23 plants were found near Miller Peak and 12 plants in Middle Valley. The two plant populations are separated by a distance of approximately 0.4 km (0.25 mi).<ref name="DNLR"/>


==Taxonomy==
[[Image:Nihoa map lrg.gif|thumb|Plant populations were found on Miller's Ridge and on the west ridge of Middle Valley<ref>Conant 1985, p. 143</ref>]]
The species was first collected during a ten-day visit to the island of Nihoa by the [[Tanager Expedition]].<ref name="Chris1931">{{cite book|last=Christophersen|first=Erling|author2=Caum, Edward L.|title=Vascular plants of the Leeward Islands, Hawaii|publisher=[[Bishop Museum|Bishop Museum Press]]|series=Bernice P. Bishop Museum Bulletin No. 81; Tanager Expedition Publication No. 7|pages=25–26|year=1931|location=Honolulu, Hawaii|quote=Nihoa: fl, fr, June 17, 1923, E. L. Caum No. 73. Type, B. P. Bishop Mus.}}</ref> Botanist [[Edward Leonard Caum]] collected the first specimen on June 17, 1923, and a second was collected by cartographer Charles S. Judd{{Ref label|B|b|none}} on June 20, 1923.<ref name="Clapp1977">{{cite journal|last=Clapp|first=Roger B.|author2=Eugene Krindler|author3=Robert R. Fleet|editor=F. R. Fosberg|editor2=M.-H. Sachet|editor3=D. R. Stoddart|year=1977|url=http://www.botany.hawaii.edu/faculty/duffy/arb/206-207/207.pdf|title=The Natural History of Nihoa Island, Northwestern Hawaiian Islands|journal=Atoll Research Bulletin|issue=207|publisher=[[Smithsonian Institution]]|location=Washington, D. C.|access-date=2011-05-12|pages=31–32|archive-url=https://web.archive.org/web/20110611162438/http://www.botany.hawaii.edu/faculty/duffy/arb/206-207/207.pdf|archive-date=2011-06-11|url-status=dead}}</ref> [[Forest B. H. Brown]], botanist of the [[Bayard Dominick Expedition]] to the [[Marquesas Islands]] (1921–1922),<ref>{{cite journal|last=Sherff|first=Earl E.|year=1932|title=Review: Monocotyledons of Southeastern Polynesia|journal=Botanical Gazette |publisher=University of Chicago Press|volume=93|issue=1|pages=107–108|doi=10.1086/334237|s2cid=85804033}}</ref> helped provide descriptions and comments for some of the species described by [[Erling Christophersen]] and Caum.<ref>Christophersen & Caum 1931, p. 4: "At the time when we started work on the "Tanager" collections Dr. F. B. H. Brown had already described some of the new species and varieties which they contained, and his descriptions are here included He has also passed his opinion on some of the other plants for which we express our indebtedness."</ref> They named ''A. brownii'' after Brown in 1931<ref name="FWS996">{{cite journal|author=United States Fish and Wildlife Service|year=1996|url=http://ecos.fws.gov/docs/federal_register/fr2983.pdf|title=Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Three Plants from the Island of Nihoa, Hawaii|journal=Federal Register|volume=61|issue=163|publisher=United States Government Printing Office|pages=43178–43184|access-date=2011-05-12}}</ref> 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.<ref>Christophersen & Caum 1931, p. 5; Clapp 1977, p. 147.</ref> The FWS does not recognize a common name.<ref name="FWS2007"/>
''A. brownii'' grows during the moist, winter season from December through July. It can be found in shallow soil on rocky [[outcrop]]s in exposed areas between 120 and 215 m (390 and 700 ft).<ref name="FWS003"/> At least ten other native plant species can be found in its habitat,{{Ref_label|B|b|none}} including 'aheahea, kakonakona, kupala, and a non-native amaranth species.<ref name="DNLR"/>


==Description==
Major threats are [[invasive species]], fire, and possible [[Hybrid (biology)|hybridization]] with other ''Amaranthus'' species.<ref name="DNLR">{{cite web
''Amaranthus brownii'' was the only endemic species of Hawaiian ''Amaranthus'' in the Hawaiian Islands.{{Ref label|A|a|none}} It was an herbaceous [[annual plant]] that grows to a height of {{convert|30|-|90|cm|ft|abbr=on}} and had narrow, linear leaves, small green flowers, and fruit that held a single, dark red seed. ''A. brownii'' was [[monoecious]]; that is, the male and female flowers were found together on the same plant.<ref name="FWS003">United States Fish and Wildlife Service (2003). "[http://ecos.fws.gov/docs/federal_register/fr4103.pdf 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.</ref> It differed from other Hawaiian species of ''Amaranthus'' with its spineless leaf axils, linear leaves, and [[indehiscent]] fruits (fruit which does not open to release seeds when ripe).<ref name="FWS003"/> The fruits were ovoid and between 0.8–1&nbsp;mm long and 0.6–0.8&nbsp;mm wide.<ref name="Wagner-1999">{{cite book|last=Wagner|first=Warren L.|author2=Derral R. Herbst |author3=Sy H. Sohmer |year=1999|title=Manual of the Flowering Plants of Hawaiʻi|publisher=[[University of Hawaii Press]]|volume=1|edition=Revised|location=Honolulu, Hawaii|pages=185–189|isbn=0-8248-2166-1}}</ref> The plant is thought to have been [[anemophilous]] (pollinated by wind).<ref name=Evenhuis_Eldredge/>
| title = Amaranthus brownii fact sheet
| work = Hawaii's Comprehensive Wildlife Conservation Strategy
| publisher = [[Hawai'i Department of Land and Natural Resources]]
| date = October 1, 2005
| url = http://www.state.hi.us/dlnr/dofaw/cwcs/files/Flora%20fact%20sheets/Ama_bro%20plant%20NTBG_OK.pdf
| format = PDF
| doi =
| accessdate = 2008-03-31
}}</ref> Inbreeding is a serious threat, as the small plant population is forced to reproduce within its own circle resulting in genetic defects.<ref name=Evenhuis_Eldredge/> In 2002 and 2004, the invasive ''[[Schistocerca nitens]]'', a nonnative grasshopper, presented an even larger threat to ''A. brownii''. Although it was first recorded on the island in the early 1980s, ''S. nitens'' has increased its population density, resulting in massive defoliation of Nihoa's vegetation, leaving ''A. brownii'' at greater risk of [[predation]].<ref name="FWS2007"/> ''A. brownii'' is also forced to compete with non-native ''[[Portulaca oleracea]]'' (pigweed), the plant's main alien species threat.<ref name="FWS003"/>
{{clear}}


==Distribution and habitat==
==Conservation==
[[File:NHINM.png|thumb|left|Papahānaumokuākea Marine National Monument]]
''A. brownii'' shares the island with 27 [[vascular plant]] species, 21 of which are native and 12 endemic. The plant is one of three endemic and endangered species only found on Nihoa, along with ''[[Pritchardia remota]]'' and ''[[Schiedea verticillata]]''.<ref name=FWS2008>{{cite web
''Amaranthus brownii'' had a very limited [[Range (biology)|range]]; it was found only on the {{convert|0.65|km2|sqmi|2|abbr=on}} island of Nihoa, located {{convert|275|km|mi|0|abbr=on}} northwest of [[Kauai]]. It is thought that this endemic plant had probably always been rare and restricted to Nihoa.<ref name="FWS998">{{cite report|author=United States Fish and Wildlife Service|date=March 1998|url=http://ecos.fws.gov/docs/recovery_plan/980331a.pdf|title=Final Recovery Plan for Three Plant Species on Nihoa Island|publisher=United States Fish and Wildlife Service|pages=10547–10550|access-date=2011-05-12}}</ref> Its former habitat is managed by the [[United States Fish and Wildlife Service]] and protected as part of the [[Hawaiian Islands National Wildlife Refuge]] in the [[Papahānaumokuākea Marine National Monument]].<ref>{{cite web|author=State of Hawaii|author2=National Oceanic and Atmospheric Administration|author3=Office of Hawaiian Affairs|author4=U.S. Fish and Wildlife Service|year=2009|url=http://www.papahanaumokuakea.gov/management/worldheritage.html|title=The United States of America's Nomination of Papahānaumokuākea Marine National Monument for Inscription on the World Heritage List|publisher=Papahānaumokuākea Marine National Monument|location=Honolulu, Hawaii|access-date=2011-05-10}}</ref> ''A. brownii'' was one of three endemic and endangered species only found on Nihoa, along with the Nihoa fan palm (''[[Pritchardia remota]]'') and the Nihoa carnation (''[[Schiedea verticillata]]'').<ref name="FWS008">Pacific Islands Fish and Wildlife Office (2008). "[http://www.fws.gov/pacificislands/flora/nihoaplants.html 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.</ref> At least nine other native plant species can be found in its habitat, including Hawaiian goosefoot (''[[Chenopodium oahuense]]''), lovegrass (''[[Eragrostis variablis]]''), ''koali {{okina}}awa'' (''[[Ipomoea indica]]''), goat's foot (''[[Ipomoea pes-caprae]]'' subsp. ''brasiliensis''), ''[[Panicum torridum]]'', ''naupaka'' (''[[Scaevola sericea]]''), ''[[Sicyos pachycarpus]]'', ''{{okina}}ilima'' (''[[Sida fallax]]''), and Nelson's horsenettle (''[[Solanum nelsonii]]'').<ref name="FWS003"/>
|author = Pacific Islands Fish and Wildlife Office
|title = Recovery Plan for Three Plant Species on Nihoa Island
|work = Endangered Species in the Pacific Islands
|publisher = [[United States Fish and Wildlife Service]]
|date = 2008
|url = http://www.fws.gov/pacificislands/flora/nihoaplants.html
|accessdate = 2008-09-05
}}</ref> ''A. brownii'' was federally listed as an [[endangered species]] in 1996,<ref name="BruegmannFWS">{{cite journal
| last = Bruegmann
| first = M. M.
| coauthors = Z. E. Ellshoff
| title = Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Three Plants from the Island of Nihoa, Hawaii
| journal = [[Federal Register]]
| volume = 61
| issue = 163
| pages = 43178–43184
| publisher = [[United States Government Printing Office]]
| date = 1996
| url = http://www.fws.gov/Endangered/pdfs/FR/f960821.pdf
| format =
| accessdate = 2008-04-12
}}</ref> and internationally classified as [[critically endangered]] on the [[IUCN Red List]] in 2003.<ref name="Redlist">{{cite web
| last = Bruegmann
| first = M.M.
| coauthors = Caraway, V.
| title = ''Amaranthus brownii''
| work = 2007 IUCN Red List of Threatened Species
| publisher = [[World Conservation Union|International Union for Conservation of Nature and Natural Resources]]
| date = 2003
| url = http://www.iucnredlist.org/search/details.php/44077/summ2007
| accessdate = 2008-04-03
}}</ref> On May 22, 2003, 171 acres (69 hectares) on the island of Nihoa were designated by the United States Fish and Wildlife Service as a [[Endangered_Species_Act#Critical_Habitat|critical habitat]] for ''A. brownii'', as well as ''Pritchardia remota'', ''Schiedea verticillata'', and two other species found on Nihoa and other Hawaiian islands, ''[[Sesbania tomentosa]]'' (''{{okina}}ohai'') and ''[[Mariscus pennatiformis]]''.<ref name="FWS003">{{cite journal
| author = United States Fish and Wildlife Service
| title = Endangered and Threatened Wildlife and Plants; Designation of Critical Habitat for Five Plant Species From the Northwestern Hawawiian Islands, Hawai
| journal = [[Federal Register]]
| volume = 68
| issue = 99
| pages = 28054–28075
| publisher = [[United States Government Printing Office]]
| date = May 22, 2003
| url = http://www.fws.gov/pacificislands/CHRules/nwhifinal.pdf
| format = PDF
| accessdate = 2008-03-31
}}</ref>


[[Image:Nihoa aerial.jpg|thumb|left|Aerial view of Nihoa. In 1983, W. Gagné recorded 12 plants living on the island.<ref name="Wagner-1999">{{cite book
[[File:Nihoa aerial.jpg|thumb|Aerial view of Nihoa. In 1983, W. Gagné recorded 12 plants living on the island.<ref name="Wagner-1999"/>]]
The plant grew during the moist, winter season from December through July in Nihoa's [[Northwestern Hawaii scrub|coastal dry shrubland]] habitat<ref name="FWS998"/> in shallow soil on rocky [[outcrop]]s in exposed areas between {{convert|120|-|215|m|ft|abbr=on}}.<ref name="FWS003"/> At the time of its discovery in 1923, botanists first observed the plant growing in great quantity on the [[ridge]]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, [[Ethnobotany|ethnobotanist]] Douglas E. Yen of the [[Bernice P. Bishop Museum|Bishop Museum]] collected specimens near Miller Peak.<ref name="Clapp1977"/> Derral R. Herbst and Wayne Takeuchi of the FWS collected the last known specimen{{Ref label|C|c|none}} on July 27, 1980.<ref name="BISH86">{{cite journal|last=Wagner|first=Warren L.|author2=Derral R. Herbst |author3=Sy H. Sohmer |year=1986|title=Contributions to the flora of Hawaiʻi. I. Acanthaceae—Asteraceae|journal=Bishop Museum Occasional Papers|publisher=Bernice Pauahi Bishop Museum|volume=26–27|pages=102–122}}</ref> Carl C. Christensen also visited Nihoa in 1980 to reevaluate endemic species last observed on the Tanager Expedition.<ref name="Conant1994">{{cite book| last = Conant| first = Sheila| editor = E. Alison Kay| author2 = Carl C. Christensen| author3 = Patrick Conant| author4 = Wayne C. Gagné| author5 = M. Lee Goff| chapter = The Unique Terrestrial Biota of the Northwestern Hawaiian Islands| chapter-url = http://www.botany.hawaii.edu/faculty/morden/BotZool450/Conant1984.pdf| title = A Natural History of the Hawaiian Islands: Selected Readings II| publisher = [[University of Hawaii Press]]| year = 1994| location = Honolulu| pages = [https://archive.org/details/naturalhistoryof00kaye/page/378 378–390]| isbn = 978-0-8248-1659-9| url-access = registration| url = https://archive.org/details/naturalhistoryof00kaye/page/378}}</ref> 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:<ref name="Conant-1985">{{cite journal |last=Conant |first=Sheila |title=Recent Observations on the Plants of Nihoa Island, Northwestern Hawaiian Islands |journal=Pacific Science |volume=39 |issue=2 |pages=135–149 |publisher=[[University of Hawaii Press]] |year=1985 |hdl=10125/921 |url=http://scholarspace.manoa.hawaii.edu/handle/10125/921 |access-date=2011-05-17}}</ref> 23 plants were found near Miller Peak and 12 plants in Middle Valley. The two plant populations were separated by a distance of approximately {{convert|0.4|km|mi|abbr=on}}.<ref name="DNLR"/>
| last = Wagner
| first = W.L.
| coauthors = Herbst, D.R.; Sohmer, S.H.
| title = Manual of the flowering plants of Hawai'i
| publisher = [[University of Hawaii Press]]
| volume = I
| pages = 185-189
| edition = Revised
| date = 1999
| location = Honolulu, Hawaii
| isbn = 0824821661
}}</ref>]]
Wildlife refuge staff visited Nihoa during the dry season at least 21 times between 1983-1996.<ref name="FWS2007">{{cite paper
| author = Pacific Islands Fish and Wildlife Office
| authorlink =
| coauthors =
| title = ''Amaranthus brownii'' (No common name); 5-Year Review; Summary and Evaluation
| version =
| publisher = [[United States Fish and Wildlife Service]]
| date = 2007
| url = http://www.fws.gov/pacific/ecoservices/endangered/recovery/Documents/Amaranthusbrownii.pdf
| format = PDF
| accessdate = 2008-04-04
}}</ref> In more than a decade of field surveys, no living plants were identified. Because winter surveys of Nihoa tend to be difficult and dangerous due to poor landing conditions,<ref name="FWS003"/> surveys were not conducted during the moist, winter growing season from December through March when the plant is easiest to find.<ref name=Evenhuis_Eldredge/> Most of the surveys have been completed during the summer months, when the stems of ''A. brownii'' dry up and cannot be distinguished from other [[herbaceous plant]]s.<ref name="FWS2007"/> As a result, additional winter surveys of ''A. brownii'' on Nihoa are required to accurately assess its current conservation status.<ref name=Evenhuis_Eldredge/> A recent seven day visit to the island in April of 2006 still did not find any specimens. Nevertheless, as of 2007, botanists are optimistic that the species has survived.<ref name="FWS2007"/>


[[Nihoa#Old human habitation|Prehistoric Polynesian habitation]] on Nihoa{{Ref label|D|d|none}} may have initially led to a decrease in the plant population of ''A. brownii''.<ref name="Beacham-2000">{{cite book|last=Beacham|first=Walton|title=Beacham's Guide to the Endangered Species of North America|author2=Frank Vincent Castronova|author3=Suzanne Sessine|publisher=Gale Group|year=2000|isbn=0-7876-5028-5|series=Arachnids and Crustaceans, Insects, Lichens, Fern Allies, True Ferns, Conifers, Dicots|volume=3|location=Detroit|pages=1641–1642}}</ref> Major threats to the plant included [[invasive species]], fire, and [[Hybrid (biology)|hybridization]] with other ''Amaranthus'' species.<ref name="DNLR">{{cite web |author=Hawai'i Department of Land and Natural Resources|author-link=Hawai'i Department of Land and Natural Resources |year=2005|url=http://www.state.hi.us/dlnr/dofaw/cwcs/files/Flora%20fact%20sheets/Ama_bro%20plant%20NTBG_OK.pdf|title=Amaranthus brownii fact sheet|work=Hawaii's Comprehensive Wildlife Conservation Strategy|publisher=State of Hawaii|access-date=2011-05-12|archive-url=https://web.archive.org/web/20110517072006/http://www.state.hi.us/dlnr/dofaw/cwcs/files/Flora%20fact%20sheets/Ama_bro%20plant%20NTBG_OK.pdf|archive-date=17 May 2011|url-status=dead|df=dmy-all}}</ref> Inbreeding was a serious threat, as the small plant population must reproduce within its own circle resulting in genetic defects.<ref name=Evenhuis_Eldredge/> ''A. brownii wa''s also forced to compete with the non-native weed (''[[Portulaca oleracea]]''), the plant's main alien species threat.<ref name="FWS003"/> In 2002 and 2004, the invasive gray bird grasshopper (''[[Schistocerca nitens]]'') presented an even larger threat to ''A. brownii''.{{Ref label|E|e|none}} First recorded on the island in 1977,<ref name="Conservation" /> the increasing population density of gray bird grasshoppers led to massive defoliation on the island, leaving ''A. brownii'' at greater risk of [[predation]].<ref name="FWS2007"/> In 2004 alone, an estimated 400,000 gray bird grasshoppers destroyed almost 90% of the vegetation on Nihoa.<ref name="Conservation">Lockwood, Jeffrey A.; Alexandre V. Latchininsky (2008). "[http://www.conservationmagazine.org/articles/v9n3/confessions-of-a-hit-man/ Confessions of a Hit Man]". ''Conservation Magazine'' (Seattle, WA: Society for Conservation Biology) '''9''' (3). Retrieved 2011-05-12.</ref><ref>{{cite journal|last=Latchininsky|first=Alexandre V.|year=2008|title=Grasshopper Outbreak Challenges Conservation Status of a Small Hawaiian Island|journal=Journal of Insect Conservation|publisher=Springer Netherlands|volume=12|issue=3–4|pages=343–357|doi=10.1007/s10841-008-9143-8|s2cid=12374828}}</ref> All these threats may have contributed to its ultimate extinction.<ref name="iucn status 19 November 2021" />
[[Ex-situ conservation]] efforts to propagate ''A. brownii'' by seed in [[botanical garden]]s have been unsuccessful.<ref name=Evenhuis_Eldredge>{{cite book

| last = Evenhuis
==Conservation==
| first = Neal L. (ed.)
[[File:Nihoa map lrg.gif|thumb|left|Plant populations were found on Miller's Ridge and on the west ridge of Middle Valley<ref name=Conant-1985/>]]
| coauthors = Eldredge, Lucius G. (ed.)
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."<ref name="Conant1994"/> 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.<ref name="FWS2007">{{cite report| author = Pacific Islands Fish and Wildlife Office | title = ''Amaranthus brownii'' (No common name); 5-Year Review; Summary and Evaluation| publisher = [[United States Fish and Wildlife Service]]| year = 2007 | url = http://www.fws.gov/pacific/ecoservices/endangered/recovery/Documents/Amaranthusbrownii.pdf| access-date = 2010-01-10 }}</ref> Originally, botanists thought the absence of the plant in field surveys might have been explained by the time of visits <ref name=Evenhuis_Eldredge/> During the summer months, the stems of ''A. brownii'' dried up and could not be distinguished from other [[herbaceous plant]]s.<ref name="FWS2007"/> However, a seven-day visit to the island in April 2006 still did not find any specimens.<ref name="FWS2007"/> Additional winter surveys were performed, but none were successful.<ref name="iucn status 19 November 2021" />
| title = Natural History of Nihoa and Necker Islands
| publisher = [[Bishop Museum|Bishop Museum Press]]
| series = Bishop Museum Bulletin in Cultural and Environmental Studies; No. 1
| date = 2004
| location = Honolulu, Hawaii
| isbn = 1-58178-029-X
}}</ref> During the 1981 expedition, ''A. bronwii'' seeds were collected by Sheila Conant and presented to the [[Waimea Arboretum]] on the Hawaiian island of [[Oahu]] and the [[Royal Botanic Gardens, Kew|Kew Gardens]] in [[London, England]]. Although the seeds at the Waimea Arboretum [[Germination|germinated]] and grew for a while, no plants survived beyond the stage of [[seedling]] development. Information about the outcome of the seeds sent to Kew Gardens is unavailable.<ref name="Conant"/>


[[Ex-situ conservation|''Ex situ'' conservation]] efforts to propagate ''A. brownii'' by seed in [[botanical garden]]s were also unsuccessful.<ref name=Evenhuis_Eldredge>{{cite book| editor-last = Evenhuis| editor-first = Neal L. | editor2= Eldredge, Lucius G. | title = Natural History of Nihoa and Necker Islands| publisher = [[Bishop Museum|Bishop Museum Press]]| series = Bishop Museum Bulletin in Cultural and Environmental Studies; No. 1| year = 2004| pages = 56–57| location = Honolulu, Hawaii| isbn = 1-58178-029-X}}</ref> During the 1981 expedition, ''A. bronwii'' seeds were collected by Sheila Conant and presented to the [[Waimea Arboretum]] on the Hawaiian island of [[Oahu]] and the [[Royal Botanic Gardens, Kew|Kew Gardens]] in [[London, England]]. Although the seeds at the Waimea Arboretum [[Germination|germinated]] and grew for a while, no plants survived beyond the stage of [[seedling]] development. Information about the outcome of the seeds sent to Kew Gardens is unavailable.<ref name="Conant-1985"/>
==Nomenclature==
The species was named in 1931 by botanists Erling Christophersen and Edward Leonard Caum after the botanist Forest Brown (F.B.H. Brown).<ref name="BruegmannFWS"/> The plant is sometimes referred to as Brown's Amaranth,<ref name="USDA_NRCS">{{cite web
| title = PLANTS Profile: ''Amaranthus brownii'' Christoph. & Caum (Brown's amaranth)
| work = USDA PLANTS Database
| publisher = [[Natural Resources Conservation Service]]
| url = http://plants.usda.gov/java/profile?symbol=AMBR5
| accessdate = 2008-08-24
}}</ref> Brown's Pigweed,<ref name="FWS_NWRS">{{cite web
| title = Threatened and Endangered Plant Species Found on the National Wildlife Refuge System
| work = America's National Wildlife Refuge System
| publisher = [[United States Fish and Wildlife Service]]
| url = http://www.fws.gov/refuges/habitats/EndSpPlants.html
| accessdate = 2008-08-24
}}</ref> or Browns Amaranth.<ref name="ZipcodeZoo">{{cite web
| title = Amaranthus brownii (Brown's Amaranth, Brown's Pigweed, Browns Amaranth)
| publisher = ZipcodeZoo
| url = http://zz4k.com/Plants/A/Amaranthus_brownii/default.asp
| accessdate = 2008-08-24
}}</ref><!-- As of August 2007, Fish and Wildlife Service does not recognize a common name - NCN (no common name) -->


A proposal for listing ''A. brownii'' under the U.S. [[Endangered Species Act]] was originally submitted on June 16, 1976, but was withdrawn on December 10, 1979, as out of date and incomplete.<ref name="NatureServe">{{cite web |author=NatureServe |author-link=NatureServe |year=2009 |title=Amaranthus brownii |website=NatureServe Explorer | publisher=NatureServe |location=Arlington, Virginia |url=http://explorer.natureserve.org/servlet/NatureServe?searchName=Amaranthus+brownii |access-date=2010-01-10}}</ref> It was proposed again on March 24, 1993,<ref name="NatureServe" /> and was federally listed as an [[endangered species]] on August 21, 1996.<ref name="FWS996"/> On May 22, 2003, the FWS designated {{convert|171|acre|ha}} on the island of Nihoa as a [[Endangered Species Act#Critical habitat|critical habitat]] for ''A. brownii'', as well as ''Pritchardia remota'', ''Schiedea verticillata'', and two other species also found on Nihoa and other Hawaiian islands, ''{{okina}}ohai'' (''[[Sesbania tomentosa]]'') and ''[[Mariscus pennatiformis]]''.<ref name="FWS003"/> Also in 2003, ''A. brownii'' was internationally classified as [[critically endangered]] on the [[IUCN Red List]].<ref name="iucn status 19 November 2021" /> As of 2010, ''A. brownii'' was one of 51 Hawaiian plant species listed as endangered or threatened under the Endangered Species Act.<ref name="FWS010">{{cite journal|author=United States Fish and Wildlife Service|url=http://www.gpo.gov/fdsys/pkg/FR-2010-04-08/pdf/2010-7915.pdf#page=1|title=Endangered and Threatened Wildlife and Plants; 5-Year Status Reviews of 69 Species in Idaho, Washington, Hawaii, Guam, and the Commonwealth of the Northern Mariana Islands|journal=Federal Register|volume=75|issue=67|pages=17947–17950|publisher=United States Government Printing Office|date=2010-04-08|access-date=2011-03-13}}</ref>
==See also==
*[[List of species of the Northwestern Hawaiian Islands]]


==Notes==
==Notes==
{{refbegin|}}
<div class="references-small" style="-moz-column-count:1; column-count:1;">
'''a.''' {{Note_label|A|a|none}} Wagner and Herbst list five naturalized species of ''[[Amaranthus]]'' 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.<ref name="Wagner-1999"/> ''[[Amaranthus graecizans|A. graecizans]]'', ''[[Amaranthus retroflexus|A. retroflexus]]'', and a third unknown species (possibly extinct or reclassified) have been proposed as additional naturalized candidates.
'''a.''' {{Note label|A|a|none}} Wagner and Herbst list five naturalized species of ''[[Amaranthus]]'' 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.<ref name="Wagner-1999"/> ''[[Amaranthus graecizans|A. graecizans]]'', ''[[Amaranthus retroflexus|A. retroflexus]]'', and a third unknown species (possibly extinct or reclassified) have been proposed as additional naturalized candidates.


{| class="wikitable" align="center" style="margin-left:1em"
{| class="wikitable"
|-
|-
! style="background:white" align="right" width="12%" | Genus
| colspan=5 style="background:white" | Species of ''[[Amaranthus]]'' found in the Hawaiian Islands:
! style="background:white" align="center" width="15%"| Species
! style="background:white" align="center" width="16%"| Common name
! style="background:white" align="center" width="15%"| Status
! style="background:white" align="center" width="40%"| Range
|-
|-
| rowspan=6 style="background:Gainsboro" valign="top" | [[Amaranthus]]
! style="background:white" align="right" | Genus
! style="background:white" align="center" | Species
| ''[[Amaranthus brownii|A. brownii]]''
| align="left" | Brown's amaranth
! style="background:white" align="center" | Common name
! style="background:white" align="center" | Status
| align="left" | Critically Endangered
! style="background:white" align="center" | Range
|-
| rowspan=6 style="background:Gainsboro" valign="top" | ''[[Amaranthus]]''
| ''A. brownii''
| align="left" | none
| align="left" | Extinct
| align="left" | [[Nihoa]] ([[Endemism|end]])
| align="left" | [[Nihoa]] ([[Endemism|end]])
|-
|-
Line 184: Line 63:
| align="left" | Spleen amaranth
| align="left" | Spleen amaranth
| align="left" | n/a
| align="left" | n/a
| align="left" | [[Kauai]], [[Oahu]], [[Lanai]], [[Hawaii (island)|Hawaii]]
| align="left" | [[Kauai]], [[Oahu]], [[Lanai]], [[Hawaii (island)|Hawaii]]
|-
|-
| ''[[Amaranthus hybridus|A. hybridus]]''
| ''[[Amaranthus hybridus|A. hybridus]]''
| align="left" | Green amaranth
| align="left" | Green amaranth
| align="left" | n/a
| align="left" | n/a
| align="left" | [[Oahu]], [[Maui]]
| align="left" | [[Oahu]], [[Maui]]
|-
|-
| ''[[Amaranthus lividus|A. lividus]]''
| ''[[Amaranthus lividus|A. lividus]]''
| align="left" | Purple amaranth
| align="left" | Purple amaranth
| align="left" | n/a
| align="left" | n/a
| align="left" | [[Kauai]], [[Oahu]], [[Maui]], [[Hawaii (island)|Hawaii]]
| align="left" | [[Kauai]], [[Oahu]], [[Maui]], [[Hawaii (island)|Hawaii]]
|-
|-
| ''[[Amaranthus spinosus|A. spinosus]]''
| ''[[Amaranthus spinosus|A. spinosus]]''
| align="left" | Spiny amaranth (''pakai kuku'')
| align="left" | Spiny amaranth (''pakai kuku'')
| align="left" | n/a
| align="left" | n/a
| align="left" | [[Kure Atoll]], [[Kauai]], [[Oahu]], [[Maui]], [[Hawaii (island)|Hawaii]]
| align="left" | [[Kure Atoll]], [[Kauai]], [[Oahu]], [[Maui]], [[Hawaii (island)|Hawaii]]
|-
|-
| ''[[Amaranthus viridis|A. viridis]]''
| ''[[Amaranthus viridis|A. viridis]]''
| align="left" | Slender amaranth (''pakai'', ''{{okina}}aheahea, pakaikai, pakapakai)
| align="left" | Slender amaranth (''pakai'', ''{{okina}}aheahea, pakaikai, pakapakai)
| align="left" | n/a
| align="left" | n/a
| align="left" | [[Kure Atoll]], [[Kaʻula]], [[Kauai]], [[Oahu]], [[Lanai]], [[Maui]], [[Kahoolawe]], [[Hawaii (island)|Hawaii]]
| align="left" | [[Kure Atoll]], [[Kaʻula]], [[Kauai]], [[Oahu]], [[Lanai]], [[Maui]], [[Kahoolawe]], [[Hawaii (island)|Hawaii]]
|-
| colspan=5 style="background:white" | ''A. brownii'' is one of 12 endemic flowering plant species in the family [[Amaranthaceae]] found in the Hawaiian Islands:<ref name="Lilleeng-Rosenberger-2005">{{cite book|last=Lilleeng-Rosenberger|first=Kerin E.|year=2005|title=Growing Hawaii's Native Plants: A Simple Step-by-Step Approach for Every Species|publisher=Mutual Publishing|isbn=1-56647-716-6}}</ref>
|-
! style="background:white" align="right" | Genus
! style="background:white" align="center" | Species
! style="background:white" align="center" | Common name
! style="background:white" align="center" | Status
! style="background:white" align="center" | Range
|-
| rowspan=3 style="background:Gainsboro" valign="top" | ''[[Achyranthes]]''
| ''[[Achyranthes atollensis|A. atollensis]]''
| align="left" | Hawai{{okina}}i chaff flower
| align="left" | Extinct
| align="left" | [[Kure Atoll]], [[Midway Atoll]], [[Pearl and Hermes Atoll]], [[Laysan]] ([[Extinction|ex.]])
|-
| ''[[Achyranthes mutica|A. mutica]]''
| align="left" | Blunt chaff flower
| align="left" | Critically endangered
| align="left" | [[Kauai]] ([[Extinction|ex.]]), [[Hawaii (island)]]
|-
| ''[[Achyranthes splendens|A. splendens]]''
| align="left" | Maui chaff flower
| align="left" | Vulnerable
| align="left" | [[Oahu]], [[Molokai]] ([[Extinction|ex.]]), [[Lanai]], [[Maui]]
|-
| rowspan=1 style="background:Gainsboro" valign="top"| ''[[Amaranthus]]''
| ''A. brownii''
| align="left" |
| align="left" | Extinct
| align="left" | [[Nihoa]]
|-
| rowspan=5 style="background:Gainsboro" valign="top" | ''[[Charpentiera]]''
| ''[[Charpentiera densiflora|C. densiflora]]''
| align="left" | Dense-flowered pāpala (''pāpala'')
| align="left" | Critically endangered
| align="left" | [[Kauai]], [[Maui]]
|-
| ''[[Charpentiera elliptica|C. elliptica]]''
| align="left" | ''Pāpala''
| align="left" |
| align="left" | [[Kauai]]
|-
| ''[[Charpentiera obovata|C. obovata]]''
| align="left" | Broadleaf pāpala (''pāpala'')
| align="left" |
| align="left" | [[Kauai]], [[Oahu]], [[Molokai]], [[Lanai]], [[Maui]], [[Hawaii (island)]]
|-
| ''[[Charpentiera ovata|C. ovata]]''
| align="left" | Koʻolau Range pāpala (''pāpala'')
| align="left" |
| align="left" | [[Oahu]], [[Molokai]], [[Maui]], [[Hawaii (island)]]
|-
| ''[[Charpentiera tomentosa|C. tomentosa]]''
| align="left" | Waiʻanae Range pāpala (''pāpala'')
| align="left" |
| align="left" | [[Oahu]], [[Molokai]], [[Lanai]], [[Maui]], [[Hawaii (island)]]
|-
| rowspan=3 style="background:Gainsboro" valign="top" | ''[[Nototrichium]]''
| ''[[Nototrichium divaricatum|N. divaricatum]]''
| align="left" | Nā Pali rockwort (''kuluʻī'')
| align="left" |
| align="left" | [[Kauai]]
|-
| ''[[Nototrichium humile|N. humile]]''
| align="left" | Kaʻala rockwort (''kuluʻī'')
| align="left" | Endangered
| align="left" | [[Oahu]], [[East Maui]]
|-
| ''[[Nototrichium sandwicense|N. sandwicense]]''
| align="left" | Hawaiʻi rockwort (''kuluʻī'')
| align="left" |
| align="left" | [[Hawaiian Islands|All eight southeastern Hawaiian Islands]]
|-
|-
|}
|}
{{-}}


'''b.''' {{Note label|B|b|none}} Nihoa: fl, fr, June 20, 1923, C. S. Judd No. 2<ref name="Chris1931"/>
'''b.''' {{Note_label|B|b|none}} Native plants found in and around the habitat of ''A. brownii'' include: ''[[Chenopodium oahuense]]'' (Hawaiian Goosefoot), ''[[Eragrostis variablis]]'', ''[[Ipomoea indica]]'', ''[[Ipomoea pes-caprae]]'' ssp. ''brasiliensis'', ''[[Panicum torridum]]'', ''[[Scaevola sericea]]'', ''[[Schiedea verticillata]]'' (Nihoa Carnation), ''[[Sicyos pachycarpus]]'', ''[[Sida fallax]]'', and ''[[Solanum nelsonii]]'' (Nelson's Horsenettle).<ref name="FWS003"/>

</div>
'''c.''' {{Note label|C|c|none}} Herbst & Takeuchi 6545; BISH.<ref name="BISH86"/> Also see the [http://botany.si.edu/pacificislandbiodiversity/hawaiianflora/specs.cfm 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.''' {{Note label|D|d|none}} According to Mark J. Rauzon, anthropologist [[Kenneth Emory]], 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.<ref name=Rauzon-2001>{{cite book| last = Rauzon| first = Mark J.| title = Isles of Refuge: Wildlife and History of the Northwestern Hawaiian Islands | publisher = University of Hawaii Press| year = 2001| page = 12| isbn = 0-8248-2330-3}}</ref>

'''e.''' {{Note label|E|e|none}} Compare the destruction of vegetation on Nihoa in 2004 to that of [[Laysan]] island. In 1894, [[Max Schlemmer]] introduced rabbits to Laysan. Eventually, the rabbit problem and bird [[poaching]] led U.S. President [[Theodore Roosevelt]] 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]] had become extinct. The Tanager Expedition arrived in 1923 and exterminated the last of the surviving rabbits.<ref>{{cite book |title=Max Schlemmer, Hawaii's King of Laysan Island |last=Unger |first=Tom E. |year=2004 |chapter=The Tanager Expedition |publisher=[[iUniverse]] |isbn=978-0-595-29988-1}}</ref>
{{refend}}


==References==
==References==
{{reflist}}
{{Reflist|2}}


==Further reading==
==Further reading==
*{{cite journal|last=Carr|first=Gerald D.|year=1985|hdl=10125/936|title=Additional Chromosome numbers of Hawaiian flowering plants|journal=Pacific Science|publisher=University of Hawaii Press|volume=39|issue=3|pages=302–306|issn=0030-8870}}
*{{cite web|last=Carr|first=Gerald D.|year=2006|url=http://www.botany.hawaii.edu/faculty/carr/natives.htm|title=''Hawaiian Native Plant Genera''|work=Hawaiian Native Plant Propagation Database|publisher=University of Hawaiʻi at Mānoa|access-date=2011-05-12| archive-url= https://web.archive.org/web/20110514231734/http://www.botany.hawaii.edu/faculty/carr/natives.htm| archive-date= 14 May 2011 | url-status= live}}
*{{cite journal|last=Eliasson|first=Uno H.|title=The Evolutionary Patterns of the Plant Family Amaranthaceae on the Galápagos and Hawaiian Islands|journal=Journal of the Torrey Botanical Society|volume=131|issue=2|pages=105–109|publisher=Torrey Botanical Society|year=2004|jstor =4126911|doi=10.2307/4126911}}{{Subscription required}}
*{{cite journal|last=Gagné|first=Wayne C.|author2=Sheila Conant|year=1983|title=Nihoa: Biological Gem of the Northwest Hawaiian Islands|journal=Bishop Museum News|issue=7|publisher=Bernice P. Bishop Museum|pages=3–5}}
*{{cite web|author=Germplasm Resources Information Network (GRIN)|author-link=Germplasm Resources Information Network|title=Taxon: ''Amaranthus brownii'' Christoph. & Caum|work=Taxonomy for Plants |publisher=United States Department of Agriculture, [[Agricultural Research Service|ARS]], National Genetic Resources Program, National Germplasm Resources Laboratory, Beltsville, Maryland| year = 2008|url=https://www.ars-grin.gov/cgi-bin/npgs/html/taxon.pl?415055|access-date=2009-05-29}}
*{{cite book|last=Kartesz|first=John T.|author2=Rosemarie Kartesz|year=1980|title=A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland|url=https://archive.org/details/synonymizedcheck0000kart|url-access=registration|publisher=University of North Carolina Press|isbn=0-8078-1434-2}}
*{{cite news|last=Leone|first=Diana|year=2002|title=Plan to protect NW Hawaiian Isle plants|newspaper=Honolulu Star-Bulletin|url=http://archives.starbulletin.com/2002/05/17/news/story17.html|access-date=2009-05-31}}
*{{cite journal|author=United States Fish and Wildlife Service|year=2002|title=Endangered and Threatened Wildlife and Plants; Proposed Determinations of Prudency and Proposed Designations of Critical Habitat for Plant Species From the Northwestern Hawaiian Islands, HI; Proposed Rule|journal=Federal Register|volume=67|issue=93|pages=34522–34545| publisher=United States Government Printing Office|url=http://frwebgate.access.gpo.gov/cgi-bin/getdoc.cgi?dbname=2002_register&docid=fr14my02-19.pdf|access-date=2009-05-31}}
*{{cite book|last=Yen|first=Douglas E.|year=1969|title=Nihoa — 1969: A Preliminary Report|publisher=Bernice Pauahi Bishop Museum|quote=Unpublished Department of Anthropology Manuscript}}

==External links==
{{Wikispecies|Amaranthus brownii}}
{{Wikispecies|Amaranthus brownii}}
*{{FWS profile|spcode=Q01G |sci=Amaranthus brownii |access-date=May 17, 2011}}
*''[https://web.archive.org/web/20160810235351/http://nsdb.bishopmuseum.org/?w=BPBM&id=-1185059026 Amaranthus brownii Christoph. & Caum]''. ''Hawaii Biological Survey''. [[Bishop Museum]]. Retrieved May 20, 2011.


{{Taxonbar|from=Q4740157}}
<div class="references-2column">

*{{cite book
[[Category:Amaranthus|brownii]]
| last = Carr
[[Category:Endemic flora of Nihoa]]
| first = Gerald D.
[[Category:Extinct flora of Hawaii]]
| authorlink =
[[Category:Plants described in 1931]]
| coauthors =
[[Category:Tanager Expedition]]
| chapter = Chromosome evolution and speciation in Hawaiian flowering plants
[[Category:Plant extinctions since 1500]]
| editor = Tod F. Stuessy, Mikio Ono
| title = Evolution and Speciation of Island Plants
| publisher = [[Cambridge University Press]]
| date = 1998
| location =
| pages =
| url = http://www.cambridge.org/catalogue/catalogue.asp?isbn=0521496535
| isbn = 0521496535
}}
*{{cite book
| last = Christophersen
| first = Erling
| coauthors = Caum, Edward L.
| title = Vascular plants of the Leeward Islands, Hawaii
| publisher = [[Bishop Museum|Bishop Museum Press]]
| series = Bernice P. Bishop Museum Bulletin No. 81; Tanager Expedition Publication No. 7
| date = 1931-07-01
| location = Honolulu, Hawaii
| isbn =
}}
*{{cite news
| last = Gagné
| first = W.C.
| coauthors = S. Conant
| title = Nihoa: biological gem of the Northwest Hawaiian Islands
| publisher = Newslett. of the B.P. Bish. Mus. Assoc.
| date = July 1983
}}
</div>
[[Category:Amaranthus]]
[[Category:Species endemic to the Northwestern Hawaiian Islands]]
[[Category:Nihoa]]

Latest revision as of 08:35, 19 May 2024

Amaranthus brownii
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Amaranthaceae
Genus: Amaranthus
Species:
A. brownii
Binomial name
Amaranthus brownii

Amaranthus brownii was an annual herb in the family Amaranthaceae. The plant was found only on the small island of Nihoa in the Northwestern Hawaiian Islands, growing on rocky outcrops at altitudes of 120–215 m (394–705 ft). It was one of nine species of Amaranthus in the Hawaiian Islands, as well as the only endemic Hawaiian species of the genus. It is now considered extinct.

A. brownii was first discovered during the Tanager Expedition in 1923 by botanist Edward Leonard Caum. It differed from other Hawaiian species of Amaranthus with its spineless leaf axils, linear leaves, and indehiscent 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 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 (FWS), which administers the island of Nihoa as part of the Hawaiian Islands National Wildlife Refuge in the Papahānaumokuākea Marine National Monument. 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 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.[1]

Taxonomy[edit]

The species was first collected during a ten-day visit to the island of Nihoa by the Tanager Expedition.[2] Botanist Edward Leonard Caum collected the first specimen on June 17, 1923, and a second was collected by cartographer Charles S. Judd[b] on June 20, 1923.[3] Forest B. H. Brown, botanist of the Bayard Dominick Expedition to the Marquesas Islands (1921–1922),[4] helped provide descriptions and comments for some of the species described by Erling Christophersen and Caum.[5] They named A. brownii after Brown in 1931[6] 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.[7] The FWS does not recognize a common name.[8]

Description[edit]

Amaranthus brownii was the only endemic species of Hawaiian Amaranthus in the Hawaiian Islands.[a] It was an herbaceous annual plant that grows to a height of 30–90 cm (0.98–2.95 ft) and had narrow, linear leaves, small green flowers, and fruit that held a single, dark red seed. A. brownii was monoecious; that is, the male and female flowers were found together on the same plant.[9] It differed from other Hawaiian species of Amaranthus with its spineless leaf axils, linear leaves, and indehiscent fruits (fruit which does not open to release seeds when ripe).[9] The fruits were ovoid and between 0.8–1 mm long and 0.6–0.8 mm wide.[10] The plant is thought to have been anemophilous (pollinated by wind).[11]

Distribution and habitat[edit]

Papahānaumokuākea Marine National Monument

Amaranthus brownii had a very limited range; it was found only on the 0.65 km2 (0.25 sq mi) island of Nihoa, located 275 km (171 mi) northwest of Kauai. It is thought that this endemic plant had probably always been rare and restricted to Nihoa.[12] Its former habitat is managed by the United States Fish and Wildlife Service and protected as part of the Hawaiian Islands National Wildlife Refuge in the Papahānaumokuākea Marine National Monument.[13] A. brownii was one of three endemic and endangered species only found on Nihoa, along with the Nihoa fan palm (Pritchardia remota) and the Nihoa carnation (Schiedea verticillata).[14] At least nine other native plant species can be found in its habitat, including Hawaiian goosefoot (Chenopodium oahuense), lovegrass (Eragrostis variablis), koali ʻawa (Ipomoea indica), goat's foot (Ipomoea pes-caprae subsp. brasiliensis), Panicum torridum, naupaka (Scaevola sericea), Sicyos pachycarpus, ʻilima (Sida fallax), and Nelson's horsenettle (Solanum nelsonii).[9]

Aerial view of Nihoa. In 1983, W. Gagné recorded 12 plants living on the island.[10]

The plant grew during the moist, winter season from December through July in Nihoa's coastal dry shrubland habitat[12] in shallow soil on rocky outcrops in exposed areas between 120–215 m (394–705 ft).[9] At the time of its discovery in 1923, botanists first observed the plant growing in great quantity on the ridges 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 Douglas E. Yen of the Bishop Museum collected specimens near Miller Peak.[3] Derral R. Herbst and Wayne Takeuchi of the FWS collected the last known specimen[c] on July 27, 1980.[15] Carl C. Christensen also visited Nihoa in 1980 to reevaluate endemic species last observed on the Tanager Expedition.[16] 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:[17] 23 plants were found near Miller Peak and 12 plants in Middle Valley. The two plant populations were separated by a distance of approximately 0.4 km (0.25 mi).[18]

Prehistoric Polynesian habitation on Nihoa[d] may have initially led to a decrease in the plant population of A. brownii.[19] Major threats to the plant included invasive species, fire, and hybridization with other Amaranthus species.[18] Inbreeding was a serious threat, as the small plant population must reproduce within its own circle resulting in genetic defects.[11] A. brownii was also forced to compete with the non-native weed (Portulaca oleracea), the plant's main alien species threat.[9] In 2002 and 2004, the invasive gray bird grasshopper (Schistocerca nitens) presented an even larger threat to A. brownii.[e] First recorded on the island in 1977,[20] the increasing population density of gray bird grasshoppers led to massive defoliation on the island, leaving A. brownii at greater risk of predation.[8] In 2004 alone, an estimated 400,000 gray bird grasshoppers destroyed almost 90% of the vegetation on Nihoa.[20][21] All these threats may have contributed to its ultimate extinction.[1]

Conservation[edit]

Plant populations were found on Miller's Ridge and on the west ridge of Middle Valley[17]

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."[16] 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.[8] Originally, botanists thought the absence of the plant in field surveys might have been explained by the time of visits [11] During the summer months, the stems of A. brownii dried up and could not be distinguished from other herbaceous plants.[8] However, a seven-day visit to the island in April 2006 still did not find any specimens.[8] Additional winter surveys were performed, but none were successful.[1]

Ex situ conservation efforts to propagate A. brownii by seed in botanical gardens were also unsuccessful.[11] During the 1981 expedition, A. bronwii seeds were collected by Sheila Conant and presented to the Waimea Arboretum on the Hawaiian island of Oahu and the Kew Gardens in London, England. Although the seeds at the Waimea Arboretum germinated and grew for a while, no plants survived beyond the stage of seedling development. Information about the outcome of the seeds sent to Kew Gardens is unavailable.[17]

A proposal for listing A. brownii under the U.S. Endangered Species Act was originally submitted on June 16, 1976, but was withdrawn on December 10, 1979, as out of date and incomplete.[22] It was proposed again on March 24, 1993,[22] and was federally listed as an endangered species on August 21, 1996.[6] On May 22, 2003, the FWS designated 171 acres (69 ha) on the island of Nihoa as a critical habitat for A. brownii, as well as Pritchardia remota, Schiedea verticillata, and two other species also found on Nihoa and other Hawaiian islands, ʻohai (Sesbania tomentosa) and Mariscus pennatiformis.[9] Also in 2003, A. brownii was internationally classified as critically endangered on the IUCN Red List.[1] As of 2010, A. brownii was one of 51 Hawaiian plant species listed as endangered or threatened under the Endangered Species Act.[23]

Notes[edit]

a. ^ Wagner and Herbst list five naturalized species of Amaranthus 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.[10] A. graecizans, A. retroflexus, and a third unknown species (possibly extinct or reclassified) have been proposed as additional naturalized candidates.

Species of Amaranthus found in the Hawaiian Islands:
Genus Species Common name Status Range
Amaranthus A. brownii none Extinct Nihoa (end)
A. dubius Spleen amaranth n/a Kauai, Oahu, Lanai, Hawaii
A. hybridus Green amaranth n/a Oahu, Maui
A. lividus Purple amaranth n/a Kauai, Oahu, Maui, Hawaii
A. spinosus Spiny amaranth (pakai kuku) n/a Kure Atoll, Kauai, Oahu, Maui, Hawaii
A. viridis Slender amaranth (pakai, ʻaheahea, pakaikai, pakapakai) n/a Kure Atoll, Kaʻula, Kauai, Oahu, Lanai, Maui, Kahoolawe, Hawaii
A. brownii is one of 12 endemic flowering plant species in the family Amaranthaceae found in the Hawaiian Islands:[24]
Genus Species Common name Status Range
Achyranthes A. atollensis Hawaiʻi chaff flower Extinct Kure Atoll, Midway Atoll, Pearl and Hermes Atoll, Laysan (ex.)
A. mutica Blunt chaff flower Critically endangered Kauai (ex.), Hawaii (island)
A. splendens Maui chaff flower Vulnerable Oahu, Molokai (ex.), Lanai, Maui
Amaranthus A. brownii Extinct Nihoa
Charpentiera C. densiflora Dense-flowered pāpala (pāpala) Critically endangered Kauai, Maui
C. elliptica Pāpala Kauai
C. obovata Broadleaf pāpala (pāpala) Kauai, Oahu, Molokai, Lanai, Maui, Hawaii (island)
C. ovata Koʻolau Range pāpala (pāpala) Oahu, Molokai, Maui, Hawaii (island)
C. tomentosa Waiʻanae Range pāpala (pāpala) Oahu, Molokai, Lanai, Maui, Hawaii (island)
Nototrichium N. divaricatum Nā Pali rockwort (kuluʻī) Kauai
N. humile Kaʻala rockwort (kuluʻī) Endangered Oahu, East Maui
N. sandwicense Hawaiʻi rockwort (kuluʻī) All eight southeastern Hawaiian Islands

b. ^ Nihoa: fl, fr, June 20, 1923, C. S. Judd No. 2[2]

c. ^ Herbst & Takeuchi 6545; BISH.[15] Also see the 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, 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.[25]

e. ^ Compare the destruction of vegetation on Nihoa in 2004 to that of Laysan island. In 1894, Max Schlemmer introduced rabbits to Laysan. Eventually, the rabbit problem and bird poaching led U.S. President Theodore Roosevelt 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 had become extinct. The Tanager Expedition arrived in 1923 and exterminated the last of the surviving rabbits.[26]

References[edit]

  1. ^ a b c d e Bruegmann, M.; Caraway, V.L.; Keir, M. (2018). "Amaranthus brownii". IUCN Red List of Threatened Species. 2018: e.T44077A83794603. doi:10.2305/IUCN.UK.2018-2.RLTS.T44077A83794603.en. Retrieved 19 November 2021.
  2. ^ a b Christophersen, Erling; Caum, Edward L. (1931). Vascular plants of the Leeward Islands, Hawaii. Bernice P. Bishop Museum Bulletin No. 81; Tanager Expedition Publication No. 7. Honolulu, Hawaii: Bishop Museum Press. pp. 25–26. Nihoa: fl, fr, June 17, 1923, E. L. Caum No. 73. Type, B. P. Bishop Mus.
  3. ^ a b Clapp, Roger B.; Eugene Krindler; Robert R. Fleet (1977). F. R. Fosberg; M.-H. Sachet; D. R. Stoddart (eds.). "The Natural History of Nihoa Island, Northwestern Hawaiian Islands" (PDF). Atoll Research Bulletin (207). Washington, D. C.: Smithsonian Institution: 31–32. Archived from the original (PDF) on 2011-06-11. Retrieved 2011-05-12.
  4. ^ Sherff, Earl E. (1932). "Review: Monocotyledons of Southeastern Polynesia". Botanical Gazette. 93 (1). University of Chicago Press: 107–108. doi:10.1086/334237. S2CID 85804033.
  5. ^ Christophersen & Caum 1931, p. 4: "At the time when we started work on the "Tanager" collections Dr. F. B. H. Brown had already described some of the new species and varieties which they contained, and his descriptions are here included He has also passed his opinion on some of the other plants for which we express our indebtedness."
  6. ^ a b United States Fish and Wildlife Service (1996). "Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Three Plants from the Island of Nihoa, Hawaii" (PDF). Federal Register. 61 (163). United States Government Printing Office: 43178–43184. Retrieved 2011-05-12.
  7. ^ Christophersen & Caum 1931, p. 5; Clapp 1977, p. 147.
  8. ^ a b c d e Pacific Islands Fish and Wildlife Office (2007). Amaranthus brownii (No common name); 5-Year Review; Summary and Evaluation (PDF) (Report). United States Fish and Wildlife Service. Retrieved 2010-01-10.
  9. ^ a b c d e f 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.
  10. ^ a b c Wagner, Warren L.; Derral R. Herbst; Sy H. Sohmer (1999). Manual of the Flowering Plants of Hawaiʻi. Vol. 1 (Revised ed.). Honolulu, Hawaii: University of Hawaii Press. pp. 185–189. ISBN 0-8248-2166-1.
  11. ^ a b c d Evenhuis, Neal L.; Eldredge, Lucius G., eds. (2004). Natural History of Nihoa and Necker Islands. Bishop Museum Bulletin in Cultural and Environmental Studies; No. 1. Honolulu, Hawaii: Bishop Museum Press. pp. 56–57. ISBN 1-58178-029-X.
  12. ^ a b United States Fish and Wildlife Service (March 1998). Final Recovery Plan for Three Plant Species on Nihoa Island (PDF) (Report). United States Fish and Wildlife Service. pp. 10547–10550. Retrieved 2011-05-12.
  13. ^ State of Hawaii; National Oceanic and Atmospheric Administration; Office of Hawaiian Affairs; U.S. Fish and Wildlife Service (2009). "The United States of America's Nomination of Papahānaumokuākea Marine National Monument for Inscription on the World Heritage List". Honolulu, Hawaii: Papahānaumokuākea Marine National Monument. Retrieved 2011-05-10.
  14. ^ 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.
  15. ^ a b Wagner, Warren L.; Derral R. Herbst; Sy H. Sohmer (1986). "Contributions to the flora of Hawaiʻi. I. Acanthaceae—Asteraceae". Bishop Museum Occasional Papers. 26–27. Bernice Pauahi Bishop Museum: 102–122.
  16. ^ a b Conant, Sheila; Carl C. Christensen; Patrick Conant; Wayne C. Gagné; M. Lee Goff (1994). "The Unique Terrestrial Biota of the Northwestern Hawaiian Islands" (PDF). In E. Alison Kay (ed.). A Natural History of the Hawaiian Islands: Selected Readings II. Honolulu: University of Hawaii Press. pp. 378–390. ISBN 978-0-8248-1659-9.
  17. ^ a b c Conant, Sheila (1985). "Recent Observations on the Plants of Nihoa Island, Northwestern Hawaiian Islands". Pacific Science. 39 (2). University of Hawaii Press: 135–149. hdl:10125/921. Retrieved 2011-05-17.
  18. ^ a b Hawai'i Department of Land and Natural Resources (2005). "Amaranthus brownii fact sheet" (PDF). Hawaii's Comprehensive Wildlife Conservation Strategy. State of Hawaii. Archived from the original (PDF) on 17 May 2011. Retrieved 12 May 2011.
  19. ^ Beacham, Walton; Frank Vincent Castronova; Suzanne Sessine (2000). Beacham's Guide to the Endangered Species of North America. Arachnids and Crustaceans, Insects, Lichens, Fern Allies, True Ferns, Conifers, Dicots. Vol. 3. Detroit: Gale Group. pp. 1641–1642. ISBN 0-7876-5028-5.
  20. ^ a b 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.
  21. ^ Latchininsky, Alexandre V. (2008). "Grasshopper Outbreak Challenges Conservation Status of a Small Hawaiian Island". Journal of Insect Conservation. 12 (3–4). Springer Netherlands: 343–357. doi:10.1007/s10841-008-9143-8. S2CID 12374828.
  22. ^ a b NatureServe (2009). "Amaranthus brownii". NatureServe Explorer. Arlington, Virginia: NatureServe. Retrieved 2010-01-10.
  23. ^ United States Fish and Wildlife Service (2010-04-08). "Endangered and Threatened Wildlife and Plants; 5-Year Status Reviews of 69 Species in Idaho, Washington, Hawaii, Guam, and the Commonwealth of the Northern Mariana Islands" (PDF). Federal Register. 75 (67). United States Government Printing Office: 17947–17950. Retrieved 2011-03-13.
  24. ^ Lilleeng-Rosenberger, Kerin E. (2005). Growing Hawaii's Native Plants: A Simple Step-by-Step Approach for Every Species. Mutual Publishing. ISBN 1-56647-716-6.
  25. ^ Rauzon, Mark J. (2001). Isles of Refuge: Wildlife and History of the Northwestern Hawaiian Islands. University of Hawaii Press. p. 12. ISBN 0-8248-2330-3.
  26. ^ Unger, Tom E. (2004). "The Tanager Expedition". Max Schlemmer, Hawaii's King of Laysan Island. iUniverse. ISBN 978-0-595-29988-1.

Further reading[edit]

External links[edit]