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{{Short description|Genus of flowering plant}}
{{Short description|Genus of flowering plant}}
{{hatnote|For other plants called saltbush, see [[Saltbush]]}}
{{Hatnote|For other plants called saltbush, see [[Saltbush]]}}
{{Automatic taxobox
{{Automatic_taxobox
|image = Atriplex hortensis cleaned Sturm.png
|image = Atriplex hortensis cleaned Sturm.png
|image_caption = [[Garden orache]] (''[[Atriplex hortensis]]'')<br/>From Sturm & [[Jacob Sturm|Sturm]] (1796): ''Deutschlands Flora in Abbildungen.''
|image_caption = [[Garden orache]] (''[[Atriplex hortensis]]'')<br/>From Sturm & [[Jacob Sturm|Sturm]] (1796): ''Deutschlands Flora in Abbildungen.''
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}}
}}


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'''''Atriplex''''' ({{IPAc-en|ˈ|æ|t|r|ᵻ|p|l|ɛ|k|s}}<ref>''Sunset Western Garden Book,'' 1995:606–607</ref>) is a plant [[genus]] of about 250 species, known by the common names of '''saltbush''' and '''orache''' ({{IPAc-en|ˈ|ɒ|r|ɪ|tʃ|,_|-|ə|tʃ}};<ref>{{Cite book|title=The Chambers Dictionary|publisher=Chambers|year=2003|isbn=0-550-10105-5|edition=9th|chapter=orach}}</ref><ref>{{cite encyclopedia | encyclopedia=[[Oxford English Dictionary]] third edition |title=orache |url=http://www.oed.com/view/Entry/132134 |access-date=7 December 2018| date=June 2004 |publisher=Oxford University Press}}</ref> also spelled '''orach'''). It belongs to the [[subfamily]] [[Chenopodioideae]] of the [[family (biology)|family]] [[Amaranthaceae]] [[sensu|''s.l.'']].

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'''''Atriplex''''' ({{IPAc-en|ˈ|æ|t|r|ᵻ|p|l|ɛ|k|s}}<ref>''Sunset Western Garden Book,'' 1995:606–607</ref>) is a plant [[genus]] of 250–300 [[species]], known by the common names of '''saltbush''' and '''orache''' ({{IPAc-en|ˈ|ɒ|r|ɪ|tʃ|,_|-|ə|tʃ}};<ref>{{Cite book|title=The Chambers Dictionary|publisher=Chambers|year=2003|isbn=0-550-10105-5|edition=9th|chapter=orach}}</ref><ref>{{cite encyclopedia | encyclopedia=[[Oxford English Dictionary]] third edition |title=orache |url=http://www.oed.com/view/Entry/132134 |access-date=7 December 2018| date=June 2004 |publisher=Oxford University Press}}</ref> also spelled '''orach'''). It belongs to the [[subfamily]] [[Chenopodioideae]] of the [[family (biology)|family]] [[Amaranthaceae]] [[sensu|''s.l.'']].
The genus is quite variable and widely distributed. It includes many [[desert]] and [[shore|seashore]] plants and [[halophyte]]s, as well as plants of moist environments.
The genus is quite variable and widely distributed. It includes many [[desert]] and [[shore|seashore]] plants and [[halophyte]]s, as well as plants of moist environments.
The generic name originated in [[Latin]] and was applied by [[Pliny the Elder]] to the edible oraches.<ref>{{cite book |url=https://books.google.com/books?id=esMPU5DHEGgC |first=Umberto |last=Quattrocchi |title=CRC World Dictionary of Plant Names |volume=I: A-C |year=2000 |publisher=CRC Press |isbn=978-0-8493-2675-2 |page=232}}</ref> The name saltbush derives from the fact that the plants retain salt in their leaves; they are able to grow in areas affected by [[soil salination]].
The generic name originated in [[Latin]] and was applied by [[Pliny the Elder]] to the edible oraches.<ref>{{cite book |url=https://books.google.com/books?id=esMPU5DHEGgC |first=Umberto |last=Quattrocchi |title=CRC World Dictionary of Plant Names |volume=I: A-C |year=2000 |publisher=CRC Press |isbn=978-0-8493-2675-2 |page=232}}</ref> The name saltbush derives from the fact that the plants retain salt in their leaves; they are able to grow in areas affected by [[soil salination]].


== Description ==
== Description ==
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Species of plants in genus ''Atriplex'' are [[Annual plant|annual]] or [[perennial]] herbs, [[subshrub]]s, or shrubs. The plants are often covered with bladderlike hairs, that later collapse and form a silvery, scurfy or mealy surface, rarely with elongate [[trichome]]s. The leaves are arranged alternately along the branches, rarely in opposite pairs, either [[Sessility (botany)|sessile]] or on a [[Petiole (botany)|petiole]], and are sometimes deciduous. The leaf blade is variably shaped and may be entire, tooth or lobed.<ref name="FlNAmerica" /><ref name="FlChina" /><ref name="RBGV" /><ref name="RBGS" /><ref name="FloraBase" />
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[[File:Atriplex patula (5129939806).jpg|thumb|''[[Atriplex patula]]'', female flower with [[Bract#bracteole|bracteoles]] and [[ovule]]]]
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The species in genus ''Atriplex'' are [[Annual plant|annual]] or [[perennial]] herbs, [[subshrub]]s, or [[shrubs]]. The plants are often covered with bladderlike hairs, that later collapse and form a silvery, scurfy or mealy surface, rarely with elongate [[trichome]]s. The alternate or rarely opposite [[leaves]] are petiolate or sessile, often persistent or tardily deciduous. The flat or slightly fleshy leaf blades are either entire, or serrate, or lobed and very variable in shape.<ref name="FlNAmerica" /><ref name="FlChina" />


The [[inflorescences]] consist of axillary or terminal spikes or spicate panicles, or axillary clusters of glomeruled flowers. The flowers are [[unisexual]], some species are [[monoecious]], others [[dioecious]]. Male flowers consist of 3-5 [[perianth]] lobes and 3-5 [[stamens]]. Female flowers are usually lacking a perianth, but are enclosed by 2 foliaceous [[bracteole]]s, and contain an ovary with a short style and 2 [[Stigma (botany)|stigmas]].<ref name="FlNAmerica" /><ref name="FlChina" />
The flowers are borne in leaf axils or on the ends of branches, in spikes or spike-like [[panicle]]s . The flowers are [[unisexual]], some species [[monoecious]], others [[dioecious]]. Male flowers have 3-5 [[perianth]] lobes and 3-5 [[stamens]]. Female flowers are usually lacking a perianth, but are enclosed by 2 leaf-like [[bracteole]]s, have a short [[Style (botany)|style]] and 2 [[Stigma (botany)|stigmas]].<ref name="FlNAmerica" /><ref name="FlChina" /><ref name="RBGV" /><ref name="RBGS" /><ref name="FloraBase" />


In fruit, the bracteoles can enlarge, thicken or become appendaged. They enclose the fruit tightly, without becoming connate to it. The pericarp is adnate to the vertically orientated, flattened seed. The seed coat is thick, leathery or hardening. The annular embryo surrounds the perisperm.<ref name="FlNAmerica" /><ref name="FlChina" />
After flowering, the bracteoles sometimes enlarge, thicken or become appendaged, enclosing the fruit but without adhering to it.<ref name="FlNAmerica" /><ref name="FlChina" /><ref name="RBGV" /><ref name="RBGS" /><ref name="FloraBase" />


The [[chromosome]] base number is x = 9, except for ''Atriplex lanfrancoi'', which is x=10.<ref name="Kadereit" />
The [[chromosome]] base number is x = 9, except for ''Atriplex lanfrancoi'', which is x=10.<ref name="Kadereit" />


A few ''Atriplex'' species are [[C3 carbon fixation|C<sub>3</sub>-plants]], but most species are [[C4 carbon fixation|C<sub>4</sub>-plants]], with a characteristic leaf anatomy (kranz anatomy).<ref name="Kadereit" />
A few ''Atriplex'' species are [[C3 carbon fixation|C<sub>3</sub>-plants]], but most species are [[C4 carbon fixation|C<sub>4</sub>-plants]], with a characteristic leaf anatomy, known as kranz anatomy.<ref name="Kadereit" />

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<gallery mode="packed">
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Atriplex elegans.jpg|''[[Atriplex elegans]]''
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Atriplex hymenelytra bracts.jpg|''[[Atriplex hymenelytra]]''
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Starr 050516-1394 Atriplex lentiformis.jpg|''[[Atriplex lentiformis]]''
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Atriplex nummularia.JPG|''[[Atriplex nummularia]]''
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Starr 040125-0020 Chenopodium murale.jpg|''[[Atriplex suberecta]]''
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Atriplex patula (5129939806).jpg|''[[Atriplex patula]]'', female flower with [[bracteoles]] and [[ovule]]
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</gallery>


==Taxonomy==
==Taxonomy==
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{{Further|List of Atriplex species|label1=List of ''Atriplex'' species}}
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The genus ''Atriplex'' was first formally described in 1753 by [[Carl Linnaeus]] in''[[Species Plantarum]]''.<ref name="POWO" /><ref name="L.">{{cite book |last1=Linnaeus |first1=Carl |title=Species Plantarum |date=1753 |volume=2 |pages=1052–1053 |url=https://www.biodiversitylibrary.org/item/84236#page/498/mode/1up |access-date=19 June 2022}}</ref> The genus name was used by [[Pliny the Elder|Pliny]] for orach, or mountain spinach (''[[Atriplex hortensis|A. hortensis]]'').<ref name="Sharr">{{cite book |last1=Sharr |first1=Francis Aubi |last2=George |first2=Alex |title=Western Australian Plant Names and Their Meanings |date=2019 |publisher=Four Gables Press |location=Kardinya, WA |isbn=9780958034180 |page=61 |edition=3rd}}</ref>


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The genus ''Atriplex'' was first formally described in 1753 by [[Carl Linnaeus]] in ''[[Species Plantarum]]''.<ref name="POWO" /><ref name="L.">{{cite book |last1=Linnaeus |first1=Carl |title=Species Plantarum |date=1753 |volume=2 |pages=1052–1053 |url=https://www.biodiversitylibrary.org/item/84236#page/498/mode/1up |access-date=19 June 2022}}</ref><ref name="SpPl" /> The genus name was used by [[Pliny the Elder|Pliny]] for orach, or mountain spinach (''[[Atriplex hortensis|A. hortensis]]'').<ref name="Sharr">{{cite book |last1=Sharr |first1=Francis Aubi |last2=George |first2=Alex |title=Western Australian Plant Names and Their Meanings |date=2019 |publisher=Four Gables Press |location=Kardinya, WA |isbn=9780958034180 |page=61 |edition=3rd}}</ref>
===Species list===
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See [[List of Atriplex species|List of ''Atriplex'' species]]


=== Phylogeny ===
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== Distribution and evolution ==
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The genus evolved in Middle [[Miocene]], the [[C4 carbon fixation|C<sub>4</sub>-photosynthesis]] pathway developed about 14.1–10.9 [[million years ago]] (mya), when the climate became increasingly dry. The genus diversified rapidly and spread over the continents. The C<sub>4</sub> ''Atriplex'' colonized North America probably from Eurasia during the Middle/Late Miocene, about 9.8–8.8 mya, and later spread to South America. Australia was colonized twice by two C<sub>4</sub> lineages, one from Eurasia or America about 9.8–7.8 mya, and one from Central Asia about 6.3–4.8 mya. The last lineage diversified rapidly, and became the ancestor of most Australian ''Atriplex'' species.<ref name="Kadereit" />
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The genus ''Atriplex'' is distributed nearly worldwide from [[subtropical]] to [[temperate]] and to [[subarctic]] regions. Most species-rich are [[Australia]], [[North America]], [[South America]] and [[Eurasia]]. Many species are [[halophyte]]s and are adapted to dry environments with salty soils.<ref name="Kadereit" />

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The genus evolved in Middle [[Miocene]], the [[C4 carbon fixation|C<sub>4</sub>]]-[[photosynthesis]] pathway developed about 14.1-10.9 million years ago, when the climate became increasingly dry. The genus diversified rapidly and spread over the continents. The C<sub>4</sub>''Atriplex'' colonized North America probably from Eurasia during the Middle/Late Miocene, about 9.8-8.8 million years ago, and later spread to South America. Australia was colonized twice by two C<sub>4</sub> lineages, one from Eurasia or America about 9.8-7.8 million years ago, and one from Central Asia about 6.3-4.8 million years ago. The last lineage diversified rapidly, and became the ancestor of most Australian ''Atriplex'' species.<ref name="Kadereit" />

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== Ecology ==
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''Atriplex'' species are used as food plants by the [[larva]]e of some [[Lepidoptera]] species; see the [[list of Lepidoptera that feed on Atriplex|list of Lepidoptera which feed on ''Atriplex'']].<ref>{{Cite web|title=HOSTS - The Hostplants and Caterpillars Database at the Natural History Museum|url=https://www.nhm.ac.uk/our-science/data/hostplants/search/list.dsml?searchPageURL=index.dsml&PGenus=Atriplex&sort=Taxon|access-date=2020-09-28|website=www.nhm.ac.uk}}</ref> For [[spider]]s such as ''[[Phidippus californicus]]'' and other [[arthropod]]s, saltbush plants offer opportunities to hide and hunt in [[habitat]] that is otherwise often quite barren.

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It has been proposed that genus ''Atriplex'' was a main food source in the diet of the extinct giant kangaroo ''[[Procoptodon goliah]]''.<ref>{{Cite journal|last1=Prideaux|first1=Gavin J.|last2=Ayliffe|first2=Linda K.|last3=DeSantis|first3=Larisa R. G.|last4=Schubert|first4=Blaine W.|last5=Murray|first5=Peter F.|last6=Gagan|first6=Michael K.|last7=Cerling|first7=Thure E.|date=2009-07-14|title=Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo|journal=Proceedings of the National Academy of Sciences|language=en|volume=106|issue=28|pages=11646–11650|doi=10.1073/pnas.0900956106|issn=0027-8424|pmid=19556539|pmc=2710660|bibcode=2009PNAS..10611646P|s2cid=19904871|doi-access=free}}</ref> Stable isotopic data suggested that their diet consisted of plants that used the [[C4 photosynthesis|C4 photosynthetic pathway]], and due to their semi-arid distribution, chenopod saltbushes were likely responsible.<ref>{{Cite journal|last1=Kadereit|first1=Gudrun|last2=Mavrodiev|first2=Evgeny V.|last3=Zacharias|first3=Elizabeth H.|last4=Sukhorukov|first4=Alexander P.|date=2010|title=Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis|url=https://bsapubs.onlinelibrary.wiley.com/doi/abs/10.3732/ajb.1000169|journal=American Journal of Botany|language=en|volume=97|issue=10|pages=1664–1687|doi=10.3732/ajb.1000169|pmid=21616801|issn=1537-2197}}</ref><ref>{{cite journal |last1=Prideaux |first1=G. |title=Systematics and Evolution of the Sthenurine Kangaroos |journal= UC Publications in Geological Sciences |date=2004 |doi=10.1525/california/9780520098459.001.0001 |publisher=University of California Press|isbn=9780520098459 |url=https://escholarship.org/uc/item/9z42c7t1 }}</ref><ref>{{Cite journal|last1=DeSantis|first1=Larisa R. G.|last2=Field|first2=Judith H.|last3=Wroe|first3=Stephen|last4=Dodson|first4=John R.|date=May 2017|title=Dietary responses of Sahul (Pleistocene Australia–New Guinea) megafauna to climate and environmental change|journal=Paleobiology|language=en|volume=43|issue=2|pages=181–195|doi=10.1017/pab.2016.50|s2cid=13134989|issn=0094-8373|doi-access=free}}</ref>

== Use by humans ==
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The favored species for human consumption is now usually [[garden orache]] (''A. hortensis''),<ref>{{Cite web|title=Orach facts and health benefits|date=4 February 2020|url=https://www.healthbenefitstimes.com/orach/|access-date=2020-09-28|language=en-US}}</ref><ref>{{Cite web|title=Atriplex hortensis Orach, Garden orache PFAF Plant Database|url=https://pfaf.org/user/Plant.aspx?LatinName=Atriplex+hortensis|access-date=2020-09-28|website=pfaf.org}}</ref><ref>{{Cite web|title=Atriplex hortensis (garden orache)|url=https://www.cabi.org/ISC/datasheet/87637450|access-date=2020-09-28|website=www.cabi.org|language=en}}</ref> but many species are edible and the use of ''Atriplex'' as food is known since at least the late [[Epipaleolithic]] (Mesolithic).

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Common orache (''[[Atriplex patula|A. patula]]'') is attested as an [[archaeophyte]] in northern Europe, and the [[Ertebølle culture]] is presumed to have used it as a food.<ref>{{Cite web|url=http://home3.inet.tele.dk/evakoch/veg-uk.htm|archive-url=https://web.archive.org/web/20060717232907/http://home3.inet.tele.dk/evakoch/veg-uk.htm|url-status=dead|archive-date=2006-07-17|title=Prehistoric plant food of Denmark|last=Koch|first=Eva|date=December 1999}}</ref> Its seed has been found among apparent evidence of cereal preparation and cooking at Late Iron Age villages in Britain.<ref>{{citation|title=North West Cambridge Archaeology: University of Cambridge 2013 Excavations, The Traveller's Rest Sub-site|author=Christopher Evans|publisher=Cambridge Archaeological Unit University Of Cambridge|pages=100–113|year=2015|url=http://www-cau.arch.cam.ac.uk/1271%20North%20West%20Cambridge%20Part%208.pdf}}</ref> In the [[Hebrew Bible|biblical]] [[Book of Job]], ''mallûaḥ'' (מַלּ֣וּחַ, probably [[Mediterranean saltbush]], ''A. halimus'', the major culinary saltbush in the region) is mentioned as food eaten by social outcasts ({{bibleverse||Job|30:4|31}}<ref>Mistranslated as "[[Malva|mallow]]s" in the [[King James Bible]] and as ''Nesseln'' ([[Urtica|nettles]]) in the [[Luther Bible]]</ref>). [[Grey saltbush]] (''[[Atriplex cinerea|A. cinerea]]'') has been used as [[bushfood]] in [[Australia]] since prehistoric times.
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[[Chamiso]] (''A. canescens'') and [[shadscale]] (''A. confertifolia'') were eaten by [[Indigenous peoples of the Americas|Native Americans]], and spearscale (''A. hastata'') was a food in rural [[Eurasia]].

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Studies on ''Atriplex'' species demonstrated their potential use in agriculture. Meat from sheep which have grazed on saltbush has surprisingly high levels of [[vitamin E]], is leaner and more hydrated than regular [[Domestic sheep|lamb]] and has consumer appeal equal to grain-fed lamb.<ref name="Pearce" /> The vitamin E levels could have animal health benefits while extending the shelf-life and maintaining the fresh red colour of saltbush lamb. This effect has been demonstrated for old man saltbush (''[[Atriplex nummularia|A. nummularia]]'') and river saltbush (''[[Atriplex amnicola|A. amnicola]]''). For reasons unknown, sheep seem to prefer the more fibrous, less nutritious river saltbush.<ref name="Norman" /><ref name="Pearce" />

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A study on ''[[Atriplex nummularia]]'' discovered the species have a nitrogen content of 2.5–3.5%, and could potentially be used as a protein supplement for grazing if palatable.<ref>{{cite journal |last1=Wilson, A.D., Harrington |first1=G.N. |title=Nutritive value of Australian browse plants |journal=International Livestock Centre for Africa |date=1980 |page=291 |location=Addis Ababa}}</ref> A subsequent study allowed sheep and goats to voluntarily feed on ''[[Atriplex halimus]]'' and aimed to determine if the saltbush was palatable, and if so, did it provide enough nutrients to supplement the diet of these animals.<ref>{{cite journal |last1=Valderrabano, J., Munoz, F., Delgado |first1=I. |title=Browsing ability and utilisation by sheep and goats of Atriplex hamilus L. shrubs. |journal=Small Ruminant Research |date=1996 |volume=19 |issue=2 |pages=131–136 |url=https://www.academia.edu/10164599 |accessdate=13 April 2020 |location=Zaragoza, Spain|doi=10.1016/0921-4488(95)00754-7 }}</ref> In this study they determined when goats and sheep are given as much ''[[Atriplex halimus|A. halimus]]'' as they like, they do obtain enough nutrients to supplement their diet{{Snd}}unless the animal requirements are higher during pregnancy and milk production.

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Saltbushes are also used as an [[ornamental plant]] in [[landscape|landscaping]] and can be used to prevent [[soil erosion]] in coastal areas.<ref>{{Cite web|title=Saltbushes for dryland salinity management in Western Australia|url=https://www.agric.wa.gov.au/soil-salinity/saltbushes-dryland-salinity-management-western-australia|access-date=2020-09-28|website=www.agric.wa.gov.au|language=en}}</ref><ref>{{Cite web|title=Atriplex semibaccata (Australian saltbush)|url=https://www.cabi.org/isc/datasheet/7903|access-date=2020-09-28|website=www.cabi.org|language=en}}</ref> Old man saltbush (''[[Atriplex nummularia]]'') has also been successfully used to rehabilitate old mining sites around [[Lightning Ridge, New South Wales|Lightning Ridge]] (Australia).

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<gallery mode=packed>
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File:Atriplex elegans.jpg|''Atriplex elegans''
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File:Atriplex hymenelytra bracts.jpg|''Atriplex hymenelytra''
File:Atriplex latifolia.jpg|''Atriplex latifolia''
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File:Starr 050516-1394 Atriplex lentiformis.jpg|''Atriplex lentiformis''
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File:Atriplex nummularia.JPG|''Atriplex nummularia''
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File:Starr 040125-0020 Chenopodium murale.jpg|''Atriplex suberecta''
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</gallery>


== Systematics ==
=== Systematics ===
The genus ''Atriplex'' was first described in 1753 by [[Carl Linnaeus]].<ref name="SpPl" /> The type species ([[lectotype]]) is ''[[Atriplex hortensis]]''.<ref name="Tropicos" /> The name is derived from [[Ancient Greek]] ἀτράφαξυς (''atraphaxys''), "orach", itself a [[Pre-Greek substrate]] loanword.
The type species ([[lectotype]]) is ''Atriplex hortensis''.<ref name="Tropicos" /> The name is derived from [[Ancient Greek]] ἀτράφαξυς (''atraphaxys''), "orach", itself a [[Pre-Greek substrate]] loanword.


''Atriplex'' is an extremely species-rich genus and comprises about 250<ref name="FlChina" />-300<ref name="Kadereit" /> species, with new species still being discovered. An example includes ''Atriplex yeelirrie'', formally described in 2015.<ref>{{cite journal |last1=Shepherd |first1=Kelly |last2=Thiele |first2=Kevin |last3=Sampson |first3=Jane |last4=Coates |first4=David |last5=Bryne |first5=Margaret |title=A rare, new species of Atriplex (Chenopodiaceae) comprising two genetically distinct but morphologically cryptic populations in arid Western Australia: implications for taxonomy and conservation |journal=Australian Systematic Botany |date=2015 |volume=28 |issue=4 |page=235 |doi=10.1071/SB15029 |s2cid=85903596 |url=https://www.publish.csiro.au/sb/SB15029 |access-date=8 September 2021}}</ref>
''Atriplex'' is an extremely species-rich genus and comprises about 250<ref name="FlChina" />-300<ref name="Kadereit" /> species, with new species still being discovered. An example includes ''Atriplex yeelirrie'', formally described in 2015.<ref>{{cite journal |last1=Shepherd |first1=Kelly |last2=Thiele |first2=Kevin |last3=Sampson |first3=Jane |last4=Coates |first4=David |last5=Bryne |first5=Margaret |date=2015 |title=A rare, new species of Atriplex (Chenopodiaceae) comprising two genetically distinct but morphologically cryptic populations in arid Western Australia: implications for taxonomy and conservation |url=https://www.publish.csiro.au/sb/SB15029 |journal=Australian Systematic Botany |volume=28 |issue=4 |page=235 |doi=10.1071/SB15029 |access-date=8 September 2021 |s2cid=85903596}}</ref>


Traditional [[taxonomy (biology)|taxonomy]] of [[Atripliceae]] based on morphological features has been controversial.<ref name ='Cladistic'>{{Cite journal|title = A Cladistic Analysis of Atripliceae (Chenopodiaceae) Based on Morphological Data|url = https://www.researchgate.net/publication/274809339|journal = Journal of the Torrey Botanical Society|volume = 128|issue = 3|doi = 10.2307/3088719|first1 = Hilda|last1 = Flores|first2 = Jerrold I.|last2 = Davis|year = 2001|page = 297|jstor = 3088719}}</ref> Molecular studies have found that many genera are not true [[clade]]s. One such study found that Atripliceae could be divided into two main clades, ''Archiatriplex'', with a few, scattered species, and the larger ''Atriplex'' clade, which is highly diverse and found around the world.<ref name ='Cladistic'/> After [[phylogenetic]] research, Kadereit et al. (2010) excluded ''[[Halimione]]'' as a distinct sister genus. The remaining ''Atriplex'' species were grouped into several clades.<ref name="Kadereit" />
Traditional [[taxonomy (biology)|taxonomy]] of [[Atripliceae]] based on morphological features has been controversial.<ref name="Cladistic">{{Cite journal |last1=Flores |first1=Hilda |last2=Davis |first2=Jerrold I. |year=2001 |title=A Cladistic Analysis of Atripliceae (Chenopodiaceae) Based on Morphological Data |url=https://www.researchgate.net/publication/274809339 |journal=Journal of the Torrey Botanical Society |volume=128 |issue=3 |page=297 |doi=10.2307/3088719 |jstor=3088719}}</ref> Molecular studies have found that many genera are not true [[clade]]s. One such study found that Atripliceae could be divided into two main clades, ''Archiatriplex'', with a few, scattered species, and the larger ''Atriplex'' clade, which is highly diverse and found around the world.<ref name="Cladistic" /> After [[phylogenetic]] research, Kadereit et al. (2010) excluded ''[[Halimione]]'' as a distinct sister genus. The remaining ''Atriplex'' species were grouped into several clades.<ref name="Kadereit" />


The following is a cladogram with estimated divergence times for the tribe Atripliceae. To infer the phylogeny, an ITS matrix composed of spacer ITS-1, the 5.8S subunit, and spacer ITS-2 were amplified and sequenced for each specimen. Not all species in the genus ''Atriplex'' are presented in the cladogram (based on page 7 of <ref name="unexpected insights">{{cite journal |last1=Brignone |first1=Nicolás F. |last2=Pozner |first2=Raúl E. |last3=Denham |first3=Silvia S. |title=Origin and evolution of Atriplex (Amaranthaceae s.l.) in the Americas: Unexpected insights from South American species |journal= Taxon|date=October 2019 |volume=68 |issue=5 |page=1027 |doi=10.1002/tax.12133 |s2cid=214371753 }}</ref>). This work suggested that the Americas were colonised by C4 ''Atriplex'' from Eurasia or Australia. Furthermore, that in the Americas ''Atriplex'' first appeared in South America, where two lineages underwent ''in situ'' diversification and evolved sympatrically. North America was then colonised by ''Atriplex'' from South America, then one lineage later moved back to South America.<ref name="unexpected insights" />
The following is a cladogram with estimated divergence times for the tribe Atripliceae. To infer the phylogeny, an ITS matrix composed of spacer ITS-1, the 5.8S subunit, and spacer ITS-2 were amplified and sequenced for each specimen. Not all species in the genus ''Atriplex'' are presented in the cladogram (based on page 7 of <ref name="unexpected insights">{{cite journal |last1=Brignone |first1=Nicolás F. |last2=Pozner |first2=Raúl E. |last3=Denham |first3=Silvia S. |date=October 2019 |title=Origin and evolution of Atriplex (Amaranthaceae s.l.) in the Americas: Unexpected insights from South American species |journal=Taxon |volume=68 |issue=5 |page=1027 |doi=10.1002/tax.12133 |s2cid=214371753}}</ref>). This work suggested that the Americas were colonised by C4 ''Atriplex'' from Eurasia or Australia. Furthermore, that in the Americas ''Atriplex'' first appeared in South America, where two lineages underwent ''in situ'' diversification and evolved sympatrically. North America was then colonised by ''Atriplex'' from South America, then one lineage later moved back to South America.<ref name="unexpected insights" />
{{collapse top|title=Cladogram of estimated divergences within the genus ''Atriplex''}}
{{collapse top|title=Cladogram of estimated divergences within the genus ''Atriplex''}}
[[File:World distribution of Atriplex.svg|thumb|upright=2|right|
[[File:World distribution of Atriplex.svg|thumb|upright=2|right|
Line 492: Line 470:
** ''[[Atriplex vesicaria]]'' Heward ex Benth. &ndash; [[Bladder saltbush]]: in Australia.
** ''[[Atriplex vesicaria]]'' Heward ex Benth. &ndash; [[Bladder saltbush]]: in Australia.
{{collapse bottom}}
{{collapse bottom}}

⚫
== Distribution and habitat ==
⚫
The genus ''Atriplex'' is distributed nearly worldwide from [[subtropical]] to [[temperate]] and to [[subarctic]] regions. Most species-rich are [[Australia]], [[North America]], [[South America]] and [[Eurasia]]. Many species are [[halophyte]]s and are adapted to dry environments with salty soils.<ref name="Kadereit" />

⚫
== Ecology ==
⚫
''Atriplex'' species are used as food plants by the [[larva]]e of some [[Lepidoptera]] species; see the list of Lepidoptera which feed on ''Atriplex''.<ref>{{Cite journal|title=HOSTS - The Hostplants and Caterpillars Database at the Natural History Museum|url=https://www.nhm.ac.uk/our-science/data/hostplants/search/list.dsml?searchPageURL=index.dsml&PGenus=Atriplex&sort=Taxon|access-date=2020-09-28|website=www.nhm.ac.uk| date=2023 | doi=10.5519/havt50xw | last1=Robinson | first1=Gaden S. | last2=Ackery | first2=Phillip R. | last3=Kitching | first3=Ian | last4=Beccaloni | first4=George W. | last5=Hernández | first5=Luis M. }}</ref> They are also sometimes consumed by camels.<ref>{{Cite journal |last=Shawket |first=S. M. |date=1999 |title=Fattening of camel calves on saltbush, Atriplex halimas with different energy sources |url=https://scholar.google.com/scholar_lookup?title=Fattening+of+camel+calves+on+saltbush%2C+Atriplex+halimas+with+different+energy+sources&author=Shawket%2C+S.M.+%28Ministry+of+Agriculture%2C+Cairo+%28Egypt%29.+Desert+Research+Centre%29&publication_year=1999 |journal=Mansoura University, Journal of Agricultural Sciences (Egypt) |language=English |issn=1110-0346}}</ref> For [[spider]]s such as ''[[Phidippus californicus]]'' and other [[arthropod]]s, saltbush plants offer opportunities to hide and hunt in [[habitat]] that is otherwise often quite barren.

⚫
It has been proposed that genus ''Atriplex'' was a main food source in the diet of the extinct giant kangaroo ''[[Procoptodon goliah]]''.<ref>{{Cite journal|last1=Prideaux|first1=Gavin J.|last2=Ayliffe|first2=Linda K.|last3=DeSantis|first3=Larisa R. G.|last4=Schubert|first4=Blaine W.|last5=Murray|first5=Peter F.|last6=Gagan|first6=Michael K.|last7=Cerling|first7=Thure E.|date=2009-07-14|title=Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo|journal=Proceedings of the National Academy of Sciences|language=en|volume=106|issue=28|pages=11646–11650|doi=10.1073/pnas.0900956106|issn=0027-8424|pmid=19556539|pmc=2710660|bibcode=2009PNAS..10611646P|s2cid=19904871|doi-access=free}}</ref> Stable isotopic data suggested that their diet consisted of plants that used the [[C4 photosynthesis|C4 photosynthetic pathway]], and due to their semi-arid distribution, chenopod saltbushes were likely responsible.<ref>{{Cite journal|last1=Kadereit|first1=Gudrun|last2=Mavrodiev|first2=Evgeny V.|last3=Zacharias|first3=Elizabeth H.|last4=Sukhorukov|first4=Alexander P.|date=2010|title=Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis|url=https://bsapubs.onlinelibrary.wiley.com/doi/abs/10.3732/ajb.1000169|journal=American Journal of Botany|language=en|volume=97|issue=10|pages=1664–1687|doi=10.3732/ajb.1000169|pmid=21616801|issn=1537-2197}}</ref><ref>{{cite journal |last1=Prideaux |first1=G. |title=Systematics and Evolution of the Sthenurine Kangaroos |journal= UC Publications in Geological Sciences |date=2004 |doi=10.1525/california/9780520098459.001.0001 |publisher=University of California Press|isbn=9780520098459 |url=https://escholarship.org/uc/item/9z42c7t1 }}</ref><ref>{{Cite journal|last1=DeSantis|first1=Larisa R. G.|last2=Field|first2=Judith H.|last3=Wroe|first3=Stephen|last4=Dodson|first4=John R.|date=May 2017|title=Dietary responses of Sahul (Pleistocene Australia–New Guinea) megafauna to climate and environmental change|journal=Paleobiology|language=en|volume=43|issue=2|pages=181–195|doi=10.1017/pab.2016.50|bibcode=2017Pbio...43..181D |s2cid=13134989|issn=0094-8373|doi-access=free}}</ref>

== Uses ==
⚫
The favored species for human consumption is now usually [[garden orache]] (''A. hortensis''),<ref>{{Cite web|title=Orach facts and health benefits|date=4 February 2020|url=https://www.healthbenefitstimes.com/orach/|access-date=2020-09-28|language=en-US}}</ref><ref>{{Cite web|title=Atriplex hortensis Orach, Garden orache PFAF Plant Database|url=https://pfaf.org/user/Plant.aspx?LatinName=Atriplex+hortensis|access-date=2020-09-28|website=pfaf.org}}</ref><ref>{{Cite web|title=Atriplex hortensis (garden orache)|url=https://www.cabi.org/ISC/datasheet/87637450|access-date=2020-09-28|website=www.cabi.org|language=en}}</ref> but many species are edible and the use of ''Atriplex'' as food is known since at least the late [[Epipaleolithic]] (Mesolithic).

⚫
Common orache (''[[Atriplex patula|A. patula]]'') is attested as an [[archaeophyte]] in northern Europe, and the [[Ertebølle culture]] is presumed to have used it as a food.<ref>{{Cite web|url=http://home3.inet.tele.dk/evakoch/veg-uk.htm|archive-url=https://web.archive.org/web/20060717232907/http://home3.inet.tele.dk/evakoch/veg-uk.htm|url-status=dead|archive-date=2006-07-17|title=Prehistoric plant food of Denmark|last=Koch|first=Eva|date=December 1999}}</ref> Its seed has been found among apparent evidence of cereal preparation and cooking at Late Iron Age villages in Britain.<ref>{{citation|title=North West Cambridge Archaeology: University of Cambridge 2013 Excavations, The Traveller's Rest Sub-site|author=Christopher Evans|publisher=Cambridge Archaeological Unit University Of Cambridge|pages=100–113|year=2015|url=http://www-cau.arch.cam.ac.uk/1271%20North%20West%20Cambridge%20Part%208.pdf}}</ref> In the [[Hebrew Bible|biblical]] [[Book of Job]], ''mallûaḥ'' (מַלּ֣וּחַ, probably [[Mediterranean saltbush]], ''A. halimus'', the major culinary saltbush in the region) is mentioned as food eaten by social outcasts ({{bibleverse||Job|30:4|31}}<ref>Mistranslated as "[[Malva|mallow]]s" in the [[King James Bible]] and as ''Nesseln'' ([[Urtica|nettles]]) in the [[Luther Bible]]</ref>). [[Grey saltbush]] (''[[Atriplex cinerea|A. cinerea]]'') has been used as [[bushfood]] in [[Australia]] since prehistoric times.
⚫
[[Chamiso]] (''A. canescens'') and [[shadscale]] (''A. confertifolia'') were eaten by [[Indigenous peoples of the Americas|Native Americans]], and spearscale (''A. hastata'') was a food in rural [[Eurasia]].

⚫
Studies on ''Atriplex'' species demonstrated their potential use in agriculture. Meat from sheep which have grazed on saltbush has surprisingly high levels of [[vitamin E]], is leaner and more hydrated than regular [[Domestic sheep|lamb]] and has consumer appeal equal to grain-fed lamb.<ref name="Pearce" /> The vitamin E levels could have animal health benefits while extending the shelf-life and maintaining the fresh red colour of saltbush lamb. This effect has been demonstrated for old man saltbush (''[[Atriplex nummularia|A. nummularia]]'') and river saltbush (''[[Atriplex amnicola|A. amnicola]]''). For reasons unknown, sheep seem to prefer the more fibrous, less nutritious river saltbush.<ref name="Norman" /><ref name="Pearce" />

⚫
A study on ''A. nummularia'' discovered the species have a nitrogen content of 2.5–3.5%, and could potentially be used as a protein supplement for grazing if palatable.<ref>{{cite journal |author1=Wilson, A.D.|author2= Harrington, G.N. |title=Nutritive value of Australian browse plants |journal=International Livestock Centre for Africa |date=1980 |page=291 |location=Addis Ababa}}</ref> A subsequent study allowed sheep and goats to voluntarily feed on ''[[Atriplex halimus]]'' and aimed to determine if the saltbush was palatable, and if so, did it provide enough nutrients to supplement the diet of these animals.<ref>{{cite journal |author1=Valderrabano, J.|author2=Munoz, F. |author3=Delgado, I. |title=Browsing ability and utilisation by sheep and goats of Atriplex hamilus L. shrubs. |journal=Small Ruminant Research |date=1996 |volume=19 |issue=2 |pages=131–136 |url=https://www.academia.edu/10164599 |access-date=13 April 2020 |location=Zaragoza, Spain|doi=10.1016/0921-4488(95)00754-7 }}</ref> In this study they determined when goats and sheep are given as much ''[[Atriplex halimus|A. halimus]]'' as they like, they do obtain enough nutrients to supplement their diet{{Snd}}unless the animal requirements are higher during pregnancy and milk production.

⚫
Saltbushes are also used as an [[ornamental plant]] in [[landscape|landscaping]] and can be used to prevent [[soil erosion]] in coastal areas.<ref>{{Cite web|title=Saltbushes for dryland salinity management in Western Australia|url=https://www.agric.wa.gov.au/soil-salinity/saltbushes-dryland-salinity-management-western-australia|access-date=2020-09-28|website=www.agric.wa.gov.au|language=en}}</ref><ref>{{Cite web|title=Atriplex semibaccata (Australian saltbush)|url=https://www.cabi.org/isc/datasheet/7903|access-date=2020-09-28|website=www.cabi.org|language=en}}</ref> Old man saltbush (''[[Atriplex nummularia]]'') has also been successfully used to rehabilitate old mining sites around [[Lightning Ridge, New South Wales|Lightning Ridge]] (Australia).


== See also ==
== See also ==
*[[Barbara Hulme]], produced Atriplex hybrids
*[[Barbara Hulme]], producer of Atriplex hybrids


==References ==
==References ==
Line 500: Line 498:
<ref name="FlNAmerica">Stanley L. Welsh: [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=103110 ''Atriplex'' - online], In: Flora of North America Editorial Committee (ed.): ''Flora of North America North of Mexico'', Volume 4: ''Magnoliophyta: Caryophyllidae, part 1.'', Oxford University Press, New York. 2003, {{ISBN|0-19-517389-9}}.</ref>
<ref name="FlNAmerica">Stanley L. Welsh: [http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=103110 ''Atriplex'' - online], In: Flora of North America Editorial Committee (ed.): ''Flora of North America North of Mexico'', Volume 4: ''Magnoliophyta: Caryophyllidae, part 1.'', Oxford University Press, New York. 2003, {{ISBN|0-19-517389-9}}.</ref>
<ref name="FlChina">Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants: ''Chenopodiaceae'': [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=103110 ''Atriplex'' - online], In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (ed.): ''Flora of China'', Volume 5: ''Ulmaceae through Basellaceae.'', Science Press und Missouri Botanical Garden Press, Beijing und St. Louis, 2003, {{ISBN|1-930723-27-X}}.</ref>
<ref name="FlChina">Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants: ''Chenopodiaceae'': [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=103110 ''Atriplex'' - online], In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (ed.): ''Flora of China'', Volume 5: ''Ulmaceae through Basellaceae.'', Science Press und Missouri Botanical Garden Press, Beijing und St. Louis, 2003, {{ISBN|1-930723-27-X}}.</ref>
<ref name="RBGV">{{cite web |last1=Walsh |first1=Neville G. |title=''Atriples'' |url=https://vicflora.rbg.vic.gov.au/flora/taxon/c41cd3e8-c729-4e74-b251-293ce28c8557 |publisher=Royal Botanic Gardens Victoria |access-date=16 July 2022}}</ref>
<ref name="RBGS">{{cite web |last1=Jacobs |first1=Surrey W.L. |title=''Atriplex'' |url=https://plantnet.rbgsyd.nsw.gov.au/cgi-bin/NSWfl.pl?page=nswfl&showsyn=&dist=&constat=&lvl=gn&name=Atriplex |publisher=Royal Botanic Garden Sydney |access-date=16 July 2022}}</ref>
<ref name=FloraBase>{{FloraBase|name=''Atriplex''|id=21347}}</ref>
<ref name="Kadereit">Gudrun Kadereit, Evgeny V. Mavrodiev, Elizabeth H. Zacharias & Alexander P. Sukhorukov: Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 Photosynthesis. - ''American Journal of Botany'' 97(10): 1664-1687, 2010.</ref>
<ref name="Kadereit">Gudrun Kadereit, Evgeny V. Mavrodiev, Elizabeth H. Zacharias & Alexander P. Sukhorukov: Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 Photosynthesis. - ''American Journal of Botany'' 97(10): 1664-1687, 2010.</ref>
<ref name="Norman">{{aut|Norman, Hayley C.; Freind, Colby; Masters, David G.; Rintoul, Allan J.; Dynes, Robyn A. & Williams, Ian H.}} (2004): Variation within and between two saltbush species in plant composition and subsequent selection by sheep. ''[[Australian Journal of Agricultural Research|Aust. J. Agr. Res.]]'' '''55'''(9): 999–1007. {{doi|10.1071/AR04031}} (HTML abstract)</ref>
<ref name="Norman">{{aut|Norman, Hayley C.; Freind, Colby; Masters, David G.; Rintoul, Allan J.; Dynes, Robyn A. & Williams, Ian H.}} (2004): Variation within and between two saltbush species in plant composition and subsequent selection by sheep. ''[[Australian Journal of Agricultural Research|Aust. J. Agr. Res.]]'' '''55'''(9): 999–1007. {{doi|10.1071/AR04031}} (HTML abstract)</ref>
<ref name="Pearce">{{aut|Pearce, Kelly & Jacob, Robin}} (2004): Saltbush lifts sheep meat vitamin content. ''Farming Ahead'' '''153'''(October): 63. [http://www.csiro.au/proprietaryDocuments/Saltbush_vitamin.pdf PDF fulltext] {{webarchive |url=https://web.archive.org/web/20050723234637/http://www.csiro.au/proprietaryDocuments/Saltbush_vitamin.pdf |date=July 23, 2005 }}</ref>
<ref name="Pearce">{{aut|Pearce, Kelly & Jacob, Robin}} (2004): Saltbush lifts sheep meat vitamin content. ''Farming Ahead'' '''153'''(October): 63. [http://www.csiro.au/proprietaryDocuments/Saltbush_vitamin.pdf PDF fulltext] {{webarchive |url=https://web.archive.org/web/20050723234637/http://www.csiro.au/proprietaryDocuments/Saltbush_vitamin.pdf |date=July 23, 2005 }}</ref>
<ref name="SpPl">{{citation |chapter-url=https://www.biodiversitylibrary.org/page/359073 |volume=2 |chapter=''Atriplex'' |title= ''[[Species Plantarum]]'' |author=Carl Linnaeus |publisher=Lars Salvius/Biodiversity Heritage Library |date=1753|accessdate=19 May 2015 |pages=1052–1054}}</ref>
<ref name="SpPl">{{citation |chapter-url=https://www.biodiversitylibrary.org/page/359073 |volume=2 |chapter=''Atriplex'' |title= ''[[Species Plantarum]]'' |author=Carl Linnaeus |publisher=Lars Salvius/Biodiversity Heritage Library |date=1753|access-date=19 May 2015 |pages=1052–1054}}</ref>
<ref name="Tropicos">[http://www.tropicos.org/Name/40022497 ''Atriplex''] at Tropicos, accessed 2013-07-11</ref>
<ref name="Tropicos">[http://www.tropicos.org/Name/40022497 ''Atriplex''] at Tropicos, accessed 2013-07-11</ref>
</references>
</references>
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{{Taxonbar|from=Q157801}}
{{Taxonbar|from=Q157801}}
{{Authority control}}


[[Category:Atriplex| ]]
[[Category:Atriplex| ]]

Latest revision as of 20:47, 18 October 2023

For other plants called saltbush, see Saltbush

Atriplex
Garden orache (Atriplex hortensis)
From Sturm & Sturm (1796): Deutschlands Flora in Abbildungen.
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Amaranthaceae
Subfamily: Chenopodioideae
Tribe: Atripliceae
Genus: Atriplex
L.[1]
Type species
Atriplex hortensis
L.
Species

See List of Atriplex species

Synonyms[1]
List
  • Armola (Kirschl.) Montandon
    Blackiella Aellen
    Cremnophyton Brullo & Pavone
    Halimione Aellen
    Halimus Wallr.
    Haloxanthium Ulbr.
    Lophocarya Nutt. ex Moq.
    Morrisiella Aellen
    Neopreissia Ulbr.
    Obione Gaertn.
    Pachypharynx Aellen
    Phyllocarpa Nutt. ex Moq.
    Pterocarya Nutt. ex Moq.
    Pterochiton Torr. & Frém.
    Schizotheca (C.A.Mey.) Lindl.
    Senniella Aellen
    Sukhorukovia Vasjukov
    Teutliopsis (Dumort.) Celak.
    Theleophyton Moq.

Atriplex (/ˈætrɪplɛks/[2]) is a plant genus of about 250 species, known by the common names of saltbush and orache (/ˈɒrɪtʃ, -ətʃ/;[3][4] also spelled orach). It belongs to the subfamily Chenopodioideae of the family Amaranthaceae s.l.. The genus is quite variable and widely distributed. It includes many desert and seashore plants and halophytes, as well as plants of moist environments. The generic name originated in Latin and was applied by Pliny the Elder to the edible oraches.[5] The name saltbush derives from the fact that the plants retain salt in their leaves; they are able to grow in areas affected by soil salination.

Description

[edit]

Species of plants in genus Atriplex are annual or perennial herbs, subshrubs, or shrubs. The plants are often covered with bladderlike hairs, that later collapse and form a silvery, scurfy or mealy surface, rarely with elongate trichomes. The leaves are arranged alternately along the branches, rarely in opposite pairs, either sessile or on a petiole, and are sometimes deciduous. The leaf blade is variably shaped and may be entire, tooth or lobed.[6][7][8][9][10]

The flowers are borne in leaf axils or on the ends of branches, in spikes or spike-like panicles . The flowers are unisexual, some species monoecious, others dioecious. Male flowers have 3-5 perianth lobes and 3-5 stamens. Female flowers are usually lacking a perianth, but are enclosed by 2 leaf-like bracteoles, have a short style and 2 stigmas.[6][7][8][9][10]

After flowering, the bracteoles sometimes enlarge, thicken or become appendaged, enclosing the fruit but without adhering to it.[6][7][8][9][10]

The chromosome base number is x = 9, except for Atriplex lanfrancoi, which is x=10.[11]

A few Atriplex species are C3-plants, but most species are C4-plants, with a characteristic leaf anatomy, known as kranz anatomy.[11]

Taxonomy

[edit]
Further information: List of Atriplex species

The genus Atriplex was first formally described in 1753 by Carl Linnaeus in Species Plantarum.[1][12][13] The genus name was used by Pliny for orach, or mountain spinach (A. hortensis).[14]

Phylogeny

[edit]

The genus evolved in Middle Miocene, the C4-photosynthesis pathway developed about 14.1–10.9 million years ago (mya), when the climate became increasingly dry. The genus diversified rapidly and spread over the continents. The C4 Atriplex colonized North America probably from Eurasia during the Middle/Late Miocene, about 9.8–8.8 mya, and later spread to South America. Australia was colonized twice by two C4 lineages, one from Eurasia or America about 9.8–7.8 mya, and one from Central Asia about 6.3–4.8 mya. The last lineage diversified rapidly, and became the ancestor of most Australian Atriplex species.[11]

Systematics

[edit]

The type species (lectotype) is Atriplex hortensis.[15] The name is derived from Ancient Greek ἀτράφαξυς (atraphaxys), "orach", itself a Pre-Greek substrate loanword.

Atriplex is an extremely species-rich genus and comprises about 250[7]-300[11] species, with new species still being discovered. An example includes Atriplex yeelirrie, formally described in 2015.[16]

Traditional taxonomy of Atripliceae based on morphological features has been controversial.[17] Molecular studies have found that many genera are not true clades. One such study found that Atripliceae could be divided into two main clades, Archiatriplex, with a few, scattered species, and the larger Atriplex clade, which is highly diverse and found around the world.[17] After phylogenetic research, Kadereit et al. (2010) excluded Halimione as a distinct sister genus. The remaining Atriplex species were grouped into several clades.[11]

The following is a cladogram with estimated divergence times for the tribe Atripliceae. To infer the phylogeny, an ITS matrix composed of spacer ITS-1, the 5.8S subunit, and spacer ITS-2 were amplified and sequenced for each specimen. Not all species in the genus Atriplex are presented in the cladogram (based on page 7 of [18]). This work suggested that the Americas were colonised by C4 Atriplex from Eurasia or Australia. Furthermore, that in the Americas Atriplex first appeared in South America, where two lineages underwent in situ diversification and evolved sympatrically. North America was then colonised by Atriplex from South America, then one lineage later moved back to South America.[18]

Cladogram of estimated divergences within the genus Atriplex
Background colour in cladogram represents the region where a species is endemic.
  South America
  North America
  Australia
  Eurasia
Brignone et al. (2019) hypothesis for the evolution and movement of Atriplex species globally.[18]
Atripliceae
Atriplex
Halimione
Archiatriplex clade

Extriplex californica

(13 other taxa)

Clades of Atriplex
  • Atriplex lanfrancoi/cana-Clade:
    • Atriplex lanfrancoi (Brullo & Pavone) G. Kadereit et Sukhor. (Syn.: Cremnophyton lanfrancoi Brullo & Pavone): endemic to Malta and Gozo.
    • Atriplex cana C.A. Mey.: from Eastern European Russia to western China.
  • Atriplex section Atriplex: annual C3-plants.
  • Atriplex section Teutliopsis Dumort.: annual C3-plants.
    • Atriplex australasica Moq.
    • Atriplex calotheca (Rafn) Fr.: in Northern Europe.
    • Atriplex davisii Aellen: from southern Europe to Egypt.
    • Atriplex glabriuscula Edmondston – Northeastern saltbush, Babington's orache, smooth orache, Scotland orache, glabrous orache: In central and northern Europe.
    • Atriplex gmelinii C.A. Mey. ex Bong. – Gmelin's saltbush: in Asia and North America.
    • Atriplex intracontinentalis Sukhor.: from Central Europe to Asia.
    • Atriplex laevis C.A. Mey.: in Asia, naturalized in eastern Europe.
    • Atriplex latifolia Wahlenb.: in Eurasia.
    • Atriplex littoralis L. – Grass-leaved orache: in Eurasia and North Africa.
    • Atriplex longipes Drejer – Long-stalked orache: in northern Europe.
    • Atriplex micrantha C.A. Mey.: in Asia, naturalized in Europe.
    • Atriplex nudicaulis Boguslaw – Baltic saltbush: in Eurasia.
    • Atriplex patula L. – Common orache, spreading orache: in Eurasia and North Africa.
    • Atriplex praecox Hülph. – Early orache: in northern Europe.
    • Atriplex prostrata Moq. – Spear-leaved orache, thin-leaved orache, triangle orache, fat hen: in Eurasia and North Africa.
  • C4-Atriplex-Clade: containing the majority of species. The traditional classification into sections (sect. Obione, sect. Pterochiton, sect. Psammophila, sect. Sclerocalymma, sect. Stylosa) did not reflect the phylogenetical relationships and was rejected by Kadereit et al. (2010).[11]
    • Atriplex acanthocarpa (Torr.) S. Watson: in North America.
    • Atriplex acutibractea Anderson: in Australia.
    • Atriplex altaica Sukhor.: in Asia.
    • Atriplex angulata Benth.: in Australia.
    • Atriplex billardierei (Moq.) Hook. f.: in Australia.
    • Atriplex canescens (Pursh) Nutt. – Chamiso, chamiza, four-winged saltbush, grey sagebrush: in North America.
    • Atriplex centralasiatica Iljin: in Asia.
    • Atriplex cinerea Poir. – Grey saltbush, truganini:[19] in Australia
    • Atriplex codonocarpa P.G. Wilson: in Australia.
    • Atriplex conduplicata F. Muell.: in Australia.
    • Atriplex confertifolia (Torr. & Frém.) S. Watson – Shadscale (saltbush): in North America.
    • Atriplex cordobensis Gand. & Stuck.: in South America.
    • Atriplex deserticola Phil.: in South America.
    • Atriplex dimorphostegia Kar. & Kir.: in North Africa.
    • Atriplex eardleyae Aellen: in Australia
    • Atriplex elachophylla F. Muell.: in Australia.
    • Atriplex fissivalvis F. Muell.: in Australia
    • Atriplex flabellum Bunge ex Boiss.: in Eurasia.
    • Atriplex gardneri (Moq.) D. Dietr. – Gardner's saltbush, moundscale: in North America
    • Atriplex glauca L.: in Portugal, Spain and in North Africa.
    • Atriplex halimus L. – Mediterranean saltbush, sea orache, shrubby orache: in south Europe, North Africa and southwest Asia.
    • Atriplex herzogii Standl.: in North America.
    • Atriplex holocarpa F. Muell.: in Australia.
    • Atriplex hymenelytra (Torr.) S. Watson – Desert holly: in North America.
    • Atriplex hymenotheca Moq.: in Australia.
    • Atriplex imbricata (Moq.) D. Dietr.: in South America.
    • Atriplex inamoena Aellen: in Eurasia.
    • Atriplex intermedia Anderson: in Australia.
    • Atriplex isatidea Moq.: in Australia.
    • Atriplex laciniata L. – Frosted orache: In western and northern Europe.
    • Atriplex lampa (Moq.) Gillies ex Small: in South America.
    • Atriplex lehmanniana Bunge: in Eurasia.
    • Atriplex lentiformis (Torr.) S. Watson – Quail bush: in North America.
    • Atriplex leptocarpa F. Muell.: in Australia.
    • Atriplex leucoclada Boiss.: in Eurasia.
    • Atriplex leucophylla (Moq.) D. Dietr.: in North America
    • Atriplex lindleyi Moq.: in Australia.
    • Atriplex moneta Bunge ex Boiss.: in Eurasia.
    • Atriplex muelleri Benth.: in Australia.
    • Atriplex nessorhina S.W.L. Jacobs: in Australia.
    • Atriplex nummularia Lindl. – Old man saltbush, giant saltbush: in Australia.
    • Atriplex obovata Moq.: in North America.
    • Atriplex pamirica Iljin: in Eurasia.
    • Atriplex parishii S. Watson: in North America
    • Atriplex parryi S. Watson: in North America
    • Atriplex parvifolia Kunth: in South America.
    • Atriplex patagonica (Moq.) D. Dietr.: in South America.
    • Atriplex phyllostegia (Torr. ex S. Watson) S. Watson: in North America.
    • Atriplex polycarpa (Torr.) S. Watson – Allscale (saltbush), desert saltbush, cattle saltbush, cattle spinach: in North America.
    • Atriplex powellii S. Watson – Powell's saltbush: in North America.
    • Atriplex pseudocampanulata Aellen: in Australia.
    • Atriplex quinii F. Muell.: in Australia.
    • Atriplex recurva d'Urv.: in Eurasia, endemic to areas around the Aegean.
    • Atriplex rhagodioides F. Muell.: in Australia.
    • Atriplex rosea L. – Tumbling orache: in Eurasia and North Africa.
    • Atriplex rusbyi Britton ex Rusby: in South America.
    • Atriplex schugnanica Iljin: in Asia.
    • Atriplex semibaccata R. Br. – Australian saltbush, berry saltbush, creeping saltbush: in Australia.
    • Atriplex semilunaris Aellen: in Australia.
    • Atriplex serenana A. Nelson ex Abrams: in North America
    • Atriplex sibirica L.; in Asia, naturalized in Europe.
    • Atriplex sphaeromorpha Iljin: in Russia, Ukraine and Caucasus.
    • Atriplex spinibractea Anderson: in Australia.
    • Atriplex spongiosa F. Muell.: in Australia.
    • Atriplex stipitata Benth.: in Australia.
    • Atriplex sturtii S.W.L. Jacobs: in Australia.
    • Atriplex suberecta I. Verd. – Sprawling saltbush, lagoon saltbush: in Australia.
    • Atriplex tatarica Aellen: in Europe, North Africa and Asia.
    • Atriplex turbinata (Anderson) Aellen: in Australia.
    • Atriplex undulata (Moq.) D. Dietr.: in South America.
    • Atriplex velutinella F. Muell.: in Australia.
    • Atriplex vesicaria Heward ex Benth. – Bladder saltbush: in Australia.

Distribution and habitat

[edit]

The genus Atriplex is distributed nearly worldwide from subtropical to temperate and to subarctic regions. Most species-rich are Australia, North America, South America and Eurasia. Many species are halophytes and are adapted to dry environments with salty soils.[11]

Ecology

[edit]

Atriplex species are used as food plants by the larvae of some Lepidoptera species; see the list of Lepidoptera which feed on Atriplex.[20] They are also sometimes consumed by camels.[21] For spiders such as Phidippus californicus and other arthropods, saltbush plants offer opportunities to hide and hunt in habitat that is otherwise often quite barren.

It has been proposed that genus Atriplex was a main food source in the diet of the extinct giant kangaroo Procoptodon goliah.[22] Stable isotopic data suggested that their diet consisted of plants that used the C4 photosynthetic pathway, and due to their semi-arid distribution, chenopod saltbushes were likely responsible.[23][24][25]

Uses

[edit]

The favored species for human consumption is now usually garden orache (A. hortensis),[26][27][28] but many species are edible and the use of Atriplex as food is known since at least the late Epipaleolithic (Mesolithic).

Common orache (A. patula) is attested as an archaeophyte in northern Europe, and the Ertebølle culture is presumed to have used it as a food.[29] Its seed has been found among apparent evidence of cereal preparation and cooking at Late Iron Age villages in Britain.[30] In the biblical Book of Job, mallûaḥ (מַלּ֣וּחַ, probably Mediterranean saltbush, A. halimus, the major culinary saltbush in the region) is mentioned as food eaten by social outcasts (Job 30:4[31]). Grey saltbush (A. cinerea) has been used as bushfood in Australia since prehistoric times. Chamiso (A. canescens) and shadscale (A. confertifolia) were eaten by Native Americans, and spearscale (A. hastata) was a food in rural Eurasia.

Studies on Atriplex species demonstrated their potential use in agriculture. Meat from sheep which have grazed on saltbush has surprisingly high levels of vitamin E, is leaner and more hydrated than regular lamb and has consumer appeal equal to grain-fed lamb.[32] The vitamin E levels could have animal health benefits while extending the shelf-life and maintaining the fresh red colour of saltbush lamb. This effect has been demonstrated for old man saltbush (A. nummularia) and river saltbush (A. amnicola). For reasons unknown, sheep seem to prefer the more fibrous, less nutritious river saltbush.[33][32]

A study on A. nummularia discovered the species have a nitrogen content of 2.5–3.5%, and could potentially be used as a protein supplement for grazing if palatable.[34] A subsequent study allowed sheep and goats to voluntarily feed on Atriplex halimus and aimed to determine if the saltbush was palatable, and if so, did it provide enough nutrients to supplement the diet of these animals.[35] In this study they determined when goats and sheep are given as much A. halimus as they like, they do obtain enough nutrients to supplement their diet – unless the animal requirements are higher during pregnancy and milk production.

Saltbushes are also used as an ornamental plant in landscaping and can be used to prevent soil erosion in coastal areas.[36][37] Old man saltbush (Atriplex nummularia) has also been successfully used to rehabilitate old mining sites around Lightning Ridge (Australia).

See also

[edit]

References

[edit]
  1. ^ a b c "Atriplex L." Plants of the World Online. Retrieved 19 June 2022.
  2. ^ Sunset Western Garden Book, 1995:606–607
  3. ^ "orach". The Chambers Dictionary (9th ed.). Chambers. 2003. ISBN 0-550-10105-5.
  4. ^ "orache". Oxford English Dictionary third edition. Oxford University Press. June 2004. Retrieved 7 December 2018.
  5. ^ Quattrocchi, Umberto (2000). CRC World Dictionary of Plant Names. Vol. I: A-C. CRC Press. p. 232. ISBN 978-0-8493-2675-2.
  6. ^ a b c Stanley L. Welsh: Atriplex - online, In: Flora of North America Editorial Committee (ed.): Flora of North America North of Mexico, Volume 4: Magnoliophyta: Caryophyllidae, part 1., Oxford University Press, New York. 2003, ISBN 0-19-517389-9.
  7. ^ a b c d Gelin Zhu, Sergei L. Mosyakin & Steven E. Clemants: Chenopodiaceae: Atriplex - online, In: Wu Zhengyi, Peter H. Raven, Deyuan Hong (ed.): Flora of China, Volume 5: Ulmaceae through Basellaceae., Science Press und Missouri Botanical Garden Press, Beijing und St. Louis, 2003, ISBN 1-930723-27-X.
  8. ^ a b c Walsh, Neville G. "Atriples". Royal Botanic Gardens Victoria. Retrieved 16 July 2022.
  9. ^ a b c Jacobs, Surrey W.L. "Atriplex". Royal Botanic Garden Sydney. Retrieved 16 July 2022.
  10. ^ a b c "Atriplex". FloraBase. Western Australian Government Department of Biodiversity, Conservation and Attractions.
  11. ^ a b c d e f g Gudrun Kadereit, Evgeny V. Mavrodiev, Elizabeth H. Zacharias & Alexander P. Sukhorukov: Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 Photosynthesis. - American Journal of Botany 97(10): 1664-1687, 2010.
  12. ^ Linnaeus, Carl (1753). Species Plantarum. Vol. 2. pp. 1052–1053. Retrieved 19 June 2022.
  13. ^ Carl Linnaeus (1753), "Atriplex", Species Plantarum, vol. 2, Lars Salvius/Biodiversity Heritage Library, pp. 1052–1054, retrieved 19 May 2015
  14. ^ Sharr, Francis Aubi; George, Alex (2019). Western Australian Plant Names and Their Meanings (3rd ed.). Kardinya, WA: Four Gables Press. p. 61. ISBN 9780958034180.
  15. ^ Atriplex at Tropicos, accessed 2013-07-11
  16. ^ Shepherd, Kelly; Thiele, Kevin; Sampson, Jane; Coates, David; Bryne, Margaret (2015). "A rare, new species of Atriplex (Chenopodiaceae) comprising two genetically distinct but morphologically cryptic populations in arid Western Australia: implications for taxonomy and conservation". Australian Systematic Botany. 28 (4): 235. doi:10.1071/SB15029. S2CID 85903596. Retrieved 8 September 2021.
  17. ^ a b Flores, Hilda; Davis, Jerrold I. (2001). "A Cladistic Analysis of Atripliceae (Chenopodiaceae) Based on Morphological Data". Journal of the Torrey Botanical Society. 128 (3): 297. doi:10.2307/3088719. JSTOR 3088719.
  18. ^ a b c Brignone, Nicolás F.; Pozner, Raúl E.; Denham, Silvia S. (October 2019). "Origin and evolution of Atriplex (Amaranthaceae s.l.) in the Americas: Unexpected insights from South American species". Taxon. 68 (5): 1027. doi:10.1002/tax.12133. S2CID 214371753.
  19. ^ Tasmanian name, also transcribed trucanini, trucaninny, trugannini, trugernanna, etc. The plant was the namesake for Truganini, among the last of her people.
  20. ^ Robinson, Gaden S.; Ackery, Phillip R.; Kitching, Ian; Beccaloni, George W.; Hernández, Luis M. (2023). "HOSTS - The Hostplants and Caterpillars Database at the Natural History Museum". www.nhm.ac.uk. doi:10.5519/havt50xw. Retrieved 2020-09-28.
  21. ^ Shawket, S. M. (1999). "Fattening of camel calves on saltbush, Atriplex halimas with different energy sources". Mansoura University, Journal of Agricultural Sciences (Egypt). ISSN 1110-0346.
  22. ^ Prideaux, Gavin J.; Ayliffe, Linda K.; DeSantis, Larisa R. G.; Schubert, Blaine W.; Murray, Peter F.; Gagan, Michael K.; Cerling, Thure E. (2009-07-14). "Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo". Proceedings of the National Academy of Sciences. 106 (28): 11646–11650. Bibcode:2009PNAS..10611646P. doi:10.1073/pnas.0900956106. ISSN 0027-8424. PMC 2710660. PMID 19556539. S2CID 19904871.
  23. ^ Kadereit, Gudrun; Mavrodiev, Evgeny V.; Zacharias, Elizabeth H.; Sukhorukov, Alexander P. (2010). "Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): Implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis". American Journal of Botany. 97 (10): 1664–1687. doi:10.3732/ajb.1000169. ISSN 1537-2197. PMID 21616801.
  24. ^ Prideaux, G. (2004). "Systematics and Evolution of the Sthenurine Kangaroos". UC Publications in Geological Sciences. University of California Press. doi:10.1525/california/9780520098459.001.0001. ISBN 9780520098459.
  25. ^ DeSantis, Larisa R. G.; Field, Judith H.; Wroe, Stephen; Dodson, John R. (May 2017). "Dietary responses of Sahul (Pleistocene Australia–New Guinea) megafauna to climate and environmental change". Paleobiology. 43 (2): 181–195. Bibcode:2017Pbio...43..181D. doi:10.1017/pab.2016.50. ISSN 0094-8373. S2CID 13134989.
  26. ^ "Orach facts and health benefits". 4 February 2020. Retrieved 2020-09-28.
  27. ^ "Atriplex hortensis Orach, Garden orache PFAF Plant Database". pfaf.org. Retrieved 2020-09-28.
  28. ^ "Atriplex hortensis (garden orache)". www.cabi.org. Retrieved 2020-09-28.
  29. ^ Koch, Eva (December 1999). "Prehistoric plant food of Denmark". Archived from the original on 2006-07-17.
  30. ^ Christopher Evans (2015), North West Cambridge Archaeology: University of Cambridge 2013 Excavations, The Traveller's Rest Sub-site (PDF), Cambridge Archaeological Unit University Of Cambridge, pp. 100–113
  31. ^ Mistranslated as "mallows" in the King James Bible and as Nesseln (nettles) in the Luther Bible
  32. ^ a b Pearce, Kelly & Jacob, Robin (2004): Saltbush lifts sheep meat vitamin content. Farming Ahead 153(October): 63. PDF fulltext Archived July 23, 2005, at the Wayback Machine
  33. ^ Norman, Hayley C.; Freind, Colby; Masters, David G.; Rintoul, Allan J.; Dynes, Robyn A. & Williams, Ian H. (2004): Variation within and between two saltbush species in plant composition and subsequent selection by sheep. Aust. J. Agr. Res. 55(9): 999–1007. doi:10.1071/AR04031 (HTML abstract)
  34. ^ Wilson, A.D.; Harrington, G.N. (1980). "Nutritive value of Australian browse plants". International Livestock Centre for Africa. Addis Ababa: 291.
  35. ^ Valderrabano, J.; Munoz, F.; Delgado, I. (1996). "Browsing ability and utilisation by sheep and goats of Atriplex hamilus L. shrubs". Small Ruminant Research. 19 (2). Zaragoza, Spain: 131–136. doi:10.1016/0921-4488(95)00754-7. Retrieved 13 April 2020.
  36. ^ "Saltbushes for dryland salinity management in Western Australia". www.agric.wa.gov.au. Retrieved 2020-09-28.
  37. ^ "Atriplex semibaccata (Australian saltbush)". www.cabi.org. Retrieved 2020-09-28.
  • Davidson, Alan (1999): Orach. In: Oxford Companion to Food: 556. ISBN 0-19-211579-0
  • Everitt, J.H.; Lonard, R.L.; Little, C.R. (2007). Weeds in South Texas and Northern Mexico. Lubbock: Texas Tech University Press. ISBN 0-89672-614-2
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