{{short description|Subfamily of plants within the borage family (Boraginaceae)}} {{Automatic taxobox |image = Ogórecznik.JPG |image_caption = ''[[Borago officinalis]]'' |taxon = Boraginoideae |authority = Arn. |type_genus = ''[[Borago]]'' |type_genus_authority = L. |subdivision_ranks = Tribes |subdivision = * [[Boragineae]] * [[Lithospermeae]] }}

'''Boraginoideae''' is a [[subfamily]] of the plant family [[Boraginaceae]] ''{{abbr|s.s|in the strict sense}}'', with about 42 genera. That family is [[Circumscription (taxonomy)|defined]] in a much broader sense (Boraginaceae ''{{abbr|s.l.|in the broad sense}}'') in the [[Angiosperm Phylogeny Group]] (APG) system of classification for flowering plants. The APG has not specified any subfamilial structure within Boraginaceae ''s.l.''

== Taxonomy ==

Some [[taxonomist]]s placed the genera ''[[Codon (plant)|Codon]]'' and ''[[Wellstedia]]'' in Boraginoideae. Others place one or both of these in separate, [[monogeneric]] subfamilies. ''Codon'' was long regarded as an odd member of [[Hydrophylloideae]], but in 1998, a [[molecular phylogenetic]] study suggested that it is closer to Boraginoideae.{{sfn|Ferguson|1998}} Neither is included n more modern classifications.{{sfn|Chacon et al|2016}}

Some authors proposed a revision of earlier APG systems, in which Boraginaceae had been included as an unplaced family (i.e. not included in a specified [[Order (biology)|order]]) within the [[lamiid]] clade of [[eudicots]]. In that system. Boraginaceae was defined broadly (Boraginaceae ''sensu lato'' or ''s.l.''). Instead they proposed recognizing five to eight families in a separate order, the [[Boraginales]].{{sfn|Weigand et al|2014}}{{sfn|Chacon et al|2016}} In that system Boraginaceae is treated in a narrow sense (''[[sensu stricto]]'' or ''s.s.''). Subsequently the order Boraginales was added to the 2016 revision ([[APG IV]]) to include Boraginaceae at the ordinal level, in which it was the sole family. The consensus was to continue the broad usage rather than split it into separate families, based on its [[monophyletic]] composition.<ref name="apweb">Peter F. Stevens. (2001 onwards). "Boraginaceae". At: [[Angiosperm Phylogeny Website]]. At: [[Missouri Botanical Garden]] Website. (see ''External links'' below)</ref> The subfamilial structure applies solely to Boraginaceae ''s.s.'', which includes around 1,600 to 1,700 species divided into some 90 genera.{{cn|date=January 2023}}

=== Subdivision ===

Comparisons of [[DNA sequence]]s by [[cladistic]] methods suggested the division of Boraginoideae into four [[Tribe (biology)|tribes]]: Echiochileae, Boragineae, Lithospermeae, and Cynoglosseae.{{sfn|Cohen|2014}} This was subsequently resolved into only two tribes, Boragineae and Lithospermeae (25 genera). Boragineae was then further subdivided into two subtribes, Boragininae (15 genera) and Moritziinae (2).{{sfn|Chacon et al|2016}}

=== Genera ===

Genera and approximate number of species in the system of Chacon et al (2016):{{sfn|Chacon et al|2016}} * Tribe Boragineae * Subtribe Boragininae (140 species) ''Anchusa'', ''Anchusella'', ''Borago'', ''Brunnera'', ''Cynoglottis'', ''Gastrocotyle'', ''Hormuzakia'', ''Lycopsis'', ''Melanortocarya'', ''Nonea'', ''Pentaglottis'', ''Phyllocara'', ''Pulmonaria'', ''Symphytum'', ''Trachystemon'' * Subtribe Moritziinae (6 species) ''Moritzia'', ''Thaumatocaryon'' * Tribe Lithospermeae (460 species) ''Aegonychon'', ''Alkanna'', ''Ancistrocarya'', ''Arnebia'', ''Buglossoides'', ''Cerinthe'', ''Cystostemon'', ''Echiostachys'', ''Echium'', ''Glandora'', ''Halacsya'', ''Huynhia'', ''Lithodora'', ''Lithospermum'', ''Lobostemon'', ''Maharanga'', ''Mairetis'', ''Moltkia'', ''Moltkiopsis'', ''Neatostema'', ''Onosma'', ''Paramoltkia'', ''Podonosma'', ''Pontechium'', ''Stenosolenium''.<ref>Armen Takhtajan {{google books|oumyfO-NHuUC|Flowering Plants (2009) |page=542}}</ref>

The following list of genera consists of ''Codon'' plus the genera listed for Boraginoideae at the [[Germplasm Resources Information Network]]<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2089 |title=Genera of ''Boraginaceae'' subfam. ''Boraginoideae'' |work=[[Germplasm Resources Information Network]] |publisher=[[United States Department of Agriculture]] |access-date=2009-04-02 |url-status=dead |archive-url=https://web.archive.org/web/20150924052400/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2089 |archive-date=2015-09-24 }}</ref>

{{columns-list|colwidth=17em| * ''[[Actinocarya]]'' <small>Benth.</small> * ''[[Adelocaryum]]'' <small>Brand</small> * ''[[Afrotysonia]]'' <small>Rauschert</small> * ''[[Alkanna]]'' <small>Tausch</small> * ''[[Amblynotus]]'' <small>(A.DC.) I. M. Johnst.</small> * ''[[Amphibologyne]]'' <small>Brand</small> * ''[[Amsinckia]]'' <small>Lehm.</small> * ''[[Anchusa]]'' <small>L.</small> * ''[[Ancistrocarya]]'' <small>Maxim.</small> * ''[[Anoplocaryum]]'' <small>Ledeb.</small> * ''[[Antiotrema]]'' <small>Hand.-Mazz.</small> * ''[[Antiphytum]]'' <small>DC. ex [[Carl Meissner|Meisn.]]</small> * ''[[Arnebia]]'' <small>Forssk.</small> * ''[[Asperugo]]'' <small>L.</small> * ''[[Auxemma]]'' <small>Miers</small> * ''[[Borago]]'' <small>L.</small> * ''[[Bothriospermum]]'' <small>Bunge</small> * ''[[Brachybotrys]]'' <small>Maxim. ex Oliv.</small> * ''[[Brunnera]]'' <small>Steven</small> * ''[[Buglossoides]]'' <small>Moench</small> * ''[[Caccinia]]'' <small>Savi</small> * ''[[Carmona (plant)|Carmona]]'' <small>Cav.</small> * ''[[Cerinthe]]'' <small>L.</small> * ''[[Chionocharis]]'' <small>I. M. Johnst.</small> * ''[[Choriantha]]'' <small>Riedl</small> * ''[[Codon (plant)|Codon]]'' <small>[[Carl Linnaeus|L.]]</small> * ''[[Craniospermum]]'' <small>Lehm.</small> * ''[[Cryptantha]]'' <small>Lehm. ex G.Don</small> * ''[[Cynoglossopsis]]'' <small>Brand</small> * ''[[Cynoglossum]]'' <small>L.</small> * ''[[Cynoglottis]]'' <small>(Gusul.) Vural & Kit Tan</small> * ''[[Cystostemon]]'' <small>Balf.f.</small> * ''[[Dasynotus]]'' <small>I.M.Johnst.</small> * ''[[Decalepidanthus]]'' <small>Riedl</small> * ''[[Echiochilon]]'' <small>Desf.</small> * ''[[Echiostachys]]'' <small>Levyns</small> * ''[[Echium]]'' <small>L.</small> * ''[[Elizaldia]]'' <small>Willk.</small> * ''[[Embadium]]'' <small>J.M.Black</small> * ''[[Eritrichium]]'' <small>Schrad. ex Gaudin</small> * ''[[Gastrocotyle (plant)|Gastrocotyle]]'' <small>Bunge</small> * ''[[Gyrocaryum]]'' <small>Valdés</small> * ''[[Hackelia]]'' <small>Opiz.</small> * ''[[Halacsya]]'' <small>Dörfl.</small> * ''[[Heliocarya]]'' <small>Bunge</small> * ''[[Heterocaryum]]'' <small>A.DC.</small> * ''[[Huynhia]]'' <small>Greuter</small> * ''[[Ivanjohnstonia]]'' <small>Kazmi</small> * ''[[Lacaitaea]]'' <small>Brand</small> * ''[[Lappula]]'' <small>Moench</small> * ''[[Lasiarrhenum]]'' <small>I.M.Johnst.</small> * ''[[Lasiocaryum]]'' <small>I.M.Johnst.</small> * ''[[Lepechiniella]]''<small>Popov</small> * ''[[Lindelofia]]'' <small>Lehm.</small> * ''[[Lithodora]]'' <small>Griseb.</small> * ''[[Lithospermum]]'' <small>L.</small> * ''[[Lobostemon]]'' <small>Lehm.</small> * ''[[Macromeria]]'' <small>D.Don</small> * ''[[Maharanga]]'' <small>A.DC.</small> * ''[[Mairetis]]'' <small>I.M.Johnst.</small> * ''[[Mattiastrum]]'' <small>(Boiss.) Brand</small> * ''[[Mertensia]]'' <small>Roth</small> * ''[[Metaeritrichium]]'' <small>W.T.Wang</small> * ''[[Microcaryum]]'' <small>I.M.Johnst.</small> * ''[[Microula]]'' <small>Benth.</small> * ''[[Mimophytum]]'' <small>Greenm.</small> * ''[[Moltkia (plant)|Moltkia]]'' <small>Lehm.</small> * ''[[Moltkiopsis]]'' <small>I.M.Johnst.</small> * ''[[Moritzia]]'' <small>DC. ex Meisn.</small> * ''[[Myosotidium]]'' <small>Hook.</small> * ''[[Myosotis]]'' <small>L.</small> * ''[[Neatostema]]'' <small>I.M.Johnst.</small> * ''[[Nesocaryum]]'' <small>I.M.Johnst.</small> * ''[[Nogalia]]'' <small>Verdc.</small> * ''[[Nomosa]]'' <small>I.M.Johnst.</small> * ''[[Nonea]]'' <small>Medik.</small> * ''[[Ogastemma]]'' <small>Brummitt</small> * ''[[Omphalodes]]'' <small>Mill.</small> * ''[[Omphalolappula]]'' <small>Brand</small> * ''[[Omphalotrigonotis]]'' <small>W.T.Wang</small> * ''[[Onosma]]'' <small>L.</small> * ''[[Oxyosmyles]]'' <small>Speg.</small> * ''[[Paracaryum]]'' <small>(A.DC.) Boiss.</small> * ''[[Pardoglossum]]'' <small>E.Barbier & Mathez</small> * ''[[Pectocarya]]'' <small>DC. ex Meisn.</small> * ''[[Pentaglottis]]'' <small>Tausch</small> * ''[[Perittostema]]'' <small>I.M. Johnst.</small> * ''[[Plagiobothrys]]'' <small>Fisch. & C.A.Mey.</small> * ''[[Pseudomertensia]]'' <small>Riedl</small> * ''[[Psilolaemus]]'' <small>I.M.Johnst.</small> * ''[[Pulmonaria]]'' <small>L.</small> * ''[[Rindera]]'' <small>Pall.</small> * ''[[Rochelia]]'' <small>Rchb.</small> * ''[[Scapicephalus]]'' <small>Ovcz. & Czukav.</small> * ''[[Selkirkia (plant)|Selkirkia]]'' <small>Hemsl.</small> * ''[[Sericostoma (plant)|Sericostoma]]'' <small>Stocks</small> * ''[[Sinojohnstonia]]'' <small>Hu</small> * ''[[Solenanthus]]'' <small>Ledeb.</small> * ''[[Stenosolenium]]'' <small>Turcz.</small> * ''[[Stephanocaryum]]'' <small>Popov</small> * ''[[Suchtelenia]]'' <small>Kar. ex Meisn.</small> * ''[[Symphytum]]'' <small>L.</small> * ''[[Thaumatocaryon]]'' <small>Baill.</small> * ''[[Thyrocarpus]]'' <small>Hance</small> * ''[[Tianschaniella]]'' <small>B.Fedtsch. ex Popov</small> * ''[[Trachelanthus (plant)|Trachelanthus]]'' <small>Kunze</small> * ''[[Trachystemon]]'' <small>D.Don</small> * ''[[Traxara]]'' <small>Raf.</small> * ''[[Trichodesma]]'' <small>R.Br.</small> * ''[[Trigonocaryum]]'' <small>Trautv.</small> * ''[[Trigonotis]]'' <small>Steven</small> * ''[[Ulugbekia]]'' <small>Zakirov</small> * ''[[Valentiniella]]'' <small>Speg.</small> * ''[[Wellstedia]]'' <small>Balf.f.</small> }}

==References== {{Reflist|30em}}

== Bibliography == {{refbegin|30em}} * {{cite journal |last1=Chacón |first1=Juliana |last2=Luebert |first2=Federico |last3=Hilger |first3=Hartmut H. |last4=Ovchinnikova |first4=Svetlana |last5=Selvi |first5=Federico |last6=Cecchi |first6=Lorenzo |last7=Guilliams |first7=C. Matt |last8=Hasenstab-Lehman |first8=Kristen |last9=Sutorý |first9=Karel |last10=Simpson |first10=Michael G. |last11=Weigend |first11=Maximilian |title=The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera |url=https://www.researchgate.net/publication/304454123 |journal=[[Taxon (journal)|Taxon]] |date=24 June 2016 |volume=65 |issue=3 |pages=523–546 |doi=10.12705/653.6 |ref={{harvid|Chacon et al|2016}} |hdl=2158/1062785 |hdl-access=free }} * {{cite journal |last1=Cohen |first1=James I. |title=A phylogenetic analysis of morphological and molecular characters of Boraginaceae: evolutionary relationships, taxonomy, and patterns of character evolution |journal=[[Cladistics (journal)|Cladistics]] |date=April 2014 |volume=30 |issue=2 |pages=139–169 |doi=10.1111/cla.12036|pmid=34784688 |s2cid=25399963 }} * {{cite journal |last1=Ferguson |first1=Diane M. |title=Phylogenetic Analysis and Relationships in Hydrophyllaceae Based on ndhF Sequence Data |journal=[[Systematic Botany]] |date=July 1998 |volume=23 |issue=3 |pages=253–268 |doi=10.2307/2419504|jstor=2419504 }} * {{cite journal |last1=Weigend |first1=Maximilian |last2=Luebert |first2=Federico |last3=Gottschling |first3=Marc |last4=Couvreur |first4=Thomas L.P. |last5=Hilger |first5=Hartmut H. |last6=Miller |first6=James S. |title=From capsules to nutlets-phylogenetic relationships in the Boraginales |journal=[[Cladistics (journal)|Cladistics]] |date=October 2014 |volume=30 |issue=5 |pages=508–518 |doi=10.1111/cla.12061|s2cid=11954615 |ref={{harvid|Weigand et al|2014}}|doi-access=free |pmid=34794245 }} * [http://www.mobot.org/MOBOT/Research/APweb/maps/boraginaceae.gif Distribution Map] {{color|green|And}} [http://www.mobot.org/MOBOT/Research/APweb/genera/boraginaceaegen.html Genus list] {{color|green|At}} [http://www.mobot.org/MOBOT/Research/APweb/orders/boraginalesweb.htm#Boraginaceae Boraginaceae] {{color|green|At}} [http://www.mobot.org/MOBOT/Research/APweb/orders/boraginalesweb.htm#Boraginales Boraginales] {{color|green|At:}} [http://www.mobot.org/MOBOT/Research/APweb/treeapweb2map.html Trees] {{Webarchive|url=https://web.archive.org/web/20150521153254/http://www.mobot.org/mobot/research/apweb/treeapweb2map.html |date=2015-05-21 }} {{color|green|At:}} [http://www.mobot.org/MOBOT/Research/APweb/welcome.html APweb] {{color|green|At:}} [http://www.missouribotanicalgarden.org/ Missouri Botanical Garden] {{refend}}

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[[Category:Boraginoideae| ]] [[Category:Asterid subfamilies]]