Hippeastreae

{{Short description|Tribe of flowering plants}} {{use dmy dates|cs1-dates=ly|date=June 2021}} {{Automatic taxobox |taxon = Hippeastreae |image = Starr 080327-3845 Hippeastrum striatum.jpg |image_caption = ''Hippeastrum striatum'' |authority = Herb. ex Sweet{{sfn|Garcia et al|2019}} |synonyms_ref = |synonyms = |subdivision_ranks = Subtribes |subdivision = *''Hippeastrinae'' Walp. *''Traubiinae'' D. & U. Müll.-Doblies |type_genus = ''Hippeastrum'' |type_genus_authority = (L.) Herb. }}

'''Hippeastreae''' is a tribe of plants belonging to the subfamily Amaryllidoideae of the Amaryllis family (Amaryllidaceae). Species in this tribe are distributed in South America. Flowers are large and showy, zygomorphic, with the stamens in varying lengths, inflorescence bracts are often fused basally (along one side). The seeds are flattened, winged or D-shaped. Reported basic chromosome numbers are ''x''= 8-13, 17, and higher. All the species in this tribe present a remarkable aesthetic interest and horticultural value.{{sfn|Meerow et al|2000}}{{sfn|Vigneron|2008}}

== Taxonomy == Meerow et al. (1999) provide a history of the treatment of the genera of Amaryllidaceae, including Hippeastreae, from the mid-twentieth century.{{sfn|Meerow et al|1999}} While morphological phylogeny has been frustrated by the perversive homoplasy typical of the Amaryllidaceae,{{sfn|Garcia et al|2014}} application of molecular phylogenetics to the Amaryllidaceae did not indicate clear tribal divisions but rather broad biogeographical clades. However the American clade resolved the tribe Hippeastreae.{{sfn|Meerow et al|1999}} A later examination of the deeper relationships of the American genera suggested the two subclades, Andean and hippeastroid and within the latter separated the Brazilian Griffineae as sister to the remaining hippeastroids. The larger and more diverse grouping of hippeastroids formed two smaller monophyletic groups. The smaller contained ''Hippeastrum'' (with the exception of ''Hippeastrum blumenavium''),{{efn|''Hippeastrum blumenavium'', or ''Hippeastrum blumenavia'', was earlier known as ''Griffinia blumenavia'' and is an unusual species more closely resembling ''Rhodophiala''. Ultimately Meerow ''et al.'' recommended reassigning it to a monotypic genus, where it is now known as ''Eithea''.}} but also a ''Rhodophiala''. With the exception of ''Rodophilia'' (Brazil) all specimens were from Chile and Argentina. The second group corresponded to those genera variously included in tribe Zephyrantheae (Traub) or subtribe Zephyranthinae (Müller-Doblies), but only including some ''Zephyranthes'' species. The hippeastroid clade is predominantly diploid and extra-Andean by comparison to the Andean clade which is predominantly tetraploid, and contain those genera traditionally included in Hippeastreae. The precise position of Griffineae remained unresolved since its sister status to Hippeastrae was weak, leaving the possibility that it could be sister to the whole American clade.{{sfn|Meerow et al|2000}} The tribe consists of 10–13 genera and about 180 species.{{sfn|Garcia et al|2014}}

=== Phylogeny === The placement of Hippeastreae within subfamily Amaryllidoideae is shown in the following cladogram, where this tribe is shown as a sister group to the Griffineae, forming the Hippeastroid subclade, of two American clades:{{sfn|Meerow et al|2000}}

{{cladogram | title= {{anchor|Clad3}}Cladogram: Tribes of subfamily Amaryllidoideae | align=left | cladogram={{clade|style=font-size:92%;line-height:100%;width:700px;

| label1 = Subfamily '''Amaryllidoideae'''

|1= {{Clade | label1= Africa | 1= Tribe Amaryllideae | 2= {{Clade | 1= {{clade | label1=Africa | 1= Tribe Cyrtantheae | 2= {{clade | label1=Africa | 1= Tribe Haemantheae | label2= Australasia | 2= Tribe Calostemmateae }} }} | 2= {{clade | label1= Eurasian clade | label2= American clade | 1= {{clade | label1= Asia | 1= Tribe Lycorideae | label2= Mediterranean | 2= {{clade | 1= Tribe Galantheae | 2= {{clade | 1= Tribe Pancratieae | 2= Tribe Narcisseae }} }} }} | 2= {{clade | label1= Hippeastroid clade | label2= Andean clade | 1= {{clade | label1=Brazil | 1= Tribe Griffineae | 2= Tribe '''Hippeastreae'''

}} | 2= {{clade | 1= {{clade | 1= Tribe Eustephieae }} | 2= {{clade | label1=Petiolate | 1= Tribe Eucharideae/Stenomesseae

| 2= {{clade | 1= {{clade | 1= Tribe Clinantheae | 2= Tribe Hymenocallideae }} }} }} }} }} }} }} }} }} }} {{Clear}}

=== Subdivision ===

The genera of the tribe Hippeastreae have been treated in a number of different ways over the years. Traub (1963){{sfn|Traub|1963}} in his monograph on the Amaryllidacea distributed those genera now considered in this tribe over a number of other tribes (see Table, below), while Dahlgren ''et al.'' included them all under Hippeastrae for the first time.{{sfn|Dahlgren|Clifford|Yeo|1985}} The concept of subtribes came from the Müller-Doblies' (1996), who had three subtribes, Griffinineae, Hippeastrinae and Zephyranthinae.{{sfn|Müller-Doblies|Müller-Doblies|1996}}

Consequently, it has been customary to describe the tribe Hippeastreae as consisting of two subtribes: * subtribe Hippeastrinae - includes species of medium height and often with many flowers in each inflorescence and inflorescence bracts are different in size and fused basally. Genera that have been placed in this subtribe include ''Placea'', ''Hippeastrum'', ''Phycella'' (includes ''Famatina''), ''Eithea'', ''Rhodophiala'', and ''Traubia''. (Some authors use the spelling Hippeastrineae){{sfn|Vigneron|2008}} * subtribe Zephyranthinae - includes species of small height with solitary flowers. Inflorescence bracts are fused forming a tube surrounding the pedicel of the flower. Genera in this subtribe are ''Sprekelia'', ''Habranthus'', ''Cooperia'', and ''Zephyranthes''.{{sfn|Vigneron|2008}}{{sfn|GRIN|2016|loc=[https://npgsweb.ars-grin.gov/gringlobal/taxonomygenus.aspx?id=14092 Cooperia]}} (Some authors use the spelling Zephyranthinea){{sfn|Vigneron|2008}}

In the study of Meerow ''et al.'' (2000) based on molecular markers Zephyranthinae (Zephyranthae of Traub){{sfn|Traub|1963}} were clearly polyphyletic, largely due to the polyphyly of ''Zephyranthes'' itself. This subgroup has been broadly characterised as having a chromosome number, x=6, but with considerable variation. Other polyphyletic genera included ''Rodophiala'' and ''Habranthus''.{{sfn|Meerow et al|2000}} Hippeastreae also include ''Haylockia'', ''Rhodolirion'' and ''Tocantinia''.{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}}

A more focused study of Hippeastreae alone in 2014, resolved two major clades:{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}} * Clade A: ''Traubia'', ''Placea'', ''Phycella'', ''Rhodolirium'', and ''Famatina maulensis'' (now ''Phycella maulensis'') * Clade B: ''Rhodophiala'', ''Habranthus'', ''Haylockia'', ''Hippeastrum'', ''Sprekelia'', ''Zephyranthes'', and the remainder of ''Famatina''.

However it also showed that of the 13 genera, two are monotypic (''Haylockia'' and ''Traubia''). Of the remaining 11 genera, based on Internal transcribed spacer (ITS) sequences the only monophyletic non-monotypic genera were ''Hippeastrum'' (about 60 species) and ''Sprekelia'' (2 species). But on chloroplast DNA (cpDNA) analysis, not even these genera were monophyletic. This brings into question the existing generic classification within Hippeastreae. Consequently, the authors proposed the following nomenclature:{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}} * Subtribe Traubiinae (Clade A) D.Müll.-Doblies & U.Müll.-Doblies * Subtribe Hippeastrinae (Clade B)

A major generic recircumscription was published in 2019, although the basic subdivision into the two above subtribes remained unchanged.{{sfn|Garcia et al|2019}}

==== Subtribes ====

Subtribe Traubiinae D. & U. Müll.-Doblies (Clade A) includes about 20 endemic Chilean taxa, but only about 10% of the species within tribe Hippeastreae. Characterisation includes a haploid chromosome number, x=8, lack of polyploidy and a capitate stigma.{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}} This subtribe has 4 genera.{{sfn|Garcia et al|2019}}

Subtribe Hippeastrinae Walp. (Clade B), by contrast has a variable chromosome number, x=6–11, with frequent aneuploidy and polyploidy. Although there are no unique synapomorphies, most taxa exhibit a trifid or trilobed stigma, although in a few it is capitate.{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}} This subtribe has 2 genera.{{sfn|Garcia et al|2019}}

==== Historical circumscription ====

Tribe Hippeastreae has included various numbers of genera over the years, some of which were monotypic.{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}}

{| class="wikitable sortable" border="1" |+ <big> Historical distribution of Hippeastreae (''sensu'' Meerow 1999) genera by tribes and ''subtribes''</big> ! style="background: #ccf;" |Genus (alphabetical)||Traub 1963{{sfn|Traub|1963}}||Dahlgren 1985{{sfn|Dahlgren|Clifford|Yeo|1985}}||Müller-Doblies 1996{{sfn|Müller-Doblies|Müller-Doblies|1996}} ||Meerow 1998{{sfn|Meerow|Snijman|1998}}||Garcia 2014{{sfn|Garcia et al|2014}}{{sfn|Garcia et al|2017}}||Species |- | ''Eithea'' formerly ''Hippeastrum blumenavium''|| || || || ||Hippeastreae ''Hippeastrinae'' ||1–2 |- | ''Famatina'' =''Rhodophiala'', ''Phycella''|| || || || || ||4 |- | ''Griffinia''||Euchareae|| ||Hippeastreae ''Griffiniinae''||Hippeastreae||Griffineae||21 |- |''Habranthus''||Zephyrantheae||Hippeastreae||Hippeastreae ''Zephyranthinae''||Hippeastreae||Hippeastreae ''Hippeastrinae''||40 |- | ''Haylockia''||Zephyrantheae|| ||Hippeastreae ''Zephyranthinae''||=''Zephyranthes''||=''Zephyranthes''||1 |- | ''Hippeastrum''||=''Amaryllis''||Hippeastreae|| Hippeastreae ''Hippeastrinae''||Hippeastreae||Hippeastreae ''Hippeastrinae''||60 |- | ''Phycella''||Eustephieae|| || ||Hippeastreae||Hippeastreae ''Traubiinae''||6 |- | ''Placea'' ||Amarylleae||Hippeastreae ||Hippeastreae ''Hippeastrinae''||Hippeastreae||Hippeastreae ''Traubiinae''||6 |- | ''Sprekelia''||Zephyrantheae||Hippeastreae||Hippeastreae ''Zephyranthinae''||Hippeastreae||Hippeastreae ''Hippeastrinae''||2 |- | ''Rhodolirium''||=''Rhodophiala'' || || || ||Hippeastreae ''Traubiinae''||5 |- |''Rhodophiala''||Zephyrantheae||Hippeastreae||Hippeastreae ''Hippeastrinae''||Hippeastreae||Hippeastreae ''Hippeastrinae''||8 |- | ''Tocantinia''|| || || || ||Hippeastreae ''Hippeastrinae''||1–2 |- | ''Traubia''||Traubieae|| ||Hippeastreae Traubiinae||Hippeastreae||Hippeastreae ''Traubiinae''||1 |- | ''Worsleya''||Amarylleae||Hippeastreae|| ||Hippeastreae||Griffineae||1 |- | ''Zephyranthes''||Zephyrantheae||Hippeastreae||Hippeastreae ''Zephyranthinae''||Hippeastreae||Hippeastreae ''Hippeastrinae''||50 |}

====Genera==== {{As of|2022|March}}, the Angiosperm Phylogeny Website (APweb) accepted only six genera in the tribe Hippeastrae:<ref name=APweb>Stevens, P.F. (2001 onwards). [https://www.mobot.org/MOBOT/research/APweb/genera/amaryllidaceaegen.html "Amaryllidaceae genera"]. ''Angiosperm Phylogeny Website''. Retrieved 2022-03-30.</ref> shown here as amended by Garcia et al 2019.{{sfn|Garcia et al|2019}} * Subtribe Traubiinae: ** ''Eremolirion'' Nic.Garcia (now ''Paposoa'') ** ''Phycella'' Lindlley ** ''Pyrolirion'' Herbert (now part of a separate tribe, Eustephieae) ** ''Rhodolirium'' ''s.str.'' Phil. ** ''Traubia'' Moldenke

* Subtribe Hippeastrinae ** ''Hippeastrum'' Herbert ** ''Zephyranthes'' Herbert

Genera that have been synonymized with other genera in the tribe according to APweb include:<ref name=APweb/> *''Cooperia'' Herbert = ''Zephyranthes'' *''Famatina'' Ravenna = ''Zephyranthes'' *''Habranthus'' Herbert = ''Zephyranthes'' *''Haylockia'' Herbert = ''Zephyranthes'' *''Placea'' Miers = ''Phycella'' *''Rhodolirium'' Philippi = ''Phycella'' *''Rhodophiala'' C. Presl = ''Phycella'' *''Sprekelia'' Heister = ''Zephyranthes'' *''Tocantina'' Ravenna = ''Hippeastrum''

== Distribution and habitat ==

Hippeastreae have a major center of diversification in central Chile and western (Andean) Argentina, together with minor centres in eastern Brazil, the north east of Argentina and with more distant centers in Mexico, the Greater Antilles and southern United States (''Habranthus'', ''Zephyranthes'').{{sfn|Garcia et al|2014}}

== Uses == The economic significance of the tribe lies in its horticultural usage.{{sfn|Garcia et al|2014}}

== Notes == {{notelist}}

== References == {{Reflist|20em}}

== Bibliography == {{Refbegin|30em}} === Books ===

* {{cite book |last1=Dahlgren |first1=R.M. |last2=Clifford |first2=H.T. |last3=Yeo |first3=P.F. |author-link1=Rolf Dahlgren|title=The families of the monocotyledons |year=1985 |publisher=Springer-Verlag |location=Berlin |url=https://books.google.com/books?id=3iGndTFY0skC|isbn= 978-3-642-64903-5| access-date=10 February 2014}} * {{cite thesis|last1=García Berguecio|first1=Nicolás|title=Systematics and evolution of Amaryllidaceae tribe Hippeastreae (Asparagales)|date=2015|publisher=University of Florida |type=PhD Thesis |url=http://www.iapt-taxon.org/downloads/grants/Garcia_Report_2013.pdf|access-date=2016-02-14|archive-url=https://web.archive.org/web/20150907225614/http://www.iapt-taxon.org/downloads/grants/Garcia_Report_2013.pdf|archive-date=2015-09-07}} * {{cite book|last=Hutchinson|first=John|author-link=John Hutchinson (botanist)|title=The families of flowering plants, arranged according to a new system based on their probable phylogeny |edition=1st |publisher=Macmillan}} ** Volume 1: Monocotyledonae (1926) ** Volume 2: Dicotyledonae (1934) * {{cite book|editor-last=Kubitzki|editor-first=K.|editor-link=Klaus Kubitzki|title=Flowering Plants · Monocotyledons: Lilianae (except Orchidaceae) |series=The families and genera of vascular plants. Vol. 3 |year=1998 |publisher=Springer-Verlag|location=Berlin|isbn=978-3-540-64060-8 |url=https://books.google.com/books?id=FyPVYzL76sMC|access-date=14 January 2014}} * {{harvc|last1=Meerow|first1=A.W.|last2=Snijman|first2=D.A.|author-link1=Alan Meerow|c=Amaryllidaceae |pages=83–110 |year=1998 |in=Kubitzki}} {{doi|10.1007/978-3-662-03533-7_11}} [https://books.google.com/books?id=FyPVYzL76sMC&pg=PA85 (excerpts)] * {{cite book|last1=Traub|first1=H.P.|author-link=Hamilton Traub|title=Genera of the Amaryllidaceae |date=1963|publisher=American Plant Life Society|location=La Jolla, California}}

=== Articles ===

* {{cite journal|last1=García|first1=Nicolás|last2=Meerow|first2=Alan W.|last3=Soltis|first3=Douglas E. |last4=Soltis|first4=Pamela S.|author-link2=Alan Meerow|author-link4 = Pamela S. Soltis|title=Testing Deep Reticulate Evolution in Amaryllidaceae Tribe Hippeastreae (Asparagales) with ITS and Chloroplast Sequence Data|journal=Systematic Botany|date=1 March 2014|volume=39|issue=1|pages=75–89 |doi=10.1600/036364414X678099 |bibcode=2014SysBo..39...75G |s2cid=86117335 |ref={{harvid|Garcia et al|2014}}}} * {{cite journal |last1=García |first1=Nicolás |last2=Folk |first2=Ryan A. |last3=Meerow |first3=Alan W. |last4=Chamala |first4=Srikar |last5=Gitzendanner |first5=Matthew A. |last6=Oliveira |first6=Renata Souza de |last7=Soltis |first7=Douglas E. |last8=Soltis |first8=Pamela S. |author-mask1=1|title=Deep reticulation and incomplete lineage sorting obscure the diploid phylogeny of rain-lilies and allies (Amaryllidaceae tribe Hippeastreae) |journal=Molecular Phylogenetics and Evolution |date=June 2017 |volume=111 |pages=231–247 |doi=10.1016/j.ympev.2017.04.003|pmid=28390909 |ref={{harvid|Garcia et al|2017}}|doi-access=free |bibcode=2017MolPE.111..231G }} * {{cite journal |last1=García |first1=Nicolás |last2=Meerow |first2=Alan W. |last3=Arroyo-Leuenberger |first3=Silvia |last4=Oliveira |first4=Renata S. |last5=Dutilh |first5=Julie H.|last6=Soltis |first6=Pamela S. |last7=Judd |first7=Walter S. |author-mask1=1|title=Generic classification of Amaryllidaceae tribe Hippeastreae |journal=Taxon |date=June 2019 |volume=68 |issue=3 |pages=481–498 |doi=10.1002/tax.12062|bibcode=2019Taxon..68..481G |s2cid=202854432 |ref={{harvid|Garcia et al|2019}}}}{{link note|note=With [https://doi.org/10.1002/tax.12208 corrections] published 2020}} * {{cite journal |last1=Meerow |first1=A.W. |last2=Fay |first2=M.F. |last3=Guy |first3=C.L. |last4=Li |first4=Q.-B. |last5=Zaman |first5=F.Q. |last6=Chase |first6=M.W. |year=1999 |title=Systematics of Amaryllidaceae based on cladistic analysis of plastid rbcL and trnL-F sequence data |journal=Am. J. Bot. |volume=86 |issue=9 |pages=1325–1345 |doi=10.2307/2656780 |pmid=10487820 |jstor=2656780 |ref={{harvid|Meerow et al|1999}}}} * {{cite journal |last1=Meerow |first1=A.W. |last2=Guy |first2=C.L. |last3=Li |first3=Q.-B. |last4=Yang |first4=S.-L. |author-link1=Alan Meerow |author-mask1=1 |title=Phylogeny of the American Amaryllidaceae Based on nrDNA ITS Sequences |journal=Systematic Botany |year=2000 |volume=25 |issue=4 |pages=708–726 |doi=10.2307/2666729 |jstor=2666729 |bibcode=2000SysBo..25..708M |s2cid=20392462 |url=http://www.bulbsociety.org/meerow/Meerow%20et%20al-American.pdf |access-date=2015-01-25 |archive-url=https://web.archive.org/web/20150923195144/http://www.bulbsociety.org/meerow/Meerow%20et%20al-American.pdf |archive-date=2015-09-23 |ref={{harvid|Meerow et al|2000}}}} * {{cite journal |last1=Meerow |first1=Alan W. |last2=Gardner |first2=Elliot M. |last3=Nakamura |first3=Kyoko |author-link1=Alan Meerow|author-mask1=1|title=Phylogenomics of the Andean Tetraploid Clade of the American Amaryllidaceae (Subfamily Amaryllidoideae): Unlocking a Polyploid Generic Radiation Abetted by Continental Geodynamics |journal=Frontiers in Plant Science |date=5 November 2020 |volume=11 |article-number=582422 |doi=10.3389/fpls.2020.582422|pmid=33250911 |pmc=7674842 |bibcode=2020FrPS...1182422M |doi-access=free }} * {{cite journal|last1=Müller-Doblies|first1=U.|last2=Müller-Doblies|first2=D.|author-link2=Dietrich Müller-Doblies|author-link1=Ute Müller-Doblies|title=Tribes and subtribes and some species combinations in Amaryllidaceae J St Hil R Dahlgren & al. 1985|journal=Feddes Repertorium|date=1996|volume=107|issue=5–6 |pages=S.c.1–S.c.9}}

=== Websites ===

* {{citation |last=Stevens |first=P.F. |date=2001{{ndash}}2012 |title=Angiosperm Phylogeny Website: Amaryllidoideae |url=https://www.mobot.org/mobot/research/apweb/orders/asparagalesweb.htm#Amaryllidaceae}} * {{cite web|last1=Vigneron|first1=Pascal|title=Amaryllidaceae|url=http://www.amaryllidaceae.org//index.htm |website=Amaryllidaceae.org|date=2008|access-date=23 October 2014|language=fr|archive-url= https://web.archive.org/web/20150104114211/http://www.amaryllidaceae.org/index.htm|archive-date=2015-01-04}} * {{cite web|last=GRIN|title=GRIN Taxonomy for Plants|publisher=Germplasm Resources Information Network |date=2016 |url=https://npgsweb.ars-grin.gov/gringlobal/taxon/abouttaxonomy.aspx|access-date=2016-02-15}} * {{cite web|title=Amaryllidaceae: A taxonomic tool for the Amaryllidaceae of the world|publisher=eMonocot |url=http://amaryllidaceae.e-monocot.org/|access-date=2016-02-14|archive-url= https://web.archive.org/web/20160304161517/http://amaryllidaceae.e-monocot.org/|archive-date=2016-03-04}} {{Refend}}

== External links == * [http://amaryllidaceae.e-monocot.org/taxonomy/term/17198 Amaryllidaceae project at eMonocot] {{Webarchive|url=https://web.archive.org/web/20160304060507/http://amaryllidaceae.e-monocot.org/taxonomy/term/17198 |date=2016-03-04 }}

Category:Amaryllidoideae Category:Asparagales tribes