Editing Bromeliaceae

{{Short description|Family of monocot flowering plants}}
{{Redirect|Bromeliad|the trilogy of children's books|The Nome Trilogy}}
{{Automatic taxobox
 | fossil_range = {{Fossil range|100|0}}
 | image = Pineapple1.JPG
 | image_caption = [[Pineapple]] (''Ananas comosus''), a bromeliad of economic importance
 | taxon = Bromeliaceae
 | authority = [[Antoine Laurent de Jussieu|Juss.]]<ref name="apgiii">{{Citation |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free }}</ref>
 | range_map = WorldBromeliadDistribution.PNG
 | subdivision_ranks = Subfamilies
 | subdivision = 
*[[Brocchinioideae]]
*[[Bromelioideae]]
*[[Hechtioideae]]
*[[Lindmanioideae]]
*[[Navioideae]]
*[[Pitcairnioideae]]
*[[Puyoideae]]
*[[Tillandsioideae]]
}}

The '''Bromeliaceae''' (the '''bromeliads''') are a [[Family (biology)|family]] of [[monocot]] [[flowering plant]]s of about 80 genera and 3700 known species,<ref name=EOBs>{{citation |mode=cs2 |last1=Gouda |first1=E.J. |last2=Butcher |first2=D. |last3=Gouda |first3=C.S. |date=2022 |title=Species and Infra Species Counts |website=Encyclopaedia of Bromeliads |publisher=Utrecht University Botanic Gardens |url=https://bromeliad.nl/encyclopedia/brome.php?action=counts |access-date=2022-11-24}}</ref> native mainly to the [[Tropics|tropical]] [[Americas]], with several species found in the American [[subtropics]] and one in tropical [[west Africa]], ''[[Pitcairnia feliciana]]''.<ref name=Mabberley>{{cite book |first=D.J. |last=Mabberley |year=1997 |title=The Plant Book |url=https://archive.org/details/plantbookportabl00mabb |url-access=registration |publisher=[[Cambridge University Press]] |location=[[Cambridge]]|isbn=9780521414210 }}</ref>

It is among the [[basal (phylogenetics)|basal]] families within the [[Poales]] and is the only family within the order that has [[Septal nectary|septal nectaries]] and [[Ovary (plants)|inferior ovaries]].<ref name=Judd>Judd, Walter S. Plant systematics a phylogenetic approach. 3rd ed. Sunderland, MA: Sinauer Associates, Inc., 2007.</ref> These [[Ovary (plants)|inferior ovaries]] characterize the [[Bromelioideae]], a subfamily of the Bromeliaceae.<ref name=Sajo>{{cite journal | last1 = Sajo | first1 = M. G. | year = 2004 | title = Floral anatomy of Bromeliaceae, with particular reference to the epigyny and septal nectaries in commelinid monocots | journal = Plant Systematics and Evolution | volume = 247 | issue = 3–4| pages = 215–31 | doi=10.1007/s00606-002-0143-0| s2cid = 20457047 }}</ref> The family includes both [[epiphyte]]s, such as [[Spanish moss]] (''[[Tillandsia usneoides]]''), and [[Terrestrial plant|terrestrial]] species, such as the [[pineapple]] (''[[Ananas comosus]]''). Many bromeliads are able to store water in a structure formed by their tightly overlapping [[leaf]] bases. However, the family is diverse enough to include the tank bromeliads, grey-leaved [[epiphytic|epiphyte]] ''[[Tillandsia]]'' species that gather water only from leaf structures called [[trichome]]s, and many desert-dwelling [[succulent]]s.

The largest bromeliad is ''[[Puya raimondii]]'', which reaches {{convert|3–4|metre}} tall in vegetative growth with a flower spike {{convert|9–10|metre}} tall,<ref>{{Cite web|url=http://www.llifle.com/Encyclopedia/BROMELIADS/Family/Bromeliaceae/34205/Puya_raimondii|title=Llifle|website=Encyclopaedia of living things}}</ref><ref>{{Cite web|url=http://www.strangewonderfulthings.com/186.htm|title=Puya Raimondii - World's Largest Bromeliad|website=Strange Wonderful Things}}</ref> and the smallest is Spanish moss.{{citation needed|date=September 2016}}

==Description==
[[Image:Bromeliad3.jpg|thumb|Bromeliad]]
Bromeliads are mostly [[Herbaceous plant|herbaceous]] [[Perennial plant|perennials]], although a few have a more tree-like habit. Many are more or less [[Succulent plant|succulent]] or have other adaptations to resist drought. They may be terrestrial or [[Epiphyte|epiphytic]], rarely climbing (e.g. ''[[Pitcairnia]]'' species).<ref name=WatsDall92/> Some species of ''[[Tillandsia]]'' (e.g. [[Spanish moss]], ''Tillandsia usneoides'') are  [[Raunkiær plant life-form#Aerophytes|aerophyte]]s, which have very reduced root systems and absorb water directly from the air.<ref name=GaláHageVice99>{{Cite journal |last1=Galán de Mera |first1=A. |last2=Hagen |first2=M.A. |last3=Vicente Orellana |first3=J.A. |date=1999 |title=Aerophyte, a New Life Form in Raunkiaer's Classification? |journal=Journal of Vegetation Science |volume=10 |issue=1 |pages=65–68 |doi=10.2307/3237161 |jstor=3237161 |name-list-style=amp }}</ref> Many terrestrial and epiphytic bromeliads have their leaves in the form of vase-shaped rosettes which accumulate water. These rosettes, called "tanks", can hold as much as ten liters (eighteen pints) of water, and be little biotic communities unto themselves. One individual tank was found to contain the following: four [[harvestmen]], a spider, three species of [[wood lice]], a centipede, a "jumping millipede"[sic], a [[pseudoscorpion]], "various metallic beetles", [[earwigs]], a tree seedling, [[Chironomidae]] fly larva, an ant colony, an earthworm, numerous mites, and a small frog.<ref>{{cite book | editor= Silcock, Lisa |date=1992 | title= The rainforests - A Celebration | location= San Francisco | publisher= Chronicle Books | page= 155 | isbn= 0-8118-0155-1 }}</ref> Individual leaves are not divided and have parallel veins without cross connections. The epidermis of the leaf contains silica. Bromeliad flowers are aggregated into [[inflorescence]]s of various forms. The flowers have bracts, often brightly coloured, and distinct calyces of three [[sepal]]s and corollas of three [[petal]]s. The flowers have [[Nectar#Floral nectaries|nectaries]]. They are [[Pollination|pollinated]] by insects, birds (often [[hummingbird]]s) or bats, or more rarely (in ''[[Navia (plant)|Navia]]'') they are wind-pollinated. Fruits are variable, typically taking the form of a capsule or a berry.<ref name=WatsDall92>{{Citation |mode=cs1 |last1=Watson |first1=L. |last2=Dallwitz |first2=M.J. |date=1992–2021 |title=Bromeliaceae Juss. in ''The families of flowering plants: descriptions, illustrations, identification, and information retrieval'' |website=delta-intkey.com |url=https://www.delta-intkey.com/angio/www/bromelia.htm |archive-url=https://web.archive.org/web/20211030053016/https://www.delta-intkey.com/angio/www/bromelia.htm |archive-date=2021-10-30 |name-list-style=amp }}</ref>

Bromeliads are able to live in an array of environmental conditions due to their many adaptations. [[Trichome]]s, in the form of scales or hairs, allow bromeliads to capture water in cloud forests and help to reflect sunlight in desert environments.<ref name= Schulte>{{cite journal | last1 = Schulte | first1 = Katharina | last2 = Barfuss | first2 = Michael H. | last3 = Zizka | first3 = Georg | year = 2009 | title = Phylogeny of Bromelioideae (Bromeliaceae) inferred from nuclear plastid DNA loci reveals the evolution of the tank habit within the subfamily | journal = Molecular Phylogenetics and Evolution | volume = 51 | issue = 2| pages = 327–39 | doi=10.1016/j.ympev.2009.02.003| pmid = 19236934 }}</ref> Bromeliads with leaf vases can capture water and nutrients in the absence of a well-developed root system.<ref name="Schulte"/> Many bromeliads also use [[crassulacean acid metabolism]] (CAM) [[photosynthesis]] to create sugars. This adaptation allows bromeliads in hot or dry climates to open their [[stoma]]ta at night rather than during the day, which reduces water loss.<ref name= Rex>{{cite journal | last1 = Rex | first1 = Martina | last2 = Patzolt | first2 = Kerstin | last3 = Schulte | first3 = Katharina | last4 = Zizka | first4 = Georg | last5 = Vasquuez | first5 = Roberto | last6 = Ibisch | first6 = Pierre L. | last7 = Weising | first7 = Kurt | year = 2007 | title = AFLP analysis of genetic relationships in the genus Fosterella L.B. Smith (Pitcairnioideae, Bromeliaceae) | doi = 10.1139/g06-141 | pmid = 17546075 | journal = Genome | volume = 50 | issue = 1| pages = 90–105 }}</ref> Both CAM and epiphytism have evolved multiple times within the family, with some taxa reverting to [[C3 carbon fixation|C3 photosynthesis]] as they radiated into less arid climates.<ref name=":1" />

==Evolution==
Bromeliads are among the more recent plant groups to have emerged. They are thought to have originated in the [[tepui]]s of the [[Guiana Shield]] approximately 100 million years ago. The greatest number of extant [[Basal (phylogenetics)|basal]] species are found in the [[Andes|Andean]] highlands of South America.<ref name= Givnish>{{cite journal | last1 = Givnish | first1 = Thomas J. | last2 = Millam | first2 = Kendra C. | last3 = Evans | first3 = Timothy M. | last4 = Hall | first4 = Jocelyn C. | last5 = Pires | first5 = J. C. | last6 = Berry | first6 = Paul E. | last7 = Sytsma | first7 = Kenneth J. | year = 2004 | title = Ancient vicariance or recent long-distance dispersal? Inferences about phylogeny and South American-African disjunctions in Raptaceae and Bromeliaceae based on ndhf sequence data | journal = International Journal of Plant Sciences | volume = 165 | issue = 4| pages = 35–54 | doi=10.1086/421067| s2cid = 18808651 }}</ref> However, the family did not diverge into its extant subfamilies until 19 million years ago. The long period between the origin and diversification of bromeliads, during which no extant species evolved, suggests that there was much [[speciation]] and extinction during that time, which would explain the genetic distance of the Bromeliaceae from other families within the Poales.<ref name= Barfuss/>

Based on [[Molecular phylogenetics|molecular phylogenetic]] studies, the family is divided into eight subfamilies. The relationship among them is shown in the following [[cladogram]].<ref name=Givnish/>
{{clade
|label1=Bromeliaceae
|newick1=([[Brocchinioideae]], ([[Lindmanioideae]], ([[Tillandsioideae]], ([[Hechtioideae]], ([[Navioideae]], ([[Pitcairnioideae]], ([[Puyoideae]], [[Bromelioideae]])))))))
}}

The most basal genus, ''[[Brocchinia]]'' (subfamily Brocchinioideae), is endemic to the Guiana Shield, and is placed as the [[Cladistics|sister group]] to the remaining [[Genus|genera]] in the family.<ref name=Barfuss>{{cite journal | last1 = Barfuss | first1 = Michael H. | last2 = Samuel | first2 = Rosabelle | last3 = Till | first3 = Walter | last4 = Stuessy | first4 = Todd F. | year = 2005 | title = Phylogenetic relationships in subfamily Tillandsioideae (Bromeliaceae) based on DNA sequence data from seven plastid regions | journal = American Journal of Botany | volume = 92 | issue = 2| pages = 337–51 | doi=10.3732/ajb.92.2.337| pmid = 21652410 | doi-access = free }}</ref> The subfamilies  [[Lindmanioideae]] and [[Navioideae]] are endemic to the Guiana Shield as well.<ref>{{cite journal |last1=Yardeni |first1=Gil |last2=Viruel |first2=Juan |last3=Paris |first3=Margot |last4=Hess |first4=Jaqueline |last5=Groot Crego |first5=Clara |last6=de La Harpe |first6=Marylaure |last7=Rivera |first7=Norma |last8=Barfuss |first8=Michael H. J. |last9=Till |first9=Walter |last10=Guzmán-Jacob |first10=Valeria |last11=Krömer |first11=Thorsten |last12=Lexer |first12=Christian |last13=Paun |first13=Ovidiu |last14=Leroy |first14=Thibault |title=Taxon-specific or universal? Using target capture to study the evolutionary history of a rapid radiation |journal=[[Molecular Ecology Resources]] |date=2021-05-22 |volume=22 |issue=3 |pages=927–945 |doi=10.1111/1755-0998.13523 |pmid=34606683 |pmc=9292372 |s2cid=238357548 | doi-access=free}}</ref>

The [[West Africa]]n species ''[[Pitcairnia feliciana]]'' is the only bromeliad not [[Endemism|endemic]] to the Americas, and is thought to have reached Africa via long-distance dispersal about 12 million years ago.<ref name="Givnish" />

=== Radiation of Tillandsioideae and ''Hechtia'' ===
The first groups to leave the Guiana Shield were the subfamily [[Tillandsioideae]], which spread gradually into northern South America, and the genus ''[[Hechtia]]'' (Hechtioideae), which spread to Central America via long-distance dispersal. Both of these movements occurred approximately 15.4 million years ago. When it reached the Andes mountains, the speciation of Tillandsioideae occurred quite rapidly, largely due to the [[Andean orogeny|Andean uplift]], which was also occurring rapidly from 14.2 to 8.7 million years ago. The uplift  greatly altered the region's geological and climatic conditions, creating a new mountainous environment for the epiphytic tillandsioids to colonize. These new conditions directly drove the speciation of the Tillandsioideae, and also drove the speciation of their animal pollinators, such as [[hummingbird]]s.<ref name=":0" /><ref name=":1" /><ref>{{Cite journal|last=Bleiweiss|first=Robert|date=September 1998|title=Tempo and mode of hummingbird evolution|journal=Biological Journal of the Linnean Society|language=en|volume=65|issue=1|pages=63–76|doi=10.1111/j.1095-8312.1998.tb00351.x|doi-access=free}}</ref><ref>{{Cite journal|last1=Hoorn|first1=C.|last2=Wesselingh|first2=F. P.|last3=ter Steege|first3=H.|last4=Bermudez|first4=M. A.|last5=Mora|first5=A.|last6=Sevink|first6=J.|last7=Sanmartin|first7=I.|last8=Sanchez-Meseguer|first8=A.|last9=Anderson|first9=C. L.|last10=Figueiredo|first10=J. P.|last11=Jaramillo|first11=C.|date=2010-11-12|title=Amazonia Through Time: Andean Uplift, Climate Change, Landscape Evolution, and Biodiversity |journal=Science|language=en|volume=330|issue=6006|pages=927–931|doi=10.1126/science.1194585|pmid=21071659|bibcode=2010Sci...330..927H|s2cid=206528591|issn=0036-8075|url=https://www.zora.uzh.ch/id/eprint/42535/10/Figure2.pdf}}</ref>

=== Evolution of the Bromelioideae ===
Around 5.5 million years ago, a clade of epiphytic [[Bromelioideae|bromelioids]] arose in [[Serra do Mar]], a lush mountainous region on the coast of Southeastern Brazil. This is thought to have been caused not only by the uplift of Serra do Mar itself at that time, but also because of the continued uplift of the distant Andes mountains, which impacted the circulation of air and created a cooler, wetter climate in Serra do Mar.<ref name=":1" /> These epiphytes thrived in this humid environment, since their trichomes rely on water in the air rather than from the ground like terrestrial plants. Many epiphytic bromeliads with the tank habit also speciated here.

Even before this, a few other bromelioids had already dispersed to the Brazilian shield while the climate was still arid, likely through a gradual process of short-distance dispersal. These make up the terrestrial members of the Bromelioideae, which have highly [[xeromorphic]] characters.<ref name=":1" />

==Classification ==
The family Bromeliaceae is currently placed in the order [[Poales]].

===Subfamilies===
The family Bromeliaceae is organized into eight subfamilies:<ref name=":0">{{cite journal|last1=Givnish|first1=Thomas|last2=Millam|first2=Kendra|last3=Berry|first3=Paul|last4=Sytsma|first4=Kenneth|title=Phylogeny, Adaptive Radiation, and Historical Biogeography of Bromeliaceae Inferred from ndhF Sequence Data|url=https://www.researchgate.net/publication/237426562|journal=Aliso|volume=23|issue=1|year=2007|pages=3–26|issn=2327-2929|doi=10.5642/aliso.20072301.04|doi-access=free}}</ref>

*[[Brocchinioideae]]
*[[Lindmanioideae]]
*[[Tillandsioideae]]
*[[Hechtioideae]]
*[[Navioideae]]
*[[Pitcairnioideae]]
*[[Puyoideae]]
*[[Bromelioideae]]

Bromeliaceae were originally split into three subfamilies based on morphological seed characters: Bromelioideae (seeds in [[Berry|baccate]] fruits), Tillandsioideae (plumose seeds), and Pitcairnioideae (seeds with wing-like appendages).<ref>{{Cite book |title = Flora Neotropica: Monograph 14 |vauthors=Smith LB, Downs RJ |date = 1974|editor= New York Botanical Garden |volume=2 |publisher=Hafner Press |location=New York}}</ref> However, molecular evidence has revealed that while Bromelioideae and Tillandsioideae are monophyletic, Pitcairnioideae as traditionally defined is [[Paraphyly|paraphyletic]]<ref>{{Cite journal|url =http://repository.uwyo.edu/cgi/viewcontent.cgi?article=1010&context=botany_facpub |title = Examination of subfamilial phylogeny in Bromeliaceae using comparative sequencing of the plastid locus ndhF|last = Terry|first = Randall|date = 1997|journal = American Journal of Botany|doi = 10.2307/2445903|pmid =21708619|volume=84|issue = 5|pages = 664–670|jstor = 2445903|hdl = 20.500.11919/753|hdl-access = free}}</ref> and should be split into six subfamilies: Brocchinioideae, Lindmanioideae, Hechtioideae, Navioideae, Pitcairnioideae, and Puyoideae.<ref>{{Cite journal|title = Genetics, evolution and conservation of Bromeliaceae|last = Zanella|first = Camila|date = 2012|journal = Genetics and Molecular Biology|doi = 10.1590/s1415-47572012000600017|pmid = 23412953|volume=35|issue = 4 suppl 1|pages=1020–1026|pmc = 3571438}}</ref>

Brocchinioideae is defined as the most basal branch of Bromeliaceae based on both morphological and molecular evidence, namely genes in chloroplast DNA.<ref>{{Cite journal|title = Molecular phylogenetics of Bromeliaceae: evidence from trnL (UAA) intron sequences of the chloroplast genome|last = Horres|first = Ralf|date = 2000|journal = Plant Biology|doi =10.1055/s-2000-3700|volume=2|issue = 3|pages=306–315}}</ref>

Lindmanioideae is the next most basal branch distinguished from the other subfamilies by convolute sepals and chloroplast DNA.<ref name=":1">{{Cite journal|url = http://repository.uwyo.edu/cgi/viewcontent.cgi?article=1011&context=botany_facpub|title = Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: insights from an eight-locus plastid phylogeny|last = Givnish|first = Thomas|date = 2011|journal = American Journal of Botany|doi = 10.3732/ajb.1000059 |pmid =21613186|volume=98|issue = 5|pages=872–895|hdl = 2027.42/142109|hdl-access = free}}</ref>

Hechtioideae is also defined based on analyses of chloroplast DNA; similar morphological adaptations to arid environments also found in other groups (namely the genus ''[[Puya (plant)|Puya]])'' are attributed to [[convergent evolution]].<ref name=":0" />

Navioideae is split from Pitcairnioideae based on its cochlear sepals and chloroplast DNA.<ref>{{Cite journal|title = Multiple origins of crassulacean acid metabolism and the epiphytic habit in the Neotropical family Bromeliaceae|last = Crayn|first = Darren|date = 2004|journal = Proceedings of the National Academy of Sciences|doi = 10.1073/pnas.0400366101 |pmid =14982989|volume=101|issue = 10|pages=3703–3708|pmc =373526|bibcode = 2004PNAS..101.3703C|doi-access = free}}</ref>

Puyoideae has been re-classified multiple times and its monophyly remains controversial according to analyses of chloroplast DNA.<ref name=":1" />

===Genera===
{{As of|2025|February}}, [[Plants of the World Online]] (PoWO) accepted 76 genera, as listed below.<ref name="POWO_30000083-2">{{cite web |title=''Bromeliaceae'' Juss. |work=Plants of the World Online |publisher=Royal Botanic Gardens, Kew|url=https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30000083-2 |access-date=2025-02-04 }}</ref> A few more genera were accepted by the Encyclopaedia of Bromeliads, including ''Josemania'' and ''Mezobromelia'', which PoWO sinks into ''Cipuropsis''.
{{div col}}
*''[[Acanthostachys]]'' <small>[[Johann Friedrich Klotzsch|Klotzsch]] </small>
*''[[Aechmea]]'' <small>[[Hipólito Ruiz López|Ruiz]] & [[Pav.]]</small>
*''[[Alcantarea]]'' <small>[[E.Morren ex Mez|Harms]]</small>
*''[[Ananas]]'' <small>[[Mill.]]</small>, including ''Pseudananas'' <small>Hassl. ex [[Hermann August Theodor Harms|Harms]]</small> (includes the [[pineapple]])
*''[[Androlepis]]'' <small>[[Brongn.]] ex [[Houllet]] </small>
*''[[Araeococcus]]'' <small>Brongn.</small>
*''[[Bakerantha]]'' <small>[[L.B.Sm.]]</small>
*''[[Barfussia]]'' <small>[[Manzan. & W.Till]]</small>
*''[[Billbergia]]'' <small>[[Thunb.]]</small>
*''[[Brewcaria]]'' <small>L.B.Sm., [[Steyerm.]] & [[H.Rob]]</small>,<ref name=EOB_Brewcaria>{{citation |mode=cs1 |last1=Gouda |first1=E.J. |last2=Butcher |first2=D. |last3=Gouda |first3=C.S. |date=2022 |title=genus ''Brewcaria'' L.B.Sm., Steyerm. & H.Rob. |website=Encyclopaedia of Bromeliads |publisher=Utrecht University Botanic Gardens |url=https://bromeliad.nl/species/Brewcaria |access-date=2022-11-01}}</ref> synonym of ''Navia'' in PoWO
*''[[Brocchinia]]'' <small>[[Schult.f.]]</small>
*''[[Bromelia]]'' <small>[[Carl Linnaeus|L.]]</small>
*''[[Canistropsis]]'' <small>(Mez) Leme</small>
*''[[Canistrum]]'' <small>[[Charles Jacques Édouard Morren|E.Morren]]</small>
*''[[Catopsis]]'' <small>[[August Grisebach|Griseb.]]</small>
*''[[Cipuropsis]]'' <small>[[Ernst Heinrich Georg Ule|Ule]]</small>
*''[[Connellia]]'' <small>[[N.E.Br.]]</small>
*''[[Cottendorfia]]'' <small>[[Schult.f.]]</small>
*''[[Cryptanthus]]'' <small>[[Christoph Friedrich Otto|Otto]] & [[A.Dietr.]]</small>
*''[[Deinacanthon]]'' <small>Mez</small>
*''[[Deuterocohnia]]'' <small>[[Carl Christian Mez|Mez]]</small>
*''[[Disteganthus]]'' <small>[[Lem.]]</small>
*''[[Dyckia]]'' <small>[[Schult.f.]]</small>
*''[[Edmundoa]]'' <small>Leme</small>
*''[[Eduandrea]]'' <small>Leme, W.Till, G.K.Br., J.R.Grant & Govaerts</small>
*''[[Encholirium]]'' <small>[[Mart.]] ex [[Schult.f.]]</small>
*''[[Fascicularia]]'' <small>Mez</small>
*''[[Fernseea]]'' <small>[[John Gilbert Baker|Baker]]</small>
*''[[Forzzaea]]'' <small>Leme, S.Heller & Zizka</small>
*''[[Fosterella]]'' <small>L.B.Sm.</small>
*''[[Glomeropitcairnia]]'' <small>Mez</small>
*''[[Goudaea]]'' <small>W.Till & Barfuss</small>
*''[[Gregbrownia]]'' <small>W.Till & Barfuss</small>
*''[[Greigia]]'' <small>[[Eduard August von Regel|Regel]]</small>
*''[[Guzmania]]'' <small>[[Hipólito Ruiz López|Ruiz]] & [[Pav.]]</small>
*''[[Hechtia]]'' <small>[[Johann Friedrich Klotzsch|Klotzsch]]</small>
*''[[Hohenbergia]]'' <small>[[Schult.f.]]</small>
*''[[Hohenbergiopsis]]'' <small>L.B.Sm. & [[Read (botanical)|Read]]</small>
*''[[Hoplocryptanthus]]'' <small>(Mez) Leme, S.Heller & Zizka</small>
*''[[Hylaeaicum]]'' <small>(Ule ex Mez) Leme, Forzza, Zizka & Aguirre-Santoro</small>
*''[[Jagrantia]]'' <small>Barfuss & W.Till</small>
*''[[Josemania]]'' <small>W.Till & Barfuss</small>
*''[[Karawata]]'' <small>J.R.Maciel & G.M.Sousa</small>
*''[[Krenakanthus]]'' <small>(Leme, S.Heller & Zizka) Leme, Zizka & Paule</small>
*''[[Lapanthus]]'' <small>Louzada & Versieux</small>
*''[[Lemeltonia]]'' <small>Barfuss & W.Till</small>
*''[[Lindmania]]'' <small>Mez</small>
*''[[Lutheria]]'' <small>Barfuss & W.Till</small>
*''[[Lymania]]'' <small>Read</small>
*''[[Mezobromelia]]'' <small>L.B.Sm.</small>
*''[[Navia (plant)|Navia]]'' <small>[[Schult.f.]]</small>
*''[[Neoglaziovia]]'' <small>Mez</small>
*''[[Neoregelia]]'' <small>L.B.Sm. </small>
*''[[Nidularium]]'' <small>[[Lem.]]</small>
*''[[Ochagavia]]'' <small>[[Phil.]]</small>
*''[[Orthocryptanthus]]'' <small>(Leme, S.Heller & Zizka) Leme, Zizka & Paule</small>
*''[[Orthophytum]]'' <small>Beer</small>
*''[[Pitcairnia]]'' <small>[[L'Her.]]</small>, including [[Pitcairnia subg. Pepinia|subgenus ''Pepinia'']]<ref name=EOB_Pepinia>{{citation |mode=cs2 |last1=Gouda |first1=E.J. |last2=Butcher |first2=D. |last3=Gouda |first3=C.S. |date=2022 |title=''Pepinia'' (subgen. of ''Pitcairnia'') (Brongniart ex André) Baker |website=Encyclopaedia of Bromeliads |publisher=Utrecht University Botanic Gardens |url=https://bromeliad.nl/species/Pepinia/(subgen./of/Pitcairnia) |access-date=2022-11-01}}</ref>
*''[[Portea]]'' <small>[[K. Koch]]</small>
*''[[Pseudaechmea]]'' <small>L.B.Sm. & Read</small>,<ref name=EOB_Pseudaechmea>{{citation |mode=cs1 |last1=Gouda |first1=E.J. |last2=Butcher |first2=D. |last3=Gouda |first3=C.S. |date=2022 |title=genus ''Pseudaechmea'' L.B.Sm. & Read |website=Encyclopaedia of Bromeliads |publisher=Utrecht University Botanic Gardens |url=https://bromeliad.nl/species/Pseudaechmea |access-date=2022-11-01}}</ref> synonym of ''Billbergia'' in PoWO
*''[[Pseudalcantarea]]'' <small>(Mez) Pinzón & Barfuss</small>
*''[[Pseudaraeococcus]]'' <small>(Mez) R.A.Pontes & Versieux</small>
*''[[Puya (plant)|Puya]]'' <small>[[Juan Ignacio Molina|Molina]]</small>
*''[[Quesnelia]]'' <small>[[Gaudich.]]</small>
*''[[Racinaea]]'' <small>M.A.Spencer & L.B.Sm.</small>
*''[[Rokautskyia]]'' <small>Leme, S.Heller & Zizka</small>
*''[[Ronnbergia]]'' <small>E.Morren & [[Édouard André|André]]</small>
*''[[Sequencia]]'' <small>Givnish</small>
*''[[Sincoraea]]'' <small>Ule</small>
*''[[Siqueiranthus]]'' <small>Leme, Zizka, E.H.Souza & Paule</small>
*''[[Steyerbromelia]]'' <small>L.B.Sm.</small>
*''[[Stigmatodon]]'' <small>[[Leme, G.K.Br. & Barfuss]]</small>
*''[[Tillandsia]]'' <small>[[Carl Linnaeus|L.]]</small>
*''[[Ursulaea]]'' <small>Read & H.U.Baensch</small>,<ref name=EOB_Ursulaea>{{citation |mode=cs1 |last1=Gouda |first1=E.J. |last2=Butcher |first2=D. |last3=Gouda |first3=C.S. |date=2022 |title=genus ''Ursulaea'' Read & H.U.Baensch|website=Encyclopaedia of Bromeliads |publisher=Utrecht University Botanic Gardens |url=https://bromeliad.nl/species/Ursulaea |access-date=2022-11-01}}</ref> synonym of ''Aechmea'' in PoWO
*''[[Vriesea]]'' <small>[[Lindl.]]</small>
*''[[Wallisia]]'' <small>(Regel) É.Morren</small>
*''[[Waltillia]]'' <small>Leme, Barfuss & Halbritt.</small>
*''[[Werauhia]]'' <small>[[J.R.Grant]]</small>
*''[[Wittmackia]]'' <small>Mez</small>
*''[[Wittrockia]]'' <small>[[Lindm.]]</small>
*''[[Zizkaea]]'' <small>W.Till & Barfuss</small>
{{div col end}}

===Hybrid genera===
Intergeneric hybrid genera accepted by Plants of the World Online include:
*× ''Cryptbergia'' <small>R.G.Wilson & C.L.Wilson</small> =  ''Cryptanthus'' × ''Billbergia''
*× ''Guzlandsia'' <small>Gouda</small> = ''Guzmania'' × ''Tillandsia'' 
*× ''Hohenmea'' <small>B.R.Silva & L.F.Sousa</small> = ''Hohenbergia'' × ''Aechmea'' 
*× ''Niduregelia'' <small>Leme</small> = ''Nidularium'' × ''Neoregelia''

==Gallery==
<gallery>
File:Billbergia pyramidalis in Hyderabad Nursery W IMG 0428.jpg|''[[Billbergia pyramidalis]]''
File:條紋水塔花 20190712192104.jpg|''[[Billbergia vittata]] Brongniart''
File:Tillandsia usneoides04.jpg|''[[Tillandsia usneoides]]'' hanging from branches
File:Puya berteroana.jpg|''[[Puya alpestris]]''
File:Tillandsia tower arp.jpg|''[[Tillandsia]]'' airplants mounted on the bark of a cork oak
File:Del Monte Pineapple field at Camp Philips, Bukidnon, Philippines 03.jpg|A [[Del Monte Foods|Del Monte]] [[plantation]] of [[pineapple]] (''Ananas comosus'') plants in [[Bukidnon]], the Philippines
</gallery>

==Distribution and habitat==
[[File:Tillandsia sp. telephone line (codiferous).jpg|thumb|right|Bromeliads growing on telephone lines in [[Bolivia]]]]
Plants in the Bromeliaceae are widely represented in their natural climates across the Americas. One species (''[[Pitcairnia feliciana]]'') can be found in Africa.<ref>{{Cite journal|last1=Porembski|first1=Stefan|last2=Barthlott|first2=Wilhelm|date=1999|jstor=41761298|journal=Harvard Papers in Botany|volume=4|issue=1|pages=175–184|title=Pitcairnia Feliciana: The Only Indigenous African Bromeliad}}</ref> They can be found at altitudes from sea level to 4,200 meters, from [[Tropical and subtropical moist broadleaf forests|rainforests]] to [[desert]]s. 1,814 species are [[epiphyte]]s, some are [[lithophyte]]s, and some are terrestrial. Accordingly, these plants can be found in the [[Andean]] highlands, from northern Chile to Colombia, in the [[Sechura Desert]] of coastal Peru, in the [[cloud forest]]s of Central and South America, in southern United States from southern [[Virginia]] to [[Florida]] to [[Texas]], and in far southern [[Arizona]].

== Ecology ==
Bromeliads often serve as [[phytotelma]]ta, accumulating water between their leaves. One study found 175,000 bromeliads per hectare (2.5 acres) in one forest; that many bromeliads can sequester 50,000 liters (more than 13,000 gallons) of water.<ref name="nyt">"[https://opinionator.blogs.nytimes.com/2008/03/18/pineapple-dreams Pineapple Dreams]", The Wild Side, Olivia Judson, ''[[The New York Times]]'', March 18, 2008</ref> The aquatic habitat created as a result is host to a diverse array of [[invertebrate]]s, especially aquatic insect larvae,<ref>{{Cite journal|last1=Frank|first1=J. H.|last2=Lounibos|first2=L. P.|date=2009-02-01|title=Insects and allies associated with bromeliads: a review|journal=Terrestrial Arthropod Reviews|volume=1|issue=2|pages=125–153|doi=10.1163/187498308X414742|issn=1874-9836|pmc=2832612|pmid=20209047}}</ref><ref>Picado, C. (1913). Les broméliacées épiphytes considérées comme milieu biologique. Bulletin scientifique de la France et de la Belgique 5: 215-360</ref> including those of mosquitos.<ref>[https://www.sun-sentinel.com/health/fl-reg-miami-worst-mosquitoes-cities-20180820-story.html] Life in South Florida can be an itch – but other places are worse</ref> These bromeliad [[invertebrate]]s benefit their hosts by increasing [[nitrogen]] uptake into the plant.<ref>{{Cite journal|last1=Ngai |first1=Jacqueline T.| last2=Srivastava |first2=Diane S. |author-link2=Diane Srivastava |date=2006-11-10 |title=Predators Accelerate Nutrient Cycling in a Bromeliad Ecosystem |journal=Science |language=en |volume=314 |issue=5801|pages=963 |doi=10.1126/science.1132598|issn=0036-8075|pmid=17095695|s2cid=27072688}}</ref><ref>{{Cite journal|last1=Leroy|first1=Céline|last2=Corbara|first2=Bruno|last3=Dejean|first3=Alain|last4=Céréghino|first4=Régis|date=2009-09-01|title=Ants mediate foliar structure and nitrogen acquisition in a tank-bromeliad|journal=New Phytologist|language=en|volume=183|issue=4|pages=1124–1133|doi=10.1111/j.1469-8137.2009.02891.x|issn=1469-8137|pmid=19500265|doi-access=free}}</ref><ref>{{Cite journal|last1=Romero|first1=Gustavo Q.|last2=Srivastava|first2=Diane S. |author-link2=Diane Srivastava |date=2010-09-01|title=Food-web composition affects cross-ecosystem interactions and subsidies|journal=Journal of Animal Ecology|language=en|volume=79|issue=5|pages=1122–1131|doi=10.1111/j.1365-2656.2010.01716.x|pmid=20584097|issn=1365-2656|doi-access=free}}</ref> A study of 209 plants from the Yasuní Scientific Reserve in [[Ecuador]] identified 11,219 animals, representing more than 350 distinct species,<ref>{{Cite journal |last1=Armbruster |first1=Peter |last2=Hutchinson |first2=Robert A. |last3=Cotgreave |first3=Peter |date=February 2002 |title=Factors influencing community structure in a South American tank bromeliad fauna |url=https://doi.org/10.1034/j.1600-0706.2002.960204.x |journal=Oikos |volume=96 |issue=2 |pages=225–234 |doi=10.1034/j.1600-0706.2002.960204.x |issn=0030-1299}}</ref> many of which are found only on bromeliads. Examples include some species of [[ostracod]]s, small [[salamander]]s about {{convert|2.5|cm|in|0|abbr=on}} in length, and [[tree frog]]s. [[Jamaica]]n bromeliads are home to ''[[Metopaulias|Metopaulias depressus]]'', a reddish-brown [[crab]] {{convert|2|cm|in|1|abbr=on}} across, which has evolved social behavior to protect its young from predation by ''[[Diceratobasis macrogaster]]'', a species of [[damselfly]] whose [[larva]]e live in bromeliads. Some bromeliads even form homes for other species of bromeliads.<ref name="nyt"/>

Trees or branches that have a higher incidence of sunlight tend to have more bromeliads. In contrast, the sectors facing west receive less sunlight and therefore fewer bromeliads. In addition, thicker trees have more bromeliads, possibly because they are older and have greater structural complexity.<ref>Gename, K., & Monge-Nájera, J. (2012). How organisms reach and colonize bromeliads: a field experimental test of two of Picado’s hypotheses, and the effect of tree age and cardinal distribution on bromeliads in Cartago, Costa Rica. UNED Research Journal, 4(2), 181-186.</ref><ref>López, L. C. S., Alves, R. R. D. N., & Ríos, R. I. (2009). Micro-environmental factors and the endemism of bromeliad aquatic fauna. Hydrobiología, 625(1), 151-156.</ref>

==Cultivation and uses==
[[File:Bromeliaceae mixed collection.jpg|thumb|Bromeliaceae mixed cultivated collection]]
Humans have been using bromeliads for thousands of years. The [[Incas]], [[Aztecs]], [[Maya civilization|Maya]] and others used them for food, protection, fiber and ceremony, just as they are still used today. European interest began when Spanish conquistadors returned with [[pineapple]], which became so popular as an exotic food that the image of the pineapple was adapted into European art and sculpture. In 1776, the species ''[[Guzmania lingulata]]'' was introduced to Europe, causing a sensation among gardeners unfamiliar with such a plant. In 1828, ''[[Aechmea fasciata]]'' was brought to Europe, followed by ''[[Vriesea splendens]]'' in 1840. These transplants were so successful, they are still among the most widely grown bromeliad varieties.

In the 19th century, breeders in Belgium, France and the Netherlands started [[Hybrid (biology)#Hybrid plants|hybridizing]] plants for wholesale trade. Many exotic varieties were produced until World War I, which halted breeding programs and led to the loss of some species. The plants experienced a resurgence of popularity after World War II. Since then, [[Netherlands|Dutch]], [[Belgium|Belgian]] and North American nurseries have greatly expanded bromeliad production.

Only one bromeliad, the pineapple (''Ananas comosus''), is a commercially important food crop. [[Bromelain]], a common ingredient in meat tenderizer, is extracted from pineapple stems. Many other bromeliads are popular [[ornamental plant]]s, grown as both garden and [[houseplant]]s.

Bromeliads are important food plants for many peoples. For example, the [[Pima people|Pima]] of Mexico occasionally consume flowers of ''[[Tillandsia erubescens]]'' and ''[[Tillandsia recurvata|T. recurvata]]'' due to their high sugar content; in Argentina and Bolivia, the shoot apices of ''[[Tillandsia rubella|T. rubella]]'' and ''[[Tillandsia maxima|T. maxima]]'' are consumed; in Venezuela, indigenous coastal tribes eat a sour-tasting but sweet-smelling berry, known as 'Maya', of ''[[Bromelia chrysantha]]'' as a fruit or in [[fermented beverage]]s; in Chile, the sweet fruit of ''[[Greigia sphacelata]],'' known as 'chupones', is consumed raw.<ref name = bromeliads2011polibotanica/>

===Collectors===
[[Édouard André]] was a French collector/explorer whose many discoveries of bromeliads in the Cordilleras of South America would be influential on horticulturists to follow. He served as a source of inspiration to 20th-century collectors, in particular [[Mulford B. Foster]] and [[Lyman Bradford Smith|Lyman Smith]] of the United States and [[Werner Rauh]] of Germany and Michelle Jenkins of Australia.<ref>André, Édouard François. "Bromeliaceae Andreanae. Description et histoire des Bromeliacees recoltees dans La Colombie, L'Ecuador et Le Venezuela". Paris: Librairie Agricole; G. Masson, 1889</ref>

==See also==
*[[List of foliage plant diseases (Bromeliaceae)]]

==References==
{{Reflist|30em| refs =

<ref name = bromeliads2011polibotanica>{{cite journal | title = Bromeliads: Traditional Plant Food in Latin America Since pre-Hispanic Times | url = https://www.academia.edu/866074 | date = 2011 | last1 = Hornung-Leoni | journal = Polibotánica | volume = 32 | pages = 219–229 | access-date = 30 Mar 2020}}</ref>

}}

==External links==
{{Commons category|Bromeliaceae}}
{{Wikispecies|Bromeliaceae}}
* [https://bromeliad.nl/taxonList.php The New Bromeliad Taxon List] A constantly updated list of current Bromeliad names and synonyms.
* Luther, H. E. (2008) [http://selby.org/wp-content/uploads/Bromeliad_Binomial_List_For_Web.pdf An Alphabetical List of Bromeliad Binomials, Eleventh Edition] [http://www.selby.org The Marie Selby Botanical Gardens], [[Sarasota, Florida]]. Published by [http://www.bsi.org/ The Bromeliad Society International].
* [http://www.bromeliads.info/ Bromeliad care information]
* [https://registry.bsi.org/ Bromeliad cultivar registry]
* [http://www.chileflora.com/Florachilena/FloraEnglish/PIC_FAMILIES_SIMPLE_Bromeliaceae.php Bromeliads of Chile] in Chileflora
* [https://web.archive.org/web/20161203151338/http://palm-trees.org/ Palm trees, small palms, Cycads, Bromeliads and tropical plants] Photos of Bromeliads and associated flora, with information on habitat and cultivation.

{{Angiosperm families}}
{{Taxonbar|from=Q156529}}
{{Authority control}}

[[Category:Bromeliaceae| ]]
[[Category:Poales families]]