Editing Bromeliaceae (section)

==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" />