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The Bromeliaceae
Bromeliaceae
(the bromeliads) are a family of monocot flowering plants of 51 genera and around 3475 known species[2] native mainly to the tropical Americas, with a few species found in the American subtropics and one in tropical west Africa, Pitcairnia
Pitcairnia
feliciana.[3] They are among the basal families within the Poales
Poales
and are the only family within the order that has septal nectaries and inferior ovaries.[4] These inferior ovaries characterize the Bromelioideae, a subfamily of the Bromeliaceae.[5] The family includes both epiphytes, such as Spanish moss
Spanish moss
( Tillandsia
Tillandsia
usneoides), and terrestrial species, such as the pineapple ( Ananas
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 epiphyte Tillandsia
Tillandsia
species that gather water only from leaf structures called trichomes, and a large number of desert-dwelling succulents. The largest bromeliad is Puya raimondii, which reaches 3–4 m tall in vegetative growth with a flower spike 9–10 m tall, and the smallest is Spanish moss.[citation needed]

Contents

1 Description 2 Distribution 3 Ecology 4 Evolution

4.1 Adaptations

5 Classification

5.1 Subfamilies 5.2 Genera

6 Gallery 7 Cultivation and uses

7.1 Collectors

8 See also 9 References 10 External links

Description[edit]

Bromeliad

Bromeliads are plants that are adapted to various climates. Foliage takes different shapes, from needle-thin to broad and flat, symmetrical to irregular, spiky to soft. The foliage, which usually grows in a rosette, is widely patterned and colored. Leaf
Leaf
colors range from maroon, through shades of green, to gold. Varieties may have leaves with red, yellow, white and cream variations. Others may be spotted with purple, red, or cream, while others have different colors on the tops and bottoms of the leaves. The inflorescences produced by bromeliads are also regarded as considerably more diverse than any other plant family. Some flower spikes may reach 10 meters tall, while others only measure 2–3 mm across. Upright stalks may be branched or simple with spikes retaining their color from two weeks up to 12 months, depending on species. In some species, the flower remains unseen, growing deep in the base of the plants. Root systems vary according to plant type. Terrestrial bromeliad species have complex root systems that gather water and nutrients, while epiphytic bromeliads only grow hard, wiry roots to attach themselves to trees and rocks.

An epiphytic bromeliad

Bromeliad at US Botanic Garden

Some bromeliads are faintly scented, while others are heavily perfumed. Blooms from the species Tillandsia
Tillandsia
cyanea have a fragrance resembling that of clove spice. 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.[6] A wide variety of organisms takes advantage of the pools of water trapped by bromeliads. A study of 209 plants from the Ecuadorian lowlands identified 11,219 animals, representing more than 300 distinct species, many of which are found only on bromeliads. Examples include some species of ostracods, small salamanders about 2.5 cm (1 in) in length, and tree frogs. Jamaican bromeliads are home to Metopaulias
Metopaulias
depressus, a reddish-brown crab 2 cm (0.8 in) across, which has evolved social behavior to protect its young from predation by Diceratobasis macrogaster, a species of damselfly whose larvae live in bromeliads. Some bromeliads even form homes for other species of bromeliads.[6] Distribution[edit]

Bromeliads growing on telephone lines in Bolivia

Plants in the Bromeliaceae
Bromeliaceae
are widely represented in their natural climates across the Americas. One species can be found in Africa.[7] They can be found at altitudes from sea level to 4200 meters, from rainforests to deserts. 1814 species are epiphytes, some are lithophytes, and some are terrestrial. Accordingly, these plants can be found in the Andean
Andean
highlands, from northern Chile to Colombia, in the Sechura Desert
Desert
of coastal Peru, in the cloud forests of Central and South America, in southern United States from southern Virginia
Virginia
to Florida
Florida
to Texas, and in far southern Arizona. Ecology[edit] Bromeliads often serve as phytotelmata, accumulating water between their leaves. The aquatic habitat created as a result is host to a diverse array of invertebrates, especially aquatic insect larvae.[8] These bromeliad invertebrates benefit their hosts by increasing nitrogen uptake into the plant.[9][10][11] Evolution[edit] Bromeliads are among the more recent plant groups to have emerged. The greatest number of primitive species resides in the Andean
Andean
highlands of South America, where they originated in the tepuis of the Guyana Shield.[12] The most basal genus, Brocchinia, is endemic to these tepuis, and is placed as the sister group to the remaining genera in the family.[13] The west African species Pitcairnia feliciana is the only bromeliad not endemic to the Americas, and is thought to have reached Africa via long-distance dispersal about 12 million years ago.[12] Adaptations[edit] Bromeliads are able to live in a vast array of environmental conditions due to their many adaptations. Trichomes, in the form of scales or hairs, allow bromeliads to capture water in cloud forests and help to reflect sunlight in desert environments.[14] Some bromeliads have also developed an adaptation known as the tank habit, which involves them forming a tightly bound structure with their leaves that helps to capture water and nutrients in the absence of a well-developed root system.[14] Bromeliads also use crassulacean acid metabolism (CAM) photosynthesis to create sugars. This adaptation allows bromeliads in hot or dry climates to open their stomates at night rather than during the day, which reduces water loss.[15] Classification[edit] The family Bromeliaceae
Bromeliaceae
is currently placed in the order Poales. Subfamilies[edit] The family Bromeliaceae
Bromeliaceae
is organized into eight subfamilies:[16]

Brocchinioideae Lindmanioideae Tillandsioideae Hechtioideae Navioideae Pitcairnioideae Puyoideae Bromelioideae

Bromeliaceae
Bromeliaceae
were originally split into three subfamilies: Bromelioideae, Tillandsioideae, and Pitcairnioideae
Pitcairnioideae
based on morphological characters.[17] However, molecular evidence has revealed that while Bromelioideae
Bromelioideae
and Tillandsioideae
Tillandsioideae
are monophyletic, Pitcairnioideae
Pitcairnioideae
is, in fact, paraphyletic[18] and should be split into six subfamilies: Brocchinioideae, Lindamanioideae, Hechtioideae, Navioideae, Pitcairnioideae, and Puyoideae.[19] Brocchinioideae is defined as the most basal branch of Bromeliaceae based on both morphological and molecular evidence, namely genes in chloroplast DNA.[20] Lindmanioideae is the next most basal branch distinguished from the other subfamilies by convolute sepals and chloroplast DNA.[21] Hechtioideae is also defined based on analyses of chloroplast DNA; similar morphological adaptations to arid environments also found in other groups are attributed to convergent evolution.[16] Navioideae is split from Pitcairnioideae
Pitcairnioideae
based on its cochlear sepals and chloroplast DNA.[22] Puyoideae has been re-classified multiple times and its monophyly remains controversial according to analyses of chloroplast DNA.[21] Genera[edit]

Acanthostachys
Acanthostachys
Klotzsch Aechmea
Aechmea
Ruiz & Pav. Alcantarea
Alcantarea
Harms Ananas
Ananas
Mill. — Includes the pineapple. Androlepis
Androlepis
Brongn.
Brongn.
ex Houllet Araeococcus
Araeococcus
Brongn. Barfussia Manzan. & W. Till Billbergia
Billbergia
Thunb. Brewcaria L.B.Sm., Steyerm. & H.Rob Brocchinia
Brocchinia
Schult.f. Bromelia
Bromelia
L. Canistropsis Canistrum
Canistrum
E.Morren Catopsis
Catopsis
Griseb. Cipuropsis
Cipuropsis
Ule Connellia N.E.Br. Cottendorfia Schult.f.

Cryptanthus
Cryptanthus
Otto & A.Dietr. Deinacanthon Deuterocohnia
Deuterocohnia
Mez Disteganthus
Disteganthus
Lem. Dyckia
Dyckia
Schult.f. Edmundoa Eduandrea Encholirium
Encholirium
Mart.
Mart.
ex Schult.f. Fascicularia
Fascicularia
Mez Fernseea Baker Fosterella
Fosterella
L.B.Sm. Glomeropitcairnia
Glomeropitcairnia
Mez Goudaea W. Till & Barfuss Greigia
Greigia
Regel Gregbrownia W. Till & Barfuss Guzmania
Guzmania
Ruiz & Pav. Hechtia
Hechtia
Klotzsch

Hohenbergia
Hohenbergia
Schult.f. Hohenbergiopsis
Hohenbergiopsis
L.B.Sm. & Read Jagrantia Barfuss & W. Till Josemania W. Till & Barfuss Lemeltonia Barfuss & W. Till Lindmania Mez Lutheria Barfuss & W. Till Lymania
Lymania
Read Mezobromelia
Mezobromelia
L.B.Sm. Navia Schult.f. Neoglaziovia
Neoglaziovia
Mez Neoregelia
Neoregelia
L.B.Sm. Nidularium
Nidularium
Lem. Ochagavia
Ochagavia
Phil. Orthophytum Pepinia Pitcairnia
Pitcairnia
L'Her. Portea
Portea
K. Koch Pseudaechmea L.B.Sm. & Read

Pseudalcantarea (Mez) Pinzón & Barfuss Pseudananas
Pseudananas
Hassl.
Hassl.
ex Harms Puya Molina Quesnelia
Quesnelia
Gaudich. Racinaea Ronnbergia E.Morren & André Sequencia Steyerbromelia L.B.Sm. Stigmadoton Leme, G.K.Br. & Barfuss Tillandsia
Tillandsia
L. Ursulaea Vriesea
Vriesea
Lindl. Wallisia E. Morren Werauhia
Werauhia
J.R.Grant Wittrockia
Wittrockia
Lindm. Zizkaea W. Till & Barfuss

Gallery[edit]

Billbergia
Billbergia
pyramidalis

Billbergia
Billbergia
pyramidalis

Tillandsia usneoides
Tillandsia usneoides
hanging from branches

The flower of a Billbergia
Billbergia
sp.

Puya alpestris

Flower close-up

A bromeliad

Tillandsia
Tillandsia
airplants mounted on the bark of a cork oak

Cultivation and uses[edit] Humans have been using bromeliads for thousands of years. The Incas, Aztecs, 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
Aechmea
fasciata was brought to Europe, followed by Vriesea
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 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, Dutch, Belgian and North American nurseries have greatly expanded bromeliad production. Only one bromeliad, the pineapple ( Ananas
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 plants, grown as both garden and houseplants. Collectors[edit] Édouard André
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
Mulford B. Foster
and Lyman Smith of the United States and Werner Rauh of Germany and Michelle Sullivan of Australia.[23] See also[edit]

List of foliage plant diseases (Bromeliaceae)

References[edit]

^ Angiosperm Phylogeny Group (2009), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III", Botanical Journal of the Linnean Society, 161 (2): 105–121, doi:10.1111/j.1095-8339.2009.00996.x, retrieved 2010-12-10  ^ Christenhusz, M. J. M. & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. Magnolia Press. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.  ^ Mabberley, D.J. (1997). The Plant
Plant
Book. Cambridge: Cambridge University Press.  ^ Judd, Walter S. Plant
Plant
systematics a phylogenetic approach. 3rd ed. Sunderland, MA: Sinauer Associates, Inc., 2007. ^ Sajo, M. G. (2004). "Floral anatomy of Bromeliaceae, with particular reference to the epigyny and septal nectaries in commelinid monocots". Plant
Plant
Systematics and Evolution. 247: 215–31. doi:10.1007/s00606-002-0143-0.  ^ a b " Pineapple
Pineapple
Dreams", The Wild Side, Olivia Judson, The New York Times, March 18, 2008 ^ Porembski, Stefan; Barthlott, Wilhelm (1999). "PITCAIRNIA FELICIANA: THE ONLY INDIGENOUS AFRICAN BROMELIAD". Harvard Papers in Botany. 4 (1): 175–184. JSTOR 41761298.  ^ Frank, J. H.; Lounibos, L. P. (2009-02-01). "Insects and allies associated with bromeliads: a review". Terrestrial Arthropod Reviews. 1 (2): 125–153. doi:10.1163/187498308X414742. ISSN 1874-9836. PMC 2832612 . PMID 20209047.  ^ Ngai, Jacqueline T.; Srivastava, Diane S. (2006-11-10). "Predators Accelerate Nutrient Cycling in a Bromeliad Ecosystem". Science. 314 (5801): 963–963. doi:10.1126/science.1132598. ISSN 0036-8075. PMID 17095695.  ^ Leroy, Céline; Corbara, Bruno; Dejean, Alain; Céréghino, Régis (2009-09-01). "Ants mediate foliar structure and nitrogen acquisition in a tank-bromeliad". New Phytologist. 183 (4): 1124–1133. doi:10.1111/j.1469-8137.2009.02891.x. ISSN 1469-8137. PMID 19500265.  ^ Romero, Gustavo Q.; Srivastava, Diane S. (2010-09-01). "Food-web composition affects cross-ecosystem interactions and subsidies". Journal of Animal Ecology. 79 (5): 1122–1131. doi:10.1111/j.1365-2656.2010.01716.x. ISSN 1365-2656.  ^ a b Givnish, Thomas J.; Millam, Kendra C.; Evans, Timothy M.; Hall, Jocelyn C.; Pires, J. C.; Berry, Paul E.; Sytsma, Kenneth J. (2004). "Ancient vicariance or recent long-distance dispersal? Inferences about phylogeny and South American-African disjunctions in Raptaceae and Bromeliaceae
Bromeliaceae
based on ndhf sequence data". International Journal of Plant
Plant
Sciences. 165 (4): 35–54.  ^ Barfuss, Michael H.; Samuel, Rosabelle; Till, Walter; Stuessy, Todd F. (2005). "Phylogenetic relationships in subfamily Tillandsioideae (Bromeliaceae) based on DNA sequence data from seven plastid regions". American Journal of Botany. 92 (2): 337–51. doi:10.3732/ajb.92.2.337.  ^ a b Schulte, Katharina; Barfuss, Michael H.; Zizka, Georg (2009). "Phylogeny of Bromelioideae
Bromelioideae
(Bromeliaceae) inferred from nuclear plastid DNA loci reveals the evolution of the tank habit within the subfamily". Molecular Phylogenetics and Evolution. 51: 327–39. doi:10.1016/j.ympev.2009.02.003.  ^ Rex, Martina; Patzolt, Kerstin; Schulte, Katharina; Zizka, Georg; Vasquuez, Roberto; Ibisch, Pierre L.; Weising, Kurt (2007). "AFLP analysis of genetic relationships in the genus Fosterella
Fosterella
L.B. Smith (Pitcairnioideae, Bromeliaceae)". Genome. 50: 90–105. doi:10.1139/g06-141.  ^ a b Givnish, Thomas (2007). "Phylogeny, adaptive radiation, and historical biogeography of Bromeliaceae
Bromeliaceae
inferred from ndhF sequence data". Aliso: A Journal of Systematic and Evolutionary Botany.  ^ Smith LB, Downs RJ (1974). "Flora neotropica: monograph. 14.(Bromeliaceae)". New York Botanical Garden.  ^ Terry, Randall (1997). "Examination of subfamilial phylogeny in Bromeliaceae
Bromeliaceae
using comparative sequencing of the plastid locus ndhF". American Journal of Botany. 84: 664. doi:10.2307/2445903.  ^ Zanella, Camila (2012). "Genetics, evolution and conservation of Bromeliaceae". Genetics and Molecular Biology. 35: 1020–1026. doi:10.1590/s1415-47572012000600017.  ^ Horres, Ralf (2000). "Molecular phylogenetics of Bromeliaceae: evidence from trnL (UAA) intron sequences of the chloroplast genome". Plant
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Biology. 2: 306–315. doi:10.1055/s-2000-3700.  ^ a b Givnish, Thomas (2011). "Phylogeny, adaptive radiation, and historical biogeography in Bromeliaceae: insights from an eight-locus plastid phylogeny". American Journal of Botany. 98: 872–895. doi:10.3732/ajb.1000059.  ^ Crayn, Darren (2004). "Multiple origins of crassulacean acid metabolism and the epiphytic habit in the Neotropical family Bromeliaceae". Proceedings of the National Academy of Sciences. 101: 3703–3708. doi:10.1073/pnas.0400366101.  ^ André, Édouard François. " Bromeliaceae
Bromeliaceae
Andreanae. Description et histoire des Bromeliacees recoltees dans La Colombie, L' Ecuador
Ecuador
et Le Venezuela". Paris: Librairie Agricole; G. Masson, 1889

External links[edit]

Wikimedia Commons has media related to Bromeliaceae.

Wikispecies
Wikispecies
has information related to Bromeliaceae

Bromeliad care information Puya raimondii
Puya raimondii
photos The World Botanical Gardens Bromeliads of Chile in Chileflora The Brom-L Bromeliad Gallery The Photo Gallery of the (Virtual) World Wide Web Bromeliad Society The New Bromeliad Taxon List A constantly updated list of current Bromeliad names and synonyms. Palm trees, small palms, Cycads, Bromeliads and tropical plants Photos of Bromeliads and associated flora, with information on habitat and cultivation. Luther, H. E. (2008) An Alphabetical List of Bromeliad Binomials, Eleventh Edition The Marie Selby Botanical Gardens, Sarasota, Florida. Published by The Bromeliad Society International. Bromeliaceae
Bromeliaceae
in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval. Published by Delta-intkey (2002-06-18)

Taxon identifiers

Wd: Q156529 EoL: 8198 EPPO: 1BROF FNA: 10123 FoC: 10123 Fossilworks: 55818 GBIF: 3740 GRIN: 169 IPNI: 30000083-2 ITIS: 42330 NCBI: 4613 Tropicos: 42000361 Watson &

.