Bromeliaceae - WikiHQ

Bromeliaceae, members of which are called bromeliads (), is a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, Pitcairnia feliciana.

It is the sister group to the Typhaceae, and together this clade is sister to the remaining lineages in the Poales. Bromeliaceae is the only family within the order that has septal nectaries and inferior ovaries. These inferior ovaries characterize the Bromelioideae, a subfamily of the Bromeliaceae. The family includes both epiphytes, such as Spanish moss (Tillandsia usneoides), and terrestrial species, such as the pineapple (Ananas comosus). Many bromeliads, colloquially called "tank bromeliads", are able to store water in a structure (a "tank") formed by their tightly overlapping leaf bases. However, the family is diverse enough to include the tank bromeliads, grey-leaved epiphyte Tillandsia species that gather water only from leaf structures called trichomes, and many desert-dwelling succulents.

The largest bromeliad is Puya raimondii, which reaches tall in vegetative growth with a flower spike tall, and the smallest are some Tillandsia species.

Description

thumb|[[Pitcairnia bifrons, a terrestrial bromeliad]]

Bromeliads are mostly herbaceous perennials, although a few have a more tree-like habit. Many are more or less succulent or have other adaptations to resist drought. They may be terrestrial or epiphytic, rarely climbing (e.g. Pitcairnia species). Many terrestrial and epiphytic bromeliads have their leaves in the form of vase-shaped rosettes which accumulate water. These rosettes can hold as much as of water, and be little biotic communities unto themselves. One individual tank was found to contain four harvestmen, a spider, three species of wood lice, a centipede, a millipede, a pseudoscorpion, various metallic beetles, earwigs, a tree seedling, chironomid fly larva, an ant colony, an earthworm, numerous mites, and a small frog. Individual leaves are not divided and have parallel veins without cross connections. The epidermis of the leaf contains silica. Bromeliad flowers are aggregated into inflorescences of various forms. The flowers have bracts, often brightly coloured, and distinct calyces of three sepals and corollas of three petals. The flowers have nectaries. They are pollinated by insects, birds (often hummingbirds) or bats, or more rarely (in Navia) they are wind-pollinated. Fruits are variable, typically taking the form of a capsule or a berry.

Bromeliads are able to live in an 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. Bromeliads with leaf vases can capture water and nutrients in the absence of a well-developed root system. Both CAM and epiphytism have evolved multiple times within the family, with some taxa reverting to C3 photosynthesis as they radiated into less arid climates. 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. The subfamilies Lindmanioideae and Navioideae are endemic to the Guiana Shield as well.

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.

Evolution of the Bromelioideae

Around 5.5 million years ago, a clade of epiphytic 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.

Brocchinioideae
Lindmanioideae
Tillandsioideae
Hechtioideae
Navioideae
Pitcairnioideae
Puyoideae
Bromelioideae

Bromeliaceae were originally split into three subfamilies based on morphological seed characters: Bromelioideae (seeds in baccate fruits), Tillandsioideae (plumose seeds), and Pitcairnioideae (seeds with wing-like appendages). However, molecular evidence has revealed that while Bromelioideae and Tillandsioideae are monophyletic, Pitcairnioideae as traditionally defined is paraphyletic and should be split into six subfamilies: Brocchinioideae, Lindmanioideae, Hechtioideae, Navioideae, Pitcairnioideae, and Puyoideae.

Brocchinioideae is defined as the most basal branch of Bromeliaceae based on both morphological and molecular evidence, namely genes in chloroplast DNA.

Lindmanioideae is the next most basal branch distinguished from the other subfamilies by convolute sepals and chloroplast DNA.

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) are attributed to convergent evolution.

Puyoideae has been re-classified multiple times and its monophyly remains controversial according to analyses of chloroplast DNA. A few more genera were accepted by the Encyclopaedia of Bromeliads, including Josemania and Mezobromelia, which PoWO sinks into Cipuropsis.

Acanthostachys Klotzsch
Aechmea Ruiz & Pav.
Alcantarea Harms
Ananas Mill., including Pseudananas Hassl. ex Harms (includes the pineapple)
Androlepis Brongn. ex Houllet
Araeococcus Brongn.
Bakerantha L.B.Sm.
Barfussia Manzan. & W.Till
Billbergia Thunb.
Brewcaria L.B.Sm., Steyerm. & H.Rob, synonym of Navia in PoWO
Brocchinia Schult.f.
Bromelia L.
Canistropsis (Mez) Leme
Canistrum E.Morren
Catopsis Griseb.
Cipuropsis Ule
Connellia N.E.Br.
Cottendorfia Schult.f.
Cryptanthus Otto & A.Dietr.
Deinacanthon Mez
Deuterocohnia Mez
Disteganthus Lem.
Dyckia Schult.f.
Edmundoa Leme
Eduandrea Leme, W.Till, G.K.Br., J.R.Grant & Govaerts
Encholirium Mart. ex Schult.f.
Fascicularia Mez
Fernseea Baker
Forzzaea Leme, S.Heller & Zizka
Fosterella L.B.Sm.
Glomeropitcairnia Mez
Goudaea W.Till & Barfuss
Gregbrownia W.Till & Barfuss
Greigia Regel
Guzmania Ruiz & Pav.
Hechtia Klotzsch
Hohenbergia Schult.f.
Hohenbergiopsis L.B.Sm. & Read
Hoplocryptanthus (Mez) Leme, S.Heller & Zizka
Hylaeaicum (Ule ex Mez) Leme, Forzza, Zizka & Aguirre-Santoro
Jagrantia Barfuss & W.Till
Josemania W.Till & Barfuss
Karawata J.R.Maciel & G.M.Sousa
Krenakanthus (Leme, S.Heller & Zizka) Leme, Zizka & Paule
Lapanthus Louzada & Versieux
Lemeltonia Barfuss & W.Till
Lindmania Mez
Lutheria Barfuss & W.Till
Lymania Read
Mezobromelia L.B.Sm.
Navia Schult.f.
Neoglaziovia Mez
Neoregelia L.B.Sm.
Nidularium Lem.
Ochagavia Phil.
Orthocryptanthus (Leme, S.Heller & Zizka) Leme, Zizka & Paule
Orthophytum Beer
Pitcairnia L'Her., including subgenus Pepinia
Portea K. Koch
Pseudaechmea L.B.Sm. & Read, synonym of Billbergia in PoWO
Pseudalcantarea (Mez) Pinzón & Barfuss
Pseudaraeococcus (Mez) R.A.Pontes & Versieux
Puya Molina
Quesnelia Gaudich.
Racinaea M.A.Spencer & L.B.Sm.
Rokautskyia Leme, S.Heller & Zizka
Ronnbergia E.Morren & André
Sequencia Givnish
Sincoraea Ule
Siqueiranthus Leme, Zizka, E.H.Souza & Paule
Steyerbromelia L.B.Sm.
Stigmatodon Leme, G.K.Br. & Barfuss
Tillandsia L.
Ursulaea Read & H.U.Baensch, synonym of Aechmea in PoWO
Vriesea Lindl.
Wallisia (Regel) É.Morren
Waltillia Leme, Barfuss & Halbritt.
Werauhia J.R.Grant
Wittmackia Mez
Wittrockia Lindm.
Zizkaea W.Till & Barfuss

Hybrid genera

Intergeneric hybrid genera accepted by Plants of the World Online include:

× Cryptbergia R.G.Wilson & C.L.Wilson = Cryptanthus × Billbergia
× Guzlandsia Gouda = Guzmania × Tillandsia
× Hohenmea B.R.Silva & L.F.Sousa = Hohenbergia × Aechmea
× Niduregelia Leme = Nidularium × Neoregelia

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:Starr 040518-0085 Tillandsia usneoides.jpg|Tillandsia usneoides hanging from branches

File:Puya chilensis 16.jpg|Puya chilensis

File:Tillandsia sp. telephone line (codiferous).jpg|Tillandsia growing on telephone lines in Bolivia

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 plantation of pineapple (Ananas comosus) plants in Bukidnon, the Philippines

</gallery>

Distribution and habitat

Plants in the Bromeliaceae are widely represented in their natural climates across the Americas. One species (Pitcairnia feliciana) can be found in Africa. They can be found at altitudes from sea level to 4,200 meters, from rainforests to deserts. 1,814 species are epiphytes, some are lithophytes, 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 forests 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 phytotelmata, accumulating water between their leaves. One study found 175,000 bromeliads per hectare (2.5 acres) in one forest; that many bromeliads can hold of water. The aquatic habitat created as a result is host to a diverse array of invertebrates, especially aquatic insect larvae, including those of mosquitos. These bromeliad invertebrates benefit their hosts by increasing nitrogen uptake into the plant. A study of 209 plants from the Yasuní Scientific Reserve in Ecuador identified 11,219 animals, representing more than 350 distinct species, many of which are found only on bromeliads. Examples include some species of ostracods, small salamanders about in length, and tree frogs. Jamaican bromeliads are home to Metopaulias depressus, a reddish-brown crab 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.

Cultivation and uses

thumb|Bromeliaceae mixed cultivated collection

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 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 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 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.

Bromeliads are important food plants for many peoples. For example, the Pima of Mexico occasionally consume flowers of Tillandsia erubescens and T. recurvata due to their high sugar content; in Argentina and Bolivia, the shoot apices of T. rubella and 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 beverages; in Chile, the sweet fruit of Greigia sphacelata, known as 'chupones', is consumed raw.

References

External links

The New Bromeliad Taxon List A constantly updated list of current Bromeliad names and synonyms.
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.
Bromeliad care information
Bromeliad cultivar registry
Bromeliads of Chile in Chileflora
Palm trees, small palms, Cycads, Bromeliads and tropical plants Photos of Bromeliads and associated flora, with information on habitat and cultivation.