Bromeliaceae

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Bromeliaceae
Temporal range: 100–0  Ma
Pineapple1.JPG
Pineapple (Ananas comosus), a bromeliad of economic importance
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Poales
Family: Bromeliaceae
Juss. [1]
Subfamilies
WorldBromeliadDistribution.PNG

The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, [2] native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, Pitcairnia feliciana . [3]

Contents

It is among the basal families within the Poales and is 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 ( Tillandsia usneoides ), and 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 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 3–4 metres (9.8–13.1 ft) tall in vegetative growth with a flower spike 9–10 metres (30–33 ft) tall, [6] [7] and the smallest is Spanish moss.[ citation needed ]

Description

Bromeliad Bromeliad3.jpg
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). [8] Some species of Tillandsia (e.g. Spanish moss, Tillandsia usneoides) are aerophytes, which have very reduced root systems and absorb water directly from the air. [9] 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, Chironomia fly larva, an ant colony, an earthworm, numerous mites, and a small frog. [10] 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. [8]

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. [11] Bromeliads with leaf vases can capture water and nutrients in the absence of a well-developed root system. [11] Many bromeliads also use crassulacean acid metabolism (CAM) photosynthesis to create sugars. This adaptation allows bromeliads in hot or dry climates to open their stomata at night rather than during the day, which reduces water loss. [12] 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. [13]

Evolution

Bromeliads are among the more recent plant groups to have emerged. They are thought to have originated in the tepuis of the Guiana Shield approximately 100 million years ago. The greatest number of extant basal species are found in the Andean highlands of South America. [14] 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. [15]

Based on molecular phylogenetic studies, the family is divided into eight subfamilies. The relationship among them is shown in the following cladogram. [14]

Bromeliaceae

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 sister group to the remaining genera in the family. [15] The subfamilies Lindmanioideae and Navioideae are endemic to the Guiana Shield as well. [16]

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. [14]

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 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 hummingbirds. [17] [13] [18] [19]

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. [13] 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. [13]

Classification

The family Bromeliaceae is currently placed in the order Poales.

Subfamilies

The family Bromeliaceae is organized into eight subfamilies: [17]

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). [20] However, molecular evidence has revealed that while Bromelioideae and Tillandsioideae are monophyletic, Pitcairnioideae as traditionally defined is paraphyletic [21] and should be split into six subfamilies: Brocchinioideae, Lindmanioideae, Hechtioideae, Navioideae, Pitcairnioideae, and Puyoideae. [22]

Brocchinioideae is defined as the most basal branch of Bromeliaceae based on both morphological and molecular evidence, namely genes in chloroplast DNA. [23]

Lindmanioideae is the next most basal branch distinguished from the other subfamilies by convolute sepals and chloroplast DNA. [13]

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. [17]

Navioideae is split from Pitcairnioideae based on its cochlear sepals and chloroplast DNA. [24]

Puyoideae has been re-classified multiple times and its monophyly remains controversial according to analyses of chloroplast DNA. [13]

Genera

As of December 2022, Plants of the World Online (PoWO) accepted 72 genera, as listed below. [25] A few more genera were accepted by the Encyclopaedia of Bromeliads, including Josemania and Mezobromelia, which PoWO sinks into Cipuropsis.

Hybrid genera

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

Distribution and habitat

Bromeliads growing on telephone lines in Bolivia Tillandsia sp. telephone line (codiferous).jpg
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. [30] 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 sequester 50,000 liters (more than 13,000 gallons) of water. [31] The aquatic habitat created as a result is host to a diverse array of invertebrates, especially aquatic insect larvae, [32] [33] including those of mosquitos. [34] These bromeliad invertebrates benefit their hosts by increasing nitrogen uptake into the plant. [35] [36] [37] A study of 209 plants from the Yasuní Scientific Reserve in Ecuador identified 11,219 animals, representing more than 350 distinct species, [38] 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 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. [31]

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. [39] [40]

Cultivation and uses

Bromeliaceae mixed cultivated collection Bromeliaceae mixed collection.jpg
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. [41]

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 Smith of the United States and Werner Rauh of Germany and Michelle Jenkins of Australia. [42]

See also

Related Research Articles

<i>Tillandsia</i> Genus of flowering plants

Tillandsia is a genus of around 650 species of evergreen, perennial flowering plants in the family Bromeliaceae, native to the forests, mountains and deserts of the Neotropics, from northern Mexico and the southeastern United States to Mesoamerica and the Caribbean to central Argentina. Their leaves, more or less silvery in color, are covered with specialized cells (trichomes) capable of rapidly absorbing water that gathers on them.

<i>Brocchinia</i> Genus of carnivorous plants

Brocchinia is a genus of the botanical family Bromeliaceae, and is the sole genus of the subfamily Brocchinioideae, containing 20 species. The genus is named for Giovanni Battista Brocchi, Italian naturalist (1772–1826). Brocchinia species are native primarily to the ancient Guayana Shield in southern Venezuela and Guyana, with some species extending into Colombia and northern Brazil. Its species are generally restricted to areas of sand and sandstone of the Roraima Formation; a few occur on granite.

<i>Neoregelia</i> Genus of flowering plants

Neoregelia is a genus of epiphytic flowering plants in the family Bromeliaceae, subfamily Bromelioideae, native to South American rainforests. The genus name is for Eduard August von Regel, Director of St. Petersburg Botanic Gardens in Russia (1875–1892).

<i>Vriesea</i> Genus of epiphytes

Vriesea is a genus of flowering plants in the botanical family Bromeliaceae, subfamily Tillandsioideae. The genus name is for Willem Hendrik de Vriese, Dutch botanist, physician (1806–1862). Its species are widespread over Mexico, Central America, South America and the West Indies.

<i>Aechmea</i> Genus of flowering plants

Aechmea is a genus of flowering plants in the family Bromeliaceae. The name comes from the Greek aichme, meaning "spear". Suggested pronunciations include EEK-me-ə and eek-MEE-ə. Aechmea comprises eight subgenera and around 250 species distributed from Mexico through South America and the Caribbean. Most of the species in this genus are epiphytes.

<i>Cryptanthus</i> Genus of flowering plants

Cryptanthus is a genus of flowering plants in the family Bromeliaceae, subfamily Bromelioideae. The genus name is from the Greek cryptos (hidden) and anthos (flower). The genus formerly had two recognized subgenera: the type subgenus and subgenus HoplocryptanthusMez which has been raised to the separate genus Hoplocryptanthus. All species of this genus are endemic to Brazil. The common name for any Cryptanthus is "Earth star".

<span class="mw-page-title-main">Bromelioideae</span> Subfamily of bromeliad flowering plants

Bromelioideae is a subfamily of the bromeliads (Bromeliaceae). This subfamily is the most diverse in the family, represented by the greatest number of genera with about 40. Most of the plants in this group are epiphytes, though some have evolved in, or will adapt to, terrestrial conditions. This subfamily features the most plant types which are commonly cultivated by people, including the pineapple.

<span class="mw-page-title-main">Pitcairnioideae</span> Species of flowering plant

Pitcairnioideae is a subfamily of the bromeliad family, Bromeliaceae. Traditionally, it was a large subfamily, comprising all those species with winged or more rarely naked seeds. Molecular phylogenetic studies showed that traditional Pitcairnioideae was not monophyletic, and the subfamily was more narrowly circumscribed. As of November 2022, the Encyclopaedia of Bromeliads placed five genera in the subfamily. Members of the subfamily are found from the Andes to the coast of Brazil, with one genus (Fosterella) found northwards to Mexico.

<span class="mw-page-title-main">Tillandsioideae</span> Subfamily of family Bromeliaceae

Tillandsioideae is a subfamily of plants in the bromeliad family Bromeliaceae. This subfamily contains the greatest number of species. Most are epiphytic or lithophytic, growing in trees or on rocks where they absorb water and nutrients from the air. Spanish moss of the genus Tillandsia is a well-known species. Bromeliads in the genera Guzmania and Vriesea are the more commonly cultivated members of this subfamily.

<i>Pitcairnia</i> Species of flowering plant

Pitcairnia is a genus of plants in the family Bromeliaceae, subfamily Pitcairnioideae. It was named for William Pitcairn, Scottish physician and gardener (1711–1791). The genus Pitcairnia ranks as the second most prolific of the bromeliad family. They are most abundant in Colombia, Peru and Brazil, but can also be found in areas from Cuba and Mexico south to Argentina. One species, Pitcairnia feliciana, is found in tropical West Africa and is the only member of the family Bromeliaceae not native to the Americas.

<i>Orthophytum</i> Genus of flowering plants

Orthophytum is a genus in the plant family Bromeliaceae, subfamily Bromelioideae.

<i>Encholirium</i> Genus of flowering plants

Encholirium is a genus of plants in the family Bromeliaceae, subfamily Pitcairnioideae. The entire genus is endemic to Brazil. The genus name is from the Greek “enchos” (spear) and “leiron” (lily).

<i>Racinaea</i> Genus of flowering plants

Racinaea is a genus of the flowering plants in the family Bromeliaceae, subfamily Tillandsioideae. The genus is named for Racine Foster, wife of Mulford B. Foster and co-founder of the BSI. It is found in tropical regions of the Americas.

<i>Pitcairnia <span style="font-style:normal;">subg.</span> Pepinia</i> Genus of flowering plants

Pitcairnia subg. Pepinia is a subgenus of plants in the family Bromeliaceae. It has at times been treated as the separate genus Pepinia, but is now included again in the genus Pitcairnia. The name is for Pierre Denis Pépin, French member of the Imperial and Central Society of Agriculture (c.1802-1876).

<i>Hylaeaicum</i> Genus of flowering plants

Hylaeaicum is a genus of flowering plant in the family Bromeliaceae, native to tropical northern South America. The taxon was first described by Ernst Heinrich Georg Ule in 1935 as a subgenus of "Aregelia". It was later treated as a subgenus of Neoregelia, before being raised to a full genus in 2021, a status accepted by both Plants of the World Online and the Encyclopaedia of Bromeliads.

<span class="mw-page-title-main">Navioideae</span> Subfamily of plants

Navioideae is a subfamily of the bromeliad family, Bromeliaceae. It contains four or five genera, formerly placed in a more broadly defined subfamily Pitcairnioideae.

<i>Tillandsia <span style="font-style:normal;">subg.</span> Tillandsia</i> Subgenus of flowering plants

Tillandsia subg. Tillandsia is a subgenus of the genus Tillandsia.

Josemania is a genus of flowering plant in the family Bromeliaceae, first described in 2016.

<i>Rokautskyia</i> Genus of plants

Rokautskyia is a genus of flowering plant in the family Bromeliaceae, native to eastern Brazil. The genus was first established in 2017, and is placed in subfamily Bromelioideae.

References

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  36. ↑ 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.
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  39. ↑ 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.
  40. ↑ 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.
  41. ↑ Hornung-Leoni (2011). "Bromeliads: Traditional Plant Food in Latin America Since pre-Hispanic Times". Polibotánica. 32: 219–229. Retrieved 30 Mar 2020.
  42. ↑ 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
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