Arecaceae

The arecaceae (family Arecaceae) are a family of monocotyledonous plants, the only family in the order Arecales.

They are commonly referred to as palm trees or palms. Individuals from this important family are easy to recognize visually, although there may be confusion with species from the Cycadaceae and Zamiaceae families due to morphological similarities. They are woody plants (but no secondary growth from the trunk, only primary). Despite being monocotyledonous, many of them are arborescent, with large crown leaves at the end of the stem, generally pinnate (pinnatisectal) or palmate (palmatisectal). Its flowers have 3 sepals and 3 petals, and are arranged in inflorescences with one or more spathes. The fruit is fleshy - a berry or a drupe. They are widely distributed in tropical to temperate regions, but mainly in warm regions.

The family was recognized by modern classification systems such as the APG III classification system (2009) and the APWeb (2001 onwards). Traditionally it was also recognized in other classification systems due to its distinctive morphological characters. In these classification systems, they are placed in their own monotypic order Arecales, in the subclass Commelinidae.

Among the palm trees there are species of economic importance and species of ornamental value, in addition to others such as coconut, oil palm, date palm, heart of palm, rattan, carnauba wax, raffia, among others.

In the world they grow as species typical of tropical zones, there are concentrations of them in countries like Madagascar. Colombia is the country with the largest number of varieties and one of them is the national tree. Additionally there are several botanical gardens specialized in palm trees and they are often called palmetum. Some of these collections include the Palmetum in Santa Cruz de Tenerife and the Palmeral in Elche, in Spain, the El Palmar National Park in Entre Ríos, Argentina, and to a lesser extent, the Caracas Botanical Garden in Venezuela, and the National Botanical Garden of Cuba in Havana, Cuba. There are also important collections of palm trees in the Molino de Inca Botanical Garden in Torremolinos (Málaga), the La Concepción Botanical Garden in Malaga and the Barcelona Botanical Garden.

Description

Oncosperma filamentosum.

Trees or shrubs with unbranched or rarely branching stems, occasionally long rhizomatous herbs, or non-liana climbing palms (eg Calamus). The sex of the plant is variable. Secondary growth is absent. Stem apex with a large apical meristem, leaves develop helically. Tannins and polyphenols often present. Variegated hairs, and plants sometimes spiny due to modified leaf segments, exposed fibers, pointed roots, or petiole growths.

The root system is always adventitious originating from the hypocotyl and lower nodes of the stem, since the primary seminal root is replaced at an early stage of seedling development, for which the root of the palms is fasciculated (fibrous), with abundant ramifications, generally short and very dense, forming a bulb at the base of the trunk, which provides mechanical support in addition to absorbing water and mineral elements. Palms can present mycorrhizal symbiosis.

Stem is usually arborescent with a single unbranched trunk (dichotomously branched in Hyphaene) or in a tufted clump of erect stems, or in an erect dichotomously branched Nypa rhizome, or in a slender stem elongated similar to that of bamboo, but supportive (rattans). In some species it reaches more than 30 m in height.

Typical leaves are quite large, alternate and spiral (rarely distichous or tristicate), often grouped in a terminal crown (acrocaulis), but sometimes well separated, entire, sheathing at the base, with an elongated, erect petiole (sometimes referred to as pseudopetiole) between sheathing base and lamina. In arborescent taxa the sheathing bases of adjacent leaves may overlap each other, forming a crownshaft at the apex of the trunk. The leaves may be simple, usually divided pinnately or palmately as the leaf expands, and at maturity appearing palmately lobed (with segments radiating from a single point), costapalmately lobate (with more or less palmate segments diverging from a short central axis, or "costa"), pinnately lobed or compound (with a well-developed central axis bearing pinnate segments), or rarely twice pinnately compound. Sometimes forked. With leaflets turned into spines present in some taxa. Lamina "plicated", and segments either induplicate (V-shaped in cross section) or reduplicated (Λ-shaped in cross section), each segment with more or less parallel to divergent veins.

Exemplary Livistona chinensis.

Determinate or indeterminate inflorescences, panicles or spikes of solitary flowers or cymose units, typically axillary or also terminal, with small to large persistent and deciduous bracts. The inflorescences emerge from below (infrafoliar) or between (interfoliar) or above (suprafoliar) the leaves. The peduncle has a often large profile below with 1 to numerous spathes.

Flowers bisexual or unisexual (and then monoecious to dioecious plants), radial, usually sessile, with perianth usually differentiated into calyx and corolla, hypogynous.

Sepals usually 3, separated to connate, usually imbricate.

Petals usually 3, usually separate to connate, imbricate to valvate.

Stamens 3 or 6 to numerous, filaments separate to connate, free or adnate to petals. Staminodia present in some species. Longitudinal anthers, rarely poricidal in dehiscence.

Pollen usually monosulcate.

Carpels usually 3, but occasionally as many as 10, sometimes appearing to be only one, connately separated. Ovary superior, usually with axillary placentation, but placentation variable. Styles, if present, separate or connate, stigmata sessile or at the tip of styles, stigma varied. Ovules 1 per locule, anatropous to orthotropic, bitegmic.

Nectaries in the ovary septa or without nectaries.

The fruit is a drupe, usually single-seeded, often fibrous, or rarely a berry. rarely dehiscent. Some have external scales (Calamoideae), hairs, stingers, or other protective structures.

Seeds usually one per fruit and with an endosperm rich in oils or carbohydrates (hemicelluloses), sometimes ruminated. Absent starch.

Ecology

Washingtonia filifera

Widely distributed in tropical and subtropical regions, mainly places with high humidity, with more than 2400 mm of average annual precipitation, more than 160 days with rain and more than 21 °C. Because of their abundance, they are often ecologically important where they occur. It also has representatives in temperate zones (for example Chamaerops), they survive in desert environments (Phoenix spp.), from tropical forests to mangroves (Nypa fruticans), and from sea level (Cocos nucifera) to very high altitudes (Trachycarpus).

There are more than 2,400 species worldwide, belonging to 27 tribes in five subfamilies. Around 790 species grow wild in the Neotropics (Dransfield et al. 2008). The most species-rich Neotropical regions are found in the Chocó region, where up to 83 species can be found in a grid of around 10,000 km², followed by the Mesoamerican Isthmus region (Panama and Costa Rica) (Bjorholm et al. 2005). Colombia alone has 289 species grouped into 66 genera, which is why it is considered the richest country in palms on the American continent, it also has the largest number of endemic species with a total of 33 species, equivalent to 15% of all palms. Of the territory; unfortunately Colombia is also the country with the largest number of threatened palms in America with 30 species that are located in some category of danger of which 17 are endemic). The most representative genera in Colombia are: Astrocaryum, Bactris, Chamaedorea, Desmoncus, Euterpe, Geonoma, Mauritia, Oenocarpus and Syagrus

The flowers of palm trees are usually pollinated by insects, especially beetles, bees, and flies. Many times the nectar is used as a pollination reward (Henderson 1986).

The fruits of palm trees are usually fleshy and dispersed by a wide variety of mammals and birds, although some (such as Nypha and Cocos) are dispersed by water and they float on ocean currents (Zona and Henderson 1989).

Phylogeny

A large body of work over a period of some 30 years has clarified our understanding of palms (see for example Dransfield 1986, Dransfield and Uhl 1998, Henderson 1995, Henderson et al. 1995, Moore 1973, Moore and Uhl 1982, Tomlinson 1990, Uhl and Dransfield 1987, Zona 1997). Dransfield et al. (2005) presents a clarification of the family based on molecular relationships (see especially Asmussen et al. 2006).

Arecaceae is easy to recognize and monophyletic. The palms are easily identified although there would be no consistent synapomorphies for the family. Uhl and Dransfield (1987) and Uhl et al. (1995) would have identified two main diagnostic characters: 1) "woody" (due to the presence of fibrous sclerenchyma, not secondary growth), and 2) leaves pleated at the buds and subsequent division in most clumps.

Calamoideae have palmately pinnate leaves and distinctive fruits that are covered with imbricate reflex scales (a synapomorphic character). Notable genera are Raphia, Mauritia, Lepidocaryum, Metroxylon, and Calamus.

Nypa (Nypoideae) has a prostrate stem that divides dichotomously, and erect, pinnate leaves, and undifferentiated tepals. Fossils are known in Europe and America early in the Tertiary.

Phylogenetic analyzes of multiple DNA sequences show that the subfamily Calamoideae is sister to all other palms. Nypa (the only genus of Nypoideae), a distinctive genus of mangrove communities in Asia and the western Pacific, would remain sister to all other palms (except Calamoideae). Then Nypa and Calamoideae form a paraphyletic complex, with usually pinnate and reduplicate leaves, while the rest of the genera, with usually costapalmate or palmate and induplicate leaves —the Coryphoideae— form a monophyletic group (Hahn 2002, Uhl et al. 1995).

Arecoideae have pinnate leaves and flowers in groups of 3 (triads), with one carpellate flower surrounded by two staminate flowers (probably a synapomorphy, but lost in some subgroups). Within the Arecoideae, a few well-defined monophyletic groups are evident.

Hyophorbeae (which has for example Chamaedorea, Hyophorbe), have imperfect flowers in lines.

Cocoseae has the inflorescence associated with a persistent, large, woody bract, and fruits with a triporate, bone-like endocarp, and includes genera such as Elaeis, Cocos, Syagrus, Attalea, Bactris, Desmoncus and Jubaea.

Iriarteae (having eg Iriartea, Socratea) have "stilt" roots, and leaf segments with blunt apices and diverging veins.

Most of the Arecoideae are located within a heterogeneous Areceae (Baker et al. 2006), representative genera include Areca, Dypsis, Wodyetia, Veitchia, Ptychosperma and Dictyosperma. These palms sometimes have a structure made up of a series of large or overlapping leaf bases, which looks like a vertical extension of the stem.

Coryphoideae are traditionally divided into the first 3 tribes below, a fourth one is included here:

• The Phoeniceae monogenerics (Phoenix, the datiles) have distinctive pine leaves den which the basal segments are like thorns.
• Borasseae (which contains for example a Latania, Borassus, Lodoicea, and Hyphaene) have staminated flowers embedded in axis of thickened inflorescence.
• Most Coryphoideae genres are located in a third tribe, the heterogeneous "Corypheae". Remarkable genders include Chamaerops, Rhapis, Licuala, Copernicia, Corypha, Washingtonia, Serenoa, Livistona, Rhapidophyllum, and Acoelorrapheand are difficult to characterize together. Genres like Sabal, Thrinaxand Coccothrinax they are phenothetically similar, and have been included here, but their inclusion would make the non-monophytic tribe.
• Caryoteae (fish tails) includes, for example, Caryota and Arenga, form a distinctive nail within Coryphoideae (Asmussen et al. 2000, Asmussen and Chase 2001, Hahn 2002) due to their flower triads (which evolved in parallel with those of Arecoideae). This group has unduplicated, pointed sheet segments with divergent veins.

Taxonomy

The family was recognized by the APG III (2009), the Linear APG III (2009) assigned it the family number 76. The family had already been recognized by the APG II (2003).

In the APG III system it is found in the monotypic order Arecales, subclass Commelinidae, class Monocotyledoneae. The APG II classification system, which does not use formal names above the order level, placed it within the comelinid clade, which had been maintained since the publication of the APG system in 1998 to the present.

Previous systems placed the group in the subclass Arecidae (Cronquist, 1981) or in the superorder Arecanae (Dahlgreen, Thorne).

The name Arecales, formed according to the rules of the International Code of Botanical Nomenclature from the type genus Areca (which includes the betel palm, Areca catechu) It is of relatively recent use. Traditional nomenclatures used the descriptive name Principales, from the Latin the first ones.

As an exception to the rule of priority, the old names used by Linnaeus can still be used for this family: Palmae for the family and Palmales for the order. The names palmáceas or Palmaceae are rejected by the International Code of Botanical Nomenclature. They are called palm trees in Spain, Peru, Uruguay, Argentina and Chile, and palms in most of the countries of America.

The extensive family has 200 genera, 2780 species. The most represented genera are Calamus (370 species), Bactris (200 species), Daemonorops (115 species), Licuala (100 species), and Chamaedorea (100 species).

Here are some genera of the family, with some of their species, their botanical authorship and their common names, as they are used in different Spanish-speaking countries.

• Aiphanes

Aiphanes caryotifolia (H.B.K.) J.C.Wendl. - Mararay (Venezuela, Ecuador and Colombia)

• Areca
• Arenga
• Astrocaryum

Astrocaryum aculeatum G.Mey. - cumare (Venezuela and Colombia)

Astrocaryum standleyanum - Guerregue, Wérregue, Chunga (Chocó).

• Attalea
• Bactris

Bactris gasipaes Kunth - chontaduro (Peru, Colombia and Venezuela)

Bactris brongniartii - Cubarro, Palma de Cubarro, Albarico.

Bactris guineensis (L.) H.E.Moore

• Borassus
• Ceroxylon
• Nuciferous coconuts L. - Coconut.
• Copernicia

Copernicia tectorum-Sará, Palmiche (Colombia and Venezuela)

• Chamaedorea

Chamaedorea pauciflora -Iakake (Miraña)

• Chamaerops

Chamaerops humilis L.: Palmito, Margalló (Spain, Spanish and Catalan)

• Desmoncus

Desmoncus polyacanthos -Matamba, Bejuco Mayor (Venezuela and Colombia)

Desmoncus mitis-Atajadanta (Amazonas), Bejuco mayor

• Elaeis

Elaeis guineensis Jacq. - Oil palm, Oil palm, African palm

Elaeis oleifera (H.B.K.) Courts - Nolí (Venezuela and Colombia)

• Euterpe

Euterpe oleracea - Murrapo, Palmito, Naidí, Palmiche, Manaca (Colombia and Venezuela)

Precatory euterpe - Asaí, Palmiche, Palma Manaca (Colombia and Venezuela)

• Geonoma

Reverse geonoma - Goguire de centro de monte (Uitoto), Tataba (Miraña)

Geonoma macrostachys - Ucsha (Ecuador), Palmiche (Peru)

• Iriartea

Iriartea deltoidea - Bombona, Bombonaje, Corneto, Trompeto, Barrigona, Cachudo (Venezuela and Colombia)

• Jessenia H.Karst.

Jessenia bataua Burret Right name: Oenocarpus bataua

• Jubaea

Jubaea chilensis (Molina) Baill. (Chile)

• Jubaeopsis

Jubaeopsis caffra Becc.

• Howea

Howea forsterana (C.Moore & F.Muell.) Becc.

• Leopoldinia

Leopoldinia piassaba- Chiquichiqui (Colombia), Málama (Vaupés), Piassava (Brazil), Fibra

• Lepidocaryum

Lepidocaryum tenue - Pui, Caraná, Erere (Uitoto)

• Livistona

Livistona australis (R.Br.) Mart.

• Lodoicea

Lodoice maldivica Coco de mer (Seychelles), Coco de mar or Coco de las Seicheles)

• Manicaria

Manicaria saccifera Gaertn. - Napa (Panama and Colombia) or Cabecinegro (Colombia)

• Mauritia

Mauritia minor Burret

Mauritia flexuosa L.f. - Moriche (Venezuela), Canangucha (Colombia) or Aguaje (Ecuador)

• Oenocarpus Mart.

Oenocarpus bataua- Seje, Milpesos (Colombia), Milpé, Komaña (Uitoto), Ungurahua (Peru)

• Orbignya Mart. ex Endl.
• Phytelephas Cook

Phytelephas seemanii - Tagua, Palma Tagua. Ivory palm.

• Phoenix

Phoenix canariensis Chabaud- Canarian palm tree

Phoenix dactylifera L. - datilera palm.

• Rhapis L.f.or
• Roystonea O.F.Cook
  • Roystonea regia H.B.K. - royal palm, real Cuban Palma
  • Roystonea oleracea O.F.Cook - Chaguaramo (Venezuela), royal palm (other countries)
• Sabal Adans.
• Salacca Reinw.
• Syagrus
• Trachycarpus L.

Trachycarpus fortunei (Hook.) H.Wendl.

• Veitchia H.Wendl. in Seem.
• Wallichia Roxb.
• Washington H.Wendl.

Washingtonia filifera (Linden) H.Wendl.

Washingtonia robust

• Weittinia

The following classification was proposed by N.W.Uhl and J.Dransfield in 1987 in Genera palmarum: a classification based on the work of Harold E. Moore, Jr. (but see newer classifications such as that of Asmussen et al. 2006):

The list of all the genera of the botanical family Arecaceae in Annex:Genus and tribes of Arecaceae.

Synonymy

Synonymy, according to APWeb (visited January 2009):

• of Arecales: Cocosales Dumort. - Arecanae Takht. - Arecidae Takht. - Phoenicopsida Brongn.
• of Calamoideae: Calamaceae Perleb, Lepidocaryaceae O.F.Cook
• of Nypoideae: Nypaceae Le Maout " Decne.
• de Coryphoideae: Borassaceae Schultz Sch., Coryphaceae Schultz Sch., Phoeniciaceae Burnett, Sabalaceae Schultz Sch.
• Ceroxyloideae: Phytelephaceae Perleb,
• of Arecoideae: Acristaceae O.F.Cook, Ceroxylaceae O.F.Cook, Chamaedoraceae O.F.Cook, Cocosaceae Schultz Sch., Geonomataceae O.F.Cook, Iriarteaceae O.F.Cook " Doyle, Malortieaceae O.F.Cook, Manicariaceae O.F.Cook, Pseudophoeniciaceae O.F.Cook, Synechanthaceae O.F.Cook

Economic importance

El Palmar National Park, Entre Ríos, Argentina, is with 8500 hectares a vast protected territory with plenty of palm trees alreadytay. The park attracts thousands of tourists annually.

In general, palm groves are one of the most important elements for Amazonian communities due to their economic, cultural, and ecological value, since from this resource they obtain their food, housing, and multiple items that satisfy their material needs. Many species of palms have great current and potential value as sources of food, oil, fiber, medicine, and other products, including their value as ornamental plants; All of the previous potentialities (used in a sustainable way) can become a valuable source of resources for the economy.

It is one of the most economically important botanical families.

Food plants come from Areca, Butia, Attalea, Bactris, Cocos (coconut tree, Cocos nucifera), Elaeis (oil-bearing, eg Elaeis oleifera ), Metroxylon (which provide starch), and Phoenix (date). Many genera have an edible apical bud. Euterpe edulis is the edible palm heart in the Southern Cone, replaced by Bactris gasipaes which allows sustainable exploitation.

Other economically important palms are Calamus (and others called rattan), Butia (the leaves have various uses), Copernicia (wax carnaúba, Copernicia cerifera: wax palm), Phytelephas (coot), Raphia (raffia), and many genera that provide straw or fiber for moored.

Finally, the family includes a large number of ornamentals, Caryota, Chamaerops, Butia, Livistona, Phoenix, Roystonea, Sabal, Syagrus, Washingtonia, Chamaedorea, Trithrinax, Rhapidophyllum, Thrinax, Coccothrinax, Licuala, Veitchia, Acoelorraphe, Butia, Copernicia, Dypsis, and Wodyetia. Among the ornamentals, Phoenix canariensis (Canary Islands palm) and Roystonea regia (Cuban royal palm) and many others stand out.

Some species are cultivated in large areas, such as the coconut palm Cocos nucifera, the oil palm Elaeis guineensis and the date palm Phoenix dactylifera.

The sap of some species is concentrated or fermented to make "honeys" and "wines" of palm tree

The fruit of various species of the genus Areca is chewed in Asia as a stimulant and is known as betel.

Other species confused with Arecaceae

Some species are also commonly known as palms, although they do not belong to the Arecaceae:

• Cordyline australis(family Asparagaceae) and other representatives of the genus Cordyline and also Dracaena with whom Cordyline It can be confused.
• Stresses Cycas revoluta, but usually many species of the Cycadopsida class are similar and confused with arecaceae.
• Ravenala(family Strelitziaceae)
• Pandanus spiralisand maybe other species of Pandanus
• Cyathea cunninghamiiand other arborescent ferns (Cyatheaceae and Dicksoniaceae families) can also be confused with palm trees.
• Setaria palmifoliaA gram.
• Carludovica palmataand other members of the Cyclanthaceae family.
• Pachypodium lamerei^{[chuckles]required]} (Family Apocynaceae) and perhaps other species of the genus Pachypodium
• Yucca elephantipes^{[chuckles]required]} and other species of the genus Yucca and the subfamily Agavoideae
• Various species of the subfamily Nolinoideae^{[chuckles]required]}
• Various species of gender Kingia^{[chuckles]required]}
• Various species of gender Cyperus^{[chuckles]required]}
• Various species of the family Pandanaceae^{[chuckles]required]}
• Various species of gender Beaucarnea^{[chuckles]required]}
• Various species of the subfamily Xanthorrhoeoideae^{[chuckles]required]}
• Various species of the genus Wilkesia^{[chuckles]required]}
• The Puya raimondii^{[chuckles]required]}

References Cited