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Attalea (genus)
Encyclopedia
Attalea is a large genus
of palms
native to Mexico
, the Caribbean
, Central
and South America
. This pinnate
ly leaved, non-spiny genus includes both small palms lacking an aboveground stem and large trees. The genus has a complicated taxonomic history, and has often been split into four or five genera based on differences in the male flowers. Since the genera can only be distinguished on the basis of their male flowers, the existence of intermediate flower types and the existence of hybrids between different genera has been used as an argument for keeping them all in the same genus. This has been supported by a recent molecular phylogeny.
Somewhere between 29 and 67 species are recognised in the genus, with estimates of as many as 100. Incomplete herbarium
collections make it difficult to determine whether certain groups represent single species, or groups of similar species. Attalea species have a long history of human use, and include economically important sources of palm oil
and fibre. Many species are fire tolerant and thrive in disturbed habitats. Their seeds are animal dispersed, including some which are thought to have been adapted for dispersal by now-extinct Pleistocene megafauna
.
The genus Attalea has pinnate
ly compound leaves—rows of leaflets emerge on either side of the axis of the leaf in a feather-like or fern-like pattern. Species are also non-spiny
palms and includes both large trees with stout stems up to 30 metres (98.4 ft) tall and acaulescent palms (one which lack an aboveground stem). The number of leaves per individual varies from about three to thirty-five; larger plants tend to have more and longer leaves.
Inflorescence
s are large, branched and borne among the leaves. The inflorescence consists of a main axis—the peduncle
and the rachis
—and a series of smaller branches, the rachillae. The rachillae, which bear the flowers, emerge from the rachis. The peduncle is the main stalk, connecting the rachis with the stem. Inflorescences either consist entirely of male flowers, or are predominantly female with a few male flowers. Fruit usually have two or three seeds, although fewer or more are present in some species, and are usually brown, yellow, orange-brown or purple when mature.
Four different types of male flowers exist. On the basis of these flower types, the genus has often been split into four genera—a more narrowly defined Attalea, Orbignya, Maximiliana and Scheelea. The species sometimes referred to Orbignya have coiled anthers, while the other groups have straight ones. The petals of those placed in Maximiliana are much shorter than the stamen
s, while those placed in Scheelea and a more narrowly defined Attalea have petals that are longer than the stamens. Five species do not fit easily into any of these groups; this fact has been used as an argument in favour of considering this group a single genus.
Arecoideae, the tribe
Cocoseae and the subtribe Attaleinae, together with the genera Allagoptera
, Beccariophoenix
, Butia
, Cocos, Jubaea
, Jubaeopsis
, Lytocaryum
, Parajubaea
, Syagrus
and Voanioala
. Within this subtribe, Attalea has been found to be a monophyletic group, and sister to the clade containing Allagoptera, Polyandrococos
, Parajubaea, Butia and Jubaea.
Disagreement exists as to whether Attalea should be considered a single genus, or a group of related genera. In their 1996 Field Guide to the Palms of the Americas, Andrew Henderson
, Gloria Galeano
and Rodrigo Bernal
combined all the species in the subtribe Attaleinae (as it was then defined) into a single genus, Attalea. In his 1999 Taxonomic Treatment of Palm Subtribe Attaleinae, American botanist Sidney F. Glassman divided the group into five genera—a more narrowly defined Attalea, Orbignya, Maximiliana, Scheelea and Ynesa. Rafäel Govaerts and John Dransfield
recognised a single genus in their 2005 World Checklist of Palms
, and Jean-Christophe Pintaud continued this usage in his 2008 review of the genus.
The multi-genus approach is based solely on the structure of the male flowers; no other characters could be consistently associated with one genus or another. Four of the genera—Attalea (in a narrow sense), Orbignya, Maximiliana and Scheelea—correspond to four different types of male flowers found within the genus. However, a few species have flowers that are intermediate between these four types, including A. colenda (which Glassman placed in its own genus, Ynesa) and this has been used as an argument for the single-genus approach. In addition, there are several hybrids between species that would be considered different genera under Glassman's five-genus system, which has also been used as an argument for placing them in a single genus. In 2009 Alan Meerow and colleagues published a molecular phylogeny of the subtribe which found that some species placed in Orbignya were actually more closely related to species placed in Scheelea than they were to other members of that genus (if the five-genus approach was used), while A. crassispatha
, placed in Orbignya by Glassman, was actually a sister to both Scheelea and Orbignya.
in 1816 based on specimens collected by Alexander von Humboldt
and Aimé Bonpland
, although older, pre-Linnaean descriptions exist, including Charles Plumier
's 1703 description of A. crassispatha. The genus was named for Attalus III Philometor
, king of Pergamon
, known for his interest in medicinal plants. The type species
is A. amygdalina, a Colombia
n endemic. The genera Maximiliana and Orbignya were described by Carl Friedrich Philipp von Martius
in 1826 and 1837 respectively. Scheelea was described by Hermann Karsten
in 1857, and Ynesa by Orator F. Cook
in 1942.
estimated that there may be as many as 100 species in the genus. In their 1996 Field Guide to the Palms of the Americas Andrew Henderson and coauthors recognised 29 species in the genus, while Sidney Glassman recognised 65 species in his 1999 treatment of the group. Largely following Glassman's lead, Rafaël Govaerts and John Dransfield recognised 67 species in their 2005 World Checklist of Palms. An important element of this disagreement is the decision by Glassman to define species more narrowly than Henderson. As a result, what Henderson interpreted as variation within species, Glassman took as differences between morphologically
similar species. This problem is complicated by the fact that many of these species are poorly represented in herbarium
collections. The large size of the leaves, inflorescences and fruit of many Attalea species makes them difficult to collect. In addition, many important collections, including type specimen, have been lost or destroyed. Sparse or incomplete collections make it difficult to differentiate variation within a single species from variation between different species.
The three recent treatments (Henderson and coauthors, Glassman, and Govaerts and Dransfield) recognised a total of 73 species, but only 20 species are accepted by all of them. The remainder account for either nine species or more than 40. For example, what Andrew Henderson considered a single species, Attalea attaleoides, other authors have considered a species complex
consisting of four or five species. Glassman doubted the validity of A. attaleoides as a species, and described four new species from material that had previously been attributed to A. attaleoides—A. camopiensis, A. degranvillei, A. guianensis and A. maripensis. Govaerts and Dransfield accepted both Glassman's four species and A. attaleoides. However, Jean-Christophe Pintaud was of the opinion that A. guianensis, A. maripensis and A. attaleoides were all very similar, and thought it likely that they all represented the same species.
Another species complex in Attalea includes A. speciosa
and related species. Henderson (1995) recognised A. speciosa and A. spectabilis, considering the latter to either be an acaulescent form of A. speciosa or a hybrid between it and A. microcarpa. Govaerts and Dransfield accepted A. spectabilis, but Glassman considered it a dubious
taxon. Attalea vitrivir was recognised as a distinct species by Michael Balick and coauthors; Glassman and Govaerts and Dransfield concurred, but Henderson considered it part of A. speciosa. Glassman also described a fourth member of this group, A. brejinhoensis, and it is accepted by Govaerts and Dransfield.
—male and female flowers are separate, but are borne by the same plant. Various species have been described as being insect pollinated, including A. phalerata, while pollination in A. colenda and A. speciosa
, has been attributed both to insects and wind.
Seed germination
is remote tubular—during germination, as the cotyledon
expands it pushes the young shoot away from the seed. After germination, the stem initially grows downward before turning to grow upward and produce the aboveground stem. This produces a "saxophone shaped" belowground portion of the stem. The fact that the shoot tips of Attalea seedlings are underground it likely to contribute to their fire-tolerance.
in the north to Bolivia
, Paraguay
, and southern Brazil
in the south. According to Govaerts and coauthors, three species are found in Mexico, four in Central America
, and 62 in South America
. Three species are present in the Caribbean
—two in Trinidad and Tobago
, along the southern edge of the region, and one in Haiti.
Attalea includes both large trees and small acaulescent palms which occupy a number of different ecological niches. Dense stands of some of the larger species are conspicuous elements on the landscape, while smaller species are found in both in the forest understorey and in savannas.
Disturbance has been implicated in the formation of vegetation dominated by large Attalea species. In seasonally dry Amazonian forests the density of large adult A. maripa palms was correlated with canopy openness; the species also dominates savannas formed by repeated forest fires in Trinidad and Tobago
. Attalea speciosa
forms pure stands in many parts of Brazil where natural forest vegetation has been cleared. Similarly, stands of A. funifera in Bahia
, Brazil (which are cultivated for piassava
fibre) are managed using fire—the seedlings survive cutting and burning, and are able to dominate burned forest patches.
The fruit are dispersed by animals; fruit which are not dispersed frequently suffer seed predation by bruchid beetles
. Certain species of Attalea have been mentioned as examples of "anachronistic" species which are adapted for dispersal by now-extinct Pleistocene megafauna
. On Maracá Island, Roraima
, in the Brazilian Amazon, Attalea maripa fruit were consumed by tapirs
, collared peccaries
, deer and primates. Rodents, including agouti
s, fed upon the fruit and, as the fruit availability declined, they fed on the seeds. Other dispersers of Attalea fruit include Crested Caracaras
which consume the fruit and disperse the seeds of A. phalerata in the Brazilian Pantanal
.
. A variety of species remain important sources of edible oil, thatch, edible seeds and fibre. The leaves of Attalea butyracea and A. maripa are used extensively for thatching. Several species are oil palms, with A. speciosa
among the most important economically. Products extracted from A. speciosa were reported to support over 300,000 households in the Brazilian state of Maranhão
in 2005, and in 1985 it was estimated to support over 450,000 households throughout the Brazil. Piassava
fibres, extracted from the leaf bases of A. funifera, are commercially important, and generated about US $
20 million in annual income to Brazilian farmers in 1996.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of palms
Arecaceae
Arecaceae or Palmae , are a family of flowering plants, the only family in the monocot order Arecales. There are roughly 202 currently known genera with around 2600 species, most of which are restricted to tropical, subtropical, and warm temperate climates...
native to Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
, the Caribbean
Caribbean
The Caribbean is a crescent-shaped group of islands more than 2,000 miles long separating the Gulf of Mexico and the Caribbean Sea, to the west and south, from the Atlantic Ocean, to the east and north...
, Central
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
and South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
. This pinnate
Pinnate
Pinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
ly leaved, non-spiny genus includes both small palms lacking an aboveground stem and large trees. The genus has a complicated taxonomic history, and has often been split into four or five genera based on differences in the male flowers. Since the genera can only be distinguished on the basis of their male flowers, the existence of intermediate flower types and the existence of hybrids between different genera has been used as an argument for keeping them all in the same genus. This has been supported by a recent molecular phylogeny.
Somewhere between 29 and 67 species are recognised in the genus, with estimates of as many as 100. Incomplete herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...
collections make it difficult to determine whether certain groups represent single species, or groups of similar species. Attalea species have a long history of human use, and include economically important sources of palm oil
Palm oil
Palm oil, coconut oil and palm kernel oil are edible plant oils derived from the fruits of palm trees. Palm oil is extracted from the pulp of the fruit of the oil palm Elaeis guineensis; palm kernel oil is derived from the kernel of the oil palm and coconut oil is derived from the kernel of the...
and fibre. Many species are fire tolerant and thrive in disturbed habitats. Their seeds are animal dispersed, including some which are thought to have been adapted for dispersal by now-extinct Pleistocene megafauna
Pleistocene megafauna
Pleistocene megafauna is the set of species of large animals — mammals, birds and reptiles — that lived on Earth during the Pleistocene epoch and became extinct in a Quaternary extinction event. These species appear to have died off as humans expanded out of Africa and southern Asia,...
.
Description
Pinnate
Pinnate is a term used to describe feather-like or multi-divided features arising from both sides of a common axis in plant or animal structures, and comes from the Latin word pinna meaning "feather", "wing", or "fin". A similar term is pectinate, which refers to a comb-like arrangement of parts...
ly compound leaves—rows of leaflets emerge on either side of the axis of the leaf in a feather-like or fern-like pattern. Species are also non-spiny
Thorns, spines, and prickles
In botanical morphology, thorns, spines, and prickles are hard structures with sharp, or at least pointed, ends. In spite of this common feature, they differ in their growth and development on the plant; they are modified versions of different plant organs, stems, stipules, leaf veins, or hairs...
palms and includes both large trees with stout stems up to 30 metres (98.4 ft) tall and acaulescent palms (one which lack an aboveground stem). The number of leaves per individual varies from about three to thirty-five; larger plants tend to have more and longer leaves.
Inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
s are large, branched and borne among the leaves. The inflorescence consists of a main axis—the peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...
and the rachis
Rachis
Rachis is a biological term for a main axis or "shaft".-In zoology:In vertebrates a rachis can refer to the series of articulated vertebrae, which encase the spinal cord. In this case the rachis usually form the supporting axis of the body and is then called the spine or vertebral column...
—and a series of smaller branches, the rachillae. The rachillae, which bear the flowers, emerge from the rachis. The peduncle is the main stalk, connecting the rachis with the stem. Inflorescences either consist entirely of male flowers, or are predominantly female with a few male flowers. Fruit usually have two or three seeds, although fewer or more are present in some species, and are usually brown, yellow, orange-brown or purple when mature.
Four different types of male flowers exist. On the basis of these flower types, the genus has often been split into four genera—a more narrowly defined Attalea, Orbignya, Maximiliana and Scheelea. The species sometimes referred to Orbignya have coiled anthers, while the other groups have straight ones. The petals of those placed in Maximiliana are much shorter than the stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...
s, while those placed in Scheelea and a more narrowly defined Attalea have petals that are longer than the stamens. Five species do not fit easily into any of these groups; this fact has been used as an argument in favour of considering this group a single genus.
Taxonomy
Attalea has been placed in the subfamilyFamily (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
Arecoideae, the tribe
Tribe (biology)
In biology, a tribe is a taxonomic rank between family and genus. It is sometimes subdivided into subtribes.Some examples include the tribes: Canini, Acalypheae, Hominini, Bombini, and Antidesmeae.-See also:* Biological classification* Rank...
Cocoseae and the subtribe Attaleinae, together with the genera Allagoptera
Allagoptera
Allagoptera is a monoecious genus of flowering plant in the palm family found in South America consisting of four similar-looking species. Compared to other genera within the Cocoeae it is described as particularly specialized...
, Beccariophoenix
Beccariophoenix
Beccariophoenix is a genus of 3 species of Arecaceae , native to Madagascar. The genus is closely related to the Cocos, or coconut genus, and Beccariophoenix species, notably Beccariophoenix alfredii are very similar in appearance to the coconut palm, although they are somewhat cold hardy, making...
, Butia
Butia
Butia,also known as a Pindo Palm is a genus of palms in the family Arecaceae, native to South America in Brazil, Paraguay, Uruguay and Argentina...
, Cocos, Jubaea
Jubaea
Jubaea chilensis is the sole extant species in the genus Jubaea in the palm family Arecaceae. It is native to southwestern South America, where it is endemic to a small area of central Chile, between 32°S and 35°S in southern Coquimbo, Valparaíso, Santiago, O'Higgins and northern Maule regions...
, Jubaeopsis
Jubaeopsis
The Pondoland Palm is a flowering plant species in the palm family . It belongs to the monotypic genus Jubaeopsis....
, Lytocaryum
Lytocaryum
Lytocaryum is a monoecious genus of flowering plant in the palm family endemic to Brazil where two species are known. Palms once classified as Microcoelum are herein included; the genus is closely related to Syagrus, from which it is differentiated only by abundant tomentum, strongly versatile...
, Parajubaea
Parajubaea
Parajubaea is a genus of flowering plant in the Arecaceae family.It contains the following species:* Sunkha Palm * Palma De Pasobaya * Coquitos...
, Syagrus
Syagrus
Syagrus is a genus of 30 to 42 species of Arecaceae , native to South America, with one species endemic to the Lesser Antilles. The genus is closely related to the Cocos, or coconut genus, and many Syagrus species produce edible seeds similar to the coconut.-Description:Palms in this group have...
and Voanioala
Voanioala
Voanioala gerardii, commonly known as the Forest Coconut, is a species of flowering plant in the Arecaceae family. It is a large Coconut relative that is monotypic within the genus Voanioala.It is endemic to Madagascar....
. Within this subtribe, Attalea has been found to be a monophyletic group, and sister to the clade containing Allagoptera, Polyandrococos
Polyandrococos
Polyandrococos is a monotypic genus of flowering plant in the palm family endemic to Brazil, where it is known as buri palm. The name combines the Greek words for "many" and "anther" with the name of another palm genus Cocos, and the epithet is Latin for "bearlike", referring to the hairy...
, Parajubaea, Butia and Jubaea.
Disagreement exists as to whether Attalea should be considered a single genus, or a group of related genera. In their 1996 Field Guide to the Palms of the Americas, Andrew Henderson
Andrew Henderson (botanist)
Andrew James Henderson is a palm-systematist and Curator of the Institute of Systematic Botany at the New York Botanical Garden. He has authored taxonomic descriptions of 140 species, subspecies and varieties of plants, especially in the palm family...
, Gloria Galeano
Gloria Galeano
Gloria A. Galeano Garcés is a Colombian botanist and agronomist. Galeano is a faculty member at the Institute of Natural Sciences, National University of Colombia. She received her Ph.D...
and Rodrigo Bernal
Rodrigo Bernal
Rodrigo Bernal González is a Colombian botanist who specialises in the palm family. Bernal is a faculty member at the Institute of Natural Sciences, National University of Colombia. He received his Ph.D. from the University of Aarhus in 1996.Bernal has published over sixty works on palm...
combined all the species in the subtribe Attaleinae (as it was then defined) into a single genus, Attalea. In his 1999 Taxonomic Treatment of Palm Subtribe Attaleinae, American botanist Sidney F. Glassman divided the group into five genera—a more narrowly defined Attalea, Orbignya, Maximiliana, Scheelea and Ynesa. Rafäel Govaerts and John Dransfield
John Dransfield
John Dransfield is an honorary research fellow and former head of palm research at the Royal Botanic Gardens, Kew, United Kingdom, as well as being an authority on the phylogenetic classification of palms....
recognised a single genus in their 2005 World Checklist of Palms
, and Jean-Christophe Pintaud continued this usage in his 2008 review of the genus.
The multi-genus approach is based solely on the structure of the male flowers; no other characters could be consistently associated with one genus or another. Four of the genera—Attalea (in a narrow sense), Orbignya, Maximiliana and Scheelea—correspond to four different types of male flowers found within the genus. However, a few species have flowers that are intermediate between these four types, including A. colenda (which Glassman placed in its own genus, Ynesa) and this has been used as an argument for the single-genus approach. In addition, there are several hybrids between species that would be considered different genera under Glassman's five-genus system, which has also been used as an argument for placing them in a single genus. In 2009 Alan Meerow and colleagues published a molecular phylogeny of the subtribe which found that some species placed in Orbignya were actually more closely related to species placed in Scheelea than they were to other members of that genus (if the five-genus approach was used), while A. crassispatha
Attalea crassispatha
Attalea crassispatha is a palm which is endemic to southwest Haiti. The most geographically isolated member of the genus, it is considered a critically endangered species and has been called one of the rarest palms in the Americas.-Description:...
, placed in Orbignya by Glassman, was actually a sister to both Scheelea and Orbignya.
History
The genus Attalea was first described Carl Sigismund KunthCarl Sigismund Kunth
Carl Sigismund Kunth , also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist...
in 1816 based on specimens collected by Alexander von Humboldt
Alexander von Humboldt
Friedrich Wilhelm Heinrich Alexander Freiherr von Humboldt was a German naturalist and explorer, and the younger brother of the Prussian minister, philosopher and linguist Wilhelm von Humboldt...
and Aimé Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...
, although older, pre-Linnaean descriptions exist, including Charles Plumier
Charles Plumier
Charles Plumier was a French botanist, after whom the Frangipani genus Plumeria is named. Plumier is considered one of the most important of the botanical explorers of his time...
's 1703 description of A. crassispatha. The genus was named for Attalus III Philometor
Attalus III
Attalus III Philometor Euergetes was the last Attalid king of Pergamon, ruling from 138 BC to 133 BC....
, king of Pergamon
Pergamon
Pergamon , or Pergamum, was an ancient Greek city in modern-day Turkey, in Mysia, today located from the Aegean Sea on a promontory on the north side of the river Caicus , that became the capital of the Kingdom of Pergamon during the Hellenistic period, under the Attalid dynasty, 281–133 BC...
, known for his interest in medicinal plants. The type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...
is A. amygdalina, a Colombia
Colombia
Colombia, officially the Republic of Colombia , is a unitary constitutional republic comprising thirty-two departments. The country is located in northwestern South America, bordered to the east by Venezuela and Brazil; to the south by Ecuador and Peru; to the north by the Caribbean Sea; to the...
n endemic. The genera Maximiliana and Orbignya were described by Carl Friedrich Philipp von Martius
Carl Friedrich Philipp von Martius
Carl Friedrich Philipp von Martius was a German botanist and explorer.Martius was born at Erlangen, where he graduated M.D. in 1814, publishing as his thesis a critical catalogue of plants in the botanic garden of the university...
in 1826 and 1837 respectively. Scheelea was described by Hermann Karsten
Gustav Karl Wilhelm Hermann Karsten
Gustav Karl Wilhelm Hermann Karsten was a German botanist and geologist. Born in Stralsund, he followed the example of Alexander von Humboldt and traveled 1844-56 to the north of South America. He died 1908 in Berlin-Grunewald....
in 1857, and Ynesa by Orator F. Cook
Orator F. Cook
Orator Fuller Cook was an American botanist, entomologist, and agronomist. Cook, born in Clyde, New York in 1867, graduated from Syracuse University in 1890. He worked for one year as an instructor at Syracuse. In 1891 Cook became a special agent of the New York State Colonization Society. He...
in 1942.
Species
Experts disagree about the number of species in the genus Attalea (broadly defined). In 1965, Dutch taxonomist Jan Gerard Wessels BoerJan Gerard Wessels Boer
Jan Gerard Wessels Boer is a Dutch plant taxonomist best known for his work on palms. He has described 37 species, subspecies and varieties of plants, especially in the palm family-References:...
estimated that there may be as many as 100 species in the genus. In their 1996 Field Guide to the Palms of the Americas Andrew Henderson and coauthors recognised 29 species in the genus, while Sidney Glassman recognised 65 species in his 1999 treatment of the group. Largely following Glassman's lead, Rafaël Govaerts and John Dransfield recognised 67 species in their 2005 World Checklist of Palms. An important element of this disagreement is the decision by Glassman to define species more narrowly than Henderson. As a result, what Henderson interpreted as variation within species, Glassman took as differences between morphologically
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
similar species. This problem is complicated by the fact that many of these species are poorly represented in herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...
collections. The large size of the leaves, inflorescences and fruit of many Attalea species makes them difficult to collect. In addition, many important collections, including type specimen, have been lost or destroyed. Sparse or incomplete collections make it difficult to differentiate variation within a single species from variation between different species.
The three recent treatments (Henderson and coauthors, Glassman, and Govaerts and Dransfield) recognised a total of 73 species, but only 20 species are accepted by all of them. The remainder account for either nine species or more than 40. For example, what Andrew Henderson considered a single species, Attalea attaleoides, other authors have considered a species complex
Species complex
A species complex is a group of closely related species, where the exact demarcation between species is often unclear or cryptic owing to their recent and usually still incomplete reproductive isolation. Ring species, superspecies and cryptic species complex are example of species complex...
consisting of four or five species. Glassman doubted the validity of A. attaleoides as a species, and described four new species from material that had previously been attributed to A. attaleoides—A. camopiensis, A. degranvillei, A. guianensis and A. maripensis. Govaerts and Dransfield accepted both Glassman's four species and A. attaleoides. However, Jean-Christophe Pintaud was of the opinion that A. guianensis, A. maripensis and A. attaleoides were all very similar, and thought it likely that they all represented the same species.
Another species complex in Attalea includes A. speciosa
Attalea speciosa
Attalea speciosa is a palm native to the Amazon Rainforest region in South America...
and related species. Henderson (1995) recognised A. speciosa and A. spectabilis, considering the latter to either be an acaulescent form of A. speciosa or a hybrid between it and A. microcarpa. Govaerts and Dransfield accepted A. spectabilis, but Glassman considered it a dubious
Nomen dubium
In zoological nomenclature, a nomen dubium is a scientific name that is of unknown or doubtful application...
taxon. Attalea vitrivir was recognised as a distinct species by Michael Balick and coauthors; Glassman and Govaerts and Dransfield concurred, but Henderson considered it part of A. speciosa. Glassman also described a fourth member of this group, A. brejinhoensis, and it is accepted by Govaerts and Dransfield.
Reproduction and growth
Attalea species are monoeciousPlant sexuality
Plant sexuality covers the wide variety of sexual reproduction systems found across the plant kingdom. This article describes morphological aspects of sexual reproduction of plants....
—male and female flowers are separate, but are borne by the same plant. Various species have been described as being insect pollinated, including A. phalerata, while pollination in A. colenda and A. speciosa
Attalea speciosa
Attalea speciosa is a palm native to the Amazon Rainforest region in South America...
, has been attributed both to insects and wind.
Seed germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...
is remote tubular—during germination, as the cotyledon
Cotyledon
A cotyledon , is a significant part of the embryo within the seed of a plant. Upon germination, the cotyledon may become the embryonic first leaves of a seedling. The number of cotyledons present is one characteristic used by botanists to classify the flowering plants...
expands it pushes the young shoot away from the seed. After germination, the stem initially grows downward before turning to grow upward and produce the aboveground stem. This produces a "saxophone shaped" belowground portion of the stem. The fact that the shoot tips of Attalea seedlings are underground it likely to contribute to their fire-tolerance.
Distribution
Species range across the Neotropics from MexicoMexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
in the north to Bolivia
Bolivia
Bolivia officially known as Plurinational State of Bolivia , is a landlocked country in central South America. It is the poorest country in South America...
, Paraguay
Paraguay
Paraguay , officially the Republic of Paraguay , is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to the east and northeast, and Bolivia to the northwest. Paraguay lies on both banks of the Paraguay River, which runs through the center of the...
, and southern Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
in the south. According to Govaerts and coauthors, three species are found in Mexico, four in Central America
Central America
Central America is the central geographic region of the Americas. It is the southernmost, isthmian portion of the North American continent, which connects with South America on the southeast. When considered part of the unified continental model, it is considered a subcontinent...
, and 62 in South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...
. Three species are present in the Caribbean
Caribbean
The Caribbean is a crescent-shaped group of islands more than 2,000 miles long separating the Gulf of Mexico and the Caribbean Sea, to the west and south, from the Atlantic Ocean, to the east and north...
—two in Trinidad and Tobago
Trinidad and Tobago
Trinidad and Tobago officially the Republic of Trinidad and Tobago is an archipelagic state in the southern Caribbean, lying just off the coast of northeastern Venezuela and south of Grenada in the Lesser Antilles...
, along the southern edge of the region, and one in Haiti.
Habitat and ecology
Disturbance has been implicated in the formation of vegetation dominated by large Attalea species. In seasonally dry Amazonian forests the density of large adult A. maripa palms was correlated with canopy openness; the species also dominates savannas formed by repeated forest fires in Trinidad and Tobago
Trinidad and Tobago
Trinidad and Tobago officially the Republic of Trinidad and Tobago is an archipelagic state in the southern Caribbean, lying just off the coast of northeastern Venezuela and south of Grenada in the Lesser Antilles...
. Attalea speciosa
Attalea speciosa
Attalea speciosa is a palm native to the Amazon Rainforest region in South America...
forms pure stands in many parts of Brazil where natural forest vegetation has been cleared. Similarly, stands of A. funifera in Bahia
Bahia
Bahia is one of the 26 states of Brazil, and is located in the northeastern part of the country on the Atlantic coast. It is the fourth most populous Brazilian state after São Paulo, Minas Gerais and Rio de Janeiro, and the fifth-largest in size...
, Brazil (which are cultivated for piassava
Piassava
Piassava, which is also called Piaçaba and Piasaba or Pissaba, or Piassaba, is a fibrous product of two Brazilian palms: Attalea funifera and Leopoldinia piassaba. It is often used in making brooms, and for other purposes....
fibre) are managed using fire—the seedlings survive cutting and burning, and are able to dominate burned forest patches.
The fruit are dispersed by animals; fruit which are not dispersed frequently suffer seed predation by bruchid beetles
Bean weevil
The bean weevils or seed beetles are a subfamily of beetles, now placed in the family Chrysomelidae, though they have historically been treated as a separate family. They are granivores, and typically infest various kinds of seeds or beans, living for most of their lives inside a single seed...
. Certain species of Attalea have been mentioned as examples of "anachronistic" species which are adapted for dispersal by now-extinct Pleistocene megafauna
Pleistocene megafauna
Pleistocene megafauna is the set of species of large animals — mammals, birds and reptiles — that lived on Earth during the Pleistocene epoch and became extinct in a Quaternary extinction event. These species appear to have died off as humans expanded out of Africa and southern Asia,...
. On Maracá Island, Roraima
Roraima
Roraima is the northernmost and least populated state of Brazil, located in the Amazon region. It borders the states of Amazonas and Pará, as well as the nations of Venezuela and Guyana. The population is 400,000 and the capital is Boa Vista...
, in the Brazilian Amazon, Attalea maripa fruit were consumed by tapirs
Brazilian Tapir
The South American Tapir , or Brazilian Tapir or Lowland Tapir or Anta, is one of four species in the tapir family, along with the Mountain Tapir, the Malayan Tapir, and Baird's Tapir...
, collared peccaries
Collared Peccary
The collared peccary is a species of mammal in the family Tayassuidae that is found in North, Central, and South America. They are commonly referred to as javelina, saíno or báquiro, although these terms are also used to describe other species in the family...
, deer and primates. Rodents, including agouti
Common agouti
The popular term Agouti designates several rodent species of the genus Dasyprocta that inhabit areas of Middle America, the West Indies, and northern South America. They are related to guinea pigs and look quite similar but have longer legs. The species vary in color from tawny to dark brown with...
s, fed upon the fruit and, as the fruit availability declined, they fed on the seeds. Other dispersers of Attalea fruit include Crested Caracaras
Caracara (genus)
Caracara is a genus of birds of prey in the family Falconidae found throughout a large part of the Americas. They are part of a group collectively referred to as caracaras...
which consume the fruit and disperse the seeds of A. phalerata in the Brazilian Pantanal
Pantanal
The Pantanal is a tropical wetland and one of the world's largest wetland of any kind. Most of it lies within the Brazilian state of Mato Grosso do Sul, but it extends into Mato Grosso and portions of Bolivia and Paraguay, sprawling over an area estimated at between and...
.
Uses
Attalea species have a long history of human utilisation. Carbonised Attalea maripa seeds have been found in archaeological sites in Colombia dating back to 9000 BPBefore Present
Before Present years is a time scale used in archaeology, geology, and other scientific disciplines to specify when events in the past occurred. Because the "present" time changes, standard practice is to use AD 1950 as the origin of the age scale, reflecting the fact that radiocarbon...
. A variety of species remain important sources of edible oil, thatch, edible seeds and fibre. The leaves of Attalea butyracea and A. maripa are used extensively for thatching. Several species are oil palms, with A. speciosa
Attalea speciosa
Attalea speciosa is a palm native to the Amazon Rainforest region in South America...
among the most important economically. Products extracted from A. speciosa were reported to support over 300,000 households in the Brazilian state of Maranhão
Maranhão
Maranhão is a northeastern state of Brazil. To the north lies the Atlantic Ocean. Maranhão is neighbored by the states of Piauí, Tocantins and Pará. The people of Maranhão have a distinctive accent...
in 2005, and in 1985 it was estimated to support over 450,000 households throughout the Brazil. Piassava
Piassava
Piassava, which is also called Piaçaba and Piasaba or Pissaba, or Piassaba, is a fibrous product of two Brazilian palms: Attalea funifera and Leopoldinia piassaba. It is often used in making brooms, and for other purposes....
fibres, extracted from the leaf bases of A. funifera, are commercially important, and generated about US $
United States dollar
The United States dollar , also referred to as the American dollar, is the official currency of the United States of America. It is divided into 100 smaller units called cents or pennies....
20 million in annual income to Brazilian farmers in 1996.