Attalea is a large genus of palms native to Mexico , the Caribbean , Central and South America . This pinnately 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 four generations. 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 generations 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 recognizable 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 fiber. Many species are fire tolerant and thrive in disturbed habitats. Their seeds are animal dispersed, Including Some are thought to qui-have-been adapté for dispersal by now-extinct Pleistocene megafauna .
Description
The genus Attalea has pinnately 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 include both large trees with stout up to 30 meters (98 ft) tall and acaulescent palms (which lack an aboveground stem). [3] The number of leaves per individual varies from one to three to thirty-five; larger plants tend to have more leaves. [3]
Inflorescences are broad, branched and bound among the leaves. [3] The inflorescence consists of a main axis-the peduncle and the rachis -and a series of smaller branches, the rachillae. The rachillae, which bears the flowers, emerge from the rachis. The peduncle is the main stalk, connecting the rachis with the stem. [4] Inflorescences or consist of male flowers, or are predominantly female with a few male flowers. [3] Fruit usually have two or three seeds, and are usually brown, yellow, orange-brown or purple when mature. [3]
Four different types of male flowers On the basis of these flower types, the genus has often been divided into two types: Attalea , Orbignya , Maximiliana and Scheelea . [3] The species sometimes referred to as Orbignya have coiled anthers , while the other groups have straight ones. The petals Of Those Placed in Maximiliana are much go short than the stamens , while Those Placed in Scheelea and a more narrowly defined Attalea -have That petals are along than the stamens. [4]Five species do not fit into any of these groups; This fact has been used as an argument in favor of this single genus. [4]
Taxonomy
Attalea has been placed in the subfamily Arecoideae , the tribe Cocoseae and the subtribe Attaleinae , together with the genera Allagoptera , Beccariophoenix , Butia , Cocos , Jubaea , Jubaeopsis , Lytocaryum , Parajubaea , Syagrus and Voanioala . [1] Within this subtribe, Attalea has been found to be a monophyletic group, and sister to the clade containing Allagoptera,Polyandrococos , [6] Parajubaea , Butia and Jubaea . [5]
Disagreement exists as to whether Attalea shoulds be regarded 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 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 andYnesa . [4] Rafäel Govaerts and John Dransfield ( 2005). The World Checklist of Palms , [7] and Jean-Christophe Pintaud continue this use in his review of the genus. [4]
The multi-genus approach is based on the structure of the male flowers; no other characters could be consistently associated with one genus or another. [8] Four of the genera – Attalea (in a narrow sense), Orbignya , Maximiliana and Scheelea – correspond to different types of male flowers found in the genus. However, a few species have been identified by 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 under glassman’s five-genus system, which has also been used as an argument for placing them in a single genus. [4] In 2009 Alan Meerow and colleagues published a molecular phylogeny of the subtribe qui found That Some species Placed in Orbignya Were Actually more étroitement 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. [5]
History
The genus Attalea was first described Carl Sigismund Kunth in 1816 based on specimens by Alexander von Humboldt and Aimé Bonpland , [2] even older, pre- Linnaean descriptions exist, including Charles Plumier’s 1703 description of A. crassispatha . [9] The genus was named for Attalus III Philometor , king of Pergamon , known for his interest in medicinal plants . [3] The species species is A. amygdalina , a Colombian endemic. [10] The genera Maximiliana and Orbignya were described by Carl Friedrich Philipp von Martius in 1826 [11] and 1837 [12]respectively. Scheelea was described by Hermann Karsten in 1857, [13] and Ynesa by Orator F. Cook in 1942. [14]
Species
Experts disagree about the number of species in the genus Attalea (broadly defined). In 1965, Dutch taxonomist Jan Gerard Wessels Boerestimated that there are many species in the genus. In their 1996 Field Guide to the Palms of the Americas Andrew Henderson and Coauthors Recognized 29 species in the genus, while Sidney Glassman recognized 65 species in his 1999 treatment of the group. Largely following Glassman’s Lead, Rafael Govaerts and John Dransfield Recognized 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 construed as variation Within species, Glassman Took have differences entre 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 the fruit of many Attalea species makes them difficult to collect. In addition, many important collections, including type specimen , have been lost or destroyed. [4]Sparse or incomplete collections make it difficult to differentiate variation within species. [4] [8]
The three recent treatments (Henderson and coauthors, Glassman, and Govaerts and Dransfield) are all of a kind. For example, what Andrew Henderson considered as a single species, Attalea attaleoides , [15] other authors have considered a species complex consisting of four species or species. Glassman doubted the validity of A. attaleoides as a species, and new species from oven Described material That HAD beens Previously Attributed To A. attaleoides – A. camopiensis , A. degranvillei , A. guianensis andA. maripensis . Govaerts and Dransfield accepted both Glassman ‘s oven 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. [4]
Another species complex in Attalea includes A. speciosa and related species. Henderson (1995), A. speciosa and A. spectabilis , which is one of the acaulescent forms 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 has been recognized as a separate species by Michael Balick and coauthors; [16] Glassman and Govaerts and Dransfield competed, 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. [4]
Reproduction and growth
Attalea species are monoecious -male and female flowers are separate, but are bound by the same plant. [3] Various species have been described as being insect-pollinated, including A. phalerata , [17] while pollination in A. colenda [18] and A. speciosa , [19] has been attributed to both insects and wind.
Seed germination is tubular remote [20] -during germination, as the cotyledon expands it pushes the young shoot away from the seed. [21] After germination, the stem first grows downward before turning to grow upward and produce the aboveground stem. [22] This produces a “saxophone shaped” belowground portion of the stem. [23] The fact that the shoot tips of Attaleaseedlings are under the influence of their fire-tolerance. [3]
Distribution
Species range across the Neotropics from Mexico to Bolivia , Paraguay , and southern Brazil in the south. [4] 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. [24]
Habitat and ecology
Attalea includes both large 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 the forest understorey and in savannas. [4]
Disturbance has been implicated in the formation of vegetation dominated by large Attalea species. [3] In seasonally dry Amazonian forests the density of adults A. maripa palms was correlated with canopy openness; [23] The species also dominates savannas by repeated forest fires in Trinidad and Tobago . [25] Attalea speciosa forms pure stands in many parts of Brazil where natural forest vegetation has been cleared. [26] Similarly, stands of A. funifera in Bahia , Brazil (which are cultivated for piassavafiber) are managed using fire-the seedlings, and are able to dominate burned forest patches. [27]
The fruit are dispersed by animals ; fruit qui arent frequently dispersed Suffer seed predation by bruchid beetles . [3] Some species of Attalea -have-been MENTIONED as examples of “anachronistic” species are qui adapté for dispersal by now-extinct Pleistocene megafauna . [28] [29] On Maracá Island, Roraima , in the Amazon Amazon , Attalea maripa fruit were consumed by tapirs , collared peccaries , deer and primates. Rodents, including agoutis, fed upon the fruit, and the fruit supply declined, they fed on the seeds. [30] Other dispersers of Attalea fruit include Crested Caracaras which consumes the fruit and disperses the seeds of A. phalerata in the Brazilian Pantanal . [31]
Uses
Attalea species have a long history of human use. Carbonized Attalea maripa seeds have been found in archaeological sites in Colombia dating back to 9000 BP . [32] A variety of species remains important sources of edible oil , thatch , edible seeds and fiber. 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. [3] Products extracted from A. speciosa were reported to support over 300,000 households in the Brazilian state ofMaranhão in 2005, [33] and in 1985 it was estimated to support over 450,000 households throughout the Brazil. [26] Piassava fibers, extracted from the leaf bases of A. funifera , are commercially important, [3] and generated about US $ 20 million in annual income to Brazilian farmers in 1996. [34]
References
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