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Attalea(plant)

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Attalea
Tall, single-stemmed palm standing alone in a field of grass with scattered shrubs and palms. The low hills in the background are forested. The leaves are all angled above horizontal, and are shorter than the stem. A single brownish-yellow infructescence is visible just below the leaves. The lower half of the stem is bare, but the upper half has old leaf-based still attached. Ferns and a strangler fig grow on the upper part of the stem just below the leaves, rooted in the old leaf bases.
Attalea brasiliensis
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Arecales
Family: Arecaceae
Subfamily: Arecoideae
Tribe: Cocoseae
Genus: Attalea
Kunth
Type species
Attalea amygdalina
Kunth
Diversity
Between 29 and 67 species
Synonyms[1]

Attaleais a largegenusofpalmsnative to Mexico, the Caribbean, Central and South America. Thispinnately-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 recent molecular phylogenies.

Between 29 and 67 species are recognised in the genus, with estimates of as many as 100. Incompleteherbariumcollections make it difficult to determine whether certain groups represent single species, or groups of similar species.Attaleaspecies have a long history of human use, and include economically important sources ofpalm oiland 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-extinctPleistocene megafauna.

Description

[edit]
A view of the crown of a palm tree from below: Dark green leaves emerge in a radial pattern from the trunk of the tree, above old, dried leaf bases from which the leaves have been cut off. Small ferns grow on them, wedged between the old leaf bases and the trunk of the palm. Between the dried leaf bases and the green leaves there are several brown inflorescences, each of which lies below a reddish-brown bract which is larger than the inflorescence.
Detail of the crown ofAttalea maripashowing leaf arrangement and inflorescences

Attaleais a genus ofnon-spinypalms with 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 range from large trees with stout stems up to 30 metres (98 ft) tall toacaulescentpalms (ones which lack an aboveground stem).[2]The number of leaves per individual varies from about three to thirty-five; larger plants tend to have more and longer leaves.[2]

Inflorescencesare large, branched and borne among the leaves.[2]The inflorescence consists of a main axis—thepeduncleand therachis—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.[3]Inflorescences either consist entirely of male flowers, or are predominantly female with a few male flowers.[2]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.[2]

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 definedAttalea,Orbignya,Maximiliana,andScheelea.[2]The species sometimes referred toOrbignyahave coiledanthers,while the other groups have straight ones. The petals of those placed inMaximilianaare much shorter than thestamens,while those placed inScheeleaand a more narrowly definedAttaleahave petals that are longer than the stamens.[3]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.[3]

Taxonomy

[edit]

Cocos nucifera

Syagrus

Lytocaryum(nested withinSyagrus)

Simplified phylogeny of members of the subtribeAttaleinae,based on seven WRKY gene loci.[4]

Attaleahas been placed in thesubfamilyArecoideae,thetribeCocoseaeand the subtribeAttaleinae,together with the generaAllagoptera,Beccariophoenix,Butia,Cocos,Jubaea,Jubaeopsis,Parajubaea,Syagrus,andVoanioala.[5][6]Within this subtribe,Attaleahas been found to be amonophyleticgroup, and sister to the clade containingAllagoptera,Polyandrococos,[7]Parajubaea,Butia,andJubaea.[4]

Disagreement exists as to whetherAttaleashould be considered a single genus, or a group of related genera. In their 1996Field Guide to the Palms of the Americas,Andrew Henderson,Gloria Galeano,andRodrigo Bernalcombined all the species in the subtribe Attaleinae (as it was then defined) into a single genus,Attalea.In his 1999Taxonomic Treatment of Palm Subtribe Attaleinae,American botanistSidney F. Glassmandivided the group into five genera—a more narrowly definedAttalea,Orbignya,Maximiliana,ScheeleaandYnesa,[3]although he thought it likely thatYnesa colenda,the only member of that genus, was actually a hybrid.[8]Rafäel GovaertsandJohn Dransfieldrecognised a single genus in their 2005World Checklist of Palms,[9]and Jean-Christophe Pintaud continued this usage in his 2008 review of the genus.[3]

The multigenus approach is based solely on the structure of the male flowers; no other characters could be consistently associated with one genus or another.[10]Four of the genera—Attalea(in a narrow sense),Orbignya,MaximilianaandScheelea—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, includingA. colenda(which Glassman placed in its own genus,Ynesa) and this has been used as an argument for the single-genus approach. The fact that there are several hybrids between species that would be considered different genera under Glassman's five-genus system was also used as an argument for placing them in a single genus.[3]Molecular phylogenetic work byAlan Meerowand colleagues concluded that multi-genus approach did not producemonophyleticgroups, but treatingAttaleaas a single genus did.[6]

Cintia Freitas and colleagues identified three main clades within the genus based on the nuclearWRKY gene family.The first of these, a group of species from the coastalAtlantic Forestregion in Brazil all of which had been placed inAttaleain the narrow sense, was termed theAttalea-like clade. This group was a sister to the other two clades. The second group, which they called theScheelea-like clade, consisted of most of the species formerly placed inScheelea,together with several that had been placed inAttalea(narrowly defined) andOrbigyna.The third group consisted mainly of species formerly placed inOrbigynaandMaximiliana;they called this theOrbigyna-like clade. Despite the existence of three well-supported clades, Freitas and colleagues concluded that the concept ofAttaleaas a single genus was best supported by their evidence.[6]

History

[edit]

The genusAttaleawas first describedCarl Sigismund Kunthin 1816 based on specimens collected byAlexander von HumboldtandAimé Bonpland,[1]although older, pre-Linnaeandescriptions exist, includingCharles Plumier's 1703 description ofA. crassispatha.[11]The genus was named forAttalus III Philometor,king ofPergamon,known for his interest inmedicinal plants.[2]Thetype speciesisA. amygdalina,a Colombianendemic.[12]The generaMaximilianaandOrbignyawere described byCarl Friedrich Philipp von Martiusin 1826[13]and 1837[14]respectively.Scheeleawas described byHermann Karstenin 1857,[15]andYnesabyOrator F. Cookin 1942.[16]

Species

[edit]
Main article:List of Attalea species
Five young palm trees planted together in a group, with a wooden bench below them: The trunks of the palms are marked with alternative pale and dark rings, and are only one-quarter to one-half the length of the leaves.
Attalea crassispatha,a Haitianendemic,is the most geographically isolated species in the genus.

Experts disagree about the number of species in the genusAttalea(broadly defined). In 1965, Dutch taxonomistJan Gerard Wessels Boerestimated that as many as 100 species may be in the genus. In their 1996Field Guide to the Palms of the AmericasAndrew 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 2005World 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 betweenmorphologicallysimilar species. This problem is complicated by the fact that many of these species are poorly represented inherbariumcollections. The large size of the leaves, inflorescences and fruit of manyAttaleaspecies makes them difficult to collect. In addition, many important collections, includingtype specimens,have been lost or destroyed.[3]Sparse or incomplete collections make it difficult to differentiate variation within a single species from variation between different species.[3][10]Taxonomic uncertainty is exacerbated by frequent hybridisation between species.[6]

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,[17]other authors have considered aspecies complexconsisting of four or five species. Glassman doubted the validity ofA. attaleoidesas a species, and described four new species from material that had previously been attributed toA. attaleoides—A. camopiensis,A. degranvillei,A. guianensisandA. maripensis.Govaerts and Dransfield accepted both Glassman's four species andA. attaleoides.However, Jean-Christophe Pintaud was of the opinion thatA. guianensis,A. maripensisandA. attaleoideswere all very similar, and thought it likely that they all represented the same species.[3]

Another species complex inAttaleaincludesA. speciosaand related species. Henderson (1995) recognisedA. speciosaandA. spectabilis,considering the latter to either be an acaulescent form ofA. speciosaor a hybrid betweenA. microcarpaand it. Govaerts and Dransfield acceptedA. spectabilis,but Glassman considered it adubioustaxon.Attalea vitrivirwas recognised as a distinct species by Michael Balick and coauthors;[18]Glassman and Govaerts and Dransfield concurred, but Henderson considered it part ofA. speciosa.Glassman also described a fourth member of this group,A. brejinhoensis,and it is accepted by Govaerts and Dransfield.[3]

Reproduction and growth

[edit]
Male inflorescence ofAttaleasp. swarming with insects.
Attalea sp.–MHNT

Attaleaspecies aremonoecious—male and female flowers are separate, but are borne by the same plant.[2]Various species have been described as being insect-pollinated, includingA. phalerata,[19]while pollination inA. colendaandA. speciosa,has been attributed both to insects and wind.[20][21]The fruit areanimal-dispersed.[2]

Seedgerminationisremote tubular[22]—during germination, as thecotyledonexpands it pushes the young shoot away from the seed.[23]After germination, the stem initially grows downward before turning to grow upward and produce the aboveground stem.[24]This produces a "saxophone shaped" belowground portion of the stem.[25]The fact that theshoot tipsofAttaleaseedlings are underground it likely to contribute to their fire-tolerance.[2]

Distribution

[edit]

Species range across the Neotropics from Mexico in the north to Bolivia, Paraguay, and southern Brazil in the south,[3]from low elevations in coastal Brazil to up to 1,600 m (5,200 ft) above sea level in the Andes.[6]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.[26]

Habitat and ecology

[edit]
Scarlet macawfeeding onAttaleafruit.

Attaleaincludes 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.[3]

Disturbance has been implicated in the formation of vegetation dominated by largeAttaleaspecies.[2]In seasonally dry Amazonian forests, the density of large adultA. maripapalms was correlated with canopy openness;[25]the species also dominates savannas formed by repeated forest fires in Trinidad and Tobago.[27]A. speciosaforms pure stands in many parts of Brazil where natural forest vegetation has been cleared.[28]Similarly, stands ofA. funiferainBahia,Brazil (which are cultivated forpiassavafibre) are managed using fire—the seedlings survive cutting and burning, and are able to dominate burned forest patches.[29]

The fruit are dispersed by animals; fruit which are not dispersed frequently suffer seed predation bybruchid beetles.[2]Certain species ofAttaleahave been mentioned as examples of anachronistic species which are adapted for dispersal by now-extinctPleistocene megafauna.[30][31]On Maracá Island,Roraima,in theBrazilian Amazon,A. maripafruit were consumed bytapirs,collared peccaries,deer, and primates. Rodents, includingagoutis,fed upon the fruit, and as the fruit availability declined, they fed on the seeds.[32]Other dispersers ofAttaleafruit includecrested caracaras,which consume the fruit and disperse the seeds ofA. phaleratain the BrazilianPantanal.[33]

Uses

[edit]

Attaleaspecies have a long history of human use. CarbonisedAttalea maripaseeds have been found in archaeological sites in Colombia dating back to 9000BP.[34]Several species remain important sources ofedible oil,thatch,edible seeds, and fibre. The leaves ofAttalea butyraceaandA. maripaare used extensively for thatching. Several species areoil palms,withA. speciosaamong the most important economically.[2]Products extracted fromA. speciosawere reported to support over 300,000 households in the Brazilian state ofMaranhãoin 2005,[35]and in 1985 it was estimated to support over 450,000 households throughout the Brazil.[28]Piassava fibres, extracted from the leaf bases ofA. funifera,are commercially important,[2]and generated aboutUS$20 million in annual income to Brazilian farmers in 1996.[36]

References

[edit]
  1. ^ab"AttaleaKunth in F.W.H.von Humboldt, A.J.A.Bonpland & C.S.Kunth, Nov. Gen. Sp. 1: 309 (1816) ".World Checklist of Selected Plant Families.Archived fromthe originalon 2020-08-05.Retrieved2007-02-24.
  2. ^abcdefghijklmnHenderson, Andrew;Gloria Galeano(1995).Field Guide to the Palms of the Americas.Princeton, New Jersey: Princeton University Press. pp. 153–164.ISBN0-691-08537-4.
  3. ^abcdefghijklPintaud, Jean-Christophe (2008)."An overview of the taxonomy of Attalea (Arecaceae)"(PDF).Revista Peruana de Biología.15(supl. 1): 055–063.
  4. ^abMeerow, Alan W.;Larry Noblick; James W. Borrone; Thomas L. P. Couvreur; Margarita Mauro-Herrera; William J. Hahn; David N. Kuhn; Kyoko Nakamura; Nora H. Oleas; Raymond J. Schnell (2009). Joly, Simon (ed.)."Phylogenetic Analysis of Seven WRKY Genes across the Palm Subtribe Attaleinae (Arecaceae) Identifies Syagrus as Sister Group of the Coconut".PLOS ONE.4(10): e7353.Bibcode:2009PLoSO...4.7353M.doi:10.1371/journal.pone.0007353.PMC2752195.PMID19806212.
  5. ^Dransfield, John; Natalie W. Uhl; Conny B. Asmussen; William J. Baker; Madeline M. Harley; Carl E. Lewis (2005). "A New Phylogenetic Classification of the Palm Family, Arecaceae".Kew Bulletin.60(4): 559–69.JSTOR25070242.
  6. ^abcdeFreitas, Cintia; Meerow, Alan W.; Pintaud, Jean-Christophe; Henderson, Andrew; Noblick, Larry; Costa, Flavia R. C.; Barbosa, Carlos E.; Barrington, David (2016)."Phylogenetic analysis of Attalea (Arecaceae): insights into the historical biogeography of a recently diversified Neotropical plant group".Botanical Journal of the Linnean Society.182(2): 287–302.doi:10.1111/boj.12466.
  7. ^Dransfieldet al.listed this genus as a synonym ofAllagoptera,butMeerowet al.recognised it as a distinct genus in their analysis.
  8. ^Glassman, Sidney F. (1999)."A Taxonomic Treatment of the Palm SubtribeAttaleinae(TribeCocoeae) ".Illinois Biological Monographs.59:1–414.
  9. ^"WCSP".World Checklist of Selected Plant Families.Archivedfrom the original on May 17, 2008.Retrieved2010-04-02.
  10. ^abAnderson, Anthony B.; Michael J. Balick (1988)."Taxonomy of the Babassu Complex (Orbignyaspp.: Palmae) "(PDF).Systematic Botany.13(1): 32–50.doi:10.2307/2419239.JSTOR2419239.
  11. ^Henderson, Andrew; Michael Balick (1991). "Attalea crassispatha, a rare and endemic Haitian palm".Brittonia(JSTORsubscription required).43(3): 189–194.doi:10.2307/2807059.JSTOR2807059.S2CID12177917.
  12. ^"Attalea amygdalina".World Checklist of Selected Plant Families.Retrieved2010-02-17.
  13. ^"Maximiliana".World Checklist of Selected Plant Families.Retrieved2010-02-17.
  14. ^"Orbignya".World Checklist of Selected Plant Families.Retrieved2010-02-17.
  15. ^"Scheelea".World Checklist of Selected Plant Families.Retrieved2010-02-17.
  16. ^"Ynesa".World Checklist of Selected Plant Families.Retrieved2010-02-17.
  17. ^Henderson, Andrew (1995).The palms of the Amazon.Oxford University Press.ISBN978-0-19-508311-8.
  18. ^Balicket al..1987, Anderson and Balick 1988
  19. ^Moraes R., Mónica; Finn Borchsenius; Ulla Blicher-Mathiesen (1996). "Notes on the Biology and Uses of the Motacú Palm (Attalea phalerata,Arecaceae) from Bolivia ".Economic Botany.50(4): 423–428.doi:10.1007/BF02866525.JSTOR4255886.S2CID19066490.
  20. ^Feil, Jan Peter (1996). "Fruit Production of Attalea colenda (Arecaceae) in Coastal Ecuador: An Alternative Oil Resource?".Economic Botany.50(3): 300–309.doi:10.1007/BF02907337.JSTOR4255847.S2CID22885187.
  21. ^Anderson, Anthony B.; William L. Overal; Andrew Henderson (1988). "Pollination Ecology of a Forest-Dominant Palm (Orbignya phalerataMart.) in Northern Brazil ".Biotropica.20(3): 192–205.doi:10.2307/2388234.JSTOR2388234.
  22. ^Uhl, Natalie E.; John Dransfield (1987).Genera Palmarum: a classification of palms based on the work of Harold E. Moore Jr.Lawrence, Kansas: The L. H. Bailey Hortorium and the International Palm Society. pp. 503–512.ISBN0-935868-30-5.
  23. ^Pinheiro, Claudio Urbano B. (2001). "Germination strategies in palms: the case ofSchippia concolorin Belize ".Brittonia.53(4): 519–527.doi:10.1007/BF02809652.S2CID45335410.
  24. ^Salm, Rudolfo (2004)."Tree species diversity in a seasonally-dry forest: the case of the Pinkaití site, in the Kayapó Indigenous Area, Southeastern limits of the Amazon".Acta Amazonica.34(3): 435–443.doi:10.1590/S0044-59672004000300009.
  25. ^abSalm, Rudolfo (2005)."The importance of forest disturbance for the recruitment of the large arborescent palm Attalea maripa in a seasonally dry Amazonian forest".Biota Neotropica.5(1).
  26. ^Govaerts, R.; J. Henderson; S.F. Zona; D.R. Hodel; A. Henderson (2010)."World Checklist of Arecaceae".The Board of Trustees of theRoyal Botanic Gardens, Kew.Archived fromthe originalon February 21, 2007.Retrieved2010-02-27.
  27. ^Beard, J. S.(1953). "The Savanna Vegetation of Northern Tropical America".Ecological Monographs.23(2): 149–215.doi:10.2307/1948518.JSTOR1948518.
  28. ^abMay, Peter H.; Anthony B. Anderson; Michael J. Balick; José Mário F. Frazão (1985)."Subsistence Benefits from the Babassu Palm (Orbignya martiana)"(PDF).Economic Botany.39(2): 113–129.doi:10.1007/BF02907831.JSTOR4254727.S2CID43133375.
  29. ^Voeks, Robert A.; Sergio G. da Vinha (1988)."Fire Management of the Piassava Fiber Palm (Attalea funifera) in Eastern Brazil "(PDF).Yearbook, Conference of Latin Americanist Geographers.14:7–13. Archived fromthe original(PDF)on 2011-07-20.Retrieved2010-03-03.
  30. ^Janzen, Daniel H.; Paul S. Martin (1982)."Neotropical Anachronisms: The Fruit the Gomphotheres Ate".Science.215(4528): 19–27.Bibcode:1982Sci...215...19J.doi:10.1126/science.215.4528.19.PMID17790450.S2CID19296719.[permanent dead link]
  31. ^Guimarães, Paola R. Jr.; Mauro Galetti; Pedro Jordano (2008). Hansen, Dennis Marinus (ed.)."Seed Dispersal Anachronisms: Rethinking the Fruits Extinct Megafauna Ate".PLOS ONE.3(3): e1745.Bibcode:2008PLoSO...3.1745G.doi:10.1371/journal.pone.0001745.PMC2258420.PMID18320062.
  32. ^Fragoso, Jose M. V. (1997). "Tapir-Generated Seed Shadows: Scale-Dependent Patchiness in the Amazon Rain Forest".Journal of Ecology.85(4): 519–29.doi:10.2307/2960574.JSTOR2960574.
  33. ^Galetti, Mauro; Paulo R. Guimarães Jr. (2004)."Seed dispersal ofAttalea phalerata(Palmae) by Crested caracaras (Caracara plancus) in the Pantanal and a review of frugivory by raptors "(PDF).Ararajuba.12(2): 133–135.[permanent dead link]
  34. ^Morcote-Ríos, Gaspar;Rodrigo Bernal(2001). "Remains of palms (Palmae) at archaeological sites in the New World: A review".Botanical Review.67(3): 309–50.doi:10.1007/BF02858098.S2CID46582757.
  35. ^Porro, Roberto (2005). "Palms, Pastures, and Swidden Fields: The Grounded Political Ecology of" Agro-Extractive/Shifting-cultivator Peasants "in Maranhão, Brazil".Human Ecology.33(1): 17–56.doi:10.1007/s10745-005-1654-2.S2CID153617434.
  36. ^Voeks, Robert A. (2002). "Reproductive ecology of the piassava palm (Attalea funifera) of Bahia, Brazil ".Journal of Tropical Ecology.18:121–136.doi:10.1017/s0266467402002079.S2CID84211050.
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