Aiphanes
Aiphanes | |
---|---|
Aiphanes horrida at Jena botanical garden | |
Scientific classification | |
Kingdom: | Plantae |
Clade: | Tracheophytes |
Clade: | Angiosperms |
Clade: | Monocots |
Clade: | Commelinids |
Order: | Arecales |
Family: | Arecaceae |
Subfamily: | Arecoideae
|
Tribe: | Cocoseae |
Genus: | Aiphanes Willd. |
Type species | |
Aiphanes horrida | |
Diversity | |
About 26 species | |
Native distribution of Aiphanes | |
Synonyms[1] | |
Martinezia (sensu |
Aiphanes is a
Carl Ludwig Willdenow coined the name Aiphanes in 1801. Before that, species belonging to the genus had been placed in Bactris or Caryota. The name Martinezia had also been applied to the genus, and between 1847 and 1932 it was generally used in place of Aiphanes. Max Burret resurrected the name Aiphanes in 1932 and laid the basis for the modern concept of the genus. Aiphanes is most closely related to several other genera of spiny palms—Acrocomia, Astrocaryum, Bactris and Desmoncus. Two species are widely planted as ornamentals and the fruit, seeds or palm heart of several species have been eaten by indigenous peoples of the Americas for millennia.
Description
Aiphanes is a genus of spiny palms ranging from 20-metre (66 ft) tall subcanopy trees to small shrubs with subterranean stems growing in the forest understorey.[3] Its name combines the Ancient Greek ai, meaning "always", with phaneros, meaning "evident", "visible" or "conspicuous".[4] In their 1996 monograph on the genus, botanists Finn Borchsenius and Rodrigo Bernal pointed out that "ironically, species of Aiphanes are generally very hard to spot and find in dense vegetation and, accordingly, are among the most poorly collected neotropical palms".[4]
Stems
While some species are single-stemmed, others form multi-stemmed (
Two species are characterised by an acaulescent growth habit—A. acaulis and A. spicata. Two other species—A. ulei and A. weberbaueri—occur in both acaulescent populations and those which produce above-ground stems. Several species are single-stemmed understorey palms, an unusual growth form. Aiphanes grandis and A. minima are single-stemmed palms which grow to be more than 10 metres (33 ft) tall, while the remainder are multi-stemmed understorey species. Multi-stemmed palms range from plants with a single main stem and a few basal suckers to caespitose clumps of 20 densely packed stems. A variety of growth forms can exist within a single species and this appears to be influenced by habitat and environmental conditions.[3]
Leaves
The leaves of Aiphanes species are usually pinnately divided—rows of leaflets emerge on either side of the axis of the leaf in a feather-like or fern-like pattern. The sole exception to this is
Leaves are spiny but the degree varies both within and among species. Leaf sheaths are always densely spiny, but the spines usually become smaller and sparser towards the ends of the leaves.[5]
Spines
Spines are characteristic of Aiphanes and other members of the subtribe Bactridinae. They are found almost everywhere on the plants and are especially well-developed on the stem, leaf bases, and the peduncle. In Aiphanes, the spines are formed from the outer tissues of the plant and are not derived through the modification of other plant organs. They range from less than 1 millimetre (0.04 in) to more than 25 centimetres (9.8 in) long.[6]
Flowers
Aiphanes species are pleonanthic—they flower repeatedly over the course of their lifespan—and
Flowers are usually borne in groups of three—one female flower together with two male flowers. In some species groups of four flowers (two male and two female) have been reported. At the far end of the inflorescence, away from the axis of the tree, pairs of male flowers replace the triads of male and female flowers. Flower colour is poorly known. It must be recorded from live plants, since preserved flowers lose their colour over time, and records of these species in the wild are incomplete. Male flowers tend to fall into two groups—those with cream or yellow flowers and those with some amount of purple in the flowers. Female flowers are even less well known than male flowers.[7]
Pollen grains are usually spherical to ellipsoid in shape, sometimes triangular, about 20 to 30 micrometres along their long axis and 20 to 30 μm in diameter. They are typically
Fruit
The fruit of Aiphanes species is usually a red, spherical, single-seeded
Karyotype
Published
Taxonomy
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Relationship between members of the subtribe Bactridinae, based on plastid DNA phylogeny.[13] *Aiphanes aculeata is a botanical synonym of A. horrida. |
Aiphanes is placed in the
In his 1932 revision of the genus, German botanist Max Burret recognised 32 species. Seventeen of these were new species, mostly based on collections made by German botanist Wilhelm Kalbreyer in northern Colombia between 1877 and 1881. Working with a very narrow species concept, and not being familiar with the variation present in natural populations, Burret placed almost every specimen into a distinct species. The bombing of the Berlin Herbarium during the Second World War destroyed the only known collections for 13 of these 32 species, further complicating the situation.[15]
The
Burret divided Aiphanes into two subgenera, Brachyanthera and Macroanthera. Eleven species were placed in Macroanthera, while the remainder were placed in Brachyanthera.[15] In their 1996 monograph, Borchsenius and Bernal questioned the applicability of these subgenera. They recognised that if Macroanthera was reduced to three species (A. horrida, A. eggersii and A. minima) it could form a viable grouping, but that this would leave Brachyanthera overly heterogeneous. Consequently, they abandoned Burret's use of subgenera.[19]
In the three decades following Burret's delimitation of the genus a further 15 species were described, bringing the total species count to 47.
Burret divided Aiphanes into two subgenera, Brachyanthera and Macroanthera. Eleven species were placed in Macroanthera, with the remainder in Brachyanthera.[15] In their 1996 monograph, Borchsenius and Bernal questioned the applicability of these subgenera. They recognised that if Macroanthera was reduced to three species (A. horrida, A. eggersii and A. minima) it could form a viable grouping, but that this would leave Brachyanthera overly heterogeneous. Consequently, they abandoned Burret's use of subgenera.[19]
History
The earliest botanical description of a species in the genus was made by French botanist Charles Plumier, who described two species based on his visits to the West Indies between 1689 and 1695. Both of Plumier's species are now considered to be Aiphanes minima. The same species was described by Dutch botanist Nikolaus Joseph von Jacquin in 1763. Spanish botanist José Celestino Mutis produced a detailed description of A. lindeniana and illustrations of that species and what is thought to be A. horrida in 1779.[4]
In 1791 Joseph Gaertner included a species of Aiphanes in his De Fructibus et Seminibus Plantarum, calling it Bactris minima. This is the oldest validly published name for any member of the genus. The name Aiphanes was coined by German botanist Carl Ludwig Willdenow in 1801. He described a single species, A. aculeata, in 1806.[4]
Jacquin had used the name Caryota horrida to describe a plant that belonged to the same species (and may have been the same individual) described by Willdenow. Borchsenius and Bernal cite an 1809 publication date for Jacquin's description, which gave precedence to Willdenow's name.[4] However, the more recent World Checklist (2006) gives an 1801 publication date for Jacquin's description, making A. horrida the correct name for the species.[23]
In 1816 Alexander von Humboldt, Aimé Bonpland and Carl Sigismund Kunth described Martinezia caryotifolia, adding another name to the list of synonyms for A. horrida. Since the original diagnostic characters of Martinezia did not fit any existent species, it was redefined by Kunth to fit M. caryotifolia. Consequently, Martinezia came to replace Aiphanes and the latter name was rarely used between 1847 and 1932. In 1857 Hermann Karsten created a new genus, Marara, to accommodate two Colombian species, M. bicuspidata (later shown to be a synonym for A. horrida) and M. erinacea (now A. erinacea). Hermann Wendland attempted to resurrect Aiphanes in 1878, merging Martinezia and Marara into it, but his proposal was ignored. In 1901 Orator F. Cook created two new genera—Curima, into which he put A. minima, and Tilmia, which housed A. horrida.[24] In 1932, after publishing a species in Martinezia, Burret changed his mind about the genus and synonymised it with Aiphanes. This led to the current delimitation of the genus.[15]
Species
Species accepted by the World Checklist of Selected Plant Families:[2]
- Aiphanes acanthophylla(Mart.) Burret – Puerto Rico, Dominican Republic
- Aiphanes acaulis Galeano & R.Bernal – Colombia
- Aiphanes argos R.Bernal, Borchs. & Hoyos-Gómez – Colombia[25]
- Aiphanes bicornis Cerón & R.Bernal – Ecuador
- Aiphanes buenaventurae R.Bernal & Borchs. – Valle del Cauca in Colombia
- Aiphanes chiribogensis Borchs. & Balslev – Ecuador
- Aiphanes deltoidea Burret – Colombia, Peru, northwestern Brazil
- Aiphanes duquei Burret – Colombia
- Aiphanes eggersii Burret – Ecuador, Peru
- Aiphanes erinacea (H.Karst.) H.Wendl. – Colombia, Ecuador
- Aiphanes gelatinosa H.E.Moore – Colombia, Ecuador
- Aiphanes graminifolia Galeano & R.Bernal – Colombia
- Aiphanes grandis Borchs. & Balslev – Ecuador
- Aiphanes hirsuta Burret – Colombia, Ecuador, Panama, Costa Rica
- Aiphanes horrida (Jacq.) Burret – Trinidad, Colombia, Venezuela, Peru, northwestern Brazil, Bolivia
- Aiphanes leiostachys Burret – Antioquia in Colombia
- Aiphanes lindeniana (H.Wendl.) H.Wendl. – Colombia
- Aiphanes linearis Burret – Antioquia and Valle del Cauca in Colombia
- Aiphanes macroloba Burret – Colombia, Ecuador
- Aiphanes minima (Gaertn.) Burret – Saint Lucia, Barbados
- Aiphanes multiplex R.Bernal & Borchs. – Valle del Cauca in Colombia
- Aiphanes parvifolia Burret – Colombia
- Aiphanes pilaris R.Bernal – Colombia
- Aiphanes simplex Burret – Colombia
- Aiphanes spicata Borchs. & R.Bernal – Peru
- Aiphanes stergiosii S.M.Niño – State of Portuguesa in western Venezuela
- Aiphanes tricuspidata Borchs., M.Ruíz & Bernal – Colombia, Ecuador
- Aiphanes ulei (Dammer) Burret – Colombia, Ecuador, Peru, northwestern Brazil
- Aiphanes verrucosa Borchs. & Balslev – Ecuador
- Aiphanes weberbaueri Burret – Ecuador, Peru
Distribution and status
The genus Aiphanes ranges from Hispaniola (the Dominican Republic) and Panama in the north, to Trinidad and Tobago in the east, across Colombia and down along the Andes to Bolivia. In Brazil it only occurs along the border with Peru. Aiphanes is primarily South American—one species (A. hirsuta) is present in Panama and two others (A. horrida and A. minima) are found in the Caribbean. Aiphanes minima, which is endemic to the insular Caribbean, is the only species absent from the South American mainland. Although A. horrida has been reported from Guyana and southern Venezuela these reports have not been verified with herbarium vouchers.[26]
Aiphanes horrida is the most widely distributed species. It ranges from
Habitat and ecology
Aiphanes species are palms of the forest understorey and subcanopy. The most widely distributed species, A. horrida, occurs both in
Records of visits by
The fruit of A. horrida is rich in vitamins and energy and likely to be eaten by many animals.
Several species show clumped distributions. Dispersal limitation has been invoked to explain the clumped distribution of adults and limited recruitment of seedlings in both A. erinacea in Ecuador[39] and A. minima in Puerto Rico.[38] Similarly, the rarity of A. lindeniana and A. simplex in Colombian forests may be linked to limited seed production and the limited effectiveness of seed dispersal by avian and mammalian frugivores.[40]
Uses
Aiphanes species have a long history of human use. The remains of carbonised seeds thought to belong to A. horrida have been found in archaeological sites in
Notes
- ^ Borchsenius & Bernal (1996), pp. 33–34
- ^ a b Kew World Checklist of Selected Plant Families
- ^ a b c d Borchsenius & Bernal (1996), p. 4
- ^ a b c d e Borchsenius & Bernal (1996), p. 2
- ^ a b Borchsenius & Bernal (1996), pp. 8–9
- ^ Borchsenius & Bernal (1996), pp. 6–8
- ^ a b Borchsenius & Bernal (1996), pp. 11–14
- ^ a b Borchsenius & Bernal (1996), pp. 16–22
- ^ Hoen, Peter (1999). "Glossary of Pollen and Spore Terminology". Laboratory of Palaeobotany and Palynology, Michigan Technological University. Archived from the original on 18 May 2010. Retrieved 17 March 2010.
- S2CID 28497230.
- ^ a b Borchsenius & Bernal (1996), pp. 14–15
- ^ Borchsenius & Bernal (1996), p. 26
- .
- JSTOR 25070242.
- ^ a b c d e Borchsenius & Bernal (1996), p. 3
- ^ "Article 7". International Code of Botanical Nomenclature (Vienna Code). Retrieved 20 March 2010.
- ^ JSTOR 1220905.
- ^ Borchsenius & Bernal (1996)
- ^ a b Borchsenius & Bernal (1996), p. 33
- ^ George R. Proctor, in Acevedo-Rodríguez & Strong (2005), pp. 138–139
- ^ Borchsenius & Bernal (1996), p. 87
- ^ Govaerts, R.; J. Henderson; S.F. Zona; D.R. Hodel; A. Henderson (2006). "Search for Aiphanes". World Checklist of Arecaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. Retrieved 2010-01-05.
- ^ "Aiphanes horrida". Royal Botanic Gardens, Kew: World Checklist of Selected Plant Families. Retrieved 2010-01-04.
- ^ Borchsenius & Bernal (1996), pp. 2–3
- ^ "Aiphanes argos". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2 March 2023.
- ^ a b c Borchsenius & Bernal (1996), pp. 26–30
- ^ Montúfar, R.; N. Pitman (2003). "Aiphanes grandis". IUCN Red List of Threatened Species. 2003: e.T43757A10826566. Retrieved 4 February 2020.
- ^ Bernal, R. (1998). "Aiphanes leiostachys". IUCN Red List of Threatened Species. 1998: e.T38942A10158301. Retrieved 4 February 2020.
- ^ Montúfar, R.; N. Pitman (2003). "Aiphanes verrucosa". IUCN Red List of Threatened Species. 2003: e.T38069A10095781. Retrieved 4 February 2020.
- ^ Montúfar, R.; N. Pitman (2003). "Aiphanes chiribogensis". IUCN Red List of Threatened Species. 2003: e.T38752A10147801. Retrieved 4 February 2020.
- ^ Bernal, R. (1998). "Aiphanes duquei". IUCN Red List of Threatened Species. 1998: e.T38941A10158248. Retrieved 4 February 2020.
- ^ Bernal, R. (1998). "Aiphanes lindeniana". IUCN Red List of Threatened Species. 1998: e.T38943A10158352. Retrieved 4 February 2020.
- .
- S2CID 34175807.
- S2CID 32827272.
- ^ a b Borchsenius & Bernal (1996), pp. 30–32
- ^ Culzac-Wilson, Lystra (2005). Species Conservation Plan for the St. Vincent Parrot Amazona guildingii. Puerto de la Cruz, Tenerife: Loro Parque Fundación.
- ^ .
- S2CID 83543567.
- ^ Correa-Gómez, Diego F.; Orlando Vargas-Ríos (2009). "Regeneración de palmas in bosques nativos y plantaciones del Sanctuario de Fauna y Flora Otún-Quimbaya (Risaralda, Colombia)". Caldasia (in Spanish). 31 (2): 195–212. Archived from the original on 2011-09-27.
- S2CID 46582757.
- ^ Borchsenius & Bernal (1996), p. 47
- ISBN 0-691-08537-4.
- ISBN 978-0-88192-776-4.
- ^ Borchsenius & Bernal (1996), p. 52
- ^ Borchsenius & Bernal (1996), p. 54
- ^ Borchsenius & Bernal (1996), p. 69
- ^ Borchsenius & Bernal (1996), p. 72
- ^ Borchsenius & Bernal (1996), p. 71
- PMID 12956057.
- ^ Goodsell, David S. (2001-05-01). "Cyclooxygenase". RCSB Protein Data Bank. Archived from the original on 2009-11-23. Retrieved 2010-01-17.
References
- Acevedo-Rodríguez, Pedro; Mark T. Strong (2005). Monocotyledons and Gymnosperms of Puerto Rico and the Virgin Islands. Contributions from the United States National Herbarium. Vol. 52. Washington, D.C.: Smithsonian Institution. pp. 1–415.
- Borchsenius, Finn; JSTOR 4393869.