Aiphanes
Encyclopedia
Aiphanes is a genus
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 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
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...

 which is native to tropical
Tropics
The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately  N and the Tropic of Capricorn in the southern hemisphere at  S; these latitudes correspond to the axial tilt of the Earth...

 regions of South and Central America and the Caribbean. There are about 26 species in the genus, ranging in size from understorey
Understory
Understory is the term for the area of a forest which grows at the lowest height level below the forest canopy. Plants in the understory consist of a mixture of seedlings and saplings of canopy trees together with understory shrubs and herbs...

 shrubs with subterranean stems to subcanopy trees as much as 20 metres (65.6 ft) tall. Most have 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 compound leaves (leaves which are divided into leaflets arranged feather-like, in pairs along a central axis); one species has entire
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...

 leaves. Stems, leaves and sometimes even the fruit are covered with spines. Plants flower repeatedly over the course of their lifespan and have separate male and female flowers, although these are borne together on the same 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...

. Although records of pollinators are limited, most species appear to be pollinated by insects. The fruit are eaten by several birds and mammals, including at least two species of Amazon parrot
Amazon parrot
Amazon parrot is the common name for a parrot of the genus Amazona. These are medium-size parrots native to the New World ranging from South America to Mexico and the Caribbean....

s.

Carl Ludwig Willdenow
Carl Ludwig Willdenow
Carl Ludwig Willdenow was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants...

 coined the name Aiphanes in 1801. Before that, species belonging to the genus had been placed in Bactris
Bactris
Bactris is a genus of about 240 species in the palm family, Arecaceae, native to Central and South America, and the Caribbean. They are trees growing to 4-20 m tall. The leaves are up to 5 m long, and pinnate with numerous leaflets...

or Caryota
Caryota
Caryota is a genus of palm trees. They are often known as fishtail palms because of the shape of their leaves. There are about 13 species native to Asia and the South Pacific. One of the more widely known species is Caryota urens, which yields sap used to make an unrefined sugar called jaggery, and...

. 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
Max Burret
Karl Ewald Maximilian Burret, commonly known as Max Burret was a German botanist.Burret was born in Saffig near Andernach in the Prussian Rhine Province. He originally studied law at Lausanne and Munich at the instigation of his father...

 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
Acrocomia
Acrocomia aculeata is a species of palm native to tropical regions of the Americas, from southern Mexico and the Caribbean south to Paraguay and northern Argentina. Common names include Grugru Palm, Macaúba Palm, Coyol Palm, and Macaw Palm; synonyms include A. lasiospatha, A. sclerocarpa, A. totai,...

, Astrocaryum
Astrocaryum
Astrocaryum is a genus of about 36 to 40 species of palms native to Central and South America and Trinidad.-Description:Astrocaryum is a genus of spiny palms 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. Some...

, Bactris and Desmoncus
Desmoncus
Desmoncus is a genus of spiny palms native to the Neotropics. The genus extends from Mexico in the north to Brazil and Bolivia in the south, with two species, D. orthacanthos and D...

. Two species are widely planted as ornamentals
Ornamental plant
Ornamental plants are plants that are grown for decorative purposes in gardens and landscape design projects, as house plants, for cut flowers and specimen display...

 and the fruit, seeds or palm heart
Heart of palm
Heart of palm, also called palm heart, palmito, burglar's thigh, chonta, palm cabbage or swamp cabbage, is a vegetable harvested from the inner core and growing bud of certain palm trees Heart of palm, also called palm heart, palmito, burglar's thigh, chonta, palm cabbage or swamp cabbage, is a...

 of several species have been eaten by indigenous peoples of the Americas for centuries.

Description

Aiphanes is a genus of spiny palms ranging from 20 metres (65.6 ft) tall subcanopy trees to small shrubs with subterranean stems growing in the forest understorey. Its name combines the Ancient Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 ai, meaning "always", with phaneros, meaning "evident", "visible" or "conspicuous". In their 1996 monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...

 on the genus, botanists Finn Borchsenius 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...

 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".

Stems

While some species are single-stemmed, others for multi-stemmed (caespitose) clumps. Coupled with variation in stem size, this produces a diversity of growth forms in the genus—solitary (single-stemmed) palms that grow into the subcanopy of the forest, solitary or caespitose palms that grow in the forest understorey and acaulescent palms which lack an aboveground stem.

Two species are characterised by an acaulescent growth habit—A. acaulis
Aiphanes acaulis
Aiphanes acaulis is a spiny palm endemic to western Colombia. it is acaulescent, as the specific epithet "acaulis" implies, – that is, the stem is short to the extent that it is difficult to see at all, and mostly subterranean. Plants have 8 to 10 leaves which are up to 1.5 metres long...

and A. spicata. Two other species—A. ulei
Aiphanes ulei
Aiphanes ulei is a species of flowering plant in the Arecaceae family.It is found in Brazil, Ecuador, and Peru.-References:* Henderson, A. 1998. . Downloaded on 20 July 2007....

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
Aiphanes grandis
Aiphanes grandis is a species of flowering plant in the Arecaceae family.It is found only in Ecuador.Its natural habitats are subtropical or tropical moist lowland forests and subtropical or tropical moist montanes.It is threatened by habitat loss....

and A. minima
Aiphanes minima
Aiphanes minima is a spiny palm tree which is native to the insular Caribbean from Hispaniola to Grenada, and widely cultivated elsewhere. Usually tall, it sometimes grows as an understorey tree and only in height.-Description:...

are single-stemmed palms which grow to be more than 10 metres (32.8 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 sucker
Basal shoot
A basal shoot, root sprout, adventitious shoot, water sprout or sucker is a shoot or cane which grows from a bud at the base of a tree or shrub or from its roots. This shoot then becomes, or takes the form of, a singular plant. A plant that produces suckers is referred to as surculose...

s 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.

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 A. macroloba which has entire leaves. They are usually spirally arranged, but some species have a distichous leaf arrangement, a condition that is normal in palm seedlings but uncommon among adults. Old leaf bases detach cleanly from the stem, except in A. hirsuta subsp. fosteriorum
Aiphanes hirsuta
Aiphanes hirsuta is a species of flowering plant in the Arecaceae family.It is found in Colombia, Costa Rica, Ecuador, and Panama.-References:* Bernal, R. 1998. . Downloaded on 20 July 2007....

, which often has old leaf bases attached to the newer portions of the stem.

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.

Spines

Spines
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...

 are characteristic of Aiphanes and other members of the subtribe
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...

 Bactridinae. They are found almost everywhere on the plants and are especially well-developed on the stem, leaf bases, and 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...

. 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 (0.0393700787401575 in) to more than 25 centimetres (9.8 in) long.

Flowers

Aiphanes species are pleonanthic—they flower repeatedly over the course of their lifespan—and monoecious
Plant 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....

, meaning that there are separate male and female flowers, but individuals plants bear both types of flowers. In Aiphanes, male and female flowers are borne together on the same 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...

. Usually only a single inflorescence is borne at each node
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...

, although A. gelatinosa often bears then in groups of three at a single node. The inflorescence usually consists of a main axis consisting of a peduncle and a 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...

. The rachis bears rachillae, which are smaller branches which themselves bear the flowers, while the peduncle is the main stalk connecting the rachis with the stem of the plant. In some species there is second-order branching—the rachillae themselves are branched and the flowers are borne on these branches.

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.

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 monosulcate, meridionosulcate or more rarely trichotomosulcate. The sulcus
Sulcus (anatomy)
A sulcus is a depression or fissure in the surface of an organ, especially the brain.-Elsewhere:* anterior interventricular sulcus* calcaneal sulcus* coronal sulcus* gingival sulcus* gluteal sulcus* interlabial sulci...

 is a furrow which runs along the surface of the pollen grain and is usually the site at which pollination occurs. Monosulcate pollen has a single furrow that runs along the pole of the pollen grain. Meridionosulcate pollen have a furrow that runs along the equator of the pollen grain. Trichotomosulcate pollen, on the other hand, has three furrows. The outer layer of the pollen is covered to a greater or lesser extent with ridges, spines or warts. This "sculpting" tends to be more pronounced in species that are fly-pollinated and less pronounced in those that are pollinated by beetles or bees.

Fruit

The fruit of Aiphanes species is usually a red, spherical, single-seeded drupe
Drupe
In botany, a drupe is a fruit in which an outer fleshy part surrounds a shell of hardened endocarp with a seed inside. These fruits develop from a single carpel, and mostly from flowers with superior ovaries...

. A thin skin (or epicarp), which can be either smooth or spiny, covers the fleshy mesocarp, which is typically orange and sweet. The mesocarp of A. horrida
Aiphanes horrida
Aiphanes horrida is a palm native to northern South America and Trinidad and Tobago. Aiphanes horrida is a solitary, spiny tree. In the wild it grows 3–10 metres tall tall with a stem diameter of 6–10 centimetres ; cultivated trees may be as much as 15 m tall with a 15 cm diameter...

has one of the highest reported carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

 contents of any plant product and is also rich in protein. The endocarp, which encases the seed, is brown or black and very hard at maturity. Seeds are light brown with a thin seed coat (or testa) and white endosperm
Endosperm
Endosperm is the tissue produced inside the seeds of most flowering plants around the time of fertilization. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This makes endosperm an important source of nutrition in human diet...

, which is sweet and tastes somewhat like coconut
Coconut
The coconut palm, Cocos nucifera, is a member of the family Arecaceae . It is the only accepted species in the genus Cocos. The term coconut can refer to the entire coconut palm, the seed, or the fruit, which is not a botanical nut. The spelling cocoanut is an old-fashioned form of the word...

.

Karyotype

Published chromosome counts
Karyotype
A karyotype is the number and appearance of chromosomes in the nucleus of an eukaryotic cell. The term is also used for the complete set of chromosomes in a species, or an individual organism.p28...

 exist for two species, Aiphanes minima and A. horrida; haploid chromosome counts vary from 15 to 18. Borchsenius and Bernal report that it is difficult to get accurate chromosome counts in palms and that differences in chromosome counts may reflect these difficulties.

Taxonomy

Aiphanes is placed in the subfamily
Family (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 Bactridinae, together with the genera Desmoncus
Desmoncus
Desmoncus is a genus of spiny palms native to the Neotropics. The genus extends from Mexico in the north to Brazil and Bolivia in the south, with two species, D. orthacanthos and D...

, Bactris
Bactris
Bactris is a genus of about 240 species in the palm family, Arecaceae, native to Central and South America, and the Caribbean. They are trees growing to 4-20 m tall. The leaves are up to 5 m long, and pinnate with numerous leaflets...

, Acrocomia
Acrocomia
Acrocomia aculeata is a species of palm native to tropical regions of the Americas, from southern Mexico and the Caribbean south to Paraguay and northern Argentina. Common names include Grugru Palm, Macaúba Palm, Coyol Palm, and Macaw Palm; synonyms include A. lasiospatha, A. sclerocarpa, A. totai,...

and Astrocaryum
Astrocaryum
Astrocaryum is a genus of about 36 to 40 species of palms native to Central and South America and Trinidad.-Description:Astrocaryum is a genus of spiny palms 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. Some...

.

In his 1932 revision of the genus, German botanist Max Burret
Max Burret
Karl Ewald Maximilian Burret, commonly known as Max Burret was a German botanist.Burret was born in Saffig near Andernach in the Prussian Rhine Province. He originally studied law at Lausanne and Munich at the instigation of his father...

 recognised 32 species. Seventeen of these were new species, mostly based on collections made by German botanist Wilhelm Kalbreyer
Guillermo Kalbreyer
Wilhelm Kalbreyer was a German plant collector who was sent by James Veitch & Sons of Chelsea, London to collect new plants in West Africa and South America....

 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.

The International Code of Botanical Nomenclature requires each species to be represented by a type collection. The destruction of Burret's type collections left many species only known from his original descriptions, which generally lacked illustrations. Other specimens (called neotypes) were designated to replace these, either by Rodrigo Bernal and colleagues in 1989 or by Borchsenius and Bernal in their 1996 monograph of the genus. Bernal and colleagues attempted to retrace Kalbreyer's travels in northern Colombia and collect specimens from as close as possible to the location of the original collections.

Burret divided Aiphanes into two subgenera, Brachyanthera and Macroanthera. Eleven species were placed in Macroanthera, while the remainder were placed in Brachyanthera. 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
Aiphanes eggersii
Aiphanes eggersii, known locally as Corozo, is a species of spiney, pinnately leaved palm which is endemic to the coastal plain of Ecuador.-Description:...

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.

In the three decades following Burret's delimitation of the genus a further 15 species were described, bringing the total species count to 47. Borchsenius and Bernal determined that many of these names were synonyms, although American botanist George Proctor disagreed with their decision to lump A. acanthophylla into A. minima. Borchsenius and Bernal also described one new species, Aiphanes spicata, bringing the total number of accepted species to 22. In two cases the destruction of the only known collections made it impossible to be absolutely certain that a name was a synonym. The current World Checklist of Selected Plant Families, maintained by Rafaël Govaerts at the Royal Botanic Gardens, Kew
Royal Botanic Gardens, Kew
The Royal Botanic Gardens, Kew, usually referred to as Kew Gardens, is 121 hectares of gardens and botanical glasshouses between Richmond and Kew in southwest London, England. "The Royal Botanic Gardens, Kew" and the brand name "Kew" are also used as umbrella terms for the institution that runs...

, recognises 26 species, including four species described since the publication of Borchsenius and Bernal's monograph.

Burret divided Aiphanes into two subgenera, Brachyanthera and Macroanthera. Eleven species were placed in Macroanthera, with the remainder in Brachyanthera. 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
Aiphanes eggersii
Aiphanes eggersii, known locally as Corozo, is a species of spiney, pinnately leaved palm which is endemic to the coastal plain of Ecuador.-Description:...

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.

History

The earliest botanical description of a species in the genus was made by French botanist 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...

, 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
Nikolaus Joseph von Jacquin
Nikolaus Joseph Freiherr von Jacquin or Baron Nikolaus von Jacquin. was a scientist who studied medicine, chemistry and botany....

 in 1763. Spanish botanist José Celestino Mutis
José Celestino Mutis
-External links:*** at The Catholic Encyclopedia official site...

 produced a detailed description of A. lindeniana
Aiphanes lindeniana
Aiphanes lindeniana is a species of palm that is endemic to Colombia. Although widespread in the Cordillera Occidental and Cordillera Central, it is threatened by habitat loss and forest management practices.-Description:...

and illustrations of that species and what is thought to be A. horrida in 1779.

In 1791 Joseph Gaertner
Joseph Gaertner
Joseph Gaertner was a German botanist, best known for his work on seeds, De Fructibus et Seminibus Plantarum ....

 included a species of Aiphanes in his De Fructibus et Seminibus Plantarum
De Fructibus et Seminibus Plantarum
De Fructibus et Seminibus Plantarum, also known by its standard botanical abbreviation Fruct. Sem. Pl., is a three-volume botanic treatise by Joseph Gaertner. The first volume was published in December 1788. The second volume was published in four parts, in 1790, 1791, 1791 and 1792 respectively...

, 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
Carl Ludwig Willdenow
Carl Ludwig Willdenow was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants...

 in 1801. He described a single species, A. aculeata, in 1806.

Jacquin had used the name Caryota
Caryota
Caryota is a genus of palm trees. They are often known as fishtail palms because of the shape of their leaves. There are about 13 species native to Asia and the South Pacific. One of the more widely known species is Caryota urens, which yields sap used to make an unrefined sugar called jaggery, and...

 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. 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.

In 1816 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...

, 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...

 and Carl Sigismund Kunth
Carl Sigismund Kunth
Carl Sigismund Kunth , also Karl Sigismund Kunth or anglicized as Charles Sigismund Kunth, was a German botanist...

 described Martinezia caryotifolia, adding another name to the list of synonyms
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

 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
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....

 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
Hermann Wendland
Hermann Wendland was a German botanist and gardener.He was a noted authority on the family Arecaceae , on which he published a major monograph which formed the basis for the modern classification of the family, including many of the generic names currently in use.The South American palm genus...

 attempted to resurrect Aiphanes in 1878, merging Martinezia and Marara into it, but his proposal was ignored. In 1901 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...

 created two new genera—Curima, into which he put A. minima, and Tilmia, which housed A. horrida. 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.

Distribution and status

The genus Aiphanes ranges from the Dominican Republic
Dominican Republic
The Dominican Republic is a nation on the island of La Hispaniola, part of the Greater Antilles archipelago in the Caribbean region. The western third of the island is occupied by the nation of Haiti, making Hispaniola one of two Caribbean islands that are shared by two countries...

 and Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...

 in the north, to 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...

 in the east, across 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...

 and down along the Andes
Andes
The Andes is the world's longest continental mountain range. It is a continual range of highlands along the western coast of South America. This range is about long, about to wide , and of an average height of about .Along its length, the Andes is split into several ranges, which are separated...

 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...

. In 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...

 it only occurs along the border with Peru
Peru
Peru , officially the Republic of Peru , is a country in western South America. It is bordered on the north by Ecuador and Colombia, on the east by Brazil, on the southeast by Bolivia, on the south by Chile, and on the west by the Pacific Ocean....

. Aiphanes is primarily South American—one species (A. hirsuta
Aiphanes hirsuta
Aiphanes hirsuta is a species of flowering plant in the Arecaceae family.It is found in Colombia, Costa Rica, Ecuador, and Panama.-References:* Bernal, R. 1998. . Downloaded on 20 July 2007....

) 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
Guyana
Guyana , officially the Co-operative Republic of Guyana, previously the colony of British Guiana, is a sovereign state on the northern coast of South America that is culturally part of the Anglophone Caribbean. Guyana was a former colony of the Dutch and of the British...

 and southern Venezuela these reports have not been verified with 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...

 vouchers.

Aiphanes horrida is the most widely distributed species. It ranges from Trinidad
Trinidad
Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...

 to Bolivia but is absent from Ecuador
Ecuador
Ecuador , officially the Republic of Ecuador is a representative democratic republic in South America, bordered by Colombia on the north, Peru on the east and south, and by the Pacific Ocean to the west. It is one of only two countries in South America, along with Chile, that do not have a border...

 and northern Peru. Other species have narrower ranges with one centre of diversity
Center of diversity
A center of diversity is an area that has a high degree of genetic variation for a particular species or genus of plants that can also be the center of origin for that species. The two areas often, but not always, coincide; the degree of coincidence remains the subject of debate...

 in western Colombia and Ecuador and another minor one in northeastern Peru. The 2006 IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

 includes three species which are endangered by habitat destruction—A. grandis, A. leiostachys
Aiphanes leiostachys
Aiphanes leiostachys is a species of palm that is endemic to Colombia. Known from only a few forest fragments in the Cordillera Central, it is threatened by habitat loss and forest management practices.-Description:...

and A verrucosa
Aiphanes verrucosa
Aiphanes verrucosa is a species of flowering plant in the Arecaceae family.It is found only in Ecuador.Its natural habitat is subtropical or tropical moist montanes.It is threatened by habitat loss.-References:...

—and three others considered vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

 to the same threat—A. chiribogensis
Aiphanes chiribogensis
Aiphanes chiribogensis is a species of palm which is endemic to western Ecuador. Its natural habitats are subtropical or tropical moist lowland forests and subtropical or tropical moist montane forests...

, A. duquei
Aiphanes duquei
Aiphanes duquei is a species of palm that is endemic to Colombia. Known from only a small area in the Cordillera Occidental, it is threatened by habitat loss and forest management practices.-Description:...

and A. lindeniana. Rodrigo Bernal and 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...

 expanded this list in a 2005 review of the status of Colombian palms. They listed two species as critically endangered—A. graminifolia, a species that was first described in 2002, and A. leiostachys (which was classified as endangered in the IUCN Red List). They classified two species as endangered—A. acaulis and A. parvifolia—and two species as vulnerable—A. gelatinosa and A. pilaris. They also classified six species as near threatened
Near Threatened
Near Threatened is a conservation status assigned to species or lower taxa that may be considered threatened with extinction in the near future, although it does not currently qualify for the threatened status...

A. erinacea, A. hirsuta, A. lindeniana (vulnerable according to the IUCN Red List), A. linearis
Aiphanes linearis
Aiphanes linearis is a species of flowering plant in the Arecaceae family.It is found only in Colombia.-References:* Bernal, R. 1998. . Downloaded on 20 July 2007....

, A. macroloba and A. simplex. The threats to these species were not listed, but Jens-Christian Svenning reported that A. erinacea was threatened by logging given its limited distribution and poor ability to regenerate in disturbed forests. In addition to these, A. deltoidea
Aiphanes deltoidea
Aiphanes deltoidea, known as shicashic in Peru, is a species of palm which is native to northeastern South America.-Description:Aiphanes deltoidea is a small palm tall with either a single stem or two large and several smaller stems, about in diameter...

, which is widely distributed across the western Amazon Rainforest
Amazon Rainforest
The Amazon Rainforest , also known in English as Amazonia or the Amazon Jungle, is a moist broadleaf forest that covers most of the Amazon Basin of South America...

, is present at such low densities that it was classified as a rare species by Francis Kahn and Farana Moussa in 1994.

Habitat and ecology

Aiphanes species are palms of the forest understorey and subcanopy. The most widely distributed species, A. horrida, occurs both in tropical dry forest and in more humid forest types, but there is a gap in its distribution which coincides with the wettest forests of the upper Amazon Basin
Amazon Basin
The Amazon Basin is the part of South America drained by the Amazon River and its tributaries that drains an area of about , or roughly 40 percent of South America. The basin is located in the countries of Bolivia, Brazil, Colombia, Ecuador, Guyana, Peru, and Venezuela...

. Two other species, A. minima and A. eggersii, are also found in drier environments; A. eggersii is found in areas receiving as little as 500 mm (19.7 in) of precipitation annually. The remaining species are found in montane
Cloud forest
A cloud forest, also called a fog forest, is a generally tropical or subtropical evergreen montane moist forest characterized by a persistent, frequent or seasonal low-level cloud cover, usually at the canopy level. Cloud forests often exhibit an abundance of mosses covering the ground and...

 forests at high elevations or in wet—often very wet—lowland forests, including areas receiving as much as 9000 mm (354.3 in) of annual precipitation.

Records of visits by pollinator
Pollinator
A pollinator is the biotic agent that moves pollen from the male anthers of a flower to the female stigma of a flower to accomplish fertilization or syngamy of the female gamete in the ovule of the flower by the male gamete from the pollen grain...

s exist for only a few species, but most of these suggest that the species are pollinated by insects. Flowers of A. chiribogensis produce small quantities of nectar, but lack a scent. Fruit flies (Drosophilidae
Drosophilidae
Drosophilidae is a diverse, cosmopolitan family of flies, which includes fruit flies. Another family of flies called Tephritidae also includes fruit flies. The best known species of Drosophilidae is Drosophila melanogaster, within the genus Drosophila, and this species Is used extensively for...

), fungus gnat
Fungus gnat
Fungus gnats are small, dark, short-lived flies, of the families Sciaridae, Diadocidiidae, Ditomyiidae, Keroplatidae, Bolitophilidae and Mycetophilidae , sometimes placed in the superfamily Mycetophiloidea, whose larvae feed on plant roots or fungi and aid in the decomposition of organic matter...

s (Mycetophilidae
Mycetophilidae
Mycetophilidae is a family of small flies, forming the bulk of those species known as fungus gnats. There are approximately 3000 described species in 150 genera but the true number of species is undoubtedly much higher...

, Sciaridae
Sciaridae
Sciaridae is a family of flies, commonly known as dark-winged fungus gnats. Commonly found in moist environments, they are known to be a pest of mushroom farms and are commonly found in household plant pots. This is one of the least studied of the large Diptera families, probably due to the small...

), midges (Cecidomyiidae
Cecidomyiidae
Cecidomyiidae is a family of flies known as gall midges or gall gnats. As the name implies, the larvae of most gall midges feed within plant tissue, creating abnormal plant growths called galls.These are very fragile small insects usually only 2–3 mm in length and many are less than...

, Ceratopogonidae
Ceratopogonidae
Ceratopogonidae, or biting midges , are a family of small flies in the order Diptera...

) and micromoths (Lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...

) were recorded visiting these flowers, but bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s and hover flies were not. Aiphanes eggersii was thought to be pollinated by bees and possibly by wind. Fruit flies (Drosophilidae
Drosophilidae
Drosophilidae is a diverse, cosmopolitan family of flies, which includes fruit flies. Another family of flies called Tephritidae also includes fruit flies. The best known species of Drosophilidae is Drosophila melanogaster, within the genus Drosophila, and this species Is used extensively for...

), hover flies
Hoverfly
Hoverflies, sometimes called flower flies or syrphid flies, make up the insect family Syrphidae. As their common name suggests, they are often seen hovering or nectaring at flowers; the adults of many species feed mainly on nectar and pollen, while the larvae eat a wide range of foods...

 (Syrphidae), biting midges (Ceratopogonidae
Ceratopogonidae
Ceratopogonidae, or biting midges , are a family of small flies in the order Diptera...

) and leaf beetle
Leaf beetle
Beetles in the family Chrysomelidae are commonly known as leaf beetles. This is a family of over 35,000 species in more than 2,500 genera, one of the largest and most commonly encountered of all beetle families....

s (Chrysomelidae) were recorded visiting the flowers of A. erinacea, but bees were not. Aiphanes horrida was reportly pollinated by wind, bees (Meliponidae), weevils (Curculionidae
Curculionidae
Curculionidae is the family of the "true" weevils . It was formerly recognized in 1998 as the largest of any animal family, with over 40,000 species described worldwide at that time...

) and bugs (Hemiptera
Hemiptera
Hemiptera is an order of insects most often known as the true bugs , comprising around 50,000–80,000 species of cicadas, aphids, planthoppers, leafhoppers, shield bugs, and others...

). Flies and weevils were observed on the flowers of A. simplex.

The fruit of A. horrida is rich in vitamins and energy and likely to be eaten by many animals. Oilbird
Oilbird
The Oilbird , also known as Guácharo, is a bird found in the northern areas of South America . They are nocturnal feeders on the fruits of the Oil Palm and tropical laurels, and are the only nocturnal fruit eating birds in the world...

s are reported to eat its fruit and disperse its seeds. Squirrels are also reported to consume the fruit, despite the spiny nature of the tree. The fruit, flowers and seeds of A. minima are consumed by the vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

 St. Vincent Amazon (Amazona guildingii) and is also considered a potentially important food species for the critically endangered
Critically endangered
Version 2010.3 of the IUCN Red List of Threatened Species identified 3744 Critically Endangered species, subspecies and varieties, stocks and subpopulations.Critically Endangered by kingdom:*1993 Animalia*2 Fungi*1745 Plantae*4 Protista-References:...

 Puerto Rican Amazon (Amazona vittata).

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 and A. minima in Puerto Rico. 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.

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 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...

 dating back to about 2800 BP
Before 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...

; seeds of this species are still consumed and are traded in local markets. Aiphanes horrida is also widely planted as an ornamental, as is A. minima. The fruit or seeds of A. deltoidea, A. eggersii, A. linearis and A. minima are all consumed locally. The palm heart
Heart of palm
Heart of palm, also called palm heart, palmito, burglar's thigh, chonta, palm cabbage or swamp cabbage, is a vegetable harvested from the inner core and growing bud of certain palm trees Heart of palm, also called palm heart, palmito, burglar's thigh, chonta, palm cabbage or swamp cabbage, is a...

 of A. macroloba is consumed by the Coaiquer
Awa-Cuaiquer language
Awa or Awa pit, also known as Cuaiquer, is a Barbacoan language spoken in northwestern South America.It has 21,000 speakers nearly all in the Colombian Pacific slopes, with 1,000 speakers in an adjacent area of Ecuador...

 people of northwestern South America. Aiphanol, a compound isolated from A. horrida, has shown significant inhibitory
Enzyme inhibitor
An enzyme inhibitor is a molecule that binds to enzymes and decreases their activity. Since blocking an enzyme's activity can kill a pathogen or correct a metabolic imbalance, many drugs are enzyme inhibitors. They are also used as herbicides and pesticides...

 activity against cyclooxygenase
Cyclooxygenase
Cyclooxygenase is an enzyme that is responsible for formation of important biological mediators called prostanoids, including prostaglandins, prostacyclin and thromboxane. Pharmacological inhibition of COX can provide relief from the symptoms of inflammation and pain...

s; inhibition of these enzymes can provide relief from the symptoms of inflammation
Inflammation
Inflammation is part of the complex biological response of vascular tissues to harmful stimuli, such as pathogens, damaged cells, or irritants. Inflammation is a protective attempt by the organism to remove the injurious stimuli and to initiate the healing process...

and pain.
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