Platypodium Elegans
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''Platypodium elegans'', the graceful platypodium, is a large leguminous tree found in the Neotropics that forms part of the forest canopy. It was first described by
Julius Rudolph Theodor Vogel Julius Rudolph Theodor Vogel (30 July 1812 – 17 December 1841) was a German botanist.Biographical sketch by Gottfried Reinhold Treviranus, translated by Miles Joseph Berkeley. In Life He was born in Berlin, and studied at the Friedrich-Wilhelm ...
in 1837 and is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus. The tree has been known to grow up to 30 metres in height and have a trunk with a diameter up to 1 m at breast height. Its trunk has large holes in it, sometimes making it possible to see through the trunk. The holes provide a habitat for giant damselflies and other insects both when alive and once the tree has died and fallen over. It has
compound leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
each of which is made up of 10–20 leaflets. Three new chemical compounds have been isolated from the leaves and they form part of the diet of several monkeys and the squirrel ''
Sciurus ingrami ''Sciurus ingrami'', usually called Ingram's squirrel in English, is a squirrel found in South America. It is known as ''serelepe'' in southeastern Brazil. It is found in the Atlantic Forest Biome of Brazil and Misiones Province, Argentina. It i ...
''. In Panama it flowers from April to June, the flowers contain only four ovules, but normally only one of these reaches maturity forming a winged seed pod around 10 cm long and weighing 2 g. During the dry season around a year after the flowers are fertilised, the seeds are dispersed by the wind and the tree loses it leaves. The seeds are eaten by agoutis and by bruchid beetle larvae. The majority of seedlings are killed by damping off fungi in the first few months of growth, with seedlings that grow nearer the parent trees being more likely to die. The seedlings are relatively unable to survive in deep shade compared to other species in the same habitat. Various
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s are known to grow on ''P. elegans'' with the cactus ''
Epiphyllum phyllanthus ''Epiphyllum phyllanthus'', commonly known as the climbing cactus, is a species of epiphytic cacti. It has no leaves, instead having stems that photosynthesise. It is thought to be pollinated by hawkmoths, as the flowers only open at night and pr ...
'' being the most abundant in Panama. Despite having holes in its trunk which should encourage debris and seeds to collect,
hemiepiphyte A hemiepiphyte is a plant that spends part of its life cycle as an epiphyte. The seeds of primary hemiepiphytes germinate in the canopy and initially live epiphytically. They send roots downward, and these roots eventually make contact with t ...
s are relatively uncommon, meaning that animals are not attracted to it to feed and then defecate. It has no known uses in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
and although it can be used for timber, the wood is of poor quality.


Description

''Platypodium elegans'' is a large forest tree, which forms part of the forest canopy. It can grow up to 30 m in height, with mature trees having an average crown diameter of 16 m and a diameter at breast height (dbh) of 75–100 cm. Trees over 20 cm dbh grow at a rate of around 0.5 cm per year, as measured by how their dbh increases. Its trunk is
fenestrated A fenestra (fenestration; plural fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomical s ...
, having large and conspicuous holes in it, even so much so that it is possible to see through the trunk, meaning it can be mistaken for a
strangler fig Strangler fig is the common name for a number of tropical and subtropical plant species in the genus ''Ficus'', including those that are commonly known as banyans. Some of the more well-known species are: * ''Ficus altissima'' * ''Ficus aurea'', ...
. The bark is soft and dark brown and contains a foul-smelling sap. It has
compound leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
which grow up to 25 cm in length, each having 10–20 leaflets, which are 2.5–7.5 cm long, 1–3 cm wide,
hairless Hairless, also known as H, is a well-characterized Drosophila gene. Since Hairless is a dominant loss of function mutation, many mutations to Hairless are embryonic lethal, but there are several viable hairless mutants. This specific Drosophila ...
on the upper side and positioned not quite opposite each other. Three new compounds have been isolated from the leaves of ''P. elegans''; two ''seco''-lupane triterpenes (canaric acid and dihydrocanaric acid), as well as a coumarin, 6,7,8-trimethoxycoumarin. The mature leaves are relatively tough, requiring more than 100 g/mm2 to be applied to them to puncture them. It is deciduous, losing its leaves during the dry season, when its seeds are dispersed. The diameter of the conducting vessels in the roots are on average 69 
μm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
, up to 98 μm and in the shoots 57 μm up to 87 μm. The trees (over 20 cm dbh) are relatively stiff compared to other trees on Barro Colorado Island (BCI), having a
Young's modulus Young's modulus E, the Young modulus, or the modulus of elasticity in tension or compression (i.e., negative tension), is a mechanical property that measures the tensile or compressive stiffness of a solid material when the force is applied leng ...
of 180,000 kg/cm2. In Panama, it flowers from April to June. It was first reported to flower only every other year, but this is now known to be incorrect, although seed production can vary considerably from year to year. The flowers of ''P. elegans'' contain four ovules, but normally only the most distal ovule develops into a seed, with other seeds being aborted before they mature. The
pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
s are 8–12 mm long, the
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
s around 2 mm long and the calyx around 4 mm long. After being fertilised a winged fruit (a
samara Samara ( rus, Сама́ра, p=sɐˈmarə), known from 1935 to 1991 as Kuybyshev (; ), is the largest city and administrative centre of Samara Oblast. The city is located at the confluence of the Volga and the Samara (Volga), Samara rivers, with ...
) develops quickly, but it takes around one year for the seed to mature. The fruit remains on the tree whilst the seeds develop and is thought to photosynthesise during this time. The samaras dry out during the dry season, before detaching from the tree and being dispersed by the wind over 2–3 months. In Panama the seeds are dispersed between February and April, just under a year after the flowers formed. Each samara normally only contains one seed, but sometimes they contain two instead, which affects their dispersal. The samaras vary in size and shape between trees, but are generally similar on each individual tree. On average, they weigh around 2 g when dry, and are around 10 cm long but can grow up to 16 cm. Samaras containing two seeds are heavier, have a larger surface area and fall faster from the tree than those containing only one seed. Each seed weighs around one third of a gram, making them relatively large compared to other trees in its habitat, but seeds that are the result of
self-fertilisation Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species of ...
are significantly lighter (by 0.03 g). The
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The numb ...
s, of the seedlings remain underground after germination and only serve as a stored source of nutrients, they detach within 8 weeks of germination. File:Flickr - João de Deus Medeiros - Platypodium elegans (1).jpg, leaves File:Flickr - João de Deus Medeiros - Platypodium elegans (2).jpg, seeds File:Flickr - João de Deus Medeiros - Platypodium elegans.jpg, trunk


Taxonomy

''Platypodium elegans'' was first described by
Julius Rudolph Theodor Vogel Julius Rudolph Theodor Vogel (30 July 1812 – 17 December 1841) was a German botanist.Biographical sketch by Gottfried Reinhold Treviranus, translated by Miles Joseph Berkeley. In Life He was born in Berlin, and studied at the Friedrich-Wilhelm ...
in 1837 and it is the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus '' Platypodium''. In 1862 George Bentham described a variety, ''Platypodium elegans'' var. ''major''. In 1917
Henri François Pittier Henri François Pittier de Fabrega (August 13, 1857 in Bex, Switzerland – January 27, 1950 in Caracas, Venezuela) was a Swiss-born geographer and botanist who started Venezuelan National Park history. Biography He graduated as an engineer ...
described ''Platypodium maxonianum'' from Chiriquí, Panama, noting that it differed from Vogel's description of ''P. elegans'' as it had larger leaves and fruits. He named the species after William Maxon, then a curator of the
United States National Herbarium The United States National Herbarium is a collection of five million preserved plant specimens housed in the Department of Botany at the National Museum of Natural History, which is part of the Smithsonian Institution. It represents about 8% of the ...
. ''P. maxonianum'' is now considered to be a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''P. elegans''. According to
The Plant List The Plant List was a list of botanical names of species of plants created by the Royal Botanic Gardens, Kew and the Missouri Botanical Garden and launched in 2010. It was intended to be a comprehensive record of all known names of plant species ...
, ''P. elegans'' is currently one of only two accepted species of ''Platypodium'', the other being '' P. viride''.


Vernacular names

There are many common names for ''P. elegans'' in different languages. In English it is known as the graceful platypodium and similarly, in French as ''platypodium graceiux''. In Brazil it has several names: ''amendoim-do-campo'', ''amendoim-bravo'', ''jacarandá-branco'', ''jacarandá-bana'', ''jacarandá-do-campo'', ''jacarandazinho'', ''jacarandá-tã'', ''faviero'', ''secupiruna'' and ''uruvalheira''. Several names are also used in Panama: ''carcuera'', ''costilla'', ''arbol soga'', ''canalua'', ''canaleto'' and ''tigre''. In Paraguay it is known as ''desconocido'' and in Colombia as ''lomo de caimán''.


Distribution

''Platypodium elegans'' is found in the
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
s and
savannah A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the Canopy (forest), canopy does not close. The open canopy allows sufficient light to rea ...
of the Neotropics, ranging from
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
in the North, to
Paraguay Paraguay (; ), officially the Republic of Paraguay ( es, República del Paraguay, links=no; gn, Tavakuairetã Paraguái, links=si), is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to th ...
in the South. It is also found in Bolivia, Brazil, Colombia and Venezuela. It is not found in the central and northern parts of Brazil, but is found in the cerrado,
Mato Grosso Mato Grosso ( – lit. "Thick Bush") is one of the states of Brazil, the third largest by area, located in the Central-West region. The state has 1.66% of the Brazilian population and is responsible for 1.9% of the Brazilian GDP. Neighboring ...
,
Minas Gerais Minas Gerais () is a state in Southeastern Brazil. It ranks as the second most populous, the third by gross domestic product (GDP), and the fourth largest by area in the country. The state's capital and largest city, Belo Horizonte (literally ...
and around
São Paulo São Paulo (, ; Portuguese for 'Saint Paul') is the most populous city in Brazil, and is the capital of the state of São Paulo, the most populous and wealthiest Brazilian state, located in the country's Southeast Region. Listed by the GaWC a ...
. On Barro Colorado Island (BCI) ''P. elegans'' is found at a moderate abundance in both old and young forest, each hectare may contain several mature trees, but it is not unusual to find isolated individuals. Generally, only large individuals are found, with saplings being rare, except in forest gaps.


Ecology


Reproduction

''Platypodium elegans'' is pollinated by
bees Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfamil ...
. In Panama pollen is moved on average between 368 and 419 m from the parent tree and commonly over 1 km away. The population that regularly share genes (termed the
deme In Ancient Greece, a deme or ( grc, δῆμος, plural: demoi, δημοι) was a suburb or a subdivision of Athens and other city-states. Demes as simple subdivisions of land in the countryside seem to have existed in the 6th century BC and ear ...
) is estimated to be between 25 and 50 hectares around each tree. 92% of seeds that mature result from flowers that have been pollinated with pollen from other individuals but
self-fertilisation Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species of ...
is actually much higher than this would suggest. The difference between these values is explained by the fact that many fruits are aborted after being fertilised, but before dispersal occurs. Hufford and Hamrick suggested that they abort fruit for two reasons; they could have a set amount of resources to invest in their seeds in one year but produce extra flowers which then compete between each other, with only some surviving to maturity. Alternatively, the tree may detect which fruits are the result of self-fertilisation and selectively abort them, but this is considered less likely. The seeds are dispersed by wind. In Panama the seeds germinate at the start of the rainy season in May. Damping off fungi account for 64–95% of the deaths of seedlings during their first three months of growth, with deaths being more common for seedlings near their parents, as the Janzen-Connell hypothesis predicts. Falling leaf litter and digging mammals are also significant causes of seedling mortality. An artificial experiment found that if seedlings have their leaves removed or are placed in deep shade (0.08% of full sunlight) they die within 60 days, whereas around 80% of the seedlings of '' Lacmellea panamensis'' are able to survive these treatments. The seedlings are able to grow slowly in 0.8% sunlight, but the seedlings are at the low end of the spectrum in terms of being able to tolerate shade. Another experiment has shown that seedlings die faster if either their cotyledons or leaves are removed, those that have had their cotyledons removed die more quickly.


Habitat

As in all legumes, the roots of ''P. elegans'' are colonised by
nitrogen fixing Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the Atmosphere of Earth, air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but al ...
bacteria, in this case from the genus ''
Bradyrhizobium ''Bradyrhizobium'' is a genus of Gram-negative soil bacteria, many of which fix nitrogen. Nitrogen fixation is an important part of the nitrogen cycle. Plants cannot use atmospheric nitrogen (N2); they must use nitrogen compounds such as nitrat ...
''. Genetic analysis of the bacteria has shown that different
genotype The genotype of an organism is its complete set of genetic material. Genotype can also be used to refer to the alleles or variants an individual carries in a particular gene or genetic location. The number of alleles an individual can have in a ...
s colonise the roots of the same tree and are strains of '' Bradyrhizobium japonicum''. The
epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
cactus ''
Epiphyllum phyllanthus ''Epiphyllum phyllanthus'', commonly known as the climbing cactus, is a species of epiphytic cacti. It has no leaves, instead having stems that photosynthesise. It is thought to be pollinated by hawkmoths, as the flowers only open at night and pr ...
'' is particularly abundant in the canopies of ''P. elegans'' on BCI particularly growing in cavities in the trunk. Another cactus, '' Rhipsalis baccifera'', and the ferns ''
Niphidium crassifolium ''Niphidium crassifolium'', commonly known as the graceful fern, is a species of fern in the family Polypodiaceae found in Central and South America. It is predominantly epiphytic, growing on other plants—for example, in the canopies of trees ...
'' and '' Campyloneurum phyllitidis'' are also found growing on ''P. elegans''. After rainfall, the bark stores around one third of a gram of water per cm² which epiphytes can then absorb, a moderate amount compared to other trees. Todzia found that despite having a trunk with deep invaginations that collect debris and which should encourage the germination of seeds,
hemiepiphyte A hemiepiphyte is a plant that spends part of its life cycle as an epiphyte. The seeds of primary hemiepiphytes germinate in the canopy and initially live epiphytically. They send roots downward, and these roots eventually make contact with t ...
s (plants which germinate on the tree and then send down roots into the soil) are relatively rare on ''P. elegans'' on BCI. It is thought that this is because its seeds are wind dispersed, and the tree therefore attracts relatively few animals which could deposit the seeds of hemiepiphytes whilst feeding on seeds. Todzia noted that '' Hura crepitans'' disperses it seeds explosively, yet is heavily laden with hemiepiphytes however. A survey of 20 trees on BCI with a diameter at breast height of 20 cm or more found that 75% had
liana A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...
s growing on them. Various invertebrates live in water-filled holes which form in ridges of the trunk of ''P. elegans'' when they die and fall over, and in tree hollows that exist when the tree is alive. Leaf litter collects in the water and as it decomposes animals feed on the debris. An experiment, where leaves of ''P. elegans'' were added to an artificial pool containing 650 ml of water in the rainforest, found that 17 species lived in them, with the mosquito ''
Culex mollis ''Culex'' is a genus of mosquitoes, several species of which serve as Vector (epidemiology), vectors of one or more important diseases of birds, humans, and other animals. The diseases they vector include arbovirus infections such as West Nile ...
'' being the most abundant. The pools contained a greater species diversity and abundance of animals than similar experiments using leaves of '' Ceiba pentandra'', ''
Dipteryx panamensis ''Dipteryx oleifera'' (syns. ''Dipteryx panamensis'' and ''Coumarouna panamensis''), the eboe, choibá or almendro (almond in Spanish), is a species of flowering plant in the family Fabaceae, native to Honduras, Nicaragua, Costa Rica, Panama, Co ...
'' and ''
Ficus yoponensis ''Ficus yoponensis'' is a species of fig tree found in Central and South America. It can grow to heights of tall, having a trunk diameter of . The trunk is buttressed, light grey in colour and reasonably smooth. Its petioles are long, the stip ...
'', species that also contain water pools in their trunks. Yanoviak suggested that this indicates that the leaves are a relatively higher-quality nutrient source than those of the other species. On BCI, Fincke found that trees had between 1 and 10 water-filled holes, more than any other tree species investigated, each containing on average 2 litres of water. Yanoviak found that the holes contained only 400 ml of water on average however. The pools are an important habitat for the larvae of giant damselflies. As a coloniser of new habitats, ''P. elegans'' may provide an ideal habitat for giant damselflies in
secondary forest A secondary forest (or second-growth forest) is a forest or woodland area which has re-grown after a timber harvest or clearing for agriculture, until a long enough period has passed so that the effects of the disturbance are no longer evident. ...
. The beetle '' Microvelia cavicola'' also lives in the water-filled holes, with the
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
of the species being found in one.


Food

The embryos of immature fruit are eaten by agoutis (''
Dasyprocta punctata The Central American agouti (''Dasyprocta punctata'') is a species of agouti from the family Dasyproctidae. The main portion of its range is from Chiapas and the Yucatan Peninsula (southern Mexico), through Central America, to northwestern Ecuad ...
'') once they have fallen to the forest floor. Mature seeds can be infected by fungi and are also eaten by bruchid beetle larvae. The squirrel, ''
Sciurus ingrami ''Sciurus ingrami'', usually called Ingram's squirrel in English, is a squirrel found in South America. It is known as ''serelepe'' in southeastern Brazil. It is found in the Atlantic Forest Biome of Brazil and Misiones Province, Argentina. It i ...
'' eats the leaves of ''P. elegans'' on BCI, but not in Southeastern Brazil. Woolly spider monkeys in Brazil feed extensively on the leaves in October, prior to the beginning of their mating season. The leaves are thought to be low in
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'', ...
s and other secondary metabolites which hinder protein digestion, making them an ideal food before the mating season. They also contain
phytoestrogen A phytoestrogen is a plant-derived xenoestrogen (see estrogen) not generated within the endocrine system, but consumed by eating plants or manufactured foods. Also called a "dietary estrogen", it is a diverse group of naturally occurring nonstero ...
s which can change the monkey's
estrogen Estrogen or oestrogen is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three major endogenous estrogens that have estrogenic hormonal acti ...
levels, possibly affecting their fertility. Both young and mature leaves of ''P. elegans'' are eaten by howler monkeys, as the leaves mature, the protein content decreases, the cell wall content increases, but the proportion of non-structural carbohydrates remains equal. When fresh, the leaves contain between 100 and 200 mg of
ascorbic acid Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) an ...
(
Vitamin C Vitamin C (also known as ascorbic acid and ascorbate) is a water-soluble vitamin found in citrus and other fruits and vegetables, also sold as a dietary supplement and as a topical 'serum' ingredient to treat melasma (dark pigment spots) an ...
) per 100 g, like humans some primates on BCI require in their diet, since they do not possess the gene for L-gulonolactone oxidase, the enzyme required to convert glucose to ascorbic acid.


Uses

''Platypodium elegans'' is not known to have any uses in traditional medicine. The Smithsonian Tropical Research Institute herbarium report its wood is used for timber, being described as "white, knotty, light and fragile". Pittier remarked in 1931 that the wood is little used in Panama however since the mature trunks are normally hollow and filled with an oily liquid.


References


External links


Photographs of ''Platypodium elegans''
from the
Smithsonian Tropical Research Institute The Smithsonian Tropical Research Institute (STRI, es, Instituto Smithsonian de Investigaciones Tropicales) is located in Panama and is the only bureau of the Smithsonian Institution based outside of the United States. It is dedicated to understa ...
Herbarium
Herbarium specimens
from JSTOR plants. {{Taxonbar, from=Q7202696 Dalbergieae Plants described in 1887 Trees of Panama Trees of Paraguay Trees of Bolivia Trees of Colombia Trees of Brazil Trees of the Amazon rainforest Trees of Venezuela