Philodendron Propinquum
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''Philodendron'' is a large
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s in the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
. , the
Plants of the World Online Plants of the World Online (POWO) is an online taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...
accepted 621 species; other sources accept different numbers. Regardless of number of species, the genus is the second-largest member of the family Araceae, after genus ''
Anthurium ''Anthurium'' (; Schott, 1829) is a genus of about 1,000Mantovani, A. and T. E. Pereira. (2005)''Anthurium'' (section ''Urospadix''; subsection ''Flavescentiviridia'').''Rodriguesia'' 56(88), 145–60. species of flowering plants, the largest g ...
''. Taxonomically, the genus ''Philodendron'' is still poorly known, with many
undescribed species In taxonomy, an undescribed taxon is a taxon (for example, a species) that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. U ...
. Many are grown as ornamental and indoor plants. The name derives from the Greek words ''philo-'' 'love, affection' and ''dendron'' 'tree'. The generic name, ''Philodendron'', is often used as the English name.


Description


Growth habit

Compared to other genera of the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
, philodendrons have an extremely diverse array of growth methods. The habits of growth can be
epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another 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 epiphyt ...
, hemiepiphytic, or rarely
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
. Others can show a combination of these growth habits depending on the environment. Hemiepiphytic philodendrons can be classified into two types: primary and secondary hemiepiphytes. A primary hemiepiphytic philodendron starts life high up in the canopy where the seed initially sprouts. The plant then grows as an epiphyte. Once it has reached a sufficient size and age, it will begin producing aerial roots that grow toward the forest floor. Once they reach the forest floor, nutrients can be obtained directly from the soil. In this manner, the plant's strategy is to obtain light early in its life at the expense of nutrients. Some primary epiphytic species have a
symbiotic relationship Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
with ants. In these species, the ants' nest is grown amongst the plant's roots, which help keep the nest together. Philodendrons have
extrafloral nectaries Nectar is a viscous, sugar-rich liquid produced by plants in glands called nectaries, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists, ...
, glands that secrete nectar to attract the ants. The philodendron, in turn, obtains nutrients from the surrounding ant nest, and the aggressive nature of the ants serves to protect the plant from other insects which would eat it. Secondary hemiepiphytes start life on the ground or on part of a tree trunk very close to the ground, where the seeds sprout. These philodendrons have their roots in the ground early in their lives. They then begin climbing up a tree and eventually may become completely epiphytic, doing away with their subterranean roots. Secondary hemiepiphytes do not always start their lives close to a tree. For these philodendrons, the plant will grow with long
internodes A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, engages in pho ...
along the ground until a tree is found. They find a suitable tree by growing towards darker areas, such as the dark shadow of a tree. This trait is called scototropism. After a tree has been found, the scototropic behavior stops and the philodendron switches to a
phototropic In biology, phototropism is the growth of an organism in response to a light stimulus. Phototropism is most often observed in plants, but can also occur in other organisms such as fungi. The cells on the plant that are farthest from the ligh ...
growth habit and the internodes shorten and thicken. Usually, however, philodendrons germinate on trees. A few species, such as '' Philodendron fragrantissimum'' switch repeatedly between globular clumps of leaves with very short
internodes A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, engages in pho ...
and naked internodes of uncommon length (up to a yard (meter) or more) followed by another clump and so on. '' Philodendron linnaei'' is another such species. A few species show three peaks in temperature during flowering, which stimulates beetles within the spathe and increasing the likelihood they will be sufficiently coated with pollen. A sticky resin is also produced in drops attached to the spadix which help to keep the pollen attached to the beetles. This resin producing quality is unique to ''Philodendron'' and '' Monstera'', as other genera of
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
do not produce it on their spadices. The resin is also found on the stems, leaves, and roots of philodendrons. Its color can be red, orange, yellow, or colorless when it is first produced. Yet, over time, it will turn brown as it is exposed to air. Also, some evidence suggests the thermogenesis triggers the beetles to mate. It also appears to distribute the pheromones into the air. The reason for the spadix being held at 45° relative to the spathe may be to maximize the heat's ability to waft the pheromones into the air. Oxidizing stored carbohydrates and lipids has been found to be the energy source for thermogenesis. The part of the spadix that heats up is the sterile zone. As it heats up, carbohydrates are used, but once the spadix has reached its maximum temperature, lipids are oxidized. The lipids are not first converted to carbohydrates, but rather are directly oxidized. The thermogenic reaction is triggered when concentrations of acetosalicytic acid form in the sterile zone. The acid sets off the
mitochondria A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
in the cells that make up the sterile zone to switch to an
electron transport chain An electron transport chain (ETC) is a series of protein complexes and other molecules which transfer electrons from electron donors to electron acceptors via redox reactions (both reduction and oxidation occurring simultaneously) and couples th ...
called the cyanide-resistant pathway, which results in the production of heat. Philodendrons consume
oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...
during thermogenesis. The rate at which oxygen is used is remarkably high, close to that of
hummingbirds Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
and sphinx moths. The spadix has been shown to generate
infrared radiation Infrared (IR; sometimes called infrared light) is electromagnetic radiation (EMR) with wavelengths longer than that of visible light but shorter than microwaves. The infrared spectral band begins with the waves that are just longer than those ...
.


Leaves

The leaves are usually large and imposing, often lobed or deeply cut, and may be more or less
pinnate Pinnation (also called pennation) is the arrangement of feather-like or multi-divided features arising from both sides of a common axis. Pinnation occurs in biological morphology, in crystals, such as some forms of ice or metal crystals, and ...
. They can also be oval (''Philodendron'' 'White Princess'), spear-shaped, divided ('' Philodendron tripartitum'') or in many other possible shape variations. The leaves are borne alternately on the stem. A quality of philodendrons is that they do not have a single type of leaf on the same plant. Instead, they have juvenile leaves and adult leaves, which can be drastically different from one another. The leaves of seedling philodendrons are usually heart-shaped early in the life of the plant. But after it has matured past the seedling stage, the leaves will acquire the typical juvenile leaf's shape and size. Later in the philodendron's life, it starts producing adult leaves, a process called
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth transformation or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and different ...
. Most philodendrons go through metamorphosis gradually; there is no immediately distinct difference between juvenile and adult leaves. Aside from being typically much bigger than the juvenile leaves, the shape of adult leaves can be significantly different. In fact, considerable taxonomic difficulty has occurred in the past due to these differences, causing juvenile and adult plants to mistakenly be classified as different species. The trigger for the transformation to adult leaves can vary considerably. One possible trigger is the height of the plant. Secondary hemiepiphytes start off on the dark forest floor and climb their way up a tree, displaying their juvenile type leaves along the way. Once they reach a sufficient height, they begin developing adult type leaves. The smaller juvenile leaves are used for the darker forest floor where light is in scarce supply, but once they reach a sufficient height in the canopy the light is bright enough that the bigger adult leaves can serve a useful purpose. Another possible trigger occurs in primary hemiepiphytes. These philodendrons typically send their aerial roots downward. Once their roots have reached the ground below, the plant will begin taking up nutrients from the soil, of which it had been previously deprived. As a result, the plant will quickly morph into its adult leaves and gain in size dramatically. Another quality of philodendrons leaves is they are often quite different in shape and size even between two plants of the same species. As a result of all these different possible leaf shapes, it is often difficult to differentiate natural variations from
morphogenesis Morphogenesis (from the Greek ''morphê'' shape and ''genesis'' creation, literally "the generation of form") is the biological process that causes a cell, tissue or organism to develop its shape. It is one of three fundamental aspects of deve ...
.


Cataphylls

Philodendrons also produce cataphylls, which are modified leaves that surround and protect the newly forming leaves. Cataphylls are usually green, leaf-like, and rigid while they are protecting the leaf. In some species, they can even be rather
succulent In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
. Once the leaf has been fully formed, the cataphyll usually remains attached where the stem and base of the leaf meet. In philodendrons, cataphylls typically fall into two categories:
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
and persistent types. A deciduous cataphyll curls away from the leaf once it has formed, eventually turning brown and drying out, and finally falling off the plant, leaving a scar on the stem where it was attached. Deciduous cataphylls are typically found on vining philodendrons, whereas persistent cataphylls are typical of epiphytic philodendrons or appressed climbers. In the latter, the cataphylls are prevented from falling off in a timely manner due to the short
internodes A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, engages in pho ...
of the plant. The cataphylls will remain attached, drying out and becoming nothing more than fibers attached at the nodes. In some philodendrons, the cataphylls build up over time and eventually form a wet mass at the nodes. This may keep emerging roots moist and provide some form of lubrication to new leaves.


Roots

Philodendrons have both aerial and subterranean roots. The aerial roots occur in many shapes and sizes and originate from most of the plant's nodes or occasionally from an internode. The size and number of aerial roots per node depends on the presence of a suitable substrate for the roots to attach themselves. Aerial roots serve two primary purposes. They allow the philodendron to attach itself to a tree or other plant, and they allow it to collect water and nutrients. As such, the roots are divided morphologically into these two categories. Aerial roots used for attaching to trees tend to be shorter, more numerous, and sometimes have a layer of root hairs attached; those used for collecting water and nutrients tend to be thicker and longer. These feeder roots tend to attach flush with the substrate to which the philodendron is attached, and make their way directly downwards in search of soil. In general, feeder roots tend to show both positive hydrotropic and negative heliotropic behaviors. Characteristic of roots in philodendrons is the presence of a sclerotic hypodermis, which are cylindrical tubes inside the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
that can be one to five cells long. The cells that line the sclerotic hypodermis are elongated and tend to be hardened. Underneath the epidermis is a unique layer of cells in a pattern of long cells followed by short cells.


Extrafloral nectaries

Some philodendrons have extrafloral nectaries (nectar-producing glands found outside of the flowers). The nectar attracts ants, with which the plant enjoys a protective symbiotic relationship. Nectaries can be found in a variety of locations on the plant, including the stalks, sheaths, lower surfaces of the leaves, and
spathe In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
s. The nectaries produce a sweet, sticky substance the ants like to eat and which provides an incentive for them to build their nests amongst the roots of the given philodendron. In some cases, the amount of
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
produced can be quite extensive, resulting in the surface becoming entirely covered with it.


Reproduction


Sexual

When philodendrons are ready to reproduce, they will produce an
inflorescence In botany, an inflorescence is a group or cluster of flowers arranged on a plant's Plant stem, stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a mai ...
which consists of a leaf-like hood called a
spathe In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
within which is enclosed a tube-like structure called a spadix. Depending on the species, a single inflorescence can be produced or a cluster of up to 11
inflorescences In botany, an inflorescence is a group or cluster of flowers arranged on a plant's stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a main axis ( pe ...
can be produced at a single time on short peduncles. The spathe tends to be waxy and is usually bicolored. In some philodendrons, the color of the base of the spathe contrasts in color with the upper part, and in others, the inner and outer surfaces of the spathe differ in coloration. The paler color tends to be either white or green, and the darker usually red or crimson.
Pelargonidin Pelargonidin is an anthocyanidin, a type of plant pigment producing a characteristic orange color used in food and industrial dyes. Natural occurrences Presence in flowers Pelargonidin can be found in red geraniums (Geraniaceae). It is the pr ...
is the predominant
pigment A pigment is a powder used to add or alter color or change visual appearance. Pigments are completely or nearly solubility, insoluble and reactivity (chemistry), chemically unreactive in water or another medium; in contrast, dyes are colored sub ...
causing the red coloration in the spathes. The upper portion of the spathe is called the limb or blade, while the lower portion is called the convolute tube or chamber due to its tubular structure at the base. The spadix is more often than not white and shorter than the spathe. On the spadix are found fertile female, fertile male, and sterile male flowers. The fertile male and female flowers are separated on the spadix by a sterile zone or staminodal region composed of sterile male flowers. This barrier of sterile male flowers ensures fertile male flowers do not fertilize the female flowers. The arrangement tends to be vertical, with fertile male flowers at the top of the spadix followed by sterile male flowers, and fertile female flowers very close to the bottom in the region known as the spathe tube or chamber. In some philodendrons, an additional region of sterile male flowers is found at the very top of the spadix. The fertile female flowers are often not receptive to fertilization when the fertile males are producing
pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...
, which again prevents
self-pollination Self-pollination is a form of pollination in which pollen arrives at the stigma of a flower (in flowering plants) or at the ovule (in gymnosperms) of the same plant. The term cross-pollination is used for the opposite case, where pollen from ...
. The pollen itself is thread-like and appears to project out from the region where the fertile male flowers are located.
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
is achieved by means of
beetles Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...
, with many philodendron species requiring the presence of a specific beetle species to achieve pollination. The reverse is not always the case, as many beetle species will pollinate more than one philodendron species. These same beetles could also pollinate other genera outside of philodendron, as well as outside of the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
. The pollinating beetles are males and members of the subfamily Rutelinae and
Dynastinae Dynastinae or rhinoceros beetles are a subfamily of the scarabaeidae, scarab beetle family (biology), family (Scarabaeidae). Other common names – some for particular groups of rhinoceros beetles – include Hercules beetles, unicorn be ...
, and to date the only beetles seen to pollinate the inflorescence are in the genera ''
Cyclocephala ''Cyclocephala'' is a genus of scarab beetles from the subfamily Dynastinae (Coleoptera, Scarabaeidae). Beetles of this genus occur from southeastern Canada to Argentina, India and the West Indies. Adults of this genus are nocturnal or crepuscu ...
'' or '' Erioscelis''. Other smaller types of beetles in the genus'' Neelia'' visit the inflorescences, as well, but they are not believed to be involved in pollinating philodendrons. To attract the beetles, the sterile male flowers give off
pheromones A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavi ...
to attract the male beetles, usually at dusk. This process, female
anthesis Anthesis is the period during which a flower is fully open and functional. It may also refer to the onset of that period. The onset of anthesis is spectacular in some species. In ''Banksia'' species, for example, anthesis involves the extension ...
, is followed by male anthesis, in which the pollen is produced. Female anthesis typically lasts up to two days and includes the gradual opening of the spathe to allow the beetles to enter. Some evidence suggests the timing of opening of the spathe is dependent on light levels, where cloudy, darker days result in the spathe opening up earlier than on clear days. During female anthesis, the spadix will project forward at roughly 45° relative to the spathe. Once female anthesis is nearing its end and the female flowers have been pollinated, the spathe will be fully open and male anthesis begins. In the beginning of male anthesis, the fertile male flowers complete the process of producing the pollen and the female flowers become unreceptive to further pollination. Additionally, the spadix moves from its 45° position and presses up flush to the spathe. Towards the end of male anthesis, the spathe begins to close from the bottom, working its way up and forcing the beetles to move up and across the upper region of the spathe, where the fertile male flowers are located. In doing so, the philodendron controls when the beetles come and when they leave and forces them to rub against the top of the spadix where the pollen is located as they exit, thus ensuring they are well-coated with pollen. One would expect the beetles to stay indefinitely if they could due to the very favorable conditions the inflorescence provides. After male anthesis, the males will go off and find another philodendron undergoing female anthesis, so will pollinate the female flowers with the pollen it had collected from its previous night of mating.


Fruit

Botanically, the fruit produced is a
berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit although many pips or seeds may be present. Common examples of berries in the cul ...
. The berries develop later in the season; berry development time varies from species to species from a few weeks to a year, although most philodendrons take a few months. The
spathe In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
will enlarge to hold the maturing berries. Once the fruit are mature, the spathe will begin to open again, but this time it will break off at the base and fall to the forest floor. Additionally, the berries are edible, although they contain calcium oxalate crystals, and have a taste akin to bananas. Many botanical sources will indicate that the berries are poisonous, probably due to the oxalate crystals. Many tropical plants contain oxalates in varying amounts. Sometimes proper preparation can render these harmless, and in many cases eating minor amounts causes most people no distress or minor gastric irritation. However, care should be taken to verify the toxicity of any particular species before ingesting these berries, particularly regularly or in large amounts. The color of the berries can vary depending on the species, but most produce a white berry with slight tones of green. Some produce orange berries and others yellow berries, though. Still others will produce berries that start off white, but then change to another color with time. Philodendrons that produce orange berries tend to be members of the section ''Calostigma''. Contained within the berries are the seeds which are extremely small compared to other members of the family Araceae. The berries often give off odors to attract animals to eat and disperse them. For example, '' Philodendron alliodorum'' berries are known to emit an odor similar to that of garlic. The animals that distribute the seeds depends on the species, but some possible dispersers include bats and monkeys. Insects also may be responsible for dispersing seeds, as beetles and wasps have been seen feeding on philodendron berries. Eurytomid wasps also seek out philodendrons and are known to lay their eggs in the ovaries of many ''Philodendron'' species, resulting in galled inflorescences.


Hybridization

Philodendrons exhibit extremely few physical reproductive barriers to prevent hybridization, but very few natural hybrids are found in nature. This may be because philodendrons have many geographic and time barriers to prevent any such cross pollination. For example, it is rare for more than one philodendron species to be flowering at the same time or to be pollinated by the same species of beetles. The beetles have also been observed to be selective to the height of the plant they pollinate, which would serve as an additional preventive measure to make hybrids less likely. Because of these outside barriers, philodendrons may not have had to evolve physical mechanisms to prevent cross-pollination. Hybrids in nature are only rarely reported. When found, these hybrids often can show remarkable genetic relationships. Crosses between two philodendrons in different sections can occur successfully.


Taxonomy


History

Philodendrons are known to have been collected from the wild as early as 1644 by Georg Marcgraf, but the first partly successful scientific attempt to collect and classify the genus was done by Charles Plumier. Plumier collected approximately six species from the islands of
Martinique Martinique ( ; or ; Kalinago language, Kalinago: or ) is an island in the Lesser Antilles of the West Indies, in the eastern Caribbean Sea. It was previously known as Iguanacaera which translates to iguana island in Carib language, Kariʼn ...
,
Hispaniola Hispaniola (, also ) is an island between Geography of Cuba, Cuba and Geography of Puerto Rico, Puerto Rico in the Greater Antilles of the Caribbean. Hispaniola is the most populous island in the West Indies, and the second-largest by List of C ...
, and St. Thomas. Since then, many exploration attempts have been made to collect new species by others. These include those by N.J. Jacquin who collected new species in the
West Indies The West Indies is an island subregion of the Americas, surrounded by the Atlantic Ocean, North Atlantic Ocean and the Caribbean Sea, which comprises 13 independent island country, island countries and 19 dependent territory, dependencies in thr ...
, Colombia, and Venezuela. At this time in history, the names of the philodendrons they were discovering were being published with the genus name ''Arum'', since most aroids were considered to belong to this same genus. The genus ''Philodendron'' had not yet been created. Throughout the late 17th, 18th century, and early 19th centuries, many plants were removed from the genus'' Arum'' and placed into newly created genera in an attempt to improve the classification.
Heinrich Wilhelm Schott Heinrich Wilhelm Schott (7 January 1794 – 5 March 1865) was an Austrian botanist. He is known for his extensive work on aroids ( Araceae). Biography Schott was born on 7 January 1794 in Brno, Moravia. He studied botany, agriculture and chemi ...
addressed the problem of providing improved taxonomy and created the genus ''Philodendron'' and described it in 1829. The genus was first spelled as 'Philodendrum', but in 1832, Schott published a system for classifying plants in the family
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also ...
titled ''Meletemata Botanica'' in which he provided a method of classifying philodendrons based on flowering characteristics. In 1856, Schott published a revision of his previous work titled ''Synopsis aroidearum'', and then published his final work ''Prodromus Systematis Aroidearum'' in 1860, in which he provided even more details about the classification of ''Philodendron'' and described 135 species.


Modern classification

Philodendron are usually extremely distinctive and not usually confused with other genera, although a few exceptions in the genera ''
Anthurium ''Anthurium'' (; Schott, 1829) is a genus of about 1,000Mantovani, A. and T. E. Pereira. (2005)''Anthurium'' (section ''Urospadix''; subsection ''Flavescentiviridia'').''Rodriguesia'' 56(88), 145–60. species of flowering plants, the largest g ...
'' and '' Homalomena'' resemble ''Philodendron''. The genus Philodendron has been subdivided into three subgenera: ''Meconostigma'', ''Pteromischum'', and ''Philodendron''. In 2018, it was proposed that ''Philodendron'' subg. ''Meconostigma'' be recognized as a separate genus, ''Thaumatophyllum''. The genus ''Philodendron'' can also be subdivided into several sections and subsections. Section '' Baursia'', section ''Philopsammos'', section ''Philodendron'' (subsections ''Achyropodium'', ''Canniphyllium'', ''Macrolonchium'', ''Philodendron'', ''Platypodium'', ''Psoropodium'' and ''Solenosterigma''), section ''Calostigma'' (subsections ''Bulaoana'', ''Eucardium, Glossophyllum'', ''Macrobelium'' and ''Oligocarpidium''), section ''Tritomophyllum'', section ''Schizophyllum'', section ''Polytomium'', section ''Macrogynium'' and section ''Camptogynium''. Typically, the inflorescence is of great importance in determining the species of a given philodendron, since it tends to be less variable than the leaves. The genus ''Philodendron'' could be classified further by means of differentiating them based on the pattern of thermogenesis observed, although this is not currently used.


Selected species

* '' Philodendron acutatum'' Schott * '' Philodendron alliodorum'' Croat & Grayum * ''
Philodendron appendiculatum ''Philodendron appendiculatum'', also known as güembé, is a perennial species in the genus '' Philodendron'', belonging to the family Araceae. It lives in the jungles, wetlands, and moist forests of South America. Distribution and habitat ...
'' Nadruz & Mayo * '' Philodendron auriculatum'' Standl. & L. O. Williams * '' Philodendron balaoanum'' Engl. * '' Philodendron bipennifolium'' Schott * '' Philodendron carinatum'' E.G.Gonç. * '' Philodendron chimboanum'' Engl. * '' Philodendron consanguineum'' Schott - rascagarganta * '' Philodendron cordatum'' (Vell.) Kunth * '' Philodendron crassinervium'' Lindl. * '' Philodendron cruentospathum'' Madison * '' Philodendron davidsonii'' Croat * '' Philodendron devansayeanum'' L. Linden * '' Philodendron domesticum'' G. S. Bunting * '' Philodendron duckei'' Croat & Grayum * '' Philodendron ensifolium'' Croat & Grayum * '' Philodendron erubescens'' K. Koch & Augustin * '' Philodendron eximium'' Schott * '' Philodendron fragrantissimum'' (Hook.) G. Don in Sweet * '' Philodendron ferrugineum'' Croat * ''
Philodendron giganteum ''Philodendron giganteum'' is a species of plant in the Araceae family. It is found in the Caribbean and South America. Heinrich Wilhelm Schott first described it in 1856. ''P. giganteum'' inflorescences are thermogenetic and emit a sweet odor. I ...
'' Schott - giant philodendron * '' Philodendron gigas'' Croat * '' Philodendron gloriosum'' André * '' Philodendron gualeanum'' Engl. * '' Philodendron hastatum'' K. Koch & Sello - silver philodendron, also known incorrectly as ''Philodendron glaucophyllum'' * '' Philodendron hederaceum'' (Jacq.) Schott - vilevine, heartleaf philodendron, velvet-leaved philodendron * '' Philodendron herbaceum'' Croat & Grayum * '' Philodendron hooveri'' Croat & Grayum * '' Philodendron imbe'' Schott ex Endl. - philodendron * '' Philodendron jacquinii'' Schott * '' Philodendron lacerum'' (Jacq.) Schott * '' Philodendron lingulatum'' (L.) K. Koch - treelover * '' Philodendron mamei'' André * '' Philodendron marginatum'' Urban - Puerto Rico philodendron * '' Philodendron martianum'' Engl.- also known incorrectly as Philodendron cannifolium * '' Philodendron mayoi'' Symon Mayo * '' Philodendron maximum'' K. Krause * '' Philodendron melanochrysum'' Linden & André * '' Philodendron microstictum'' Standl. & L. O. Williams * '' Philodendron musifolium'' Engl. * '' Philodendron nanegalense'' Engl. * '' Philodendron opacum'' Croat & Grayum * '' Philodendron ornatum'' Schott * '' Philodendron pachycaule'' K. Krause * '' Philodendron panduriforme'' (Kunth) Kunth * '' Philodendron pedatum'' (Hook.) Kunth * '' Philodendron pinnatifidum'' (Jacq.) Schott * '' Philodendron pogonocaule'' Madison * '' Philodendron pterotum'' K.Koch & Augustin * '' Philodendron quitense'' Engl. * '' Philodendron radiatum'' Schott * '' Philodendron recurvifolium'' Schott * '' Philodendron renauxii'' Reitz * '' Philodendron riparium'' Engl. * '' Philodendron robustum'' Schott * '' Philodendron rugosum'' Bogner & G.S.Bunting * '' Philodendron sagittifolium'' Liebm. * '' Philodendron santa leopoldina'' Liebm. * '' Philodendron sphalerum'' Schott * '' Philodendron squamiferum'' Poepp. * '' Philodendron standleyi'' Grayum * '' Philodendron tatei'' Krause * '' Philodendron tripartitum'' (Jacq.) Schott * '' Philodendron validinervium'' Engl. * '' Philodendron ventricosum'' Madison * '' Philodendron verrucosum'' L. Mathieu ex Schott * '' Philodendron victoriae'' G.S. Bunting * ''
Philodendron warszewiczii ''Philodendron warszewiczii'' is a species of flowering plant in the family Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually ac ...
'' K. Koch & C. D. Bouché * '' Philodendron wendlandii'' Schott


Evolution

''Philodendron'' diverged from '' Adelonema'' and diversified during the late
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
, 25 million years ago, in the New World.


Distribution and habitat

Philodendron species can be found in many diverse habitats in the tropical Americas and the
West Indies The West Indies is an island subregion of the Americas, surrounded by the Atlantic Ocean, North Atlantic Ocean and the Caribbean Sea, which comprises 13 independent island country, island countries and 19 dependent territory, dependencies in thr ...
. Most occur in humid tropical forests, but can also be found in swamps and on river banks, roadsides and rock outcrops. They are also found throughout the diverse range of elevations from sea level to over 2000 m above sea level. Species of this genus are often found clambering over other plants, or climbing the trunks of
trees In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only p ...
with the aid of aerial roots. Philodendrons usually distinguish themselves in their environment by their large numbers compared to other plants, making them a highly noticeable component of the
ecosystems An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
in which they are found. They are found in great numbers in road clearings. Philodendrons can also be found in Australia, some
Pacific islands The Pacific islands are a group of islands in the Pacific Ocean. They are further categorized into three major island groups: Melanesia, Micronesia, and Polynesia. Depending on the context, the term ''Pacific Islands'' may refer to one of several ...
, Africa and Asia, although they are not indigenous and were introduced or accidentally escaped.


Ecology

The leaves of philodendrons are also known to be eaten by Venezuelan red howler monkeys, making up 3.1% of all the leaves they eat. The resins produced during the flowering of '' Monstera'' and ''Philodendron'' are known to be used by ''
Trigona ''Trigona'' is one of the largest genera of stingless bees, comprising about 32 species, exclusively occurring in the New World, and formerly including many more subgenera than the present assemblage; many of these former subgenera have been ele ...
'' stingless bees in the construction of their nests. The spathe provides a safe breeding area for beetles. As such, male beetles are often followed there by female beetles. The philodendrons benefit from this
symbiotic relationship Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
because the males will eventually leave the spathe covered in pollen and repeat the process at another philodendron, pollinating it in the process. Females that see a male beetle headed for a philodendron flower know he does so with intention of mating, and females which are sexually receptive and need to mate know that they can find males if they follow the pheromones produced by the philodendron flowers. As a result, the male beetles benefit from this relationship with the philodendrons because they do not have to produce pheromones to attract females. Additionally, male beetles benefit because they are ensured of mating with only sexually receptive females. Pheromones produced by the philodendrons may be similar to those produced by female beetles when they wish to attract males to mate. Also, the pheromones have a sweet, fruity smell in many species and no noticeable smell for others. Additionally, the beetles consume pollen from the fertile male flowers throughout the night, in addition to the sterile male flowers which are rich in
lipids Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins Vitamin A, A, Vitamin D, D, Vitamin E, E and Vitamin K, K), monoglycerides, diglycerides, phospholipids, and others. The fu ...
. Typically, five to 12 beetles will be within the spathe throughout the night. Rarely, cases of 200 beetles at a time have been observed and almost always the beetles are of the same species. Another feature of this symbiotic relationship, less well understood, is the series of events in which the spadix begins to heat up prior to the spathe opening up for the beetles. This process is known as
thermogenesis Thermogenesis is the process of heat production in organisms. It occurs in all warm-blooded animals, and also in a few species of thermogenic plants such as the Eastern skunk cabbage, the Voodoo lily ('' Sauromatum venosum''), and the giant w ...
. By the time the spathe is open and the beetles have arrived, the spadix is usually quite hot; up to around 46 °C in some species, but usually around 35 °C. The thermogenesis coincides with the arrival of the beetles and appears to increase their presence. The maximum temperature reached by the spadix remains about 20 °C higher than the outside ambient temperature. The time dependence of the temperature can vary from species to species. In some species, the temperature of the spadix will peak on the arrival of the beetles, then decrease, and finally increase reaching a maximum once again when the philodendron is ready for the beetles to leave. Other species, though, only show a maximum temperature on the arrival of the beetles, which remains roughly constant for about a day, and then steadily decreases. As the beetles home in on the inflorescence, they first move in a zig-zag pattern until they get reasonably close, when they switch to a straight-line path. The beetles may use scent to find the inflorescence when they are far away, but once within range, they find it by means of the infrared radiation, accounting for the two types of paths the beetles follow.


Cultivation


Growing

Philodendrons can be grown outdoors in mild climates in shady spots. They thrive in moist soils with high organic matter. In milder climates, they can be grown in pots of soil or in the case of ''Philodendron oxycardium'' in containers of water. Indoor plants thrive at temperatures between 15 and 18 °C and can survive at lower light levels than other house plants. Although philodendrons can survive in dark places, they much prefer bright lights. Wiping the leaves off with water will remove any dust and insects. Plants in pots with good root systems will benefit from a weak fertilizer solution every other week.


Propagation

New plants can be grown by taking stem cuttings with at least two joints. Cuttings then can be rooted in pots of sand and
peat moss ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
mixtures. These pots are placed in greenhouses with bottom heat of 21–24 °C. During the rooting, cuttings should be kept out of direct sunlight. Once rooted, the plants can be transplanted to larger pots or directly outside in milder climates. Stem cuttings, particularly from trailing varieties, can be rooted in water. In four to five weeks, the plant should develop roots and can be transferred to pots. Philodendrons can also propagate through air layering which is a more advanced method of propagation that involves creating a new plant on the stem of an existing plant. Hybridizing philodendrons is quite easy if flowering plants are available, because they have very few barriers to prevent hybridization. However, some aspects of making crosses can make philodendron hybridization more difficult. Philodendrons often flower at different times and the time when the spathe opens up varies from plant to plant. The pollen and the inflorescence both have short lives, which means a large collection of philodendrons is necessary if crossbreeding is to be done successfully. The pollen life can be extended to a few weeks by storing it in film canisters in a
refrigerator A refrigerator, commonly shortened to fridge, is a commercial and home appliance consisting of a thermal insulation, thermally insulated compartment and a heat pump (mechanical, electronic or chemical) that transfers heat from its inside to ...
. Artificial pollination is usually achieved by first mixing the pollen with water. A window is then cut into the spathe and the water-pollen mixture is rubbed on the fertile female flowers. The entire spathe is then covered in a plastic bag so the water–pollen mixture does not dry out; the bag is removed a few days later. If the inflorescence has not been fertilized, it will fall off, usually within a few weeks.


Toxicity

Philodendrons can contain as much as 0.7% of
oxalates Oxalate (systematic IUPAC name: ethanedioate) is an anion with the chemical formula . This dianion is colorless. It occurs naturally, including in some foods. It forms a variety of salts, for example sodium oxalate (), and several esters such as ...
in the form of
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula or . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydr ...
crystals as raphides. The risk of death, if even possible, is extremely low if ingested by an average adult, although its consumption is generally considered unhealthy. In general, the calcium oxalate crystals have a very mild effect on humans, and large quantities have to be consumed for symptoms to even appear. Possible symptoms include increased salivation, a sensation of burning of the mouth, swelling of the
tongue The tongue is a Muscle, muscular organ (anatomy), organ in the mouth of a typical tetrapod. It manipulates food for chewing and swallowing as part of the digestive system, digestive process, and is the primary organ of taste. The tongue's upper s ...
,
stomatitis Stomatitis is inflammation of the mouth and lips. It refers to any inflammatory process affecting the mucous membranes of the mouth and lips, with or without oral ulceration. In its widest meaning, stomatitis can have a multitude of different c ...
,
dysphagia Dysphagia is difficulty in swallowing. Although classified under " symptoms and signs" in ICD-10, in some contexts it is classified as a condition in its own right. It may be a sensation that suggests difficulty in the passage of solids or l ...
, an inability to speak, and
edema Edema (American English), also spelled oedema (British English), and also known as fluid retention, swelling, dropsy and hydropsy, is the build-up of fluid in the body's tissue (biology), tissue. Most commonly, the legs or arms are affected. S ...
. Cases of mild
dermatitis Dermatitis is a term used for different types of skin inflammation, typically characterized by itchiness, redness and a rash. In cases of short duration, there may be small blisters, while in long-term cases the skin may become thickened ...
due to contact with the leaves have also been reported, with symptoms including vesiculation and
erythema Erythema (, ) is redness of the skin or mucous membranes, caused by hyperemia (increased blood flow) in superficial capillaries. It occurs with any skin injury, infection, or inflammation. Examples of erythema not associated with pathology inc ...
. The chemical derivatives of alkenyl resorcinol are believed to be responsible for the dermatitis in some people. Contact with philodendron oils or fluids with the eyes have also been known to result in
conjunctivitis Conjunctivitis, also known as pink eye or Madras eye, is inflammation of the conjunctiva, the thin, clear layer that covers the white surface of the eye and the inner eyelid. It makes the eye appear pink or reddish. Pain, burning, scratchiness ...
. Fatal poisonings are extremely rare; one case of an infant eating small quantities of a philodendron resulting in hospitalization and death has been reported. This one case study, however, was found to be inconsistent with the findings from a second study. In this study, 127 cases of children ingesting philodendrons were studied, and they found only one child showed symptoms; a 10-month-old had minor upper lip swelling when he chewed on a philodendron leaf. The study also found the symptoms could subside without treatment and that previously reported cases of severe complications were exaggerated. In a 1961 study, 72 cases of cat poisonings were examined, of which 37 resulted in the death of the cat. The symptoms of the poisoned cats included excitability,
spasm A spasm is a sudden involuntary contraction of a muscle, a group of muscles, or a hollow organ, such as the bladder. A spasmodic muscle contraction may be caused by many medical conditions, including dystonia. Most commonly, it is a musc ...
s,
seizures A seizure is a sudden, brief disruption of brain activity caused by abnormal, excessive, or synchronous neuronal firing. Depending on the regions of the brain involved, seizures can lead to changes in movement, sensation, behavior, awareness, o ...
, kidney failure, and
encephalitis Encephalitis is inflammation of the Human brain, brain. The severity can be variable with symptoms including reduction or alteration in consciousness, aphasia, headache, fever, confusion, a stiff neck, and vomiting. Complications may include se ...
. In a 1978 study, three cats (two adults and one
kitten A kitten is a Juvenile (organism), juvenile cat. After being born, kittens display primary altriciality and are fully dependent on their mothers for #Establishing immunity, survival. They normally do not open their eyes for seven to ten days. A ...
) were tube-fed a puréed leaf and water mixture of '' P. cordatum'', then euthanized. A necropsy showed no signs or symptoms of acute poisoning or toxicity. Dosages of 2.8, 5.6, and 9.1 g/kg were used, the highest dose being much more than a house cat could consume. Earlier
epidemiological Epidemiology is the study and analysis of the distribution (who, when, and where), patterns and Risk factor (epidemiology), determinants of health and disease conditions in a defined population, and application of this knowledge to prevent dise ...
studies (which did not necessarily invoke a purée-and-water feeding method) were suggested to be wrong based on the premise that many of the sick cats in those studies may have had conditions and merely consumed philodendrons in an attempt to alleviate their illnesses. Some philodendrons are known to be toxic to mice and rats. In one study, 100 mg of ''P. cordatum'' leaves suspended in distilled water were fed to six mice. Three of the mice died. The same experiment was done with 100 mg of ''P. cordatum'' stems on three mice and none of them died. Leaves and flowers of ''P. sagittifolium'' were also orally administered in 100-mg doses to the mice. Three mice were used for each of the leaves and flowers; none of the mice died. A similar experiment was done on rats with the leaves and stems of ''P. cordatum'', but instead of oral administration of the dose, it was injected intraperitoneally using 3 g of plant
extract An extract (essence) is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol, oil or water. Extracts may be sold as tinctures or absolutes or dried and powdered. The aromatic principles of ma ...
from either the leaves or stems. Six rats were injected with the leaf extract and five of them died. Eight rats were injected with the stem extract and two of them died.


Uses

Indigenous people from South America use the resin from bees' nests (made from the species) to make their
blowgun A blowgun (also called a blowpipe or blow tube) is a simple ranged weapon consisting of a long narrow tube for shooting light projectiles such as darts. It operates by having the projectile placed inside the pipe and using the force created by ...
s air- and watertight. Though they contain calcium oxalate crystals, the berries of some species are eaten by the locals. For example, the sweet white berries of ''
Thaumatophyllum bipinnatifidum ''Thaumatophyllum bipinnatifidum'' (common names: split-leaf philodendron, lacy tree philodendron, selloum, horsehead philodendron, guaimbé) is a plant in the genus ''Thaumatophyllum'', in the family Araceae. Previously it was classified in the ...
'' are known to be used. Additionally, the aerial roots are also used for
rope A rope is a group of yarns, Plying, plies, fibres, or strands that are plying, twisted or braided together into a larger and stronger form. Ropes have high tensile strength and can be used for dragging and lifting. Rope is thicker and stronger ...
in this particular species. Also, in the making of a particular recipe for
curare Curare ( or ; or ) is a common name for various alkaloid arrow poisons originating from plant extracts. Used as a paralyzing agent by indigenous peoples in Central and South America for hunting and for therapeutic purposes, curare only ...
by the Amazonian Taiwanos, the leaves and stems of an unknown philodendron species are used. The leaves and stems are mixed with the bark of ''Vochysia ferruginea'' and with some parts of a species in the genus ''
Strychnos ''Strychnos'' is a genus of flowering plants, belonging to the family Loganiaceae (sometimes Strychnaceae). The genus includes about 200 accepted species of trees and lianas. The genus is widely distributed around the world's tropics and is n ...
''. Yet another use of philodendrons is for catching fish. A tribe in the Colombian Amazon is known to use '' P. craspedodromum'' to poison to the water, temporarily stunning the fish, which rise up to the surface, where they can be easily scooped up. To add the poison to the water, the leaves are cut into pieces and tied together to form bundles, which are allowed to
ferment Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic compound, Organic molecules, such as glucose or other sugars, are Catabo ...
for a few days. The bundles are crushed and added to the water into which the poison will dissipate. Although the toxicity of ''P. craspedodromum'' is not fully known, active ingredients in the poisoning of the fish possibly are
coumarins Coumarin derivatives are derivatives of coumarin and are considered phenylpropanoids. Among the most important derivatives are the 4-hydroxycoumarins, which exhibit anticoagulant properties, a characteristic not present for coumarin itself. ...
formed during the fermentation process. Some philodendrons are also used for ceremonial purposes. Among the Kubeo tribe, native to Colombia, '' P. insigne'' is used by
witch doctor A witch doctor (also spelled witch-doctor), or witchcraft doctor, is a kind of magical healer who treats ailments believed to be caused by witchcraft. The term is often misunderstood, and they could more accurately be called "anti-witch doctors ...
s to assuage ill patients. They use the juice of the
spathe In botany, a bract is a modified or specialized leaf, associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves in size, color, shape or texture. They also look ...
to stain their hands red, since many such tribes view the color red as a sign of power.


Notes


References

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External links


International Aroid Society: The genus ''Philodendron''

GardeningNorm - How to propagate philodendron: Step-by-step Guide
{{Authority control Araceae genera Epiphytes Low light plants