Orchidaceae (/ɔːrkəˈdeɪʃiː/ or-kə-DAY-shee[citation needed]), commonly called the orchid family, is a diverse and widespread family of flowering plants, with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering plants. The Orchidaceae have about 28,000 currently accepted species, distributed in about 763 genera. The determination of which family is larger is still under debate, because verified data on the members of such enormous families are continually in flux. Regardless, the number of orchid species is nearly equal to the number of bony fishes, more than twice the number of bird species, and about four times the number of mammal species.
The family encompasses about 6–11% of all seed plants. The largest genera are Bulbophyllum (2,000 species), Epidendrum (1,500 species), Dendrobium (1,400 species) and Pleurothallis (1,000 species). It also includes Vanilla (the genus of the vanilla plant), the type genus Orchis, and many commonly cultivated plants such as Phalaenopsis and Cattleya. Moreover, since the introduction of tropical species into cultivation in the 19th century, horticulturists have produced more than 100,000 hybrids and cultivars.
Description
Orchids are easily distinguished from other plants, as they share some very evident derived characteristics or synapomorphies. Among these are: bilateral symmetry of the flower (zygomorphism), many resupinate flowers, a nearly always highly modified petal (labellum), fused stamens and carpels, and extremely small seeds.
Stem and roots
All orchids are perennial herbs that lack any permanent woody structure. They can grow according to two patterns:
Monopodial: The stem grows from a single bud, leaves are added from the apex each year, and the stem grows longer accordingly. The stem of orchids with a monopodial growth can reach several metres in length, as in Vanda and Vanilla.
Sympodial: Sympodial orchids have a front (the newest growth) and a back (the oldest growth). The plant produces a series of adjacent shoots, which grow to a certain size, bloom and then stop growing and are replaced. Sympodial orchids grow laterally rather than vertically, following the surface of their support. The growth continues by development of new leads, with their own leaves and roots, sprouting from or next to those of the previous year, as in Cattleya. While a new lead is developing, the rhizome may start its growth again from a so-called 'eye', an undeveloped bud, thereby branching. Sympodial orchids may have visible pseudobulbs joined by a rhizome, which creeps along the top or just beneath the soil.
Neotinea lactea, collected in Sardinia; the small size, compared to a one-Euro coin, and the two globose tuberoids typical of the Neotinea genus are highlighted
Terrestrial orchids may be rhizomatous or form corms or tubers. The root caps of terrestrial orchids are smooth and white.
Some sympodial terrestrial orchids, such as Orchis and Ophrys, have two subterranean tuberous roots. One is used as a food reserve for wintry periods, and provides for the development of the other one, from which visible growth develops.
In warm and constantly humid climates, many terrestrial orchids do not need pseudobulbs.
Epiphytic orchids, those that grow upon a support, have modified aerial roots that can sometimes be a few meters long. In the older parts of the roots, a modified spongy epidermis, called a velamen, has the function of absorbing humidity. It is made of dead cells and can have a silvery-grey, white or brown appearance. In some orchids, the velamen includes spongy and fibrous bodies near the passage cells, called tilosomes.
The cells of the root epidermis grow at a right angle to the axis of the root to allow them to get a firm grasp on their support. Nutrients for epiphytic orchids mainly come from mineral dust, organic detritus, animal droppings and other substances collecting among on their supporting surfaces.
The pseudobulb of Prosthechea fragrans
The base of the stem of sympodial epiphytes, or in some species essentially the entire stem, may be thickened to form a pseudobulb that contains nutrients and water for drier periods.
The pseudobulb has a smooth surface with lengthwise grooves, and can have different shapes, often conical or oblong. Its size is very variable; in some small species of Bulbophyllum, it is no longer than two millimeters, while in the largest orchid in the world, Grammatophyllum speciosum (giant orchid), it can reach three meters. Some Dendrobium species have long, canelike pseudobulbs with short, rounded leaves over the whole length; some other orchids have hidden or extremely small pseudobulbs, completely included inside the leaves.
With ageing the pseudobulb sheds its leaves and becomes dormant. At this stage it is often called a backbulb. Backbulbs still hold nutrition for the plant, but then a pseudobulb usually takes over, exploiting the last reserves accumulated in the backbulb, which eventually dies off, too. A pseudobulb typically lives for about five years. Orchids without noticeable pseudobulbs are also said to have growths, an individual component of a sympodial plant.
Leaves
Like most monocots, orchids generally have simple leaves with parallel veins, although some Vanilloideae have reticulate venation. Leaves may be ovate, lanceolate, or orbiculate, and very variable in size on the individual plant. Their characteristics are often diagnostic. They are normally alternate on the stem, often folded lengthwise along the centre ("plicate"), and have no stipules. Orchid leaves often have siliceous bodies called stegmata in the vascular bundle sheaths (not present in the Orchidoideae) and are fibrous.
The structure of the leaves corresponds to the specific habitat of the plant. Species that typically bask in sunlight, or grow on sites which can be occasionally very dry, have thick, leathery leaves and the laminae are covered by a waxy cuticle to retain their necessary water supply. Shade-loving species, on the other hand, have long, thin leaves.
The leaves of most orchids are perennial, that is, they live for several years, while others, especially those with plicate leaves as in Catasetum, shed them annually and develop new leaves together with new pseudobulbs.
The leaves of some orchids are considered ornamental. The leaves of the Macodes sanderiana, a semiterrestrial or rock-hugging ("lithophyte") orchid, show a sparkling silver and gold veining on a light green background. The cordate leaves of Psychopsis limminghei are light brownish-green with maroon-puce markings, created by flower pigments. The attractive mottle of the leaves of lady's slippers from tropical and subtropical Asia (Paphiopedilum), is caused by uneven distribution of chlorophyll. Also, Phalaenopsis schilleriana is a pastel pink orchid with leaves spotted dark green and light green. The jewel orchid (Ludisia discolor) is grown more for its colorful leaves than its white flowers.
Some orchids, such as Dendrophylax lindenii (ghost orchid), Aphyllorchis and Taeniophyllum depend on their green roots for photosynthesis and lack normally developed leaves, as do all of the heterotrophic species.
Orchids of the genus Corallorhiza (coralroot orchids) lack leaves altogether and instead wrap their roots around the roots of mature trees and use specialized fungi to harvest sugars.
Flowers
The Orchidaceae are well known for the many structural variations in their flowers.
Some orchids have single flowers, but most have a racemose inflorescence, sometimes with a large number of flowers. The flowering stem can be basal, that is, produced from the base of the tuber, like in Cymbidium, apical, meaning it grows from the apex of the main stem, like in Cattleya, or axillary, from the leaf axil, as in Vanda.
As an apomorphy of the clade, orchid flowers are primitively zygomorphic (bilaterally symmetrical), although in some genera, such as Mormodes, Ludisia, and Macodes, this kind of symmetry may be difficult to notice.
The orchid flower, like most flowers of monocots, has two whorls of sterile elements. The outer whorl has three sepals and the inner whorl has three petals. The sepals are usually very similar to the petals (thus called tepals, 1), but may be completely distinct.
The medial petal, called the labellum or lip , which is always modified and enlarged, is actually the upper medial petal; however, as the flower develops, the inferior ovary or the pedicel usually rotates 180°, so that the labellum arrives at the lower part of the flower, thus becoming suitable to form a platform for pollinators. This characteristic, called resupination, occurs primitively in the family and is considered apomorphic, a derived characteristic all Orchidaceae share. The torsion of the ovary is very evident from the longitudinal section shown (below right). Some orchids have secondarily lost this resupination, e.g. Epidendrum secundum.
The normal form of the sepals can be found in Cattleya, where they form a triangle. In Paphiopedilum (Venus slippers), the lower two sepals are fused into a synsepal, while the lip has taken the form of a slipper. In Masdevallia, all the sepals are fused.
Orchid flowers with abnormal numbers of petals or lips are called peloric. Peloria is a genetic trait, but its expression is environmentally influenced and may appear random.
Orchid flowers primitively had three stamens, but this situation is now limited to the genus Neuwiedia. Apostasia and the Cypripedioideae have two stamens, the central one being sterile and reduced to a staminode. All of the other orchids, the clade called Monandria, retain only the central stamen, the others being reduced to staminodes . The filaments of the stamens are always adnate (fused) to the style to form cylindrical structure called the gynostemium or column . In the primitive Apostasioideae, this fusion is only partial; in the Vanilloideae, it is more deep; in Orchidoideae and Epidendroideae, it is total. The stigma is very asymmetrical, as all of its lobes are bent towards the centre of the flower and lie on the bottom of the column.
Pollen is released as single grains, like in most other plants, in the Apostasioideae, Cypripedioideae, and Vanilloideae. In the other subfamilies, which comprise the great majority of orchids, the anther carries two pollinia.
A pollinium is a waxy mass of pollen grains held together by the glue-like alkaloid viscin, containing both cellulosic strands and mucopolysaccharides. Each pollinium is connected to a filament which can take the form of a caudicle, as in Dactylorhiza or Habenaria, or a stipe, as in Vanda. Caudicles or stipes hold the pollinia to the viscidium, a sticky pad which sticks the pollinia to the body of pollinators.
At the upper edge of the stigma of single-anthered orchids, in front of the anther cap, is the rostellum , a slender extension involved in the complex pollination mechanism.
As mentioned, the ovary is always inferior (located behind the flower). It is three-carpelate and one or, more rarely, three-partitioned, with parietal placentation (axile in the Apostasioideae).
In 2011, Bulbophyllum nocturnum was discovered to flower nocturnally.
Asexuality
Some species, such as in the genera Phalaenopsis, Dendrobium, and Vanda, produce offshoots or plantlets formed from one of the nodes along the stem, through the accumulation of growth hormones at that point. These shoots are known as keiki.
Evolution
A study in the scientific journal Nature has hypothesised that the origin of orchids goes back much longer than originally expected. An extinct species of stingless bee, Proplebeia dominicana, was found trapped in Miocene amber from about 15-20 million years ago. The bee was carrying pollen of a previously unknown orchid taxon, Meliorchis caribea, on its wings. This find is the first evidence of fossilised orchids to date and shows insects were active pollinators of orchids then. This extinct orchid, M. caribea, has been placed within the extant tribe Cranichideae, subtribe Goodyerinae (subfamily Orchidoideae). An even older orchid species, Succinanthera baltica, was described from the Eocene Baltic amber by Poinar & Rasmussen (2017).
Genetic sequencing indicates orchids may have arisen earlier, 76 to 84 million years ago during the Late Cretaceous.According to Mark W. Chase et al. (2001), the overall biogeography and phylogenetic patterns of Orchidaceae show they are even older and may go back roughly 100 million years.
Using the molecular clock method, it was possible to determine the age of the major branches of the orchid family. This also confirmed that the subfamily Vanilloideae is a branch at the basal dichotomy of the monandrous orchids, and must have evolved very early in the evolution of the family. Since this subfamily occurs worldwide in tropical and subtropical regions, from tropical America to tropical Asia, New Guinea and West Africa, and the continents began to split about 100 million years ago, significant biotic exchange must have occurred after this split (since the age of Vanilla is estimated at 60 to 70 million years).
Genome duplication occurred prior to the divergence of this taxon
