Laurales
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- Related Topics:
- Lauraceae
- Monimiaceae
- Amborellaceae
- Siparunaceae
- Atherospermataceae
Laurales, the laurel order of flowering plants, containing 7 families, 91 genera, and about 2,900 species. Members of Laurales are trees, shrubs, or woody vines. Most are found in tropical or warm temperate climates, and they are especially abundant in regions with moist equable climates. Lumber, medicinal extracts such as camphor, and essential oils for perfume are derived from some Laurales species, and several are important ornamentals.
Members of Laurales are characterized by woodiness, aromatic parts, and a single strand of conducting tissues continuing from the stem into the leaf. Along with the orders Magnoliales, Piperales, and Canellales, Laurales forms the magnoliid clade, which is an early evolutionary branch in the angiosperm tree; the clade corresponds to part of the subclass Magnoliidae under the old Cronquist botanical classification system. The families in Laurales are Atherospermataceae, Calycanthaceae, Gomortegaceae, Hernandiaceae, Lauraceae, Monimiaceae, and Siparunaceae. Lauraceae and Monimiaceae together constitute most of the genera in this order.
Distribution and abundance
Lauraceae, or the laurel family, contains 50 genera, more than half the genera in the order, and about eight-ninths of the species (2,500). Lauraceae is distributed throughout tropical and subtropical regions; principally Southeast Asia and tropical America, particularly Brazil. The majority of the species occur in only 7 genera: Ocotea has about 400 species in tropical America, South Africa, and the Mascarene Islands; Litsea has more than 200 species in Asia, Australasia, and America; Cryptocarya and Cinnamomum (the source of camphor and the spice cinnamon) contain about 350 species and 250 species, respectively; Persea (including the avocado plant) has about 100 species; and Beilschmiedia contains about 250 species throughout many tropical regions as well as Australia and New Zealand. Persea and Cryptocarya are found in many tropical regions, and Cinnamomum is widely distributed in all the major tropical and subtropical regions. The roughly 100 species of the largely tropical and subtropical genus Lindera include the North American spicebush (L. benzoin).
Cassytha, a rootless vinelike stem parasite with vestigial scalelike leaves, is the most unusual member of the family; the genus contains 15–20 species native to the Old World. Laurus (laurel) consists of two species, one of which is L. nobilis (sweet bay tree, or bay laurel), a native of the Mediterranean. The leaves of the bay laurel were once formed into laurel crowns by the ancient Greeks. Sassafras, one of the few economically important genera of the family, has two species in eastern Asia and one in eastern North America; oil of sassafras was once used medicinally, and Amerindians made a tea from the bark and twigs. The family is of great importance in the tropics for its valuable lumber, derived from many different species. Some of the wood remains fragrant for decades after it is cut.
The second largest family, Monimiaceae, has 22 genera and 200 species, less than 10 percent of Laurales species. This family is also found in tropical and subtropical regions, but it is less extensively distributed and occurs mainly in the warmer areas of the Southern Hemisphere. The type genus, Monimia, is restricted to the Mascarene Islands.
The family Siparunaceae includes 75 species in two genera. Glossocalyx, from tropical West Africa, has four species. The remainder of the species in the family are in the genus Siparuna, found in Mexico, Central America, and tropical South America.
The remaining four families have a combined total of 83 species. Hernandiaceae (55 species) is a pantropical family of trees, shrubs, and some lianas. The largest genus, Hernandia (22 species), is distributed in Central and South America, the West Indies, West Africa, Indo-Malaysia (a region comprising India, South China, and Southeast Asia), and the Pacific Islands. Atherospermataceae includes 6 or 7 genera and 16 species, which are native to Australia, New Guinea, New Zealand, New Caledonia, and Chile. Calycanthaceae, or the strawberry shrub family, has a discontinuous distribution: Calycanthus (strawberry shrub, sweet shrub, or Carolina allspice) is found in California and in the southeastern United States, and Chimonanthus and Sinocalycanthus occur in China. The single species of Idiospermum is a very rare evergreen species from Queensland, Austl. Gomortegaceae, or the queule family, consists of a single species, Gomortega keule, which is a rare species native to central Chile.
Economic and ecological importance
Lauraceae
Lauraceae is by far the most economically important family in Laurales. Persea americana (avocado) is a highly nutritious fruit, rich in proteins and fats and low in sugar. The total food value of avocado is high; it provides nearly twice the energy of an equivalent weight of meat and an abundance of several vitamins, such as A, B, C, D, and E. There are several wild species of Persea in Central America. Cultivated varieties were developed by people in the region of modern Mexico and Guatemala thousands of years ago. (Seeds found in the caves of the Tehuacán Valley, south of Mexico City, have been determined to be nearly 10,000 years old and are cited as proof of the early use of the avocado fruit by humans.)
Avocado trees are of medium size, generally not exceeding about 20 metres (65 feet) in height, with simple evergreen elliptical leaves 15 to 20 cm (6 to 8 inches) long. Mature fruits can be spherical and about 8 cm (3 inches) long or pear-shaped and up to 22 cm (9 inches) long. The fruit has a large central woody seed, typically the size of a hen’s egg. There are a number of cultivars of avocados, each of which can be placed into one of three groups. Fruits of Mexican species have dark, smooth skin, and the trees are hardy, capable of withstanding cold weather to − 6 °C (21 °F) and poor growing conditions. Guatemalan species are a little less resistant, withstanding temperatures only to about − 4.5 °C (24 °F), and produce large fruits with thick, rough skins. The West Indian species are the most susceptible of all to cold weather, succumbing to temperatures below − 2 °C (28 °F); they produce large fruits with smooth, tough skins. Some species are picked when the fruits are beginning to soften; others, like the Hass and Fuerte cultivars, remain hard until picked.
The largest avocado plantations are in California and Florida, where a number of varieties have been developed. The United States produces about 10 percent of the world’s supply of avocados. A serious disease of avocado trees caused by the fungus Phytophthora cinnamomi affects trees grown in soils with a high degree of moisture. The fungus invades the vascular system of the roots, and, in most cases, the entire tree eventually dies.
The leaves of the Mediterranean Laurus nobilis (bay laurel) are dried and used as a flavouring for cooking, particularly for meat and fish dishes. A fat extracted from the seeds is used to make soap. Cinnamon spice is derived from the inner bark of Cinnamomum zeylanicum, the cinnamon tree, a native of Sri Lanka and southern India. The bark is removed from two-year-old shoots during the monsoon season, as at that time the vascular cambium is actively growing and the bark can be taken off more easily. Extraneous outer tissue is removed, and the bark is dried to form quills or ground to make powder. Several thousand tons are produced annually, mostly from Sri Lanka, Madagascar, and the Seychelles. Cinnamon oil is distilled from bark chips and used to alleviate stomach upset. Cinnamon was used by the ancient Egyptians during the embalming process. Eugenol, an oil distilled from the green leaves, is used as a substitute for clove oil, as an ingredient in some perfumes, and as a flavouring for sweets, foods, and toothpaste. Camphor is derived from Cinnamomum camphora, the camphor tree, of China, Taiwan, and Japan. It is obtained by steam distillation of wood chips. The wood of the camphor tree may contain up to 5 percent of the crude oil, and a single tree can yield up to 3 tons of the oil, which settles from the distillate and crystallizes. The oil can be redistilled to yield other compounds, notably safrole, which is used in perfumes and for flavourings. Camphor was one of the raw materials used in making celluloid, which has now been replaced by other plastics. Camphor is employed in pharmaceuticals, especially liniments, and in insecticides.
Many other species of Cinnamomum have uses as spices and medicines. Cinnamomum cambodianum bark is used to make joss sticks, which are burned as incense. Oil of sassafras, as much as 80 percent composed of the compound safrole, was previously distilled in large quantities from the bark enclosing the roots of Sassafras albidum (also called S. officinale), a plant native to Canada and the United States. This oil once served as a flavouring for sweets, medicines, toothpastes, root beer, and sarsaparilla, a drink derived from the genus Smilax (family Smilacaceae). However, the U.S. Food and Drug Administration (FDA) banned the use of sassafras oil when it was realized that the substance is a mild carcinogen.
To say that the wood of all trees of Lauraceae is suitable for industrial purposes seems to be only a slight exaggeration. Most of the best-known timbers of Lauraceae have been depleted through overexploitation, however, and are not likely to remain economically important in the future unless serious conservation efforts are undertaken. Many species of the widespread genus Ocotea have been utilized for timber. Chlorocardium rodiei (formerly Ocotea rodiei), commonly known as greenheart, an olive-green to black wood from northern South America, is a very durable, strong, dense wood ideally suited to underwater applications, such as boats and wharf pilings. Bebeerine, a highly poisonous alkaloid produced as a secondary compound, has been extracted from several species of Ocotea, as well as from greenheart. Ocotea venenosa is a source of a poison used for the tips of arrows by Brazilian natives. Because alkaloids are present in many woods of Lauraceae, timber workers who process them are susceptible to dermatitis and serious irritations of the respiratory tract.
Other families
Calycanthus floridus (Carolina allspice) and C. occidentalis (California allspice), both members of Calycanthaceae, are grown as ornamental shrubs and valued for their sweetly fragrant summer flowers. The aromatic bark of C. floridus is used as a spice. Chimonanthus praecox (also called C. fragrans, and commonly known as wintersweet) is a cultivated shrub that flowers in winter before the leaves are produced. The light yellow flowers are popular for their spicy fragrance. The beautiful creamy, pink infused flowers of Sinocalycanthus have captured the interest of horticulturists.
Various members of the family Monimiaceae are important locally for their timber and fruits and in making perfumes, medicine, and dyes. Peumus boldus, native to Chile, is the source of boldo wood, a hardwood used in cabinetmaking. A dye is obtained from its bark, and the leaves contain an essential oil and the alkaloid boldine, which are employed medicinally as a digestive aid and stimulant. The leaves of Doryphora sassafras and D. aromatica, both known in eastern Australia as sassafras, produce a sarsaparilla-like odour when crushed. An essential oil containing safrole is distilled from the leaves and bark of D. sassafras and used in perfumery, and the fragrant wood is used in furniture making and wood turning.
A decoction of the bark of Siparuna cujabana (family Siparunaceae) from Brazil is used by local residents to induce sweating and as an abortifacient.
The South American species Laurelia sempervirens (sometimes called L. aromatica), from the family Atherospermataceae, is known as Chile laurel or Peruvian nutmeg, and its seeds are ground up and used as a spice. Laurelia novae-zelandiae is used in New Zealand for boat building and furniture making. It yields a light, hard wood that is difficult to split and that dents rather than breaks upon impact. The bark contains an alkaloid, pukateine (after pukatea, the Maori name for the plant), that has strong pain-killing properties, similar to morphine. At one time the bark was boiled in water and used to treat ulcers, skin ailments (including boils and ulcers), toothache, and neuralgia.
Characteristic morphological features
Despite the great diversity of structure among families of the order, some structural features common to all distinguish Laurales from other orders. Except for the twining, rootless stem parasite Cassytha (family Lauraceae), all members of the Laurales order are woody, with a primitive nodal anatomy (arrangement of vascular bundles at the juncture of leaf and stem) of the type called unilacunar, and all have ethereal (aromatic) oil cells and pollen grains having either two apertures or no apertures. Members of Laurales characteristically have perigynous or epigynous flowers. In perigynous flowers the semi-inferior ovary region is surrounded by the hypanthium, a cup-shaped extended receptacle, on the rim of which the perianth and stamens are inserted. In epigynous flowers the ovary is enclosed by the hypanthium and fused to it, and the perianth and stamens arise from the top of the hypanthium above the inferior ovary. The stamens of many members have nectar-bearing appendages, and, in most species, the anthers release pollen by means of valves. Staminodia, reduced stamens that do not produce pollen, are commonly present between the stamens and carpels. The female structures usually have only a single carpel. Laurales is closely related to the order Magnoliales. However, unlike members of Magnoliales, which generally have primitive leaflike carpels and stamens, most Laurales species have more-specialized floral organs.
Lauraceae
The vast majority of species of Lauraceae differ from the other families of Laurales in possessing leaves that are alternately arranged or whorled, although a few have opposite leaves. They resemble members of Calycanthaceae in having a seed with a large embryo and no endosperm at maturity. Pollen of Lauraceae species is inaperturate and surrounded by a reduced exine; it is, therefore, seldom found in the fossil record because it decays so readily. Leaves of Lauraceae are usually leathery and evergreen with numerous ethereal oil cavities, which accounts for the aromatic nature of many species. The generally small green, yellow, or white flowers are usually arranged in clusters, and the floral parts develop in multiples of three. The perianth is not differentiated into sepals and petals. There are between 3 and 12 stamens per flower, and the filament of each stamen often has paired nectariferous appendages attached near the base, as in many Monimiaceae species. Stamens may have two (Beilschmiedia) or four (Litsea) pollen sacs, each with valvular flap dehiscence, again in common with various members of Monimiaceae. Unlike the latter family, however, the flowers of Lauraceae have a single carpel, and the hypanthium is short. The single-seeded fruits are mostly fleshy berries or drupes, and they often have a smooth cupule surrounding the base akin to the cap of an acorn. Most of the species are strongly aromatic because of ethereal oil cells in the leaves, wood, and bark.
Monimiaceae
Members of Monimiaceae are evergreen trees or shrubs, rarely woody vines (lianas). The leaves are simple and mostly oppositely arranged. The flowers are unisexual or bisexual and are usually perigynous with a well-developed receptacle. The tepals are inconspicuous and rarely differentiated into sepals and petals. The stamens have two or four pollen sacs that open either by longitudinal slits or by the outward bending and lifting upward of oval flaps of tissue, hinged at the tip of each sac (valvular dehiscence). Paired ear-shaped appendages, often attached near the base of the short filaments, act as nectaries. The female flowers may have sterile stamens (staminodes), with attached nectaries to attract pollinators. There are numerous carpels (as many as 2,000), each with a single ovule, and the outer carpels of female flowers are sometimes sterile. After fertilization, an enlarged perigynous receptacle may enclose the fruits; this aggregate fruit splits open irregularly in a number of species to expose the individual drupelets (small fleshy fruits with a single seed inside).
Other families
Members of Siparunaceae are trees or woody vines with opposite, mostly serrate leaves. The flowers are unisexual; pollen-bearing flowers and ovule-bearing flowers occur either on the same plant or different plants, depending on the species. In this family, glands are not present at the base of the stamens, and the stamen number varies from one to many. The hypanthium becomes woody and splits when mature, exposing the fleshy fruits (drupes). Glossocalyx species have two forms of leaves, differing in shape and size, at the same nodes. The flowers are small and are either bisexual or unisexual.
Atherospermataceae species also have opposite, serrate leaves. There are as many stamens as perianth parts. The hypanthium becomes woody and splits when mature. The dry fruits (achenes) have a tuft of hair.
Gomortega keule, the only member of the family Gomortegaceae, has an inferior ovary and bisexual flowers with only two or three carpels that are fused to form a compound ovary. As in many Monimiaceae species, the pollen sacs of the stamens have valvular dehiscence.
The members of Calycanthaceae differ from most of the other families in Laurales in having seeds with a large embryo and little if any endosperm at maturity. Except for Idiospermum, the leaves of Calycanthaceae species tend to be thinner and softer than other members of Laurales because they are deciduous plants of the temperate zone. The pollen sacs on the numerous stamens dehisce by longitudinal slits, and the pollen is biaperturate. There are 1 to 35 carpels per flower. Except in Idiospermum, the hypanthium becomes woody as it matures, and dry fruits (achenes) fall from the open top. In Idiospermum the embryo has three or four large, fleshy cotyledons.
Hernandiaceae shares a number of features with Lauraceae, including alternate leaves (which are sometimes lobed or palmately compound) and a single carpel per flower. Members of the family also have inaperturate pollen and develop stamens with valvular dehiscence and nectariferous appendages. Hernandiaceae differ in having an inferior ovary and indehiscent dry fruits (which are found in a very few Lauraceae).
Evolution
Although closely related to Magnoliales, most members of Laurales are more advanced than the majority of Magnoliales species in several respects. Floral evolution has advanced to perigyny in most members of Laurales and even to epigyny (inferior ovaries) and fused carpels in some members. Instead of being predominantly uniaperturate, pollen is inaperturate or biaperturate. The number of ovules per carpel has been reduced; in fact, there is only a single functional ovule in many Laurales. The stamens are not leaflike, and many have the unusual feature of valvular dehiscence. In addition, many stamens have paired appendages near the base of the stamen filament, which function as nectaries in most plants. The morphological nature of these appendages is unclear—they may have arisen de novo (Latin for “anew,” a term used in biology to refer to a genetic mutation), or each may be the remains of a sterile stamen.
Pollination ecology in Laurales is similar to that in Magnoliales. Insect pollination predominates, although there is evidence that wind pollination occurs in some of the members of Monimiaceae. Bird pollination does not seem to take place. As in Magnoliales, the habit has not evolved beyond trees, shrubs, and vines. The leaves of Laurales species are almost all simple, sometimes toothed, but seldom lobed; compound leaves are rare. A feature common to all members of Laurales is unilacunar nodal anatomy; trilacunar or multilacunar nodal anatomy occurs in Magnoliales. The two orders are so closely related that several families have been shifted from one order to the other by various authorities.
F. Bruce Sampson Paul E. Berry