Lauraceae, or the laurels, is a plant family that includes the true laurel and its closest relatives. This family comprises about 2850 known species in about 45 genera worldwide.[3] They are dicotyledons, and occur mainly in warm temperate and tropical regions, especially Southeast Asia and South America. Many are aromatic evergreen trees or shrubs, but some, such as Sassafras, are deciduous, or include both deciduous and evergreen trees and shrubs, especially in tropical and temperate climates. The genus Cassytha is unique in the Lauraceae in that its members are parasiticvines. Most laurels are highly poisonous.
The family has a worldwide distribution in tropical and warm climates. The Lauraceae are important components of tropical forests ranging from low-lying to montane. In several forested regions, Lauraceae are among the top five families in terms of the number of species present.
The Lauraceae give their name to habitats known as laurel forests, which have many trees that superficially resemble the Lauraceae, though they may belong to other plant families such as Magnoliaceae or Myrtaceae. Laurel forests of various types occur on most continents and on many major islands.
Although the taxonomy of the Lauraceae is still not settled, conservative estimates suggest some 52 genera worldwide, including 3,000 to 3,500 species.[4] Compared to other plant families, the taxonomy of Lauraceae still is poorly understood. This is partly due to its great diversity, the difficulty of identifying the species, and partly because of inadequate investment in taxonomic work.[4]
Recent monographs on small and medium-sized genera of Lauraceae (up to about 100 species) have revealed many new species.[4] Similar increases in the numbers of species recognised in other larger genera are to be expected.[4]
The fruits of Lauraceae are drupes, one-seeded fleshy fruit with a hard layer, the endocarp, surrounding the seed. However, the endocarp is very thin, so the fruit resemble a one-seeded berry.[5] The fruit in some species (particularly in the genera Ocotea) are partly immersed or covered in a cup-shaped or deep thick cupule, which is formed from the tube of the calyx where the peduncle joins the fruit; this gives the fruit an appearance similar to an acorn. In some Lindera species, the fruit have a hypocarpium at the base of the fruit.[citation needed]
Distribution and uses
Because the family is so ancient and was so widely distributed on the Gondwanasupercontinent, modern species commonly occur in relict populations isolated by geographical barriers, for instance on islands or tropical mountains. Relict forests retain endemicfauna and flora in communities of great value in inferring the palaeontological succession and climate change that followed the breakups of the supercontinents.
Many Lauraceae contain high concentrations of essential oils, some of which are valued for spices and perfumes. Within the plants, most such substances are components of irritant or toxic sap or tissues that repel or poison many herbivorous or parasitic organisms.
Conversely, some species, though commercially valuable in some countries, are regarded as aggressive invaders in other regions. For example, Cinnamomum camphora, though a valued ornamental and medicinal plant, is so invasive as to have been declared a weed in subtropical forested areas of South Africa.[10]
Ecology
Lindera melissifolia: This endangered species is native to the southeastern United States, and its demise is associated with habitat loss from extensive drainage of wetlands for agriculture and forestry.
Lauraceae flowers are protogynous, often with a complex flowering system to prevent inbreeding. The fruits are an important food source for birds, on which some Palaeognathae are highly dependent.[citation needed] Other birds that rely heavily on the fruit for their diets include members of the families Cotingidae, Columbidae, Trogonidae, Turdidae, and Ramphastidae, amongst others. Birds that are specialised frugivores tend to eat the whole fruit and regurgitate seeds intact, thereby releasing the seeds in favourable situations for germination (ornithochory). Some other birds that swallow the fruit pass the seed intact through their guts.
The leaves of some species in the Lauraceae have domatia in the axils of their veins. The domatia are home to certain mites. Other lauraceous species, members of the genus Pleurothyrium in particular, have a symbiotic relationship with ants that protect and defend the tree. Some Ocotea species are also used as nesting sites by ants, which may live in leaf pockets or in hollowed-out stems.[13]
Defense mechanisms that occur among members of the Lauraceae include irritant or toxic sap or tissues that repel or poison many herbivorous organisms.
Trees of the family predominate in the world's laurel forests[citation needed] and cloud forests,[citation needed] which occur in tropical to mild temperate regions of both northern and southern hemispheres. Other members of the family however, occur pantropically in general lowland and Afromontane forest, and in Africa for example there are species endemic to countries such as Cameroon, Sudan, Tanzania, Uganda and Congo. Several relict species in the Lauraceae occur in temperate areas of both hemispheres. Many botanical species in other families have similar foliage to the Lauraceae due to convergent evolution, and forests of such plants are called laurel forest. These plants are adapted to high rainfall and humidity, and have leaves with a generous layer of wax, making them glossy in appearance, and a narrow, pointed-oval shape with a 'drip tip', which permits the leaves to shed water despite the humidity, allowing transpiration to continue. Scientific names similar to Daphne (e.g., Daphnidium, Daphniphyllum)[14] or "laurel" (e.g.,Laureliopsis, Skimmia laureola) indicate other plant families that resemble Lauraceae.
Some Lauraceae species have adapted to demanding conditions in semiarid climates, but they tend to depend on favorable edaphic conditions, for example, perennial aquifers, periodic groundwater flows, or periodically flooded forests in sand that contains hardly any nutrients. Various species have adapted to swampy conditions by growing pneumatophores, roots that grow upward, that project above the levels of periodic floods that drown competing plants which lack such adaptations.[15]
Paleobotanists have suggested the family originated some 174±32 million years ago (Mya), while others[16] do not believe they are older than the mid-Cretaceous. Fossil flowers attributed to this family occur in Cenomanian clays (mid-Cretaceous, 90-98 Mya) of the Eastern United States (Mauldinia mirabilis). Fossils of Lauraceae are common in the Tertiary strata of Europe and North America, but they virtually disappeared from central Europe in the Late Miocene.[17] Because of their unusual fragility, the pollens of Lauraceae do not keep well and have been found only in relatively recent strata.
Deciduous Lauraceae lose all of their leaves for part of the year depending on variations in rainfall. The leaf loss coincides with the dry season in tropical, subtropical, and arid regions.
Laurel wilt disease, caused by the virulent fungal pathogen Raffaelea lauricola, a native of southern Asia, was found in the southeast United States in 2002. The fungus spreads between hosts via a wood-boring beetle, Xyleborus glabratus, with which it has a symbiotic relationship. Several Lauraceae species are affected. The beetle and disease are believed to have arrived in the US via infected solid wood packing material, and have since spread to several states.[18]
Classification within the Lauraceae is not fully resolved. Multiple classification schemes based on a variety of morphological and anatomical characteristics have been proposed, but none are fully accepted. According to Judd et al. (2007),[19] the suprageneric classification proposed by van der Werff and Richter (1996)[20] is currently the authority. However, due to an array of molecular and embryological evidence that disagrees with the groupings, it is not fully accepted by the scientific community. Their classification is based on inflorescence structure and wood and bark anatomy. It divides Lauraceae into two subfamilies, Cassythoideae and Lauroideae. The Cassythoideae comprise a single genus, Cassytha, and are defined by their herbaceous, parasitic habit. The Lauroideae are then divided into three tribes: Laureae, Perseeae, and Cryptocaryeae.[citation needed]
The subfamily Cassythoideae is not fully supported. Backing has come from matK sequences of chloroplast genes[21] while a questionable placement of Cassytha has been concluded from analysis of intergenetic spacers of chloroplast and nuclear genomes.[22] Embryological studies also appear contradictory. One study by Heo et al. (1998)[23] supports the subfamily. It found that Cassytha develops an ab initio cellular-type endosperm and the rest of the family (with one exception) develops a nuclear-type endosperm. Kimoto et al. (2006)[24] suggest Cassytha should be placed in the tribe Cryptocaryeae because it shares a glandular anther tapetum and an embryo sac protruding from the nucellus with other members of the Cryptocaryeae.
The tribes Laureae and Perseeae are not well supported by any molecular or embryological studies. Sequences of the matK chloroplast gene,[21] as well as sequences of chloroplast and nuclear genomes,[22] reveal close relationships between the two tribes. Embryological evidence does not support a clear division between the two tribes, either. Genera such as Caryodaphnopsis and Aspidostemon that share embryological characteristics with one tribe and wood and bark characteristics or inflorescence characteristics with another tribe blur the division of these groups.[23] All available evidence, except for inflorescence morphology and wood and bark anatomy, fails to support separate tribes Laureae and Perseeae.
The tribe Cryptocaryeae is partially supported by molecular and embryological studies. Chloroplast and nuclear genomes support a tribal grouping that contains all the genera circumscribed by van der Weff and Richter (1996),[20] as well as three additional genera.[22] Partial support for the tribe is also attained from the matK sequences of chloroplast genes[21] as well as embryology.[25]
Challenges in Lauraceae classification
The knowledge of the species comprising the Lauraceae is incomplete. In 1991, about 25-30% of neotropical Lauraceae species had not been described.[26] In 2001, embryological studies had only been completed on individuals from 26 genera yielding a 38.9% level of knowledge, in terms of embryology, for this family.[25] Additionally, the huge amount of variation within the family poses a major challenge for developing a reliable classification.[20][26]
Phytochemistry
The adaptation of Lauraceae to new environments has followed a long evolutionary journey which has led to many specializations, including defensive or deterrent systems against other organisms.
A laurel wreath, a round or horseshoe-shaped wreath made of connected laurel branches and leaves, is an ancient symbol of triumph in classical Western culture originating in Greek mythology, and is associated in some countries with academic or literary achievement.
Related Research Articles
Laurel forest, also called laurisilva or laurissilva, is a type of subtropical forest found in areas with high humidity and relatively stable, mild temperatures. The forest is characterized by broadleaf tree species with evergreen, glossy and elongated leaves, known as "laurophyll" or "lauroid". Plants from the laurel family (Lauraceae) may or may not be present, depending on the location.
Ocotea is a genus of flowering plants belonging to the family Lauraceae. Many are evergreen trees with lauroid leaves.
Cryptocarya is a genus of about 360 species of flowering plants in the laurel family, Lauraceae. Most species are trees, occasionally shrubs, distributed through the Neotropical, Afrotropical, Indomalayan, and Australasian realms. Most plants in the genus Cryptocarya have leaves arranged alternately along the stems, small flowers with 6 tepals, stamens in 2 rows, the inner row alternating with staminodes, and the fruit is a drupe.
Cassytha is a genus of some two dozen species of obligately parasitic vines in the family Lauraceae. Superficially, and in some aspects of their ecology, they closely resemble plants in the unrelated genus Cuscuta, the dodders. When fruit and flowers are absent in the field, the physical resemblance is so close that few people without technical training can discern the difference. In this respect and in their ecology the two genera present a spectacular example of convergent evolution. Nonetheless, Nickrent comments that "Cassytha is uneqivocally assigned to Lauraceae based on (both) morphological and molecular data." In its divergence from habits typical of the Lauraceae, Cassytha also presents examples of mosaic evolution
Neocinnamomum is a genus of flowering plants belonging to the family Lauraceae. They are evergreen shrubs or small trees, indigenous to Bhutan, China, India, Indonesia (Sumatra), Myanmar, Nepal, Thailand, and Vietnam.
Caryodaphnopsis is a genus of 16 species belonging to the flowering plant family Lauraceae, distributed in tropical areas in southern North America, northern South America, and East and Southeast Asia.
Nectandra is a genus of plant in the family Lauraceae. They are primarily Neotropical, with Nectandra coriacea being the only species reaching the southernmost United States. They have fruit with various medical effects. Sweetwood is a common name for some plants in this genus.
Ocotea foetens, commonly called til or stinkwood is a species of tree in the family Lauraceae. It is evergreen and grows up to 40 m tall. It is a common constituent of the laurisilva forests of Madeira and the Canary Islands. Leaf fossils of this species are known from the Mio-Pleistocene of Madeira Island.
Ocotea rugosa is a species of evergreen tree to 10 m (33 ft) tall in the plant genus Ocotea, in the family Lauraceae. It is endemic to Andean Ecuador at an altitude of 1,700 to 2,500 m. Its natural habitat is subtropical or tropical moist montane forests and cloud forest. This species requires moisture and protection of other trees for growing. The principal threats are fires, grazing, and the conversion of forest to farmland.
Magnoliids, Magnoliidae or Magnolianae are a clade of flowering plants. With more than 10,000 species, including magnolias, nutmeg, bay laurel, cinnamon, avocado, black pepper, tulip tree and many others, it is the third-largest group of angiosperms after the eudicots and monocots. The group is characterized by trimerous flowers, pollen with one pore, and usually branching-veined leaves.
Apollonias barbujana, the Canary laurel or barbusano, is perhaps the only species of flowering plants belonging to the genus Apollonias of the laurel family, Lauraceae. It is endemic to the Macaronesian islands of Madeira and the Canary Islands. Molecular phylogenies have found that the species is nested within the genus Persea, closely related to Persea americana (avocado).
Aspidostemon is a genus of flowering plants belonging to the family Lauraceae. It occurs in Madagascar.
Cinnadenia is a flowering plant genus belonging to the family Lauraceae. They are present in low and mountain cloud forests in Southeast Asia.
Endlicheria is a neotropical plant genus consisting of approximately 60 species, occurring mostly in northern South America and the Amazon region. Most species are medium-sized trees, sometime up to 40 metres in height, but a few species are shrubs. DNA molecular data shows that it is closely related to Rhodostemonodaphne and Ocotea.
Rhodostemonodaphne is a genus of flowering plants in the family Lauraceae. It is a neotropical genus consisting of approximately 41 species occurring in Central America and northern South America. This genus has many species that are valued for timber. The classification of the genus is unclear since the species in the genus fall into a well-supported but unresolved clade that also includes species with unisexual flowers currently placed in the genera Endlicheria and part of Ocotea.
Kubitzia is an American neotropical flowering plant genus in the family Lauraceae with two species from South America.
Potoxylon is a monotypic genus of evergreen trees belonging to the Laurel family, Lauraceae. Its only species, Potoxylon melagangai, is endemic to Borneo.
Cinnamomum kinabaluense is a species of plant in the laurel family (Lauraceae). It is endemic to Borneo.
Cinnamomum lawang is a species of tree in the laurel family (Lauraceae). It is endemic to Borneo.
Kuloa ikonyokpe is a species of tree in the laurel family (Lauraceae). It is endemic to Cameroon.
↑ Little, Stefan A.; Stockey, Ruth A.; Penner, Bonnie (March 2009). "Anatomy and development of fruits of Lauraceae from the Middle Eocene Princeton Chert". American Journal of Botany. 96 (3): 637–651. doi:10.3732/ajb.0800318. PMID21628220. S2CID38272445.
↑ Henderson, Lesley (2001). Alien Weeds and Invasive Plants: A Complete Guide to Declared Weeds and Invaders in South Africa, Including Another 36 Species Invasive in that Region. Plant Protection Research Institute. ISBN978-1-86849-192-6.[pageneeded]
↑ Stout, Jean (1979). "An Association of an Ant, a Mealy Bug, and an Understory Tree from a Costa Rican Rain Forest". Biotropica. 11 (4): 309–311. doi:10.2307/2387924. JSTOR2387924.
↑ Lübbe, W. A. (1991-06-01). "The Response of Ocotea bullata (Lauraceae) to Flooded Conditions". South African Forestry Journal. 157 (1): 32–37. doi:10.1080/00382167.1991.9629097. ISSN0038-2167.
↑ Judd, Walter S.; Campbell, Christopher S.; Kellogg, Elizabeth Anne (2008). Plant Systematics: A Phylogenetic Approach. W. H. Freeman. ISBN978-0-87893-407-2.[pageneeded]
1 2 Kimoto, Y.; H. Tobe (2001). "Embryology of Laurales: a review and perspectives". Journal of Plant Research. 114 (3): 247–261. doi:10.1007/PL00013988. S2CID31592261.
Meisner, C. F. (1864). "Lauraceae". Prodromus Systematis Naturalis Regni Vegetabilis. 15 (1): 1–260. NAID10007643152.
Mez, Carl Christian (1889). "Lauraceae Americanae Monographice Descripsit". Jahrbuch des Königlichen Botanischen Gartens und des Botanischen Museums zu Berlin. 5: 1–556.
Rohwer, Jens G. in Kubitzki, K.(Editor) 1993. The Families and Genera of Vascular Plants, Vol.2: K. Kubitzki, J. G. Rohwer & V. Bittrich, 366-390. ISBN3-540-55509-9
Wagner, W. L., D. R. Herbst, and S. H. Sohmer. 1990. Manual of the Flowering Plants of Hawai'i. Spec. Publ. 83. University of Hawaii Press and Bishop Museum Press. Bishop Museum. 1854 pp.
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