Rhododendron Barbatum
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''Rhododendron'' (; from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
''rhódon'' "rose" and ''déndron'' "tree") is a very large
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of about 1,024
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of woody plants in the heath family (Ericaceae). They can be either
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
or
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 leaves, usually in the autumn; to the shedding of petals, aft ...
. Most species are native to eastern Asia and the
Himalaya The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 100 ...
n region, but smaller numbers occur elsewhere in Asia, and in North America, Europe and Australia. It is the
national flower In a number of countries, plants have been chosen as symbols to represent specific geographic areas. Some countries have a country-wide floral emblem; others in addition have symbols representing subdivisions. Different processes have been used to ...
of
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, the
state flower This is a list of U.S. state, federal district, and territory flowers. See also *List of U.S. state trees * Lists of U.S. state insignia References External linksList of state flowers {{USStateLists * U.S. state flowers Flowers ...
of
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and
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in the United States, the state flower of
Nagaland Nagaland () is a landlocked state in the northeastern region of India. It is bordered by the Indian states of Arunachal Pradesh to the north, Assam to the west, Manipur to the south and the Sagaing Region of Myanmar to the east. Its capital cit ...
in India, the provincial flower of
Jiangxi Jiangxi (; ; formerly romanized as Kiangsi or Chianghsi) is a landlocked province in the east of the People's Republic of China. Its major cities include Nanchang and Jiujiang. Spanning from the banks of the Yangtze river in the north int ...
in China and the
state tree This is a list of U.S. state, federal district, and territory trees, including official trees of the following of the states, of the federal district, and of the territories. Table See also * List of U.S. state, district, and territorial insign ...
of
Sikkim Sikkim (; ) is a state in Northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Province No. 1 of Nepal in the west and West Bengal in the south. Sikkim is also close to the Siligur ...
and
Uttarakhand Uttarakhand ( , or ; , ), also known as Uttaranchal ( ; the official name until 2007), is a state in the northern part of India. It is often referred to as the "Devbhumi" (literally 'Land of the Gods') due to its religious significance and ...
in India. Most species have brightly colored flowers which bloom from late winter through to early summer.
Azalea Azaleas are flowering shrubs in the genus ''Rhododendron'', particularly the former sections ''Tsutsusi'' (evergreen) and '' Pentanthera'' (deciduous). Azaleas bloom in the spring (April and May in the temperate Northern Hemisphere, and Octob ...
s make up two subgenera of ''Rhododendron''. They are distinguished from "true" rhododendrons by having only five
anther The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s per flower.


Species


Description

''Rhododendron'' is a genus of
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
s and small to (rarely) large
tree 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, including only woody plants with secondary growth, plants that are ...
s, the smallest species growing to tall, and the largest, ''R. protistum'' var. ''giganteum'', reported to tall. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
are spirally arranged; leaf size can range from to over , exceptionally in ''R. sinogrande''. They may be either
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
or
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 leaves, usually in the autumn; to the shedding of petals, aft ...
. In some species, the undersides of the leaves are covered with scales (lepidote) or hairs (indumentum). Some of the best known species are noted for their many clusters of large flowers. There are alpine species with small flowers and small leaves, and tropical species such as
section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sign ...
''
Vireya ''Vireya'' rhododendron ''Rhododendron'' section ''Vireya'' (vireyas) is a tropical group of ''Rhododendron'' species, numbering about 300 in all. Vireyas are native to southeastern Asia and range from Thailand to Australia. Description ''Vire ...
'' that often grow as
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s. Species in this genus may be part of the heath complex in oak-heath forests in eastern North America. They have frequently been divided based on the presence or absence of scales on the abaxial (lower) leaf surface (
lepidote A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
or elepidote). These scales, unique to subgenus ''Rhododendron'', are modified hairs consisting of a polygonal scale attached by a stalk. ''Rhododendron'' are characterised by having
inflorescences An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
with scarious (dry) perulae, a
chromosome number Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
of x=13,
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
that has a
septicidal Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that o ...
capsule, an
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
that is superior (or nearly so),
stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
that have no appendages, and agglutinate (clumped)
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
.


Taxonomy

''Rhododendron'' is the largest genus in the family
Ericaceae The Ericaceae are a family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with c.4250 known species spread across 124 genera, making it th ...
, with over 1000 species, (though estimates vary from 850 to 1200) and is morphologically diverse. Consequently, the
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
has been historically complex.


Early history

Although Rhododendrons had been known since the description of ''
Rhododendron hirsutum ''Rhododendron hirsutum'', commonly known as the hairy alpenrose is one of the species of ''Rhododendron'' native to the mountains of Europe. It occurs widely in the Alps except for the southwestern region (approximately south and west of the Ma ...
'' by
Charles de l'Écluse Charles de l'Écluse, L'Escluse, or Carolus Clusius (19 February 1526 – 4 April 1609), seigneur de Watènes, was an Artois doctor and pioneering botanist, perhaps the most influential of all 16th-century scientific horticulturists. Life Clu ...
(Clusius) in the sixteenth century, and were known to classical writers (Magor 1990), and referred to as ''Chamaerhododendron'' (low-growing rose tree), the genus was first formally described by
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
in his
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genera. It is the first work to consistently apply binomial names and was the ...
in 1753. He listed five species under ''Rhododendron'' (''
Rhododendron ferrugineum ''Rhododendron ferrugineum'', the alpenrose, snow-rose, or rusty-leaved alpenrose is an evergreen shrub that grows just above the tree line in the Alps, Pyrenees, Jura and northern Apennines, on acid soils. It is the type species for the genus ' ...
'' (
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
), '' R. dauricum'', '' R. hirsutum'', ''R. chamaecistus'' (now '' Rhodothamnus chamaecistus'' (L.) Rchb.) and '' R. maximum''). At that time he considered the then known six species of ''
Azalea Azaleas are flowering shrubs in the genus ''Rhododendron'', particularly the former sections ''Tsutsusi'' (evergreen) and '' Pentanthera'' (deciduous). Azaleas bloom in the spring (April and May in the temperate Northern Hemisphere, and Octob ...
'' that he had described earlier in 1735 in his
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
as a separate genus. Linnaeus' six species of ''Azalea'' were '' Azalea indica'', '' A. pontica'', '' A. lutea'', '' A. viscosa'', '' A. lapponica'' and ''A. procumbens'' (now ''
Kalmia procumbens ''Kalmia procumbens'', commonly known as alpine azalea or trailing azalea, is a dwarf shrub of high mountain regions of the Northern Hemisphere that usually grows no more than tall. Originally named by Linnaeus as ''Azalea procumbens'', it is al ...
''), which he distinguished from ''Rhododendron'' by having five
stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
, as opposed to ten. As new species of what are now considered ''Rhododendron'' were discovered, they were assigned to separate genera if they seemed to differ significantly from the type species. For instance ''Rhodora'' (Linnaeus 1763) for ''
Rhododendron canadense ''Rhododendron canadense'', the rhodora or Canada rosebay, is a deciduous flowering shrub that is native to northeastern North America. Classification Today's botanists consider the rhodora to be a distant relative of the other North American ...
'', ''Vireya'' (Carl Ludwig Blume, Blume 1826) and ''Hymenanthes'' (Blume 1826) for ''Rhododendron metternichii'', now Rhododendron degronianum, R. degronianum. Meanwhile, other botanists such as Richard Anthony Salisbury, Salisbury (1796) and Tate (1831) began to question the distinction between ''Azalea'' and ''Rhododendron'', and finally in 1836, ''Azalea'' was incorporated into ''Rhododendron'' and the genus divided into eight sections. Of these ''Tsutsutsi'' (''Rhododendron subg. Tsutsuji, Tsutsusi''), ''Pentanthera'', ''Pogonanthum'', ''Ponticum'' and ''Rhodora'' are still used, the other sections being ''Lepipherum'', ''Booram'', and ''Chamaecistus''. This structure largely survived till recently (2004), following which the development of molecular phylogeny led to major re-examinations of traditional morphological classifications, although other authors such as Candolle, who described six sections, used slightly different numeration. Soon, as more species became available in the nineteenth century so did a better understanding of the characteristics necessary for the major divisions. Chief amongst these were Maximovicz's ''Rhododendreae Asiae Orientali'' and Jules Émile Planchon, Planchon. Maximovicz used flower bud position and its relationship with leaf buds to create eight "Sections". Bentham and Hooker used a similar scheme, but called the divisions "Series". It was not until 1893 that Bernhard Adalbert Emil Koehne, Koehne appreciated the significance of scaling and hence the separation of lepidote and elepidote species. The large number of species that were available by the early twentieth century prompted a new approach when Isaac Bayley Balfour, Balfour introduced the concept of grouping species into Series (botany), series. ''The Species of Rhododendron'' referred to this series concept as the Balfourian system. That system continued up to modern times in Davidian's four volume ''The Rhododendron Species''.


Modern era

The next major attempt at classification was by Sleumer who from 1934 began incorporating the Balfourian series into the older hierarchical structure of subgenera and sections, according to the International Code of Botanical Nomenclature, culminating in 1949 with his "Ein System der Gattung ''Rhododendron'' L.", and subsequent refinements. Most of the Balfourian series are represented by Sleumer as subsections, though some appear as sections or even subgenera. Sleumer based his system on the relationship of the flower buds to the leaf buds, habitat, flower structure, and whether the leaves were lepidote or non-lepidote. While Sleumer's work was widely accepted, many in the United States and the United Kingdom continued to use the simpler Balfourian system of the Edinburgh group. Sleumer's system underwent many revisions by others, predominantly the Edinburgh group in their continuing Royal Botanic Garden Edinburgh notes. Cullen of the Edinburgh group, placing more emphasis on the lepidote characteristics of the leaves, united all of the lepidote species into subgenus ''Rhododendron'', including four of Sleumer's subgenera (''Rhododendron'', ''Pseudoazalea'', ''Pseudorhodorastrum'', ''Rhodorastrum''). In 1986 Philipson & Melva Philipson, Philipson raised two sections of subgenus ''Aleastrum'' (''Mumeazalea'', ''Candidastrum'') to subgenera, while reducing genus ''Therorhodion'' to a subgenus of ''Rhododendron''. In 1987 Spethmann, adding phytochemical features proposed a system with fifteen subgenera grouped into three 'chorus' subgenera. A number of closely related genera had been included together with ''Rhododendron'' in a former tribe, Rhodoreae. These have been progressively incorporated into ''Rhododendron''. Chamberlain and Rae moved the monotypic section ''Tsusiopsis'' together with the monotypic genus ''Tsusiophyllum'' into section ''Tsutsusi'', while Kron & Judd reduced genus ''Ledum'' to a subsection of section ''Rhododendron''. Then Judd & Kron moved two species (''Rhododendron schlippenbachii'', ''Rhododendron quinquefolium, R. quinquefolium'') from section ''Brachybachii'', subgenus ''Tsutsusi'' and two from section ''Rhodora'', subgenus ''Pentanthera'' (''Rhododendron albrechtii, R. albrechtii'', ''Rhododendron pentaphyllum, R. pentaphyllum'') into section ''Sciadorhodion'', subgenus ''Pentanthera''. Finally Chamberlain brought the various systems together in 1996, with 1,025 species divided into eight subgenera. For a comparison of the Sleumer and Chamberlain schemata see Table 1 of Goetsch (2005).


Phylogenetic analyses

The era of molecular analysis rather than descriptive features can be dated to the work of Kurashige (1988) and Kron (1997) who used Maturase K, matK sequencing. Later Gao ''et al.'' (2002) used Internal transcribed spacer, ITS sequences to determine a cladistic analysis. They confirmed that the genus ''Rhododendron'' was monophyletic, with subgenus ''Therorhodion'' in the Basal (phylogenetics), basal position, consistent with the matK studies. Following publication of the studies of Goetsch ''et al.'' (2005) with RPB2, there began an ongoing realignment of species and groups within the genus, based on evolutionary relationships. Their work was more supportive of Sleumer's original system than the later modifications introduced by Chamberlain ''et al.''. 2005 Annual ARS Convention The major finding of Goetsch and colleagues was that all species examined (except Rhododendron camtschaticum, ''R. camtschaticum'', subgenus ''Therorhodion'') formed three major clades which they labelled A, B and C, with the subgenera ''Rhododendron'' and ''Hymenanthes'' nested within clades A and B as monophyletic groups respectively. By contrast subgenera ''Azaleastrum'' and ''Pentanthera'' were polyphyletic, while ''R. camtschaticum'' appeared as a Sister group, sister to all other rhododendrons. The small polyphyletic subgenera ''Pentanthera'' and ''Azaleastrum'' were divided between two clades. The four sections of ''Pentanthera'' between clades B and C, with two each, while ''Azaleastrum'' had one section in each of A and C. Thus subgenera ''Azaleastrum'' and ''Pentanthera'' needed to be disassembled, and ''Rhododendron'', ''Hymenanthes'' and ''Tsutsusi'' correspondingly expanded. In addition to the two separate genera included under ''Rhododendron'' by Chamberlain (''Ledum'', ''Tsusiophyllum''), Goetsch ''et al.''. added ''Menziesia'' (Clade C). Despite a degree of paraphyly, the subgenus ''Rhododendron'' was otherwise untouched with regard to its three sections but four other subgenera were eliminated and one new subgenus created, leaving a total of five subgenera in all, from eight in Chamberlain's scheme. The discontinued subgenera are ''Pentanthera'', ''Tsutsusi'', ''Candidastrum'' and ''Mumeazalea'', while a new subgenus was created by elevating subgenus ''Azaleastrum'' section ''Choniastrum'' to subgenus rank. Subgenus ''Pentanthera'' (deciduous azaleas) with its four sections was dismembered by eliminating two sections and redistributing the other two between the existing subgenera in clades B (''Hymenanthes'') and C (''Azaleastrum''), although the name was retained in section ''Pentanthera'' (14 species) which was moved to subgenus ''Hymenanthes''. Of the remaining three sections, monotypic ''Viscidula'' was discontinued by moving ''Rhododendron nipponicum'' to ''Tsutsusi'' (C), while ''Rhodora'' (2 species) was itself polyphyletic and was broken up by moving ''
Rhododendron canadense ''Rhododendron canadense'', the rhodora or Canada rosebay, is a deciduous flowering shrub that is native to northeastern North America. Classification Today's botanists consider the rhodora to be a distant relative of the other North American ...
'' to section ''Pentanthera'' (B) and ''Rhododendron vaseyi'' to section ''Sciadorhodion'', which then became a new section of subgenus ''Azaleastrum'' (C). Subgenus ''Tsutsusi'' (C) was reduced to section status retaining the name, and included in subgenus ''Azaleastrum''. Of the three minor subgenera, all in C, two were discontinued. The single species of monotypic subgenus ''Candidastrum'' (''Rhododendron albiflorum'') was moved to subgenus ''Azaleastrum'', section ''Sciadorhodion''. Similarly the single species in monotypic subgenus ''Mumeazalea'' (''Rhododendron semibarbatum'') was placed in the new section ''Tsutsusi'', subgenus ''Azaleastrum''. Genus ''Menziesa'' (9 species) was also added to section ''Sciadorhodion''. The remaining small subgenus ''Therorhodion'' with its two species was left intact. Thus two subgenera, ''Hymenanthes'' and ''Azaleastrum'' were expanded at the expense of four subgenera that were eliminated, although ''Azaleastrum'' lost one section (''Choniastrum'') as a new subgenus, since it was a distinct subclade in A. In all, ''Hymenanthes'' increased from one to two sections, while ''Azaleastrum'', by losing one section and gaining two increased from two to three sections. (See schemata under #Subgenera, Subgenera) (Table 1.) Subsequent research has supported the revision by Goetsch, although has largely concentrated on further defining the phylogeny within the subdivisions. In 2011 the two species of ''Diplarche'' were also added to ''Rhododendron'', ''incertae sedis''.


Subdivision

This
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
has been progressively subdivided into a hierarchy of subgenus,
section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sign ...
, Subsection (botany), subsection, and
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
.


Subgenera

Terminology from the Sleumer (1949) system is frequently found in older literature, with five subgenera and is as follows; * Subgenus ''Lepidorrhodium'' Koehne: Lepidotes. 3 sections * Subgenus ''Eurhododendron'' Maxim.: Elipidotes. * Subgenus ''Pseudanthodendron'' Sleumer: Deciduous azaleas. 3 sections * Subgenus ''Anthodendron'' Rehder & Wilson: Evergreen azaleas. 3 sections * Subgenus ''Azaleastrum'' Planch.: 4 sections In the later traditional classification, attributed to Chamberlain (1996), and as used by horticulturalists and the American Rhododendron Society, ''Rhododendron'' has eight subgenera based on Morphology (biology), morphology, namely the presence of scales (lepidote), deciduousness of leaves, and the floral and vegetative branching patterns, after Sleumer (1980). These consist of four large and four small subgenera. The first two subgenera (''Rhododendron'' and ''Hymenanthes'') represent the species commonly considered as 'Rhododendrons'. The next two smaller subgenera (''Pentanthera'' and ''Tsutsusi'') represent the 'Azaleas'. The remaining four subgenera contain very few species. The largest of these is subgenus ''Rhododendron'', containing nearly half of all known species and all of the lepidote species. *Subgenus ''Rhododendron subg. Rhododendron, Rhododendron'' : Small leaf or lepidotes (scales on the underside of the leaves). 3 sections, 462 species,
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
: ''
Rhododendron ferrugineum ''Rhododendron ferrugineum'', the alpenrose, snow-rose, or rusty-leaved alpenrose is an evergreen shrub that grows just above the tree line in the Alps, Pyrenees, Jura and northern Apennines, on acid soils. It is the type species for the genus ' ...
''. *Subgenus ''Rhododendron subg. Hymenanthes, Hymenanthes'' : Large leaf or elepidotes (without scales). 1 section, 224 species, type ''Rhododendron degronianum''. *Subgenus ''Rhododendron subg. Pentanthera, Pentanthera'' : Deciduous azaleas. 4 sections, 23 species, type ''Rhododendron luteum''. *Subgenus ''Rhododendron sect. Tsutsusi, Tsutsusi'' : Evergreen azaleas. 2 sections, 80 species, type ''Rhododendron indicum''. *Subgenus ''Rhododendron subg. Azaleastrum, Azaleastrum'' : 2 sections, 16 species, type ''Rhododendron ovatum''. *Subgenus ''Candidastrum'' : 1 species, ''Rhododendron albiflorum''. *Subgenus ''Mumeazalea'' : 1 species, ''Rhododendron semibarbatum''. *Subgenus ''Therorhodion'' : 2 species (''Rhododendron camtschaticum'', ''Rhododendron redowskianun''). For a comparison of the Sleumer and Chamberlain systems, see Goetsch ''et al.'' (2005) Table 1. This division was based on a number of what were thought to be key morphological characteristics. These included the position of the inflorescence buds (terminal or lateral), whether lepidote or elepidote, deciduousness of leaves, and whether new foliage was derived from axils from previous year's shoots or the lowest scaly leaves (Table 2.). Following the cladistic analysis of Goetsch ''et al.'' (2005) this scheme was simplified, based on the discovery of three major Clade, clades (A, B, C) as follows. Clade A * Subgenus ''Rhododendron subg. Rhododendron, Rhododendron'' : Small leaf or lepidotes (scales on the underside of the leaves). 3 sections, about 400 species,
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
: ''
Rhododendron ferrugineum ''Rhododendron ferrugineum'', the alpenrose, snow-rose, or rusty-leaved alpenrose is an evergreen shrub that grows just above the tree line in the Alps, Pyrenees, Jura and northern Apennines, on acid soils. It is the type species for the genus ' ...
''. * Subgenus ''Rhododendron subg. Choniastrum, Choniastrum'' : 11 species Clade B * Subgenus ''Rhododendron subg. Hymenanthes, Hymenanthes'' : Large leaf or elepidotes (without scales), including deciduous azaleas. 2 sections, about 140–225 species, type ''Rhododendron degronianum''. Clade C * Subgenus ''Rhododendron subg. Azaleastrum, Azaleastrum'' : Evergreen azaleas. 3 sections, about 120 species, type ''Rhododendron ovatum''. Sister taxon * Subgenus ''Therorhodion'' : 2 species (''Rhododendron camtschaticum'', ''Rhododendron redowskianun'').


Sections and subsections

The larger subgenera are further subdivided into sections and subsections Some subgenera contain only a single section, and some sections only a single subsection. Shown here is the traditional classification, with species number after Chamberlain (1996), but this scheme is undergoing constant revision. Revisions by Goetsch ''et al.'' (2005) and by Craven ''et al.'' (2008) shown in (''parenthetical italics''). Older ranks such as Series (groups of species) are no longer used but may be found in the literature, but the American Rhododendron Society still uses a similar device, called Alliances *Subgenus ''Rhododendron'' L. (3 sections, 462 species: increased to five sections in 2008) **(''Discovereya (Sleumer) Argent, raised from Vireya'') **''Pogonathum'' Aitch. & Hemsl. (13 species; Himalaya and adjacent mountains) **(''Pseudovireya (C.B.Clarke) Argent, raised from Vireya'') **''Rhododendron (section), Rhododendron'' L. (149 species in 25 subsections; temperate to subarctic Northern Hemisphere) **''Rhododendron subgenus Vireya, Vireya'' (Blume) Copel.f. (300 species in 2 subsections; tropical southeast Asia, Australasia. At one time considered separate subgenus) *Subgenus ''Rhododendron subg. Hymenanthes, Hymenanthes'' (Blume) K.Koch (1 section, 224 species) (''Increased to two sections'') **''Ponticum'' (24 subsections) **(''Rhododendron sect. Pentanthera, Pentanthera'' (2 subsections – new section, moved from subgenus ''Pentanthera'') *Subgenus ''Pentanthera'' (4 sections, 23 species) (''Discontinued'') **''Rhododendron sect. Pentanthera, Pentanthera'' (2 subsections – moved to subgenus ''Hymenanthes'') **''Rhododendron canadense, Rhodora'' (L.) G. Don (2 species; ''
Rhododendron canadense ''Rhododendron canadense'', the rhodora or Canada rosebay, is a deciduous flowering shrub that is native to northeastern North America. Classification Today's botanists consider the rhodora to be a distant relative of the other North American ...
'', ''Rhododendron vaseyi'') (''Discontinued, redistributed'') **''Sciadorhodion'' Rehder & Wilson (4 species) (''Moved to subgenus Azaleastrum'') **''Viscidula'' Matsum. & Nakai (1 species; ''Rhododendron nipponicum'') (''Discontinued, added to section Tsutsusi, subgenus Azaleastrum'') *Subgenus ''Tsutsusi'' (Sweet) Pojarkova (2 sections, 80 species) (''Discontinued, reduced to section and moved to subgenus Azaleastrum'') **''Brachycalyx'' Sweet (3 alliances, 15 species) **''Tsutsusi'' (Sweet) Pojarkova (65 species) *Subgenus ''Azaleastrum'' Planch. (2 sections, 16 species) (''Increased to three sections'') **''Azaleastrum'' Planch. (5 species) **(''Choniastrum'' Franch. (11 species) (''Raised to subgenus'')) **(''Sciadorhodion Rehder & Wilson (4 species) (Moved from subgenus Pentanthera'')) **''(Tsutsusi (Sweet) Pojarkova (reduced from subgenus))'' *Subgenus ''Candidastrum'' Franch. (1 species: ''Rhododendron albiflorum'') (''Discontinued, moved to section Sciadorhodion, subgenus Azaleastrum'') *Subgenus ''Mumeazalea'' (Sleumer) W.R. Philipson & M.N. Philipson (1 species: ''Rhododendron semibarbatum'') (''Discontinued, moved to section Tsutsusi, subgenus Azaleastrum'') *Subgenus ''Therorhodion'' A. Gray (2 species) *(''Subgenus Choniastrum Franch. (11 species)'')


Distribution and habitat

Species of the genus ''Rhododendron'' are widely distributed between latitudes 80°N and 20°S and are native plant, native to areas from North America to Europe, Russia, and Asia, and from Greenland to Queensland, Australia and the Solomon Islands. The center of diversity, centres of diversity are in the Himalayas and Malaysia, with the greatest species diversity in the Sino-Himalayan region, Southwest China and northern Burma, from India – Himachal Pradesh,
Uttarakhand Uttarakhand ( , or ; , ), also known as Uttaranchal ( ; the official name until 2007), is a state in the northern part of India. It is often referred to as the "Devbhumi" (literally 'Land of the Gods') due to its religious significance and ...
,
Sikkim Sikkim (; ) is a state in Northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Province No. 1 of Nepal in the west and West Bengal in the south. Sikkim is also close to the Siligur ...
and
Nagaland Nagaland () is a landlocked state in the northeastern region of India. It is bordered by the Indian states of Arunachal Pradesh to the north, Assam to the west, Manipur to the south and the Sagaing Region of Myanmar to the east. Its capital cit ...
to
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
, northwestern Yunnan and western Sichuan and southeastern Tibet. Other significant areas of diversity are in the mountains of Korea, Japan and Taiwan. More than 90% of ''Rhododendron'' ''sensu'' Chamberlain belong to the Asian subgenera ''Rhododendron'', ''Hymenanthes'' and section ''Tsutsusi''. Of the first two of these, the species are predominantly found in the area of the Himalayas and Southwest China (Sino-Himalayan Region). The 300 Tropical climate, tropical species within the ''
Vireya ''Vireya'' rhododendron ''Rhododendron'' section ''Vireya'' (vireyas) is a tropical group of ''Rhododendron'' species, numbering about 300 in all. Vireyas are native to southeastern Asia and range from Thailand to Australia. Description ''Vire ...
'' section of subgenus ''Rhododendron'' occupy the Malay archipelago from their presumed Southeast Asian origin to Northern Australia, with 55 known species in Borneo and 164 in New Guinea. The species in New Guinea are native to Subalpine zone, subalpine moist grasslands at around 3,000 metres above sea level in the Central Highlands. Subgenera ''Rhododendron'' and ''Hymenanthes'', together with section ''Pentanthera'' of subgenus ''Pentanthera'' are also represented to a lesser degree in the Mountainous areas of North America and Western Eurasia. Subgenus ''Tsutsusi'' is found in the maritime regions of East Asia (Japan, Korea, Taiwan, East China), but not in North America or Eurasia. In the United States, native ''Rhododendron'' mostly occur in lowland and montane forests in the Pacific Northwest, California, the Northeastern United States, Northeast, and the Appalachian Mountains.


Ecology


Invasive species

''Rhododendron ponticum'' has become invasive in Republic of Ireland, Ireland and the United Kingdom. It is an introduced species, spreading in woodland areas and replacing the natural understory. ''R. ponticum'' is difficult to eradicate, as its roots can make new shoots.


Insects

A number of insects either target rhododendrons or will opportunistically attack them. Rhododendron borers and various weevils are major pests of rhododendrons, and many caterpillars will preferentially devour them. ''Rhododendron'' species are used as food plants by the larvae (caterpillars) of some butterflies and moths; see List of Lepidoptera that feed on rhododendrons.


Diseases

Major diseases include ''Phytophthora'' root rot, stem and twig fungal dieback. Rhododendron bud blast, a fungal condition that causes buds to turn brown and dry before they can open, is caused by the fungus ''Pycnostysanus azaleae'', which may be brought to the plant by the rhododendron leafhopper, ''Graphocephala fennahi''.


Cultivation

Both species and hybrid rhododendrons (including azaleas) are used extensively as ornamental plants in landscaping in many parts of the world, including both temperate and wiktionary:subtemperate, subtemperate regions. Many species and cultivars are grown commercially for the Nursery (horticulture), nursery trade. Rhododendrons can be propagated by air layering or stem cuttings. They can self-propagate by sending up shoots from the roots. Sometimes an attached branch that has drooped to the ground will root in damp mulch, and the resulting rooted plant then can be cut off the parent rhododendron. Rhododendrons are often valued in landscaping for their structure, size, flowers, and the fact that many of them are evergreen. Azaleas are frequently used around foundations and occasionally as hedges, and many larger-leafed rhododendrons lend themselves well to more informal plantings and woodland gardens, or as specimen plants. In some areas, larger rhododendrons can be pruned to encourage more tree-like form, with some species such as ''Rhododendron arboreum'' and ''Rhododendron falconeri, R. falconeri'' eventually growing to a height of or more.


Commercial growing

Rhododendrons are grown commercially in many areas for sale, and are occasionally collected in the wild, a practice now rare in most areas. Larger commercial growers often ship long distances; in the United States, most of them are on the west coast (Oregon, Washington state and California). Large-scale commercial growing often selects for different characteristics than hobbyist growers might want, such as resistance to root rot when overwatered, ability to be forced into budding early, ease of rooting or other propagation, and saleability.


Horticultural divisions

Horticulturally, rhododendrons may be divided into the following groups: * Evergreen rhododendrons - large group of evergreen shrubs that vary greatly in size. Most rhododendron flowers are bell-shaped and have 10 stamens. *
Vireya ''Vireya'' rhododendron ''Rhododendron'' section ''Vireya'' (vireyas) is a tropical group of ''Rhododendron'' species, numbering about 300 in all. Vireyas are native to southeastern Asia and range from Thailand to Australia. Description ''Vire ...
(Malesian) rhododendrons: epiphyte, epiphytic hardiness (plants), tender shrubs * Azaleas – group of shrubs which have smaller and thinner leaves than evergreen rhododendrons. They are generally medium-sized shrubs with smaller funnel-shaped flowers that usually have 5 stamens: ** Deciduous hybrid azaleas: *** Exbury hybrids – derived from the Knap Hill hybrids, developed by Lionel de Rothschild (born 1882), Lionel de Rothschild at the Exbury Gardens, Exbury Estate in England. *** Ghent (Gandavense) hybrids – Belgian raised *** Knap Hill hybrids – developed by Anthony Waterer at the Knaphill, Knap Hill Nursery in England. *** Mollis hybrids – Dutch and Belgian raised *** New Zealand Ilam hybrids – derived from Knap Hill/Exbury hybrids *** Occidentale hybrids – English raised *** Rustica Flore Pleno hybrids – sweet-scented, double-flowered ** Evergreen hybrid azaleas: *** Gable hybrids – raised by Joseph B. Gable in Pennsylvania. *** Glenn Dale hybrids – US raised complex hybrids *** Indian (Indica) hybrids – mostly of Belgian origin *** Kaempferi hybrids – Dutch raised *** Kurume hybrids – Japanese raised *** Kyushu hybrids – very hardy Japanese azaleas (to −30 °C) *** Oldhamii hybrids – dwarf hybrids raised at Exbury, England *** Satsuki hybrids – Japanese raised, originally for bonsai *** Shammarello hybrids – raised in northern Ohio *** Vuyk (Vuykiana) hybrids – raised in the Netherlands * Azaleodendrons – semi-evergreen hybrids between deciduous azaleas and rhododendrons


Planting and care

Like other Ericaceae, ericaceous plants, most rhododendrons prefer acid soils with a pH of roughly 4.5–5.5; some tropical Vireyas and a few other rhododendron species grow as
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s and require a planting mix similar to orchids. Rhododendrons have fibrous roots and prefer well-drained soils high in organic material. In areas with poorly drained or alkaline soils, rhododendrons are often grown in raised beds using media such as composted pine bark. Mulching and careful watering are important, especially before the plant is established. A new calcifuge, calcium-tolerant stock of rhododendrons (trademarked as 'Inkarho') has been exhibited at the Royal Horticultural Society, RHS Chelsea Flower Show in London (2011). Individual hybrids of rhododendrons have been grafting, grafted on to a rootstock on a single rhododendron plant that was found growing in a chalk quarry. The rootstock is able to grow in calcium-rich soil up to a pH of 7.5.


Hybrids

Rhododendrons are extensively hybridized in cultivation, and natural hybrids often occur in areas where species ranges overlap. There are over 28,000 cultivars of Rhododendron in th
International Rhododendron Registry
held by the Royal Horticultural Society. Most have been bred for their flowers, but a few are of garden interest because of ornamental leaves and some for ornamental bark or stems. Some hybrids have fragrant flowers—such as the Loderi hybrids, created by crossing ''Rhododendron fortunei'' and ''Rhododendron griffithianum, R. griffithianum''. Other examples include the PJM hybrids, formed from a cross between ''Rhododendron carolinianum'' and ''Rhododendron dauricum, R. dauricum'', and named after Peter J. Mezitt of Weston Nurseries, Massachusetts.


Uses


Pharmacology

''Rhododendron'' species have long been used in traditional medicine. Animal studies and ''in vitro'' research have identified possible anti-inflammatory and hepatoprotective activities which may be due to the antioxidant effects of flavonoids or other natural phenol, phenolic compounds and saponins the plant contains. Xiong ''et al.'' have found that the root of the plant is able to reduce the activity of NF-κB in rats.


Toxicology

Some species of rhododendron are poisonous to grazing animals because of a toxin called grayanotoxin in their
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
and nectar (plant), nectar. People have been known to become ill from eating honey made by bees feeding on rhododendron and azalea flowers. Xenophon described the odd behaviour of Greeks, Greek soldiers after having consumed honey in a village surrounded by ''Rhododendron ponticum'' during the march of the Ten Thousand (Greek), Ten Thousand in 401 BCE. Pompey's soldiers reportedly suffered lethal casualties following the consumption of honey made from ''Rhododendron'' deliberately left behind by Pontic Greeks, Pontic forces in 67 BCE during the Third Mithridatic War. Later, it was recognized that honey resulting from these plants has a slightly hallucinogenic and laxative effect. The suspect rhododendrons are ''Rhododendron ponticum'' and ''Rhododendron luteum'' (formerly ''Azalea pontica''), both found in northern Asia Minor. Eleven similar cases during the 1980s have been documented in Istanbul, Turkey. Rhododendron is extremely toxic to horses, with some animals dying within a few hours of ingesting the plant, although most horses tend to avoid it if they have access to good forage. The effects of ''R. ponticum'' were mentioned in the 2009 film ''Sherlock Holmes (2009 film), Sherlock Holmes'' as a proposed way to arrange a fake execution. It was also mentioned in the third episode of Season 2 of BBC's ''Sherlock (TV series), Sherlock'', speculated to have been a part of Sherlock's fake death scheme.


Culture


Symbolism

In the language of flowers, the Rhododendron symbolizes danger and to beware. ''Rhododendron arboreum'' (''lali guransh'') is the national flower of
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
. ''R. ponticum'' is the Floral emblem, state flower of Jammu and Kashmir (state), Indian-administered Kashmir and Azad Kashmir, Pakistan-controlled Kashmir. ''Rhododendron niveum'' is the state tree of
Sikkim Sikkim (; ) is a state in Northeastern India. It borders the Tibet Autonomous Region of China in the north and northeast, Bhutan in the east, Province No. 1 of Nepal in the west and West Bengal in the south. Sikkim is also close to the Siligur ...
in India. Rhododendron arboreum is also the state tree of the state of
Uttarakhand Uttarakhand ( , or ; , ), also known as Uttaranchal ( ; the official name until 2007), is a state in the northern part of India. It is often referred to as the "Devbhumi" (literally 'Land of the Gods') due to its religious significance and ...
, India. Pink Rhododendron (''Rhododendron campanulatum'') is the state flower of Himachal Pradesh, India. Rhododendron is also the provincial flower of Jiangxi, China and the state flower of
Nagaland Nagaland () is a landlocked state in the northeastern region of India. It is bordered by the Indian states of Arunachal Pradesh to the north, Assam to the west, Manipur to the south and the Sagaing Region of Myanmar to the east. Its capital cit ...
, the 16th state of the Indian Union. ''Rhododendron maximum'', the most widespread rhododendron of the Appalachian Mountains, is the state flower of the US state of
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the Bur ...
, and is in the Flag of West Virginia. ''Rhododendron macrophyllum'', a widespread rhododendron of the Pacific Northwest, is the state flower of the US state of
Washington Washington commonly refers to: * Washington (state), United States * Washington, D.C., the capital of the United States ** A metonym for the federal government of the United States ** Washington metropolitan area, the metropolitan area centered o ...
.


Literature

The nineteenth-century American poet and essayist Ralph Waldo Emerson in 1834 wrote a poem titled "The Rhodora, On Being Asked, Whence Is the Flower", In James Joyce, Joyce's ''Ulysses (novel), Ulysses'', rhododendrons play an important role in Leopold and Molly's early courtship: Molly remembers them in her soliloquy – "the sun shines for you he said the day we were lying among the rhododendrons on Howth head in the grey tweed suit and his straw hat the day I got him to propose to me". Jasper Fforde a British author, also uses rhododendron as a motif throughout many of his published books. See ''Thursday Next'' series, and ''Shades of Grey 1: The Road to High Saffron, Shades of Grey''. Amongst the Zomi tribes in India and Myanmar, "Rhododendrons" called "Ngeisok" is used in a poetic manner to signify a lady. In Daphne Du Maurier's novel ''Rebecca (novel), Rebecca'', the character of Rebecca is associated with "blood red" rhododendrons throughout the novel, perhaps due to the toxic roots of the plant mirroring the poisonous character of Rebecca. On the other hand, azaleas (a type of rhododendron) represent the second Mrs. De Winter. In the young adult novel ''Walk Two Moons'' by Sharon Creech, the protagonist, Sal, and her friend, Phoebe, construct an elaborate story in which they suspect Phoebe's neighbor of murdering her husband and burying his body beneath a rhododendron in her yard. Chapter 14 of the book is entitled "The Rhododendron".


Culinary

The rhododendron is the national flower of
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
, where the flower is considered edible and enjoyed for its sour taste. The pickled flower can last for months and the flower juice is also marketed. The flower, fresh or dried, is added to fish curry in the belief that it will soften the bones. The juice of rhododendron flower is used to make a Squash (drink), squash called burans (named after the flower) in the hilly regions of
Uttarakhand Uttarakhand ( , or ; , ), also known as Uttaranchal ( ; the official name until 2007), is a state in the northern part of India. It is often referred to as the "Devbhumi" (literally 'Land of the Gods') due to its religious significance and ...
. It is admired for its distinctive flavor and color.


Labrador tea

The herbal tea called Labrador tea (not a true tea) is made from one of three closely related species: * ''Rhododendron tomentosum'' (Northern Labrador tea, previously ''Ledum palustre'') * ''Rhododendron groenlandicum'', (Bog Labrador tea, previously ''Ledum groenlandicum'' or ''Ledum latifolium'') * ''Rhododendron neoglandulosum'', (Western Labrador tea, or trapper's tea, previously ''Ledum glandulosum'')


Conservation

In the UK the forerunner of the Rhododendron, Camellia and Magnolia Group (RCMG), The Rhododendron Society was founded in 1916. while in Scotland species are being conserved by the Rhododendron Species Conservation Group.


See also

* List of Award of Garden Merit rhododendrons * List of Rhododendron diseases * List of Rhododendron species * List of Sections in Subgenus Rhododendron


References


Bibliography


Books and book chapters

* (also available online a
Gallica
* * * * In four volumes: Vol. I. ''Lepidotes'' , Vol. II. ''Elepidotes. Arboreum-Lacteum'' , Vol. III. ''Elepidotes Continued, Neriiflorum-Thomsonii, Azaleastrum and Camtschaticum'' , Vol. IV. ''Azaleas'' . * . * *


Articles

* * *


Subdivisions


Azaleas

* *


Tsutsusi

* * * * *


Vireya

* . A reprint from Flora Malesiana ser. I, vol. 6, part 4. Pages 473 through 674. * * * * Yearbook of the Rhododendron Species Foundation, Federal Way, WA. * * * * *


Separate genera

*


Additional resources

Records of the Rhododendron Society of America reside at the Albert and Shirley Small Special Collections Library at the University of Virginia.


External links


History of Rhododendron Discovery & Culture












* [http://www.vireya.net/ Information on Vireyas]
Information+photos of hybrids and species

Information on Rhododendrons by Marc Colombel, founder of the Société Bretonne du Rhododendron.


* [https://web.archive.org/web/20090130005655/http://www.rhodo.citymax.com/page/page/627471.htm History of Rhododendrons] * [http://www.rododendron.cz/fotogalerie-rododendronu/pruhonicky-park-2011/ Rhododendron in botanical garden Pruhonice-Czech republic ]


Databases

* USDA Plants Database
''Rhododendron''
* ITIS Report
''Rhododendron''

''eFloras.org''
*
Flora of North America: Rhododendron
*
Flora of China: Rhododendron
*
Annotated Checklist of the Flowering Plants of Nepal: Rhododendron


Societies


American Rhododendron Society
*
The Quarterly Bulletin of the American Rhododendron Society 1947–1981
*
Journal of the American Rhododendron Society (JARS) 1982–
** Information Source: .
The Rhododendron, Camellia & Magnolia Group of the Royal Horticultural Society

Rhododendron Species Foundation and Botanical Garden



Australian Rhododendron Society

German Rhododendron Society

New Zealand Rhododendron Association

Danish Rhododendron Society

Fraser South Rhododendron Society


Botanical gardens


Royal Botanic Garden Edinburgh: Rhododendrons at the four Gardens

National Rhododendron Gardens Melbourne Australia
{{Authority control Rhododendron, Ericaceae genera Plants used in traditional Chinese medicine National symbols of Nepal Taxa named by Carl Linnaeus Extant Ypresian first appearances