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Rhododendron

Rhododendron ferrugineum
Scientific classification
Kingdom: Plantae
(unranked): Angiosperms
(unranked): Eudicots
(unranked): Asterids
Order: Ericales
Family: Ericaceae
Subfamily: Ericoideae
Genus: Rhododendron
L. Sp. Pl. i 392 (1753)
[1]
Type species
Rhododendron ferrugineum
L.
Subgenera
[2]
Azaleastrum
Choniastrum
Hymenanthes
Rhododendron
Therorhodion
- Former subgenera -
Candidastrum
Mumeazalea
Pentanthera
From Wikipedia, the free encyclopedia
Rhododendron (from Ancient Greek rhdon "rose" and
dndron "tree")
[3][4]
is a genus of 1,024 species of woody plants in the
heath family (Ericaceae), either evergreen or deciduous, and found
mainly in Asia. Most species have showy flowers. Azaleas make up two
subgenera of Rhododendron. They are distinguished from "true"
rhododendrons by having only five anthers per flower.
1 Description
2 Taxonomy
2.1 Early history
2.2 Modern era
2.3 Phylogenetic analyses
2.4 Subdivision
2.4.1 Subgenera
2.4.2 Sections and subsections
2.4.3 Species
3 Distribution and habitat
4 Ecology
4.1 Invasive species
4.2 Insects
4.3 Diseases
5 Cultivation
5.1 Commercial growing
5.2 Horticultural divisions
5.3 Planting and care
5.4 Hybrids
6 Uses
6.1 Pharmacology
6.2 Toxicology
7 Culture
7.1 Symbolism
7.2 Literature
7.3 Culinary
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Tsutsusi 7.4 Labrador tea
8 See also
9 References
10 Bibliography
10.1 Books and book chapters
10.2 Articles
10.3 Subdivisions
10.3.1 Azaleas
10.3.2 Tsutsusi
10.3.3 Vireya
10.3.4 Separate genera
11 External links
11.1 Databases
11.2 Flora
11.3 Rhododendron societies
11.4 Botanical gardens
Rhododendron is a genus characterised by shrubs and small to (rarely) large trees, the smallest species growing
to 10100 cm (3.939.4 in) tall, and the largest, R. protistum var. giganteum, reported to 30 m (98 ft) tall.
[5][6]
The leaves are spirally arranged; leaf size can range from 12 cm (0.390.79 in) to over 50 cm (20 in),
exceptionally 100 cm (39 in) in R. sinogrande. They may be either evergreen or deciduous. 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 Vireya that often grow as epiphytes. Species in this genus may be
part of the heath complex in oak-heath forests in eastern North America.
[7][8]
They have frequently been
divided based on the presence or absence of scales on the abaxial (lower) leaf surface (lepidote or elepidote).
These scales, unique to subgenus Rhododendron, are modified hairs consisting of a polygonal scale attached by
a stalk.
[2]
Rhododendron are characterised by having inflorescences with scarious (dry) perulae, a chromosome number of
x=13, fruit that has a septicidal capsule, an ovary that is superior (or nearly so), stamens that have no
appendages, and agglutinate (clumped) pollen.
[9]
The Rhododendron genus is the largest of the genera in the Ericaceae family, with 1,024 species,
[10]
though
estimates vary from 850-1000 depending on the authority used, (Fayaz 2012) and is morphologically diverse.
Consequently the taxonomy has been historically complex.
[9]
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Rhododendron wardii var. puralbum
Rhododendron in Japan
A garden with tall Rhododendrons in
Lynnwood, Washington
Rhododendron forest in Nepal
Early history
Although Rhododendrons had been known since the description of
Rhododendron hirsutum by Charles de l'cluse (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 in his Species Plantarum in 1753.
[1][11]
He listed five species under Rhododendron (Rhododendron ferrugineum
(type species), 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
[12]
that he had
described earlier in 1735 in his Systema Naturae as a separate genus.
[13][14]
Linnaeus' six species of Azalea were Azalea indica, A. pontica, A. lutea,
A. viscosa, A. lapponica and A. procumbens (now Kalmia procumbens),
which he distinguished from Rhododendron by having five stamens, as
opposed to ten. As new species of what are now considered
Rhododendron were discovered, if they seemed to differ significantly
from the type species they were assigned to separate genera. For
instance Rhodora for Rhododendron canadense (Linnaeus 1763), Vireya
(Blume 1826)
[15]
and Hymenanthes for Rhododendron metternichhii,
now R. degronianum (1826). Meanwhile other botanists such as
Salisbury (1796)
[16]
and Tate (1831)
[17]
began to question the distinction
between Azalea and Rhododendron, and finally in 1836, Azalea was
incorporated into Rhododendron (Don 1834)
[18]
and the genus divided
into eight sections. Of these Tsutsutsi (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,
[13][14]
although other authors such as
Candolle (1838), who described six sections,
[19]
used slightly different
numeration.
As more species became available in the nineteenth century a better
understanding of the characteristics necessary for the major divisions.
Chief amongst these were Maximovicz's Rhododendreae Asiae Orientali
(1870)
[20]
and Planchon. Maximovicz used flower bud position and its
relationship with leaf buds to create eight Sections.
[21]
Bentham and
Hooker (1876) used a similar scheme, but called the divisions Series.
[22]
It was not until 1893 that 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 Balfour introduced the concept of
grouping species into series, in The Species of Rhododendron (1930), referred to as the Balfourian system.
[23]
That system continued up to modern times in Davidian's four volume The Rhododendron Species
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Cladogram of genus Rhododendron
(Goetsch et al. 2005)
A
Rhododendron
Choniastrum
B
Hymenanthes
C
Azaleastrum
(1982-1995).
[24]
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 his Ein System der Gattung Rhododendron L. (1949),
[25]
and
subsequent refinements.
[26][27]
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 (1980) in Edinburgh, placing more emphasis on the lepidote
characteristics of the leaves united all of the lepidote species into subgenus Rhodendron, including four of
Sleumer's (1980) subgenera (Rhododendron, Pseudoazalea, Pseudorhodorastrum, Rhodorastrum).
[28]
Philipson
& Philipson (1986) raised two sections of subgenus Aleastrum (Mumeazalea, Candidastrum) to subgenera,
while reducing genus Therorhodion to a subgenus of Rhododendron.
[29]
In 1987 Spethmann, adding
phytochemical features proposed a system with fifteen subgenera grouped into three 'chorus' subgenera.
[30]
A number of closely related genera had been included together with Rhododendron in a former tribe,
Rhodoreae. These have been progressively incorporated into Rhododendron.
[31]
Chamberlain and Rae (1990)
moved the monotypic section Tsusiopsis together with the monotypic genus Tsusiophyllum into section
Tsutsusi,
[32]
while in the same year Kron & Judd reduced genus Ledum to a subsection of section
Rhododendron.
[33]
Then Judd & Kron (1995) moved two species (Rhododendron schlippenbachii, R.
quinquefolium) from section Brachybachii subgenus Tsutsusi and two from section Rhodora subgenus
Pentanthera (R. albrechtii, R. pentaphyllum) into section Sciadorhodion subgenus Pentanthera.
[34]
Finally
Chamberlain brought the various systems together in 1996, with 1,025 species divided into eight subgenera. For
a comparison of the Sleuner and Chamberlain schemata see Table 1 of Goetsch (2005).
[9][2][14][35][36]
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
matK sequencing, while Lian-Ming used ITS sequences
[37]
to
determine a cladistic analysis. They confirmed that the genus
Rhodendron was monophyletic, with subgenus Therorhodion in the
basal position, consistent with the matK studies. Following publication
of the studies of Goetsch et al. with RPB2 (2005).
[2]
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..
[2][38]
The major finding of Goetsch and colleagues was that all species
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Therorhodion
examined (except 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 to all other rhododendrons.
The small polyphyletic subgenera Pentanthera and Azaleastrum were divided between two clades. The four
sections of Pentanethra between clades B and C, with two each, while Azaleastrum had one section in each of
A and C.
Thus subgenera Azaleastrum and Pentanethera needed to be dissassembled, 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 Rhodendron 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 Pentanethera, Tsutsusi, Candidastrum and
Mumeazalea, while a new subgenus was created by elevating subgenus Azaleastrum section Choniastrum to
subgenus rank.
Subgenus Pentanethera (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 Pentanethera (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 to section Pentanethera (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).
[2]
(Table 1.)
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Table 1.: Taxonomic changes within genus Rhododendron
(Goetsch 2005)
Chamberlain (1996) Goetsch
Genus Subgenus Section Species Subgenus Section
Menziesa
Azaleastrum
Sciadorhodion
Candidastrum
Tsutsusi
Tsutsusi Mumeazaleum
Pentanthera
Viscidula
Pentanthera
Hymenanthes Pentanethra
Rhodora
R.canadense
R.vaseyi Azaleastrum Sciadorhodion
Azaleastrum Choniastrum Choniastrum
Subsequent research has supported the revision by Goetsch, although has largely concentrated on further
defining the phylogeny within the subdivisions.(Craven 2008) In 2011 the two species of Diplarche were also
added to Rhododendron, incertae sedis.(Craven 2011) Similar findings were reported independently the
following year by Brown et al.
[9]
Subdivision
This genus has been progressively subdivided into a hierarchy of subgenus, section, subsection, and species.
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,
[39]
Rhododendron has eight subgenera based on morphology, namely the
presence of scales (lepidote), deciduousness of leaves, and the floral and vegetative branching patterns, after
Sleumer (1980).
[9][2][27]
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.
[40]
The largest of these is subgenus Rhododendron, containing nearly half of all known
species and all of the lepidote species.
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Subgenus Rhododendron L.: Small leaf or lepidotes (scales on the underside of the leaves). 3 sections,
462 species, type species: Rhododendron ferrugineum.
Subgenus Hymenanthes (Blume) K.Koch: Large leaf or elepidotes (without scales). 1 section, 224
species, type Rhododendron degronianum.
Subgenus Pentanthera (G. Don) Pojarkova: Deciduous azaleas. 4 sections, 23 species, type
Rhododendron luteum.
Subgenus Tsutsusi (Sweet) Pojarkova: Evergreen azaleas. 2 sections, 80 species, type Rhododendron
indicum.
Subgenus Azaleastrum Planch.: 2 sections, 16 species, type Rhododendron ovatum.
Subgenus Candidastrum Franch. : 1 species, Rhododendron albiflorum.
Subgenus Mumeazalea (Sleumer) W.R. Philipson & M.N. Philipson: 1 species, Rhododendron
semibarbatum.
Subgenus Therorhodion (Maxim.) A. Gray: 2 species (Rhododendron camtschaticum, Rhododendron
redowskianun).
For a comparison of the Sleumer and Chamberlain systems, see Goetsch et al. (2005) Table 1.
[2]
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.).
Table 2.: Morphological classification of Rhododendron
(Chamberlain 1996)
[35]
Inflorescence buds Leaf scales Leaf shoots Leaves Subgenus Section
Terminal
Present Rhododendron
Absent
Previous year
Evergreen Hymenanthes
Deciduous Pentanthera
Pentanthera
Rhodora
Viscidula
Lowest leaves
Pentanthera Sciadorhodion
Tsutsusi
Lateral
Evergreen Azaleastrum
Deciduous
Candidastrum
Mumeazalea
Therorhodion
Following the cladistic analysis of Goetsch et al. (2005)
[2]
this scheme was simplified, based on the discovery
of three major clades (A,B,C) as follows.
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Clade A
Subgenus Rhododendron L.: Small leaf or lepidotes (scales on the underside of the leaves). 3 sections,
about 400 species, type species: Rhododendron ferrugineum.
Subgenus Choniastrum Franch. : 11 species
Clade B
Subgenus Hymenanthes (Blume) K.Koch: Large leaf or elepidotes (without scales), including deciduous
azaleas. 2 sections, about 140-225 species, type Rhododendron degronianum.
Clade C
Subgenus Azaleastrum Planch.: Evergreen azaleas. 3 sections, about 120 species, type Rhododendron
ovatum.
Sister taxon
Subgenus Therorhodion (Maxim.) A. Gray: 2 species (Rhododendron camtschaticum, Rhododendron
redowskianun).
Sections and subsections
The larger subgenera are further subdivided into sections and subsections
[40]
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)
[2]
and by Craven et al. (2008)
[41]
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
[39]
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 L. (149 species in 25 subsections; temperate to subarctic Northern Hemisphere)
Vireya (Blume) Copel.f. (300 species in 2 subsections; tropical southeast Asia, Australasia. At one
time considered separate subgenus
[42]
)
Subgenus Hymenanthes (Blume) K.Koch (1 section, 224 species) (Increased to two sections)
Ponticum G. Don (24 subsections)
(Pentanthera (G. Don) Pojarkova (2 subsections) new section, moved from subgenus Pentanthera)
Subgenus Pentanthera (G. Don) Pojarkova (4 sections, 23 species) (Discontinued)
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Pentanthera (G. Don) Pojarkova (2 subsections) (Moved to subgenus Hymenanthes)
Rhodora (L.) G. Don (2 species; Rhododendron canadense, 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))
Species
Species of the genus Rhododendron are widely distributed between latitudes 80N and 20S and are considered
Alpine native plants from North America to Europe, Russia, and Asia, and from Greenland to Queensland,
Australia and the Solomon Islands.
[9]
The centres of diversification are in the Himalayas and Malaysia,
[37]
with
the greatest species diversity in the Sino-Himalayan region, Southwest China and northern Burma, from
Uttarakhand, Nepal and Sikkim to northwestern Yunnan and western Sichuan and southeastern Tibet, and with
other significant areas of diversity 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).
[2]
The 300 Tropical species within the Vereya 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 moist grasslands at around 3,000 metres above
sea level in the Central Highlands.
[42]
Subgenera Rhododendron and Hymenanthes, together with section
Pentanethera of subgenus Pentanethera are also represented to a lesser degree in the Mountainous areas of
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Rhododendron fallacinum
photographed in situ on
Mount Kinabalu, Borneo
Rhododendron-Park Bremen,
Germany
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.
[2][21]
Invasive species
Some species (e.g. Rhododendron ponticum in Ireland
[43]
and the United
Kingdom) are invasive as introduced plants, spreading in woodland areas
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 of some members of the order Lepidoptera
(butterflies and moths) (See List of Lepidoptera that feed on rhododendrons).
Diseases
Major diseases include Phytophthora root rot, stem and twig fungal dieback; Ohio State University Extension
provides information on maintaining health of rhododendrons.
[44]
Rhododendrons can easily be suffocated by
other plants.
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 subtemperate regions,(Craven
2008) while many species and cultivars are grown commercially for the
nursery trade. Rhododendrons are often valued in landscaping for their
structure, size, flowers, and the fact that many of them are evergreen.
[45]
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 R. arboreum and R. falconeri eventually growing to 1015 m or more
tall.
[45]
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,
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most of them are located 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.
[46]
In the Indian state of Himachal Pradesh, rhododendron flowers have been used for some time to
make popular fruit and flower wines. The industry is promoted by the state government with tax benefits,
looking to promote this industry as a full-fledged subclass of its economy.
[47]
Horticultural divisions
Horticulturally, rhododendrons may be divided into the following groups:-
[48]
Evergreen rhododendrons: the main default category
Vireya (Malesian) rhododendrons: epiphytic tender shrubs
[49]
Azaleas (section of generally small-sized, small-leaved and small-flowered shrubs):
Deciduous hybrid azaleas:
[50]
Ghent (Gandavense) hybrids - Belgian raised
[51]
Knap Hill-Exbury hybrids - English raised
[52]
Mollis hybrids - Dutch & Belgian raised
[53]
New Zealand Ilam hybrids - derived from Knap Hill/Exbury hybrids
Occidentale hybrids - English raised
Rustica hybrids - sweet-scented, double-flowered
Evergreen hybrid azaleas:
Gable hybrids - raised by Joseph B. Gable in Pennsylvania, USA
[54]
Glenn Dale hybrids - USA 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, USA
[55]
Vuyk (Vuykiana) hybrids - raised in the Netherlands
[56]
Azaleodendrons - semi-evergreen hybrids between deciduous azaleas and rhododendrons
Planting and care
Like other 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 epiphytes 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
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
11 of 24 8/4/2014 11:29 PM
Nova Zembla Rhododendrons
growing in a nursery in New Jersey.
bark.
[57]
Mulching and careful watering are important, especially before
the plant is established.
A new calcium-tolerant stock of rhododendrons (trademarked as
'Inkarho') has been exhibited at the RHS Chelsea Flower Show in
London (2011). Individual hybrids of rhododendrons have been 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.
[58][59]
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 the International Rhododendron Registry
(http://www.rhs.org.uk/Plants/RHS-Publications/Plant-registers/Rhododendron) 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
[60]
such as the Loderi hybrids,
created by crossing R. fortunei and R. griffithianum.
[61]
Other examples include the PJM hybrids, formed from
a cross between Rhododendron carolinianum and Rhododendron dauricum, and named after Peter J. Mezitt of
Weston Nurseries, Massachusetts.
[62]
Pharmacology
Rhododendron species have long been used in traditional medicine.
[63]
Animal studies and in vitro research has
identified possible anti-inflammatory and hepatoprotective activities which may be due to the antioxidant
effects of flavonoids or other phenolic compounds and saponins the plant contains.
[64][65][66]
Xiong et al. have
found that the root of the plant is able to reduce the activity of NF-B in rats.
[67]
Toxicology
Some species of rhododendron are poisonous to grazing animals because of a toxin called grayanotoxin in their
pollen and 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 Greek soldiers after having
consumed honey in a village surrounded by Rhododendron ponticum during the march of the Ten Thousand in
401 BC. Pompey's soldiers reportedly suffered lethal casualties following the consumption of honey made from
Rhododendron deliberately left behind by Pontic forces in 67 BC during the Third Mithridatic War. Later, it was
recognized that honey resulting from these plants has a slightly hallucinogenic and laxative effect.
[68]
The
suspect rhododendrons are Rhododendron ponticum and Rhododendron luteum (formerly Azalea pontica), both
found in northern Asia Minor.A brief documented video of this occurring in the modern day involves a group of
men in Nepal foraging for this affected honey can be found here: http://eupterrafoundation.com/hallucinogenic-
honey. Eleven similar cases have been documented in Istanbul, Turkey during the 1980s.
[69]
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 was mentioned in the 2009
film Sherlock Holmes as a proposed way to arrange a fake execution.
[70]
It was also mentioned in the third
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
12 of 24 8/4/2014 11:29 PM
episode of Season 2 of BBC's Sherlock (TV series), and has been speculated to have been a part of Sherlock's
fake death scheme.
Symbolism
Rhododendron arboreum (lali guransh) is the national flower of Nepal. R. ponticum is the state flower of
Indian-administered Kashmir and Pakistan-controlled Kashmir. Rhododendron niveum is the state tree of
Sikkim in India. Rhododendron is also the state tree of the state of Uttarakhand, India. Pink Rhododendron
(Rhododendron campanulatum) is the State Flower of Himachal Pradesh, India.
Rhododendron maximum, the most widespread rhododendron of the Appalachian Mountains, is the state flower
of West Virginia, and is in the Flag of West Virginia. Rhododendron macrophyllum, a widespread rhododendron
of the Pacific Northwest, is the state flower of Washington.
Literature
In Joyce's 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,
[71]
and Shades of Grey.
[72]
Amongst the Zomi tribes in India and Myanmar,
"Rhododendrons" called "Ngeisok" is used in a poetic manner to signify a lady.
Culinary
The rhododendron is the national flower of Nepal, 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 called burans (named after the flower) in the hilly regions of Uttarakhand. It is admired for its
distinctive flavor and color.
Labrador tea
Labrador tea is an herbal tea (not a true tea) made from three closely related species:
Rhododendron tomentosum (Northern Labrador tea, previously Ledum palustre),
Rhododendron groenlandicum, (Bog Labrador tea, previously Ledum groenlandicum or Ledum latifolium)
and
Rhododendron neoglandulosum, (Western Labrador tea, or trapper's tea, previously Ledum glandulosum).
List of Award of Garden Merit rhododendrons
List of Rhododendron diseases
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
13 of 24 8/4/2014 11:29 PM
List of Rhododendron species
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rhododendron on the activation
of NF- B in a chronic
glomerulonephritis rat model".
Journal of Nanjing Medical
University 23: 73.
doi:10.1016/S1007-4376(09)600
31-9 (http://dx.doi.org
/10.1016%2FS1007-4376%2809
%2960031-9).
67.
^ U S Food & Drug
Administration Center for Food
Safety & Applied Nutrition
Food borne Pathogenic
Microorganisms and Natural
Toxins 1992 (Bad Bug Book)
68.
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^ Nurhayat Stlpmar, Afife
Mat and Yurdagl Satganoglu
(February 1993). "Poisoning by
toxic honey in Turkey"
(http://www.springerlink.com
/content/w322484735r06208/).
Archives of Toxicology 67 (2):
148150.
doi:10.1007/BF01973687
(http://dx.doi.org
/10.1007%2FBF01973687).
PMID 8481104
(https://www.ncbi.nlm.nih.gov
/pubmed/8481104).
69. ^ "Rhododendron Poison
Truth behind the science of
Sherlock Holmes"
(http://www.aschoonerofscience.
com/?p=932). A Schooner of
Science. 2009-12-27. Retrieved
2013-02-26.
70.
^ Fforde, Jasper (26 July 2005).
Something rotten
(http://books.google.com
/?id=OaHIeqT1G9kC&
pg=PT306&lpg=PT306&
dq=jasper+fforde+%22rhododen
dron%22+thursday+next#v=one
page&q=rhododendron&
f=false).
ISBN 978-0-14-303541-1.
71.
^ Jasper Fforde. "Shades of
Grey - An Interview with Jasper
Fforde about Shades of Grey"
(http://www.jasperfforde.com
/grey/interview.html).
Jasperfforde.com. Retrieved
2013-02-26.
72.
Books and book chapters
A. P. de Candolle et al. (18241873). "Rhododendron" (http://biodiversitylibrary.org/item/7156#page
/724/mode/1upurl=). Prodromus systemati naturalis regni vegetabilis sive enumeratio contracta ordinum,
generum specierumque plantarum huc usque cognitarum, juxta methodi naturalis normas digesta. p. 719.
(also available online at Gallica (http://gallica.bnf.fr/))
Sweet, Robert. The British Flower Garden (Series the Second) Volume I. Ridgway, London 1831
(http://books.google.ca/books?id=xVwCAAAAYAAJ&printsec=frontcover#v=onepage&q&f=false)
Hooker, Joseph Dalton, Fitch, W. H.,Hooker, William Jackson. The rhododendrons of Sikkim-Himalaya :
being an account, botanical and geographical, of the rhododendrons recently discovered in the mountains
of eastern Himalaya, from drawings and descriptions made on the spot, during a government botanical
mission to that country. Reeve, Benham, and Reeve, London, 1849-51.
(http://www.biodiversitylibrary.org/bibliography/11178#/summary)
James Leonard Luteyn, Mary E. O'Brien. Contributions Toward a Classification of Rhododendron:
Proceedings, International Rhododendron Conference, the New York Botanical Garden, May 15-17,
1978. New York Botanical Garden, American Rhododendron Society 1980. ISBN 0893272213
(http://books.google.ca/books/about
/Contributions_Toward_a_Classification_of.html?id=zbogAQAAIAAJ&redir_esc=y)
Davidian, H. H. The Rhododendron Species. In four volumes from 1982-1995. Timber Press. ISBN
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
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0-917304-71-3, ISBN 0-88192-109-2, ISBN 0-88192-168-8, ISBN 0-88192-311-7.
(http://books.google.ca/books/about
/The_Rhododendron_Species_Azaleas.html?id=CBQmAQAAMAAJ&redir_esc=y)
Cox, P. A. & Kenneth, N. E. The Encyclopedia of Rhododendron Species. 1997. Glendoick Publishing.
ISBN 0-9530533-0-X. (http://books.google.ca/books/about
/The_encyclopedia_of_rhododendron_species.html?id=yftGAAAAYAAJ&redir_esc=y)
James Cullen. Hardy Rhododendron Species: A Guide To Identification. Timber Press, 2005. ISBN
0881927236 (http://books.google.ca/books/about
/Hardy_Rhododendron_Species.html?id=jxY4dVOm1VUC&redir_esc=y)
Blazich, F.A. and D.B. Rowe. 2008. "Rhododendron L., rhododendron and azalea." p. 943-951. In: F.T.
Bonner and R.P. Karrfalt (eds.). The woody plant seed manual. Agr. Hdbk 727. U.S. Dept. Agr. For. Serv.,
Washington, D.C. (http://www.rhododendron.dk/rhododendron.pdf)
Articles
Walter Magor. A History of Rhododendrons. (http://scholar.lib.vt.edu/ejournals/JARS/v44n4/v44n4-
magor.htm) JARS Volume 44, Number 4, Fall 1990
GAO Lian-Ming, LI De-Zhu, ZHANG Chang-Qin, YANG Jun-Bo. Infrageneric and Sectional
Relationships in the Genus Rhododendron (Ericaceae) Inferred from ITS Sequence Data. Am Botanica
Sinica 2002, 44(11) 1351-1356 (http://www.jipb.net/pubsoft/content/2/2316/X010407(PS2).pdf)
Goetsch, Loretta; Eckert, Andrew J.; Hall, Benjamin D. (JulySeptember 2005). "The molecular
systematics of Rhododendron (Ericaceae): a phylogeny based upon RPB2 gene sequences"
(http://www.ingentaconnect.com/content/aspt/sb/2005/00000030/00000003/art00012). Systematic Botany
30 (3): 616626. doi:10.1600/0363644054782170 (http://dx.doi.org/10.1600%2F0363644054782170).
Subdivisions
Azaleas
Wilson EH, Rehder A. A MONOGRAPH OF AZALEAS RHODODENDRON SUBGENUS
ANTHODENDRON. THE UNIVERSITY PRESS, CAMBRIDGE April 15 1921. PUBLICATIONS OF
THE ARNOLD ARBORETUM, No. 9 (http://www.biodiversitylibrary.org/bibliography/55595#
/summary)
John L. Creech. An Embryological Study in the Rhododendron Subgenus Anthodendron Endl. Botanical
Gazette Vol. 116, No. 3 (Mar., 1955), pp. 234-243 (http://www.jstor.org/discover/10.2307
/2473343?uid=3739432&uid=2129&uid=2&uid=70&uid=3737720&uid=4&sid=21103875679061)
Tsutsusi
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
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Chamberlain DF. A revision of Rhododendron. IV Subgenus Tsutsusi. Edinburgh Journal of Botany
06/1990; 47(02):89 - 200. DOI: 10.1017/S096042860000319X (https://www.researchgate.net/publication
/231958128_A_revision_of_Rhododendron._IV_Subgenus_Tsutsusi)
Powell, E. Ann, Kron, Kathleen A. Molecular systematics of Rhododendron subgenus Tsutsusi
(Rhodoreae, Ericoideae, Ericaceae). Botany 2004: 147 (http://2004.botanyconference.org/engine/search
/index.php?func=detail&aid=147)
A. Kron and E. A. Powell. MOLECULAR SYSTEMATICS OF RHODODENDRON SUBGENUS
TSUTSUSI (RHODOREAE, ERICOIDEAE, ERICACEAE) Edinburgh Journal of Botany Volume 66
Issue 1 March 2009, pp 81-95 (http://journals.cambridge.org/action/displayAbstract?fromPage=online&
aid=4481360K.)
Yue-Jiao ZHANG, Xiao-Feng JIN, Bing-Yang DING and Jing-Ping ZHU. Pollen morphology of
Rhododendron subgen. Tsutsusi and its systematic implications. Journal of Systematics and Evolution.
Volume 47, Issue 2, pages 123138, March 2009 (http://onlinelibrary.wiley.com/doi/10.1111
/j.1759-6831.2009.00011.x/abstract)
Jin Xiao-Feng, Ding Bing-Yang, Zhang Yue-Jiao, and Hong De-Yuan. A Taxonomic Revision Of
Rhododendron subg. Tsutsusi sect. Brachycalyx (Ericaceae). Annals of the Missouri Botanical Garden
97(2):163-190. 2010 doi: http://dx.doi.org/10.3417/2007139 (http://www.bioone.org/doi/abs/10.3417
/2007139?journalCode=mobt)
Vireya
Sleumer, Hermann Otto. An account of rhododendron in Malesia. P. Noordhoff, Groningen 1966
David G. Leach. The Discovery of the Malaysian Rhododendrons. JARS Volume 32, Number 1 Winter
1978 (http://scholar.lib.vt.edu/ejournals/JARS/v32n1/v32n1-leach.htm)
Argent, G. Rhododendrons of subgenus Vireya. 2006. Royal Horticultural Society. ISBN 1-902896-61-0
Gillian K. Brown, Lyn A. Craven, Frank Udovicic and Pauline Y. Ladiges. Phylogenetic relationships of
Rhododendron section Vireya (Ericaceae) inferred from the ITS nrDNA region. Australian Systematic
Botany 19, 329342. August 2006. 10.1071/SB05019 (http://www.researchgate.net/publication
/235601539_Phylogenetic_relationships_of_Rhododendron_section_Vireya_(Ericaceae)_inferred_from_t
he_ITS_nrDNA_region/file/72e7e51e347034a1c1.pdf)
Hall, B.D., L.A. Craven & L.A. Goetsch. 2006. The taxonomy of subsection Pseudovireya two
distinctly different taxa within subsection Pseudovireya and their relation to the rooting of section Vireya
within subgenus Rhododendron. Rhododendron Sp. 1: 72, 9197
Craven, L.A.; Goetsch, L.A.; Hall, B.D.; Brown, G.K. (2008). "Classification of the Vireya group of
Rhododendron (Ericaceae)". Blumea - Biodiversity, Evolution and Biogeography of Plants 53 (2): 435.
doi:10.3767/000651908X608070 (http://dx.doi.org/10.3767%2F000651908X608070).
Goetsch, L.A., Craven, L.A. and Hall B.D. 2011. Major speciation accompanied the dispersal of Vireya
Rhododendrons (Ericaceae, Rhododendron sect. Schistanthe) through the Malayan archipelago: Evidence
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
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from nuclear gene sequences. Taxon 60 (4): 10151028. (http://www.ingentaconnect.com/content/iapt/tax
/2011/00000060/00000004/art00006)
Peter Adams. Evolution, Adaptive Radiation and Vireya Rhododendrons - Part I. JARS FALL 2012 pp.
201-3 (http://www.whitecloudnursery.com/Vireya%20Phylogeny%20pt%20I.pdf)
Peter Adams. Evolution, Adaptive Radiation and Vireya Rhododendrons - Part I. JARS SPRING 2013 pp.
74-76 (http://www.whitecloudnursery.com/Vireya%20Phylogeny%20pt%20II.pdf)
Biodiversity of the Vireya group of Rhododendron L. (Ericaceae) collections in New Zealand and their
potential contribution to international conservation. Fayaz A. PhD Thesis, Department of Plant Science,
Massey University, New Zealand. 2012 (http://mro.massey.ac.nz/handle/10179/4468?show=full.)
Separate genera
Craven, L.A. Diplarche and Menziesia transferred to Rhododendron (Ericaceae). Blumea Volume 56,
Number 1, April 2011, pp. 33-35(3) (http://www.ingentaconnect.com/content/nhn/blumea/2011/00000056
/00000001/art00010)
History of Rhododendron Discovery & Culture (http://www.rhodyman.net/rhodyhis.html)
Rhododendrons from Turkey, Anatolia (http://www.lahana.org/blog/Rhododendrons.htm)
German Genebank Rhododendron (http://www.lwk-niedersachsen.de/index.cfm/action
/rhodo.html?lang=gb)
Description of damage caused by Rhododendrons in the UK (http://www.countrysideinfo.co.uk
/rhododen.htm)
Information on rhododendrons at the Ericaceae web pages of Dr. Kron at Wake Forest University.
(http://www.wfu.edu/~kronka/kakbek/rhododendrons.html)
Information on Vireyas (http://www.vireya.net/)
Information+photos of hybrids and species (http://www.hirsutum.info/)
Information on Rhododendrons by Marc Colombel, founder of the Socit Bretonne du Rhododendron.
(http://www.rhodophiles.com/)
Extensive information on rhododendron species: the history of their discovery, botanical details, toxicity,
classification, cultural conditions, care for common problems, and suggestions for companion plants by
Steve Henning. (http://rhodyman.net/rahome.html)
History of Rhododendrons (http://www.rhodo.citymax.com/page/page/627471.htm)
Historical Survey of Rhododendron Collecting - With Emphasis on its Close Associations with
Horticulture (http://scholar.lib.vt.edu/ejournals/JARS/v23n4/v23n4-black.htm)
Rhododendron in botanical garden Pruhonice-Czech republic (http://www.rododendron.cz/fotogalerie-
rododendronu/pruhonicky-park-2011/)
Rhododendron - Wikipedia, the free encyclopedia http://en.wikipedia.org/wiki/Rhododendron
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Wikimedia Commons has
media related to
Rhododendron.
Wikispecies has
information related to:
Rhododendron
Databases
(http://plants.usda.gov/core/profile?symbol=RHODO)USDA: Rhododendron
(http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=23700)ITIS:
Rhododendron
Flora
Flora of China: Rhododendron (http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=128386)
Flora of North America: Rhododendron (http://www.efloras.org/florataxon.aspx?flora_id=1&
taxon_id=128386)
Rhododendron societies
American Rhododendron Society (http://rhododendron.org/)
Journal of the American Rhododendron Society (JARS) 1947-1996 (http://scholar.lib.vt.edu
/ejournals/JARS/#qbars)
"Genus Rhododendron Taxonomic Tree" (http://www.rhododendron.org/taxonomictree.asp).
American Rhododendron Society. Retrieved 18 June 2014.
The Rhododendron, Camellia & Magnolia Group of the Royal Horticultural Society
(http://www.rhodogroup-rhs.org/)
Rhododendron Species Foundation and Botanical Garden (http://www.rhodygarden.org/)
Socit Finlandaise du Rhododendron (http://honeybee.helsinki.fi/users/avainola/rhodo/club.htm)
Australian Rhododendron Society (http://www.ausrhodo.asn.au/)
German Rhododendron Society (http://www.rhodo.org/start.php?lang=gb)
New Zealand Rhododendron Association (http://www.rhododendron.org.nz/)
Danish Rhododendron Society (http://www.rhododendron.dk/alfabet.html)
Fraser South Rhododendron Society (http://frasersouthrhodos.ca/)
Botanical gardens
Royal Botanic Garden Edinburgh: Rhododendron Publications (http://www.rbge.org.uk/about-
us/publications/publications-catalogue/botanical-publications/rhododendron-publications)
Retrieved from "http://en.wikipedia.org
/w/index.php?title=Rhododendron&oldid=618571082"
Categories: Rhododendron Holarctic flora Flora of Indomalesia
Poisonous plants Plants used in traditional Chinese medicine
Flora of Korea Flora of Nepal Flora of Thailand
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