Cullenia Excelsa
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''Cullenia exarillata'' is a
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
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 ...
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 ...
species in the family
Malvaceae Malvaceae, or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include okra, cotton, cacao and durian. There are also some genera containing familiar ...
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
to the rainforests of the southern Western Ghats in
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. It is one of the characteristic trees of the mid-elevation tropical wet evergreen rainforests and an important food plant for the endemic primate, the
lion-tailed macaque The lion-tailed macaque (''Macaca silenus''), also known as the wanderoo, is an Old World monkey endemic to the Western Ghats of South India. Characteristics Lion-tailed macaques are covered in black fur, and have a striking gray or silver ...
.Kumar, A. (1987) The ecology and population dynamics of the lion-tailed macaque (''Macaca silenus'') in South India. PhD thesis, Cambridge University, UK.


Description

Tall evergreen trees with smooth greyish white bark, flaking in mature trees, with straight boles, frequently buttressed. The branches are horizontal often with series of knob-like
tubercles In anatomy, a tubercle (literally 'small tuber', Latin for 'lump') is any round nodule, small eminence, or warty outgrowth found on external or internal organs of a plant or an animal. In plants A tubercle is generally a wart-like projection ...
(for
cauliflorous Cauliflory is a botanical term referring to plants that flower and fruit from their main stems or woody trunks, rather than from new growth and shoots. This can allow trees to be pollinated or have their seeds dispersed by animals that climb o ...
attachment of flowers and fruits). The young branchlets and the underside of leaves are covered by golden brown peltate (or shield like) scales. Leaves are simple, alternate, glabrous, shiny green above and covered beneath with silvery or orangish peltate scales. The tubular,
hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have separ ...
flowers (also covered by golden brown scales) are about 4–5 cm long and cream or pinkish brown in colour. The flowers lack petals and are formed of tubular bracteoles and tube-like calyx, obscurely 5-lobed. The round fruits, about 10–13 cm in diameter and covered with spines, are clustered along the branches. The fruit is a capsule, 5-valved, containing many reddish brown seeds about 4–5 cm long and 2–3 cm wide. In the fruit, the seeds are covered by a fleshy, whitish
aril An aril (pronounced ), also called an arillus, is a specialized outgrowth from a seed that partly or completely covers the seed. An arillode or false aril is sometimes distinguished: whereas an aril grows from the attachment point of the see ...
. The fruit
dehisces 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 op ...
open when mature and dry to release seeds.


Taxonomy

The genus ''Cullenia'' was created by
Robert Wight Robert Wight Doctor of Medicine, MD Fellow of the Royal Society, FRS Linnean Society of London, FLS (6 July 1796 – 26 May 1872) was a Scottish surgeon in the East India Company, whose professional career was spent entirely in southern India, ...
and commemorates
William Cullen William Cullen FRS FRSE FRCPE FPSG (; 15 April 17105 February 1790) was a Scottish physician, chemist and agriculturalist, and professor at the Edinburgh Medical School. Cullen was a central figure in the Scottish Enlightenment: He was Dav ...
with the type species ''excelsa'' from India which Wight considered incorrectly as being identical to the Sri Lankan ''C. ceylanica'' which was earlier described under the genus ''Durio''. André Robyns examined Wight's specimen and fresh specimens from southern India and noted it as being distinct and described it as ''C. exarillata'' in 1970. ''C. ceylanica'' has the seeds covered by an aril whereas ''C. exarillata'' does not have the aril surrounding the seeds. The genus is evolutionarily close to ''Boschia'' and ''Durio''.


Distribution and habitat

The species is characteristically found and dominant in the mid-elevation (700 m to 1400 m) tropical wet evergreen rainforests, which has been called the ''Cullenia exarillata'' - ''Mesua ferrea'' - ''Palaquium ellipticum'' type. It occurs from the southern tip of the Western Ghats in Kalakad-Mundanthurai Tiger Reserve and
Agasthyamalai hills The Pothigai Hills, also known as Agasthiyar Mountain is a 1,866-metre (6,122 ft)-tall peak in the southern part of the Western Ghats of South India. The peak lies in Tirunelveli District of Tamil Nadu near the border of Kerala. The ar ...
to
Wayanad Wayanad () is a district in the north-east of Indian state Kerala with administrative headquarters at the municipality of Kalpetta. It is the only plateau in Kerala. The Wayanad Plateau forms a continuation of the Mysore Plateau, the southern ...
and
Kodagu Kodagu (also known by its former name Coorg) is an administrative district in the Karnataka state of India. Before 1956, it was an administratively separate Coorg State, at which point it was merged into an enlarged Mysore State. It occupies ...
in the
Nilgiri Biosphere Reserve The Nilgiri Biosphere Reserve is a biosphere reserve in the Nilgiri mountains of the Western Ghats in South India. It is the largest protected forest area in India, spreading across Tamil Nadu, Karnataka and Kerala. It includes the protected ar ...
.


Breeding system and dispersal

The hermaphroditic (or bisexual) flowers are mainly pollinated by bats (''
Cynopterus ''Cynopterus'' (Latin meaning: ״flying dog״) is a genus of megabats. The cynopterine section is represented by 11 genera,Andersen K. 1912. Catalogue of the chiroptera in the collection of the British Museum. Second edition, British Museum of N ...
'' and ''
Rousettus ''Rousettus'' is a genus of Old World fruit bats or megabats, referred to as rousette bats. The genus is a member of the family Pteropodidae. The genus consists of seven species that range over most of Africa to southeast Asia, and the islands o ...
'') and arboreal mammals (
Lion-tailed Macaque The lion-tailed macaque (''Macaca silenus''), also known as the wanderoo, is an Old World monkey endemic to the Western Ghats of South India. Characteristics Lion-tailed macaques are covered in black fur, and have a striking gray or silver ...
,
Brown Palm Civet The brown palm civet (''Paradoxurus jerdoni'') also called the Jerdon's palm civet is a palm civet endemic to the Western Ghats of India. Taxonomy The scientific name ''Paradoxurus jerdoni'' was introduced by William Thomas Blanford in 188 ...
,
Indian Giant Squirrel The Indian giant squirrel or Malabar giant squirrel (''Ratufa indica'') is a large multi-coloured tree squirrel species endemic to forests and woodlands in India. It is a diurnal, arboreal, and mainly herbivorous squirrel. Distribution and hab ...
,
Nilgiri Langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
, and
Indian Giant Flying Squirrel The Indian giant flying squirrel (''Petaurista philippensis''), also called the large brown flying squirrel or the common giant flying squirrel, is a species of rodent in the family Sciuridae. It is capable of gliding flight using a skin membrane ...
). These mammals and other rodents that visit the flowers such as Dusky-striped Squirrel and
Malabar Spiny Dormouse The Malabar spiny dormouse (''Platacanthomys lasiurus'') is a species of muroid rodent endemic to the Western Ghats of India. It is the only extant species in the genus ''Platacanthomys'' and although resembling a dormouse, it is not closely r ...
also act as flower predators as they consume a substantial number of flowers. The flowers are also visited by a number of bird species, including
Indian White-Eye The Indian white-eye (''Zosterops palpebrosus''), formerly the Oriental white-eye, is a small passerine bird in the white-eye family. It is a resident breeder in open woodland on the Indian subcontinent. They forage in small groups, feeding on n ...
, Square-tailed Bulbul,
Yellow-browed Bulbul The yellow-browed bulbul (''Acritillas indica''), or golden-browed bulbul, is a species of songbird in the bulbul family, Pycnonotidae. It is found in the forests of southern India and Sri Lanka. The yellow-browed bulbul is mainly yellow on the ...
,
Brown-cheeked Fulvetta The brown-cheeked fulvetta (''Alcippe poioicephala'') or brown-cheeked alcippe as the fulvettas proper are not closely related to this species,) is included in the family Alcippeidae. It was earlier also known as the quaker babbler. This specie ...
,
Common Rosefinch The common rosefinch (''Carpodacus erythrinus'') or scarlet rosefinch is the most widespread and common rosefinch of Asia and Europe. Taxonomy In a molecular phylogenetic study of the finch family published in 2012, Zuccon and colleagues found t ...
, Kerala Laughingthrush, and
White-cheeked Barbet The white-cheeked barbet or small green barbet (''Psilopogon viridis'') is a species of Asian barbet found in southern India. It is very similar to the more widespread brown-headed barbet (or large green barbet, ''Psilopogon zeylanicus''), but t ...
. However, flowers visited by birds tend to be aborted. ''Cullenia exarillata'' is an
outcrossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...
species, producing negligible fruit-set under
geitonogamy Geitonogamy (from Greek ''geiton'' (γείτων) = neighbor + ''gamein'' (γαμεῖν) = to marry) is a type of self-pollination. Geitonogamous pollination is sometimes distinguished from the fertilizations that can result from it, geitonogamy. ...
and no fruit-set under
autogamy Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species o ...
. The seeds are mechanically dispersed (gravity) as well as by Lion-tailed Macaques over short distances.


Ecology

''Cullenia exarillata'' is a dominant tree species in the mid-elevation tropical rainforests (between 700 and 1,400 m elevation) of the southern Western Ghats. It is also among the most abundant canopy trees in relatively undisturbed mature wet evergreen forests. It occurs at a density of 20 trees/ha to 59.6 trees/ha in mid-elevation wet evergreen forests of the
Anamalai Hills The Anamala or Anaimalai, also known as the Elephant Mountains, are a range of mountains in the southern Western Ghats of central Kerala (Idukki district, Ernakulam district, Palakkad district, Thrissur district) and span the border of western ...
. Individual trees were estimated to produce between 1300 and 26000 flowers (average = 8734) chiefly between February and May, followed by a fruiting peak between May and September, in a rainforest at the southern tip of the Western Ghats. In the Anamalai Hills further north in the southern Western Ghats, the species was observed flowering between October and February. As the tree flowers abundantly during the dry season, a period of fruit scarcity in the forest, the flowers attract many diurnal and
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
frugivorous A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and ...
mammals and birds, making the tree a possible
keystone species A keystone species is a species which has a disproportionately large effect on its natural environment relative to its abundance, a concept introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in maintaini ...
in these forests. The flowers are low in nectar but contain fleshy
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
s embedded with
nectaries Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
which are the main reward for animal visitors. The flowers are eaten by endemic arboreal mammals such as
Lion-tailed Macaque The lion-tailed macaque (''Macaca silenus''), also known as the wanderoo, is an Old World monkey endemic to the Western Ghats of South India. Characteristics Lion-tailed macaques are covered in black fur, and have a striking gray or silver ...
,
Brown Palm Civet The brown palm civet (''Paradoxurus jerdoni'') also called the Jerdon's palm civet is a palm civet endemic to the Western Ghats of India. Taxonomy The scientific name ''Paradoxurus jerdoni'' was introduced by William Thomas Blanford in 188 ...
,
Indian giant squirrel The Indian giant squirrel or Malabar giant squirrel (''Ratufa indica'') is a large multi-coloured tree squirrel species endemic to forests and woodlands in India. It is a diurnal, arboreal, and mainly herbivorous squirrel. Distribution and hab ...
,
Bonnet macaque The bonnet macaque (''Macaca radiata''), also known as zati,Chambers English Dictionary is a species of macaque endemic to southern India. Its distribution is limited by the Indian Ocean on three sides and the Godavari and Tapti Rivers, along wi ...
, and
Nilgiri langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
. The base of the sepals are consumed and the anthers and stigma are discarded. The seeds and flowers of this species form a major part of the diet of Lion-tailed macaque in mid-elevation rainforests: 24.8% of the annual diet in Puthuthottam forest fragment in the
Anamalai Hills The Anamala or Anaimalai, also known as the Elephant Mountains, are a range of mountains in the southern Western Ghats of central Kerala (Idukki district, Ernakulam district, Palakkad district, Thrissur district) and span the border of western ...
, 20.7% of annual diet in
Silent Valley National Park Silent Valley National Park is a national park in Kerala, India. It is located in the Nilgiri hills, has a core area of , which is surrounded by a buffer zone of . This national park has some rare species of flora and fauna. This area was explo ...
. The seeds constituted about 20% of the annual fruit diet of the Lion-tailed macaque and 7.1% of
Indian giant squirrel The Indian giant squirrel or Malabar giant squirrel (''Ratufa indica'') is a large multi-coloured tree squirrel species endemic to forests and woodlands in India. It is a diurnal, arboreal, and mainly herbivorous squirrel. Distribution and hab ...
diet in mid-elevation rainforest of Pachapal Malai or Waterfall Shola in the Anamalai Hills. Seeds were also found to be a minor food resource (<1% annual diet) for
Nilgiri langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
in this study. In lower-elevation rainforests at Varagaliar in the Anamalai Hills, ''Cullenia exarillata'' comprised a smaller percentage (0.4 – 1.2%) of the annual diet of lion-tailed macaques, with the flowers, seeds, and aril being consumed. A seed predation rate of 45% has been estimated in a wet evergreen forest of the southern Western Ghats, caused by species such as Indian Giant squirrels that feed on unripe fruits (with softer spines) and Lion-tailed macaques that feed on ripe fruits (with hard spines). Seeds fallen on the forest floor also suffer high predation by mammals and insects, with 91.6% of the seeds predated in experimental plots open to all predators and 44% predated in plots where mammals were excluded.Krishnan, A. 2020. How does fragmentation alter seed predation patterns of rainforest tree species? MSc Dissertation, National Centre for Biological Sciences, Tata Institute of Fundamental Research, Bangalore, India. 59 pp. The study also reported that seeds of ''Cullenia exarillata'' were predated by three species of rodents (
Malabar spiny dormouse The Malabar spiny dormouse (''Platacanthomys lasiurus'') is a species of muroid rodent endemic to the Western Ghats of India. It is the only extant species in the genus ''Platacanthomys'' and although resembling a dormouse, it is not closely r ...
, ''
Rattus ''Rattus'' is a genus of muroid rodents, all typically called rats. However, the term rat can also be applied to rodent species outside of this genus. Species and description The best-known ''Rattus'' species are the black rat (''R. rattus'') ...
'' sp. and
Indian crested porcupine The Indian crested porcupine (''Hystrix indica'') is a hystricomorph rodent species native to southern Asia and the Middle East. It is listed as Least Concern on the IUCN Red List. It belongs to the Old World porcupine family, Hystricidae. Des ...
), besides
Indian spotted chevrotain The Indian spotted chevrotain (''Moschiola indica'') is a species of even-toed ungulate in the family Tragulidae. It is native to India and possibly Nepal. It lives in rainforests and is nocturnal. It has a body length of with a long tail lengt ...
, sambar,
Indian muntjac The Indian muntjac or the common muntjac (''Muntiacus muntjak''), also called the southern red muntjac and barking deer, is a deer species native to South and Southeast Asia. It is listed as Least Concern on the IUCN Red List. In popular local l ...
, and a primate,
Nilgiri langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
. Seed predation is higher in rainforest fragments than in contiguous rainforests, which has been experimentally revealed to be largely due to predation by mammals.


Conservation

The species is endemic to the Western Ghats and has not yet been assessed for the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
. The tree may persist in rainforest fragments showing similar levels of flowering and fruit-set, and may even have higher fruit-set in some disturbed sites and on isolated trees.Devy, M. Soubadra (2006)
Effects of fragmentation on a keystone tree species in the rainforest of Kalakad Mundanthurai Tiger Reserve, India
Unpublished Report submitted to Rufford Foundation, Ashoka Trust for Research in Ecology and the Environment, Bangalore, India.
Highly disturbed sites, including plantations where understorey vegetation and canopy trees were removed, have lower density of ''Cullenia exarillata''.


Gallery

File:Cullenia exarillata DSCN3067.jpg, A ''Cullenia exarillata'' tree towering above others in the Anamalai Hills File:Cullenia exarillata fruiting branch P1050369.jpg, Fruiting branch in canopy File:Cullenia exarillata mature fruits.jpg, Mature fruits File:Cullenia exarillata mature fruits dehiscing.jpg, Mature fruits splitting File:Cullenia exarillata mature fruits dehisced 2.jpg, Mature fruits dehiscing open File:Cullenia exarillata DSCN3917.jpg, Dehisced fruits of ''Cullenia exarillata'' still on the tree File:Cullenia exarillata-5-bsi-yercaud-salem-India.jpg, Seeds of ''Cullenia exarillata'' File:Cullenia exarillata leaf DSC 8270.jpg, Young leaf before opening out File:Durio exarillatus 10.JPG, Upperside of leaves File:Durio exarillatus 11.JPG, Underside of leaves File:Cullenia exarillata single flower at Periya (8).jpg, Single flower File:Nilgiri Langur eating Cullenia exarillata.jpg,
Nilgiri Langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
feeding on a ''Cullenia exarillata'' seed File:Mouse Deer eating Cullenia exarillata.jpg,
Indian spotted chevrotain The Indian spotted chevrotain (''Moschiola indica'') is a species of even-toed ungulate in the family Tragulidae. It is native to India and possibly Nepal. It lives in rainforests and is nocturnal. It has a body length of with a long tail lengt ...
feeding on a ''Cullenia exarillata'' seed File:Nilgiri Langur picking Cullenia exarillata.jpg,
Nilgiri Langur The Nilgiri langur (''Semnopithecus johnii'') is a langur (a type of Old World monkey) . This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a ...
picking a ''Cullenia exarillata'' seed File:Barking Deer Feeding on Cullenia.jpg,
Indian muntjac The Indian muntjac or the common muntjac (''Muntiacus muntjak''), also called the southern red muntjac and barking deer, is a deer species native to South and Southeast Asia. It is listed as Least Concern on the IUCN Red List. In popular local l ...
approaching to feed on a ''Cullenia exarillata'' seed File:Cullenia exarillata fruit stages collage.jpg, ''Cullenia exarillata'' fruit in different stages


References

{{Taxonbar, from=Q17579748 exarillata