Memecylon Cardiophyllum

''Memecylon'' is a plant group in Melastomataceae. It consists of 350-400 species of small to medium-sized trees and shrubs occurring in the Old World tropics. ''Memecylon'' is a monophyletic group basal to the Melastomataceae clade. ''Memecylon'' taxa have more than 600 published basionyms. Diversity of this group is concentrated in tropical Africa, Madagascar, Sri Lanka, India and Malaysia.

Etymology

The name ''Memecylon'' is derived from 'memaecylon' as used by ancient Greek philosophers Dioscorides and Pliny to describe the red fruits of ''Arbutus unedo'' (oriental strawberry tree), an unrelated plant group, alluding to the pink to reddish berries often produced by ''Memecylon''. Some vernacular names in different regions of the world are given below.
English: "Blue mist plant", Hindi: ''Anjan''; Malayalam: ''Aattukanala'' ��റ്റുകനല ''Kaasaavu'': ��ാശാവ് ''Kaayaampoo'': ��ായാമ്പൂ Odia: ''Neymaru''; Sinhala: ''Korakaha'', Welikaha; Tamil: ''Kaya''.

Morphology

''Memecylon'' '' sensu lato'' can be diagnosed by exstipulate leaves, four-merous bisexual flowers, anthers opening by slits, enlarged connectives bearing terpenoid secreting glands and berries. ''Memecylon sensu stricto'' can be distinguished from other Memecyloids by obscure nervation on leaves, non-glandular roughened leaf surface having branched sclerids, imbricate calyx, unilocular ovary and large embryo with thick and convoluted cotyledons.

Stems

Species of larger stature have a characteristic brown bark with narrow and sharp furrows, most are small single stemmed trees. However, the bark of many species of smaller stature is varied and may be papery white or smooth dark red-black.

Leaves

Leaves are opposite, short-stalked, elliptic or ovate, mostly with widely spaced pinnate nerves either visible or obscure. Leaves along the twig are all the same size, shiny, glabrous, with entire margins, the node has a characteristic scar between the leaves, the twig bark is typically red, striated and flaky.

Flowers

The inflorescence is typically dense and axillary. The florets are small (usually less than 5 mm) compared to the other taxa in Melastomataceae, with short fleshy corolla parts. Cymes are bracteate, usually thyrsoid to umbel shaped, often condensed to sessile fascicles of flowers or a few-flowered heads at tips of peduncles. The florets are white or violet, the stamens blue or violet, usually obvious in aggregates, from axillary clusters. Flowers are bisexual, have inferior ovaries, but the parts are otherwise free. The calyx is valvate and there are twice as many stamens as petals.

Fruits

The fruit is from an inferior ovary, typically axillary. The calyx remnants are persistent, and are sometimes blue-black. Fruits are globose or occasionally ellipsoid, pulpy or juicy with one large seed.

Distribution

This group is distributed in approximately 53 countries of the Old World tropics and occupies a wide range of habitats. ''Memecylon'' taxa have been reported from montane forests, tropical lowland forests, grasslands, tropical rainforests with low to high rainfall, rocky mountain regions and regions with low to high temperature and a considerable overlap between ranges of different taxa. Most of the plants in this group are regionally or locally endemic. Endemic ''Memecylon'' are reported from 21 countries. The global distribution of this group is shown in the following Figure and some of the countries which have endemic ''Memecylon'' species are listed in the table.

Table1. Number of ''Memecylon'' taxa reported from different countries

Phylogeny

Phylogeny based on morphological treatments

The classification of ''Memecylon'' has switched back and forth among families Melastomataceae, Myrtaceae and Memecylaceae based on morphological, anatomical and molecular evidence. Early classifications excluded ''Memecylon'' and Mouriri from the Melastomataceae mainly based on placentation and seed characters. Subsequent treatments preferred to treat ''Memecylon'' as a member of Myrtaceae. Again, this group has been included in Melastomataceae by Cronquist . After that, morphological and anatomical character analysis of the Melastomataceae and their traditional allies by Renner identified two major lineages (Melastomataceae and Memecylaceae), and in that classification, ''Memecylon'' was placed in the Memecylaceae. Synapomorphies used in this phylogenetic analysis were anther connective and dehiscence, dorsal glands on stamen connectives, endothecium, placentation, locules, seeds, leaf venation, terminal leaf sclereids, paracytic stomata, stomata shape, leaf sclereids, indumentum, ant and mite domatia, wood and growth form characters, which excluded Memecylaceae from Melastomataceae. However, in ''Memecylon'' some characters such as seasonal flowering and small size of flowers contributed to the difficulty of assessing relationships based on the morphology. Later, several groups have been either included in broadly defined Memecylaceae or segregated from it. As a result, Memecylaceae comprised the groups ''Memecylon'', ''Lijndenia'', ''Mouriri'', and ''Votomita'.''

Phylogeny based on molecular treatments

Molecular phylogenetic analyses using ''rbcL'' sequence data showed that the Melastomataceae lineage consists of a subclade formed by Oliniaceae, Penaeaeae, Phynchocalycaceae and Alzateaceae sister to a subclade formed by Memecylaceae and Melastomataceae ''sensu stricto.'' Parsimony analysis showed distinct Myrtaceae and Melastomataceae clades. Subsequently, Bayesian analyses using chloroplast DNA sequences from the '' rbcL'' and ''ndhF'' genes, and parsimony and maximum likelihood analyses on ''rRNA, rbcL'' and '' atpB'' sequences, have shown that Memecylaceae is a sister group to Melastomataceae. Later studies using combined exon and intron sequences of the nuclear glyceraldehyde 3-phosphate dehydrogenase gene (''GapC'') has supported the monophyly of ''Memecylon''. However, the APG IV (Angiosperm Phylogeny Group IV) system of classification, a system of revised and updated classification of flowering plants, now recognizes the groups of Memecylaceae within a broader circumscription of Melastomataceae.

The most recent phylogenetic analysis of ''Memecylon'' was based on low-copy nuclear loci using representative taxa across the old world tropics, and this study revealed biogeography and ancestral states of this plant group. In addition, there is a fine-scale study associated with South African ''Memecylon'' showing the reticulate evolution of this group.

Pollination and seed dispersal

The dense and axillary showy clusters of ''Memecylon'' florets do not produce nectar. These flowers are visited by pollen-gathering bees who vibrate or manipulate the anthers. Anthers open by longitudinal slits and exposed pollen invites pollen gathering bees. Anther appendages serve as a hold for bees’ legs. These flowers have terpenoid secreting glands and bees collect terpenoids. Therefore, buzz pollination is also favored. Berries are dispersed by birds and mammals. Populations of ''Memecylon'' are widely scattered within the forests as would be expected in bird-dispersed species.

Ecology

''Memecylon'' produce flowers and fruits more regularly than many trees of the equatorial forests. It provides a food supply for wildlife as a source of fruits. Different sympatric groups appear to segregate mainly by soil moisture.

Ethnobotany

''Memecylon'' has economic, medicinal and horticultural values. A yellow dye and a mordant can be extracted from the leaves. They are traditionally used for dyeing silk in Thailand and the robes of Buddhist monks in Sri Lanka. Fruits are edible and some are used as spices. This group provides hard and valuable timber used for building houses and boats. Wood is used to make rafters, house posts, fuel wood, charcoal, tools, and handles. An infusion of leaves is used as astringent for ophthalmia. The bark is applied as a poultice to bruises. Root and heartwood decoctions are used to bring down fever associated with colds, chicken pox and measles.

A list of ''Memecylon'' species

*'' Memecylon aberrans''
*'' Memecylon accedens''
*'' Memecylon acrocarpum''
*'' Memecylon acrogenum''
*'' Memecylon acuminatissimum''
*''Memecylon acuminatum''
*'' Memecylon aenigmaticum''
*''Memecylon aequidianum''
*''Memecylon affine''
*''Memecylon afzelii''
*'' Memecylon agastyamalaianum''
*'' Memecylon alatum''
*'' Memecylon albescens''
*'' Memecylon alipes''
*'' Memecylon ambrense''
*'' Memecylon amherstianum''
*'' Memecylon amoenum''
*'' Memecylon amplexicaule''
*'' Memecylon amplifolium''
*'' Memecylon amshoffiae''
*'' Memecylon andamanicum''
*'' Memecylon angustifolium''
*'' Memecylon antseranense''
*'' Memecylon apoense''
*'' Memecylon arcuatomarginatum''
*'' Memecylon argenteum''
*'' Memecylon arnhemense''
*''Memecylon auratifolium''
*'' Memecylon australissimum''
*'' Memecylon azurinii''
*'' Memecylon bachmannii''
*'' Memecylon bakerianum''
*'' Memecylon bakossiense''
*'' Memecylon balakrishnanii''
*'' Memecylon basilanense''
*'' Memecylon batekeanum''
*'' Memecylon bernierii''
*'' Memecylon bezavonense''
*''Memecylon biokoense''
*'' Memecylon boinense''
*'' Memecylon bokorensis''
*'' Memecylon borneense''
*'' Memecylon brachybotrys''
*'' Memecylon bracteatum''
*'' Memecylon bracteolatum''
*'' Memecylon brahense''
*'' Memecylon bremeri''
*'' Memecylon bretelerianum''
*'' Memecylon buxifolium''
*'' Memecylon buxoides''
*'' Memecylon caeruleum''
*'' Memecylon calderense''
*'' Memecylon calophyllum''
*'' Memecylon calyptratum''
*'' Memecylon campanulatum''
*'' Memecylon candidum''
*'' Memecylon candolleanum''
*'' Memecylon cantleyi''
*'' Memecylon capitellatum''
*'' Memecylon capuronii''
*'' Memecylon cardiophyllum''
*'' Memecylon caudatum''
*'' Memecylon celebicum''
*'' Memecylon centrale''
*'' Memecylon ceramense''
*'' Memecylon cerasiforme''
*'' Memecylon chevalieri''
*'' Memecylon cinereum''
*'' Memecylon clarkeanum''
*'' Memecylon clavistaminum''
*'' Memecylon cogniauxii''
*''Memecylon collinum''
*'' Memecylon confertiflorum''
*'' Memecylon confusum''
*'' Memecylon conocarpum''
*'' Memecylon constrictum''
*'' Memecylon cordatum''
*'' Memecylon cordifolium''
*'' Memecylon corticosum''
*'' Memecylon corymbiforme''
*'' Memecylon cotinifolioides''
*'' Memecylon coursianum''
*'' Memecylon courtallense''
*'' Memecylon crassifolium''
*'' Memecylon crassinerve''
*'' Memecylon crassipetiolum''
*'' Memecylon cumingii''
*'' Memecylon cuneatum''
*'' Memecylon dalleizettei''
*'' Memecylon dallmannense''
*'' Memecylon dasyanthum''
*'' Memecylon delphinense''
*'' Memecylon deminutum''
*'' Memecylon densiflorum''
*'' Memecylon dichotomum''
*''Memecylon diluviorum''
*'' Memecylon discolor''
*'' Memecylon dolichophyllum''
*'' Memecylon dubium''
*'' Memecylon durum''
*''Memecylon edule''
*'' Memecylon eduliforme''
*'' Memecylon eglandulosum''
*'' Memecylon elaeagni''
*'' Memecylon elegans''
*''Memecylon ellipticum''
*'' Memecylon elliptifolium''
*'' Memecylon elongatum''
*'' Memecylon emancipatum''
*''Memecylon englerianum''
*'' Memecylon erythranthum''
*'' Memecylon excelsum''
*'' Memecylon faucherei''
*'' Memecylon fernandesiorum''
*'' Memecylon fianarantse''
*'' Memecylon flavescens''
*'' Memecylon flavovirens''
*'' Memecylon floridum''
*'' Memecylon fragrans''
*'' Memecylon fruticosum''
*'' Memecylon fugax''
*'' Memecylon fuscescens''
*'' Memecylon galeatum''
*'' Memecylon garcinioides''
*'' Memecylon geddesianum''
*'' Memecylon geoffrayi''
*'' Memecylon germainii''
*'' Memecylon gibbosum''
*'' Memecylon giganteum''
*'' Memecylon gitingense''
*'' Memecylon gopalanii''
*'' Memecylon gracile''
*'' Memecylon gracilipedicellatum''
*'' Memecylon gracilipes''
*'' Memecylon gracillimum''
*'' Memecylon grande''
*'' Memecylon grandifolium''
*'' Memecylon greenwayi''
*'' Memecylon griffithianum''
*'' Memecylon hainanense''
*'' Memecylon harmandii''
*'' Memecylon helferi''
*'' Memecylon heyneanum''
*'' Memecylon hookeri''
*'' Memecylon huillense''
*'' Memecylon hullettii''
*'' Memecylon humbertii''
*''Memecylon hylandii''
*'' Memecylon hyleastrum''
*'' Memecylon idukkianum''
*'' Memecylon impressivenum''
*'' Memecylon inalatum''
*'' Memecylon incisilobum''
*'' Memecylon infuscatum''
*'' Memecylon insigne''
*'' Memecylon insperatum''
*'' Memecylon insulare''
*'' Memecylon interjectum''
*''Memecylon intermedium''
*'' Memecylon isaloense''
*'' Memecylon ivohibense''
*'' Memecylon jadhavii''
*'' Memecylon jambosioides''
*''Memecylon klaineanum''
*'' Memecylon kollimalayanum''
*'' Memecylon korupense''
*'' Memecylon kosiense''
*'' Memecylon kunstleri''
*'' Memecylon kupeanum''
*'' Memecylon kurichiarense''
*'' Memecylon lanceolatum''
*'' Memecylon lancifolium''
*''Memecylon langbianense''
*'' Memecylon laruei''
*'' Memecylon lateriflorum''
*''Memecylon laurentii''
*'' Memecylon laureolum''
*'' Memecylon lawsonii''
*'' Memecylon leucanthum''
*'' Memecylon liberiae''
*'' Memecylon ligustrifolium''
*'' Memecylon lilacinum''
*'' Memecylon littorale''
*'' Memecylon loheri''
*'' Memecylon longicuspe''
*'' Memecylon longifolium''
*'' Memecylon longipetalum''
*'' Memecylon louvelianum''
*'' Memecylon luchuenense''
*'' Memecylon lurerii''
*'' Memecylon lushingtonii''
*'' Memecylon macneillianum''
*'' Memecylon macrocarpum''
*'' Memecylon macrodendron''
*''Memecylon macrophyllum''
*'' Memecylon madgolense''
*'' Memecylon magnifoliatum''
*'' Memecylon malaccense''
*'' Memecylon mamfeanum''
*'' Memecylon mananjebense''
*'' Memecylon mandrarense''
*'' Memecylon mangiferoides''
*'' Memecylon manickamii''
*'' Memecylon mayottense''
*'' Memecylon megacarpum''
*'' Memecylon megaspermum''
*'' Memecylon memoratum''
*'' Memecylon merguicum''
*''Memecylon minimifolium''
*'' Memecylon minutiflorum''
*'' Memecylon mocquerysii''
*'' Memecylon monchyanum''
*''Memecylon mouririoides''
*'' Memecylon multinode''
*'' Memecylon mundanthuraianum''
*'' Memecylon myrianthum''
*'' Memecylon myricoides''
*'' Memecylon myrtiforme''
*'' Memecylon myrtilloides''
*'' Memecylon natalense''
*'' Memecylon nigrescens''
*'' Memecylon nodosum''
*'' Memecylon normandii''
*''Memecylon novoguineense''
*'' Memecylon nubigenum''
*''Memecylon obscurinerve''
*'' Memecylon obtusifolium''
*'' Memecylon occultum''
*'' Memecylon ochroleucum''
*'' Memecylon octocostatum''
*'' Memecylon odoratum''
*'' Memecylon oleifolium''
*''Memecylon oligophlebium''
*'' Memecylon orbiculare''
*'' Memecylon oubanguianum''
*'' Memecylon ovatifolium''
*''Memecylon ovatum''
*'' Memecylon ovoideum''
*'' Memecylon pachyphyllum''
*'' Memecylon pallidum''
*'' Memecylon paniculatum''
*'' Memecylon papuanum''
*'' Memecylon paradoxum''
*'' Memecylon parvifolium''
*'' Memecylon pauciflorum''
*'' Memecylon pedunculatum''
*'' Memecylon pendulum''
*'' Memecylon peracuminatum''
*'' Memecylon perangustum''
*'' Memecylon perditum''
*'' Memecylon pergamentaceum''
*'' Memecylon perplexum''
*'' Memecylon perrieri''
*'' Memecylon phanerophlebium''
*'' Memecylon phyllanthifolium''
*'' Memecylon pileatum''
*'' Memecylon planifolium''
*'' Memecylon plebejum''
*'' Memecylon polyanthemos''
*'' Memecylon polyanthum''
*'' Memecylon ponmudianum''
*''Memecylon procerum''
*'' Memecylon protrusum''
*'' Memecylon pseudomegacarpum''
*'' Memecylon pseudomyrtiforme''
*'' Memecylon pterocarpum''
*''Memecylon pterocladum''
*'' Memecylon pteropus''
*'' Memecylon pubescens''
*'' Memecylon pulvinatum''
*'' Memecylon purpurascens''
*'' Memecylon pusilliflorum''
*''Memecylon ramosii''
*'' Memecylon ramosum''
*'' Memecylon randerianum''
*'' Memecylon revolutum''
*'' Memecylon rheophyticum''
*'' Memecylon rhinophyllum''
*'' Memecylon rivulare''
*''Memecylon roseum''
*'' Memecylon rostratum''
*'' Memecylon rotundatum''
*'' Memecylon rovumense''
*''Memecylon royenii''
*'' Memecylon rubiflorum''
*'' Memecylon ruptile''
*'' Memecylon sabulosum''
*'' Memecylon sahyadricum''
*'' Memecylon salicifolium''
*'' Memecylon sambiranense''
*'' Memecylon schraderbergense''
*'' Memecylon schumannianum''
*'' Memecylon scolopacinum''
*'' Memecylon scutellatum''
*'' Memecylon sejunctum''
*''Memecylon semsei''
*'' Memecylon sepicanum''
*'' Memecylon sessile''
*'' Memecylon sessilifolium''
*''Memecylon simulans''
*''Memecylon sisparense''
*''Memecylon sitanum''
*'' Memecylon sivadasanii''
*'' Memecylon sivagirianum''
*''Memecylon sorsogonense''
*'' Memecylon soutpansbergense''
*'' Memecylon stenophyllum''
*'' Memecylon strumosum''
*'' Memecylon subcaudatum''
*'' Memecylon subcordifolium''
*'' Memecylon subcuneatum''
*'' Memecylon subfurfuraceum''
*'' Memecylon subramanii''
*'' Memecylon subsessile''
*'' Memecylon sumatrense''
*''Memecylon sylvaticum''
*'' Memecylon symplociforme''
*'' Memecylon talbotianum''
*''Memecylon tayabense''
*'' Memecylon teitense''
*'' Memecylon tenuipes''
*'' Memecylon terminale''
*'' Memecylon terminaliiflorum''
*'' Memecylon tetrapterum''
*'' Memecylon thouarsianum''
*'' Memecylon thouvenotii''
*'' Memecylon tirunelvelicum''
*'' Memecylon toamasinense''
*'' Memecylon torrei''
*'' Memecylon torricellense''
*'' Memecylon travancorense''
*'' Memecylon tricolor''
*'' Memecylon trunciflorum''
*'' Memecylon tsaratananense''
*''Memecylon uapacoides''
*'' Memecylon ulopterum''
*''Memecylon umbellatum''
*'' Memecylon urceolatum''
*'' Memecylon utericarpum''
*'' Memecylon vaccinioides''
*'' Memecylon varians''
*'' Memecylon venosum''
*'' Memecylon verruculosum''
*'' Memecylon virescens''
*'' Memecylon viride''
*''Memecylon vitiense''
*'' Memecylon wallichii''
*'' Memecylon wayanadense''
*'' Memecylon wightii''
*'' Memecylon xiphophyllum''
*'' Memecylon zambeziense''
*''Memecylon zenkeri''

A number of taxa have been recently re-assigned to other species. These include:
*'' Memecylon arnottianum'' , synonym of Lijndenia capitellata
*''Memecylon bequaertii'' , synonym of Warneckea bequaertii
*'' Memecylon elegantulum'' , synonym of Memecylon rostratum
*'' Memecylon gardneri'' , synonym of Lijndenia gardneri
*''Memecylon sessilicarpum'' , synonym of Warneckea sessilicarpa
*'' Memecylon sphaerocarpum'' , synonym of Memecylon ovatifolium

Sources

* Plants of World Online ((POWO): ''Memecylon'' L.: Accepted Species.

References

{{Taxonbar, from=Q2705135

Melastomataceae genera
Taxonomy articles created by Polbot
Taxa named by Carl Linnaeus