Flora of Madagascar

Source: Wikipedia, the free encyclopedia.

traveller's tree
.

The flora of Madagascar consists of more than 12,000

extant plant groups immigrated via across-ocean dispersal
well after continental break-up.

After its continental separation, Madagascar probably experienced a dry period, and

spiny thicket in the southwest, where rainfall is lowest and the wet season shortest. Mangroves occur on the west coast, and a variety of wetland
habitats with an adapted flora are found across the island.

The first human presence in Madagascar dates only 2000–4000 years back, and settlement in the interior occurred centuries later. The

cloves, coffee, and vanilla, the latter one of the country's main export produce today. More than 1,300 introduced plants are known, of which around 600 have become naturalised, and some invasive
.

Human population growth and economic activity have put pressure on natural vegetation in the region, especially through

Tsingy de Bemaraha and the Rainforests of the Atsinanana. While historically mainly European naturalists described Madagascar's flora scientifically, today a number of national and international herbaria, botanical gardens and universities document plant diversity and engage in its conservation
.

Diversity and endemism

Madagascar has been described as "one of the most floristically unique places in the world".

Physenaceae, Sarcolaenaceae and Sphaerosepalaceae.[2] As many as 96% of Madagascan trees and shrubs are estimated to be endemic.[4]

Vascular plants

Detail of a star-shaped yellow orchid flower with six tepals
Angraecum sesquipedale (Darwin's orchid), one of over 900 orchids in Madagascar

Among the non-flowering plants, ferns, lycophytes and allies count roughly 570 described species in Madagascar. About half of these are endemic; in the scaly tree fern family Cyatheaceae, native to the humid forests, all but three of 47 species are endemic. Six conifers in genus Podocarpus – all endemic – and one cycad (Cycas thouarsii), are native to the island.[2]

In the

flowering plants, basal groups and magnoliids account for some 320 Madagascan species, around 94% of which are endemic. The families most rich in species are Annonaceae, Lauraceae, Monimiaceae, and Myristicaceae, containing mainly trees, shrubs, and lianas, and the predominantly herbaceous pepper family (Piperaceae).[2]

Orchidaceae), with over 900 species of which 85% are endemic. Palms (Arecaceae) have around 200 species in Madagascar (most in the large genus Dypsis), more than three times as many as in continental Africa; all but five are endemic. Palm genera that are endemic to Madagascar are Beccariophoenix, Bismarckia, Dypsis, Lemurophoenix, Marojejya, Masoala, Ravenea, Satranala, Tahina, and Voanioala.[5]

Other large monocot families include the Pandanaceae with 88 endemic pandan (Pandanus) species, mainly found in humid to wet habitats, and the Asphodelaceae, with most species and over 130 endemics in the succulent genus Aloe. Grasses (Poaceae, around 550 species[6]) and sedges (Cyperaceae, around 300) are species-rich, but have lower levels of endemism (40%[6] and 37%, respectively). The endemic traveller's tree (Ravenala madagascariensis), a national emblem and widely planted, is the sole Madagascan species in the family Strelitziaceae.[2]

Top of the stem of a succulent, spiny plant, with yellow flowers
Euphorbia iharanae, an endemic succulent from the north of Madagascar

The eudicots account for most of Madagascar's plant diversity. Their most species-rich families on the island are:[2]

Non-vascular plants

A checklist from 2012 records 751

hornworts. About 34% of the mosses and 19% of the liverworts are endemic. It is unknown how many of these species may have gone extinct since their discovery, and a number likely remain to be described.[7]

Fungi

Many undescribed species of

fungi are suspected in Madagascar.[8] A number of edible mushrooms are consumed in the country, especially from the genera Auricularia, Lepiota, Cantharellus (the chanterelles), and Russula (the brittlegills).[8][9] Most of the ectomycorrhizal species are found in plantations of introduced eucalyptus and pine, but also in native tapia (Uapaca bojeri) woodlands.[8] The chytrid fungus Batrachochytrium dendrobatidis, responsible for chytridiomycosis, an infectious disease threatening amphibian populations worldwide, was long considered absent from Madagascar. In 2010 it was recorded, however, and has been confirmed since in various areas and in numerous frog families, alerting scientists to a new threat to the island's already endangered frog fauna.[10]

Over 500 species of

lichens of Madagascar have been documented, but the true number was estimated to be at least twice as high. Wet tropical areas of siliceous bedrock make up approximately two-thirds of the country, and are where most of the lichens have been found. Dry tropical areas of granitic and limestone bedrock make up the other one-third of the country with just over 20 species documented in these habitats.[11]

Algae

paleoclimatic variations on the island.[12]

Vegetation types

Two maps of Madagascar, showing land cover on the left and topography on the right
Two maps of Madagascar, showing land cover on the left and topography on the right

Land cover according to the Atlas of the vegetation of Madagascar (2007)[13]

  Plateau grassland–wooded grassland mosaic (41.67%)
  Wooded grassland–bushland mosaic (22.93%)
  Humid forest (8.06%)
  Degraded humid forest (9.81%)
  Western dry forest (5.4%)
  South western dry spiny forest-thicket (3.1%)
  Cultivation (likely underestimated) (3.97%)
  Other (5.06%)

Madagascar features contrasting and unique

tsingy formations), and unconsolidated sand.[13]

The marked east–central–west distinction among Madagascan flora was already described by the English naturalist

Humid forests

Masoala Peninsula[16] – and the dry season is short or absent. The predominantly evergreen forest, up to 35 m high (115 ft), is composed of tree and understory species from various families such as Burseraceae, Ebenaceae, Fabaceae, and Myristicaceae; bamboos and lianas are frequent. Cyclones hit the east coast of Madagascar in some years and can destroy habitats.[16] The WWF classifies the eastern belt of humid forest, below 800 m (2,600 ft) elevation, in the "lowland forests" ecoregion[16] and the montane forests of the highlands in the "sub-humid forests" ecoregion.[17]

Degraded humid forest (savoka in Malagasy) covers about ten percent of the island. It spans various states of degradation and is composed of forest remnants and planted or otherwise introduced species. It is primarily the result of slash-and-burn cultivation in primary forest. Some forest fragments still harbour a considerable amount of biodiversity.[13]

Littoral forest, found in several isolated areas along the eastern coast, covers less than 1% of the land area, on mainly sandy sediments. Climate is humid, with 1,300–3,200 mm (51–126 in) annual rainfall. Littoral forest covers sandy soil forest, marsh forest, and grasslands. Its flora includes various tree families, lianas, and

epiphytic orchids and ferns; in the marsh forests, pandans (Pandanus) and the traveller's tree (Ravenala madagascariensis) are common.[13] It is part of the WWF's "lowland forests" ecoregion.[16]

An isolated area of humid forest in the south west, on the eastern slope of the Analavelona massif, is classified as "Western humid forest" by the Atlas. It occurs on lavas and sand, at 700–1,300 m (2,300–4,300 ft) elevation. The forest is maintained through condensating moisture from ascending air. It is unprotected but the local population considers it sacred.[13] The WWF includes it in the "sub-humid forests" ecoregion.[17]

Dry forests and thicket

Dry forest, accounting for roughly 5% of the surface, is found in the west, from the northern tip of the island to the

dry deciduous forest" ecoregion[18] and the southern part, including the northernmost range of Didiereaceae, as "succulent woodlands" ecoregion.[19]

"Western sub-humid forest" occurs inland in the southwest and covers less than 1% of the surface, mainly on sandstone, at 70–100 m (230–330 ft) elevation. Climate is sub-humid to sub-arid, with 600–1,200 mm (24–47 in) annual rainfall. The vegetation, up to 20 m tall (66 ft) with a closed canopy, includes diverse trees with many endemics such as baobabs (Adansonia), Givotia madagascariensis, and the palm Ravenea madagascariensis. Cutting, clearing and invasive species such as opuntias and agaves threaten this vegetation type.[13] It is part of the WWF's "sub-humid forests" ecoregion.[17]

The driest part of Madagascar in the southwest features the unique "

Didiereoideae, baobabs, and Euphorbia species. A more open coastal bushland within the region is classified separately by the Atlas. Degraded spiny forest accounts for c. 1% of the surface and is the result of cutting, clearing, and encroachment. Introduced species such as agaves and opuntias are found with remnants of the native flora.[13]

Grassland, woodland, and bushland

Grasslands dominate a large part of Madagascar, more than 75% according to some authors.[23] Mainly found on the central and western plateaus, they are dominated by C4 grasses such as the common Aristida rufescens and Loudetia simplex and burn regularly. While many authors interpret them as the result of human degradation through tree-felling, cattle raising and intentional burning, it has been suggested that at least some of the grasslands may be primary vegetation.[23][6] Grassland is often found in a mixture with trees or shrubs, including exotic pine, eucalypt, and cypress.[13]

The Atlas distinguishes a "wooded grassland–bushland mosaic" covering 23% of the surface and a "plateau grassland–wooded grassland mosaic" covering 42%. Both occur on various substrates and account for most of the WWF's "sub-humid forests" ecoregion.

hard-leaved vegetation that includes Asteraceae, Ericaceae, Lauraceae, and Podocarpaceae shrubs, among others,[13] and is singled out by the WWF as "ericoid thickets" ecoregion.[21]

An evergreen open forest or woodland type, tapia forest, is found on the western and central plateaus, at altitudes of 500–1,800 m (1,600–5,900 ft). It is dominated by the eponymous tapia tree (Uapaca bojeri) and covers less than 1% of the surface. The broad regional climate is sub-humid to sub-arid, but tapia forest is mainly found in drier microclimates. Trees other than tapia include the endemic Asteropeiaceae and Sarcolaenaceae, with a herbaceous understory. Tapia forest is subject to human pressure, but relatively well adapted to fire.[13] It falls in the WWF's "sub-humid forests" ecoregion.[17]

Wetlands

Marshes,

Peat bogs are restricted to highlands above 2,000 m (6,600 ft) elevation; their distinct vegetation includes, among others, Sphagnum moss and sundew species (Drosera). Many wetlands have been converted into rice paddies and are otherwise threatened by destruction and pollution.[13]

Mozambique channel coast, from the very north to just south of the Mangoky river delta. Eleven mangrove tree species are known from Madagascar, of which the most frequent belong to the families Acanthaceae, Lecythidaceae, Lythraceae, Combretaceae, and Rhizophoraceae. Mangrove forests are threatened by encroachment and cutting.[13] The WWF lists the mangroves of Madagascar as separate ecoregion.[22]

Examples of native vegetation in Madagascar
Typhonodorum lindleyanum

Origins and evolution

Paleogeography

Map showing Nepenthes distribution around the Indian Ocean
Yellow-green Nepenthes pitchers in close-up
Nepenthes pitcher plants mainly occur in Southeast Asia but extend west to Madagascar (below: N. madagascariensis).

Madagascar's high species richness and endemicity are attributed to its long isolation as a

Mascarenes, and Seychelles. There are however also links to other, more distant floras, such as those of India and Malesia.[27]

After their separation from Africa, Madagascar and India moved northwards, to a position south of 30° latitude. During the

subtropical ridge. This passage likely induced a dry, desert-like climate across the island, which later contracted to what is today the sub-arid spiny thicket in the southwest. Humid forests probably established since the Oligocene, when India had cleared the eastern seaway, allowing trade winds to bring in rainfall, and Madagascar had moved north of the subtropical ridge. The intensification of the Indian Ocean monsoon system after around eight million years ago is believed to have further favoured the expansion of humid and sub-humid forests in the Late Miocene, especially in the northern Sambirano region.[24] Some of the grasslands may also date to the Late Miocene, when there was a global grassland expansion.[23]

Species evolution

Several hypotheses exist as to how plants and other organisms have diversified into so many species in Madagascar. They mainly assume either that species diverged in parapatry by gradually adapting to different environmental conditions on the island, for example dry versus humid, or lowland versus montane habitats, or that barriers such as large rivers, mountain ranges, or open land between forest fragments, favoured allopatric speciation.[28] A Madagascan lineage of Euphorbia occurs across the island, but some species evolved succulent leaves, stems and tubers in adaptation to arid conditions.[29] In contrast, endemic tree ferns (Cyathea) all evolved under very similar conditions in Madagascan humid forests, through three recent radiations in the Pliocene.[30]

giant lemurs probably deprived some large-seeded plants of their seed dispersers.[33] The extinct Madagascan megafauna also included grazers such as two giant tortoises (Aldabrachelys) and the Malagasy hippopotamuses, but it is unclear to what extent their habitats resembled today's widespread grasslands.[34]

Exploration and documentation

Early naturalists

Cover page of Flacourt's book "Histoire de la grande isle Madagascar"
Histoire de la grande isle Madagascar by Étienne de Flacourt (1658) was the first detailed written account on Madagascar.

Madagascar and its natural history remained relatively unknown outside the island before the 17th century. Its only overseas connections were occasional Arab, Portuguese, Dutch, and English sailors, who brought home anecdotes and tales about the fabulous nature of Madagascar.

growing influence of the French in the Indian Ocean, it was mainly French naturalists that documented Madagascar's flora in the following centuries.[2][36]

Jean-Baptiste de Lamarck and Jean Louis Marie Poiret in France.[36]: 93–95  Sonnerat described, among others, the emblematic traveller's tree.[40] Another contemporary, Louis-Marie Aubert du Petit-Thouars, also visited Madagascar from the Isle de France; he collected on the island for six months and wrote, among others, Histoire des végétaux recueillis dans les îles australes d'Afrique[41] and a work on orchids of Madagascar and the Mascarenes.[36]: 344–345 [42]

19th to 20th century

French naturalist Alfred Grandidier was a preeminent 19th-century authority on Malagasy wildlife. His first visit in 1865 was followed by several other expeditions. He produced an atlas of the island and, in 1885, published L'Histoire physique, naturelle et politique de Madagascar, which would comprise 39 volumes.[43] Although his main contributions were in zoology, he was also a prolific plant collector; several plants were named after him, including Grandidier's baobab (Adansonia grandidieri) and the endemic succulent genus Didierea.[36]: 185–187  The British missionary and naturalist Richard Baron, Grandidier's contemporary, lived in Madagascar from 1872 to 1907 where he also collected plants and discovered up to 1,000 new species;[44] many of his specimens were described by Kew botanist John Gilbert Baker.[14] Baron was the first to catalogue Madagascar's vascular flora in his Compendium des plantes malgaches, including over 4,700 species and varieties known at that time.[44]

Sartidia perrieri, a now-extinct grass species collected only once, by Henri Perrier de la Bâthie in 1914, and described by Aimée Camus

During the

mycologists working in Madagascar.[47]

Research in the 21st century

Today, national and international research institutions are documenting the flora of Madagascar. The

University of Antananarivo has a department for plant biology and ecology.[50]

Outside the country, Royal Botanic Gardens, Kew, is one of the leading institutions in the revision of Madagascar's plant families; it also maintains the

Millennium Seed Bank project.[51] The National Museum of Natural History in Paris has traditionally been one of the centres of research on the flora of Madagascar. It holds a herbarium with roughly 700,000 Malagasy plant specimens and a seed bank and living collection, and continues to edit the Flore de Madagascar et des Comores series begun by Humbert in 1936.[47] The Missouri Botanical Garden maintains the Catalogue of the plants of Madagascar, a major online resource,[2] and also has a permanent base in Madagascar.[52]

Human impact

Madagascar was colonised rather recently compared to other landmasses, with first evidence for humans – arrived from either Africa or Asia – dating to 2,300

years before present.[54] It is assumed that humans first stayed near the coast and penetrated into the interior only several centuries later. The settlers had a profound impact on the long-isolated environment of Madagascar through land clearing and fire, introduction of zebu cattle, and probably hunting to extinction the native megafauna including, among others, elephant birds, giant lemurs and giant tortoises.[53][55] The first Europeans arrived in the 16th century, starting an age of overseas exchange. Population growth and transformation of the landscape was particularly rapid since the mid-20th century.[53]

Uses of native species

An elderly man sitting with a valiha instrument on his knees in front outside a house
The Malagasy valiha zither is made from bamboo.

The native flora of Madagascar has been and still is used for a variety of purposes by the

Malagasy people. More than a hundred plants used locally and commercially were described at the end of the 19th century by the English naturalist Richard Baron. These included many timber trees such as native ebony (Diospyros) and rosewood (Dalbergia) species, the raffia palm Raphia farinifera used for fibre, dyeing plants, as well as medicinal and edible plants.[56]

The traveller's tree has various uses in the east of Madagascar, chiefly as building material.[57] Madagascar's national instrument valiha is made from bamboo and lent its name to the endemic genus Valiha.[58] Yams (Dioscorea) in Madagascar include introduced, widely cultivated species as well as some 30 endemics, all edible.[59] Edible mushrooms, including endemic species, are collected and sold locally (see above, Diversity and endemism: Non-vascular plants and fungi).[8]

Many native plant species are used as herbal remedies for a variety of afflictions. An

ethnobotanical study in the southwestern littoral forest, for instance, found 152 native plants used locally as medicine,[60] and countrywide, over 230 plant species have been used as traditional malaria treatments.[61] The diverse flora of Madagascar holds potential for natural product research and drug production on an industrial scale; the Madagascar periwinkle (Cataranthus roseus), a source of alkaloids used in the treatment of different cancers, is a famous example.[62]

Agriculture

One of the characteristic features of agriculture in Madagascar is the widespread cultivation of

Imerina kingdom in the 19th century.[63] Land conversion for rice cultivation has been an important cause of wetlands loss.[13]

Other major crops, such as

cloves, coconut, coffee and vanilla in plantations.[67]: 107  Today Madagascar is the primary vanilla-producing country worldwide.[68]

Forestry in Madagascar involves many exotic species such as eucalypts, pines and acacias.[66] The traditional slash-and-burn agriculture (tavy), practised for centuries, today accelerates the loss of primary forests as populations grow[69] (see below, Threats and conservation).

Introduced plants

More than 1,300

Eichhornia crassipes), which spread widely through subtropical and tropical regions and is considered a serious plant pest in wetlands.[70] In general, invasive plants spread mostly in already disturbed, secondary vegetation, and the remaining primary forests of the east appear little affected.[71]

A prickly pear cactus,

biological control for the plant, which had become a nuisance; they rapidly eradicated most of the cacti. This probably led to famine among the Antandroy people, although some authors challenge the causal link between famine and cactus eradication. Today, several Opuntia species are again present mainly in the south, spreading into native vegetation in some areas.[72]

The prickly pear illustrates the dilemma of plant introductions: while many authors see exotic plants as a threat to the native flora,[13][71] others argue that they have not yet been linked directly to the extinction of a native species, and that some may actually provide economic or ecological benefits.[66] A number of plants native to Madagascar have become invasive in other regions, such as the traveller's tree in Réunion and the flamboyant tree (Delonix regia) in various tropical countries.[71]

Threats and conservation

Tree trunks piled up in front of a forest background, with workers cutting them
Illegal logging of rosewood in the north of Madagascar

Madagascar, together with its neighbouring islands, is considered a

Red List assessed 1,676 endemic vascular plant species and found over 1,000 of them to be endangered or critically endangered.[75]

Rapid human population increase and economic activity entail

Global warming is expected to reduce or shift climatically suitable areas for plant species and threatens coastal habitats, such as littoral forests, through rising sea levels.[80]

Conservation of natural habitats in Madagascar is concentrated in over six million hectares (23,000 sq mi) – about ten percent of the total land surface – of

QMM mining company include the setting aside of conservation areas and habitat restoration.[79]

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