Introduction to baobabs

Local people have carved footholds into these stunted baobabs (A. grandidieri) to help them gather the nutritious fruit and leaves. Andavadoaka. (L.J.)

DISTRIBUTION

Like many other mammal-pollinated flowers, A. grandidieri blooms are shaped like a shaving brush. (L.J.)

Madagascar drifted away from Africa around 160 million years ago, during the breakup of the prehistoric supercontinent Gondwanaland. Since about 90 million years ago, the island has remained separate from other landmasses, allowing its flora and fauna to evolve and differentiate in near isolation. Madagascar has therefore inherited an incredibly rich and divergent biodiversity, with thousands of species that are found nowhere else on Earth. Of its 11,031 native plant species, 82% are endemic to the island, along with 100% of its non-flying mammals, 99% of frogs, 92% of reptiles, 65% of freshwater fishes and 52% of birds. At higher taxonomic levels Madagascar’s endemism is even more impressive; with 24 endemic families of plant and animal (not including invertebrates), the island constitutes of one the world’s 12 centres of megadiversity. Its biodiversity plays a fundamental role in the country’s economic development and human well-being, but sadly its natural ecosystems are characterized by an extreme fragility, and habitat degradation is taking place at an increasingly alarming rate.

Baobabs illustrate the spectacular diversity, endemism and vulnerability of the Malagasy flora. Madagascar is the centre of baobab diversity: of the world’s eight species, six are unique to the island (Adansonia grandidieri, A. madagascariensis, A. perrieri, A. rubrostipa, A. suarezensis and A. za). Three of these species are considered to be Endangered by the IUCN, and a further two are Near Threatened, due to extensive habitat loss, indicating the urgent need for conservation action. In contrast, the African Baobab (A. digitata) is found throughout the semi-arid regions of sub-Saharan Africa and has been widely planted in tropical countries around the world, including Haiti, Venezuela, India, Java and Madagascar. The eighth species, the Boab (A. gregorii), grows in the drylands of northwestern Australia.

The natural distribution of all eight species of Adansonia, clearly showing how much more widespread the African Baobab (A. digitata) is compared with the other seven species.

BOTANICAL CLASSIFICATION

The flower of the African Baobab (Adansonia digitata). (L.J.)

The fruit of the African Baobab, which is known as bu hibab in Arabic, giving rise to the common name baobab. (S.V.)

Baobabs (genus Adansonia) were formerly placed within the family Bombacaceae. However, molecular studies revealed that they should be reassigned to the subfamily Bombacoideae within the huge cosmopolitan mallow family, Malvaceae. The Bombacoideae contains 27 genera and about 250 species, the majority of which occur in the New World tropics. These include some economically important species such as the South American Balsa (Ochroma pyramidale), which yields one of the world’s softest commercially traded hardwoods (used in model-making, aircraft and film props), and the Kapok (Ceiba pentandra), whose long fruits produce cotton-like fibres used as stuffing material.

Carolus Linnaeus (1707–1778) named the genus Adansonia in honour of the French botanist and explorer Michel Adanson (1727–1806), following the suggestion of Bernard de Jussieu (1699–1777). Although the fruit of the African Baobab (A. digitata) had long been known to European botanists, Adanson was the first to formally describe the tree, which he referred to as the ‘calabash tree’, in 1757. He based his description on information gathered during an exploratory voyage to Senegal while working for the French East India Company. He also collected a prodigious amount of information on the plants, animals, commerce and languages of Senegal.

Although the common name for trees of the genus Adansonia is baobab, this word is not actually derived from any African or Malagasy dialect. This name first appeared in 1592, spelled bahobab, in De Plantis Aegypti Liber, a book of Egyptian flora by the Italian doctor and botanist Prosper Alpino (1553–1617). In this volume, Alpino described fruits of A. digitata that were sold on the markets of Cairo under the name bahobab, which was likely derived from the Arabic bu hibab meaning ‘many-seeded fruit’. When Adanson described the tree, he named it baobab in light of Alpino’s work, but later this name came to be used to refer to all species of Adansonia.

Theodore de Bry (1528–1598) published the first record of a Malagasy baobab species in 1605 in Indiae Orientalis. His illustrated plates depicted peculiar, bottle-shaped trees, thought to correspond to A. rubrostipa, but it wasn’t until the end of the 19th century that Henri Baillon (1827–1895) confirmed the presence of the genus Adansonia in Madagascar. In 1952, the French botanist Henri Perrier de la Bâthie (1873–1958) published the first full description of the known baobab species of Madagascar (this paper excluded A. perrieri which was only described in 1960 after De la Bâthie’s death). Today, the seminal paper on the systematics of Adansonia remains David Baum’s revision published in 1995.

A bottle-shaped Adansonia rubrostipa putting out new leaves in preparation for the rainy season. Mikea Forest. (L.J.)

DESCRIPTION

This young A. grandidieri is branched and conical, typical for a juvenile baobab. (A.P.)

The eight species in the genus Adansonia are tropical, deciduous trees with mesic, palmate leaves and perfect (hermaphroditic) flowers. They have massive, swollen trunks reaching heights of 5–30m, which tend to narrow abruptly just beneath the canopy of branches, giving the tree a bottle-shaped appearance. The trees are usually ‘emergents’, exceeding the height of the surrounding forest or thicket. Juvenile baobabs tend to have slender, tapering trunks that are swollen at the base. The soft, spongy wood may contain up to 79% water, while the wood of more ‘typical’ trees, such as the sycamore (Acer pseudoplatanus), may contain just 27% water.

Other stem-succulents, such as Pachypodium geayi, are often confused with baobabs. (L.J.)

Among the largest stem-succulent plants in the world, baobabs, with their strange growth form, represent one of the most impressive adaptations to a semi-arid environment. Although it was long assumed that baobabs store water in their trunks for use during drought, recent research indicates they actually store this water for structural stability. If a baobab were to use too much water during the dry season, it might collapse under the weight of its branches! In fact, the tree seems to use a very small amount of water to produce new leaves at the end of the dry season, though these will remain inactive (i.e. their stomata stay closed) until the first rain falls. This strategy enables the tree to begin photosynthesizing immediately, taking full advantage of the short and unpredictable rainy season. Leaves are important organs, converting sunlight and carbon dioxide into vital nutrients for the plant, but they also lose plenty of water via transpiration. Therefore, baobabs drop their leaves shortly after the last rains, thus conserving water.

Dwarf specimens of Adansonia grandidieri in subdesert habitat near Andavadoaka. Local people harvest the nutritious leaves for cattle fodder, giving the branches a stunted appearance. (A.P. and L.J.)

An example of the tall, columnar growth form of A. grandidieri common in the Menabe region. (L.J.)

These spectacular trees are renowned for their twisted, root-like branches, which give them the appearance of having been planted upside down. Their actual root system is equally impressive, extending some 50m from the trunk. However, as in many succulent plants, the root systems of mature baobabs are relatively shallow, often running along the surface of the soil and rarely descending more than 2m deep. This allows the tree to take advantage of even the lightest rainfall, essential for survival in a dry climate.

Dwarf specimens of Adansonia grandidieri in subdesert habitat near Andavadoaka. Local people harvest the nutritious leaves for cattle fodder, giving the branches a stunted appearance. (A.P. and L.J.)

Baobabs grow in numerous different habitats, but local variations in rainfall and soil quality can drastically affect their growth rate and appearance. Under more favourable conditions, for example those that prevail in the forests near Morondava, Adansonia grandidieri achieves the massive, columnar growth form for which it is famous, becoming up to 30m tall. On the dry, rocky limestone soils and salt pans around Andavadoaka, however, this species is reduced to a stout, stumpy tree of just 4–5m in height. In extremely dry regions, such as the Mahafaly Plateau in southwestern Madagascar, some dwarf baobabs have developed highly wrinkled bark following years of repeated shrinkage and swelling of the trunk. Some specimens of A. rubrostipa and A. grandidieri have highly decorative bark, with intricate swirls and spirals in many shades of red, brown and yellow. These natural patterns are likely caused by fungal growth, though the phenomenon has yet to be studied.

The Grandmother Baobab is an ancient, stunted and wrinkled Adansonia rubrostipa from Tsimanampesotse National Park. (L.J.)

The flowers of the Brevitubae group (such as this A. grandidieri flower) are adapted to attract small mammal pollinators such as fruit bats and nocturnal lemurs. (A.P.)

The flowers are large, showy and fragrant, opening just before or after dusk. The ripe buds open remarkably quickly, usually within 30 minutes. In fact, the flowers of several species (including A. rubrostipa) can open fully in just 30 seconds – the movements of the unfurling petals are easily detected with the naked eye. Although the flowers may remain on the tree for several days, they are reproductively viable for 15 hours at the very most. The eight species in genus Adansonia can be divided into three groups according to the structure of their flowers: Brevitubae, Longitubae and Adansonia. Each group attracts specific types of animal pollinators that have adapted to feed on flowers in different ways.

A. rubrostipa flowers (like those of other species in the Longitubae group) are mainly pollinated by hawkmoths. (L.J.)

The velvety, hard-shelled fruit of Adansonia rubrostipa. (A.P.)

The Brevitubae group contains two species, A. grandidieri and A. suarezensis, whose flower buds are held aloft on short, erect stems. Their flowers are white and resemble a shaving brush, and are primarily pollinated by mammals such as the Madagascar Straw-coloured Fruit Bat (Eidolon dupreanum) and the Pale Fork-marked Lemur (Phaner pallescens). The Longitubae group comprises the other four Malagasy species (A. madagascariensis, A. perrieri, A. rubrostipa and A. za) and the Australian Boab (A. gregorii). These have long, cylindrical flower buds on erect or horizontal stems with red, yellow, orange or white petals, and are mainly pollinated by long-tongued hawkmoths. In Madagascar, the hawkmoth species Coelonia solanii and Xanthopan morgani are important pollinators of Longitubae baobabs, though the Pale Fork-marked Lemur and Fat-tailed Dwarf Lemur (Cheirogaleus medius) are also thought to play a role. Souimanga Sunbirds (Nectarinia souimanga), Ring-tailed Lemurs (Lemur catta) and various butterflies also visit the flowers to feed on nectar and pollen, but their manner of feeding makes them unlikely pollinators. The Boab is mainly pollinated by the hawkmoth Agrius convolvuli. The Adansonia group contains just one species, the African Baobab (A. digitata), whose pendulous white flowers are suspended on a long stalk and are mainly pollinated by fruit bats (such as Rousettus aegyptiacus) and, to a lesser extent, bushbabies (Galago spp.).

A Souimanga Sunbird (Nectarinia souimanga) drinks nectar from an A. grandidieri flower. Its habit of feeding at the side of the flower means it is very unlikely to transfer pollen to the stigma, which is raised above the stamens. (L.J.)

Baobabs produce quite large, round to oblong, indehiscent fruits covered with soft brown or grey hairs that feel velvety to the touch. The hard outer shell (pericarp) encloses numerous kidney-shaped seeds, which are embedded in a matrix of fibres and spongy or chalky pulp. The African Baobab is also known as the Cream of Tartar Tree, as its fruit pulp contains high levels of tartaric acid, which is used in baking. Fruit shape and size vary considerably between the different baobab species, and are also highly variable within A. digitata, A. gregorii and A. za. For example, A. za usually has oval-oblong fruits, but in some parts of its range the fruits are narrow and pointed (see first image, second row on p.100).

In Africa and Australia, baobab seeds are mainly dispersed by animals, but in Madagascar (where there is no known native animal disperser) they are likely spread via water.

A Ring-tailed Lemur (Lemur catta) consumes an A. rubrostipa flower. (L.J.)

Branches of a mature A. rubrostipa, heavily laden with ripening fruit. (A.P.)

LIFE AND DEATH

Baobab wood is composed of concentric sheets of fibre. Once dead, the wood decomposes rapidly. (A.P. and L.J.)

Baobabs can grow to an enormous size, and many look incredibly ancient – but how old are they really? During his visit to the Madeleine Islands off the coast of Senegal in 1749, Michel Adanson came across two giant baobabs whose bark sported inscriptions carved by sailors in the 15th century. He estimated their annual growth rate by comparing them with younger baobabs of a known age, and concluded that these huge trees must be more than 5,000 years old. Adanson’s calculations sparked lively controversy among the scientific community, including the notable explorer and missionary David Livingstone (1813–1873). The latter argued that no tree could have survived the biblical flood, which is said to have taken place just 3,500 years ago, and refused to believe that the giant baobabs could be as old as the Egyptian pyramids.

Those contentious trees no longer exist, but modern-day researchers continue to investigate the age of other gargantuan baobabs. Growth-rate measurements indicate that the largest living specimens must be many hundreds of years old, but for several reasons baobabs remain very difficult to age accurately using traditional methods. Firstly, while the age of most trees can be ascertained by cutting a transverse section through the trunk and counting the annual growth rings, this method cannot be used for baobabs because of the structure of their wood, which is soft and fibrous with poorly defined growth rings. These rings do not necessarily correspond to a yearly growth pattern; for example, older trees eventually stop laying down new rings, while the central rings may fuse or even rot away if the tree becomes hollow in old age. Also, when a baobab dies it rapidly collapses into an unsightly, fibrous heap and decomposes completely within a few years, making it difficult for researchers to study very old specimens.

The African Baobab (Adansonia digitata) is the longest living angiosperm tree in the world. Kruger National Park. (L.J.)

The sacred Andombiry Baobab (A. grandidieri) is the largest baobab (and possibly one of the oldest) in Madagascar with a 27.4m circumference (6m diameter). (L.J.)

In the foreground lie the dry, fibrous remains of a dead Adansonia grandidieri. Andavadoaka. (L.J.)

The largest African Baobabs were once believed to be as much as 5,000 years old, but recent research suggests 1,000–2,000 years may be a more realistic estimate. (L.J.)

Secondly, inaccuracies arise when aging a baobab based on the circumference of its trunk. The relationship between age and circumference is not necessarily linear, as the rate of growth is not constant throughout a baobab’s life, and the largest trees may not be the oldest. Growth rate is also greatly affected by local environmental conditions: if we were to plant two baobab seeds of the same species in two different regions, one receiving plentiful rain and the other suffering frequent droughts, the former would grow faster and larger than the latter. In 100 years’ time, the two trees would likely differ greatly in girth and height despite being the same age.

Although traditional methods cannot be used to gauge a baobab’s age accurately, the advent of modern chemistry and radiocarbon dating has enabled researchers to begin to resolve the debate. For example, the famous Glencoe Baobab – a large and very old African Baobab growing in Limpopo Province, South Africa – suffered a major split in 2009, providing a unique opportunity for scientists to study its age. The researchers collected and analysed several samples from the damaged trunk, and the results were startling: the oldest samples were around 1,835 years old, making the Glencoe Baobab the oldest known angiosperm tree! Similar research was carried out on the living Grandmother Baobab in southern Madagascar, which was found to be around 1,600 years old. This research indicates that many large baobab trees may indeed be extremely old, but perhaps not as ancient as many people once believed them to be.

BIOGEOGRAPHY

The ‘baobab forests’ of western Madagascar are dominated by three baobab species, including Adansonia grandidieri. Kirindy-Mite National Park. (X.V.)

The genus Adansonia has a disjointed distribution that baffled botanists for nearly two centuries. It is native to Madagascar, sub-Saharan Africa and Australia – landmasses that have been separated by hundreds of miles of open ocean for millions of years. Given that Madagascar is the centre of diversity for the genus, it has generally been considered to be the place where baobabs first evolved. However, if this is true, then how did baobabs arrive in Africa and Australia?

Initially, baobabs were thought to be relics of an ancient flora that had evolved more than 120 million years ago while Africa, Madagascar and Australia were connected to the Gondwanan supercontinent. When the great continent split apart, baobabs were supposedly carried around the globe on separate landmasses, giving rise to their strange distribution. However, recent studies using morphological analysis and molecular techniques have shown that baobabs evolved much more recently – some 35–90 million years ago – and must therefore have spread across the oceans after the breakup of Gondwanaland. It is now believed that the pioneering ancestors of modern baobabs sailed away from Africa as seeds, safely stowed away in their buoyant, hard-shelled fruits; one washed up in Madagascar and the other in Western Australia, where they produced new colonies that later diverged into discrete species.

The theory of long-distance oceanic dispersal is plausible given that the hard fruits of several baobab species are, at least partly, dispersed by water. Indeed, in 2006 several Adansonia madagascariensis trees (whose fruits float exceptionally well) were discovered growing along the southern coast of Mayotte in the Comoro Islands, just 200km from Madagascar. This finding was particularly interesting as this species is mainly found in northwestern Madagascar, the region directly facing Mayotte, so the presence of the Comorian population of baobabs is very likely the result of a successful transoceanic crossing.

In Madagascar, the genus Adansonia is distributed throughout the dry to sub-arid forests and thickets of western and southern Madagascar, from Taolagnaro in the south to Antsiranana in the north. Certain areas are particularly rich in baobabs, giving rise to their nickname ‘baobab forests’. However, they are completely absent from the central highlands and the humid east coast.

Golden-trunked A. rubrostipa growing on limestone islands in northern Madagascar. It may have been coastal trees like these that long ago dropped their fruits into the ocean, which were then swept off to start new baobab colonies on distant shores. Moramba Bay. (C.G.)

Several baobab species, including Adansonia grandidieri (pictured), A. perrieri and A. za, are often found growing along watercourses and even along coastlines and mangrove forests. The fruit of these species are at least partially dispersed by water. (C.Q.)

DISPERSAL

Nowadays, young baobab trees are scarce in Madagascar’s dry forests and thickets. Researchers are still trying to understand exactly why the regeneration process appears to have faltered, but it seems likely that a combination of fire, habitat degradation, grazing by cattle and goats, competition with invasive species and the excessive collection of fruits and seeds by people could be having a severe impact on baobab reproduction.

If Madagascar’s baobabs are no longer effectively regenerating, it may also be that an important element in their reproductive cycle has disappeared. Their seeds, like those of many other tropical trees, are thought to be adapted to dispersal by animals – a process known as zoochory. The animal intermediary consumes the seeds whole, which then pass through its digestive tract unharmed before being expelled in the animal’s faeces at some distance from the parent tree. Indeed, the African Baobab is also known as the Monkey-bread Tree, because the fruits are consumed by baboons, though other mammals such as elephants also eat and disperse the seeds. Kangaroos and wallabies are known to eat the fruits of the Australian Boab, but there are no records of Malagasy baobab fruit being consumed by any native animals. The introduced Bushpig (Potamochoerus larvatus) likely consumes baobab fruit, but its method of eating almost certainly destroys the seeds. Today, Madagascar’s baobabs are mainly spread via water dispersal, though human activities may also have a role. People collect the fruits for both local consumption and for sale in urban markets around the country, as they are used to make a tangy, nutritious juice. Domestic cattle (the humped zebu, Bos indicus), may also consume the seeds as they forage along forest paths. However, just a few hundred years ago, the island’s baobabs may have been dispersed by some very different creatures.

The different stages of germination of an Adansonia grandidieri seed. (L.J.)

It is possible that the baobabs’ key animal disperser(s) disappeared during the wave of faunal extinctions that began around 2,000 years ago after humans colonized Madagascar. The island lost at least 34 species of large vertebrate, including 17 giant lemurs (nine genera in the suborder Strepsirhini), three pygmy hippos (Hippopotamus), two giant tortoises (Aldabrachelys) and eight elephant birds (Aepyornithidae). Likely candidates for the baobab disperser include large-bodied extinct lemurs in the genera Archaeolemur and Pachylemur, which occurred in southern and western dry forests until the arrival of humans, and survived until at least 1,000 years ago. Studies indicate these species were principally frugivorous.

Other possible candidates are the two extinct giant tortoises in the genus Aldabrachelys, which once occurred in large numbers in Madagascar’s dry forests. The closely related Aldabran Giant Tortoise (A. gigantea) has been shown to readily consume baobab seeds without destroying them. The seeds then show enhanced germination once they have passed through the reptile’s gut. If, as it seems, the island’s baobabs do require animals for dispersal, the long-term future of these magnificent trees may be in jeopardy.

A two-year old A. grandidieri sapling, with a distinct water-storage tuber. (L.J.)

The seeds of Malagasy and African baobab species are able to withstand extreme dehydration, tolerating around 5% humidity. It is relatively easy to sow baobab seeds, but germination can be very slow due to the hard seed coat (tegument), which is largely impermeable to water, air and light. There are several methods by which one can break the dormancy imposed by the seed coat and speed up germination. One method is to cover the seeds with simmering water and leave them to soak for 24 hours before planting them. Alternatively, one can scarify the seeds by rubbing them with sandpaper, or by soaking them in undiluted sulphuric acid for several hours. The seeds of Adansonia madagascariensis appear to require rather more severe scarification compared with the other species.

For normal, healthy development, seedlings must be planted out in a sunny position in well-drained soil and kept moist without over-watering.

A solitary Adansonia grandidieri stands above flooded rice paddies, where once lay dense, deciduous dry forest. Morondava region. (V.V.)

CONSERVATION

Madagascar’s unique biodiversity, like that of many other floral and faunal communities around the world, is increasingly threatened by the destruction and degradation of natural habitat. Over the course of the last century, the rate of habitat destruction on the island has intensified in the wake of a population explosion. The majority of the country’s 22 million people live in rural communities dependent to a large extent on natural resources for their subsistence. Traditional agricultural methods involve shifting agriculture (known as tavy or hatsake in Malagasy), in which areas of forest are cut and then burned to create new fields. These are mainly planted with rice, maize or cassava, and may be productive for two or three years. However, the loss of forest cover results in rapid erosion and leaching of the soils by rain and wind, particularly on slopes, and the fields quickly lose their fertility. After a short space of time, the cleared lands are reduced to a dry, sterile lunar landscape unsuitable for growing food crops, so the farmers must move on to another patch of forest to begin the cycle once again.

More than ever before, Madagascar’s biodiversity is in need of protection. Fortunately, the government began to recognize the value of the island’s natural ecosystems early in the 20th century, and the conservation movement was initiated in 1927 with the creation of 10 Reserves Naturelles Integrales (RNI) or Strict Nature Reserves. These RNI were established with the intention of protecting the island’s major forest ecosystems, with access granted solely for scientific purposes. Later, several RNI were converted to national parks, along with a number of new sites, in order to protect a greater variety of ecosystems and sites of great natural beauty, while allowing for the development of ecotourism within the parks.

The silver-grey trunks of these A. za stand in stark contrast with the red tilled earth, relics of the dry spiny forest that once covered this landscape. The tree on the right contains a man-made well, accessed by the ladder lying at its base. Toliara region. (V.V.)

These A. za were once surrounded by subhumid forest, but this was cleared to make way for agricultural fields. The remains of this rare forest type are now protected within Zombitse-Vohibasia National Park, which is sadly still threatened by fire and shifting agriculture. (L.J.)

Vast expanses of baobab-dominated dry forest are cleared every year by the destructive practice of shifting agriculture (hatsake). (L.J.)

Conservation activities gathered momentum during the 1990s, as the preservation of biodiversity became a top priority for the Malagasy government, but it was during the World Parks Congress in Durban, which took place in September 2003, that things really took off. The participating countries underlined the importance of the contribution of protected areas to sustainable development, environmental services and to poverty alleviation, and it was in this spirit that the Malagasy government committed to tripling the size of the island’s protected area network, from 1.7 million hectares in 2003 to 6 million hectares by 2012, amounting to at least 12% of Madagascar’s land cover.

Across the Madagascar National Parks (MNP) protected area network, there are now 47 sites, covering 2.75 million hectares, as well as the numerous new protected areas (NPAs) that are currently being created within the framework of the Durban Vision. At the time of writing, more than 30 NPAs have been created, covering 3.34 million hectares, and a further 32 are in the planning stages, which may cover 971,000 hectares.

The dry and spiny habitats of the southern and western regions are less well represented in Madagascar’s protected area network than those of the east. In 2003, only 2.3% of remaining southern spiny forest habitat was formally protected, compared with 13.3% of eastern humid forest. However, with the implementation of the Durban Vision, the proportion of dry habitats under formal protection has begun to rise. Apart from the Avenue of Baobabs near Morondava, which is now an NPA within the Menabe Antimena Protected Landscape, there are no other sites dedicated solely to the conservation of baobabs. Nevertheless, the presence of these magnificent trees is often a key element in justifying the establishment of new protected areas, such as Kirindy-Mite National Park, whose forests boast the greatest concentration of baobabs in Madagascar, including the spectacular and Endangered Adansonia grandidieri.

The species A. grandidieri, A. rubrostipa and A. za are all well represented in Andranomena Special Reserve and Kirindy Private Reserve (both of which fall within the Menabe Antimena Protected Landscape), Kirindy-Mite and Mikea national parks, and the Mangoky-Ihotry Complex.

Adansonia za is a widespread species found in many other protected areas, including those in the west mentioned above as well as Berenty Private Reserve, Andohahela National Park and Nord-Ifotaka Protected Area in the southeast, Zombitse-Vohibasia and Tsimanampesotse national parks and Ranobe-PK32 Protected Area in the southwest.

Adansonia rubrostipa also occurs in Namoroka National Park, Ranobe-PK32 Protected Area, and in the sub-arid climate of Tsimanampesotse National Park, where stunted and often wrinkled specimens are characteristic of the spiny thicket. Towards the northern edge of its range, this species is found alongside A. madagascariensis in Anjajavy Private Reserve.

Adansonia madagascariensis and A. perrieri are both found within the Ankarana and Montagne d’Ambre national parks, Montagne des Français Protected Area and Analamerana Special Reserve. The latter two sites are also home to A. suarezensis, while the dry forests of Loky-Manambato Protected Landscape contain A. perrieri. These reserves are all of particular importance for baobab conservation, because they contain two of the rarest and most threatened species in the genus. Even though A. perrieri is likely more widespread than once believed, only one of the few sites at which it is found (Ambondromifehy) contains more than a handful of individuals, and only in Montagne d’Ambre are the baobabs sheltered from fires and deforestation. However, the long-term survival of the Montagne d’Ambre population is threatened by introduced rats, which eat the seeds and prevent regeneration. Only the fruits that fall in the river have a chance of being carried away from the rats and may succeed in germinating elsewhere.

Until recently, Adansonia suarezensis was included in just one reserve, but fortunately it is now represented within several new protected areas. However, even though it is not quite as scarce as was once thought, and despite its inclusion in the protected area network, it is still rare, with a highly restricted range, and its habitat continues to be degraded and fragmented. In addition, a recent study has predicted that climate change will have serious impacts on populations of both A. suarezensis and A. perrieri in coming decades, which, coupled with the other threats to their survival, may push them to the brink of extinction in our lifetimes.

A multiple-trunked Adansonia rubrostipa in spiny forest on the Mahafaly Plateau in Tsimanampesotse National Park. (L.J.)

Adansonia suarezensis in the Montagne des Français Protected Area and on Cap Diego. (L.J.)

Despite national efforts to establish protected areas and conserve the island’s precious fauna and flora, the main forces driving biodiversity loss persist. Not only are certain species and habitats still not represented within conservation programmes, but many protected areas also continue to be threatened by fire, logging, grazing, and shifting agriculture. One way to protect the Endangered baobabs and the island’s endemic flora from extinction is to preserve their seeds in seed banks, but it is essential to conserve their natural habitat if at all possible to enable future reintroductions.

A statuesque A. za in the subhumid forest of Zombitse-Vohibasia National Park. (L.J.)

A. madagascariensis in Ankarana National Park. (L.J.)

The Avenue of Baobabs is a world famous tourist attraction containing stunning A. grandidieri in Menabe Antimena Protected Landscape. (A.M.)

USES

Madagascar’s baobabs are much less well utilized than the African species, which is exploited locally for its leaves, bark and roots, and commercially for its fruit. This is not because the Malagasy species are less useful; in fact, they have largely the same properties as the African and Australian species, but traditionally people have harvested only their fruit, seeds and bark.

The seeds of several Adansonia species are rich in oil. According to Henri Perrier de la Bâthie, A. grandidieri seeds were exported to Marseille from 1874 until the early 20th century, where they were mixed with other oily seeds for the extraction of cooking oil. However, the supply of seeds was apparently insufficient to satisfy the market, and so the trade ceased. Villagers in the Morondava region have also been known to extract the high-quality oil from the seeds for local use. However, the oil contains fatty acids that are harmful if consumed in large quantities, which must therefore be removed or neutralized before use.

The dry fruit pulp is rich in calcium, potassium, vitamin C and tartaric acid, and can be eaten or made into a tangy, refreshing juice when mixed with water and sugar. The dried pulp of the African Baobab has recently gained popularity as a ‘superfood’ among health food enthusiasts in Europe and the United States, where it is added to smoothies and cereal bars to give a nutritious boost.

A Sakalava house with a roof made with sheets of baobab wood. Near Morondava. (T.G.)

The bark of a mature baobab tree can reach a thickness of up to 15cm, and is composed of tough, longitudinal fibres, which are often used to manufacture rope (known as hafotse in Malagasy) particularly for use on zebu-drawn carts and on the traditional outrigger canoes. The spongy, water-rich wood is composed of concentric sheets of fibre and cannot be used as fuel or for construction. This is a great advantage for the trees, which are usually left standing, even in largely deforested areas. However, in times of drought, whole trees are sometimes felled so that the watery wood can be fed to cattle. Sheets of wood are harvested from dead or living trees and then dried and sold on local markets for use as thatch. Fortunately, baobabs are fantastically resilient, and though many trees show the scars from bark and wood harvesting, most recover rapidly and regrow their bark. Edible mushrooms (Volvariella volvacea) may be found growing at the foot of old or dead baobab trunks. Sadly, some baobabs near Toliara have been deliberately felled for the cultivation of this mushroom.

This A. grandidieri bears the scars of repeated bark harvesting. Near Befandriana Sud. (A.P.)

Juice made with baobab fruit pulp is tart, refreshing, and rich in calcium and vitamin C. (L.J.)

Wooden stakes hammered into a baobab’s trunk enable people to climb to the branches and harvest the ripe fruit. (A.P.)

A thin sheet of dried baobab wood, showing its loose, fibrous structure. (L.J.)

The baobabs’ remarkable ability to survive severe wounds and regenerate bark is put to use in several ways. In Madagascar’s arid south, people carve deep holes into baobab trunks in order to create water reservoirs or wells. These collect rain water during the rainy season for use during droughts, without which locals might perish. Local people also hammer a succession of stakes into the trunks of fruiting baobabs in order to climb up to the canopy to harvest the ripe fruits before they fall and spoil. Once the stakes rot away, the tree is left with rows of footholds. Large trees, especially Adansonia digitata, A. gregorii and A. grandidieri, may become hollow as they age. Such baobabs have been used in many different ways, including as storage barns, storm shelters, bars, toilets and even prisons.

Edible mushrooms (Volvariella volvacea) growing on a dead Adansonia madagascariensis tree. Ankarafantsika. (L.J.)

The entrance to a man-made well in an Adansonia za. Between Betioky and Beheloka. (A.P.)

The interior of the same well. (A.P.)

The open fruit of A. madagascariensis, said to be the least palatable of all baobab fruit. (A.P.)

A woman selling A. grandidieri fruit on the market in Toliara. (L.J.)

African Baobab (A. digitata) at the centre of a roundabout in Mahajanga, Madagascar. (L.J.)

BAOBABS IN CULTURE

Offerings laid at the base of the great sacred African Baobab (Adansonia digitata) on a small island in Moramba Bay. Sadly, this tree collapsed in 2014. (C.G.)

In light of their outlandish appearance, it is not surprising that baobabs are the subject of folklore and superstition. Numerous legends explain the baobabs’ extraordinary shape. For example, an Arab story blames the devil for the tree’s upside-down appearance, suggesting that long ago he pulled up the tree, planted its branches in the earth and left its roots up in the air. A legend of the San people of the Kalahari tells of the time when the Great Spirit was presenting each of the first peoples and animals with a different type of tree to plant. Arriving late, the hyena received the only tree that was left, the baobab. The hyena was so furious that he planted it upside down. A widespread African legend tells us that when God planted the baobab, it continued to wander around the countryside. One day, God became annoyed and decided to plant it upside down to keep it firmly rooted in one place. Another legend describes how the baobab arrived in dry habitats: God planted the baobab next to a pool, but it complained continuously about being too wet. God became so irritated with the complaints of the ungrateful tree that he pulled it out and threw it down into the dry regions of the land.

In contrast with native African peoples, Australian aborigines do not really have a concept of God, and their legends and beliefs about baobabs tend to relate to specific trees rather than the species in general. Certain Boabs are linked to stories of the ancestor creators; some are believed to embody dark powers and are used in spells, while others sport drawings and carvings of important spirits and ancestors from the Dreamtime.

Certain baobabs in the forests of southwestern Madagascar are considered to be inhabited by the sacred spirits of the ancestors, or by forest spirits such as the kokolampo. The latter may possess passers-by, particularly elderly people, who wander too close to the tree. Giant African Land Snail (Achatina fulica) shells may be arranged at the base of sacred A. grandidieri trees to indicate their special status and receive offerings of rum. Other offerings are placed at the tree in return for blessings, such as food to ensure a good harvest, money for good fortune and rum for fertility and general prayers. Some baobabs with suggestive protuberances, such as the sacred tree at Lake Andranovorinampela in Andranomena Special Reserve, are visited by couples wishing to have a baby. The forests surrounding these sacred giants are protected by the local people, who ensure that the powerful ancestor spirits are respected.

Today, the iconic image of the baobab appears in many areas of modern life in Africa, Madagascar and Australia, including company logos, administrative emblems, postage stamps, banknotes and in marketing and commercial products. Mainland Africa in particular has an ever-increasing trade in all things baobab, including cosmetics and food supplements. The trees have also been the source of inspiration for numerous artists and writers, and are the subject of many traditional proverbs.

An old A. grandidieri on the edge of a seasonal pool near Andranomena Special Reserve. It is considered sacred by the local Sakalava people. (L.J.)

Visitors to the Avenue of Baobabs can watch the sunset from atop a baobab, or even sleep up in the canopy. Morondava. (L.J.)

Current and discontinued Malagasy banknotes. (A.P.)

The following are examples of proverbs and literary quotations from around the world:

‘Wisdom is like a baobab tree; no one individual can embrace it.’

Traditional, West Africa.

‘If a sapling grows beneath a baobab, it will die a bush.’

Traditional, Central African Republic.

‘It is at the foot of a baobab that one views a baobab best.’

Traditional, Senegal.

‘Though he may sojourn long in the branches of the baobab, the partridge will never forget the nest of lowly brush where he was hatched.’

Ahmadou Kourouma (1927–2003) in Waiting for the Wild Beasts to Vote.

‘Now there were some terrible seeds on the planet that was the home of the little prince; and these were the seeds of the baobab. The soil of that planet was infested with them. A baobab is something you will never, never be able to get rid of if you attend to it too late. It spreads over the entire planet. It bores clear through it with its roots. And if the planet is too small, and the baobabs are too many, they split it in pieces . . .’

Antoine de Saint-Exupéry, 1943, The Little Prince.

A zebu carving in the bark of an Adansonia grandidieri at Tampolove, near Andavadoaka. Baobab wood and leaves are important resources for cattle in the spiny forest where food and water may be scarce during the long dry season. (L.J.)