The long walk to treedom

A tale of the African savanna

by Glenn Moncrieff, Botany Department, University of Cape Town

he road to becoming a large tree in an African savanna is fraught with danger. Every tree that stands towering over the landscape has managed to overcome almost insurmountable odds in a battle against the most formidable of opponents. Understanding the victory a tall majestic tree represents, adds a new dimension to ones appreciation. Along

ABOVE: Oblivious to thorns and height above ground, a giraffe browses in an Umbrella Thorn, Acacia tortilis. Photo: Eugene Moll. BELOW: A stately Knob Thorn, Acacia nigrescens, amongst the diminutive grass on the African savanna. The tree has been pruned to its cylindrical shape by browsing mammals. Photo: Eugene Moll.

the course of a trees journey from seedling through sapling to adult it must survive and overcome constant bombardment from a diversity of mammalian aggressors, frequent fire and competition from their seemingly humble, yet inescapable, opponent grass. The journey begins when a seed is lucky enough to escape its numerous predators and arrive at a site where the environment is just right for it to germinate. A newly established seedling stands little chance of surviving in the brutal playing fields of the African savanna. Having access to available water is crucial if a seedling is to grow and survive. However, rainfall in savanna regions is notoriously variable, which results in water shortages that kill both seedlings and larger trees especially in the more arid savannas. As well as a natural shortage of water, seedlings face the additional stress of having to compete with grasses for any that is available. Savannas are defined by the co-existence of trees and grasses, and thus a savanna tree must necessarily live side by side with grass. Although grasses may seem rather feeble opponents, their roots can monopolize access to available water and nutrients, leaving few gaps for tree seedlings to occupy. Living alongside grasses poses another threat to trees, which challenges not only seedlings, but saplings (trees shorter than 2 m) as well. When grasses grow in environments that

experience seasonal drought, as do savannas, the inevitable fires that occur threaten the survival of trees. Fires are regular disturbances in savannas that become more frequent and of higher intensity the more grass there is. Trees with canopies below flame height inevitably succumb to fire. This means that once a seedling establishes itself, it enters a race to grow beyond the flames reach before the next fire passes through. In wetter, fertile savannas where grass growth is rapid, fires can occur as frequently as every second year, or even annually, and their flames can scorch canopies up to 3 m. Consequently, a tree may have to grow between two and three metres in just a year or so in order to survive, which means that only the fittest and the fastest manage to escape from fires. Yet it is not all over for those unlucky, or unfit, enough not to escape fire. Savanna trees are not hapless victims of the external forces that threaten to eliminate them. Many species have the remarkable ability to re-sprout from their roots and underground storage organs after the loss of stems, shoots and leaves to fire. The below-ground component of saplings can be vast, allowing trees many attempts at growing beyond the reach of fire and out of the fire trap. This spectacular battle between trees latent giants held down by the chains of fire and grass the diminutive but nonetheless potent adversary has been likened to the

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imprisonment of Jonathan Swifts famous literary figure Gulliver by a mob of tiny Lilliputians. Savanna tree saplings have thus come to be known colloquially as Gullivers. Despite this apparent fight for supremacy, savanna trees are also dependent on grass and fire for their continued existence. In the absence of fire, rather than becoming dominated by savanna trees, the environment is often invaded by fire-sensitive forest and thicket tree species. These invaders can exclude both savanna trees and grasses and, through continued thickening and expansion, can result in the phenomenon known as bush encroachment. So there is no option for a tree other than to face the challenges posed by grass competition and fire, while at the same time confronting another hurdle: herbivores. Large herbivores are a spectacular feature of African savannas and they occur in a variety of shapes and sizes, each with its own particular strategy for meeting its dietary needs. Some, such as impala and duiker, remove individual shoots and leaves, selecting only the plant parts they desire with great dexterity. Others, like giraffe and elephant, are less delicate as they need to consume vast amounts of forage to meet their nutritional requirements. This constant attrition results in trees losing large quantities of photosynthetic tissue and valuable nutrients to browsers, and acquiring densely branched hedge-like shapes as if pruned by a clumsy gardener. The upshot is that growth is slowed and trees must spend more time within the reach of fire, increasing their chances of falling victim to it. However, as anyone who has ever caressed an acacia will know, savanna trees have evolved ingenious and menacing defences to protect themselves against browsers. Long, sharp thorns protrude from the branches of many; piercing the mouth of any herbivore that approaches without caution. Other trees defend themselves chemically rather than mechanically, with toxic or unpalatable compounds in their leaves. Despite these various and many protections, most trees are not completely successful at preventing browsing, but they do however slow down and reduce herbivory sufficiently to survive. Some are successful enough to survive and grow beyond the grasp of both fire and even the tallest of herbivores. Having made the remarkable journey from seedling to sapling (perhaps many times over) and on to adult, a savanna tree is now finally safe and secure from most of the perils it faced when it was smaller. Grass no longer poses a threat to its continued survival, it has access to deeper, more reliable water sources and it is almost invulnerable to even the hottest fires. Moreover, it has grown too large to be reached by most browsers and it can now spread its branches and bathe its leaves

in sunlight. Despite this apparent fortification, even the largest tree cannot resist an assault from a fully grown African Elephant. With enough effort almost any stem can be snapped or tree toppled over. Failing that, sufficient bark could be stripped off the stem by tusks to ultimately stress the tree beyond recovery. Its then back to the beginning to attempt the journey once more. The tremendous achievement that a fully grown, towering tree rising above the savanna represents should not go unappreciated. Its existence is a most improbable event, and it deserves tremendous respect.

ABOVE: Savanna trees have evolved ingenious and menacing defences to protect themselves from browsers as this Black Monkey Thorn, Acacia burkei, demonstrates. Photo: Eugene Moll. LEFT: A Marula tree, Scelerocarya birrea, freshly ring-barked by an elephant. Photo: Eugene Moll. BELOW: A Marula tree, Scelerocarya birrea, is consumed by an intense grass fire in Hluhluwe-IMfolozi Game Reserve. Photo: Julia Wakeling. BOTTOM: Even the largest tree cannot resist an assault from a fully grown African Elephant. Photo: Glenn Moncrieff.

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