Why do grasslands have no trees?

by Julia Wakeling, with William Bond and Michael Cramer, Botany Department, University of Cape Town

T
he Highveld grasslands cover a large area south of
Johannesburg. If you have ever driven through them you may
have been amazed at the vast openness – not a single tree in
sight – until you come across the ubiquitous gum tree at a lay-by. Trees
manage to grow in towns and plantations in grassland areas, so they
can obviously survive the cool climate. Why then do the grasslands
have no trees? This was the topic of my recent Masters degree in
Botany. In my travels I spoke to many people and everyone had their
own theory, but not one that could really answer my question. Some
were correct locally, for instance, many trees do not survive in soils
that become seasonally waterlogged, and true, many of the flatter
parts of the Highveld have soils like this, but certainly not the whole
Grassland Biome.

A frosty answer
‘Ah, it’s too cold and trees are killed by frost’, is probably the most
common explanation. As I said before, trees do survive the winter, and
grow in towns, plantations, along roadsides and in forest patches high
up in Drakensberg valleys. And what about savanna trees? I wanted
to explore how frost would affect the types of trees that normally
grow within the grass layer in savannas. To do this I planted seedlings
of acacia thorn trees in grassland and savanna areas and monitored
their survival through the frosty winter of 2007. Near Piet Retief and
Volksrust almost all the seedlings survived the winter frosts, whereas
in a cooler area near Ermelo, only a few seedlings survived, but if those
surviving seedlings could become adults, the area would turn into a
wooded savanna. A few species, such as Ouhout Leucosidea sericea,
do grow on hillsides in grassland areas – highlighting yet again that
woody species can cope with the climate – but for the most part the
Highveld grasslands are wide open grassy expanses.
Cut down in their prime
Perhaps people cut them down? This is a popular idea for explaining the
tree-less nature of grasslands all over the world. But pollen cores and
soil carbon analyses are revealing that the Highveld grasslands have
been present for over 10 000 years (as have many other grasslands in
the world), much longer than the influence of modern humans. There
TOP: An exquisite grassland with indigenous trees growing in the kloof and eucalypt plantations
beyond. ABOVE: A Paperbark Thorn, Acacia sieberiana, re-sprouting after a severe frosting.
Photos: Julia Wakeling.
are certainly tracts of land that have been cleared of trees within the
last few hundred to 1000 years, but they are a small fraction of the
entire Grassland Biome from Johannesburg to the Eastern Cape and
Piet Retief to Bloemfontein!
A fiery debate
Fire, you think? I’m afraid not. Some grassland areas burn every year,
yet others burn perhaps only every 10 to 15 years. This is also true for
savannas, which are grasslands with an abundance of successful trees.
But how do savanna trees cope with fire? Seedlings are easily burnt
to the ground, but as the roots are not killed the plant can re-sprout.
Seedlings grow into saplings, which, if they grow fast before the next
fire, may grow tall enough to get their leaves above the height of the
flames. If successful, they have ‘escaped’ into adulthood. If still too
small, a sapling will burn to the ground and re-sprout from its roots
again. A sapling can survive for decades by repeated re-sprouting, but
if it is to reproduce successfully it needs to grow above flame height.
Propagation is after all a goal of all living organisms. For the little tree,
all depends on growing fast enough between fires. So what influences
this growth rate?

24 VELD&FLORA | MARCH 2010


Is it in the soil?
Soil nutrients seemed a likely candidate. The
Highveld grasslands are generally on old soils
that have been leached of their nutrients
over time, while savanna soils on the same
geological substrates are richer in nutrients.
In an experiment, I measured growth rates of
two acacia species to see if this relative lack
of nutrients in grassland areas would cause
trees to grow too slowly to ever escape the
frequent fires. I collected soils from relatively
‘nutrient poor’ Highveld grasslands (near
Ermelo, Volksrust and Piet Retief ), and the
comparatively ‘nutrient rich’ savannas (near
Vryheid, Pongola and Hluhluwe-iMfolozi
Game Reserve), and grew seedlings in these
soils. And indeed the Lowveld soils produced
faster growing seedlings than the Highveld
soils. But are the differences large enough
to explain the absence of trees in grasslands
and their presence in savannas? We know
that a tree growing in savanna soils can grow
to a height above the flames of a fire, in the
time between two fires. In this experiment,
seedlings grown in grassland soils grew only
slightly slower than those in savanna soils, but
they still grew fast enough to escape fires.
Ok, so what is it?
We are running out of explanations. Why do
savanna trees, which can survive fire, not
grow in these vast grassland areas? I should
point out that the Highveld grasslands are
far from unique. There are similar grasslands
throughout Africa: the Eastern Highlands
in Zimbabwe, the Nyika Plateau in Zambia
and Malawi, the Aberdares in Kenya; and the
rest of the world: Madagascar, India, the tall-
grass Prairies of North America, the Campos
grasslands of South America and the Balds of
South East Australia. In all these grasslands fire
is important and fire-tolerant savanna trees
occur nearby. Often there are forest patches
within the grasslands, or trees growing in
towns, highlighting that it is not too cold
for trees to exist. And yet there are no clear
answers as to why trees don’t grow there.
Keeping cool
A major aspect of the grasslands is that they
are at greater altitudes or latitudes, and thus
cooler, than adjacent savannas that do have
trees. Most living organisms grow slower
when it is cooler. I wanted to know how much
slower savanna trees would grow in grassland
climates and what this would mean for a
savanna tree trying to outgrow fires. I planted
seedlings of six acacia species at sites in cool
grassland areas and warm savanna areas.
When the experiment began, these seedlings
were no larger than a coke bottle. After a
summer of growth, seedlings at cooler sites
were about the height of a spade, and those in
warmer areas were double that, some of them
large enough to provide a shady spot for
lunch. In comparison to those in warm areas,
the growth of seedlings in cool areas was so
slow that they would not be able to grow
above the height of flames between two fires.
It is noteworthy that seedlings grew at all the
sites. However, in the presence of fire, those
that grew too slowly would be eliminated.
In summary then
Firstly, frost alone does not prevent trees from
growing in the Highveld grasslands. Indeed,
many of our grassland areas are frost-free.
Secondly, low nutrient supply slows sapling
growth as is true for the relatively nutrient-
poor grassland soils. But by far the most
important limitation on seedling growth in
my study was cool temperatures associated
with high altitudes. This slow growth in
combination with frequent fire prevents trees
from growing in the Highveld grasslands.
Implications for the future
As temperatures are increasing with climate
change, savanna trees will be able to grow
fast enough to escape fire in grassland areas
where they could not before. A reduction in
fire would also lead to tree invasions in areas
where trees grow too slowly to escape under
frequent fire regimes.
You may be thinking that I want savanna
trees to grow in the Highveld grasslands.
Quite the contrary, the grasslands are a
diverse ecosystem filled with beautiful
grasses and wildflowers. This research was
aimed at better understanding savanna and
grassland ecosystems so we know how to
manage them. Many grassland areas are used
for plantations, agriculture and cattle farming,
and the small areas of pristine grassland that
remain are threatened. What is left needs to
be conserved.
When you next drive through these vast
grasslands I encourage you to stop and
appreciate them. Enjoy the patterns of the
wind in the grass and examine the minute
intricacy of grass flowers and the tiny wild
flowers hiding between them. Ponder the
travails of a tree seedling entering this grassy
world and revel in the natural beauty of a wide
open space without a single tree.
Acknowledgements
This work was made possible by the generous provision
of land, labour, time, water, hospitality and enthusiasm
by a number of land-owners and managers, including
Robin and Sue Kirkwood, Emile Schmidt, Ian and Carol
Goss, Henry and Ria Malan, Katy and Stephan Heitland,
Clayton and Debbie Comins, Graham and Janine Ford,
Nigel, Sue, Jen and Tracy Anderson, Mark and Ruth
Armitage, Andrew and Amanda Shand, Mark, Evette,
Michael and Andrew Hitchins, the Zululand Tree Project
staff, and Mr Kosy Dongo, Lesego Bodibe, Samuel Zulu
and Jan Theron of the Agricultural Research Council. The
project was funded by the Andrew Mellon and National
Research Foundations.
ABOVE: Five month old acacia seedlings planted in warm
conditions near St Lucia, were much larger than the same type
of seedlings growing in cooler conditions near Ermelo.
BELOW: Five month old acacia seedlings planted in cool
conditions near Ermelo were smaller than similar ones planted
in warmer conditions in St Lucia. Photos: Julia Wakeling.
MARCH 2010 | VELD&FLORA 25