Exploding the myth
Fire and frost keep trees from establishing in the grasslands of the
Highveld is an oft-perpetuated myth. Back in the late 1950s, an
exceptional South African geomorphologist, Lester King, published his
book The South African Scenery in which he questioned many existing
geological paradigms, and Ken Tinley put forward his view of why the
Highveld was treeless. Tinley’s view was certainly very different from
what I had been taught, and what is still taught today – that frost and
fire keep the Highveld grasslands treeless. What rubbish!
As I remember it, the reason for King’s book was that he had submitted
a paper on the age of South African mountains and the peer-reviewed
journals (in the Northern Hemisphere) all refused to publish his work.
At that time geologists believed that there were no mountains on Earth
older than 10 million years! Yet King was telling the world that the Cape
Fold Mountains were older than 70 million years old and that we had
some land-surfaces that hark back to the break-up of Gondwana (the
Highveld). Today his pioneering work has been vindicated.
Thankfully today’s scientists have become much more open-minded
since those oppressive and restrictive days of the mid-1900s. Recently
a University of Cape Town group researched the reasons for no trees
in the Highveld grasslands (see the article ‘Why do grasslands have no
trees’ by Julia Wakeling, William Bond and Michael Cramer in the March
2009 issue of Veld & Flora). Their conclusions were inconclusive, but
they certainly demonstrated that frost is not a totally limiting factor,
nor were fires a conclusive reason for no trees but that the lack of trees
could be due to a combination of factors; one of which may be plant
available nutrients. Thus the generally held view that grasses dominate
the Highveld because it is too cold for trees and that fierce annual fires
kill woody growth has been debunked.
Ok then what are my reasons for the lack of trees? Firstly we have
to understand that most of the Highveld is a Gondwana land-surface
70  million years old. Soils have been leached of most of their important
plant nutrients. The relative quantities of aluminium (one of the most
common elements in the Earth’s crust) are very high, and this could be
a factor in restricting root exploitation, Tinley concluded. Tinley had
observed that alien trees had no problem growing in the Highveld
and were in fact invasive (Australian wattles and gums, and certain
species of pine – all of which evolved in extremely toxic aluminium soil
environments, so were pre-conditioned to survive in the Highveld).
Tinley also observed that some indigenous trees could grow well, but
only in certain places where the sub-soil drainage had been affected by,
for example, incipient dongas and roads. Thus he debunked the frost
and fire theory way back in the 1960s.
Plants get most of their nutrients from the top few centimetres
of soil where almost all the feeding action takes place. Grasses are
basically shallow rooted and trees (be they seedlings, saplings or
mature individuals) compete with grasses for 99% of their nutrients
in the organic-rich top soil. However, trees, unlike grasses, send down
anchor and water-gathering roots much deeper – into that toxic soil
horizon. Thus in relatively undisturbed grasslands the grass sward is
such an extremely competitive environment that it is difficult if not
impossible for any seedling to survive. Even grass seedlings have to
wait for some episodic event to cause a small gap. Another fact is that
unlike savannas, where there are all kinds of seeds of woody plants in
the soil waiting for an opportunity to germinate and establish, in the
Highveld there is no such soil stored seed-bank. So even if the grass
sward integrity is breached there are no tree seeds waiting to spring
into action. Today not only has the grass sward been impacted but local
drainage patterns have changed, thus the whole grass competitive top
soil and underlying toxic soil integrity has been impacted paving the
way for more tree invasion – including indigenous species.
ABOVE: Grassland in the Drakensberg foothils with a lone Cabbage Tree (Cussonia paniculata).
Photo: Eugene Moll.
If Tinley is right then one would expect to see changes in the
establishment pattern of woody plants in the Highveld grasslands
today. I have been travelling through this area since the 1960s and l see
many more indigenous trees in these grasslands. This has not surprised
me as in my ecological teaching I have always given students the ‘Tinley
view’ in opposition to the ‘classic view, now disproved’. The current rate
of invasion is more rapid, and that fits with the Tinley view as most of
the Highveld has now been disturbed and because of the planting of
indigenous trees along roads and in gardens, the seed source is not as
remote as it once was. Thus within a few years native tree species will be
a much more common sight in these Highveld grasslands, and the next
generation will not see these once vast treeless Highveld grasslands.
Just as my generation have been denied the opportunity of seeing the
vast herds of grazing antelope that one roamed these vast grasslands.
Eugene Moll, Cape Town
Julia Wakeland replies
It is fantastic that this article has stimulated this and many other
responses to one of South Africa’s most obvious yet unexplained
ecological patterns. Tinley, Feely, Acocks and others have suggested
reasons to explain the lack of trees in grasslands, but none of them
are sufficiently universal to explain the biome as a whole. For example,
adverse soil conditions such as seasonal water-logging or aluminum
toxicity may prevent trees from growing in some areas, but there are
also grasslands without seasonal water-logging or high aluminum
levels. There is scarce experimental work tackling this question, which
is unsurprising given that fieldwork over such large areas is logistically
overwhelming (speaking from experience). However, we delight in
experimental work as it provides evidence that may force us to change
our minds.
In our experiment we quantified tree growth rates in savanna and
grassland. Firstly, trees did grow in grassland soils. Nutrient differences
meant that trees grew slightly slower in grassland soils than savanna
soils. However, temperature differences between the two biomes had
a much larger effect on tree growth. In combination with frequent fires
the slow growth of trees in grassland temperatures would prevent trees
from establishing as adults. In this experiment we were interested in
the sapling to adult life stage. With our research group (the Zululand
Tree Project) we have demonstrated that seedling growth is adversely
affected by grass competition in savannas, and are currently doing
experimental work to explore the competition of grasses in grasslands.
Our focus was on savanna trees that survive grass competition and
frequent fires in savannas. There are many fire-sensitive trees such as
White Stinkwood (Celtis africana) that typically grow in forests, and
would not be expected to grow in a grassy environment. The real puzzle
is why the tree-grass mixture of savannas becomes grassland.
We feel like we are slowly homing in on the answers to this amazing
phenomenon of a savanna tree-line well below the altitude of an
alpine tree-line, but are not there yet. If you have any brilliant ideas
that we haven’t thought of yet we would love to hear them! If you’d
like to read more, a brilliant starting point is Tim O’Connor and George
Bredencamp’s ‘Grassland’ chapter in the book Vegetation of Southern
Africa, edited by R.M. Cowling, D.M. Richardson and S.M. Pierce.
MARCH 2011 | VELD&FLORA 45