Journal of Tropical Ecology (2009) 25:31–40. Copyright © 2008 Cambridge University Press
doi:10.1017/S0266467408005683 Printed in the United Kingdom
The savanna tree Acacia polyacantha facilitates the establishment
of riparian forests in Serengeti National Park, Tanzania
Gregory J. Sharam
*,1
, A. R. E. Sinclair
*
, Roy Turkington
*,
† and Aerin L. Jacob
*
*
Biodiversity Research Centre, University of British Columbia, 6270 University Blvd., Vancouver, BC, Canada, V6T 1Z4
† Department of Botany, University of British Columbia, 3529-6270 University Blvd., Vancouver, BC, Canada, V6T 1Z4
(Accepted 10 November 2008)
Abstract: Forests are being converted to grasslands and croplands across Africa and natural regeneration of forests
is typically poor. In Serengeti National Park, Tanzania, the savanna tree species Acacia polyacantha established in
riparian grasslands and forest trees subsequently established within these stands. We examined the conditions for
establishment of: (1) A. polyacantha and (2) riparian (non-Acacia) forests. Fire was excluded from three grassland areas
for 5 y allowing A. polyacantha to establish during 1999 when dry-season rainfall was high. The seedlings of forest
tree species did not establish in grasslands, but were found in large A. polyacantha stands (> 0.3 ha) with reduced
grass cover (< 10%), higher cover of herbs (> 80%) and thorny shrubs (> 90%). Seeding survival was high in large
stands (0.87 y
-1
), but declined in artificial canopy gaps due to the ingrowth of grasses (0.21 y
-1
) and subsequent fires
(0.07 y
-1
). Shrub removal also reduced seedling survival (0.46 y
-1
) due to browsing by antelope. We propose that: (1)
A. polyacantha establishes in pulses perhaps as infrequently as twice per century, and (2) riparian forests in Serengeti
have established via facilitation under larger stands where shade excludes grass, and therefore fires and thorny shrubs
exclude browsers.
Key Words: Acacia, browse, elephant, fire, forest, impala, recruitment, riparian, seedling
INTRODUCTION
Many tropical areas can support alternative vegetation
communities: open grasslands, mixtures of grasses
and savanna trees, or closed-canopy forests. The
establishment of forest trees into African grasslands
is typically poor, despite the removal of fire, leading
to ecological and conservation concerns as forests are
removed and not replaced (Chapman & Chapman 2003).
Kellman & Miyanishi (1982) proposed that savanna trees
first establish in grasslands and subsequently facilitate
the establishment of forest trees. In Serengeti National
Park, Tanzania, where establishment of forest trees is
effectively limited in grasslands (Sharam et al. 2006) we
observed stands of Acacia polyacantha Willd., a savanna
tree species, establishing as stands in grasslands adjacent
to rivers during 1978. Today, regeneration of forest trees
is associated with some of these stands. In this paper, we
examine the conditions under which stands of savanna
trees establish in grasslands and subsequently facilitate
the establishment of forests.
1
Corresponding author. Email: sharam@zoology.ubc.ca
The establishment of forest trees into African grasslands
is poor, due to repeated fires (Sharam et al. 2006), com-
petition with grasses (Chapman et al. 1999, Sharam et al.
2006), low soil fertility, poor seed dispersal (Chapman &
Chapman 1999), seed predation by rodents (Duncan &
Duncan 2000) and browsing by antelope (Sharam et al.
2006). In Serengeti, forest stands occur in riparian areas,
composed of common species from large forests elsewhere
in Africa. Establishment of these forests in Serengeti is
primarily limited in grassland by fires, browsing and
competition with grasses (Sharam et al. 2006).
In contrast, savanna tree species such as Acacia will
readily establish in grasslands when conditions are
correct – when fires (Sinclair et al. 2007) and/or browsers
(Prins & van der Jeugd 1993) are removed or dry-season
rainfall is higher than average (Higgins et al. 2000). How-
ever, the relative importance of these variables depends
on the site. In recent history, Acacia trees have undergone
two pulses of natural establishment in Serengeti. The first
occurred from 1890–1930 when the epizootic rinderpest
removed cattle and other ungulates, and thus people
and fires, from the ecosystem for ∼30 y (Sinclair et al.
2007). The second occurred in the late 1970s when a