IUBS Unesco ~h

RESPONSES OF SAVANNAS TO
STRESS AND DISTURBANCE
A Proposal for a collaborative Programme of Research

edited by
P. Frost, E. Medina, J.-C. Menaut,
O. Solbrig, M. Swift and B. Walker

Report of a W.orkshop orglanked in QolQbaratian

with The .Comrnissiop o f ~ u ~- h p
- . .- " .
, . .. : "., e aCammunitieq
*<. . 1
n
,:
(CEC),
. -.:-
;: A,: - ..
< ...-.F. , -.&... ?' - -*
<* 2
 RESPONSES OF SAVANNAS TO STRESS AND DISTURBANCE

A Proposa1 f o r a Collaborative Programme o f Research

Report o f a Meeting o f an IUBS Working Group

on
Decade o f the Tropics Programme/Tropical Savanna Ecosystems

The UNESCO Man and Biosphere (WB) Programme,

The A f r i c a n Biosciences Network (ABN),
The European Economics Commission (EEC) , and
The S c i e n t i f i c Council o f Zimbabwe (SCZ)

9 - 13 December, 1985
Harare, Zimbabwe

Edi t e d by

Peter F r o s t
Jean-Cl aude Menaut
B r i an Wal k e r
Ernesto Medi na
O t t o T. S o l b r i g
Michael Swi f t

SPECIAL ISSUE - 10

BIOLOGY INTERNATIONAL

The I n t e r n a t i o n a l Union o f B i 01ogical Sciences

News Magazine
 TABLE OF CONTENTS

SECTION 1: INTRODUCTION AND OMECTIVES 1

OVERVIEW 1

DEFINITIONS
Savanna
Stress
D i sturbance
S t a b i 1it y and Resi 1ience

RATIONALE 4

APPROACH AND KEY QUESTIONS 6

SECTION II:THE DETERMINANTS OF SAVANNAS 7

INTRODUCTION 7

SOIL MOISTURE DYNAMICS

Rainfall
Soi 1 s

SOIL NUTRIENT DYNAMICS

Soils
Organi c matter dynamics

FIRE AND HERBIVORY

F ir e
Herbi vory

INTERACTIONS
E f f e c t s o f vegetation on water and n u t r i e n t dynamics
E f f e c t s o f animals on water and n u t r i e n t dynamics
Soi1 moisture and t h e tree:grass e q u i l ibrium
Species composition and coexistence
Phenol ogy and coexistence
P l an t production
P l a n t qua1 it y
Vegetation dynamics

HUMAN INFLUENCES 36
SECTION III:HYPOTHESES ABOUT THE RESPONSES OF SAVANNAS
TO STRESS AND DISTURBANCE

INTRODUCTION

FUNCTIONAL CLASSIFICATION OF SAVANNAS

HYPOTHESES

SECTION I V : PROPOSED PROGRAMME

0 M ECTIVES

PROCEDURE
1. Improving communication between savanna researchers
2. Promotion o f short term c o l 1 aborative p r o j e c t s
3. I n t e r c o n t i n e n t a l compari sons
4. Incorporation o f research resul t s i n t o management

ORGANIZATION

FUTURE ACTIVITIES

REFERENCES

APPENDIX: ADDRESSES OF WORKING GROUP PARTICIPANTS

 RESPONSES OF SAVANNAS TO STRESS AND DISTURBANCE:
A PROPOSAL FOR A COLLABORATIVE PROGRAMME OF RESEARCH

SECTION
- - - 1: INTRODUCTION -
AND OBJECTIVES

OVERV 1EW

T h i s document represents a proposa1 f o r a programme o f c o l l a b o r a t i v e

research on t r o p i c a l savanna ecosystems. Our concern stems from t h e
c u r r e n t t r e n d o f degradation i n savannas around t h e world, i n v o l v i n g
changes i n composition and p r o d u c t i v i t y t h a t a r e adversely a f f e c t i ng
t h e c a p a c i t y o f these systems t o support humans and o t h e r oryanisms.
I n o r d e r t o a r r e s t o r reverse these changes we need t o improve Our
u n d e r s t a n d i n g o f savanna d y n a m i c s u n d e r p r e v a i l i n g and p r o j e c t e d
p a t t e r n s o f l a n d use. Therefore, t h e o b j e c t i v e o f t h i s programme i s :

To devel op a p r e d i c t i ve unders t a n d i ng --
- o f the
i n whTch savannas r e s p o n d & n a t u r a l
ways ----
and
- man-made s t r e s s e-
s -
and d i s t ! r r b a n c e s .

T h i s can b e s t be achieved t h r o u g h a c o m p a r a t i v e , in t e r c o n t i n e n t a l
a n a l y s i s o f some s e l e c t e d aspects o f t r o p i c a l savannas i n v o l v i n g a
d i v e r s i t y o f research inputs.

Over t h e p a s t few y e a r s t h e r e have been s e v e r a l s y n t h e s e s o f t h e

r e s u l t s o f r e s e a r c h on savannas i n d i f f e r e n t p a r t s o f t h e w o r l d
( H i l l s and Randall, 1968; Bourl i k e and Hadley, 1970; UNESCO, 1979;
Wal ker, 1979; Huntley and Wal k e r , 1982; Bourl i k e , 1983; Sarmiento,
1984). A l t h o u g h t h e s e a u t h o r s do n o t a g r e e on a l 1 p o i n t s , t h e
s y n t h e s e s p r o v i d e a f o u n d a t i o n on w h i c h t o b u i l d. Some o f t h e
disagreements stem from d i f f e r e n t concepts as t o what c o n s t i t u t e s a
s a v a n n a , o t h e r s f r o m t h e d i f f e r e n t a p p r o a c h e s and methods used i n
these separate research programmes. An i n t e r n a t i o n a l c o l 1a b o r a t i v e
research programme i n v o l v i n g c a r e f u l l y p l anned comparative s t u d i e s
w i t h a common a i m and a p p r o a c h o f f e r s t h e b e s t o p p o r t u n i t y f o r
r e s o l v i ng these conceptual d i f f e r e n c e s and g a i n i ng t h e i n s i g h t s t h a t
a r e so u r g e n t l y needed as a b a s i s f o r t h e r a t i o n a l u s e and
management o f savannas.

W h i l e t h e s c i e n t i f i c r e s u l t s a r e i m p o r t a n t i n t h e i r own r i g h t , i t i s
v i t a l t h a t t h e m a j o r i n s i g h t s and f i n d i n g s o f t h e programme should
u l t i m a t e l y b e a p p l i e d t o t h e p r o b l e m s o f savanna u t i l i z a t i o n and
management. It i s n o t enough t o suppose t h a t t h e f i n d i n g s pub1 i s h e d
i n t h e s c i e n t i f i c l i t e r a t u r e w i l l b e p i c k e d u p and a p p l i e d b y
d e c i s i o n makers and managers, o r t h a t they w i l l be u s e f u l t o them i n
t h a t form. A more a c t i v e extension p o l i c y i s t h e r e f o r e envisaged i n
which larid managers and extension o f f i c e r s w i l l be i n v o l v e d i n t h e
programme a t an e a r l y stage, b o t h t o h e l p evaluate t h e s i g n i f i c a n c e
o f r e s u l t s as t h e y appear, and t o a s s i s t i n d e v e l o p i n g s t r a t e g i e s
f o r t r a n s l a t i ng r e s e a r c h r e s u l t s and e i n e r g i ng c o n c e p t s in t o
management d e c i s i o n s and a c t i o n s i n t h e f i e l d .

DEFINITIONS

Savanna

F o r t h e p u r p o s e s o f t h i s programme, c o r e savannas w i l l be d e f i n e d
v e r y b r o a d l y t o i n c l ude a l 1 t h o s e t r o p i c a l and soine n e a r - t r o p i c a l
ecosystems c h a r a c t e r i z e d by a c o n t i nuous herbaceous cover consi s t i ng

+
m o s t l y o f h e l i o p h i l o u s C4 g r a s s e s and sedges t h a t show c l e a r
s e a s o n a l i t r e l a t e d t o water stress. Woody species (shrubs, trees,
palms occur b u t s e l dom form a continuous cover p a r a l l e l l i n g t h a t o f
t h e g r a s s y 1 ayer. Mary ina1 savanna s y s t e m s i n w h i c h e i t h e r one o f
these two v e g e t a t i o n components has an i n s i g n i f i c a n t e f f e c t can be
in c l uded f o r t h e i r c o m p a r a t i v e v a l ue. Savannas encompassed by Our
d e f i n i t i o n c o v e r e x t e n s i v e a r e a s o f S o u t h America, A f r i c a and
A u s t r a l i a , and a l so occur i n Central America and I n d i a (Figure 1).

Figure -
1. World d i s t r i b u t i o n o f t r o p i c a l savannas ( f r o m Lamotte
and Hadley (1984) a f t e r B o u r l i k r e (1983))
Stress
I n t h i s d o c u m e n t we u s e t h e w o r d " s t r e s s " i n i t s a c c e p t e d
p h y s i o l o g i c a 1 sense t o d e s c r i be a c o n s t r a i n i n g e n v i r o n m e n t a 1
i n f l u e n c e t h a t r e s t r i c t s t h e p r o d u c t i v i t y and e f f i c i e n c y o f an
i n d i v i d u a l and, by extension, t h e ecosystem. Such stresses usual ly
operate when an environmental variable, such as temperature, l i g h t ,
water, n u t r i e n t s o r defo lia t i on, deviates marked ly f rom it s norma 1
range o f v a l u e s i n t h e system. S t r e s s i s seldom accompanied by
m o r t a l i t y . As s t r e s s becomes more severe i t may cause a disturbance.

D i sturbance
As used here, a ' d i s t u r b a n c e " i s a change i n t h e s t r u c t u r e o f a
system w h i c h u s u a l l y a f f e c t s it s f u n c t i o n i n g . Disturbances a r e
o f t e n caused by a p e r i o d i c events, such as f.loods, p a r t i c u l a r l y
severe storms, p r o l o n g e d d r o u g h t s , e x c e s s i v e h e r b i v o r y , o r
a c t i v i t i e s such as b u s h - c l e a r i n g and c u l t i v a t i o n . Disturbance may
resu lt f r o a one, b u t more o f t e n a combination o r sequence o f extreme
v a l u e s o f e n v i r o n m e n t a l v a r i a b l e s . These may o r may n o t cause
m o r t a l i t y o f i n d i v i d u a l s . O p p o r t u n i t i e s a r e c r e a t e d f o r new
i n d i v i d u a l s o f t h e same o r d i f f e r e n t species t o become established.
Increasing ly, d i sturbances are t h e reçu lt o f human a c t i v i t i e s .

Stabi lit y and Resi l i e n c e

A s t a b l e system i s one i n which those variables d e f i n i n g the s t a t e

o f t h e system (eg. s p e c i e s c o m p o s i t i o n , r e l a t i v e abundances,
biomass, o r p r o d u c t i o n ) change l i t t l e i n response t o o u t s i d e
p r e s s u r e s such as drought, f i r e o r grazing. I f d i s t u r b e d , t h e y
r e t u r n r a p i d l y t o t h e i r o r i g i n a l v a l u e s (Walker, 1980). A s t a b l e
system t h e r e f o r e shows l i t t l e v a r i a b i l i t y t h r o u g h t i m e i n t h e
a m o u n t s o f i t s s t a t e v a r i a b l e s . The d i s t i n c t i o n b e t w e e n
composi t i o n a l s t a b i lit y ( r e l a t i v e constancy o f species composition
and abundance) and f u n c t i o n a l s t a b i l i t y ( r e l a t i v e constancy o f the
processes m a i n t a i n i n g p r i m a r y p r o d u c t i o n ) needs t o be emphasized
since the two can be negatively c o r r e l a t e d (McNaughton, 1977).

A r e s i l i e n t system, on the other hand, is usual ly n o t s t a b l e and t h e

v a l u e s o f it s s t a t e v a r i a b l e s o f t e n change c o n s i d e r a b l y when
subjected t o outside pressure (Walker, 1980). More importantly, t h e
parameters o f t h e system which i n f l u e n c e i t s dynamics a l s o change,
thereby redef i n i ng t h e boundaries w i t h i n which the system remai ns
a t t r a c t e d t o i t s equi l i b r i u m point. As a r e s u l t , f u t u r e disturbances
o f the same type are more easi ly accomodated (Walker, e t al., 1981).
I n contrast, i n a s t a b l e system, the parameters o f the system do n o t
change w i t h d i sturbance.
I n a l 1 o f these d e f i n i t i o n s the question o f scale arises. Periodic
s t r e s s , imposed a t r e g u l a r i n t e r v a l s o v e r a long p e r i o d o f ti me,
eventually assumes the c h a r a c t e r i s t i c s o f a continuous phenomenon,
w h i l e a c o n t i n u o u s s t r e s s , seen i n t h e v e r y s h o r t t e r m and a t a
l o c a l level, may behave very much as a periodic phenomenon. Whether
o r n o t a system i s perceived t o be i n equilibriurn o f t e n depends on
t h e s p a t i a l and t e m p o r a l s c a l e o f observation. F o r example,
c o n s i d e r a b l e s h o r t - t e r m f l u c t u a t i o n s i n species c o m p o s i t i o n o r
production can be observed a t a l o c a l sca le, g i v i n g the impression
o f i n s t a b i l i t y , b u t when these are viewed over a longer t i m e period,
as p a r t o f a larger landscape, they simply appear t o be v a r i a t i o n s
around sone mean value; t h e system e x h i b i t s dynamic e q u i l i b r i u m .
Simi l a r l y , changes which are regarded as i r r e v e r s i b l e a t one t i m e
scale may e x h i b i t slow recovery over longer periods.

The s p a t i a l e x t e n t o f d i f f e r e n t phenomena i s a l s o i m p o r t a n t . A
h e r b i v o r e ' s b i t e i s v e r y l o c a l i z e d , a f i r e may cover hundreds o f
hectares, whi l e drought i s experienced a i a regional l e v e l o r above.
Moreover, a t a l o c a l scale, i t i s o n l y p o s s i b l e t o d e t e c t t h e
response o f one o r a few individuals. As the s p a t i a l scale expands,
so t o o does t h e o r g a n i z a t i o n a l l e v e l a t which a response i s
observed. Given t h e marked tempora 1 and s p a t i a l heterogenei t y o f
savannas, it w i 11 be p a r t i c u l a r l y i m p o r t a n t t o c h o o s e t h e
appropriate observationa 1 scale f o r the problems being studied.

RAT1ONALE

Almost one f i f t h o f the world's population l i v e i n savannas, many o f

them i n r u r a 1 s o c i e t i e s t h a t depend on subsi stence agriculture. - Per
c a p i t a food production i s genera 1ly low, the resu lt o f a v a r i e t y o f
envi ronmenta 1, s o c i a l and economic constraints. Rainfa 11 i s high ly
seasona 1 and v a r i ab le, l e a d i ng o f t e n t o uneven and u n p r e d i c t a b l e
p a t t e r n s o f p r i m a r y p r o d u c t i o n . Many o f t h e s o i 1s a r e r e l a t i v e l y
i n f e r t i l e , p a r t i c u l a r l y i n t h e h i g h r a i n f a l l zones. T h i s i s
r e f lected i n low crop y i e l d s and i n the poor n u t r i t i o n a l q u a l i t y o f
n a t u r a l pastures, e s p e c i a l l y d u r i n g t h e d r y season. Consequently,
even i n areas w i t h a h i g h and r e l a t i v e l y c o n s t a n t l e v e l o f p l a n t
p r o d u c t i o n n o t a l 1 t h e p l a n t m a t t e r can be used p r o d u c t i v e l y by
consumers.

Compounding the e f f e c t s o f these factors i s the dual nature o f land

use i n many savanna regions. The most productive lands are o f t e n s e t
a s i d e f o r g r o w i n g cash c r o p s whi l e t h e poorer lands a r e used f o r
l o c a l f o o d p r o d u c t i o n . This, combined w i t h t h e c o n t i n u i n g r a p i d
growth o f the human population i n the tropics, i s placing increasing
p r e s s u r e on m a r g i n a l lands, r e s u l t i n g i n many cases i n degraded
envi ronments, reduced p r o d u c t i v i t y and lower c a r r y i ng capaci ties.
There i s an u r g e n t need t h e r e f o r e t o f i n d a l t e r n a t i v e models o f
deve lopment d e s i gned t o increase t h e s a t i s f a c t i o n o f human needs
w i t h o u t causing a decline i n long-term p r o d u c t i v i t y and resilience.
To be s u c c e s s f u l , t h e s e p r o g r a m m e s need t o b e b a s e d o n
s c i e n t i f i c a l ly sound t e c h n o l o g i e s which t a k e i n t o account t h e
r e l e v a n t p h y s i c a 1, b i o l o g i c a 1 and human elements o f a system.
Unfortunately, our know ledge o f these features, p a r t i c u l a r l y t h e i r
dynamic aspects, i s s t i 11 q u i t e fragmentary. T h i s p r o v i d e s one o f
t h e main motivations f o r t h i s programme.

From b o t h a s c i e n t i f i c and l a n d management v i e w p o i n t , t h e r e a r e a

v a r i e t y o f reasons why t r o p i c a l savannas are an important research
subject. Some o f these are:

1. The i n c r e a s i n g l y intense use o f savannas by an expanding human

p o p u l a t i o n i s r e s u l t i n g i n adverse changes t o t h e s o i 1 and
v e g e t a t i o n . When combined w i t h n a t u r a l s t r e s s e s such as drought,
these changes are leading t o increased erosion and a r i d i f i c a t i o n o f
t h e s o i 1, f o l lowed by f a m i n e and human misery. Research i n t o t h e
causes and consequences o f t h e s e changes i n savannas c o u l d h e l p
a l l e v i a t e some o f the problems i n the future.

2. Within the tropics, t h e coexistence and close i n t e r a c t i o n o f t h e

woody and herbaceous s t r a t a makes savannas unique. Both s t r a t a are
o f economic v a l u e and a b e t t e r u n d e r s t a n d i n g o f t h e i r c o e x i s t e n c e
WOU l d c o n t r i bute t o improved management.

3. Savannas a r e one o f t h e most seasonal o f t h e w o r l d ' s m a j o r

biomes, experiencing s t r o n g l y c o n t r a s t i n g c l i m a t i c conditions w i t h i n
a year, as w e l l as high v a r i a b i l i t y between years. They a l s o d i s p l a y
"an a p p a r e n t l y c o n f u s i n g m i x t u r e o f communities o f various sizes,
arranged i n d i f f e r e n t p a t t e r n s a t d i f f e r e n t sca les, and each o f
which may be changing i n both an o r d e r l y and d i s o r d e r l y fashion, a t
d i f f e r e n t rates" (Walker, 1981: 447). This heterogeneity creates a
constant ly v a r y i ng envi ronment f o r the b i ota and is probab l y a major
f a c t o r i n e n a b l i n g a r e l a t i v e l y l a r g e number o f s p e c i e s and
c o n t r a s t i n g l i f e f o r m s t o c o e x i s t . Savannas t h e r e f o r e a r e i d e a l
systems i n w h i c h t o s t u d y how p l a n t s and a n i m a l s cope w i t h t h e
v a r i o u s n a t u r a 1 s t r e s s e s imposed by a v a r i a b l e e n v i ronment. T h i s
W O U l d c o n t r i b u t e t o a b e t t e r u n d e r s t a n d i ng o f t h e e f f e c t s o f man-
induced stresses i n these and other ecosystems.

4. F i r e , f r e q u e n t l y a s s o c i a t e d w i t h human a c t i v i t i e s , i s a
p r o m i n e n t f e a t u r e o f most t r o p i c a l savannas. I t a f f e c t s t h e
f u n c t i o n i n g o f these systems i n a v a r i e t y o f ways and i n t e r a c t s w i t h
o t h e r processes, such as h e r b i v o r y , n u t r i e n t c y c l i n g and p l a n t
r e c r u i tment. S i n c e f i r e can be managed, a b e t t e r u n d e r s t a n d i n g o f
i t s e c o l o g i c a l effects, and how these can be integrated w i t h other,
less managab l e ecosystem processes, WOU l d be extreme ly va luab le.
5. I n a d d i t i o n t o t h e p o s s i b l e c o n t r i b u t i o n s t h a t t h i s programme
might make t o t h e b e t t e r management o f savannas, f u r t h e r advances i n
ecological theory can be expected i n view o f the uniqueness o f the
s e t o f ecologica 1 i n t e r a c t i o n s t h a t determine the existence o f these
sy stems .
APPROACH AND KEY QUESTIONS

The a p p r o a c h o f t h i s programme i s t h e r e f o r e t o d e v e l o p a
c o l l a b o r a t i v e i n t e r n a t i o n a 1 r e s e a r c h e f f o r t w h i ch, t h r o u g h
comparative and i n d i v i dua 1 studies, w i 11 improve our understandi ng
o f b o t h n a t u r a l and managed savannas and t h e i r responses t o
d i f f e r e n t kinds o f stress and disturbance. The major problems t o be
addressed by the programme are encompassed i n the f o l l o w i n g two key
questions :
1. W h a t a r e t h e s t r u c t u r a l and f u n c t i o n a l p r o p e r t i e s o f savannas
t h a t render them stable and/or r e s i l i e n t t o seasonal and aseasonal
natura 1 stresses and d i sturbance (e.g. f i re, drought)?

2. Are t h e r e c r i t i c a l l i m i t s (i.e. t h r e s h o l d s ) o f d i s t u r b a n c e
beyond which savanna ecosystems do n o t recover a f t e r the disturbance
f a c t o r i s removed?

The r e s o l u t i o n o f t h e problems c a l l s f o r answers t o two f u r t h e r

questions:
3. I n which ways (both s t r u c t u r a l l y and functionally), and by how
much, do d i f f e r e n t types o f savannas change i n response t o n a t u r a l
s t r e s s e s ( f i r e , below-average r a i n f a l l , f loods, etc.) and n a t u r a l
and anthropogenic d i sturbance (e.g. pro longed drought, over-grazing,
de-bushing, and CU l t i v a t i o n ) and t h e i r interactions?

4. What are t h e mechanisms t h a t determine the manner and t h e r a t e s

by which savannas respond t o and recover from a disturbance?
SECTION - THE DETERMINANTS OF SAVANNAS
II: -

INTRODUCTION

Recent syntheses on t h e eco l o g y o f t r o p i c a 1 savannas (Hunt l e y and

Wa l k e r , 1982; Bour l i k e , 1983; Sarmiento, 1984; T o t h i Il and M o t t ,
1985) revea 1 f o u r p r i n c i pa 1 d e t e r m i n a n t s o f savanna s t r u c t u r e and
functioning: s o i 1 moisture, s o i 1 nutrients, herbivory and f i r e . Soi 1
moisture a v a i l a b i l i t y and s o i 1 n u t r i e n t status are the key factors,
a f f e c t i n g both t h e balance between grasses and woody p l a n t s and t h e
p a t t e r n s o f p r i m a r y p r o d u c t i o n and p l a n t qua l i t y . These i n t u r n
i n f l u e n c e t h e k i n d s and e x t e n t o f h e r b i v o r y , a s s o c i a t e d a n i m a l
impacts, and t h e f r e q u e n c y and i n t e n s i t y o f f i r e . I n so doing, t h e
a v a i l a b i l i t y o f m o i s t u r e and n u t r i e n t s and t h e e f f e c t s o f t h e i r
i n t e r a c t i o n are i n d i r e c t ly modi fied.

A f i f t h determinant o f savanna s t r u c t u r e i s human a c t i v i t y . Humans

have been associated w i t h A f r i c a n savannas f o r more than two m i l l i o n
years (Harris, 1980). Likewise, i n Asia, there i s evidence o f human
occupation going back as f a r as one m i l l i o n years BP. A u s t r a l i a and
America were n o t o c c u p i e d u n t i 1 much l a t e r : about 40000 y r s BP i n
A u s t r a l i a , and about 25000 y e a r s BP i n South America. These l o n g
p e r i o d s o f o c c u p a t i o n have had a p r o f o u n d e f f e c t on b o t h t h e
s t r u c t u r e o f savannas and t h e i r geographical extent. For example, i t
i s b e l i e v e d t h a t t h e savannas o f t h e I n d i a n s u b - c o n t i n e n t a r e
d e r i v e d p r i m a r i l y f r o m woodlands t h r o u g h t h e a c t i o n o f humans
(Singh, 1976; Singh e t al., 1983; Gadgil and Meher-Homji, 1985).

Humans can a f f e c t savanna s t r u c t u r e e i t h e r d i r e c t l y , as wood-cutters

and CU l t i v a t o r s , o r i n d i r e c t ly, through t h e i r abi lity t o mani pu l a t e '

f i r e and t o i n f l u e n c e h e r b i v o r e numbers and d i s t r i b u t i o n , b o t h by

hunting and by i n t r o d u c i n g and managing domestic animals. I n more
recent times humans have developed t h e capacity t o b r i n g about r a p i d
and considerable change i n savanna s t r u c t u r e through mechanical and
c h e m i c a l means. O v e r a l l , i n many p a r t s o f t h e w o r l d today, human
a c t i v i t y c o n s t i t u t e s t h e m a j o r source o f d i s t u r b a n c e t o savanna
sys tems .
The f o l l o w i n g account r e p r e s e n t s a s e l e c t i v e r e v i e w o f savanna
f u n c t i o n i n g t o provide a background f o r t h e hypotheses which f o l low.
I n t h i s r e v i e w , we w i s h t o emphasize t h e composite n a t u r e o f t h e
main determinants, each o f which comprise a number o f r a t h e r weak
b u t high ly i n t e r a c t i v e forces capable o f producing cumulative strong
e f f e c t s (see McNaughton, 1983a, f o r discussion and examples). These
e f f e c t s are o f t e n contingent on the frequency, i n t e n s i t y and t i m i n g
o f t h e i n t e r a c t i o n s , on t h e p a s t and p r e s e n t s t a t e s o f t h e system,
and on t h e i r i n t e r a c t i o n w i t h f u t u r e events. I n t h i s respect, t h e
CO-occurrence o f two o r more events, which by themselves may have
lit t l e impact, b u t whose i n t e r a c t i o n is synergi s t i c , may strong ly
a f f e c t the system, p a r t i c u l a r ly when the probabi lity o f the events
o c c u r r i ng is low. Overa 11, t h e consequences o f these i n t e r a c t i o n s
are n o t s t r i c t l y d e t e r m i n i s t i c b u t r a t h e r the product o f a series o f
i n t e r l i n k e d probabi l i t y functions (McNaughton, 1983a).

SOIL MOISTURE DYNAMICS

S o i 1 moi s t u r e regimes i n savannas a r e i n f luenced by ( i t h e t o t a 1
amount and seasonal d i s t r i b u t i o n o f annual r a i n f a l l and t h e
p r o p o r t i o n o f t h i s t h a t e n t e r s t h e s o i 1; ( i i1 t h e w a t e r h o l d i n g
c a p a c i t y o f t h e s o i 1, which is l a r g e ly a f u n c t i o n o f s o i 1 t e x t u r e
and depth, and ( i i i ) t h e amount o f e v a p o t r a n s p i r a t i o n , which i s
r e l a t e d i n comp l e x ways t o c l i m a t e , s o i 1 t e x t u r e , s o i 1 s u r f a c e
characteristics, and the type o f vegetation a t a site.

Rainfall

Mean annual r a i n f a 11 i n savannas ranges f r o m about 300 mm t o more

t h a n 1600 mm and n o r m a l l y does n o t exceed p o t e n t i a l evapo-
t r a n s p i r a t i o n , though it may do so seasonal ly. Most o f t h e r a i n
f a 11s d u r i ng one or, i n e q u a t o r i a 1 regions, two, we 11 - d e f i ned w e t
seasons. The d r y season, which l a s t s as long as 9 months i n semi-
a r i d areas and as s h o r t as 3 months i n m o i s t regions, i s marked by a
shortage o f water f o r p l a n t growth, a t least i n the iipper 1 - 2 m o f
the s o i 1 p r o f i le.
The s p a t i a l and t e m p o r a l d i s t r i b u t i o n o f r a i n f a l l i s o f t e n h i g h l y
v a r i a b l e . Rain f a l l s m a i n l y , i n t h e f o r m o f intense, l o c a l i z e d
thundershowers, and neighbouring areas may receive d i f f e r e n t amounts
o f r a i n on any one occasion. The t i m i n g o f r a i n f a l l w i t h i n a w e t
season a l s o v a r i e s , g i v i n g r i s e t o d i f f e r e n t p a t t e r n s i n t h e
avai l a b i l i t y o f s o i 1 moi sture between years. Most. importantly, there
a r e o f t e n large year-to-year differences i n r a i n f a l l a t a s i t e ,
p a r t i c u l a r ly i n the dry areas.

Whi l e the r a i n f a l l i n any one year i s largely unpredictable, there

a r e s u g g e s t i o n s o f longer-term, q u a s i - p e r i o d i c f l u c t u a t i o n s i n
annua 1 r a i n f a 11 i n some r e g i o n s ( s o u t h e r n A f r i c a : Tyson and Dyer,
19781, b u t n o t i n o t h e r s (Serengeti: Pennycuick and Norton-
G r i f f i ths, 1976). Given the large d i fferences i n p r i mary production
t h a t occur between years i n apparent response t o f l u c t u a t i o n s i n t h e
amount and seasonal d i s t r i b u t i o n o f r a i n f a l l (Poupon and B i 1 le,
1974; R u t h e r f o r d , 1981; S i c o t and Grouzis, 1981; Dye and Spear,
19821, the possible existence and o r i g i n o f these r a i n f a l 1 cycles,
and t h e i r i m p l i c a t i o n s f o r management, needs more investigation.
Soils
The e f f e c t s of v a r i a t i o n in r a i n f a l l a r e compounded by d i f f e r e n c e s
between s o i l s i n permeabil i t y and moi s t u r e r e t e n t i o n p r o p e r t i e s .
Savanna s o i l s Vary widely i n t e x t u r e , s t r u c t u r e , p r o f i l e and depth,
r e f l e c t i n g a t one s c a l e t h e i n t e r a c t i o n of geol ogy, geomorphol ogy
and c l i m a t e , and a t a n o t h e r , t h e i n f l u e n c e o f topography, r e l i c
f e a t u r e s of p a s t landforms, the kind and e x t e n t of vegetation cover,
and animal a c t i v i t y (Young, 1976; Montgomery and Askew, 1983). On a
regional s c a l e t h e a n c i e n t , highly weathered s o i l s o f t h e moist
s a v a n n a s , exposed t o moderate t o high r a i n f a l l ( 600 - 1600 m m
p.a.), a r e l a t i v e l y s h o r t d r y s e a s o n ( < 6 months) and high s o i l
temperatures ( > 18"C), can be d i s t i n g u i s h e d from the younger, l e s s
w e a t h e r e d s o i l s of t h e d r i e r s a v a n n a s which e x p e r i e n c e low and
v a r i a b l e r a i n f a l l ( 300 - 700 m m p.a.1, and a 1 ong dry s e a s o n ( 6 -
10 months).
M o i s t savanna s o i l s : These o c c u r m a i n l y on the o l d , e x t e n s i v e
ol a t e a u x o f t h e Gondwana and mi d - T e r t i a r v ol a n a t i o n s u r f a c e s . a s
;el 1 a s on r e d i s t r i b u t e d s a n d s s i t u a t e d ;n'the p e r i p h e r y of t h e s e
p l a t e a u x and i n upland b a s i n s . Most o f t h e s o i l s a r e d e r i v e d from
f e l s i c o r i n t e r m e d i a t e r o c k s ( g r a n i t e s , gnei s s e s , p h y l l i t e s ,
s c h i s t s , s a n d s t o n e s ) of t h e u n d e r l y i n g Precambrian c o n t i n e n t a l
shields. The s o i l s a r e moderately t o highly weathered and leached,
and a r e general l y s t r u c t u r e l e s s ( o f t e n massive i n highly weathered,
c l ayey profi 1e s ) t o weakly structured. Well s t r u c t u r e d soi 1 s occur
i n 1 e s s w e a t h e r e d p r o f i l e s , p a r t i c u l a r l y ones d e r i v e d from b a s i c
i n t r u s i o n s o r from l i m e s t o n e . K a o l i n i t i c c l a y s and i r o n and
aluminium o x i d e s a r e t h e main s e c o n d a r y mi n e r a l S. Expandi ng 2: 1
1a t t i c e c l ay minera1 s a r e uncommon.
The s o i l s a r e generally sandy a t t h e surface, becoming more clayey
w i t h depth. T h i s c o n t r a s t i n t e x t u r e between t h e t o p s o i l and t h e
subsoil tends t o be gradua1 i n a c t i v e l y weathering s o i l s but where
t h e s o i l s a r e highly weathered i t i s usually abrupt, giving r i s e t o
a duplex profi le. Cateri t e i s commonly present and may form massive
sheets. Uniformly textured p r o f i l e s a r e found i n sandy s o i l s derived
from t h e weathering of sandstanes and unconsolidated sediments, and
i n some highly weathered s û i l s of the r a i n forest-savanna t r a n s i t i o n
zone. The l a t t e r s o i l s contain l a r g e amounts of f r e e iron oxides and
1:l l a t t i c e clays. The c l a y s a r e aggregated i n t o r e l a t i v e l y s t a b l e
sand-sized p a r t i c l e s t h a t impart t o the p r o f i l e the consistence and
d r a i n a g e c h a r a c t e r i s t i c s o f m o r e l o a m y s o i l s (Young, 1976;
Montgomery and Askew, 1983).
I n f i l t r a t i o n r a t e s a r e g e n e r a l l y h i g h i n sandy s o i 1s and i n c l a y s
w i t h a well-developed crack o r crumb structure. However, clays w i t h
an aggregate crumb s t r u c t u r e are susceptible t o s t r u c t u r a 1 CO 1lapse
when exposed t o r a i n d r o p i m p a c t o r t o o t h e r forms o f mechanical
pressure. These break up t h e aggregates and reduce t h e s i z e o f
s u r f a c e pores, t h e r e b y r e d u c i ng in f i1t r a t i o n r a t e s and increasing
r u n o f f ( B r i d g e e t al., 1983). I n f i l t r a t i o n i s a l s o reduced, even i n
sandy soi ls, by the presence o f water-repel l a n t seals. These can be
caused e i t h e r by dormant blue-green algae l i v i n g on t h e s o i 1
surface, o r by t h e a c c u m u l a t i o n o f secondary c h e m i c a l compounds
leached from t h e l i t t e r . Since these seals break down w i t h prolonged
wetting, t h e i r impact i s greatest e a r l y i n the wet season o r a f t e r
pro longed dry spe 11s.

The upper p a r t o f the s o i 1 p r o f i l e i n the high r a i n f a l l savannas i s

a t o r c l o s e t o f i e l d c a p a c i t y f o r much o f t h e w e t season (Young,
1976). Excess w a t e r d r a i n s r a p i d l y down t o t h e subsoi 1 which can
r e m a i n m o i s t t h r o u g h o u t t h e year, depending on t h e amount o f
r a i n f a l l and percolation, p r o f i l e depth, proximity t o groundwater,
and the r a t e o f depletion o f the accumulated water by plants. Good
a e r a t i o n and a low mechanical r e s i s t a n c e t o r o o t p e n e t r a t i o n i n
moderate t o w e l l - s t r u c t u r e d soi 1s enables deep-rooted woody p l a n t s
t o u t i l i z e t h i s moisture, o f t e n w e l l i n t o t h e d r y season and l o n g
a f t e r t h e w a t e r c o n t e n t o f t h e upper 1 -2 m o f t h e p r o f i l e has
f a 1 len t o below w i l t i n g point.
The most extreme m o i s t u r e regimes occur i n s o i 1s w i t h a s t r o n g l y
duplex p r o f i l e o r which a r e u n d e r l a i n by an i n d u r a t e d i r o n s t o n e
( l a t e r i t e ) layer. The i n t e r n a 1 drainage o f these soi 1s is poor and,
because o f t h e reduced e f f e c t i v e d e p t h o f t h e p r o f i le, t h e y a l s o
have a low w a t e r s t o r a g e c a p a c i t y . Consequently, t h e s o i l s a r e
a l t e r n a t e l y w a t e r logged and d r y (Sarmiento and Monasterio, 1975;
Tinley, 1982). However, waterlogging i s n o t confined t o such soi 1s.
S i t e d r a i n a g e on t h e a l m o s t l e v e 1 p l a t e a u x i s o f t e n poor and even
sandy soi 1s can be saturated i f they occur i n s l i g h t depressions.
Dry savanna s o i 1s: Whereas m o i s t savannas occur on p l a t e a u x and
up lands, the(lrysavannas occur on low a l t i tudes p l a i n s , i n b r o a d
downwarped basins, and i n r i f t - f a u l t e d v a l l e y s , o f t e n a t t h e
extremes o f t h e t r o p i c s . The p r e d o m i n a n t l y sandy and s i l t y s o i 1s
o r i ginate main ly from unconso lidated sediments, r e d i s t r i buted sands,
and from a c i d igneous and sedimentary rocks o f Precambrian and l a t e r
age. C l a y - r i c h p r o f i l e s a r e l e s s common but, where present, t h e y
contain higher proportions o f expanding 2:l l a t t i c e c lays than f i n e -
textured s o i 1s i n m o i s t savannas (Young, 1976). Since t h e c l i m a t e i s
dry, t h e s o i 1 p r o f il e is se ldom thorough l y wetted. Weatheri ng and
leaching r a t e s are therefore low. Where h i g h l y weathered and leached
s o i l s do occur, as i n A u s t r a l i a , t h e s e i n d i c a t e t h e occurrence o f
wetter periods i n the past.
As a r e s u l t o f the l o u e r weathering rates, s o i l s formed i n s i t u f r o i
c r y s t a l l i n e rocks, i n freely-drained s i tes, tend t o b e ç h m w and
sandy, w i t h l i t t l e o r no p r o f i l e development (Young, 1976).
I n f i l t r a t i o n and p e r c o l a t i o n r a t e s a r e high. T h i s f a v o u r s t h e
conservation o f rainwater since i t lessens the amount o f moisture
t h a t can be l o s t t h r o u g h evaporation. However, t h i s i s o n l y
e f f e c t i v e i n deep sands since most o f the s o i l s are too shallow t o
be able t o s t o r e large amounts o f water.
C layey s o i 1s i n dry savannas deve lop i n areas under l a i n by base-ri ch
r o c k s ( b a s a l t , s h a l e s , l i m e s t o n e s ) , and i n p o o r l y d r a i n e d
bottom lands and depressions where c lays and minera 1s ( p a r t i c u l a r ly
calcium and magnesium) accumulate. Self-mulching c lays ( v e r t i sols)
common ly occur and are p a r t i c u l a r l y widespread i n Austra lia, I n d i a
and i n p a r t s o f Africa. Clays w i t h t e x t u r a l B-horizons, produced by
c l a y t r a n s l o c a t i o n , a r e a l s o common. I n these s o i l s , o f t e n h i g h
leve 1s o f exchangeab l e sodium cause the dispersion o f c lays which
t h e n accumu l a t e i n t h e subsoi 1 where they r e s t r i c t i n t e r n a 1
drainage, aeration and r o o t penetration. Subsoil drainage can a l s o
be i n h i b i t e d by impermeable layers formed by the p r e c i p i t a t i o n o f
c a l c i u m carbonate and s i l i c a t e s . The gradua1 e x t e n s i o n o f t h e s e
layers towards the surface reduces the e f f e c t i v e depth o f the s o i l
and l i m i t s i t s water storage potential.

I n f i l t r a t i o n r a t e s and permeabi l i t y are considerably lower i n clayey

p r o f i les. This i s o f t e n compounded by the presence o f a l g a l c r u s t s
o r surface sea 1s resu l t ing from the dispersion o f c lay p a r t i c les by
r a i n d r o p i m p a c t (Penning de V r i e s and D j i t b e , 1982). Clayey s o i 1s
therefore tend t o be more a r i d than sandy soi 1s under equivalent low
r a i n f a 11, d e s p i t e h a v i ng a h i g h e r w a t e r - h o l d i n g c a p a c i t y . I n
c o n t r a s t t o sandy s o i l s , much o f t h e w a t e r e n t e r i n g t h e s o i 1 i s
r e t a i n e d c l o s e t o t h e s u r f a c e where i t i s s u s c e p t i b l e t o
evaporation. Furthermore, a t given water contents, clays have more
negative soi 1 water potentials, so t h a t proportionately less o f the
water i n the s o i 1 i s avai l a b l e t o plants. Soi1 t e x t u r e therefore has
o p p o s i t e e f f e c t s on s o i l m o i s t u r e r e g i m e s and p l a n t w a t e r
avai l a b i lit y i n moi s t and a r i d regions. Under high r a i n f a 11, c lays
are the w e t t e s t soi 1s but, where r a i n f a l l i s low, they are generally
the most a r i d (Walter, 1971). In sands, the s i t u a t i o n i s reversed.

The r e d i s t r i b u t i o n and concentration o f water by overland f l o u has

important l o c a l e f f e c t s on s o i 1 moisture regimes. I n some s i t e s i n
the d r i e r savannas, more than h a l f the incoming r a i n f a l l can be l o s t
as s u r f a c e r u n o f f , depending on t h e amount o f p l a n t cover, s o i l
s u r f a c e c o n d i t i o n s , topography, amount and i n t e n s i t y o f r a i n f a l l ,
and antecedant s o i 1 moi s t u r e c o n d i t i o n s (Walker and Cunningham,
1976; Penning de V r i e s and D j i t k y e , 1982). A s m a l l e r percentage o f
t o t a l r a i n f a l l i s l o s t as r u n o f f i n moist savannas, though absolute
amounts may be higher. Much o f t h i s r u n o f f flows onto adjacent areas
where i t i s absorbed. This increases t h e e f f e c t i v e r a i n f a l l o f these
s i t e s and creates refuges f o r drought-sensitive plants i n years o f
low r a i n f a l l (Macdonald, 1978). On a more l o c a l scale, t e r m i t e
mounds may have the same effect.

The i n f l u e n c e o f topography on s o i 1 p r o p e r t i e s and s o i 1 m o i s t u r e

regimes is best r e f lected i n t h e regu l a r sequence o f d i f f e r e n t s o i 1
p r o f i les which develop along a gradient from h i 1l t o p o r i n t e r f l u v e
c r e s t t o adjacent va 1ley bottom. These soi 1 catenas are p a r t i c u l a r ly
w e l l developed i n ancient, gently undu l a t i n g savanna landscapes w i t h
d i f f u s e drainage systems (Mi lne, 1935; Young, 1976). The changes i n
soi 1 properties are r e l a t e d p r i m a r i l y t o differences down the slope
i n (il t h e e x t e n t o f e r o s i o n by s u r f a c e wash, a s s o c i a t e d w i t h
changes i n s l o p e a n g l e and t h e d i s t a n c e o f o v e r l a n d f low; ( i i ) t h e
degree o f leaching, downs lope t r a n s p o r t and deposition o f c l a y s and
d i s s o l v e d s o l u t e s ; and ( i i i ) p r o x i m i t y t o groundwater and i t s
seasonal f l u c t u a t i o n s (Young, 1976). The r e s u l t i s the formation o f
w e l l - d r a i n e d and o f t e n h i g h ly- leached s o i 1s i n t h e upper p a r t s o f
t h e catena, and b a s e - r i c h g l e y s i n t h e poorly-drained, more
r e s t r i c t e d bottomlands. The changes i n soi 1 properties and moisture
r e g i mes i n t u r n g i v e r i se t o zona 1 p a t t e r n s i n t h e c o m p o s i t i o n ,
s t r u c t u r e , p r o d u c t i v i t y and q u a l i t y o f t h e vegetation, and i n t h e
c o r r e s p o n d i n g p a t t e r n s o f h e r b i v o r y (Morison e t al., 1948; Menaut
and C k a r , 1979; B e l l , 1981; T i n l e y , 1982).

SOIL NUTRIENT DYNAMICS

Savanna n u t r i e n t dynamics are the outcome over a v a r i e t y o f t i m e and

s p a t i a l sca l e s o f i n t e r a c t i o n s b e t w e e n c l i m a t e , g e o l o g y ,
geomorphology and the biota. Variations i n the physical and chemical
' properties o f savanna s o i 1s are strongly r e l a t e d t o differences i n
bedrock and the degree t o which t h i s has been exposed by weathering
and erosion. These p r o p e r t i e s have been subsequent ly modi f ied b y
i n t e r a c t i o n s between the s o i 1, c l i m a t e and geomorpho logy a f f e c t i ng
s o i 1 microclimate, secondary m i n e r a l f o r m a t i o n , and t h e e r o s i o n ,
l e a c h i n g and r e l o c a t i o n o f s o i 1 and n u t r i e n t s . Soi 1 m o i s t u r e and
temperature i n t u r n influence weathering and leaching r a t e s and t h e
l e v e l o f b i o t i c a c t i v i t y , p a r t i c u l a r l y i n the soi 1.

Soi 1s

Dystrophic savanna soi 1s: S o i l s derived from the weathering o f a c i d

c r y s t a l l i n e rocks o f t h e Precambrian continental shields, as w e l l as
those formed from ancient sedi mentary formations and r e d i s t r i buted
sands, g e n e r a l ly have a low r e s e r v e o f weatherable m i n e r a l s . They
are most widespread in, b u t are not confined t o areas r e c e i v i n g high
r a i n f a l l and so a r e o f t e n h i g h l y weathered and leached. The
predominance o f non-expanding 1:l l a t t i c e c l a y s and i r o n and
aluminium oxides resu l t s i n low e f f e c t i v e c a t i o n exchange capaci ty
and small amounts o f t o t a l exchangeable bases, p a r t i c u l a r l y Ca and
Mg (Jones and W i ld, 1975; Lopes and Cox, 1977; M o t t e t al., 1985).
The l e v e l s o f a v a i l a b l e P a r e a l s o f r e q u e n t l y v e r y low and some
s o i l s w i t h an abundance o f s e s q u i o x i d e s have a h i g h c a p a c i t y f o r
f i x i n g phosphorus. Except i n very a c i d s o i ls, the amount o f organic
m a t t e r i s t h e main d e t e r m i n a n t o f c a t i o n exchange c a p a c i t y . The
quanti ty o f soi 1 organic matter is p o s i t i v e ly corre l a t e d wi t h mean
annual r a i n f a l l and t h e l e n g t h o f t h e w e t season but, o v e r a l l ,
l e v e l s are generally low (Jones and W i ld, 1975; Lopes and Cox, 1977;
Kadeba, 1978; Montgomery and Askew, 1983; M o t t e t al., 1985).

Some h i g h l y weathered s o i l s , p a r t i c u l a r l y i n South America, have

high l e v e l s o f exchangeable aluminium (Lopes and Cox, 1977). These
pose prob lems f o r crops and introduced Pasture grasses. Indigenous
species, however, appear t o be adapted t o these high levels and some
woody species accumu l a t e t h e meta 1 i n t h e i r t i s s u e s (Haridasan,
1982). O u t s i d e South America, l i t t l e i s known about t h e i m p a c t o f
a l u m i n i u m on n u t r i e n t u p t a k e and c y c l i n g . T h i s i s a t o p i c which
needs i n v e s t i g a t i o n .
E u t r o h i c savanna s o i 1s: These s o i 1s occur i n areas u n d e r l a i n by
d s m t m c intrusions; i n i n c i sed v a l leys where less
weathered m a t e r i a l i s b e i n g exhumed; i n a l l u v i a l d e p o s i t s a l o n g
drainage lines, v a l ley bottoms o r f loodplains; and i n areas covered
by basic volcanic ash. These s o i l s are more prominent i n d r i e r areas
where the amount o f r a i n f a l l has been i n s u f f i c i e n t t o promote e i t h e r
e x t e n s i v e w e a t h e r i ng o r l e a c h i ng. Consequently, there are usual ly
adequate reserves o f weatherable minerals (Young, 1976). The s o i 1s
a r e l e s s a c i d and may even be m o d e r a t e l y a l k a l i n e . The o f t e n h i g h
p r o p o r t i o n s o f 2 : l l a t t i c e c lays, p a r t i c u l a r l y i n v e r t i sols,
c o n t r i bute t o h i gh c a t i o n exchange capacities and higher leve 1s o f
exchangeab l e bases. The amount o f s o i 1 organic matter is genera 1ly
low and therefore has l i t t l e e f f e c t on the c a t i o n exchange capacity
o f t h e s o i 1.

Organic matter dynamics

The breakdown o f o r g a n i c m a t t e r and t h e subsequent r e l e a s e o f
n u t r i e n t s , as w e l l as t h e r o l e s p l a y e d i n these processes by t h e
biota, d i f f e r considerably i n r e l a t i o n t o soi 1 moisture regimes and
s o i 1 f e r t i l i t y (Menaut e t al., 1985). I n t h e d r y savannas, p r i m a r y
production i s l i m i t e d by t h e low r a i n f a l l and short wet season. The
i n p u t o f organic matter t o the s o i 1 i s therefore also low and h i g h l y
seasonal. The q u a l i t y o f these inputs i s generally high and much o f
t h e m a t e r i a l i s r e a d i l y decomposed by t h e s o i 1 b i o t a . However,
n u t r i e n t r e l e a s e i s i n t e r m i t t e n t because m i c r o b i a 1 a c t i v i t y i s
s t r o n g l y l i m i t e d f o r much o f t h e t i m e by t h e a r i d i t y o f t h e s o i 1.
There i s a n e t m i n e r a l i z a t i o n o f nitrogen a t the s t a r t o f the rains,
though t h i s i s not sustained because the small organic matter stock
i s r a p i d l y decomposed and an i n c r e a s i n g amount o f n i t r o g e n i s
i m m o b i l i z e d i n t h e m i c r o b i a l biomass (Bernhard-Reversat, 1982;
Penning de Vries and Djitiye, 1982; Menaut e t al., 1985).

In contrast, i n the moist savannas, the higher r a i n f a l l and extended

wet season favour high p l a n t production and organic matter i n p u t t o
the soi 1, although t h i s i n p u t may be l i m i t e d by frequent fires. The
s o i 1 m i c r o c l i m a t e f a v o u r s m i c r o b i a l a c t i v i t y and t h e r e i s a
c o n s i d e r a b l e p o t e n t i a l f o r m i n e r a l i z a t i o n . However, m i c r o b i a l
a c t i v i t y i s l i m i t e d by t h e low q u a l i t y o f t h e o r g a n i c m a t t e r ,
p r i n c i p a l ly t h e low l e v e l s o f a s s i m i l a b l e carbon, h i g h C:N r a t i o s
and l i g n i n c o n t e n t s and, i n some cases, h i g h amounts o f condensed
tannins and other secondary chemicals. The release o f n u t r i e n t s i s
slow and r e s u l t s i n a low s t a n d i n g s t o c k o f a v a i l a b l e n u t r i e n t s .
Because o f t h e h i g h r a i n f a l l , t h e n u t r i e n t s a r e s u s c e p t i b l e t o
leachi ng.

M i c r o b i a l a c t i v i t y i s stimulated by exudates from p l a n t r o o t s and by

water soluble carbon compounds which are introduced i n t o the s o i l
during i t s passage through the guts o f earthworms (Lavelle e t al.,
1983; Menaut e t àl., 1985). These compounds are produced i n inverse
p r o p o r t i o n t o t h e i r l e v e 1s in t h e s o i 1, r e s u l t ing i n c o n s i derab l e
b i o l o g i c a l r e g u l a t i o n o f organic matter decomposition. Because t h e
avai l a b i lity o f assimi l a b l e carbon is l a r g e l y confined t o earthworm
c a s t s and-the neighbourhood o f r o o t s , t h e r e l e a s e o f n u t r i e n t s i s
high l y localized

Savanna p l a n t s m a i n t a i n h i g h root:shoot r a t i o s . The t u r n o v e r o f

roots, w h i c h i s r e l a t e d t o t h e l e v e l o f p l a n t production, may be
r e s p o n s i b l e f o r most o f t h e o r g a n i c m a t t e r i n savanna s o i l s ,
p a r t i c u l a r l y i n frequently burned savannas where f i r e destroys much
o f t h e l i t t e r (Sanford, 1982). Since s o i 1 o r g a n i c m a t t e r i s
i m p o r t a n t i n m a i n t a i n i n g s o i 1 s t r u c t u r e and f e r t i l i t y , more
i n f o r m a t i o n i s needed on t h e s o u r c e s , q u a l i t y and r a t e o f
decomposition o f organic material, and the factors a f f e c t i n g these.

Although t h e l e v e l s o f p l a n t - a v a i l a b l e n u t r i e n t s i n savanna s o i l s
a r e r e l a t i v e l y low, t h e g r e a t e r p r o p o r t i o n o f t h e t o t a l n u t r i e n t
p o o l l i e s i n t h e s o i 1 and s o i l o r g a n i c m a t t e r r a t h e r t h a n i n t h e
v e g e t a t i o n and l i t t e r (Nye and Green land, 1961; Abbadie, 1983;
Sarmiento, 1984; F r o s t , 1985). I n t h e d r i e r savannas, t h i s l a r g e l y
r e f l e c t s the r e l a t i v e l y low p l a n t biomass and the c o n s t r a i n t imposed
by t h e seasonal shortage o f w a t e r on p l a n t g r o w t h and n u t r i e n t
uptake. I n t h e more m o i s t savannas i t r e f l e c t s t h e s l o w r a t e o f
release o f n u t r i e n t s from s o i 1 organic matter.
The r a t e a t which n u t r i e n t s c y c l e t h r o u g h savanna v e g e t a t i o n i s
r e l a t i v e l y rapid, p a r t i c u l a r l y through the herbaceous layer where
n u t r i e n t s turnover 2 - 4 t i m e s f a s t e r t h a n through woody p l a n t s
( F r o s t , 1985). The d i f f e r e n c e i n t h e r a t e s o f n u t r i e n t t u r n o v e r i n
woody p l a n t s and grasses extends also t o l i t t e r decomposition. Grass
l i t t e r , i n t h e absence o f f i r e and under t h e same c o n d i t i o n s ,
decomposes 2-7 times f a s t e r than woody l e a f l i t t e r , and many times
f a s t e r than wood i t s e l f ( M o r r i s e t al., 1982; M o t t e t al., 1985).
Where annual f i r e s occur, much o f t h e grass and some o f t h e woody
l i t t e r g e t s b u r n t r a t h e r t h a n decomposed (Hopkins, 1966; Sanford,
1982; F r o s t , 1985). T h i s tends t o reduce b u t does n o t e n t i r e l y
e l i m i n a t e the d i f f e r e n c e i n r a t e s since most o f the grass m a t e r i a l
t h a t i s b u r n t WOU l d norma 1ly decompose w i t h i n a year, whi l e woody
l i t t e r takes much longer.

FIRE AND HERBIVORY

W hereas w a t e r and n u t r i e n t a v a i l a b i l i t y Vary continuously through

t i m e w i t h i n l i m i t s s e t by t h e p r e v a i l i n g c l i m a t e , s o i 1, and
a s s o c i a t e d b i o l o g i c a l processes, f i r e and h e r b i v o r y a r e events
o c c u r r i n g a t p a r t i c u l a r times, w i t h s p e c i f i c i n t e n s i t i e s , a t
d i s c r e t e i n t e r v a l s . These f e a t u r e s a r e i n t e r r e l a t e d and, t a k e n
together, make up the p a r t i c u l a r f i r e o r herbivory regime a t a site.
S i nce t h e s e r e g i m e s c a n be m a n i pu l a t e d d i r e c t l y , t h e y a r e
p o t e n t i a l l y important management t o o l s . However, t h e consequences
are n o t always easy t o p r e d i c t because f i r e and herbivory i n t e r a c t ,
and t h e outcome i s o f t e n c o n t i n g e n t on t h e p a r t i c u l a r times,
i n t e n s i t i e s and f r e q u e n c y o f t h e i n t e r a c t i o n s . E x t e r n a l ,
unpredictable factors, such as f u t u r e r a i n f a l l o r drought, can a l s o
intervene and influence t h e outcome.

F i r e s a n d h e r b i v o r e s b o t h consume and damage p l a n t m a t t e r and a r e

able t o k i 11 i n d i v i d u a l plants, p a r t i c u l a r l y seedlings. This reduces
t h e e x i s t i n g density and s t r u c t u r e o f a p l a n t community and a f f e c t s
i t s f u t u r e c o m p o s i t i o n and dynamics. Although changes i n p l a n t
community composition can r e s u l t from pressures exerted by f i r e o r
h e r b i v o r y a l o n e , m a j o r c h a n g e s usua 1 l y r e s u l t f r o m t h e i r
i n t e r a c t i o n . Both have r e l a t i v e l y s e l e c t i v e e f f e c t s . H e r b i v o r y i s
u s u a l l y l i m i t e d t o p a r t i c u l a r p l a n t species and p l a n t p a r t s ,
especia 1 l y those o f above-average n u t r i t i o n a 1 qua lity. Moreover,
h e r b i v o r e i m p a c t s t e n d t o be r e s t r i c t e d i n space b u t a r e more
u n i f o r m l y d i s t r i b u t e d i n time. I n c o n t r a s t , f i r e i s a p e r i o d i c
event, o f t e n o c c u r r i n g o v e r a wide area and a f f e c t i n g b o t h l i v i n g
and dead m a t e r i a l l a r g e l y i r r e s p e c t i v e o f t h e i r n u t r i t i o n a l qua lity.
However, f i r e i s s e l e c t i v e t o the degree t h a t there are considerable
d i fferences between species i n t h e i r suscepti b i lit i e s and responses
t o f i r e ( F r o s t , 1984).
M o s t savanna f i r e s o c c u r d u r i n g t h e d r y season as s u r f a c e f i r e s ,
b u r n i n g t h r o u g h t h e herbaceous v e g e t a t i o n . T h e i r i n c i d e n c e and
i n t e n s i ty depends on ( i ) t h e presence o f s u f f i c i e n t fue 1 t o support
a f i r e ; ( i i ) the moisture content o f the fuel; and ( i i i ) a source o f
i g n i t i o n . Many o f them a r e i g n i t e d by man, though i n t h e e a r l y w e t
season l i g h t n i n g can be i m p o r t a n t . I n m o i s t savannas, f i r e s
-
g e n e r a l l y o c c u r e v e r y 1 3 years, b u t as mean annual r a i n f a l l
d e c l i n e s , t h e i n t e r v a l between successive f i r e s i n c r e a s e s and
becomes more variab le.

The i n c i d e n c e o f f i r e i s l a r g e l y a f u n c t i o n o f t h e dry-season
s t a n d i n g c r o p o f grass, i t s e l f a p r o d u c t b o t h o f t h e amount o f
r a i n f a l l and p l a n t production during t h e previous wet season and t h e
e x t e n t o f herbivory. F i r e i n t e n s i t y i s v a r i a b l e and depends l a r g e l y
on t h e amount and s t r u c t u r e o f t h e f u e l , i t s degree o f c u r i n g , and
p r e v a i ling a m b i e n t c o n d i t i o n s . Because o f t h e i r h i g h e r f u e 1 load,
f i r e s i n m o i s t savannas a r e u s u a l l y more i n t e n s e t h a n t h o s e
occurring i n t h e dry savannas.

Annual b u r n i n g has l i t t l e d i r e c t e f f e c t on t h e s o i 1. Most e f f e c t s

a r e i n d i r e c t , r e s u l t i n g f r o m changes t o t h e v e g e t a t i o n , and a r e
c o n f i n e d t o t h e s u r f a c e s o i 1 (Sanford, 1982). Organic m a t t e r and
t o t a 1 n i t r o g e n a r e reduced and a v a i l a b l e phosphorus is s l i g h t l y
i n c r e a s e d i n areas exposed t o r e g u l a r , i n t e n s e , l a t e d r y season
f i r e s . E a r l y o r l e s s i n t e n s e b u r n s have much l e s s o f an e f f e c t
(Harrington and Ross, 1974; Brookman-Ami ssah e t a l., 1980).

B u r n i n g may speed up t h e r a t e o f n u t r i e n t c y c l i n g by r e d u c i n g
l i t t e r , e s p e c i a l l y components such as woody l e a f lit t e r and dead
wood which decompose s lowly. Nitrogen, carbon and sulphur are l o s t
t h r o u g h v o l a t i l i z a t i o n and removal i n smoke and ash. However, t h e
o v e r a l l s i g n i f i c a n c e o f t h e s e l o s s e s has n o t been assessed. By
reducing l i t t e r and herbaceous p l a n t cover, f i r e also bares the soi 1
surface, exposing i t t o raindrop impact, wind and Sun. The length o f
t i m e t h a t t h e soi 1 remains bare i s v a r i a b l e and depends l a r g e l y on
the time o f the f i r e , t h e r a t e o f regrowth o f the vegetation and t h e
t i m i n g o f subsequent r a i n f a 11.

F i r e k i 11s seedlings, s a p l i n g s and s m a l l trees, p a r t i c u l a r l y i n

higher r a i n f a l l areas where grass production i s substantial and t h e
annua 1 dry-season f i res are intense. Under these conditions, woody
p l a n t r e c r u i t m e n t i s g e n e r a l l y e p i s o d i c . F i r e a l s o damages
aboveground p a r t s o f p l a n t s and r e t a r d s t h e g r o w t h o f shrubs and
saplings, thereby lowering t h e biomass o f woody p l a n t s (Rutherford,
1981). The e f f e c t s on d e n s i t y a r e v a r i a b l e and depend on t h e r a t e s
o f m o r t a l i t y and coppicing. Many o f these e f f e c t s are a f u n c t i o n o f
f i r e i n t e n s i t y . The h o t t e s t f i r e s u s u a l l y occur during the l a t e dry-
season and these can reduce a woodland canopy t o coppice. Woodland
species, though, a r e a b l e t o r e g e n e r a t e under a r e g i m e o f l e s s
intense, ear ly dry-season f i res (Trapne 11, 1959; Brookman-Amissah e t
al., 1980; Sanford, 1982; Edroma, 1984).

Protection from f i r e s resu l t s i n an increase i n t r e e densi ty and a

decrease i n grass production. I n mesic areas, savanna woodland
e v e n t u a l l y develops b u t i n t h e h i g h r a i n f a l l s a v a n n a l f o r e s t
t r a n s i t i o n zone, f o r e s t species gradua 1l y estab l i s h (Trapne 11, 1959;
Menaut,1977; Brookman-Amissah etal.,1980; SanJoséand Farinas,
1983). I n the dry savannas, f i r e s seldom occur frequently enough t o
l i m i t the density o f woody plants, though when they do occur, o f t e n
a f t e r p r o l o n g e d p e r i o d s o f above-average r a i n f a 11, mature woody
p l a n t s may experience c o n s i d e r a b l e m o r t a lit y s i n c e t h e y a r e more
susceptible t o f i r e than moi s t savanna plants.

Young t r e e s grow r a p i d l y i n areas p r o t e c t e d f r o m b o t h f i r e and

herbivory, more so than i n areas protected from f i r e alone. Growth
i n p l a n t s exposed t o b o t h p r e s s u r e s i s s e v e r e l y r e s t r i c t e d
( H a r r i n g t o n and Ross, 1974; Belsky, 1984). F i r e and h e r b i v o r y
t h e r e f o r e i n t e r a c t , w i t h f r e q u e n t f i r e s keeping woody p l a n t s a t a
h e i g h t and i n a acceptable s t a t e f o r browsers, while the e f f e c t o f
the browsers i s t o reduce woody p l a n t growth and keep p l a n t s w i t h i n
t h e s i z e range a f f e c t e d by f i r e (Trollope, 1974; Pellew, 1983).

Herbi vory

Savannas make up most o f t h e w o r l d ' s t r o p i c a l g r a z i n g lands and

c u r r e n t l y s u p p o r t a l a r g e b i omass o f domestic livestock, m a i n ly
c a t t l e , goats and sheep. I n many regions, t h e biomass o f these
species c u r r e n t l y equals o r exceeds t h a t o f t h e i n d i g e n o u s
h e r b i v o r e s which used t o occur (Cumming, 1982; M o t t e t al., 1985).
Each o f the main savanna regions i n the past had a d i s t i n c t i v e large
h e r b i v o r e fauna, though t h a t o f A f r i c a appears t o have been, and
s t i l l i s , the most abundant and diverse.

F o r b i o g e o g r a p h i c reasons, A u s t r a l i a never had any proboscids,

a r t i odacty 1s o r p e r i ssodactyls. I n s t e a d , these savannas were
popu l a t e d main l y by macropods (kangaroos and wa 1l a b i e s ) , some o f
which survive today alongside f e r a 1 and domestic ungulates. South
Awerica, i n contrast, had a diverse fauna o f large ungulate and non-
ungu l a t e h e r b i v o r e s t h r o u g h o u t t h e T e r t i a r y , w i t h most o f t h e
s p e c i e s becoming e x t i n c t d u r i n g t h e l a t e P l e i s t o c e n e e i t h e r as a
r e s u l t o f concurrent c l i m a t i c change o r the a r r i v a 1 o f man ( M a r t i n
and Wright, 1967). What p r o p o r t i o n o f these species a c t u a l l y
i n h a b i t e d savannas is n o t known, b u t t h e present-day indigenous
fauna and t h e c a r r y i n g c a p a c i t y f o r domestic l i v e s t o c k on n a t u r a l
range land is r e l a t i v e l y low.
O n l y A f r i c a , and t o a l e s s e r e x t e n t I n d i a , c u r r e n t l y have
s i g n i f i c a n t p o p u l a t i o n s o f i n d i g e n o u s ungulates and o t h e r l a r g e
h e r b i v o r e s . West A f r i c a n savannas s u p p o r t l o w e r numbers and a
s m a l l e r biomass o f l a r g e h e r b i v o r e s t h a n do those i n eastern,
c e n t r a i and southern A f r i c a ( B e l l , 1982; M i l l i g a n e t al., 1982).
T h i s p a r t l y r e f l e c t s t h e h i g h h u n t i n g pressure and p a r t l y t h e
impoveri shed s o i ls, low qua lit y v e g e t a t i o n and consequent l o w
c a r r y i n g capacity o f the West A f r i c a n savannas.

S t u d i e s o f h e r b i v o r y i n savannas have focussed l a r g e l y on t h e

e f f e c t s o f w i I d l a r g e ungulates, m a i n l y i n A f r i c a n ecosystems. I t
must be emphasized however c h a t t h e s e a r e n o t t h e o n l y n o r
necessari ly the most important herbivores i n savannas. I n many areas
today, domestic l i v e s t o c k , especia 1l y c a t t l e , have e f f e c t s t h a t
override those o f indigenous species. Moreover, harvester termites,
l e a f - c u t t i n g a n t s , l e p i d o p t e r a l a r v a e , and g r a s s h o p p e r s
( p a r t i c u l a r l y l o c u s t s ) , a l 1 have m a j o r e f f e c t s i n some systems,
though i n most cases these have n o t been quantified.

There i s c o n s i d e r a b l e v a r i a t i o n among h e r b i v o r e s i n t h e degree o f

se l e c t i v i ty i n t h e i r diets, r e f l e c t i n g a comp lex interplay between
t h e k i n d o f animal, i t s body s i z e and associated energy and n u t r i e n t
requirements, and t h e g r o w t h forin, s t r u c t u r e , chemi s t r y and
phenology o f p o t e n t i a l f o o d p l a n t s . The f u n c t i o n a l responses o f
herbivores t o changes i n p l a n t abundance are also important, though
these have n o t been s u f f i c i e n t l y s t u d i e d
V e r t e b r a t e and i n v e r t e b r a t e h e r b i v o r e s i n savannas b o t h show a
marked p r e f e r e n c e f o r f e e d i n g e i t h e r on broadleafed, o f t e n woody
p l a n t s , o r on grasses and sedges. Mixed feeders a r e much l e s s
common. This s p e c i a l i z a t i o n r e f l e c t s the very d i f f e r e n t n u t r i t i o n a l
and o t h e r f e a t u r e s o f t h e s e p l a n t groups. Crude p r o t e i n l e v e l s i n
woody p l a n t s a r e g e n e r a l l y h i g h e r t h a n i n grasses, b u t t h e
d i g e s t i b i lit y o f p l a n t t i s s u e , i s reduced by h i g h l e v e l s o f
s t r u c t u r a 1 carbohydrates and, i n some instances, by secondary
chemica 1 compounds such as condensed tannins (Cooper and Owen-Smi th,
1985). Thus browsers tend t o be energy-limited whereas grazers are
often protein-limited. Moreover, browsers can experience severe food
shortages during the dry season when woody plants drop t h e i r leaves.

The e f f e c t s o f herbivory depend on ( i l t h e growth form o f the plant;

( i i ) t h e p l a n t p a r t s removed; ( i i i ) t h e i n t e n s i t y , frequency and
season o f use; ( i v ) the growth stage o f the plant; (v) s o i 1 type and
s o i 1 m o i s t u r e c o n d i t i o n s , w h i c h a f f e c t w a t e r and n u t r i e n t
a v a i l a b i l i t y and thereby the p l a n t s capacity t o regrow; and ( v i l the
h i s t o r y o f the plant, p a r t i c u l a r l y the time since a previous
occurrence o f d e f o l i a t i o n by o t h e r h e r b i v o r e s o r f i r e . S i n c e
recovery from defo 1i a t i on is n o t instantaneous, f u t u r e events such
as heavy r a i n f a l l , drought, o r f u r t h e r d e f o l i a t i o n by herbivores and
f i r e can a l s o influence the eventua 1 outcome.
Compensatory g r o w t h i s one o f t h e main responses o f p l a n t s t o
d e f o l i a t i o n . When d e f o l i a t i o n i s low, p l a n t s j u s t compensate by
r e p l a c i n g l o s t t i s s u e . Under moderate l e v e l s o f d e f o l i a t i o n , and
p r o v i d e d t h a t m o i s t u r e and n u t r i e n t c o n d i t i o n s f o r g r o w t h a r e
adequate, some p l a n t s overcompensate f o r amounts removed, leading t o
an increase i n aboveground p l a n t production (McNaughton, 1985). I f a
p l a n t i s severely defoliated, t h e amount o f regrowth i s i n s u f f i c i e n t
t o compensate f o r t h e amounts o f f o l i a g e removed, i n which case
c o n t i n u e d d e f o l i a t i o n WOU l d eventua 1 l y r e s u lt i n t h e p l a n t b e i n g
k i l l e d o r so reduced i n s i z e and s t a t u r e t h a t i t i s u l t i m a t e l y
over looked by herbivores.

Consumption by herbivores accelerates t h e processes o f energy f l o w

and n u t r i e n t c y c l i n g . D e f o l i a t i o n reduces n u t r i e n t and w a t e r
l i m i t a t i o n s on the remaining tissues and stimulates growth. I n some
cases, p h o t o s y n t h e t i c and n u t r i e n t uptake r a t e s a r e i n c r e a s e d
(McNaughton, 1979). By r e d u c i n g a p l a n t ' s t r a n s p i r a t i o n a 1 area,
d e f o l i a t i o n also contributes i n i t i a l l y t o a lower l e v e l o f water-use
and, thereby, t o the conservation o f s o i 1 moisture. This can lead t o
an extension o f the normal growing season (McNaughton, 1985).

N u t r i e n t s t h a t would otherwise be bound up i n t o standing dead p l a n t

m a t t e r and lit t e r a r e r e c y c l e d more r a p i d ly. Furthermore, by
r e c y c l i n g n u t r i e n t s i n c o n c e n t r a t e d form as dung o r u r i n e ,
herbivores contribute t o maintaining a high avai l a b i l i t y o f
n u t r i e n t s f o r plants. Where t h e p o t e n t i a l f o r leaching from the soi 1
i s high, t h i s serves t o keep the n u t r i e n t s c y c l i n g r a p i d l y through
t h e vegetation and surface s o i l , a feature which may be c r u c i a l t o
mai n t a i n i ng the longterm n u t r i e n t status o f these systems (Botkin e t
al., 1981).

Herbivory i n t e r a c t s w i t h f i r e i n both space and time. Many grazers

a r e a t t r a c t e d t o r e c e n t l y b u r n t ground t o f e e d on t h e p o s t - f i r e
regrowth o f grasses (Frost, 1984). This s t i m u l a t i o n o f regrowth a t a
t i m e o f the year when the p l a n t s are usual l y dormant, together w i t h
t h e h i g h e r n u t r i t i o n a l q u a l i t y o f t h e regrowth, i s a m a j o r reason
f o r t h e f r e q u e n t i g n i t i o n o f dry-season f i r e s by p a s t o r a l i s t s i n
t r o p i c a l savannas (Medina pers. comm., M i 1l i g a n and Su le, 1982).
Grazers i n t u r n reduce grass biomass and so l o w e r t h e s h o r t - t e r m
p r o b a b i lit y o f t h e area s u s t a i n i ng a n o t h e r burn. Patchy g r a z i ng
consequent ly causes patchy f i r e s and v i sa versa.

For browsers, dry season f i r e s can reduce the a v a i l a b i l i t y o f food

and cause the animals t o disperse t o other, unburnt, areas (Bel 1 and
Jachmann, 1984). This may increase the browsing pressure on trees i n
these areas (Harrington and Ross, 1974). The degree o f concentration
o r dispersion, and subsequent e f f e c t s such as overuse and trampling,
depend on t h e s i z e o f t h e b u r n r p a t c h i n r e l a t i o n t o t h e amount o f
unburnt vegetati on, and t h e i r r e l a t i v e a t t r a c t i veness t o herbivores.
INTERACTIONS

E f f e c t s o f vegetation on water and n u t r i e n t dynamics

The r e l a t i o n s h i p between s o i 1s and v e g e t a t i o n is i n t e r a c t i v e : t h e

nature o f the soi 1 great l y influences the type o f vegetation t h a t
occurs a t a s i t e , w h i l e t h e vegetation a f f e c t s s o i l p r o p e r t i e s both
d i r e c t l y , through the supply o f organic matter, and i n d i r e c t l y , by
a f f e c t i n g s o i 1 moi s t u r e and temperature r e g i mes, soi 1 chemi stry, and
t h e s t a b i lit y o f t h e s o i 1 surface. The e x t e n t o f p l a n t and l i t t e r
cover a f f e c t s t h e f l u x o f water between t h e atmosphere and t h e s o i l
by i n t e r c e p t i n g and r e d i s t r i b u t i n g r a i n f a l l , enhancing i n f i l t r a t i o n
r a t e s and l o w e r i n g t h e r a t e o f e v a p o r a t i o n o f w a t e r f r o m t h e s o i 1
surface ( K e l l y and Walker, 1976).

A t the same time, p l a n t s deplete the s o i l moisture s t o r e by t a k i n g

up and t r a n s p i r i n g water. The r a t e o f water loss depends p a r t l y on
s o i l w a t e r p o t e n t i a l s , p a r t l y on t h e e v a p o r a t i v e demand o f t h e
atmosphere, and p a r t l y on p l a n t c h a r a c t e r i s t i c s such as t o t a l l e a f
area, p l a n t w a t e r p o t e n t i a l s , t r a n s p i r a t i o n r a t e s and s t o m a t a l
responses m increasi ng water def i c i ts.

There are few data avai l a b l e on t h e t r a n s p i r a t i o n r a t e s o f savanna

t r e e s and grasses. I n m o i s t South American savannas, where t r e e s
have access t o w a t e r t h r o u g h o u t t h e y e a r (Sarmiento e t al., 19851,
t h e t r e e s have o n l y m a r g i n a l l y l o w e r t r a n s p i r a t i o n r a t e s t h a n
grasses (Cioldstein, pers. comm.). Therefore, t h e r e l a t i v e r a t e s a t
which t r e e s and grasses deplete t h e s o i 1 water i n such a system w i 11
depend l a r g e l y on the r e l a t i v e l e a f areas o f t h e two components.

I n t h e d r y savannas, where b o t h t r e e s and grasses e x p e r i e n c e a

seasonal w a t e r d e f i c i t , t h e r e appears t o be a d i f f e r e n c e i n t h e i r
responses t o increasing water s t r e s s (Walter, 1971; Pendle, 1982).
In grasses, t r a n s p i r a t i o n r a t e s appear t o be regulated p r i m a r i l y by
atmospheric evaporative demand. By maki ng o s m o t i c a d j u s t m e n t s t o
l e a f water p o t e n t i a 1s through concentrati ng so lutes i n t h e i r leaves
(Wi lson e t al., 19801, grasses seem able t o maintain r e l a t i v e l y high
r a t e s even a t s o i 1 moisture l e v e l s near w i l t i n g point. Whi l e t h i s
extends t h e p e r i o d o f p h y s i o l o g i c a l a c t i v i t y o f t h e grasses, i t
r e s u l t s i n a r a p i d d e p l e t i o n o f s o i 1 moisture i n the r o o t i n g zone o f
grasses and t h i s eventua 1 ly l e a d s t o l a r g e l e a f - w a t e r d e f ic i t s ,
desiccation and, often, l e a f death.

I n c o n t r a s t , t h e t r a n s p i r a t i o n r a t e s o f t r e e s i n t h e d r y savannas
appear t o be determined more by s o i 1 moisture avai l a b i l i t y than by
atmospheric demand. As t h e s o i l d r i e s out, t h e trees regulate t h e i r
water-use by lowering t r a n s p i r a t i o n r a t e s through stomata 1 c losure.
o r by r e d u c i n g l e a f area t h r o u g h shedding leaves. Trees t h e r e f o r e
deplete the s o i 1 moisture s t o r e less r a p i d l y and completely than do
t h e grasses. The resu l t i s t h a t where r a i n f a l l i s low and grasses
predominate, t h e v e g e t a t i o n aggravates t h e shortage o f w a t e r by
e x t r a c t i n g moisture from below t h e zone i n the soi 1 where i t can be
removed by evaporation a lone. T h i s h e i ghtens t h e c o n t r a s t between
t h e w e t and d r y phases o f t h e s o i 1. On t h e o t h e r hand, where
r a i n f a l l i s high and t r e e growth i s favoured, soi 1 moisture tends t o
be conserved

T r e e s a l s o i n f l u e n c e t h e s t a t u s and d i s t r i b u t i o n o f n u t r i e n t s i n
savanna soi 1s. Woodland regeneration, f o l lowi ng p r o t e c t i o n against
sustained heavy browsing and grazing, r e s u l t s i n an improvement i n
.
s o i 1 n u t r i e n t status (Hatton and Smart, 1984) Soi 1 organic matter,
pH, a v a i l a b l e phosphorus, and t h e l e v e l o f exchangeable c a t i o n s
( e x c e p t i n g Mn) a l 1 increase. T h i s i s because trees, w i t h t h e i r
r e l a t i v e l y deep r o o t systems, are able t o e x t r a c t n u t r i e n t s a t depth
i n the s o i 1 and so counter the e f f e c t s o f leaching (Kellman, 1979).

The h o r i z o n t a l l y - e x t e n s i v e r o o t systems o f t r e e s a 1low them t o

c o n c e n t r a t e n u t r i e n t s f r o m a w i d e area (Van Donselaar-Ten Bokkel
Huinink, 1966; Radwanski and Wickens 1967; F o l d a t s and R u t k i s ,
1975). Consequently, t h e s o i 1s under t r e e s i n savannas genera l l y
have a higher n u t r i e n t status which, together w i t h the moister s o i l s
and more mesic m i c r o c l i m a t e , influences the d i s t r i b u t i o n o f
herbaceous plants, favouring more mesic-adapted, palatable grasses.
This occurs i n both A f r i c a and Venezuela (Kennard and Walker, 1973;
Medina, pers. comm.) b u t n o t i n n o r t h e r n A u s t r a l i a ( M o t t e t al.,
1985) which poses an i n t e r e s t i n g question why.

E f f e c t s o f animals on water and n u t r i e n t dynamics

Animals a f f e c t the s t r u c t u r e o f the s o i 1 and therefore s o i 1 moisture
dynamics i n v a r i o u s ways. Soi 1 macrofauna i n c o r p o r a t e lit t e r and
o t h e r o r g a n i c m a t e r i a l s i n t o t h e s o i l and c o n t r i b u t e t o t h e
s y n t h e s i s o f o r g a n i c CO 1 l o i d s . These C O 1l o i d s a r e i n v o l v e d i n t h e
f o r m a t i o n o f s t a b l e s o i l aggregates w h i c h i n t u r n enhance t h e
aeration, permeabi lity and water-holdi ng capaci t y o f the soi 1. The
network o f underground g a l l e r i e s and burrows formed by soi 1 animals
presumably has s i m i l a r p o s i t i v e e f f e c t s b u t these have n o t been
t h o r o u g h l y i n v e s t i g a t e d (Woods and Sands, 1977). Earthworms and
r e r m i tes b r i n g considerable quanti t i e s o f s o i 1 t o the surface i n the
f o r m o f f a e c a l casts, g a l l e r i e s , and s u r f a c e s h e e t i n g s (Wood and
Sand, 1977; Bagine, 1984; Lave1le, 1983). This improves soi 1 surface
structure, thereby promoting h i gh water i n f i l t r a t i o n rates.
Much o f t h e m a t e r i a l b r o u g h t t o t h e s u r f a c e b y ' t e r m i t e s and
earthworms cornes from the subsoil. This counteracts t h e e f f e c t s o f
i l l u v i a t i o n and leaching, and so c o n t r i b u t e s t o the maintenance o f
u n i f o r m - t e x t u r e d s o i 1 p r o f i l e s (Boyer, 1973; Young, 1976; Josens,
1983; Lave 1le, 1983). Termi t e s a r e p a r t i c u l a r l y i m p o r t a n t i n t h i s
regard as they genera 1l y s e l e c t c lay-sized p a r t i c les i n preference
t o sand when constructing t h e i r nests and g a l l e r i e s (Wood and Sands,
1977). This can have both p o s i t i v e and negative e f f e c t s on the s o i l
moi s t u r e ba lance, depending on r a i n f a 11.

D i r e c t negative e f f e c t s o f animals on soi 1 moisture regimes r e s u l t

from compaction o f t h e surface s o i l and t h e breakdown o f aggregates
t h r o u g h tramp l i n g . T h i s i n c r e a s e s bu l k s o i 1 d e n s i ty, reduces
in f il t r a t i o n r a t e s and l o w e r s t h e w a t e r - h o l d i n g c a p a c i t y o f t h e
s o i 1. Run-off and e r o s i o n increase. I n d i r e c t n e g a t i v e e f f e c t s a r e
m e d i a t e d t h r o u g h t h e i m p a c t o f a n i m a l s on t h e v e g e t a t i o n . High
d e n s i t i e s o f l a r g e ungu l a t e s and invertebrate herbivores such as
h a r v e s t e r t e r m i t e s s i g n i f i c a n t l y reduce p l a n t and lit t e r c o v e r
( K e l l y and Walker, 1976; Lepage, 1981). This increases the exposure
o f t h e s o i 1 s u r f a c e t o r a i n d r o p i m p a c t , sun and wind, and leads t o
more extreme s o i 1 s u r f a c e temperatures, h i g h e r e v a p o r a t i o n r a t e s
and, u l t i m a t e l y , t o t h e s t r u c t u r a l c o l l a p s e and d e f l a t i o n o f t h e
s o i 1 surface. Impermeable surface sea 1s are o f t e n formed, resu l t i n g
i n a c y c l e o f degradation o f reduced i n f i l t r a t i o n rates, increased
r u n o f f , l o w e r s e e d l i ng e s t a b l i s h m e n t , l e s s p l a n t p r o d u c t i o n and
f u r t h e r exposure o f t h e s o i 1 s u r f a c e (Ke 1l y and Wa l k e r , 1976;
Macdonald, 1978; Bridges e t al., 1983; Valentin, 1985).

Local differences i n s o i l type and organic matter production r e s u l t

i n considerable s m a l l scale v a r i a t i o n s i n s o i 1 n u t r i e n t status.
These d i f f e r e n c e s a r e h e i g h t e n e d b y t h e a c t i v i t i e s o f t e r m i tes.
L i t t e r f e e d i n g t e r m i t e s a r e p a r t i c u l a r l y common i n d y s t r o p h i c
savannas where t h e y may consume u p t o 36% o f annual l i t t e r f a l l ,
i n c l u d i n g about 60% o f dead wood and g r a s s l i t t e r p r o d u c t i o n
(Wood a n d Sands, 1977; Ohiagu, 1979b; Josens, 1983). Thus
c o n s i d e r a b l e a m o u n t s o f o r g a n i c m a t t e r and n u t r i e n t s a r e
concentrated i n t e r m i t e nests, from where they are s l o w l y released
i n t o the soi 1 as t h e mounds weather. There have been r e l a t i v e l y few
s t u d i e s o f t h e t u r n o v e r t i m e s o f t e r m i t e mounds. The t u r n o v e r o f
T r i n e r v i termes ermi natus mounds i n a N i g e r i an savanna averages 6.3
y e a r s (Ohiagu, h t i n o t h e r species t h e turnover r i m e s a r e
much longer. The amount o f o r g a n i c m a t t e r and n u t r i e n t s added t o
t h e s o i l each y e a r i s g e n e r a l l y low (Wood and Sands, 1977).
Therefore, w h i l e the concentration in, and subsequent slow release
o f m a t e r i a l f r o m t e r m i t e mounds may conserve o r g a n i c m a t t e r and
secure n u t r i e n t s a g a i n s t l e a c h i n g (Menaut e t al., 19851, i t may
equally l i m i t t h e c y c l i n g o f n u t r i e n t s i n savannas.
Soi 1 moisture and the t r e e : grass equi librium

D i f f e r e n c e s i n s o i l m o i s t u r e r e g i m e s and i n t h e r e l a t i v e
a v a i l a b i l i t y o f m o i s w r e and n u t r i e n t s t o plants are major f a c t o r s
d e t e r m i n i n g t h e wide v a r i a t i o n i n t h e s t r u c t u r e o f savanna
vegetation. Savannas encompass a range o f physiognomic types, from
grassy shrublands a t t h e i n t e r f a c e w i t h deserts, t h r o u g h open
woodlands and t r e e l e s s edaphic grasslands, t o a l m o s t c l o s e d
woodlands w i t h a h e l i o p h y t i c grass understory i n the t r a n s i t i o n zone
t o semi-deciduous and evergreen f o r e s t s (Huntley and Walker, 1982).
The c h a r a c t e r i s t i c f e a t u r e s e r v i n g t o l i n k t h i s d i v e r s i t y o f
vegetation types is the norma 1ly stab l e coexistence o f grasses and
t r e e s , components which i n t h e o t h e r m a j o r biomes t e n d t o r e p l a c e
one another.

To account f o r the balance between woody plants and grasses i n the

d r y savannas, Walter (1971) suggested t h a t the s o i l consists o f two
f u n c t i o n a l l y d i s t i n c t layers: a surface layer i n which grasses, w i t h
t h e i r shallow compact r o o t systems, r e t a i n and have p r i o r access t o
the water entering the soi 1; and a lower subsoi 1 layer t o which t h e
deeper-rooted woody p l a n t s have exclusive access. I n terms o f t h i s
h y p o t h e s i s, t r e e s and grasses can c o e x i s t i n s i t u a t i o n s where t h e
amount o f w a t e r r e g u l a r l y r e a c h i n g t h e subsoi 1 i s j u s t enough t o
s u p p o r t t r e e s b u t i n s u f f i c i e n t t o enable them t o e s t a b l i s h a
c o n t i n u o u s canopy and shade o u t t h e grasses. The h y p o t h e s i s has
been f o r m a l i z e d by Walker e t al. (1981) and Walker and Noy-Meir
(1982).

The l i m i t e d i n f o r m a t i o n a v a i l a b l e on t h e r o o t systems o f savanna

p l a n t s indicates t h a t there is considerab l e v e r t i c a l over lap between
g r a s s and t r e e r o o t s , though t h e maximum d e n s i t y o f grass r o o t s
o c c u r s i n t h e t o p 10 - 20 cm o f t h e s o i l , w h i l e i n t r e e s i t i s
g e n e r a l l y below t h i s (Strang, 1969; Rutherford, 1983; Knoop and
Walker, 1985). However, b o t h have access t o s u r f a c e and subsoi 1
water (Russell, 1966; Strang, 1969; T u n s t a l l and Walker, 1975; Knoop
and Walker, 1985). D e s p i t e t h i s weakness i n one o f t h e m a j o r
assumptions o f t h e model, i t can s t i 11 be v a l i d p r o v i d e d t h a t t h e
grasses and t r e e s a r e each t h e s u p e r i o r c o m p e t i t o r i n d i f f e r e n t
p a r t s o f the s o i l p r o f i l e o r a t d i f f e r e n t tirnes (Knoop and Walker,
1985).

The h i g h e r t r a n s p i r a t i o n r a t e s and more compact r o o t systems o f

grasses s h o u l d g i v e them an advantage i n t h e upper l a y e r s o f t h e
s o i 1. On t h e o t h e r hand, once t r e e s a r e e s t a b l i s h e d , t h e y can
outcompete grasses by shadi ng them and, through bei ng longer- lived,
by gradua1 l y a p p r o p r i a t i n g space i n t h e s o i 1 when it becomes
avai lab le. The lower rates o f water-use o f trees i n t h e d r y savannas
a r e a l s o an advantage s i n c e t h e a v a i l a b l e s o i 1 moi s t u r e i n t h o s e
p a r t s o f t h e s o i 1 p r o f i l e dominated by t r e e r o o t s w i 11 n o t be
depleted so rapidly. This enables trees t o remain a c t i v e f o r longer.
Subsoil moisture levels are determined mainly by the amount o f water
draining through from the surface. The r e l a t i v e abundance o f grass
and trees w i 11 therefore depend on both the amount o f r a i n f a l l and
t h e water-holding capacity o f the topsoil. I n areas o f low rainfal.1,
p a r t i c u l a r l y where t h e w a t e r - h o l d i ng capaci t y o f t h e t o p s o i 1 is
high, most o f t h e i n c o m i n g w a t e r remains near t h e surface. T h i s
a l lows grasses t o reduce the amount o f water eventual l y reaching the
subsoi 1, thereby lim i ti ng indi r e c t ly the growth o f trees (Knoop and
Walker, 1985). Since t h e r e i s a shortage o f a v a i l a b l e m o i s t u r e ,
n e i g h b o u r i n g t r e e s a r e p o t e n t i a l competi t o r s . T h i s i s sometimes
r e f lected i n the r e g u l a r i t y o f t h e i r s p a t i a l d i s t r i b u t i o n s ( S m i t h
and Walker, 1983).

Trees are increasingly favoured as more water reaches the subsoil.

This general ly occurs on deep, porous sands, stony s lopes, fractured
l a t e r i t i c outcrops and on well-drained s o i l s i n regions w i t h high
annual r a i n f a l l (Walter, 1971; Walker and Noy-Meir, 1982; Knoop and
l i a l k e r , 1985). As t r e e d e n s i t i e s increase, grass g r o w t h dec l i n e s
both through shading and through increased cornpetition f o r water and
n u t r i e n t s . Tree removal r e s u l t s i n i n c r e a s e d grass p r o d u c t i o n
(Beale, 1973; Walker e t al., 1972; Dye and Spear, 1982), though
t o t a l abovegr'ound p l a n t production usually declines. I n most cases,
grass production is h i ghest where woody p lants have been comp l e t e ly
c leared, b u t i n some cases it reaches a maximum i n l i g h t l y wooded
communities (Kennard and Walker, 1973). The i n c r e a s e i n g r a s s
p r o d u c t i o n i s o f t e n accompanied by a change i n grass species
c o m p o s i t i o n and a r e d u c t i o n i n t h e abundance o f b e t t e r - q u a l i t y
grasses (Dye and Spear, 1982).

Whi l e competi r i o n f o r water may explain the coexistence o f grasses

and t r e e s i n t h e d r y savannas, i t does n o t adequate l y account f o r
t h e presence o f savanna v e g e t a t i o n under much h i g h e r r a i n f a l l . I n
the moist South American savannas, f o r example, the dominant trees
a r e l a r g e l y evergreen, deep-rooted and do n o t markedly reduce t h e i r
t r a n s p i r a t i o n r a t e s d u r i n g t h e d r y season (Rawitscher, 1948;
Vareschi, 1960; F o l d a t s and R u t k i s , 1975; Sarmiento e t al., 1985).
Most o f t h e species renew t h e i r leaves i n t h e m i d d l e o f t h e d r y
season when c o n d i t i o n s seem l e a s t f a v o u r a b l e f o r l e a f expansion
( M e d i n a , 1982; S a r m i e n t o , 1984). F o r t h e t r e e s t o r e m a i n
p h y s i o l o g i c a l l y a c t i v e a t t h i s time, t h e y must have access t o
adequate moiswre, e i t h e r from groundwater sources o r from moisture
which has accumulated deep i n the soi 1 p r o f i l e during the previous
wet season (Foldats and Rutkis, 1975; Sarmiento, 1984).

A s h o r t a g e o f w a t e r i s t h e r e f o r e u n l i k e l y t o be t h e main factor
lim i c i ng t r e e d e n s i t i e s i n these savannas. Soi 1 n u t r i e n t l e v e 1s
though a r e e x t r e m e l y low (Lopes and Cox, 1977; Sarmiento, 1984).
T h i s i s m a n i f e s t e d i n t h e s l o w g r o w t h and s c l e r o m o r p h i sm o f the
woody vegetation which, together w i t h the large investment made by
these p l a n t s i n t h e i r r o o t systems, l i m i t s the r a t e o f development
o f a closed t r e e canopy. Grasses can therefore c o e x i s t alongside t h e
t r e e s d e s p i t e b e i n g dormant f o r most o f t h e d r y season. Moreover,
because t h e y a r e s h a l l o w - r o o t e d and r e l a t i v e l y f a s t growing, t h e
grasses a r e a b l e t o t a k e up n u t r i e n t s r a p i d l y when t h e s e a r e
minera l i z e d i n the surface s o i 1 during the wet season. This probab ly
f u r t h e r l i m i t s the avai l a b i l i t y o f n u t r i e n t s t o trees.
F i r e may also be a f a c t o r since i t l i m i t s the establishment o f t r e e s
and shrubs and retards the development o f a closed canopy, enabling
grasses t o c o e x i s t a l o n g s i d e t r e e s . P l a n t s on i n f e r t i l e s o i 1s a r e
p a r t i c u l a r l y prone t o f i r e . The slowness w i t h which closed t r e e and
shrub canopies develop enables the grasses, which f u e l the f i r e s , t o
p e r s i s t under t r e e s f o r l o n g e r (Kellman, 1984). Most o f t h e f i r e s
o c c u r d u r i n g t h e d r y season when t h e grasses and o t h e r herbaceous
p l a n t s are dormant and therefore r e l a t i v e l y unaffected by f i r e .
I n contrast, most woody plants, even those species which tend t o be
f i r e - t o l e r a n t as a d u l t s , a r e s e n s i t i v e t o b e i n g b u r n t d u r i n g t h e
e a r l y stages o f establishment and growth, p a r t i c u l a r l y by i n t e n s e
l a t e dry-season f i r e s ( S i l v a and Castro, 1985). The biomass o f
e s t a b l i s h e d woody p l a n t s i s a l s o reduced as f i r e damages t h e
aboveground p a r t s and retards t h e i r growth. F i re, therefore, favours
g r a s s p r o d u c t i o n and t h i s , i n t u r n , f u e l s f u t u r e f i r e s , t h e r e b y
s e t t i n g up a p o s i t i v e feedback which maintains both grass and f i r e
i n t h e system. Under c o n d i t i o n s f a v o u r i n g r a p i d and s u b s t a n t i a l
grass growth, f i r e s may even be s u f f i c i e n t l y intense t o maintain an
open grass land.
Not a l 1 grasslands i n the high r a i n f a l l savannas owe t h e i r o r i g i n t o
r e g u l a r f i r e s . Pure grass lands a l s o o c c u r wherever t h e r e i s poor
s i t e d r a i n a g e and/or where a s h a l l o w s o i 1 p r o f i l e o v e r l i e s an
impermeable c l a y h o r i z o n o r l a t e r i t e l a y e r (Michelmore, 1939;
Sarmiento and Monasterio, 1975; Tinley, 1982). The moisture regime
o f t h e s e s o i 1s f l u c t u a t e s between an excess o f w a t e r d u r i n g t h e
r a i n s and exteme w a t e r d e f i c i t i n t h e d r y season. T h i s does n o t
a f f e c t s h a l l o w - r o o t e d grasses and sedges b u t i s i n i m i c a l t o t h e
growth o f the deeper-rooted trees. I n some areas, such as the llanos
o f c e n t r a l Venezuela, t h e s c a t t e r e d occurrence o f t r e e s r e f l e c t s
l o c a l differences i n soi 1 depth and t h e s t r u c t u r e o f the underlying
l a t e r i t e layer. The trees occur where the s o i 1 i s s u f f i c i e n t l y deep
and porous so t h a t waterlogging doesn't occur, o r where t h e i r r o o t s
are able t o penetrate cracks i n t h e l a t e r i t e and thereby gain access
t o groundwater d u r i n g t h e d r y season ( M o n a s t e r i o and Sarmiento,
1968; San Josb and Farinas, 1983). C o r n p e t i t i o n between t r e e s and
grasses i s n o t a f a c t o r i n t h i s case.
Changes i n t h e abundance o f t r e e s and grasses a t a s i t e i m p l y
d i f f e r e n t i a l morta l i t y o r s u r v i v a l o f i n d i v i d u a l s o f d i f f e r e n t
species. M a t e r s t r e s s , f i r e and h e r b i v o r y appear t o be t h e main
causes o f s e e d l i n g m o r t a l i t y ( S i lva, 1973; Penning de V r i e s and
D j i t k e , 1982; Belsky, 1984). L i t t l e i s known about the processes o f
seed g e r m i n a t i o n and s e e d l i ng establishment i n savannas, o r about
the r o l e o f competi tion, p a r t i c u l a r l y from estab lished plants. Both
grasses and t r e e s can i n h i b i t t h e e s t a b l i s h m e n t o f woody p l a n t
seedlings, b u t the precise mechanisms are n o t known (Strang, 1969;
Knoop and Walker, 1985). Competition f o r water and n u t r i e n t s may be
i n d i r e c t l y invo lved. Since sma 1l e r seedli ngs are general ly the most
s u s c e p t i b l e t o w a t e r s t r e s s , o r t o b e i n g b u r n t o r eaten, any
l i m i t a t i o n s on g r o w t h w i 11 i n c r e a s e t h e r i s k o f m o r t a l i t y . It i s
therefore important t o learn what f a c t o r s favour the recruitment o f
new i n d i v i d u a l s t o a p o p u l a t i o n . Does an i n c r e a s e i n p e r c o l a t i o n
f a v o u r an i n c r e a s e i n seed p r o d u c t i o n o f a d u l t p l a n t s , o r does i t
r e s u l t i n improved s e e d l i n g g r o w t h and s u r v i v a l ? S t u d i e s o f t h e
p o p u l a t i o n b i o l o g y o f s e l e c t e d t r e e and grass species w i l l be
essentia 1 i n order t o understand the mechani sms invo lved.

Species composition and coexistence

I n savannas there i s generally an inverse r e l a t i o n s h i p between p l a n t
species richness and the moisture and n u t r i e n t status o f the soil.
Well drained, sandy dystrophic s o i 1s can have upto t w i c e the number
o f s p e c i e s c o e x i s t i n g a t a s i t e compared w i t h s i t e s on e u t r o p h i c ,
poor l y drained c lays (Frost, pers. coma.). The under l y i n g mechani sms
g i v i n g r i se t o t h i s r e l a t i o n s h i p have n o t been i n v e s t i g a t e d . Two,
n o t mutua 1l y exc l u s i v e exp l a n a t i o n s seem p l a u s i b l e . F i r s t , low
n u t r i e n t a v a i l a b i l i t y r e s t r i c t s p l a n t growth and reproduction w i t h
the r e s u l t t h a t t h e r a t e a t which populations approach competitive
e q u i l i b r i u m i s reduced. T h i s may l i m i t t h e dominance o f c e r t a i n
s p e c i e s and enable a g r e a t e r number o f p o t e n t i a l c o m p e t i t o r s t o
coexi st. I n v a r i ab l e envi ronments such as savannas, t h i s coexistence
may be prolonged i n d e f i n i t e l y i f t h e competi ti veness o f d i f f e r e n t
s p e c i e s a l t e r n a t e s w i t h t h e changes i n t h e environment. I n t h i s
respect, i t i s worth noting t h a t whi l e the d i v e r s i t y o f species on
c layey s o i 1s i s o f t e n low, t h e changes i n c o m p o s i t i o n a r e more
marked, r e f l e c t i n g the greater amplitude o f change i n conditions on
clays per u n i t change i n r a i n f a l l (O'Connor, 1985).
Secondly, i f n u t r i e n t l i m i t a t i o n i s o f prime importance to p l a n t s
growi ng on dystrophi c s o i ls, then se l e c t i on shou l d favour a t t r i butes
w h i c h enhance a p l a n t ' s c a p a c i t y t o t a k e up and s t o r e n u t r i e n t s .
These may i n v o l v e s p e c i a l i z a t i o n on a narrow a r r a y o f m i c r o s i t e s
w i t h p a r t i c u l a r s o i 1 c h a r a c t e r i s t i c s and n u t r i e n t s t a t u s . Where
there is considerable heterogenei t y i n the s o i 1, t h i s WOU l d provide
more opportunities f o r coexistence and, thereby, a higher diversity.
The spatia 1 heterogenei ty o f savanna soi 1s is marked. Features such
a s -t e r m i t a r i a , l a r g e mammal a c t i v i t y s i t e s (dung s i t e s , burrows
etc.), t h e p a t c h y occurrence o f f i r e and grazing, and i n h e r e n t
d i scontinui t i e s i n the vegetation (bushc lumps, open and under-tree
sites), a l 1 contribute t o d i fferences i n soi 1 characteristics.
Marked differences i n species composition and abundance can r e a d i l y
be observed over s h o r t distances i n r e l a t i o n t o t h i s heterogeneity,
s u g g e s t i n g t h a t o p t i m a l s i t e s f o r d i f f e r e n t species are generally
l i m i t e d t o one o r a few s i t e s along an edaphic gradient. A t the same
t i m e , s i n c e e s t a b l i s h m e n t appears t o be a c r i t i c a l stage f o r many
plants, some o f these d i f f e r e n c e s may r e f l e c t processes o p e r a t i n g
p r i m a r i ly a t the seedling stage.

The marked e f f e c t t h a t changes i n s o i l c h e m i s t r y have on grass

species composi t i o n is c l e a r l y indicated by the changes which occur
i n response t o f e r t i l i z a t i o n ( M i l l s , 1964; O'Connor, 1985). The
e x t e n t o f these changes depends b o t h on s o i l type, b e i n g more
pronounced on d y s t r o p h i c sands t h a n on e u t r o p h i c c l a y s , and on
r a i n f a 11, being more apparent i n the wetter savannas, para 1l e 1l i n g
t h e observed differences i n species richness.
There a r e a l s o wide d i f f e r e n c e s i n t h e moi s t u r e r e q u i r e m e n t s o f
savanna plants. This i s r e f lected i n the d i f f e r e n t i a l d i s t r i b u t i o n
o f species along s o i 1 moisture gradients ( S i l v a and Sarmiento, 1976;
Yeaton e t al., i n prep.); i n d i f f e r e n c e s i n water-use by species
g r o w i n g under t h e same c l i m a t i c c o n d i t i o n s ( F o l d a t s and Rutkis,
1975; Medina, 1982); and by s h o r t - t e r m changes i n b o t h woody and
herbaceous community composition occurring during periods o f above-
and below-average r a i n f a 11 (Poupon, 1979; S i ngh and Krishnamurthy,
1981; O'Connor, 1985). These changes i n s p e c i e s c o m p o s i t i o n a r e
usua 1l y greater on c layey than on sandy s o i ls, r e f l e c t i n g the more
extreme soi 1 moisture regimes o f clays.

The composition o f the herbaceous layer i n dry savannas appears t o

be a f f e c t e d p r i m a r i ly by year t o year and longer-term v a r i a t i o n s i n
r a i n f a l l . The e f f e c t s o f g r a z i n g o r f i r e become more i m p o r t a n t as
mean annual r a i n f a l l i n c r e a s e s and i t s v a r i a b i lit y dec l i n e s . F o r
example, i n the Australian moi s t t r o p i c a 1 and subtropica 1 t a l l g r a s s
savannas, introduced ungu lates ( c a t t l e and sheep) and increased f i r e
frequencies have apparently caused a change from a Themeda a u s t r a l i s
dominant understory t o one dominated by Heteropogon contortus (Shaw
and Bisset, 1955).

Pheno logy and coexi stence

Seasona 1 v a r i a t i o n s i n w a t e r and n u t r i e n t a v a i l a b i 1it y , t o g e t h e r

w i t h c o œ p e t i t i o n f r o m o t h e r p l a n t s , and s t r e s s e s imposed by t h e
annual d r y season, f i r e and h e r b i v o r y , c r e a t e a s p a t i a l l y and
temporal ly s h i f t i n g mosaic o f chal lenges and opportuni t i e s , which i s
r e f l e c t e d i n t h e wide v a r i e t y o f phenological behaviour found among
savanna p l a n t s . As many as 15 f u n c t i o n a l l y d i s t i n c t p h e n o l o g i c a l
groups have been recognized, based on t h e seasonali ty/aseasona lity
o f carbon assimilation, continuous versus p e r i o d i c shoot growth, and
t h e ti me o f f lowering (Sarmiento and Monasterio, 1983).

Herbaceous p l a n t s have t h e w i dest v a r i e t y o f pheno logica 1 responses

(Menaut and c h a r , 1979; S a r m i e n t o and Monasterio, 1983; Singh e t
al., 1985). Most s p e c i e s a r e p e r e n n i a l , becoming dormant o r semi-
dormant d u r i n g t h e d r y season. Growth d u r i n g t h e w e t season i s
r a p i d, p a r t i c u l a r l y among grasses. F l o w e r i n g and seed s e t usua 1 ly
t a k e p l a c e l a t e r i n t h e w e t season b u t t h e r e a r e w i d e d i f f e r e n c e s
between s p e c i e s i n t h i s r e s p e c t (Menaut and C k a r , 1979; Singh e t
al., 19851. Some s p e c i e s f l o u e r i n response t o e a r l y r a i n s , o t h e r s
i n response t o l a t e r a i n s . Many o f t h e p e r e n n i a l f o r b s f l o w e r
t o w a r d s t h e end o f t h e d r y season o r a t b e g i n n i n g o f t h e r a i n s .
Flowering and seed dispersa1 i n some o f these species i s i n i t i a t e d
by f i r e (Coutinho, 1982; Menaut and C h a r , 1979). Most species,
though, reproduce on a f i x e d schedule, i r r e s p e c t i v e o f when the r a i n
fa I l s o r f i r e s burn. A few s p e c i e s grow c o n t i n u o u s l y ; o t h e r s a r e
opportuni sts, growi ng whenever conditions are favourab le.

A l a r g e number o f annuals o c c u r i n some savannas. These s p e c i e s

d i f f e r widely i n t h e i r t i m i n g and r a t e o f development, and include
some s p e c i e s w h i c h a r e capable o f c o m p l e t i n g t h e i r l i f e c y c l e s
r a p i d l y and opportuni s t i c a 1 ly whenever conditions are sui tab l e (van
Donse laar-Ten Bokke 1 Hui nink, 1966; Menaut and C k a r , 1979;
Sarmiento, 1984). I n a r i d savannas, the differences i n germination
t i m e r e f l e c t responses t o t w o opposing s e l e c t i o n pressures: t h e
advantage o f r a p i d germination enabling i n d i v i d u a 1s t o e x p l o i t f u 1l y
t h e b r i e f f l u s h o f w a t e r and n u t r i e n t s a t t h e s t a r t o f t h e s h o r t
g r o w i n g season, and t h e r i s k t h a t subsequent r a i n s w i 11 f a i 1 and
t h a t t h e p l a n t s w i 11 become d e s s i c a t e d b e f o r e t h e y can r e p r o d u c e
(Penning de Vries and D j i t k e , 1982). Given t h e inherent v a r i a b i l i t y
o f savanna environments, p a r t i c u l a r ly t h e stochastic nature o f t h e
r a i n f a 11, the d i v e r s i ty i n pheno logy found among herbaceous savanna
p l a n t s is probab ly a key feature promoting t h e i r coexistence.

The phenology o f woody p l a n t s i s much l e s s v a r i a b l e . Most o f t h e

species, i n A f r i c a n , A u s t r a l i a n and I n d i a n savannas a r e deciduous
(Malaisse, 1974; Menaut, 1983; Sarmiento and Monasterio, 1983; M o t t
e t al., 19851. They shed t h e i r leaves d u r i n g t h e d r y season, r e m a i n
l e a f l e s s f o r a few weeks t o months, then produce new leaves p r i o r t o
o r a t t h e b e g i n n i n g o f t h e f o l l o w i n g w e t season. The t i m e o f l e a f
f a 1 1 appears t o be r e l a t e d t o w a t e r s t r e s s , t h e t r e e s r e t a i n i n g
t h e i r leaves f o r longer i n years o f high r a i n f a l l . Some species are
o n l y b r i e f ly deciduous, the o l d leaves f a l l i n g j u s t before the new
ones emerge a t t h e s t a r t o f t h e annual rains. Leaf-out i n deciduous
species c o i n c i des w i t h an increase i n both mean dai ly temperature
and photoperiod (Rutherford and Panagos, 1982).
L e a f p r o d u c t i o n i n t h e deciduous s p e c i e s o f t h e m o i s t savannas i s
l a r g e l y determini s t i c . This and annual shoot growth are apparently
derived from carbohydrate and n u t r i e n t reserves stored i n t h e plant.
The g r e a t e r p a r t o f t h e w e t season i s devoted t o p h o t o s y n t h a t e
p r o d u c t i o n and t h e r e p l e n i s h m e n t o f t h e s e reserves, as we 11 as t o
r a d i a l growth and r o o t extension (Rutherford and Panagos, 1982). I n
contrast, i n the d r i e r savannas, shoot growth i s indeterminate and
depends on conditions during the c u r r e n t growing season.

Evergreen savanna t r e e s and shrubs a r e o n l y common i n t h e h i g h

r a i n f a l l savannas o f South America. These species, w h i c h a r e deep
r o o t e d and g e n e r a l l y appear t o have access t o an adequate w a t e r
supply, replace t h e i r f o l i a g e during the middle o f t h e dry season.
T h i s f a c i l i t a t e s t h e n u t r i e n t economy o f t h e s e t r e e s i n t w o ways.
F i r s t , i t enables n u t r i e n t s t o be withdrawn from senescing leaves
and t o be r e a 1 l o c a t e d i m m e d i a t e l y t o t h e new l y d e v e l o p i n g leaves.
Secondly, because the expansion o f the new leaves, which are h i g h l y
leachable, takes place during t h e dry season, t h e r i s k o f n u t r i e n t
loss through leaching i s reduced (Sarmiento e t al., 1985).

Flowering and f r u i x i ng are a l s o strongly seasona 1. I n Austra l i a n and

South American savannas, f lowering occurs mainly during middle o f
t h e d r y season and i s frequently associated w i t h d e f o l i a t i o n by f i r e
o r d r o u g h t (Coutinho, 1982; M o t t e t al., 1985). F r u i t s mature
r a p i d l y and t h e seeds o f most s p e c i e s a r e d i s p e r s e d d u r i n g t h e
f o 1 l o w i ng w e t season (Sarmiento, 1984). I n A f r i c a n savannas,
f l o w e r i n g i s concentrated a t the end o f t h e d r y season and beginning
o f t h e w e t season, o c c u r r i n g a t t h e same t i m e as l e a f - o u t . F r u i t s
m a t u r e d u r i n g t h e w e t season and a r e d i s p e r s e d t h r o u g h o u t t h e
f o l l o w i ng dry season (Hopkins, 1970; Ma laisse, 1974; Menaut, 1983).

P l a n t production
O r g a n i c i n a t t e r p r o d u c t i o n and p l a n t q u a l i t y i n savannas depend on
t h e t o t a l amount and seasonal d i s t r i b u t i o n o f r a i n f a l l , and on t h e
a v a i l a b i lit y o f n u t r i e n t s , p a r t i c u l a r l y n i t r o g e n and phosphorus
(San Jose and Medina, 1976; Penning de Vries and D j i t b e , 1982; M o t t
e t al., 1985). The influence o f water avai l a b i l i t y i s most apparent
i n t h e d r i e r savannas, as i n d i c a t e d by t h e p o s i t i v e c o r r e l a t i o n s
between annua 1 r a i n f a 11 and, f o r examp le, herbaceous aboveground
biomass ( W a l t e r 1971; San J o s é a n d Medina, 1976; R u t h e r f o r d , 1981;
Deshmukh, 1984; Singh e t al., 1985). However, t h e r e l a t i o n s h i p i s
o n l y very general and i s a f f e c t e d by f a c t o r s such as t h e d u r a t i o n o f
t h e w e t season, s o i 1 t y p e and t e x t u r e , n u t r i e n t a v a i l a b i l i t y ,
t e m p e r a t u r e , f i r e and s p e c i e s c o m p o s i t i o n (San Jose and Medina,
1975; Rutherford, 1981; Penning de Vries and D j i t b e , 1982; Singh e t
al., 1985).
The d i f f e r e n t patterns o f p l a n t production on clayey and sandy s o i l s
o c c u r r i ng a long r a i n f a l 1 gradients, o r i n response t o f l u c t u a t i o n s
i n annual r a i n f a 11 a t a s i te, c l e a r l y i1l u s t r a t e s the influence o f
s o i 1 t e x t u r e on water avai l a b i lity and production. When r a i n f a l l i s
low, p r o d u c t i o n on c l a y s may be as low as, o r even l o u e r t h a n t h a t
on sandy soi 1s under the same r a i n f a l l , despite the generally higher
n u t r i e n t s t a t u s o f clays. However, p r o d u c t i o n on c l a y s i n c r e a s e s
more r a p i d l y w i t h i n c r e a s i n g r a i n f a l l (Dye and Spear, 1982). The
r e s u l t i s t h a t , i n r e l a t i o n t o t h e same f l u c t u a t i o n s i n r a i n f a l l ,
production on clayey s o i 1s i s much more v a r i a b l e than on sands.

As r a i n f a l l and the length o f the n e t season increase, water becomes

less l i m i t i n g and other factors, such a low n u t r i e n t a v a i l a b i l i t y ,
begin t o l i m i t production (Mott e t al., 1985). For example, grasses
g r o w i n g on i n h e r e n t l y f e r t i l e s o i l s , o r on ones which have been
a r t i f i c i a l ly f e r t i lized, are much more productive than those growing
on i n f e r t i l e o r u n f e r t i l i z e d s i tes (Mi 1ls, 1968; San José and Garcia
Miragaya, 1981; Penning de Vries and D j i t k e , 1982). The e f f e c t s are
more pronounced when water i s n o t l i m i t i n g (Donaldson e t al., 1984).
I m p r o v e d s o i 1 f e r t i l i t y i n c r e a s e s t h e use o f w a t e r by t h e
vegetation, improves water-use e f f i c i e n c i e s , and thus contributes t o
h i gher production. However, the avai lab l e soi 1 moi sture i s used up
more r a p i d l y , t h e r e b y s h o r t e n i n g t h e p e r i o d f o r a c t i v e g r o w t h and
increasi ng the r isk o f physi O logica 1 drought.
The l e v e l o f p r o d u c t i o n i s s t o n g l y i n f luenced by t h e r e l a t i v e
a v a i l a b i l i t y o f n i t r o g e n and phosphorus. A d d i t i o n s o f n i t r o g e n o r
phosphorus alone do n o t r e s u l t i n s i g n i f i c a n t increases i n biomass
production unless the other n u t r i e n t i s already present i n excess.
N i trogen-fixing legumes, f o r examp le, show a p o s i t i v e response t o
t h e a d d i t i o n o f phosphorus a l o n e (Penning de V r i e s and D j i t b e ,
1982). However, when n i t r o g e n and p h o s p h o r u s a r e added
simultaneously there i s a massive increase i n biomass p r o d u c t i o n ,
p a r t i c u l a r l y o f grasses (Norman, 1962; Medina e t al., 1978; San José
and Garcia Miragaya, 1981; Penning de Vries and D j i t k e , 1982; M o t t
e t al., 1985).
Overall, organic matter production i n savannas i s l i m i t e d mainly by
n i t r o g e n supply. The grasses o f t h e humid, d y s t r o p h i c savannas o f
South America, f o r example, seem t o be adapted t o low phosphorus
a v a i l a b i l i t y . F e r t i l i z a t i o n e x p e r i m e n t s conducted i n A u s t r a l i a
(Norman, 19621, South A f r i c a (Weinmann, 1938) and Venezuela (Medina
e t al., 1978; San Josb and Garcia-Miragaya, 1981) show t h a t when
n i t r o g e n i s l i m i ted, f e r t i l i z a t i o n w i t h phosphorus i n c r e a s e s t h e
phosphorus content o f the newly produced aboveground biomass. This
i s n o t r e t r a n s l o c a t e d b e f o r e canopy d r i e s a t t h e end o f t h e r a i n y
season (Medina, i n press).
Low n u t r i e n t a v a i l a b i l i t y i s n o t c o n f i n e d t o t h e w e t t e r savannas.
There i s a h i g h p o t e n t i a l f o r n u t r i e n t l i m i t a t i o n i n s i t u a t i o n s
where growth must be completed i n a r e l a t i v e s h o r t period. Rapidly
g r o w i n g p l a n t s have a h i g h n u t r i e n t demand and can t e m p o r a r i l y
deplete the s o i 1 o f avai l a b l e nutrients, p a r t i c u l a r l y where n u t r i e n t
m i n e r a l i z a t i o n i s i n t e r m i t t e n t . This i s an important c o n s t r a i n t i n
a r i d savannas where the growing season i s s h o r t and where the stock
o f avai l a b l e n u t r i e n t s i s sustained by m i n e r a l i z a t i o n o f a l i m i t e d
amount o f s o i 1 organic matter (Penning de Vries and D j i t b e , 1982).
Differences i n t h e times o f peak n u t r i e n t a v a i l a b i l i t y add a f u r t h e r
c o m p l i c a t i o n . I n t h e Sahe 1, f o r examp le, n i t r o g e n m i n e r a l i z a t i o n
peaks a t the s t a r t o f the wet season when the p l a n t s are l i m i t e d by
a lack o f phosphorus. Later, as t h e wet season progresses, n i t r o g e n
i s increasingly immobilized by the soi 1 b i o t a and t h i s eventually
l i m i t s p l a n t growth (Penning de Vries and D j i t b e , 1982).

The i n c r e a s i ng n u t r i e n t c o n s t r a i n t s faced by savanna p l a n t s through

t h e growing season are c l e a r l y seen i n the seasonal changes i n p l a n t
q u a l i t y (Shaw and Bisset, 1955; Afolayan and Fafunsho, 1978; M o t t e t
al., 1981; Penning de V r i e s and D j i t b e , 1982). N i t r o g e n and
phosphorus c o n t e n t s i n leaves a r e h i g h a t t h e s t a r t o f t h e w e t
season and decline progressively as t h e p l a n t grows. P l a n t q u a l i t y
is e s p e c i a l l y low d u r i ng t h e d r y season and and t h i s becomes an
i m p o r t a n t c o n ~ t r ani t on secondary p r o d u c t i o n . Average p a s t u r e
q u a l i t y i s o f t e n below t h e minimum maintenance levels f o r domestic
l i v e s t o c k and w i l d ungulates. The a n i m a l s compensate f o r t h i s
d e c l i ne t o some e x t e n t by foraging s e l e c t i v e l y on those components
i n t h e h e r b l a y e r w h i c h have h i g h e r t h a n average f o r a g e q u a l i t y
(Bremen and de W i t , 1983). Nevertheless, t h e y s t i 11 e x p e r i e n c e a
loss i n decline i n body condition.

Attempts t o overcome these d e f i c i e n c i e s inc lude t h e use o f m i n e r a l

supp lements (urea/molasses m i x t u r e s ) , c r o p r e s idues, f o d d e r
reserves, and the improvment o f pasture q u a l i t y by i n t r o d u c i n g p l a n t
s p e c i e s ( p r i n c i p a l l y legumes) w h i c h w i 11 p r o v i d e t h e necessary
e l e m e n t s ( T o t h i 11 e t al., 1985). Supplementation generally r e s u l t s
i n heavier s t o c k i ng rates, increased natura 1 pasture u t i l i z a t i on and
reduced i n c i d e n c e o f f i r e . The i n t r o d u c t i o n o f a l i e n s p e c i e s i n t o
n a t u r a l pastures r e q u i r e s the a d d i t i o n o f non-nitrogenous
f e r t i l i z e r s , u s u a l l y superphosphate. T h i s o f t e n r e s u l t s i n h i g h e r
s o i 1 f e r t i lit y , improved f o r a g e q u a l i ty, and i n c r e a s e d c a r r y i ng
c a p a c i t i e s f o r l i v e s t o c k . I n areas w i t h a long, h a r s h d r y season,
t h i s can cause a breakdown i n the s t a b i l i t y o f the herbaceous layer
w i t h annual rasses and weeds replacing t h e n a t i v e perennia 1s (Mott
3.
e t al., 1981 The i n f l u e n c e o f t h e s e management a c t i o n s on t h e
longterm s t a b i lity o f these systems needs t o be investigated.
The e f f e c t o f f i r e on herbaceous production depends both on the t i m e
o f burni ng and on s o i 1 moi s t u r e avai l a b i lity. F i r e tends t o increase
grass production i n high r a i n f a l l areas w i t h a s h o r t dry season, b u t
i n d r i e r a r e a s p r o d u c t i o n i s g e n e r a l l y reduced r e l a t i v e t o
neighbouring unburnt areas (West, 1965; San José and Medina, 1975).
The season o f b u r n has an i m p o r t a n t b e a r i n g on t h e outcome. E a r l y
d r y season f i r e s induce p lants t o f l u s h a t a ti me w hen s o i 1 moi s t u r e
l e v e l s are already declining. This regrowth r a p i d l y depletes the
r e m a i n i ng s o i 1 moi s t u r e and t h e t i 1l e r s do n o t s u r v i v e . The
p o t e n t i a l stimulus t o production is therefore n o t sustai ned. Growth
a t the s t a r t o f t h e f o l l o w i n g wet season i s i n i t i a t e d simultaneously
i n b o t h b u r n t and u n b u r n t p l a n t s . B u r n i n g a t , o r soon a f t e r t h e
: s t a r t o f the f i r s t r a i n s general l y has the same outcome except where
growth i n unburnt p l a n t s i s l i m i t e d by an accumulation o f standing
dead
,. matter.

i n c o n t r a s t , d e f o l i a t i o n by f i r e d u r i n g t h e l a t t e r p a r t o f t h e d r y
season r e s u l t s i n e q u i l i b r a t i o n o f p l a n t and s o i 1 water potentials,
a l l o w i n g p l a n t s t o grow. P r o v i d e d t h a t t h e s o i 1 m o i s t u r e s t o r e i s
replenished by e a r l y wet season r a i n s before i t i s again depleted by
t h e grasses, t h i s e a r l y s t a r t t o t h e g r o w i n g season r e s u l t s i n a
h i g h e r p r o d u c t i o n by b u r n t p l a n t s (San Jose and Medina, 1975).
Depending on the amount o f herbivory, the dry season standing crop
o f g r a s s on r e g u l a r l y b u r n t areas can be h i g h e r t h a n on u n b u r n t
plots. This increases the probabi l i t y o f f i r e , s e t t i n g up a p o s i t i v e
feedback loop t h a t serves t o maintain a high f i r e frequency and high
g r a s s production.

Plant q u a l i t y

Primary production i n savannas has three possible fates: i t can be

consumed by herbivores, i t can be burnt, o r i t can senesce, d i e and
decompose. The amount of p l a n t matter incorporated d i r e c t l y i n t o the
l i t t e r and s o i 1 depends on the e x t e n t o f both herbivory and f i r e . I n
turn, the incidence o f f i r e i s largely a function o f the d r y season
standing crop o f grass, a product o f the l e v e l o f production during
t h e preceding wet season and the i n t e n s i t y o f herbivory. The major
d e t e r m i n a n t o f h e r b i v o r y appears t o be p l a n t q u a l i t y ( B e l 1, 1982,
1984).

P l a n t q u a l i t y , i n terms o f the s u i t a b i l i t y o f p l a n t matter as food

f o r organisms f u r t h e r along the food chain, can best be defined as
t h e r a t i o o f a s s i m i l a b l e p l a n t m a t t e r (main l y p r o t e i n - b a s e d
compounds and soluble carbohydrates) t o the amounts o f unusab l e and
i n h i b i t o r y m a t e r i a l i n a p l a n t (mainly f i b r e , l i g n i n and secondary
c h e m i c a l compounds), r a t h e r t h a n as t h e a b s o l u t e amount o f c r u d e
p r o t e i n and soluble carbohydrate present i n a p l a n t (Bell, 1981).
The a v a i l a b i l i t y o f s o i l w a t e r and n u t r i e n t s such as N and P
i n f l u e n c e p l a n t q u a l i ty by a l t e r i n g t h e r e l a t i v e p r o d u c t i o n o f
c y t o p l a s m i c and s t r u c t u r a l components.Tota1 biomass and p r o t e i n
p r o d u c t i o n b o t h i n c r e a s e w i t h i n c r e a s i n g w a t e r and n u t r i e n t
avai l a b i lity (San José and Medi na, 1976; Medi na e t al., 1978; Be 11,
1981, 1982; Penni ng de V r i e s and D j ithe, 1982; Breman and de W i t ,
1983). However, t h e p r o d u c t i o n o f carbon-based ,compounds such as
f i b r e i s l e s s a f f e c t e d by a shortage o f n u t r i e n t s t h a n i s p r o t e i n
production, so t h a t where n u t r i e n t s are l i m i t e d r e l a t i v e t o water,
p l a n t q u a l i t y i s low. On t h e o t h e r hand, p r o t e i n p r o d u c t i o n i s
s t i m u l a t e d more t h a n f i b r e p r o d u c t i o n by an i n c r e a s e i n t h e
a v a i l a b i l i t y o f n u t r i e n t s . T h i s i n c r e a s e may be a b s o l u t e (eg.
through f e r t i l i z a t i o n ) o r r e l a t i v e (eg. through a decrease i n water
avai l a b i lity.

P l a n t q u a l i t y , t h e r e f o r e , w i 11 be a f f e c t e d by any process which

a l t e r s the r e l a t i v e avai l a b i l i t y o f water and n u t r i e n t s t o plants.
F o r i n s t a n c e , low r a i n f a l l , low i n f i l t r a t i o n r a t e s caused by s o i 1
capping, low p l a n t cover, o r d e g r a d a t i o n o f t h e s o i l s u r f a c e b y
herbivore trampling, a l 1 r e s u l t i n low water avai l a b i lity. This, i n
t u r n , r e s u l t s i n t h e p r o d u c t i o n o f a sma 11 amount o f good q u a l i t y
p l a n t biomass w h i c h may r e s u l t i n a g r e a t e r p r o p o r t i o n o f t h e
v e g e t a t i on b e i ng consumed, l e s s p l a n t cover, more t r a m p ling and
i n c r e a s e d s o i 1 d e g r a d a t i o n . I n c o n t r a s t , in c r e a s i n g w a t e r
a v a i l a b i l i t y r e s u l t s i n t h e p r o d u c t i o n o f more p l a n t biomass o f
i n c r e a s i n g l y poor q u a l i t y (Breman and de W i t , 19831, l o w e r a n i m a l
consumption, and an increasi ng proportion o f the avai l a b l e n u t r i e n t s
being concentrated i n the plants. This leads t o a lower avai l a b i lity
o f n u t r i e n t s i n t h e s o i l , l o w e r p l a n t q u a l i t y and even l e s s
consumption.

Vegetati on dynamics

The r e l a t i o n s h i p between w a t e r and n u t r i e n t a v a i l a b i l i t y a t

d i f f e r e n t layers i n the soil, competition between p l a n t s f o r these
resources, and t h e r e s u l t i n g p a t t e r n s o f p l a n t p r o d u c t i o n and
q u a l i t y , a r e t h e key t o u n d e r s t a n d i n g t h e dynamics o f savanna
v e g e t a t i o n i n r e l a t i o n t o h e r b i v o r y and f i r e ( B e l l , 1981, 1982,
1984). For eastern, c e n t r a l and southern A f r i c a n savannas, f o u r main
patterns o f herbivory, f i r e and vegetation dynamics emerge, based on
the r e l a t i v e avai l a b i l i t y o f soi 1 moisture and s o i 1 n u t r i e n t s (Bell,
1984). W i t h t h e e x c e p t i o n o f t h e numbers and i m p a c t o f l a r g e
herbivore populations, which are characteri s t i c o f these savannas,
t h e patterns appear t o have counterparts i n other savanna regions.

I n areas o f low w a t e r a v a i l a b i l i t y and h i g h n u t r i e n t supply, t h e

v e g e t a t i o n c o n s i s t s l a r g e l y o f h i g h q u a l i t y g r a s s l a n d s and open
woodlands. Examples include t h e S e r e n g e t i and Amboseli r e g i o n s o f
East Africa, p a r t s o f the Deccan o f I n d i a and the Astrebla-dominated
g r a s s l a n d s on t h e Bark l y Tab l e l a n d s o f A u s t r a l i a . I n A f r i c a t h e s e
areas support a high biomass and wide d i v e r s i t y o f indigenous large
herbivores i n which smaller-bodied species r e q u i r i n g h i gh qua l i t y
d i e t s predominate. Where these species a r e now absent, t h e r e i s
usua 1ly an equiva l e n t biomass o f domestic livestock.

A l a r g e p r o p o r t i o n o f t h e annual above-ground p r o d u c t i o n i s
consumed. F o r example, on t h e S e r e n g e t i p l a i n s , an average o f 66%,
and u p t o 94%, o f n e t aboveground r i m a r y p r o d u c t i o n i s consumed
f
annual ly by large herbivores alone McNaughton, 1985). I n t h i s case,
t h e a n i m a l s a r e i n d u c i n g s u b s t a n t i a 1 compensatory g r o w t h i n t h e
p l a n t s thereby c r e a t i ng a h i gh n u t r i e n t demand and promoting h i gh
n u t r i e n t uptake r a t e s ( ~ ~ N a u g h t o n1983b, , 1985). Because o f t h i s
compensatory growth, the energy f low t o consumers is a d d i t i v e and
can, i n some cases, be m a i n t a i n e d w i t h l i t t l e o r no r e d u c t i o n i n
p l a n t biomass (McNaughton e t al., 1983).

R e g u l a r d e f o l i a t i o n a p p e a r s t o be one o f t h e m a i n f a c t o r s
maintaining the high number o f coexisting grass species. Changes i n
s p e c i e s c o m p o s i t i o n and a d e c l i n e i n d i v e r s i t y come about when an
area i s protected from grazing o r f i r e . Para1l e 1 changes, i n v o l v i n g
a d i f f e r e n t s u i t e o f species, occur i n areas t h a t a r e exposed t o
very heavy o r very f requent defo 1ia t i on (McNaughton, 1979, 1983a).
However, t h i s does n o t n e c e s s a r i l y i m p l y t h a t t h e most d i v e r s e
community is t h e most moderate ly grazed (McNaughton, 1983a1, though
t h i s can happen (Singh, 1976). O v e r a l l , f i r e i s p r o b a b l y l e s s
important than grazing, p a r t l y because i t i s a dry season phenomenon
occurring a t a t i m e when the grasses are dormant, and p a r t l y because
t h e l e v e l s o f consumption a r e u s u a l l y so h i g h t h a t t h e r e i s
i n s u f f i c i e n t g r a s s biomass l e f t d u r i n g t h e d r y season t o f u e l a
f i r e . The i n c i d e n c e o f f i r e i s t h e r e f o r e low, o t h e r t h a n a f t e r
exceptional l y wet years. Overa 11, these systems can be characterized
as being high l y v a r i a b l e b u t r e s i l i e n t (Norton-Griffiths 1979).

A t the other extreme are systems i n which water a v a i l a b i l i t y i s high

b u t t h e s o i l s a r e e x t r e m e l y n u t r i e n t - p o o r . Such systems a r e
dominated by t r e e s and shrubs and include the miombo woodlands o f
the Central A f r i c a n plateaux, the Cam O cerrados o f Brazi 1, and the
monsoonal t a l l g r a s s woodlands O7- A u s t r a l i a . Because o f t h e h i g h
r a i n f a l l and prolonged n e t season, water i s seldom l i m i t i n g , even on
s t o n y o r l a t e r i t i c s o i ls, o t h e r than on a seasonal basis. The
grasses a r e t a l l , f i b r o u s , and o f low n u t r i t i o n a l q u a l i t y . Grass
p r o d u c t i o n i s h i g h e s t i n t h e open, between trees. Grasses g r o w i n g
under t r e e s b e n e f i t f r o m t h e more mesic c o n d i t i o n s and h i g h e r
n u t r i e n t status o f the s o i l s b u t produce less biomass than grasses
growi ng i n the open. This abundance o f low-qua lity p l a n t materia 1,
i n A f r i c a a t l e a s t , s u p p o r t s o n l y a low biomass and d i v e r s i t y o f
l a r g e h e r b i v o r e s . The amount o f p l a n t m a t t e r t h a t t h e y consume
annua 1l y is genera 1ly low, though periodic outbreaks o f lepidoptera
larvae, one o f t h e c h a r a c t e r i s t i c f e a t u r e s o f these systems, can
cause severe defoliation.
Most p r i m a r y p r o d u c t i o n goes i n t o an i n c r e a s e i n s t a n d i n g c r o p
biomass o r i s r e c y c l e d as l i t t e r f a l l . L i t t e r q u a l i t y i s low and
decomposes r a t h e r s l o w l y i n t h e absence o f f i r e . The decomposer
biomass, p a r t i c u l a r l y o f t e r m i t e s and, i n t h e w e t t e r savannas,
earthworms, i s v e r y h i g h and may e q u a l o r exceed t h a t o f p r i m a r y
consumers. Dry season f i r e s occur r e g u l a r l y and are f u e l l e d by o f t e n
large amounts o f dead o r dry m a t e r i a l which has accumulated by the
end o f t h e g r o w i n g season. F i r e s a r e . s u f f i c i e n t l y i n t e n s e t o k i 11
seedlings and saplings, thereby e l i m i n a t i n g fire-sensi t i v e species.
However, f i r e i s s e l d o m a b l e t o c o n t r o l a l 1 woody p l a n t
establishment and growth. Once estab lished, many woody species are
ab l e t o s u r v i v e t h e s e f i r e s by r e s p r o u t i ng f r o m e x t e n s i v e
underground r o o t systems (Frost 1984).

The presence o f a h i g h underground biomass and t h e maintenance o f

s u b s t a n t i a l carbohydrate and n u t r i e n t reserves, t o g e t h e r w i t h t h e
a b i l i t y t o resprout i f damaged, means t h a t the woody p l a n t s i n these
systems are w e l l buffered against periodic disturbance i n the form
o f f i r e , drought and the occasional outbreak o f herbivorous insects.
The systems a r e t h e r e f o r e r e l a t i v e l y s t a b l e . I f s u b j e c t e d t o
disturbance, such as the c l e a r i n g o f woodland f o r c u l t i v a t i o n , they
tend t o r e v e r t t o woodland once the source o f disturbance has been
relieved, though t h i s may take some time, p a r t i c u l a r l y i f there has
been a loss o f woody p l a n t propagules (Robertson, pers. comm.). The
r a t e o f r e c o v e r y depends l a r g e l y on t h e e x t e n t o f d e c l i n e i n t h e
o r g a n i c m a t t e r p o o l and t h e r e f o r e on t h e n u t r i e n t s t a t u s o f t h e
system f o l lowing clearing.

Two other f u n c t i o n a l l y d i s t i n c t b u t less widespread systems occur.

The f i r s t consi s t s o f low-medium q u a l i t y grasslands w i t h few woody
plants. These are found i n areas where both water and n u t r i e n t s are
l i m i t e d . They u s u a l l y occur under r e l a t i v e l y high r a i n f a l l on s i t e s
w i t h s h a l l o w s o i 1s ( f o r example, some o f t h e u p l a n d p l a t e a u x o f
C e n t r a l A f r i c a , campo s u j o and campo l i m p o g r a s s l a n d s i n C e n t r a l
B r a z i l , and p a r t s o f t h e l l a n o s o f Venezuela), o r under l o w e r
r a i n f a l l on nutrient-poor . s a n d s o r examp le, t h e s o u t h e r n Sahel).
I n Africa, such areas support a low biomass o f fibre-tolerant, and
species- and p l a n t - p a r t s e l e c t i v e grazers. The density o f herbivores
i s generally too low t o reduce the biomass o f grass s u f f i c i e n t l y t o
exclude dry season f i r e s , where these occur. These f i r e s l i m i t t h e
e s t a b l i s h m e n t o f woody plants,. b u t have l i t t l e e f f e c t on t h e
grasses, most o f w h i c h a r e dormant a t t h e time. Because o f t h e
r e l a t i v e l y low p l a n t p r o d u c t i o n , s o i 1 o r g a n i c m a t t e r l e v e l s a r e
o f t e n low. Any reduction i n i n p u t r e s u l t i n g from excessive herbivory
o r f i r e leads t o a d e c l i n e i n t h e amount o f o r g a n i c m a t t e r which
only recovers slow l y because o f the low l e v e l o f p l a n t production.
Secondly, there are areas i n which n e i t h e r water nor n u t r i e n t s a r e
l i m i t e d These occur i n some R i f t v a l l e y s i t u a t i o n s i n Africa, and
a l o n g t h e f o o t h i 11s o f t h e m a j o r escarpment zones. The v e g e t a t i o n
consi s t s p r i m a r i ly of medium-high q u a l i t y woodland w i t h palatable
grasses. I n A f r i c a , such systems s u p p o r t a h i g h biomass and
d i v e r s i ty o f both f i bre-tolerant and selective grazers and browsers.
These a r e p o t e n t i a l l y t h e most u n s t a b l e o f savanna systems ( B e l l ,
1984). Ifd i s t u r b e d , f o r example by t h e d e s t r u c t i o n o f t h e woody
canopy, they change r a p i d l y t o one o f a number o f a l t e r n a t e States
which can be maintained by a combination o f f i r e , p l a n t cornpetition
f o r water, and the type and i n t e n s i t y o f herbivory.

For example, where woody p l a n t density i s reduced, grass production

increases, as does the amount o f water used by the grass layer. This
may reduce the amount avai lab l e t o trees and so lim i t t h e i r growth.
Subsequent changes depend on t h e q u a l i t y o f t h e grass a f t e r t r e e
c l e a r i ng. I f grass qua l i t y is low t h e n f i b r e - t o l e r a n t h e r b i v o r e s
w i 11 be favoured. However, these species seldom reduce the amount o f
g r a s s s u f f i c i e n t l y t o exclude f i r e s . The r e g e n e r a t i o n o f woody
species i s i n h i b i t e d and an open grassland maintained.

I f t h e grass q u a l i t y i s r e l a t i v e l y high, t h e n s m a l l e r , more

s e l e c t i v e g r a z e r s w i 11 be favoured. These species can reduce t h e
biomass o f grass s u f f i c i e n t l y t o l o w e r b o t h t h e frequency and
i n t e n s i t y o f f i r e s . The system may stabi l i z e a t t h i s point, w i t h a
h i g h biomass o f g r a z e r s and a h i g h p r o d u c t i o n b u t low biomass o f
grass. However, i f the competitive pressures exerted by grasses on
woody p l a n t seedlings are p e r i o d i c a l ly reduced, f o r example by the
combined e f f e c t s o f heavy grazing and drought, o r i f the c l i m a t e and
soi 1s favour the growth o f trees, then woody p l a n t s wi 11 gradua1 ly
re-estab lish and eventua 1l y suppress grass production.

Where c o n d i t i o n s do n o t f a v o u r t h e r a p i d g r o w t h o f woody p l a n t s ,
browsers a lone can someti mes check t h e i r increase. More usua 1ly,
browsers and f i r e i n t e r a c t . Regular f i r e s m a i n t a i n woody p l a n t s
w i t h i n the reach o f browsers and i n a n u t r i t i o n a l l y acceptable s t a t e
(Trollope 1974). The low biomass o f woody p l a n t s favours the growth
o f grasses which i n t u r n provide f u e l f o r l a t e r f i r e s . Occasionally,
f i r e and b r o w s i n g a r e unable t o check t h e i n c r e a s e i n woody p l a n t
biomass, f o r example, d u r i n g a s e r i e s o f v e r y d r y years. Woody
p l a n t s may then increase t o the p o i n t where they can suppress grass
growth, thereby reducing both f u e l loads and f i r e i n t e n s i t i e s .

HUMAN INFLUENCES

Humans have been a s s o c i a t e d w i t h savannas f o r a l o n g time, as

hunters, p a s t o r a lis t s and CU l t i v a t o r s . Thei r c u r r e n t i n f l u e n c e is
widespread and involves a range o f land-use practices which modify
savannas t o v a r y i n g degrees (Table 1). Some o f these a c t i v i t i e s ,
such as wood h a r v e s t i n g and t h e c l e a r i n g o f l a n d f o r c u l t i v a t i o n ,
have d i r e c t e f f e c t s on s o i l s t r u c t u r e and f e r t i l i t y , the c y c l i n g o f
n u t r i e n t s , and on t h e c o m p o s i t i o n o f v e g e t a t i o n (UNESCO, 1979).
Other a c r i v i t i e s , such as t h e use o f f i r e , t h e h u n t i n g o f w i l d
herbivores, and t h e keeping o f l i v e s t o c k , a f f e c t t h e s o i l and
vegetation i n d i r e c t l y . For examp le, increasi ng c a t t l e numbers leads
t o greater grass consumption, less p l a n t cover, increased trampling
and compaction o f the s o i 1, decreased . i n f i l t r a t i o n , greater r u n o f f
and e r o s i o n , l e s s p l a n t a v a i l a b l e m o i s t u r e , a n d i n c r e a s e d
a r i d i f i c a t i o n of the soi 1. The reduced grass growth a l s o r e s u l t s i n
fewer, o f t e n less intense f i r e s and less competition f o r woody p l a n t
seedlings, i n some cases leading t o marked increases i n the density
o f these speci es.

T a b l e 1. Types o f land-use i n savannas and t h e degree t o which

theseG d i f y savanna s t r u c t u r e and functioning.
....................................................................
Land use WILDLIFE PASTORALISM WOOD CULTIVATION
Degree CONSERVATION HARVESTING
of AND
modification UTILIZATION
---------------T----------------------------------------------------

e
MINIMUM Protected Nomadism Timber Shi f t i n g
c>
4 Areas (eg. Transhumance Fue 1 CU l t i v a t i o n
U
N a t i ona 1
Z
.#- Parks)
-
?Y
O
MEDIUM Game Ranching Timber Conventi ona 1
m
E Ranching Charcoa 1 agriculture
V)
4
- dry land
aJ
L croppi ng
U
fi MAXIMUM Domestication Cultivated Afforest- Irrigated
I
I
I
pastures: ation crop
v
I z e r o p r o d u c t i o n
g r a z i ng
....................................................................
Increasing development--------- +

Savannas on t h e d i f f e r e n t c o n t i n e n t s have been exposed t o t h e s e

impacts f o r very d i f f e r e n t lengths o f time and t h i s may be r e f lected
i n t h e i r present structure. Moreover, w i t h the sharp increase i n the
human p o p u l a t i o n , t h e i n t e n s i t y o f many o f these i m p a c t s has
increased markedly i n recent times. I n the dry savannas especially,
t h e combined e f f e c t s o f high animal numbers and reduced p l a n t cover,
 .S~IIIL~~J
k o ~ k q ~ apue q aJkJ ' i u a b ~ i n u aie^ ayz JO s l r e i a p a q i 6 ~ i p ~ P i S J a p ~ n
se ~ e k ~ se n ~mou 3 S B eaJe ue JO Â ~ o i s k qasn-pue1 J P l n 3 ~ i J P da q i
~ U ~ M O U *) s
( u~alled p a w a s q o a q i JO Âueui JO i u e u i t u J a i a p ~ o r e u ei mou
aJe suemnq z e q l az~u603aJoz aJoJaJaqi paau aM 'seuue~es Lep-iuasa~d
.JO 6 u r u o k i 3 u n ~ PUP a d n z 3 n ~ i s a q i pueisJapun oz 6 u ~ z d u i a i i e UI

*mazsÂs a q i 40 &k~!i3npOJd pue  z k ~i i ~ a g

a q i u i a u i l s a p d ~ i e q se 016 u ~ p e a lpue ' u o ~ z e i a 6 apue
~ Jaiem '1 L O S
aqa uo SaJnsSa~d~ a l e a ~uaAa 6 6 u ~ i P a J 3' 1 l e j u t e ~a 6 e ~ a ~ e - r r o l aJO
q
s ~ e a Aiuanbasqns JO s i 3 a j ~ aa q i uaiy6iaq asaql -[LOS a q i ub sa6ueq3
a l q LSJaAaJJ 1 Â ~ z u a ~ e d d0e1 p a l aAi?i( '~146110Jp6 u l ~ n p4 Ji?[n 3 k i ~ e d
 III: HYPOTHESES --
SECTION - ABOUT THE RESPONSES -
OF SAVANNAS --
TO STRESS
AND DISTURBANCE

INTRODUCTION
The underlying m o t i v a t i o n f o r t h i s programme i s t h e need t o improve
savanna management. Two issues are o f major concern: (1) a l t e r a t i o n s
t o t h e e c o l o g i c a l s t r u c t u r e o f savanna communities, i n v o l v i n g
changes i n species composition, r e l a t i v e abundance and r e l a t i o n s h i p s
between species; and, (2) changes i n f u n c t i o n i n g , p r i n c i p a l l y
d e c l i n e s i n p r o d u c t i v i t y , r e s u l t i n g f r o m changes i n w a t e r and
n u t r i e n t a v a i l a b i l i t y . To understand t h e i m p l i c a z i o n s o f t h e s e
changes f o r t h e dynamics o f savannas, and t h e r e f o r e f o r t h e i r
management, f o u r k e y q u e s t i o n s have been posed. I n a t t e m p t i n g t o
answer these questions, we propose t h a t t h e main research programme
be c o n c e n t r a t e d on t e s t i n g a number o f r e l a t e d hypotheses. These
w i l l s e r v e t o f o c u s t h e r e s e a r c h e f f o r t r a t h e r t h a n have i t
d i s s i p a t e d a c r o s s a spectrum o f u n r e l a t e d p r o j e c t s which, w h i l e
i n t e r e s t i n g i n t h e i r own r i g h t , do n o t s u b s t a n t i a l l y advance o u r
u n d e r s t a n d i n g o f savanna dynamics i n ways which would l e a d t o
improved management.
The l i s t o f hypotheses i s c l e a r l y incomplete. I t i s l i k e l y t h a t some
important aspects o f savanna eco logy have not been inc luded because
we c u r r e n t l y do n o t know enough about them. These p o o r l y understood
components and processes w i 11 need t o be i n v e s t i g a t e d w i t h i n t h e
broader frarnework o f the programme. The r e s u l t i n g knowledge can then
be used t o r e f i n e t h e e x i s t i n g hypotheses o r t o propose new ones.
The r e v i s i o n and f o r m u l a t i o n o f hypotheses i n t h e l i g h t o f new
o b s e r v a t i o n s and i n s i g h t s w i l l be one o f t h e o b j e c t i v e s o f t h e
s e r i e s o f workshops t o be h e l d during the course o f the programme.

FUNCTIONAL. CLASSIFICATION OF SAVANNAS

Savannas encoapass a w i d e v a r i e t y o f systems w i t h d i f f e r e n t

s t r u c t u r a l and f u n c t i o n a l characteristics. This makes c l a s s i f i c a t i o n
d i f f i c u l t . E x i s t i ng c l a s s i f i c a t i o n s , based p r i m a r i ly on physiognomy,
a r e u n s a t i s f a c t o r y s i n c e t h e y do n o t convey much about f u n c t i o n .
Moreover, most o f the schemes are regional or, a t best, c o n t i n e n t a l
i n scope. None a r e accepted world-wide. A g e n e r a l s t r u c t u r a l -
f u n c t i o n a l c l a s s i f i c a t i o n o f savannas i s therefore u r g e n t l y needed.

We c o n s i d e r t h a t s o i 1 m o i s t u r e and n u t r i e n t a v a i l a b i l i t y a r e t h e
primary determi nants o f savanna f u n c t i oning. Thei r wide v a r i a t i o n i s
probably a major reason f o r the d i v e r s i t y o f savanna types. However,
simple i n d i c e s o f moisture and n u t r i e n t a v a i l a b i l i t y based on, f o r
example, mean a n n u a l r a i n f a l l and s o i 1 type, do n o t a d e q u a t e l y
e x p l a i n a l 1 o f t h i s d i v e r s i ty. B e t t e r measures o f p l a n t - a v a i l a b l e
moisture and n u t r i e n t s a r e t h e r e f o r e needed b e f o r e a s t r u c t u r a l -
f u n c t i o n a l c l a s s i f i c a t i o n o f savannas can be produced. Some o f t h e
f a c t o r s t h a t should be taken i n t o account are: s o l a r radiation; a i r
temperature; s e a s o n a l i ty, p r e d i c t a b i lit y a n d v a r i a b i 1it y o f
r a i n f a l l ; s o i l t e x t u r e ; pH; c a t i o n exchange c a p a c i t y ; n u t r i e n t
minera l i z a t i o n rates; topography, and land-use h i s t o r y .

A p o s s i b l e i n d e x o f m o i s t u r e a v a i l a b i l i t y i s t h e degree t o w h i c h
e v a p o r a t i v e demand is m e t b y s o i 1 water: t h e r a t i O o f a c t u a 1 t o
p o t e n t i a l e v a p o t r a n s p i r a t i o n m i g h t be a p p r o p r i a t e . Such an i n d e x
would i n t e g r a t e s o l a r r a d i a t i o n , a i r temperature, r a i n f a l l , s o i 1
t e x t u r e , topography and s e a s o n a l i t y . An i n d e x o f s o i l n u t r i e n t
avai l a b i lit y cou l d be t h e m i n e r a l i z a b l e capaci ty o f the s o i 1, w i t h
special emphasis on n i t r o g e n and phosphorus. A l lowance may have t o
be made f o r o t h e r s o i 1 f a c t o r s which c o u l d a f f e c t t h e v e g e t a t i o n
(e.g. exchangeab l e a lumi n i um and sodium percentages, the presence o f
heavy metals, and shortages o f micronutrients).

We propose t o develop these two i n d i c e s t o produce a c l a s s i f i c a t i o n

o f t h e w o r l d ' s savannas based on an o r d i n a t i o n o f a c t u a l s i t e s i n
r e l a t i o n t o these indices. The p r e d i c t i o n t o be tested i s t h a t s i t e s
w i t h s i m i l a r m o i s t u r e and n u t r i e n t i n d i c e s w i 11 e x h i b i t s i m i l a r
s t r u c t u r a l and f u n c t i o n a l characteristics.

I n t h e c u r r e n t absence o f a s a t i s f a c t o r y c l a s s i f i c a t i o n , we have
used the f o 1l o w i ng hypothetica 1 arrangement o f savanna types w i t h i n
t h e P l a n t Avai l a b l e M o i s t u r e (PAM) and Avai l a b l e N u t r i e n t s (AN)
plane (Figure 2) as a basis f o r f o r m u l a t i n g some o f the hypotheses.

I
HIGH LOW/HIGH I HIGH/HIGH
1
(cerrado)
(monsoona i t a 1 bgra;s)
(wet miombo) ; ( v a l ley and escarpment
( 1lanos I w o d lands
( d r y miombo)
I
I

(plateau grass lands) :

1 (Serengeti p l a i n s )
I (Acacia savannas)
I (Mopane wood lands
:
LOW
1 LOW/LOY
(Sahel)
!
(Astrebla grasslands)
HIGHILOW

LOW HIGH
Avai lab l e N u t r i e n t s
Figure 2. Hypothetical d i s t r i b u t i o n o f savanna types i n r e l a t i o n t o
the m a i l determinants o f savannas (modified from Bell, 1984)
HYPOTHESES
Each o f the f o l l o w i n g hypotheses i s presented i n three parts: ( i l a
s t a t e m e n t o f t h e h y p o t h e s i s , w i t h b r i e f a m p l i f i c a t i o n where
necessary; ( i i ) a statement o f i t s i m p l i c a t i o n s f o r management; and
( i i i ) an i n d i c a t i o n as t o how the hypothesis might be tested.

Hypothesis 1

Given t h e assumption t h a t m o i s t u r e and n u t r i e n t a v a i l a b i l i t y a r e

m a j o r d e t e r m i n a n t s o f savanna f u n c t i o n i n g , t h e e x p e c t a t i o n t h a t
changes i n PAM and AN cause a p r o p o r t i o n a t e l y g r e a t e r change i n
p l a n t p r o d u c t i o n i n t h o s e systems where b o t h a r e most s t r o n g l y
l i m i t i n g i s r e l a t i v e l y s t r a i g h t f o r w a r d . I t i s l e s s obvious why a
s i m i l a r t r e n d should be expected o f changes i n species composition.
1t i s based on assumptions t h a t PAM and AN Vary more i n a r i d ,
nutrient-poor systems, and t h a t the species i n these systems d i f f e r
widely i n t h e i r c a p a c i t i e s both t o survive prolonged periods o f
w a t e r and n u t r i ' e n t s t r e s s and t o e x p l o i t o c c a s i o n a l p u l s e s i n
avai l a b i l i t y . Changes i n PAM and AN i n such systems are more l i k e l y
t o r e s u l t i n changes i n the dominant species than i n those systems
where m o i s t u r e and n u t r i e n t s a r e more r e a d i l y a v a i l a b l e and
conditions f o r growth more favourab le.

Implication
The main i m p l i c a t i o n f o r management i s t h a t actions which a f f e c t the
a v a i l a b i l i t y o f w a t e r a n d / o r n u t r i e n t s a r e g o i n g t o have
proportionately greater e f f e c t i n those systems i n which PAM and AN
are most l i m i t i n g . These e f f e c t s w i 11 r e s u l t n o t only from actions
which increase PAM and AN, such as i r r i g a t i o n and f e r t i l i z a t i o n , b u t
a lso from ones which in d i r e c t l y lower them, f o r examp le, harvesting,
the use o f f i r e , and overstocking. There are also i m p l i c a t i o n s f o r
monitoring. A t t e n t i o n needs t o be focussed on the possible e f f e c t s
o f management actions on the physical envi ronment, p a r t i c u l a r l y i n
strongly water- and n u t r i e n t - l i m i t e d systems.

Test
The hypothesis can be tested by comparing the responses o f d i f f e r e n t
savanna types i n the PAMIAN plane t o changes i n water and n u t r i e n t
avai l a b i lity (e.g. through i r r i g a t i o n and f e r t i l i z a t i o n , both s i n g l y
and i n combination) and monitoring the r e s u l t i n g changes i n species
c o m p o s i t i o n and p r o d u c t i o n i n each case. The h y p o t h e s i s p r e d i c t s
t h a t t h e magnitude of change w i 11 be g r e a t e s t i n those systems i n
which both water and n u t r i e n t s are most l i m i t i n g .
Hypothesis 2
The co-occurrence o f two o r more independant events (e
a b o v e - a v e r a g e raTnfall,frost,
synergi s t i c e f f e c t s -
- -+
an
P
f i re, h e r b i v o r
i n changi ng savanna s t r u c t u r e pro=
drought,
w i 11 have
o r

I n t e r m s o f t h i s hypothesis, marked s h i f t s i n c o m p o s i t i o n o r
abundance are o f t e n caused by the co-occurrence, o r close sequence,
o f events w i t h compoundi ng effects. For example, t h e establishment
o f woody p l a n t s may depend on t h e co-occurrence o f (ila p e r i o d o f
exceptionally high o r out-of-season r a i n f a l l ; ( i i reduced
competi t i o n f rom p e r e n n i a 1 grasses; (i i i ) low herbivore pressure;
and ( i v ) a p e r i o d w i t h o u t f i r e . From a management perspective, some
o f these events are managable (e.g f i r e , herbivory); others a r e n o t
(e.g. above-average r a i n f a 11, droughts, f r o s t ) .

Implications
I f t h i s hypothesis i s valid, then management aimed a t inducing, o r
a v o i d i n g , m a j o r changes i n community c o m p o s i t i o n may o n l y be
possible when two o r more independent events c o i n c i de. Addi t i o n a 1ly,
events w h i c h co-occur in f r e q u e n t l y a r e like l y t o i n d u c e g r e a t e r
shi f t s i n vegetation composition and production. Understanding the
e f f e c t s on key species o f i n t e r a c t i o n s between p a r t i c u l a r c l i m a t i c ,
f i r e and h e r b i v o r y events, and knowing t h e p r o b a b i l i t i e s o f co-
occurrence o f these, i s a p r e r e q u i s i t e f o r e f f e c t i v e management.

Test
A t e s t o f t h i s hypothesis would be t o impose a v a r i e t y o f stresses
o r disturbances on a communi ty, s i n g l y and i n combination, and then
moni t o r t h e subsequent changes i n s p e c i e s c o m p o s i t i o n , r e l a t i v e
abundance and production. Since any changes t h a t occur are going t o
r e f l e c t d i f f e r e n c e s between s p e c i e s i n e s t a b l i s h m e n t , growth,
r e c r u i t m e n t and m o r t a lit y , a s t u d y o f these processes w i 11 be
e s s e n t i a l f o r u n d e r s t a n d i n g t h e mechanisms o f change. I n any
between-si t e comparisons, the h i s t o r y o f each s i t e w i 11 have t o be
taken i n t o account.

Hypothesis 3

& s u b s t a n t i a l change from the r e v a i l i n g frequency, i n t e n s i t y o r

sequence o f e v e n m a m a f i b l t-i n-a marked change -
s t r u c t u r e àn-ucti C.
Implications
R e l a t i v e l y r i g i d f i r e o r h e r b i v o r y r e g i m e s a r e o f t e n a p p l i e d by
management. One consequence o f t h i s is t h a t communi t i e s eventua 1ly
become dominated by those species which are best able t o accomodate
t h e p r e v a i l i n g s t r e s s and d i s t u r b a n c e regimes. Species w h i c h a r e
b e t t e r adapted t o other regimes d e c l i ne. Any substantia 1 d e v i a t i o n
from the prevai 1ing regime genera 1l y resu l t s i n sudden changes i n
species c o m p o s i t i o n and p r o d u c t i o n , t h e magnitudes o f which a r e
r e l a t e d t o t h e degree o f change i n t h e regime. By a a i n t a i n i n g a
variable herbivory o r f i r e regime, and t h e r e b y r e g u l a r l y exposing
t h e species w i t h i n t h e system t o a w i d e r range o f c o n d i t i o n s ,
managers can i n c r e a s e t h e r e s i l i e n c e o f t h e system t o these
disturbances and minimize the r i s k o f sudden s h i f t s i n composition
and production.

Test
The hypothesis can be tested experimentally by changing v a r i a b l e and
f i xed herbi vory o r f i r e r e g i mes and monitoring the resu lting changes
i n species composition and production. The hypoth'esis p r e d i c t s t h a t
t h e g r e a t e s t change w i 11 o c c u r i n those cases where a r e g i m e has
been most r i g i d l y a p p l i e d and where t h e d i f f e r e n c e between t h e
o r i g i n a l and the new regime i s greatest.

Hypothesis 4

Implications
This introduces the element o f the t i m i n g o f events as a f a c t o r i n
savanna dynamics. The e f f e c t s o f s t r e s s e s such as f i r e , f r o s t o r
herbivory o f t e n depend on the phenological s t a t e and physiological
c o n d i t i o n o f t h e p l a n t s a t t h e time. P l a n t s which a r e dormant a r e
u s u a l l y l e s s a f f e c t e d t h a n p l a n t s which a r e a c t i v e l y growing. A
stress occurring a t one t i m e o f the year may stimu l a t e reproduction
whereas a t another t i m e i t may re'tard it. The t i m i n g o f p a r t i c u l a r
management actions therefore may be as important as t h e i r magnitude
and t h i s needs t o be taken i n t o account i n management planning. In
t h i s regard i t i s c r u c i a l t o know what are the c r i t i c a l periods i n
l i f e c y c l e s o f key s p e c i e s and hou t h e s e m i g h t be a f f e c t e d by
d i f f e r e n t management actions.

Test
A t e s t o f t h i s h y p o t h e s i s would i n v o l v e v a r y i n g i n d e p e n d e n t l y t h e
t i m i n g and i n t e n s i t y o f events such as f i r e , herbivory o r a r t i f i c i a l
drought, and m o n i t o r i n g t h e e f f e c t s o f t h i s on t h e growth,
r e p r o d u c t i o n and s u r v i v a l o f i n d i v i d u a l s o f t h e key species. The
h y p o t h e s i s p r e d i c t s t h a t changes i n species p o p u l a t i o n s w i 11 be
determined more by the t i m i n g o f the event, i n r e l a t i o n t o c r i t i c a l
periods i n the species' l i f e cycle, than by i t s magnitude.
Hypothesis 5

Increasi ng leve 1s o f herbi vor i n s i ngle-herbivore systems increases

- ü n p d s j ë c i e s --
the proporti0?lOf i n the communi tx.

Hypothesis 5(a)

An a l t e r n a t i v e t o H y p o t h e s i s 5 is: Chan es i n t h e r o o r t i o n s o f
a l a t a b l e and unpalatable s ecies depe&~ine*ix
-
&=O
herbi vory.
-- e
-l
t h e oheno oav I O t h e s o e c i e s-
x h x Z-
h a e v e - of

Implications
The u n d e r l y i n g p r i n c i p l e o f Hypothesis 5 i s t h a t i n systems
dominated by one s p e c i e s o f h e r b i v o r e (e.g. c a t t l e ) , se l e c t i v e
feedi ng by the herbivore changes the competi t i v e balance between t h e
preferred, palatable species and the unpalatable ones i n favour o f
t h e l a t t e r . T h i s e f f e c t i s t h o u g h t t o be more pronounced a t h i g h e r
g r a z i n g i n t e n s i t i e s . To c o u n t e r t h i s , a system o f c o n t r o l l e d
selective grazing i s o f t e n applied. The system i s based on l e n i e n t
use o f preferred species and no d e f o l i a t i o n o f the unpalatable ones,
and involves two main assumptions: ( i l t h a t production o f preferred
species i s s t i m u l a t e d by moderate defoliation; and ( i i ) t h a t i n t h e
absence o f d e f o l i a t i o n by g r a z i n g o r f i r e , u n p a l a t a b l e s p e c i e s
eventual ly become moribund and dec line. An a l t e r n a t i v e management
system, non-selective grazi ng, i s based on the opposi t e p r i n c i p le.
I t assumes t h a t moderate d e f o l i a t i o n is more d e t r i m e n t a 1 t o
unpalatable species than heavy u t i l i z a t i o n i s t o the palatable ones.
Accordingly, high grazing pressures are applied f o r b r i e f periods i n
o r d e r t o f o r c e t h e a n i m a l s t o graze b o t h t h e p a l a t a b l e and
unpalatable species.

The r e a s o n i n g b e h i n d H y p o t h e s i s 5(a) i s d i f f e r e n t . F o r much o f t h e

time, g r a z i n g does n o t appear t o have much o f a n e g a t i v e e f f e c t on
g r a s s e s ; t h e i r p r o d u c t i o n may e v e n be s t i m u l a t e d . However,
d e f o l i a t i o n during the e a r l y stages o f growth i s o f t e n deleterious.
The d e c l i n e o f a s p e c i e s t h e r e f o r e may be l a r g e l y t h e r e s u l t o f
d e f o l i a t i o n d u r i n g t h i s e a r l y g r o w t h phase. R e l i e v i n g t h e g r a z i n g
p r e s s u r e on p a l a t a b l e s p e c i e s a t t h i s t i m e may be necessary t o
mai n t a i n t h e i r abundance i n a sward.

I f a p p r o p r i a t e g r a z i n g s t r a t e g i e s a r e t o be d e f i n e d f o r d i f f e r e n t
savanna regions, i t i s e s s e n t i a l t h a t these d i f f e r e n t hypotheses and
t h e i r assumptions be tested across the range o f savannas. Managers
need t o know how the timing, i n t e n s i t y and frequency o f d e f o l i a t i o n
a f f e c t t h e g r o w t h and r e p r o d u c t i o n o f key Pasture species ( t h e s e
i n c l u d e b o t h s p e c i e s t h a t decrease and those t h a t i n c r e a s e under
grazing), and how t h i s i n t u r n influences t h e i r population dynamics
and i n t e r a c t i o n s .
Test
The t e s t o f these hypotheses can c a r r i e d o u t i n conjunction w i t h the
t e s t o f Hypothesis 4 by v a r y i n g i n d e p e n d e n t l y t h e i n t e n s i t y and
t i m i n g o f grazing, and m o n i t o r i n g changes i n t h e p o p u l a t i o n s o f
p r e f e r r e d and n o n - p r e f e r r e d species. Hypothesis 5 p r e d i c t s t h a t
pa l a t a b l e species are increasingly adverse ly affected by an increase
i n grazing i n t e n s i t y and t h a t t h i s favours the unpalatable species,
enabling them t o increase i n the sward. I n contrast, Hypothesis 5(a)
p r e d i c t s t h a t p l a n t s d e f o l i a t e d during the e a r l y growth period w i l l
be more s u s c e p t i b l e than those d e f o l i a t e d when t h e p l a n t s a r e
dormant o r have completed most o f t h e i r annual growth. Any changes
i n species composition which occur w i 11 be caused by differences i n
the time o f d e f o l i a t i o n r e l a t i v e t o the t i m e o f e a r l y p l a n t growth.

Hypothesis 6

The e f f e c t o f d i s t u r b a n c e on t h e r a t e and e x t e n t o f change i n t h e

s p e c i e ç m p s i t i o n o f a s a v a n n a d e n K r i n c i - a T on t h e T i =
h i story characteri s t i E - a n T a t i- o5 n 107 ogy -- -
ofw the s p e c x -

The e f f e c t s o f disturbance on the species i n a community are usual ly

s e l e c t i v e , some .species b e i n g more s u s c e p t i b l e t o a p a r t i c u l a r
d i sturbance than others. Consequently, species composition tends t o
s h i f t i n f a v o u r o f t h o s e species b e s t adapted t o s u r v i v e t h e
p a r t i c u l a r d i sturbance, r e c o v e r f r o m it s e f f e c t s and e x p l o i t t h e
post-disturbance environment. A t t r i b u t e s which enable a species to
r e c o v e r r a p i d l y a f t e r d i sturbance inc lude ( i t h e presence o f
s u b s t a n t i a l belowground reserves; (i i ) t h e capacity t o resprout;
(i ii t h e presence o f r e l a t i v e l y l a r g e r e s e r v e s o f dormant seeds
w i t h v a r i a b l e g e r m i n a t i o n r e q u i r e m e n t s or, a l t e r n a t i v e l y , t h e
c a p a c i t y t o d i s p e r s e t o and r e c o l o n i z e a s i t e r a p i d l y a f t e r
disturbance; and ( v ) t h e a b i l i t y t o e s t a b l i s h under extreme
envi ronmenta 1 conditions.

Implications
P r i o r c o n s i d e r a t i o n o f t h e ways i n w h i c h d i f f e r e n t species a r e
like 1y t o respond to p a r t i c u l a r management actions o r envi ronmenta 1
events i s c r u c i a l t o e f f e c t i v e management. An understanding o f the
main f e a t u r e s o f t h e l i f e h i s t o r y and p o p u l a t i o n b i o l o g y o f key
species is therefore necessary. For example, under what conditions
do d i f f e r e n t species e s t a b l i s h f r o m seeds, grow t o m a t u r i t y and
reproduce? What conditions cause the w i despread death o f individua 1s
o f a species? How a r e t h e s e processes a f f e c t e d by f i r e , grazing,
drought and i n t e r a c t i o n s w i t h other species? Related species do n o t
necessari l y respond i n the same way t o the same management action.
Test
The h y p o t h e s i s can be t e s t e d e i t h e r by i m p o s i n g on a community a
disturbance such as extreme d e f o l i a t i o n , o r removal o f woody plants,
o r by s t u d y i n g n a t u r a l d i s t u r b a n c e s such as extreme d r o u g h t and
m o n i t o r i n g t h e r e s u l t i n g changes i n abundance and biomass o f
p a r t i c u l a r species. The changes need t o be studied a t the population
l e v e l i n order t o understand the mechanisms involved. The species
chosen f o r s t u d y s h o u l d encompass species w i t h a w i d e a r r a y o f
c o n t r a s t i n g l i f e h i s t o r y a t t r i b u t e s and, where p o s s i b l e , known
d i fferences i n response t o the p a r t i CU l a r d i sturbance.
Since t h e r e l a t i o n s h i p s , i f any, between species' l i f e h i s t o r y
a t t r i b u t e s and kinds o f change i n the abundance and biomass o f these
species a r e mu l t i v a r i a t e , p o s s i b l e a s s o c i a t i o n s can b e s t be
d i sp l a y e d t h r o u g h correspondence analysis. The hypothesis p r e d i c t s
t h a t the direction, magnitude and manner o f change w i 11 be s i m i l a r
i n species having s i m i l a r l i f e - h i s t o r y a t t r i b u t e s , whereas species
w i t h d i f f e r e n t a t t r i b u t e s w i 11 tend t o respond d i f f e r e n t l y .

Hypothesis 7

----
b -e-
The res onses o f savanna species t o stress can be pred5ct:d.
a ç i s O the'ir-fe h i s t o r y characi%- -
apopulation
on the
ioTogr

Some o f the a t t r i b u t e s which enhance a plant's capacity t o withstand

s t r e s s i n c l u d e : ( i ) a h i g h r e p r o d u c t i v e output, p a r t i c u l a r l y one
t h a t i s stimulated by stress; ( i i ) a large root:shoot biomass r a t i o ,
which l i m i t s t h e amount o f m a t e r i a l t h a t h e r b i v o r e s o r f i r e can
consume; ( i i i ) unpa l a t a b i lity t o herbivores, o r other features which
r e s t r i c t the amount o f f o l i a g e consumed; ( i v ) compensatory growth i n
response t o d e f o l i a t i o n ; ( v ) t h e presence o f energy and n u t r i e n t
r e s e r v e s on which t h e p l a n t can draw i n t i m e s o f s t r e s s ; ( v i l t h e
a b i lit y t o r e p r o d u c e v e g e t a t i v e l y ; ( v i i m o r p h o l o g i c a l and
phenological p l a s t i c i t y , which a l l o w r a p i d adjustments t o s t r e s s ;
and ( v i i i ) physiological quiescence during c l i m a t i c a l ly unfavourable
times o f the year.

Imp 1i c a t i ons
E f f e c t i v e management depends o n b e i n g a b l e t o a s s e s s t h e
consequences o f d i f f e r e n t a c t i o n s and choose t h a t which comes
c l o s e s t t o g i v i n g t h e d e s i r e d r e s u l t . Since i t i s n o t f e a s i b l e t o
d e t e r m i n e empi r i c a 1 l y t h e response o f a 11 species t o e v e r y
management a c t i o n , sosie b a s i s is needed f o r b e i n g a b l e t o p r e d i c t
t h e p r o b a b l e responses o f t h e key species. I f t h i s can be done by
using information on the species' life-hi story a t t r i b u t e s , then the
p o t e n t i a l f o r e f f e c t i v e management w i 11 be g r e a t l y enhanced.
Test
The t e s t o f t h i s h y p o t h e s i s i s s i m i l a r t o t h a t o f Hypothesis 6
except t h a t the avai l a b l e information on the l i f e - h i s t o r y o f the key
species i s used t o p r e d i c t c h a n g e s i n t h e i r abundance andbiomass
b e f o r e t h e e x p e r i m e n t i s c a r r i e d out. ( A t o u r c u r r e n t l e v e l o f
u n d e r s t a n d i n g i t i s l i k e l y t h a t t h e p r e d i c t i o n s w i 11 o n l y be
qualitative.) The t e s t i n v o l v e s making a comparison between these
predicted changes and those induced by the p a r t i c u l a r stress. I f the
observed changes d i f f e r f r o m those p r e d i c t e d then e i t h e r t h e l i f e
h i s t o r y a t t r i b u t e s o f a species a r e n o t good p r e d i c t o r s o f
population change o r our c u r r e n t understanding o f these a t t r i butes
and t h e i r e f f e c t s i s inadequate f o r making accurate predictions.

A d i r e c t t e s t o f the hypothesi s w i 11 not be possible where there is

in s u f f i c i e n t l i f e h i s t o r y i n f o r m a t i o n a v a i l a b l e a t t h e o u t s e t .
Instead, the information on l i f e h i s t o r y a t t r i b u t e s w i l l have t o be
c o l l e c t e d f o r each species during the course o f the experiment. As
i n t h e t e s t o f Hypothesis 6, correspondence a n a l y s i s can t h e n be
used t o d e t e c t any u n d e r l y i n g a s s o c i a t i o n s between s p e c i f i c
a t t r i b u t e s and p a r t i c u l a r s t r e s s - i n d u c e d changes i n abundance and
biomass. The aim would be t o d i s t i n g u i s h between those a t t r i b u t e s
which c o r r e l a t e w i t h t h e observed p o p u l a t i o n o r biomass changes
( t h e s e w i l l be t h e a t t r i b u t e s l i k e l y t o have p r e d i c t i v e value),
those which c o r r e l a t e w i t h each other and are therefore redundant,
and those which do n o t c o r r e l a t e w i t h any o f the changes.

To d e t e r m i n e how s u c c e s s f u l l y a c t u a l changes can be p r e d i c t e d ,

e i t h e r a r e s e r v e s e t o f d a t a (i.e. d a t a f r o m one o r more o f t h e
r e p l i c a t e d p l o t s , w i t h h e l d f r o m t h e i n i t i a l a n a l y s i s ) , o r an
independent l y C O 1 l e c t e d d a t a set, can t h e n be ana l y s e d t o see t o
what e x t e n t t h e i d e n t i f i e d r e l a t i o n s h i p s r e m a i n s t a b l e and a r e
p r e d i c t a b l e . However, causa lit y i n these r e l a t i o n s h i p s cannot be
i n f e r r e d from these ana lyses. To understand the mechani sms involved
w i 11 r e q u i r e d e t a i l e d s t u d i e s o f t h e ways i n which s t r e s s a f f e c t s
key population processes i n each o f the species.

Hypothesis 8

A decrease i n the e f f e c t i v e r a i n f a l l o f a s i t e leads t o a decline i n

Implications
Management actions which reduce the i n p u t o f water t o the s o i 1 (e.g.
by overstocking t o the p o i n t where trampling and compaction o f the
soi 1 r e s u l t ) , o r which increase the r a t e o f p l a n t water-use (e.g. by
f e r t i l i z a t i o n ) , may s h o r t e n t h e l e n g t h o f t i m e t h a t w a t e r i s
a v a i l a b l e t o plants. T h i s w i l l concentrate phenological types i n
time, r e s u l t i n g i n an e v e n t u a l r e d u c t i o n i n species d i v e r s i t y
t h r o u g h c o m p e t i t i o n and drought-induced m o r t a l i t y . Management
actions which might shorten the period o f s o i 1 water a v a i l a b i l i t y
would have t o be avoided.

Test
The h y p o t h e s i s can be t e s t e d by u s i n g r a i n - o u t s h e l t e r s and
i r r i g a t i o n t o keep constant the t o t a l amount o f water avai l a b l e t o
the p l a n t s whi l e varying the period o f avai l a b i l i t y . The hypothesis
p r e d i c t s that, under t h e same t o t a l amount o f p l a n t available water,
there w i 11 be marked changes i n d i v e r s i t y , i n c l u d i n g perhaps a loss
o f species, i n t h o s e s i t u a t i o n s where t h e p e r i o d o f w a t e r
avai l a b i lit y is sharp ly reduced. Species norina 1l y developi ng i n the
middle t o l a t e r a i n y season w i 11 be most susceptible, whi l e drought-
t o l e r a n t species w i 11 be favoured.

Hypothesis 9

The s t a b i l i t y
- o f savanna ecos stems, i n t e r m s o f t h e ca a c i t o f
t h e i r c o m D o n e n t s i ë d ë ? t h r o ~ so~~ ~ t u ance. r -gr-
iI
-
s t r o n ï y ;nf l u e n i t t h e d e g r-e e o ~ v ~ e n t cao ln s t r a i n t on
3-
a t a b l i s h m e n t an growth.

Recovery from stress o r disturbance can be arrested o r r e d i r e c t e d by

the subsequent i n t e r v e n t i o n o f drought, f i r e , herbivory, etc. before
the process o f recovery has been completed. Where recovery i s slow,
the l i k e l i h o o d o f subsequent events i n f l u e n c i n g the eventual outcome
is increased.

Implications
The main i m p l i c a t i o n f o r management i s t h a t where t h e process o f
recovery from stress o r disturbance i s slow, e i t h e r because o f t h e
s e v e r i t y o f the i n i t i a l stress o r disturbance, o r because o f strong
e n v i r o n m e n t a l c o n s t r a i n t s on e s t a b l i s h m e n t and/or g r o w t h (e.g.
n u t r i e n t - p o o r s o i l s , l o w w a t e r a v a i l a b i l i t y , p o o r seedbed
conditions, etc.), subsequent events may occur which w i 11 a f f e c t the
e v e n t u a l outcome. Where these subsequent events are controllable,
managers would need t o ensure t h a t they a l l o w s u f f i c i e n t time f o r
t h e species t o r e c o v e r b e f o r e these o t h e r pressures a r e imposed.
Where subsequent events are uncontrollable and unpredictable, then
t h e i n t e n s i t y and t i m i n g o f management a c t i o n s would have t o be
a d j u s t e d so t h a t t h e p l a n t s c o u l d r e c o v e r i n t h e s h o r t e s t t i m e
possible.
Test
The hypothesis can be tested by comparing the patterns o f vegetation
change ( s p e c i e s composition, r e l a t i v e abundance, p r o d u c t i o n o f
s e l e c t e d s p e c i e s ) a f t e r d i s t u r b a n c e (e.g. a r t i f i c i a 1 drought) on
s i t e s which d i f f e r i n t h e i r s u i t a b i lity f o r p l a n t growth. The
h y p o t h e s i s p r e d i c t s t h a t where c o n d i t i o n s a r e u n f a v o u r a b l e f o r
establishment and growth, the subsequent i n t e r v e n t i o n o f events such
as f i r e , herbivory, o r f u r t h e r drought w i 11 cause greater m o r t a l i ty
and changes i n species c o m p o s i t i o n and production, t h a n w i 11 t h e
same events, occurring a t the same time, on more favourable sites.

Hypothesi s 10
I f disturbed. savanna ecosvstems t e n d t o r e t u r n t o t h e i r f o r m e r

composition -
o r production.

Hypothesis 10(a)
An a l t e r n a t i v e . t o Hypothesis 10 i s : -+
Stabi lit i s n o t a major y--
f e a t u r e o f savannas. D i s t u r b a n c e - i nduce c h a n g e s i n s p e c i e s
com o s i t ' n and p r o d u c t i v i t y - a r e accomodated & s t r z t u r a l and
&dju~,ments --
w i t h i n the community. Autogenic recovery doeS
n o t occur.
--
Implications
The concept o f s t a b l e e q u i l i b r i a i s c e n t r a l t o much c u r r e n t
ecological theory and application. I t s relevance i n the context o f
savanna dynamics can be questioned, e s p e c i a l l y i n v i e w o f t h e
s t o c h a s t i c n a t u r e o f most o f t h e d r i v i n g variables, t h e
interactiveness and n o n - l i n e a r i t y o f many o f the processes, and the
r e s u l t i n g contingency o f many o f t h e i r effects. Yet the concept i s
w ide l y app 1i e d i n savanna management. F o r instance, a t t e m p t s by
managers t o reverse undesi rab l e changes i n vegetati on composition o r
p r o d u c t i v i t y caused by, f o r example, overstocking, involve removing
the cause o f the disturbance and r e s t i n g the disturbed area.

The underlyi ng assumption i s t h a t species composition and production

a t a s i t e t e n d towards a s i n g l e , s t a b l e e q u i l i b r i u m which i s
determined p r i mari l y by the prevai ling soi 1 conditions and c l i m a t e
( m a i n l y r a i n f a l l ) . Moreover, i f c o m p o s i t i o n and p r o d u c t i o n a r e
d i s t u r b e d t h e y w i 11 r e t u r n a u t o m a t i c a l ly t o t h e i r p r e d i s t u r b a n c e
s t a t e . Hypothesis 10 i m p l i e s t h a t t h e t i m e r e q u i r e d f o r recovery,
and hence t h e p e r i o d o f r e s t t h a t i s r e q u i r e d a f t e r disturbance,
increases w i t h the increasi ng magnitude o f d i sturbance (as measured
by the i n i t i a 1 d i splacement i n species composition o r production).
I n c o n t r a s t , Hypothesis 10(a) s t a t e s t h a t once a savanna has been
d i sturbed, species composition and production do n o t automatica 1l y
return t o t h e i r predisturbance levels but, instead, the changes are
accoaodated through s t r u c t u r a l and f u n c t i o n a l rearrangements.
Therefore, i n order t o restore the previous relationships, another
disturbance, a c t i n g i n t h e o p p o s i t e d i r e c t i o n , has t o occur.
Resolving t h e q u e s t i o n o f how savannas respond t o disturbance has
important implications f o r the kinds o f management actions t h a t are
r e q u i r e d t o counter any undesi r a b l e changes brought about by
disturbance. I n b o t h cases, i t i s c l e a r t h a t regular, f r e q u e n t
monitoring i s needed i n order t o detect and respond rapidly t o any
adverse changes which might occur.

Test
Both hypotheses can be tested i n the same experiment by disturbing
a d j a c e n t areas t o d i f f e r e n t degrees (e.g. removing d i f f e r e n t
proportions o f woody biomass) and monitoring the resu l t i n g changes
i n species composition and production. Hypothesis 10 predicts t h a t
c o m p o s i t i o n and p r o d u c t i o n w i 11 e v e n t u a l l y r e t u r n t o t h e i r pre-
d i s t u r b a n c e leve ls, w i t h t h e t i m e taken b e i ng p r o p o r t i o n a 1 t o t h e
magnitude o f t h e disturbance. On t h e o t h e r hand, Hypothesis 10(a)
predicts t h a t an area, once disturbed, w i 11 not show any consistent
tendency w r e t u r n w the original, predisturbance state unless i t
i s subsequently disturbed i n the opposite direction. The system w i 11
e i t h e r remain a t t h e p o i n t t o which i t was disturbed, o r undergo
f u r t h e r change i n the sane direction.

Hypothesis 11
The r e v e r s i b i l i t o f change i n p l a n t species composition and
P T o d u- d e r ~ l rye l a t e h --
t o t h e degree -
o f change --
i n so-7
physico-chemica 1 properties.

An a l t e r n a t i v e t o Hypothesis 11 i s : No i r r e v e r s i b l e chan e i n
r ecies con o s i t i o n wi11 occur wirhout a F n c u r y e n t and i o d t i q
d k @ - - - , ~ t i c ~ l a ~ a gs o l~1 p> h - ~F i h
~ro~erties.

The a s s u i p t i o n u n d e r l y i n g b o t h hypotheses i s t h a t t h e physico-

chemical properties o f the soi1 are a major determinant o f savanna
coiposi t i o n and productivi ty. The d i s t i ngui shi ng feature between the
two hypotheses i s the degree t o which plant species composition and
production are able t o recover from disturbances t o soi 1 properties.
I n t e r m s o f H y p o t h e s i s 11, any changes t o t h e s o i 1 d u r i n g
d i sturbance r e s u l t s i n a correspondi ng change i n t h e equi l i b r i u m
Table
-- 2. Some possible e f f e c t s o f herbivores on s o i l properties and
processes
---------------------------------------------------------------------
IMPACT EFFECTS CONSEQUENCES
---------------------------------------------------------------------
1. Reductions 1.1 Increase i n area * Increased i n s o l a t i o n
i n p l a n t and o f bare soi 1 * Increased exposure u,
lit t e r cover r a i ndrop impact
* Increased s o i 1
temperature
* Reduced in f il t r a t i o n
* Increased run-off
* Increased p o t e n t i a l f o r
water erosion
1.2 Reduction i n * Lower S.O.M.
input o f l i t t e r * Less s o i 1 cohesion
1.3 Reduced r o o t * Increased p o t e n t i a l f o r
growth leaching tm groundwater
2. I n p u t o f dung 2.1 Change i n s p a t i a l * Increased heterogeneity
and u r i n e and tempora 1 i n d i s t r i bution o f s o i 1
distribution of nutrients
n u t r i e n t inputs * Increased v o l a t i l i z a t i o n
2.2 Change i n q u a l i t y * More r a p i d O.M.
' o f nutrient inputs turnover

3. Compaction 3.1 Macropore space * Reduction i n soi 1

(increase i n reduced water-ho l d i ng capaci ty
bulk density 3.2 Micropore space * Less p lant-avai lab l e
o f the s o i l ; increased water
most l i k e l y 3.2 Total pore space * Less favourable soi 1
when soi 1s reduced microclimate f o r bioca
are moist) 3.3 Reduced r a i nwater * Poor environment f o r
i n f il t r a t i o n seedling e s t a b l i s h e n t
3.4 Increased r u n - o f f and r o o t growth
* Increased p o t e n t i a l f o r
water e r o s i on

4. Trampling 4.1 Reduced s i z e o f * Increased p o t e n t i a l f o r

(break up o f soi 1 aggregates wind and water erosion
soi 1 aggregate * P o t e n t i a l loss o f
structure; surface soi 1
most l i k e l y 4.2 Formation o f soi 1 * Reduced i n f i l t r a t i o n
when soi 1s surface sea 1s * Increased run-off
are d r y ) * Loss o f s o i 1 seed s t o r e
* Unfavourable envi r o m e n t
f o r seed germination and
seedli ng estab lis h e n t
conditions f o r p l a n t growth, and t h i s i n t u r n a f f e c t s the degree t o
which the vegetation i s able t o r e t u r n t o i t s pre-disturbance state.
Thus t h e g r e a t e r t h e d i sturbance t o t h e s o i 1, t h e l e s s comp l e t e l y
the vegetation recovers t o it s pre-di sturbance state.

I n contrast, Hypothesis l l ( a ) i m p l i e s t h a t below some c r i t i c a l p o i n t

( w h i c h p r o b a b l y d i f f e r s f r o n s i t e t o s i t e , depending on t h e s o i 1
t y p e and c l i n a t e ) , any changes i n s o i 1 p r o p e r t i e s do n o t have a
l a s t i n g i n f l u e n c e on p l a n t species c o m p o s i t i o n o r production.
However, i f the soi 1 i s disturbed beyond t h i s point, the changes i n
the p l a n t community become ir r e v e r s i b le.

Iap l i c a t i o n s
Large h e r b i v o r e s a r e o f t e n i m p l i c a t e d i n changes t o p l a n t s p e c i e s
composition and production, b o t h d i r e c t l y , as p a r t i a l l y s e l e c t i v e
consuiers, and i n d i r e c t l y , through t h e i r impact on soi 1 properties
and processes. These inc lude t r a m p l i ng and compaction, reducti ons i n
p l a n t and l i t t e r cover, and t h e i n p u t o f n u t r i e n r s i n dung and
u r i n e , a l 1 o f w h i c h have many i n t e r l i n k e d consequences (Table 2).
One i m p l i c a t i o n o f t h e s e hypotheses i s t h a t t h e d i r e c t e f f e c t s o f
l a r g e h e r b i v o r e s on long-term v e g e t a t i o n change a r e l i k e l y t o be
less important these i n d i r e c t effects.

T h i s has i m p l i c a t i o n s f o r t h e assessment and m o n i t o r i n g o f range

c o n d i t i o n and c a r r y i ng capaci ty i n savannas. A t present, assessments
o f t h e c a r r y i n g c a p a c i t y o f an area a r e made a l m o s t e n t i r e l y i n
t e r m s o f t h e amount o f f o o d a v a i l a b l e f o r anima l s , r a t h e r t h a n i n
terms o f how much trampling and compaction an area can sustain and
what amount o f p l a n t c o v e r w i 11 be needed t o p r o t e c t t h e s o i 1. I n
t h i s r e s p e c t , m o r e c o n s i d e r a t i o n needs t o b e g i v e n t o t h e
d i f f e r e n c e s w h i c h e x i s t between areas i n t h e i r s u s c e p t i b i l i t y t o
degradation (e.g. clayey s o i l s are more e a s i l y degraded than sandy
s o i ls, p a r t i c u l a r l y when moist).

Test
Since large herbivores are o f t e n i m p l i c a t e d i n changes t o both s o i 1s
and vegetation, one t e s t o f Hypotheses 11 and l l ( a ) could involve a
m u l t i f a c t o r i a l e x p e r i m e n t i n which ( i s i m u l a t e d grazing; ( i i )
t r a m p l i n g and compaction; and ( i ii)t h e a d d i t i o n o f excreta, a r e
v a r i e d s i n g l y and i n c o m b i n a t i o n t o c r e a t e a s e r i e s o f changes i n
the s o i 1 and vegetation a t a site. This would also a l l o w the e f f e c t s
o f these d i f f e r e n t disturbances t o be assessed independently o f each
other. Some o f t h e p r e d i c t e d changes a r e l i s t e d i n Table 2. Key
v a r i a b l e s which need m o n i t o r i n g i n c l u d e s u r f a c e s o i 1 compaction
and/or bu l k d e n s i ty, i n f i l t r a t i o n r a t e , s o i 1 temperature, s o i 1
organic matter fractions, N and P m i n e r a l i z a t i o n rates, as ne11 as
changes i n p l a n t species composition, r o o t growth, and seed ling
establishment. These would be moni w r e d d u r i ng the d i sturbance and
a f t e r the treatments have been withdrawn, t o determine whether there
has been any recovery t o pre-treatrnent leve 1s.
The eventua 1 d i f f e r e n c e i n v e g e t a t i on "composition and p r o d u c t i o n
between t h e p r e - t r e a t m e n t and p o s t - r e c o v e r y periods, and t h e
r e l a t i o n s h i p o f t h e s e changes t o t h e degree o f change i n s o i 1
p r o p e r t i e s which has o c c u r r e d over t h e same p e r i o d , p r o v i d e s t h e
basic t e s t o f the hypotheses. Hypothesis 11 w i l l be i n v a l i d a t e d i f
t h e d i f f e r e n c e s i n v e g e t a t i o n c o m p o s i t i o n and p r o d u c t i o n a r e
u n c o r r e l a t e d w i t h t h e d e g r e e o f change i n s o i 1 p r o p e r t i e s .
Hypothesis l l ( a ) w i l l be i n v a l i d a t e d e i t h e r i f t h e r e i s no
r e l a t i o n s h i p between t h e amounts o f change i n the vegetation and i n
soi 1 properties, o r i f t h e r e l a t i o n s h i p i s p o s i t i v e and linear.

Hypothesi s 12

A reduction i n the d i v e r s i t y o f savanna

- communities r e s u l t s i n
o i 1 organic matter -
7
a decline - i n s- and g d i - f f -. e z
Tyc 1ing processes.

The d i s r u p t i o n o f n u t r i e n t c y c l i n g a r i s e s from ( i l t h e narrowing o f

the spectrum o f l i t t e r quality; (i i ) changes i n the patterns o f r o o t
growth i n t i m e and space; and ( i i i ) a reduction i n synchrony between
n u t r i e n t a v a i l a b i l i t y and demand. These changes may r e s u l t i n t h e
l o s s o f n u t r i e n t s througR leaching, p a r t i c u l a r l y on w e l l - d r a i n e d
s o i l s i n regions o f high r a i n f a l l .

Imp 1i c a t i o n s
This hypothesi s has two major i m p l i c a t i o n s f o r t h e maintenance o f
s o i 1 f e r t i l i t y . F i r s t , t h e r e is an advantage i n mai n t a i n i n g
phenologica 1 and s t r u c t u r a 1 d i v e r s i t y i n savanna p l a n t communities.
Secondly, i t i s i m p o r t a n t t o m a i n t a i n those s o i 1 b i o l o g i c a l
processes which p r e v e n t o r r e v e r s e t h e l e a c h i n g o f n u t r i e n t s ,
especially where the soi 1s are well-drained and nutrient-poor. Woody
p l a n t s are c r u c i a l i n both instances since they produce substantial
amounts o f low q u a l i t y l i t t e r , which decomposes s l o w l y and
contributes t o a bui ld-up i n surface s o i 1 organic matter. They also
maintain a c t i v e r o o t growth and n u t r i e n t uptake i n the horizonta 1
and v e r t i c a l planes o f the soi 1 p r o f i le. The removal o f woody plants
from a savanna i s therefore l i k e l y t o cause considerable disturbance
w i t h long-term e f f e c t s on n u t r i e n t status o f the system.

Test
The hypothesis can be tested by compari ng n u t r i e n t cyc l i n g processes
i n savannas w i t h v a r y i n g p r o p o r t i o n s o f t r e e s and grass. The
f o l lowi ng processes need investigating: ( i t h e t i m i n g and q u a l i ty
o f d i f f e r e n t above- and below-ground organic matter inputs; ( i i1 the
decomposi t i o n rates o f these materials and the reçu l t i n g patterns o f
n u t r i e n t r e l e a s e ; ( i i i ) t h e e x t e n t o f n u t r i e n t immobi l i z a t i o n i n ,
and release from, soi 1 organic matter; and ( i v ) the seasonal p a t t e r n
o f n u t r i e n t uptake by the vegetation. The hypothesis p r e d i c t s t h a t
those communities which are less diverse, p a r t i c u l a r l y i n respect o f