84.

and _F\ coloratum (t5.16, p<0.001) 1n the t e r in l t a r ia sites.

During the early-dry season again no s ig n i f ic a n t difference was

apparent 1n the w h it e rhinos' feeding p a t t e r n between the two
h abit at types. In the va l l e y savannas the palatable-tal 1 species
such as Heterooogon contortus (average 34.3i ) , Themeda triandra
(average 14,31), D1 gi t a r 1 a e r i a nt h a (average 8.71) and
Hypar r h e n i a h irta (average 7.1%) were h i g n l y p r e f e r r e d . The
rh in o s al so showed an Increase In the selection for the
pa l a t a b le - t a l l spec . such as Heteropogon contortus, Brachyaria
se r r a t a , D1gi t a r i a r ia n t h a , Panicum deustum and f\ deustum 1n
the secondary grasslands, t e r i n l t a r i a or Cynodon dactyl on patcnes
and t h i c k e t s .

I t is In te re st in g to note that, although the white rhinos showed

a high preference for the Cynodon dactyl on "lawns" (approximate! /
21 of the a va ila bl e gras slan d, th is p a r t i c u l a r grass species was
generally avolded.

Th e ir d ie ta ry selection during tne late-dry season was siin 11 ar to

the early-wet season and the f i r e period with a high preference
f o r the p a l a t d o l e - t a l 1 species such as Hete ro p ogon cont ort us
(HC), and p1g1t a r i a e r i a ntha (DE). However, there was an Increase
1n the s e l e c t i o n f o r both deu stum ( P0) and col oratum (PCI
during th is season as opposed to the f i r e and early-wet periods.
In both habitats the unpalatable species were avoided while the
palatable-creeper species were only s l i g h t l y selected.

Thus, although the w h it e rh in o s u t i l i z e d a wide range of grass

species, only a t h i r d of them were highly selected throughout the
y e a r (Appendix i ) . Ourlng the f i r e pe rio d, however, most of the
species represented 1n t h e i r d i e t s did not d i f f e r g r e a t l y from
t h e i r a v a i l a b i l i t y In the grasslands. The white rhinos generally
selected for the pal at able- tal l and intermediate grass species.

U n p a l a ta b le species, such as Cymbopogon excavatus, Elyonurus

argenteus and B o t h r i o c h l o a 1n s c u l p t a , were g e n e r a l l y avoided.
However they were grazed when sh or t (eg. du ring the f i r e and
 85.

early-wet periods) and occasionally during the early-dry season

when t a l l stands of Bothriochloa were u t i l i z e d .

Owen-Smith (1973) observed tnat the white rhino food selection at

the Umrolozi Game Resen was bised l a r g e l y on grassland type,
1e. dominant species, ratner than Individual species. This tends
to agree w i t h the o b s e rv a t io n s and analyses discussed in the
previous sections on the Pllanesberg rhinos. The white rhinos at
PHanesberg showed a high preference for ce rtain grass types (eg.
p a l a t a b l e - t a l 1 and 1n t e r m e d i a t e ) Irre s p e c tiv e of t h e i r
a va ila b ility. In addition, the Correspondence Analysis was able
to show t h a t botn the gra sslan d s t r u c t u r e and grass species
composition In fl u e n c e d h e r b i v o r e feeding s e l e c t i o n . There was
also a tendency for t h -* white rhinos to show a seasonal selection
for grasslands of various heights. For example, during the f i r e
p e r io d and most of the wet season, w h i t e rh in o s conc entrated
th eir g ra zi n g on the shorter grasslands generally feeding
randomly on most grass species. Most of tne grasslands selected,
eg. secondary gra ssla nd s, v a l l e y savannas and Cynodon "1 awns1*,
were under 15 cm t a l l .

As conditions became d r i e r Increasing use was made of e tallei

grasslands with a greater prefererce shown for the more palatable
ta ll species such as Heteropogon contortus, Eragrostls superba,
Panicum maximum and P. c o l o r a t u m . This supports the r e s u l t s
collected by Melton (1978), who showed that tne white rhinos feu
ma1 n l y on P^ maximum and Themeda t r i a n d r a d u ri n g both seasons,
although a greater v a r ie t y of grass species was consumed during
the d r i e r months.

In itia lly the an im a ls concentrated on the localized shor t

patches, p a r t i c u l a r l y those around t e r m i t a r l a , but as progressive
drying and food depletion occured a switch was made to the t a l l e r
grasslands g r e a t e r than 15 cm 1n h e ig ht . The s h o r t grasslands
p r o v id e food only du ring the growing season. To c a r r y tnem
through the dry season, the w h it e rh in o s are dependent on the
r e m ai n in g ta lle r grasslands. The a v a i l a b i l i t y of t h i s sub
maintenance forage du rin g the d r i e r m'-^hs t h e r e f o r e bccomes
 86.

c ritic a l.

Owen-Smith (1973) showed a s i m i l a r trend with tne white rhinos

moving into the margins of the ta 11 Themeda triandra stands thus
expanding t.ie e x t e n t of the s h o r t grass areas. He was able to
demonstrate a seasonal movement along a catena with the animals
p r o g r e s s i v e l y moving up as tne seasons became d r i e r . A s i m i l a r
s it u a t io n was not found at Pilanesberg possibly for two reasons;

(i) most of the s lo p e s a re too staep and therefo re

Inaccessible to the white rhino, and
(11) the r n l n o p o p u l a t io n at P il a n e s b e rg is a t a r e l a t i v e l y
lo w e r d e n s i t y than a t the Umfolozi Game Reserve (0.005 anim als
per ha as opposed to over 3 per ha).

4.4.3.2 Oletary Selection by the Other Large Grazers

D ietary selection by the f o u r o t h e r l a r g e gr az ers was only

recorded during the dry season for two reasons. F i r s t l y , because
of the time r e s t r i c t i o n , greater emphasis was placed on the white
rhinoceros; and secondly, because t h is period 1s believed to be
most c r i t i c a l for grazing herbivores 1n tcr.^s of food l i m i t a t i o n .
Thus d ie t a ry selection and t h e i r degree of overlap would be the
most Important during t h is period.

Table 4.8 and Appendix H give the grass s p e d es compos 1t l on 1 n

the d i e t of the o t h e r fo u r l a r g e g ra ze rs f o r the e a r l y - d r y ana
l a t e - d r y seasons. These r e s u l t s have been averaged f o r a l l the
vegetation types. Appendix H gives the percentage c ontribution in
the d i e t (PC) and the Preference Indices (PI) of the grasses
consumed 1n each h a b it a t type.

4.4.3.2.1 Har te be es t

Table 4.8 show the die tary selection by hartebeest for tne two
p a rt s of the dry season. Grass species such as Rhyncheletrum
repe ns , Hyparrheni a hi r t a , Heteropogon c o n to rtu s and Themeda
triandra were h i g h l y se le c te d du rin g both seasons, although
 Table 4.8 : Percentage Contribution in the d ie t (PC) anti Preference Indices (PI) fo r grasses eaten by the other
large herbivores, by habitat and season. A key to the grass species is given in Table 4.1

CYDA KIIRE HCCO THTR fRCU ERSU ERRI ERGU HYHI DOIN
SEASON HABITAT pc pr M M ti PC PI PC PC i PC P! PC PI PC PI PC PI
IJartebeust. 0.0 0.0
(e a rly dry) SECGLO 0.0 0.0 1 L 1 u 13.1 0.9 8J L I 1.0 0.2 5.4 0.6 3.7 0.6 L i L i 8.2 1.0

Hartebeest SECGLD u .t 0.1 i f l j AJ 0.0 0.0 S u l L fi 0.0 0.0 1J) 3.1 0.5 1 .2 10.1 16.5 LA G.O 0.0
(la te dry) VALSAV 1.4 0.2 0.1 0.1 I L J L 4 LL1 L I A J. LQ 1.4 0.6 0.1 0.1 \i a j a j L 0.0 0.0 0.0 0.0

N il debeast SECGLD 0.1 0.1 I U 2.S ix ii 0.0 0.0 L fi L i H.2 0.9 l i L ! Lfi. 0.0 0.0 0.1 0.1 0.0 0.0
(e a rly dry) VALSAV 0.0 0.0 0.0 0.0 ZflJl 5J1 0.0 0.0 iJ l L .L 0.0 0.0 0.0 0.0 0.0 0.0 1.0 0.2 0.0 0.0
PEDGLD 0.0 0.0 0.0 0.0 10.0 0.7 0.0 0.0 4.0 1 A ZL0 lO iL i 1.0 0.2 0.0 0.0 0.0 0.0

Wildebeest SECGLD 3.3 0.3 38.5 l i 4.8 0.3 1 L S L I 0.1 0.1 0.1 0.1 6.4 1.1 0.0 0.0 0.0 0.0 0.0 0.0
(la te dry) VALSAV 3.0 0.5 1.7 0.5 0.1 0.1 4.5 0.9 LS L3 0.1 0.1 1.7 0.8 U - u 12-3 LS . 0.6 0.8
PEDGLD 0.1 0.1 2.0 P .5 14,4 1.0 J0.9 10.9 0.1 0.1 l.L.1 11.1 0.0 0.0 0.0 0.0 L L i 0.8 0.0 0.0

Zebra SECGLD 0.0 0.0 0.0 0.0 28.1 2Li fi-fi. 0.0 0.0 1.0 0.1 0.0 0.0 1.5 2.1 Ul^Z L I 0.0 0.0
(e a rly dry) VALSAV 0.0 0.0 0.1 0.1 2 L S L2 LA JLi 0.0 0.0 2 L 2 U l 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
Zebra 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0
HILSAV 0.1 0.1 L3 iJ U_
( la te dry)
Impala VALSAV ??, 0 0.0 0.0 1.0 0.1 0.0 0.0 ID^Q L 0.0 0.0 LQ L .l 0.0 0.0 0.0 0.0 0.0 0.0
(dry) THICKET 6.0 H 0.0 0.0 0.0 0.0 0.0 0.0 L I 0.0 0.0 1.0 0.5 0.0 0.0 4.0 1.2 0.0 0.0

(PAL, INT and UNP are the proportions of palatable, intermediate and unpalatable grass species, re s p e c tive ly).

SECGLO * Secondary grasslands

PEDCL!) * Pediment grasslands
VALSAV V a lle y savannas
HII.SAV H ills id e savannas
THICKET Acacia and riv e rin e thickets
n Num ber nl tra n se ( l s
 Table 4.8 (Cont.)

BRSE URflO LOFL D1ER PADE PACO sppy uupa

SEASON HABITAT PC P! PC PI PC Pi PC PI PC pi PC PI PC PI PC PI PAL INI UNP n
Hartebeest 0.1 45.9 54.0 0.1 9
(ea^ly dry) SECGLD liA lid . 2.5 0.5 ' .0 0.0 2.B L I 0.0 0.0 1.0 1.0 0.0 0.0 0.1

Hartebeest SECGLD 0.0 0.0 j .6 0.7 0.0 0.0 U 0.0 0.0 6,2 L i 0.0 0.0 0.0 0.0 25.8 74.2 0.0 4
( la te dry) VALSAV 0.0 0.0 5.4 3.2 Z A 1A 2Z J i 11.1 0.0 0.0 L I L fi IA LL 0.0 0.0 49.5 50.5 0.0 5

Wildebeest SECGLD 2JJ I A . 5.6 L I 0.0 0.0 1 14 L A 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 45.1 54.9 0.0 5
(e a rly dry) VALSAV 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 LLQ I f u l 0.0 0.0 1.0 2.5 0.0 0.0 93.0 7.0 0.0 3
PEDGLD 2B.0 20.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 19.0 19.0 0.0 0.0 0.0 0.0 77.0 23.0 0.0 3

Hi ldeheest SECGLD 0.0 0.0 14.0 14 0.0 0.0 3J M A 12,7 i A LA 0.0 0.0 0.0 0.0 47.5 52.5 0.0 6
LA t l 9.6 90.4 0.0 6
(la te dry) VALSAV 0.0 0.0 1 3 0.0 0.0 0.0 0.0 0.0 0.0 L I U J 0.0 0.0
PEDGLD 2 L IL 6 . 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 lid - I L i 0.0 0.0 0.0 0.0 81.5 18.5 0.0 3

?ebra SECGLD 23,7 30,6 <1.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.1 0.1 0.0 0.0 0.0 0.0 87.4 12.0 0.0 5
(e a rly dry VALSAV U SlA 0.0 0.0 U A LLQ. L a 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 71.2 28.8 0.0 5
Zebra
( la te dry) HILSA' i ? .3 u ; 0.0 0.0 L i L I L i 0.0 0.0 0.0 0.0 8.3 0.5 0.0 0.0 74.3 25.7 0.0 6

impdla VALSAV 0.0 0.0 2.0 0.2 0.0 0.0 22.0 13.8 22.0 9.0 0.0 0.0 0.0 0.0 0.0 0.0 25.0 75.0 0.0 2
(dry) THICKET 0.0 0.0 3.0 0.3 0.0 0.0 " O (TOT 6 0 & 0.0 0.0 L L 0 L L 0.0 0.0 71.0 29.0 0.0 2
 89.

E r a g r o s t i s gumini f l ua ( t =-2.12, p<U.01) and J i gl t a r i a e r l antha

( t -3.61, p<0.01) s i g n i f i c a n t l y increased w h i l e Rhynchel etrum
rep e n s decreased (t*3.d8, p<U.01).

Section 3.3.3.2 examined the hab itat selection of the hartebeest,

depicting a high preferenc e for secondary grasslands w it h
in te rm itte n t selection for v a lley savannas. Hence, dietary
differences only occurred between these two h a b it u s .

Palatable specie:* such as Themeda triandra (TT), D1y i t a r i a

eriantha (DE), and Panicum coloratum (PC) were highly selected in
both h a b i t a t s . T h j i n t e r m e d i a t e grass species, Rhyncheletrum
repens UR), contributed over 3u* of t n e i r d i e t in the secondary
gras slands, whereas the p a l a t a b l e species such as Themeda
t r i andra (TT), Heti;ropogon contortus (HC), and O i g i t a r i a eri antha
(DE) contributed over 50i in tne va l l e y savannas.

Althougn showing d i s t i n c t h abit at differences In t h e i r diet, the

hartebeest tendod to s e le c t for th e more p a la ta b le to
Intermediate grass species (see Table 4.8).

4.4.3.2.2 W11 debees t

The wil de be e st s ' d i e t was s i m i l a r to t h a t of the har te b e es t,

except more time wis spent foraging on grass species found 1n the
v a l l e y savannas. Very l i t t l e seasonal differences (early-dry and
late-dry) 1n t h e i r d i e t could be detected (see Appendix H),
except that Heterojogon contortus (HC) showed a marked decrease
during the l a t t e r part of the dry season (t*b.78, p<0.001).

Both Rhynchy1e t r u m repens (RR) and Heter opogon co ntortus (HC)

accounted for over aot of the wildebeests' d ie t 1n the secondary
grasslands. D u r l r . j the e a r l y p a r t of the dry season p a l a t a b l e
species such as Het e r o pogon c o n t o r t u s (HC) and f\ deustum (PJ)
w e re h i g h l y s electe d , whe reas tne In te rm e d ia te s p e c ie s
Hyparrhenia hirca (HH) and to a lesser extent Eragrostis curvula
(EC) were Important during the late-dry season. In the pediment
grasslands palatable species such as Heteropogon contortus (HC),
 90 .

Brachy a r i a s e r r a t a (BS) and Pani cuin col oratum (PC/, and tne
Intermediate spedes Eragrostls superba were selected during this
season.

Hence, the p a l a t a b l e - t a l 1 species, Rhynchetrum repens (RR),

Heteropogon c o n t o r t u s (HC) and, to a l e s s e r e x t e n t Themeda
t r 1andra (TT), Br a c hy a ri a s e r r a t a (BS) and D1g1tari a er1antha
(j E), ana tne Intermediate species, Rhyncheletrum repens form the
bulk of t h e i r w in t e r d ie t. Thus more p a l a t a b l e s p e d e s were
favoured 1n a l l the v e g e t a t i o n types. S im ila r results were
obtained by Melton (1978) who showed tnat tne wildebeest in tne
Umfolozi Game Reserve selected mainly for Dig1taria macroglossa,
Heteropogon c o n t o r t u s , E r a g r o s t l s superoa, Themeda t r i andra,
Pani cum col ora turn and Sp or obo l1s smutsi i du ring the w i n t e r
months.

4.4.3.2.3 Zebra

The zeora, with t h e i r tendency to select secondary grasslands and

v a ll e y savannas during the early-dry season and h . l l s l d e (mainly
m esocllne) savannas d u r i n g tne la te- d ry season, snowed
preferences .or grasses common 1n these habitats. For Instance,
p a la ta b le - ta lI species such as Heteropogon c o n t o r t u s (HC),
Themeda t r i andra (TT) and B ra c h y a ri a serrata (BS), ard the
Intermediate species Hyparrhenla h i r t a (HH), ware favoured 1n the
secondary gr a s sl a n d s, and He te ro po go n con t o r t u s (HECO),
E r a g r o s t i s superba (ERSU) ana Loudeti a f 1avada (LuFL) in tne
v a l l e y savannas. During the late-dry season Heteropogon contortus
(HECO), Themeda t r i andra (THTR), Brachyar i a s e r r a t a (BRSE),
Loudetia fl avada (10FL), D1gi t a r i a erlantha (DIER) and Sporobolus
py rami dal 1s (SPPY) c o n t r i b u t e d over 80* of t n e l r d i e t in the
h i l l s i d e savannas. By r e f e r r in g to Table 3.1 I t Is apparent that
the grass species selected 1n t h i s habitat are tnose most readily
available. Hence, a high preference was made for the palatable-
ta ll grass species especially 1n the secondary grasslands
(t*2.92, p<0.001). The s e l e c t i o n f o r the t a l l species co nf irm s
the results obtained In the Correspondance A n a l y s is which
depicted a close relatio n sh ip between the zebra and ta ll
grasslands.

The grass species, Heteropogon co n t o rt u s ( t 3-6.04, p<0.001),

Themeda trla n dr a ( t5.66. p<0.001), B ra c h y a ri a s e r r e t a ( t. -4.34,
P<0.001), Loudetla f 1avada (t-6.89, p<0.001), and Sporobolus
pyrami dal 1s ( t-9.38, p<0.001) s< gn 1 f I c a n t l y Increased, w h i l e ,
E r a g r o s t l s superba ( t - 5.31, p<0.001) and D1g 1t a r ia er1 a nt h a
(t*4.68, o<0.001) s i g n i f i c a n t l y decreased from one season to the
next.

M e lto n (1978) found t h a t tne Umfolozl zeora m a in l y u t i l i z e d

palatable species such as Panlcum maximum, Themeda trlandra and
Heteropogon contortus. The zebra 1n the Pllanesberg Game Reserve
showed s im ila r preferences 1n t h a t the palatable species
contributed co over 70 of t n e l r dry season diet.

4.4.3.2.4 Impala

U n f o r t u n a t e l y , only f o u r feeding s i t e s were recorded f o r t n l s

species during the dry season, two In the v a l l e y savannas and two
1n the Acad a and r i v e r i n e thickets. Tnese were however, able to
give soTie Indication as to what grass species were preferred.

The p a l a t a b le - t al l species, Panlcum deustum was highly preferred

(PI of over 9.0) In both h a b i t a t types and c o n t r i b u t e d to about
501 of t h e i r diet. Palatable species such as D1g1tar1a eria n tha
(DE) and C;nodon da c t y l on (CO), and the i n t e r m e d i a t e species
Eragrostls curvula (EC) were u t i l i z e d to a greater extent 1n the
v a l l e y savannas.

Impala were observed to u t i l i z e only the highly palatable grass

speclos du rin g the dry season, esp ecially 1n tlie t h i c k e t s .
S i m i l a r res ult s were obtained by Melton (1978), who showed tnat
the Impala In the Umfolozl Game Reserve preferred mainly Panlcum
maxi mum 1n both the wet and dry seasons.
 92.

CriAPTtR FUE

HABITAT AND DIETARY OVERLAP BETWEEN THE PILANF.j BURG oRAZF.RS:

5.1 Introduction.--

The major focus In studies of co.wiuni ty structure is tne use of

pattern of resource p a r t it io n in g as a basis for explaining the
coexistence of ecologically s im il a r species (eg. Pulliam ar.d
Parker, 1979). E s s e n tia ll y ecological community theory attempts
to predict the numbers and relative abundance of coexisting
consumer species Dased on tne a v a i l a b i l i t y of resources, and the
resource u tiliza tio n patterns of tne species MacArthur and
Levins, 1967; rtacArtnur and Wilson, 1967; Levins, 1968,
MacArtnur, 1972; May, 1973; Pulliam, 19tt3). The procedure
requires the calcu latio n of firs tly , carrying capacities for
eacn species based on the a va ila b ility of resources and
secondly, competition co eff ic ie n ts for each species p a ir , based
on the sim ilarity of their resource u tiliza tio n patterns
(Pulliam, 1983).

Although the precise application and Interpre "atlon of

ecological niche overlap remain a subject of debate, many
ecologists nevertheless use various overlap measures to Indicate
degrees of ecological s i m i l a r i t y within assemblages of species.
Studies by Planka (1975) and Cody (1974) made extensive use of
niche overlap data when studying In te rs pe ci fic competition.
H ur lb er t (1978) suggested tnat niche overlap measures should
serve "as a foundation for discussion of resource u t i l i z a t i o n
st rat eg ies , competition, species packing, and so on".

The term nlcne has been variously defined by many authors. Elton
(1927) considered the niche as the fundamental role of an
organism 1n a community - what i t does, I t s re la t io n sh ip to I t s
 93.

food and to Us enemies. Gr1nnel1 (19ii4) on the othe r hand,

referred to tne niche as a subdivision of the environment
occupied by a species. Hutchinson (19o9) con ce ptu al iz ed the
n ich e as a comoination o f the f u nc t io n a l rol e of an organism in
an ecosystem as well a:, its position in time and space. He
therefore defined tne n1:he of a sp ecies as an "n -d 1inens1onal
hypervolume". Levins (196d) went f u r t h e r by s t a t i n g t h a t "ni che
dim ensionality r e f e r s not to the number of b i o l o g i c a l l y r e l e v a n t
factors In the environment, but to the number of f a c t o r s which
se rv e to separate species". Thus before attempting any nlcne
st u d y , It Is Important to ><now the .pedes concerned and to
Identify those enviromnental factors to wnlch tney respond.
Separate d ivisions or dimensions of the v a r i o u s resource types
u tilized by the sp ec ie s can then be d efi ne d. Each resource
dimension Is tnen tr e a te d as & s i n g l e niche in an e u c l i d i a n
space of many dimensions ( S l o b o J c h i k o f f and S c h u l z , 1980). It
should be noted tha t these dimensions are not g ra d ie n ts of a
resource, as Is of ten Implied ( r e f ) , out c o n s i s t of d i s c r e t e
re source e n title s. For example, a he rb iv or es d i e t cannot be
interpreted as a g ra d i e n t of grass sp ec ie s but r a t h e r a resource
dimension c o n s i s t i n g of a number of resource typos.

Pew studies on the In tersp ecific Interactions w ithin e n tire

la^ge n e rbl vo rc communlties have been attempted in A f r i c a l e g .
V e r s e y - F 1t z g e r a l d , I 960 , 1965; Lamprey, 1*60; F i e l d , 1*68; B e l l ,
1969; Field and Laws, 197u; F e r r a r , i 973; F e r r a r and Walker,
1974; Page and Walker, 197d; A t t w e l l , 1978). Thus, tne a 1in of
this s e ct io n Is to analyze the nature and the relative
Importance of those f a c t o r s which serve tc separate the s p e c i e s ,
eg. whit rh in o s and t h e i r p o t e n t i a l competitors In a southern
A f r i c a n game r e s e r v e .

5.1.1 Relationships Between o v e rl a p , S im ila rity , and

Competl 1 1on:

Befo re examining the varlors methods O'* e st im a ti n g niche

'"rlap, a clear understanding of the relationships and
 94.

differences between tne o e rla p , sim ilarity ard competition

c o e f f i c i e n t s , as exp la in e d by Lawlor ( I 9 d 0 ) , are n e cc es s a ry .

Overiap measures are designed to measure the degree to which two

species share a se t of common resources or u t i iz e the same
parts of the environment ( L a w l o r , 1^80). In other words, 1t 1s
the region of niche space ( I n tne sense of Hutchinson, 1958)
shared by two or more contiguous niches (C o lw e ll ana Futuyma,
19 71 ). Tnese ov erl ap measures are u s u a l l y sc al ed from zero to
one; zero Indicating no overlap, and one I n d i c a t i n g complete
overlap. Resource use can ce expressed as P1k, the proportion of
consumer species' ( 1) total u t i l i z a t i o n tha t i s represented by
each re so u rc e , k . The proportion of food type ( J ) depends on two
factors; the consumer s e l e c t i v i t y , and the a v a i l a b i l i t y of th a t
food type In the environment. Schoener (1974) defined e l e c t i v i t y
as tne consumer's r e l a t i v e a b i l i t y ( o r pr e fe re n ce ) to catch and
consume an item of a p a r t i c u l a r food type. I f a l l resource types
are e q u a l l y aDundant then the;** r e l a t i v e propo rtio ns 1n the d i e t
are determined only by tn e ir re la tiv e e l e c t l v i t i e s , lechovlcz
(1982) referred to the e le c tivity In d i c e s as measures of the
u tiliza tio n of food type ( r ) In r e l a t i o n to t h e i r aoundance or
a vailab ility (p) 1n the environment. Therefore, foods th a t
constitute a l a r g e r propo rtio n of the d i e t than of the a v a i l a b l e
foods are considered to be p r e f e r r e d . I f a p a r t i c u l a r food typ->
1s r e l a t i v e l y s c a re * i t may re pr e s e n t only a small proportion c f
the consumer's d iet, even though the consumer has a high
e l e c t i v i t y f o r th a t food type ( L a w l o r , 1980).

Measures of sim ilarity are often used in comparisons between

closely related sp ec ie s livin g in d ifferent communities or
envlronments. However, the problem with t h i s measure I s tha t tne
re so u rce s may d i f f e r 1n abundance between the environments thus
creating two major problems. F i r s t l y , s i m i l a r species may not
appear alike 1f livin g in t o t a l l y d i f f e r e n t environments, and
s e c o n d ly , the converse s i t u a t i o n may a r i s e where sp ecies which
are very d issim ilar may appear q uit e s i m i l a r because of the
species - environment I n t e r a c t i o n s ( L a w l o r , 19dO). Consequently,
conclusions about species' s i m i l a r i t i e s based on t h e i r ov erl ap s
may be misleading. It i s th e r e f o r e important to be quite sur'e
 95.

whether one i s i n t e r e s t e d in comparing tne sp ec ie s themselves or

the specie s-e n/ iro nme nt relationship s. Lawlor 11980), when
considering consumer s i m i l a r i t i e s , concluded t h a t they should be
b i se o on patterns of resource use r a t n e r tnan p a t t e r n s or the
r es ou rce s used. Tnus, the use of electivitie s rather tnan
pr o po rti on a l u tiliza tio n s should be used in any measive of
sim ilarity . This ensures th a t species with s i m i l a r e l e c t i v l t y
patterns w ill appear sim ilar even if compared In d i f f e r e n t
environments with d ifferent resource abundances, l a w l o r (198u)
mo difie d the Symmetric MacArtnur-Levins formula ( d i sc u s se d
late r) to measure sp eci es sim ilaritie s based on consumer
e le c tivitie s;

S1j * ___ ( a 1k x a j k ) ...................5.1

( a 1k T x (a jk )*

where a 1s the e l e c t i v i t y of resource k Dy sp ec ie s 1 and j , and

S 1s the slmi l a r i t y Inciex.

Tne main emphasis of t h i s chapter and the next 1s to Inco rpora te

competition Indices, using o v er la p measures, to calcu late
s to c k i n g ae n sltie s, whether at carrying capacity or at a
p a rticu lar malntalnec le v e l of u tiliza tio n . The theory of
com pet itio n Im p l i e s t t a t the g r e a t e r the o v er la p 1n resource use
between two organisms, the g r e a t e r the competition c o e f f i c i e n t ,
and hence the grtater the intensity of com pet iti on . By
d efin itio n , competition occurs when two or more organisms
Interfere with or 1nh1o1t one another (Planka, 1981).
Competition can be q u it e d i r e c t as 1n the case of I n t e r s p e c i f i c
te rrito ria lity (eg. Brown, 1964), and is then termed
"Interference Competl :1on" (Elton and M ille r, 1954; M ille r,
196 7). More indirect competition may a l s o o c c u r , such as th a t
arising through the jo in t use of the same l i m i t e d re so u rc e s,
th is oeing termed " E x p l o i t a t i o n Competition" ( E l t o n and M i l l e r ,
1954; M ille r, 1967). ;n t n l s c a s e , although the resource pool I s
 96.

reduced in ava ila b ility to the competing I n d i v i d u a l s , no d i r e c t

i n t e r a c t i o n s are apparent (Madaughton and Wolf, 1973).

As ntloned e arlie r.tn e degree of resource o ver la p has often

be *ed as a measure of tne Intensity of Interspecific
'i (eg. Schoener, 196d; L e v i n s , 1968; Cody 1974, C u lv e r
-inka 197J, 1975; May, 1975; whereby a positive
p has been assumed between the two measures. Planka
(19 72, 1974) on the otherhand, po st u la te d an I n ve rs e
relationship between the i n t e n s i t y of competition and the degree
of niche overla p (termed tne "Niche Overlap H y p o t h e s i s " ) , rty
reasoned that if resources are not 1n sh ort supp ly, " e x t e n s i v e
nlcne o v er la p may actu ally oe correlated with reduced
co m p e t i t i o n " . "D isjunct niches on the otherhand may often
Indicate avoidance of competition where 1t could have been
p o tentially severe". However, equating the degree of ov erl ap to
the Intensity jf competition can be dubious and m isleading
(Colwell and Futuyma, 1971). Mere overlap in resource use I s no
guarantee that competition Is taking place. T h i s 1s l a r g e l y
because:

( 1) tne resource may not be l i m i t i n g and ov erl ap th e r e f o r e

does not r e s u l t in com pet iti on ,
( 1 1 ) competition may be a complex fu nc ti on of o v e r l a p , and
( 111) low current o v er la p may be the result of past
competition (Col we ll and Futuyma 1971, H u r l b e r t 197a, P e t r i a t l s
1975, Abrams 1980, Lawlor 1980, Sio b od ch lk of f and S c h u l tz 1980).

Hence the theory of niche o v er la p and competition can only fce

a p p l ie d to systems which have l i m i t i n g r e s o u r c e s , eg. systems
at, or close to c a r r y i n g c a p a c i t y . I f one wishes to maintain a
system at a level of u t i l i z a t i o n well below c a r r y i n g c a p a c i t y
then competition is negllgable - resources are not l i m i t i n g
hen re, negligible Interspecific Interactions. However, I f the
objectives of a r e s e rv e I s to allow the d e n s i t y of one of the
>pec1es to in c re a s e w h ile maint ainin g a specified l e v e l of
u tiliz a tio n , then tne species overlap coefficients become
Im porta nt. Those sp eci es which de p ic t high ov erlap with t h i s
species w ill most likely be the ones which modify the grazing
le ve l. For example, by i n c r e a s i n g sp ecies A, sp ecies B, which
has sim ilar requirements, must L.e reduced othe rw ise the o v e r a l l
g ra zi n g pr e s su re w i l l exceed the s p e c i f i e d l e v e l .

Rosenzweig (1974, 1979, 1981) and P1mm and Rose, ^weig (1981)
have presented an interesting model with regards to h a b i t a t
selection ana com pe tit io n. Tney propose th a t sympatric s p e c i e s ,
ro ra g 1ng optim ally, avoid competition by u t 1 11zing a if f e r e n t
habitats, ie. d iffe re n tia l h atitat selection reduces
competition, sometimes to z e r o . The Important p o in t a r i s i n g from
this model 1s th a t although co m pe ti tiv e i n t e r a c t i o n s do not nave
to prevail at every possible d e n s i ty pair, tney do have to
produce an c\.?rall depression o f the d e n s i t i e s . Consequently,
i elating niche ov erl ap to tne degree of competition becomes
m ea ni n g le s s , bemuse at the stable node competition w i l l De
zero. However, Rosenzweig (lid l) arguea th a t 1ns*aa of t r y i n g
to relate competition to niche ov erl ap one should rather
discover whether the p o s i t i o n of the sp e c i e s ' e q u i l i b r i u m s are
beiow t h e i r c a r r y i n g c a p a c i t i e s .

The major crltlsm of h i s model I s t l v . t 1t depenos e n t i r e l y on

the theory of optimal foraging (Schoener, 1974; Pyke, e t a l ,
19 7 7 ), of which to date few convincing s t u d i e s are a v a i l a b l e .
Nevertheless, a number of Important Im plications can be
e x t r a c t e d from t h e i r ;nodel. They a re :

( 1) there are th re sh o ld s at which one sp ec ie s w ill

outcompete ano the r.
( 11) when a t moderate d e n s i t i e s , the I n c r e a s e of one sp ecies
may not cause the decrease of another.

Altnough 1t seems likely t h a t zero competition at e q u i l i b r i u m

may e xist when one or more of the resources are l i m i t i n g , 1t<
Importance 1n s t r u c t u r i n g communities seems d o u b tf u l . Posenzwe j
(197$, 1981) believes, as do many others ( e . g . MacArthur and
Levins, 1967; MacArthur ana Wilson, 196"; Le* u i s , 1968; May,
1y 73; Cody, 1974; P ia n k a , 1975; P u l l i a m , 1983), th a t competition
Itse lf g ive s rise to community e q u ilib ria . In n-her words,
 98.

competition m od ifi es the h a b i t a t requlrements or pr e f e re n ce s of

Ihe species, ie. competition behaves as a d i r e c t i v e fo rce in
structuring ecosystems. They b e l i e v e tha t co n tin u al adaptations
Of the components w i t n i n an ecosystem occur as a r e s u l t of t h e i r
response to interspecific competition. An a ltern ative
explanation Is proposed whereby a sp ec ie s' habitat selections
are determined Indepenaently of each other during s p e c i a t l o n ,
and thus overla p r e s u l t s i n c i d e n t a l l y to t h e i r past e v o l u t i o n a r y
h isto ry. Communities would t h e re fo r e c o n s i s t , to a .arge e x t e n t ,
of randomly assembled species. This has app ar ent ly been
explained using models which randomly s t r u c t u r e a known sp ecies
assemblage and then te st f o r s i g n i f i c a n t d i f f e r e n c e s between the
randomly structured an<i tne nat ur al communities (eg. Sale 1974,
Tompson and Rusterholz, 1982). If tne random models are
e sse n tia lly sim ilar to real communities, then one has to favour
the theory tha t community structure is b u i l t up o f a random
a c c r e t i o n of components adapted In a l l o p a t r y ( r e f ) .

F in a lly , It should be stresssed th a t when concerned w'th

estimating sto ck in g densities or carrying capacities 1n
m ulti sp ec ie s systems, the processes which structured the
communities In the p a s t , eg. competition and p r e d a t i o n , are not
as Important as the ecological sim ilaritie s or differences
presently taki ng place. Interest was th e re fo re placed only on
the pre se n t 1 n t e r a c t l o n s , ">r the contempory niche p a t t e r n s , and
not whether competition played an important r o l e in s t r u c t u r i n g
communities 1n the evolutionary sense. In addition, most
communities, especially natjre re se rvti, are comprised of a
number of Introduced and p o s s i b l y a l i e n sp eci es which have oeen
translocated I n to areas which they would n a t u r a l l y not occur
hence, unnatural processes were used to s t r u c t u r e the community.

Thus t h i s study was developed not t e s t whether competition i s

presently occurring between the Pil a n e sb e rg g r a z e r s , but r a t h e r
to relate niche theory 1n e sti m ati ng sto cking d e n s i t i e s 1n a-
m u lt i s p e c ie s system.
 99.

5. 2 Methods:-

5 . 2 . 1 F i e l d Data C o l l e c t i o n :

In order to f u l f i l tne o b j e c t i v e s of t h i s study 1t was necessary

to system atically search for, and tnen measure and record the
c h a ra cte ristic site attributes of each g ra ss l a n d s i t e se le c te d
by the foraging herbivores. From these s i t e s , in formation on
both the habitat structure and dietary selection should be
c o l l e c t e d (see chapter 4 ) .

Based on previous I n v e s t i g a t i o n s (eg. F e r r a r , 1973; F e r r a r and

Walker, 1974; Page and WaWer, 1978; A t t w e l l , 1978; Melton,
1978) and prelimlnary observations, the f o llo w in g dimensions
were found to be important in descr iDing the feeding l i c h e s of
the f i v e la rg e g r i z e r s in the r e s e r v e ;
(i) Habitat ty pe, eg. whether in Secondary Grasslands,
Pediment Grasslands, Valley Savannas, Mesocline Savannas, or
Th1CKe*.s.
( 1i ) u 1e r ary component, 1e. grass s p e c i e s .

The feeding n ic hes of the h e rb iv or es can ow be described along

two dimensions In terms of Hutchinson's multidimensional
hyperspace (Hut ch in son , 1957). A third component, the
subdivisions of the herbaceous compon i t w ithin the major
habitat t y p e s , was analyzed using a m u l t i v a r i a t e techniq known
as Discriminant Fun ction A nalysis. T h is technique e s s e n t i a l l y
reduces the multidimensional herbaceous component to a few
dimensions. Overlap along the h a b i t a t niche dimension could be
measured In mor.> d e t a i l .
 ICO.

5 , 2 . 2 C a l c u l a t i o n o f Niche Overlap

5 . 2 . 2 . 1 S in g l e Dimensions

Overlap has been quantified in numerous ways (MacArtnur and

Levins, i9o7; Schoener, 19bd; Colwell and Futuyma, 1971; P ia n k a ,
197 J ; ) . The p a r t i c u l a r overlap Index used i s somewhat a r b i t r a r y ,
since s i m i l a r q u a l i t a t i v e r e s u l t s are ootained witn a wide a rr a y
of Indices. L i t t l e or no e xp la n a ti o n I s given in the l i t e r a t u r e
a to why a p a r t i c u l a r o v er la p Index was used or not. An attempt
1s therefore made to examine the v a ri o u s methods in d e t a i l ,
emphasising t h e i r pros ana cons.

The two most w id e ly used methods are the Symmetrical and the
Asymmetrical methods. Tne Symmetrical method was f i r s t presented
by Pianka ( 1 9 7 3 ) . Tne equation produce; a symmetrical m a t r i x of
* s so t h a t

^ 1j * aJ 1

1e , the effect of sp ec ie s J on s p e d c 1 1s e q u i v a l e n t to the

e f f e c t of sp e ci e s 1 on sp eci es j and takes the form:

a1j (P 1 k x Pjk) ....5 .2

A i P1 k: )( i P j k 1)

Slo b o d c h l k o ff ana Schulz (19a0) have pointed out three

advantages of this method: (1) a ready geometrical
1n t e r p r e t a t l o r , , (2 ) the symmetrical nature of the m a t r i x , and
( 3 ) the f a c t th a t alpha cannot exceed one.

The Asymmetrical method as proposed by Levins (1968) and

MacArtnur (1 Q7 2) d iffers 1n th-it the ov erl ap Is expressed
r e l a t i v e tc only one of the s p e d e s as f o l lo w s :
 101.

a lj ( P1k x P j k ) ........... 5.3

H P l k )1

1e. it measures how much a sp e c i e s ' u t i l i z a t i o n curve ov er la p s

t h a t of another sp ec ie s c u r v e .

Accjrding to May (1 9 7 5 ) , the alpha m a tri x produc'd by t h i s

equation Is not sym m e tri c a l, 1e; olj w i l l g e n e r a l l y not equal
aj 1. This I s because P1x, the denominator, I s a property o f only
the 1th s p e ci e s (refer to Slo b od ch lk o ff and S c h u l z , 19dO). In
o th e r words, pairwise ov erl ap ( a lj) 1s c a l c u l a t e d only with
re s p e c t to species 1. Values g re a te r t l a n one may now r e s u l t
depending on whether ;pecies j u t i l i z e s resource type k to a
greater e xt e n t tnan sp ec ie s 1, I . e . P ik i s g re a t e r than F j k . Its
major disadvantage 1s th a t one must be extremely c a r e f u l to
count either a ll I n d i v i d u a l s of sp ecies 1 and j , or to count the
same proportion of each sp eci es (Slobodcnlkoff and Scnulz,
198 0). For example, i f sp ecies j !s more s u c c e s s f u l at eludin g
the Investigator than sp e c ie s 1 , i t w i l l skew the overla p value
(u lj), indicating , tha t l e s s com petitiv e pressure from j i s being
e x e r te d on 1 than I t r e a l l y I s . However, the Asymmetrical metnod
remains the most popular f o r c a l c u l a t i n g niche o v e r l a p . T h i s I s
mainly because it does not assume th a t both sp eci es have equal
effects on e*ch other and a l s o , because I t 1 s c l o s e l y r e l a t e o to
the competition c o e f f i c i e n t of tne Lotka - V o l t e r r a fo rmulatlon
of In terspecific competition (MacArthur, 1968; 197<!; Schoener,
1974).

Tne overlap measures are be s t exp la in e d g r a p h i c a l l y using f i g u r e

5.1. If one co n s i d e rs a simple system of two s p e c i e s , one a
sp ecialist and one a g eneralist, foraging along a resource
gradient, then t^e fo l lo w i n g becomes apparent. When using
Planka's Symmetrical method, ov erlap I s c a l c u l a t e d I r r e s p e c t i v e
of the proportional resource use of each species. In other
words, one I s I n t e r e s t e d only 1n the e xte n t to which both share
common r e s o u r c e s , 1e , the degree of s i m i l a r i t y (r e p re s en te d by A
1n F ig u re 5 . 1 ) .
 10 2 .

Frequency

F ig u r e 5.1 : Frequency d i s t r i b u t i o n s o f two s p e c i e s , i and j , f o r the

i t i l i z a t i o n of re so u rce type a .
 103.

If interested 1n the niche overlap with re s pe c t to only one

species, or the influence of one sp ecies on another, then the
Asymmetrical method should be chosen. Tne measure th e r e f o r e has
two different values depending on whether one c o n s i d e rs the
effect of sp e ci e s j on sp ec ie s 1, or v i c e v e r s a . In the f i r s t
instance, ot j i would be c l o s e to or g r e a t e r than I , whereas in
the l a t t e r case a j 1 would oe l e s s than 1 .

Species 1, being more s p e c i a l i z e d iri I t s feeding requlrements,

shar es only a small pro portion of sp ecies J ' s t o t a l reso ur ct
u tlH za tr* Sp ecies j, on the otherhand, 1s more of a
genera'll , and c o n se qu e n tl y , snares a la r g e proportion
of spect total resources. (The resource u tilizatio n
strategies used here are those defined by Schoener, 1977;
MacArtnur, 1972 and Maynard Smith, 1974). When c on si d er in g only
tnose food types which are represented as a , in F ig u re 5 . 1 , then
species 1 u t i l i z e s them to a much g re a te r e x t e n t than sp eci es j .
Therefore, a j1 (overlap with re spe ct to s p e c ie s 1, the
sp ecialist) has a much g r e a t e r value than a 1j ( o v e r l a p with
respect to sp ec ie s J, the g e n e r a l i s t ) . T h i s t h e r e f o r e e x p l a i n s
wny the two ov e rl a p values d iffer. It f o r example, the two
sp ec ie s forage on a broad range of food types then the two
values w ill d iffe r only s l i g h t l y . Th is occurs because both Pik
and P jk have sim ilar values. The use of t h i s method 1s best
Illustrated by r e f e r r i n g to the fol lo win g hypothec c a l example.
The pro portio nal use of four resourc s u; two sp ecies I s iOwn
below;

Species II III IV

1 0.6 0.4 0.0 0.0

2 0.2 0.3 0.4 0. 1

 104.

Spec 1vs 1 being more a s p e c i a l i s t feeder consequently, u t i l i z e s

re so u rce s I and II to a g r e a t e r e xte n t than sp ec ie s 2. T h i s i s
re ve a le d in the ov erl ap measures using the Asymmetrical method,
a 21 * J.80 and a l 2 * U . 4 6 . Species 1, because of i t s g r e a t e r
use of resources I and I I , e x e r t a g re a te r Impact on sp eci es 2,
than vice versa. P i a n k a ' s method, on the otherhand, p r e d i c t s a
s i n g l e (mutual) ov erl ap of 0 . 6 2 .

Tne Asymmetrical method was tn e re fo r e preferred to P i a n k a ' s

method, and was suose^antly used to determine the r e l a t i v e
overlaps, along each dimension ( h a b i t a t and d i e t ) between the
w hit e rhinos and t h e i r a s s o c i a t e d g r a z e r s .

5 . 2 . 2 . 2 Multidimensional Overlap

Once the d e c i s i o n on which measure to use f o r e sti m a ti n g o v erl a p

along single resource g r a d i e n t s ( e . g . d i e t or h a b i t a t ) has been
made, tne next problem l i e s 1n a ss e ss in g the combined e f f e c t s of
n1ch<; ov erlap along more thon one dimension. Cody (1974) and May
(1975) have both pointed out t h a t e i t h e r the Product ( c a l c u l a t e s
the product of the ov e rl a p measures) or the Summation
(calculates the mean of the overla p measures) methods may be
used, depending on whether the resources or dimensions are
dependent or independent. The Product method 1s g e n e r a l l y used
when re sources are Independent or orthogonal to each o t h e r , fo r
example, when a gra ss s p e ci e s or prey Item has an equal chance
of being grazed or captured in any h a b i t a t (May, 1975). The
Summation method Is used fo* dependent resources eg. where any
jrass sp eci es or prey Item occurs In only one p articular
habitat, hence, there Is a d irect relationship between the
presence of grass sp ec ie s and a h a b i t a t type (May, 1975). Thus
the choice o f method depends l a r g e l y on the type of resources or
nlc^ie dimensions th a t one 1s c o n s i d e r i n g .

The Product method calculates the product of the < s along

each dimension,
 105.

a 1jP- TT 011j ( k ) .................... 5 . 4

where, ot1j p Is the product f o r sp eci es 1 and j , and

..1 j (k) 1s the a 1j f o r the kth r e so u rc e . One simply m u l t i p l i e s
the two ov erl ap coefficients for each d ifferent resource
dimenslon.

The Summation method c a l c u l a t e s the average of the P i k ' s along

each dimension and takes the form;

a 1j
* ( a 1j ( k ) ........... 5.5
r

where a 1j s 1s tne summation f o r sp ec ie s 1 and j , and

a ij(k ) Is the a1 j computed f o r tne k ' t h re s ou rc e . T h i s method
generally o v er e st im a te s true niche overlap, whereas
m u ltiplicative overall ov er la p s often underestimate t^ue
m ul tidimensional ov erlap (Pianka, 1974). Furthermore, It is
of te n d ifficu lt tc judge whether resources are dependent or
Independent. Secondly, some resources may have su bt le
Interdependence, such as use of time, use o f food, and use of
space (Slobodchikoff and S c h u l z , 1980). T h i r d l y , the degree of
dependence or independence of p a r t i c u l a r resources m<.y d i f f e r
between the different species. The fou rth problem, which was
pointed out by May (1 9 7 5 ) , concerns tne In accuracy of e st im a ti n g
multidimensional overla p coefficients from single resource
dimensions. Th is 1s best I l l u s t r a t e d by Fig ur e 5.2 which d e p ic ts
two (continuous) resource axes a and b, eg; food and foraging
height, and two - dimensional u t i l i z a t i o n f u n c t io n s f o r the two
species, 1 and 2. It Is c l e a r th a t although the sp eci es are
non-overlapping when viewed along the s i n g l e resource axes and
o v er la p 1s dep icted. Overlap measures for multidimensional
r e s c '""ces may tn e re fo re be overestimated.

F in a lly , another problem which has r.*t been mentioned 1n the

literatu re, 1s the complex phenomenon of e l e c t l v i t l e s for a
 106.

b.
type

12 (b)
Resource

Resource type a.

Figure 5. 2 : Two non-conpetinq s p e c ie s , 1 and 2, depicting overlap

along the two s i n g l e resource dimensions. (Adapted from
Mav, 1975).
 107.

p a rticu lar prey type d lfferir.,, 1n different habitat. For

example, 1t has been observed t h * t the white rhino population a t
P il a n e s b e r g exhibit d i f f e r e n t d i e t a r y e l e c t i v l t i e s 1n d i f f e r e n t
habitats (refer to section a . 3 . 3 ). Consequently, the use of
either the Summation or the Product methods 1n t h i s case 1s
In app ro pla te for determining ov erl ap ac ross se veral h a b i t a t s .
Rusterholz (19dO), a ls o re alizin g their lim itatio ns, used a
combined method whereby he c a l c u l a t e d the mean o v er la p of those
dimensions which c o r r e l a t e d wit h f.-acn other and m u l t i p l i e d t h i s
by the niche ov erlap along the potentially Independent
dimension. Although his method improves the e sti m ate of true
m u l tl d e n s l o n a l o verlap 1t I s by no means p e r f e c t . A m o d if i c a ti o n
of both the Asymmetrical and the Product methods has been
proposed 1n an attempt to circumvent these proDlems.

When using the Asymmetrical or the Symmetrical methods, alphas

for Doth the haoitat and dietary dimensions are c a l c u l a t e d
separately. In other words, the d i e t a r y o v er la p c o e f f i c i e n t s
between sp ec ie s 1 and sp ec ie s 2 are assumed to be equal
throughout each h a b i t a t type. Th is i s not c o r r e c t i f the d i e t a r y
e le c tivltie s or eacn h er bi vo re sp eci es d i f f e r s In each h a b i t a t .
The m odiflcatlons must t h e re fo re first c a l c u l a t e the d i e t a r y
overlaps within each habitat type, and then in cor por ate the
haoitat o v e rl a p s f or each h erolvore (consumer) s p e c i e s . The
two-dimensional ov erl ap equation would th e re fo r e take the form;

01 1j * (* PlkH x PjkH j x / elH x ejH \ x art . . . 5 5 .6

l\ ( (P1krt)i / \ ( e l H )2 /

where 1 , j * h er bi vo re s p e c i e s ,
H habitat, l . . . n ,
k p l a n t sp eci es 1n d i e t , 1 . . . S ,
P1kH a proportion c f p l a n t sp ecies k 1n sp ec ie s 1 's u i e t 1n
h a b i t . i t H,
 108.

eiH a proportion of time th a t he rbivore s p e c i e s i spent in

h a b i t a t H, and
aH * proportion of the to t a l are^ made up by h a b i t a t rl.

The first part of the equation c a l c u l a t e s tne d i e t a r y overlap

w ithin each h a b i t a t using the Asymmetrical method as proposed by
Pianka (1 9 7 3 ) . Tn is value I s then m odlficu or w e i g h t ' d , f i r s t l y
by tne h a b i t a t , ov erl ap and secondly by the pro portio nal area of
that h a b i t a t . T h i s method, termed tne "Weighted Overlap" method,
calculates v a lu e s from 0 to 1 , again 0 i n d i c a t i n g no o v e r l a p ,
and 1 I n d i c a t i n g complete o v e r l a p .

By >mg this approach one I s I n co rp or a tin g the r e l a t i o n s h i p

between boln the habitat and d i e t a r y s e l e c t i o n . For example,
s p e c ie s 1 may ov erl ap greatly 1n diet w1tn sp ec ie s j but,
sp e c i e s j may spend l e s s time in tne h a b i t a t , thereby decreasing
the potential ov erl ap between them. This value 1s then modified
by the proportional area of h a b i t a t ' H' 1n the r e s e r v e . Hence,
the t o ta l overla p in the re la tive ly s m a ll er habitats are
weighted l e s s than those 1n the l a r g e r h a b i t a t s . The new metnod
proposed Is, therefore, not so much a modification of the
mathematical formulae as an a l t e r n a t i v e In the approach used to
c a l c u l a t e resource overla p along m u l t i p l e niche dimensions.

5.2.2 .3 Reduced Dimensional 1ty 1n the Measurement of H a bi ta t

ov e rl a p

Hu tch in so n 's (1950 concept of the ecological niche as a n -

dimensional hypervolume Invites the use of m ultivariate
sta tistica l tech niqu es . Rote.iberry and Wiens (198U) st re s s e d the
advantage of such methods 1n being able to d i s p l a y hi ghl y
correlated variab les along Independent axes re ; esenting what
they termed, "proximate" niche dimensions. Two w id e ly accepted
tec h n iq u e s, namely, P rin cip le Component A n a l y s i s ( P . C . A . ) and
Discriminant f u n c t io n A n a l y s i s ( D. F . A. ' have been used in niche
analyses.
 109.

The major advantage of D . F . A . 1s th a t the D i s c r i m i n a n t Functions

can be used, f i r s t l y , to I d e n t i f y the axes of m : h s e p a r a t i o n ,
and se con dly, the d i s p e r s i o n o f l o c a t i o n s along these axes can
be I n t e r p r e t e d as re pr e se n tin g niche width or s p e c i a l i z a t i o n .

Levins (1968) i n i t i a l l y sta te d th a t "niche d i m e n s i o n a l i t y r e f e r s

not to the number of b io lo g ically relevant factors on the
environment........... but to the numoer of factors which serve to
s e pa ra te species". T n i s led Cody ( I 9 6 0 ) and James (1971) to use
D .F.A. to separate bird sp eci es according to habitat
cn a ra cte ristics. However, re ce nt a p p l i c a t i o n s to niche an aly se s
appear to have been based on the papers by Green (1971, 1974)
and Shugart and Pitten (1 9 7 2 ) . Green (1 9 7 1 ) , used D . F . A . to
Id en tify sig nifican t e co l o g i c a l f a c t o r s se par at in g sp ec ie s In a
multi dim en si on al space. Tne standard d e v i a t i o r of d i s c r i m i n a n t
sc or e s as a measure of niche breadtn, and the percent ov erlap of
50 p ro o jb llity e l l i p s e s a neasure of niche o v e r l a p . Dueser and
Shugart (197d, 1979), on the other hand, used the average
distance of tne ob se r v a ti o n s fo r a sp ec ie s from the o r i g i n of
d i s c - 1m1nant spacc* as a measure of niche exp loitatio n or
position re la tive to the average m lc ro h aD it at a v a i l a b l e . The
co e fficie n t of v a r i a t i o n of the d i s t a n c e s from the d i ' c r u . l n a n t
orig in was used as a measure of niche breadth. I n d i c e s of
ov erl ap were calculated using the p la n a r area o f ov erlap of
co n c e n t r a t i o n e llip se s r e l a t i v e to t n e i r t o t a l area (Dueser and
S h u g a r t, 1979).

A controversial problem which arises when c o n si d er in g

m ultivariate a n a ly s e s of h a b U a t use or niche space i s whetner
to sample environmental v a r i a b l e s ut s i t e s determined randomly,
or to record the v a r i a b l e s only where the organisms o f I n t e r e s t
were sited. Rotenberry and Wiens (1980, 1 9 b i ) , used the former
method f o r generating a s e t of h a o i t a t v e c t o r s to be analysed by
P .C .A . Cody ( 19C8, 1978), James (1 9 7 1 ) , Grten (19 71, 1974),
Shugart and Patteri (197^), Ferrar and Walker (1 9 7 4 ) , A t t w f l l
(1975), M'Closky (1 9 7 6 ) , and Dueser and Shugart (1978, 1979),
however, a ll used the la tte r method (1 e , c e n t e r s or singing
perches w ithin t e r r i t o r i e s f o r b i r d s , and su c c e s s f u l c o l l e c t i n g
sites for mollus cs and mammals) to Identify locations fo r
 110.

habitat sampling. Green (1 9 7 1 ) , when j u s t i f y i n g h i s method of

re cor ding environmenta1 v a r i a b l e s only vhere the organisms ( i e ,
molluscs) were found, c i t e d Hutchinson's (1968) reasoning that
the " p rt - e n c e of a '. p e d e s in a sample was more i n f o r m a t i v e than
its a bse nc *" . Presence indicates th a t a sp ec ie s could l i v e a t
tne site , whereas absence could a r i s e Decause, e i t h e r the s i t e
was unsuitable, tne sp ec ie s had lot y e t col on iz ed the a r e a , or
because the sp e c ie s was misstd in the sample even though i t was
a favou rable s U e (Hutch inso n, 1968). Carey (1981) and Wiens and
Rotenoerr> (1981), however, argued t h a t t n l s technique may only
be meaningful for sedentary forms, or when ap p lie d on a
macroscopic scale. Mobile animals may 1n f a c t be found not only
in marginal habitats but also, during high densities, In
completely unsuitable haoltats. Presence Is tn e r e f o r e not
Indicative of haoitat su itab ility. However, the author argues
that If one records the sites only where the animals are
observed to forage then a l l the s i t e s measured are p a rt of the
s p e c i e s feeding n i c h e .

The major d i f f e r e n c e between these two sampling techniques l i e s

in the Interpretations of the canonical space. I f one accepts
that the origin of canonical space rep re se n ts the mean of the
dlsrim lnant (or factor) scores, or the mean of a ll sites
sampled, then the niche m e tr i c s w i l l d i f f e r f o r each sampling
method. In the f 1 method, the o r i g i n g ives a o e t t e r estimate
of tne averac, ;rohab1tat available. The second method,
however, es tim ate., only the average of the m i c r o h a b i t a t u t i l i z e d
by the species, and consequently, 1s In fl u e n c e d by their
relative abundances (Carnes and slade, 1982). Common sp eci es
would contribute more to the determination of the mean than the
rare species, and t h e r e f o r e , are more l i k e l y to be lo ca ted near
the o r i g i n of the can on ical space.

When Information 1s re quired on niche s p e c i a l i z a t i o n s or niche

b re a d th s, samples of sp ec ie s absence must be i n cl u d e d . However,
measurements of the s i t e s where the organisms were p re se nt i.eed
only be recorded If i n t e r e s t e d In niche o v e r l a p . In t h i s case
one Is prim arily Interested 1n the r e l a t i o n s h i p s between the
different species, rather than between the s p e c ie s and t h e i r
envi ronment.

During this study samples were measured only a t the s i t e s where

the h e r bi v or es were present in order to d e s c ri b e the
m i c r o n a b i t a t s e l e c t i o n w i t h i n the major h a b i t a t s .

The data collected consisted of se veral groups ( h e rb i v o re

species) each comprising a numoer of individual sites
characterized by a numuer of s i t e a t t r i o u t e s . The v a r i a b l e s take
the form of e i t h e r measured (eg. grass height) estimated scores
( e g . percentage herbaceous c o e r ) .

5 . 2 . 2 . 3 . 1 01s c r i m i n a n t Function A n a l y s i s

Discriminant Functio n Analysis (DFA) is a w ell known

m ultivariate sta tistica l technique orig fnally developed by
Fisher ( 19) as a means of c l a s s i f i c a t i o n . The method se parates
popu latio ns or groups which are known to be d i f f i c u l t , on the
basis of se v e ra l characters or a t t r i b u t e s . It is particu larly
u se fu l 1n niche analysis because it reduces the number of
c rite ria or dimensions ustu fo r discrim ination before the
1n t e r p r e t a l o n of sp ec ie s se par ati on 1s necessary (Green, 1971;
F e r r a r , 1973).

The first step In t h i s a n a l y s i s i s the combination o f F - r a t l o s

for each variab le so as to I d e n t i f y those v a r i a b l e s f o r which
the r a t i o of between sp ec ie s v ar ia n ce to w i t h i n sp e c i e s v a ri a n c e
Is s u f f i c i e n t l y la rg e to enable d i s c r i m i n a t i o n between h er bi vo re
species (Ferrar and Walker, 1974). Secondly, a s e r i e s o f l a t e n t
ro ots (discrim inant functions) and v e c t o rs are e x t r a c t e d from
the to ta l data set. These d i s c r i m i n a n t f u nc t io n s e xp res s the
difference between sp e c ie s In terms of a few common g r a d i e n ts of
variation In ste a d of the g ra d ie n ts represented by each of the
Individual variab les. The l a t e n t roots then give the proportion
of the t o ta l v ar ia n ce e x t r a c t e d with the vector o f the v a r i a b l e
weights indicating the degree to which each v ariab le is
a s s o c i a t e d with tne d i s c r i m i n a n t f u n c t i o n .

Discrim ination between groups i s th e re fo re expressed in terms of

a ^ew common g r a d i e n t s c h ic h can be i d e n t i f i e d by those v a r i a b l e
weights most s t r o n g l y a s s o c i a te d w i M each d i s c r i m i n a n t fu nc tio n
(Ferrar and Walker, 1974). The next step then c a l c u l a t e s a
d iscrim inant score f o r each s i t e (feeding l o c a l i t y ) , by summing
the products of the variable weights and their respective
values. The mean of the d i s c r i m i n a n t scores f or a l l s i t e s for
each sp eci es 1s known as the group or species c e n t r o s J ( F e r r a r
and w a l k e r , 1974).

D ls c r i m a n t Function A nalysis fin ally provides a posteriori

cla ssifica tio n f u n c t io n which can be used to a s se s s the e x te n t
of group overla p based on tne s i t e c h a r a c t e r i s t i c s ( T e l f o r d ,
1982).

The program algor ith m used f o r t h i s a n a l y s i s i s tn a t of Kleck a

( 1975) and was performed using an S . P . S . S package ( N i e , e t a l ,
1975).
Author Borthwick Michael Robert
Name of thesis Habitat Use By The White Rhinoceros In Relation To Other Grazing Ungulates In Pilanesberg Game
Reserve, Bophuthatswana. 1986

PUBLISHER:
University of the Witwatersrand, Johannesburg
2013

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