nutrients and toxins of plants in amboseli, kenyacomposition of amboseli plants 28 1 the solution....
TRANSCRIPT
Afr. J . Ecol. 1987, Volume 25, pages 279-293
Nutrients and toxins of plants in Amboseli, Kenya
S . A. A L T M A N N * , D. G . P O S T t a n d D . F . K L E I N $ Department of Biology, University of Chicago, U.S.A. and Institute of Primate Research, National Museum of Kenya, ?Department of Anthropology, Columbia University, and $Department oj’Anthropology, New York University, U.S.A.
Summary Nutrient and toxin analyses were carried out on wild foods that are eaten by the baboon (Papio cynocephalus (L.)) and black-faced vervet (Cercopithecus aethiops (L.)) and other animals in Amboseli National Park, Kenya. Data are provided on proximate composition (water, ash, fat, protein, fibre, other carbohydrates), fourteen minerals, four vitamins (folic acid, ascorbic acid, riboflavin, P-carotine) and a toxin (trypsin inhibitor). In total, 864 analyses were carried out on ninety-one foods from thirty-seven species of plants.
Resume On a realist des analyses des nutriments at toxines dans I’alimentation du babouin (Papio cynocephalus (L.)), du cercopitheque grivet (Cercopithecus aethiops (L.)) et d’autres animaux du Parc National d’Amboseli, Kenya. Les donnees fournies concernent la composition approximative en eau, carbone, graisse, protkine, fibres et autres carbohydrates. Quatorze mintraux, quatre vitamines (acide folique, acide ascorbique, riboflavine, carotene p) et un toxine (inhibitrice de la trypsin). Au total, 864 analyses furent realistes sur nonante et un echantillons protant sur trente-sept especes de plantes.
Introduction As part of our researches on the behavioural ecology of primates (baboons and vervets) in the Amboseli region of southern Kenya, the nutritional compositions of many plant foods in the area are being analysed. Our results to date are summarized in this report. These data may be useful to others who are studying the foraging of animals in Amboseli or elsewhere.
Nutrients and toxins reported here include those obtained in a standard ‘proximate analysis’ (water, ash, lipids, proteins, fibres and non-fibre carbo- hydrates), several vitamins (folic acid, ascorbic acid, riboflavin, provitamin A), a toxin (trypsin inhibitor) and several mineral elements (P, K, Ca, Mg, Na, Al, Ba, Fe, Sr, B, Cu, Zn, Mn and Cr).
Materials and methods All of the plant material was collected within the area formerly designated as the Maasai-Amboseli Game Reserve in southern Kenya, Africa. The habitat consists
*Correspondence: Prof. S. Altmann, 940 East 57th Street, Chicago, IL 60637, U.S.A. Present addresses: t3225 33rd Place NW, Washington, DC 2008, U.S.A. and SPratt Institute, Brooklyn,
NY 11205, U S A .
279
280 S. A . Altmann, D . G. Post and D . F. Klein
primarily of savanna with scattered acacia trees. Permanent waterholes and swamps in the area have their own distinctive associated flora. The collection area of about 100 km2 lies primarily within the boundaries of Amboseli National Park.
The collection area is bounded by Enkongo Narok Swamp on the east and north-east, by Nairabala Hill and the dry bed of Lake Amboseli on the west and north-west, and extends southward about 1 km south of Endoinyo Ositet Hill. The area is shown on the Kenya 1 : 50,000 map Series Y73 1, Sheet 18 1 / 1, Edition 2-SK (Survey of Kenya, 1963), in Squares BH and CH. Longitude 37’10’E and latitude 2’40‘s run through the approximate centre of the area.
All of the samples were collected in 1973-1976 or 1980 during the phenophases in which they were being eaten by baboons or vervets, from within the home ranges of these animals. For each food, we attempted to collect at least 20g each of representative samples of exactly what these primates eat, paying attention to plant part, size, colour, degree of ripeness, rejected components (e.g. rind, calyx, seeds, insect-infested material) and so forth. In some cases, material was analysed because the animals overtly avoided or discarded it, e.g. the seed coats of green seeds from umbrella trees (Acacia tortilis).
Further details on the role of these materials in the diets of baboons and vervets can be found in Altmann (1987), Klein (1978), Post (1978), Post et a/ . (1980). Many of these plant foods are eaten by other animals of the region, although for none of these other animals has foraging been investigated as intensively.
All samples with a number ending with /74 and all of the form M**/75 (where ** indicates two or three digits) were collected by D.P. All other samples whose references end with /75 and all of those ending /76 were collected by D.K. All others (designated 1 A, 1 B, 2A, 2B, etc.) were collected by S.A. except for 28B, collected by D.K. In many cases, one of the other co-authors or various other colleagues (see Acknowledgments) also helped.
The samples were preserved in one of three ways. For proximate analyses, most of S.A.’s and D.K.’s samples were frozen. This was done as soon after collecting as practical, usually within 1-2 h. These samples were collected in plastic-lined bags designed to hold freeze-dried foods (Foil Pak no. 6, Champion Packages Co., Columbus, GA). Particular care was taken with foods that were likely to desiccate rapidly after picking, e.g. the ovaries of Trianthema ceratosepala. The collecting bags were kept in the shade and were kept closed. The samples were collected, weighed, and frozen as quickly as possible.
After being sealed, the collecting bags were in turn sealed into clear polythene bags, to avoid changes in weight from surface condensation, then placed in a freezer or the freezing compartment of a refrigerator at 01 Tukai, in Amboseli. At about monthly intervals, these samples were packed with a bag of ice, wrapped in newspaper or other insulating material, and taken to Nairobi. There the samples were kept in a freezer until they were analysed. Other samples used for proximate analysis, including all of D.P.’s samples and the few samples used for mineral analysis, were air-dried, either in the sun (D.K.) or above an outdoor ‘oven’ (a heavy-gauge, steel ammunition box) that was kept warm by the coals of our campfire (D.P.).
For analysis of vitamins and trypsin inhibitor, samples were stabilized by placing them immediately after picking into 3-5% oxalic acid, in UV-proof jars. Very small items, such as the ovaries of T. ceratosepala, were placed directly into
Composition of Amboseli plants 28 1
the solution. Larger items, such as the berries of Azima tetracantha, were first lanced, or cut into pieces. These wet-preserved samples were kept in a dark cabinet at our base camp at 01 Tukai until our next trip (usually monthly) to Nairobi, from which they were shipped by air freight to the Nutrition Division of the WARF Institute, in Madison, Wisconsin, where all of the vitamin analyses were carried out. Unfortunately, we did not know, at the time that we made our field collections, that Vitamin A and folicin are unstable in acid media (Altmann, 1979). Therefore, our values for these vitamins are probably underestimates. The preservation method that was used for each sample is indicated in the tabulated results below.
Quantitative analyses were carried out by three laboratories: the WARF Institute Inc., in Madison, Wisconsin, the National Agricultural Laboratories, Nairobi, and the Nutrition Laboratory, Department of Animal Production. University of Nairobi, Kabete. The methods of analysis, where indicated to us by the laboratories, are referenced in the Tables.
Except where data on water content were inadvertently missed, all analyses were converted to a fresh-matter basis, in order to compute the actual nutrient intakes of animals (Altmann, 1987).
At present, we have only limited tests of the consistency (replicability) of these chemical analyses and no tests of their accuracy (Altmann, unpubl). Although Table 1 lists several foods that were analysed more than once, the dates and places of collection for most of these materials were usually quite remote and the material was gathered by different people, so that one cannot rule out heterogeneity (sample-to-sample differences) as an explanation for any differences in composition. In one case, however, an attempt was made to check on consistency by analysing matched samples. Dry seeds of fever trees (Acacia xanthophloea) were divided into three piles. One pile (Sample 20B-2) was sent to WARF, the other two (21 B, 22B), to the Nutrition Laboratory of the University of Nairobi. In each case, a proximate analysis was requested but the laboratories were not notified that these samples were in any way special. The difference between the largest and smallest of the three values for each nutrient are as follows: water 0%, minerals 0.2%, lipids 0.5%, proteins 1.0%, fibres 8.1%, soluble carbohydrates 7.6%. There is a large discrepancy for fibre, and therefore also for soluble carbohydrates, since the amount of the latter is determined ‘by difference’, i.e. by subtracting the sum of all other values from 100%. The discrepancy is primarily between laboratories: the two samples analysed in Kenya differ in fibre content by only 0.9%.
Results
The results of proximate analyses (water, ash, lipids, proteins, fibres and soluble carbohydrates) are given in Table 1 and those for several vitamins and trypsin inhibitor are given in Table 2. Results for mineral elements are given in Table 3.
In these Tables, each food is given a four-letter code. These codes are used to facilitate computerized data analysis of the data in these Tables (Altmann, 1987). The first two letters represent the taxon (species, where known). The third letter indicates the part of the plant: leaves, flowers, etc. The fourth letter represents the ‘condition’, e.g. degree of ripeness.
In Table 1, four assumed values are explained in the footnotes. One requires further comment. Our one laboratory report (721-75) for protein in dry A . tortilis
Tab
le 1.
Pro
xim
ate c
ompo
sitio
ns
Dat
e of
Sp
ecie
s co
llect
ion
Ref
. no.
(m
o/da
y/yr
) C
ode
Uni
ts
gof
Oth
er
Pres
erva
tion,
Pl
ant
in
fres
h go
f ca
rbo-
la
bora
tory
5 pa
rt
sam
ple
sam
ple
unit
Wat
er
Ash
L
ipid
s Pr
otei
n Fi
bre
hydr
ates
an
dmet
hods
**
Abu
tilon
mau
ritia
num
64
2176
05
/16/
76
AB
BU
A
caci
a to
riili
s 13
B 03
/07/
76
TOB
S M
55/7
5 02
/07/
75
TOB
S 12
/76
01/2
8/76
TO
BS
13/7
6 01
/28/
76
TO
BU
2B
08/0
2/74
TO
DU
+Y
U
38B
07
/20/
76
TO
DU
M
331/
75
07/1
3/75
T
OD
U
6531
76
07/2
2/76
T
OG
U
7171
75
10/0
3/75
T
OL
U
7181
75
10/0
3/75
T
OL
U
7191
75
IO/O
3/75
T
OL
U
39B
07
/20/
76
TO
YU
40
B
07/2
0/76
T
OPU
M
332/
75
07/1
3/75
T
OPU
M
333/
75
07/1
3/75
TO
PU+
DU
36
5176
05
/04/
76
TOQ
U
Aca
cia
xanf
hoph
loea
18
92/7
4 10
/24/
74
FTB
R
5B
09/2
8/75
FT
BS
7201
75
-09/
25/7
5 FT
BS
20
52
04
/29/
76
FTD
R
21B
04
/29/
76
FTD
R
22B
04
/29/
76
FTD
R
1899
174
10/2
4/74
FT
DR
7211
75
-09/
25/7
5 T
OD
R
Flow
er b
uds
Flow
ers
Flow
ers
Flow
ers
Flow
er b
uds
Dry
seed
s G
reen
seed
s+co
ats
Gre
en se
eds-
coat
s G
reen
seed
s -co
ats
Sap
You
ng g
reen
leav
es
You
ng g
reen
leav
es
You
ng gr
een
leav
es
Coa
ts o
f gre
en see
ds
Gre
en p
ods - se
eds
Gre
en p
ods - se
eds
Gre
en p
ods +
seeds
G
reen
thor
ns
Det
ache
d flo
wer
s Fl
ower
s Fl
ower
s D
ry se
eds
Dry
seed
s D
ry se
eds
Dry
seed
s-co
ats
200
15.9
8 0.
080
84.3
361
3240
0.090
700
54.3
0 0.
078
485
24.7
0 0.
051
125
30.0
0 0.
24
260
17.5
5 0.
068
1429
59
.78
0,04
2 49
7 38
.10
0.07
7 22
4.15
478
3030
0.
063
422
16.4
5 0.
039
1425
69
.03
0,04
8 15
5 24
1.50
1.
56
51
75.5
0 1.
48
158
352.
90
2.23
37
5 21
.18
0.05
6
80.8
81
.0
78.9
80
.5
14.3
69
.9
65.8
69
.9
94.0
69
.9
74.2
69
.9
56.7
67
.1
68.7
72
.4
78.5
1500
81
.00
0.05
4 -
142
24.9
5 0.
18
74.1
28
1 35
.10
0.12
75
.0
34.6
5 8.
9 51
31
24.9
5 0.
005
8.9
2460
8.
9 5 1
.20
12.0
1.3
0.6 1.7
I .2
1.1
3.2
1.2
1.8
1.3
2.0 1.6
1.9
1 .o
2.3
2.1 1.9
0.6
(7.0
) I .
3 1.
8 3.
6 3.
7 3.
8 4.
3
0.5
2.2 1.5
2.2
1.9
3.6
0.6
0.4
0.9
0.2
1.3
1 .o
1.3
0.6
0.4
0.7
0.8
0.9
(4.9
) 0.
8 1.
2 4.
8 4.
3 4.
3 2.
4
3.5
3.7
4.2
4.4
4.8
20.8
7.
3 3.
3 7.
3 1.8
5.2
3.3
2.8
3.0
4.0
6.0
(1 1.
3)
1 .o
1.2
19.1
18
.1
18.9
18
.8
3.2
3.1
4.4
3.7
3.5
17.7
4.
6 4.
2 5.
7 0.
5 3.
6
3.8
12.6
10
.3
11.3
8.
6 3.
6
(1 5.
0)
3.9
4.5
15.6
22
.8
23.7
23
.0
7.2
9.7
7.2
9.7
8.2
16.3
(b)
24.4
14
43
1.5
4@4(
a)S
18.0
25
.9
17.2
14
.2
12.3
10
.5
(61.
8)
19.0
16
.3
48.0
42
.3
40.4
39
.5
F, U
NN
Lt
F, U
NN
L
D, U
NN
L
F, U
NN
L
F, U
NN
L
N, U
NN
L
F, U
NN
L
D, U
NN
L
N, U
NN
L
F, U
NN
L
F, U
NN
L
F, U
NN
L
F, U
NN
L
F, U
NN
L
D, U
NN
L
D, U
"L
F,
UN
NL
D, U
NN
L
F, U
NN
L
F, U
NN
L
N, U
NN
L
N, U
NN
L
N, U
NN
L
D, U
NN
L
D, W
AR
F-I
5017
6 02
/20/
76
FTD
U
1981
.3
12/1
5/81
FT
DU
14
B
03/0
7/76
FT
GR
15
B
03/0
7/76
FT
GU
18
9417
4 10
/24/
74
FTG
X
2101
174
12/1
2/74
FT
GX
24
4176
<0
4/13
/76
FTLU
64
7176
07
/27/
76
FTLU
32
B
06/0
1/76
FT
PR
1981
.4
12/ 1
5/81
FT
PU
1981
.1
12/1
5/81
FT
PU+D
U
2094
174
12/0
9/74
FTP
U +
DU
781/
75
12/0
3/75
FTP
U +
DU
51/7
6 <0
4/13
/76
FTQ
U
M33
8/75
07
/13/
75
FTW
C
Ach
yran
thes
asp
era
3601
76
04/2
5/76
A
PLU
A
mar
anth
us g
raec
izan
s 7/
76
12/2
0/75
AZW
U +
LU
3631
76
04/2
5/76
AZW
U +
LU
Azi
ma
tetr
acan
tha
9B
061 1
2/76
A
TFR
77
7175
11
/03/
75
ATF
R
1895
174
10/2
4/74
A
TFS
34B
06
1 12/
76
AT
FU
M32
5/75
07
/12/
75
AT
FU
Chl
orop
hytu
m s
p. n
r. ba
keri
19
B
04/2
9/76
C
BL
U
Com
mic
arpu
s plu
mba
gine
us
30B
05
/29/
76
SFFR
53
2176
05
/05/
76
SFLU
C
ordi
a gha
raf
M45
/75
02/0
2/75
C
FLU
Gre
en s
eeds
-coa
ts
Gre
en s
eeds
+ coa
ts
Bro
wn/
brow
n-bl
ack
gum
C
lear
lligh
t am
ber g
um
Gum
G
um
You
ng g
reen
leav
es
Mat
ure
gree
n le
afle
ts
Gre
en p
ods +
seeds
G
reen
pod
s - se
eds
You
ng g
reen
pod
s+ s
eeds
G
reen
pod
s + see
ds
Gre
en p
ods+
see
ds
Gre
en th
orns
C
ambi
um
You
ng le
aves
Leav
es + s
tem
s Le
aves
+ ste
ms
Rip
e fr
uits
Ripe fr
uits
Se
mi-r
ipe f
ruits
U
nrip
e fr
uits
U
nrip
e fr
uits
Gre
en le
aves
Rip
e fr
uits
Y
oung
gre
en le
aves
Gre
en le
aves
551
939
338
200
334
143
I86
500 97
950
19.9
5 0.
036
39.7
0 0.
042
136.
50
41.9
5 86
.40
130.
90
33.7
0 0-
10
25.5
5 0.
13
86.7
5 0.
26
56.2
5 0.
39
62.9
0 0.
34
144.
00
0.29
24
.15
0.25
51
.30
0.05
4 27
.50
75.3
2.
0
18.0
4.
3 24
.8
2.0
31.6
15
.1
11.2
3-
1 72
.2
1.9
68.8
1.
8
-
-
(5.4
) -
-
-
-
-
77.3
2.
0 75
.3
I .6
(2.7
) 60
.1
4.9
-
2.6
1.12
2.
1 0.
2 0.
5 1.
6 0.
7 0.
5 (1
.7)
0.15
0.90
0.9
1.4
0.5
(9.1
)
10.7
10
.8
1.4
1.2
1.7
0.6
5.9
7.5
4.19
6.
8 1
4.1
3.5
(20.
9)
I .9
(9.7
)
800
83.3
0 0.
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86-3
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3
450H
51
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2.2
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4 89
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2.8
0.2
2.6
360
117.
43 0
.33
75.4
2.
7 3.
0 4.
9 55
0 12
8.35
0.2
3 73
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2.7
2.2
4.6
350
90.8
0 0.
26
76.5
3.
2 2.8
5.
3 33
5 76
.28
0.23
72
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3.6
3.7
6.9
158
31.2
0 0.
20
70.7
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4 3.
9 7.
2
50
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8 0.
52
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3988
32
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8 57
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9.6
13.0
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0.12
65
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6.5
1.2
12.4
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88
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2.22
79
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1.3
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18
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3.5
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0.
85
10.3
6.
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13
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(39.
4)
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c)
55.8
70
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42.8
82
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15.6
18
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8)
-
-
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0.4)
19
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4.4
12.1
3-
3
11.1
12
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9.1
8.8
7.9
8.6
10.8
10
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8.1
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NN
L
F, N
AL
N
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RF-
3 D
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NL
D
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NL
D
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NL
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UN
NL
N
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NL
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NA
L
F, N
AL
D
, UN
NL
F.
UN
NL
D
, UN
NL
D
. UN
NL
F, U
NN
L
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L
9
F,U
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L
'Q 9
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NN
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F,U
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L
%.
D,U
NN
L
5 F
,UN
NL
9
D,U
NN
L
b
F?,U
NN
L
& F,
UN
NL
3 2
F,U
NN
L
'Q 2
D,U
NN
L
$ C
on!.
h,
00
W
Dat
e of
Sp
ecie
s co
llect
ion
Ref
. no.
(m
oida
y yr
) C
ode
Uni
ts
gof
Oth
er
Pres
erva
tion,
Pl
ant
in
fres
h go
f ca
rbo-
la
bora
tory
5 pa
rt
sam
ple
sam
ple
unit
Wat
er
Ash
Li
pids
Pr
otei
n Fi
bre
hydr
ates
an
d m
etho
ds**
Cyno
don
dai t
I Io
n M
90 7
5 03
15
75
CD
DU
8B
II
29/
75
CD
LU
8B
S 12
09
75
CD
LU
24
2/76
<0
4 13
76
CD
DU
+L
U
1893
174
10!2
4;74
53
I !76
05
I 15/
76
6431
76
07/0
8/76
C
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NL
D
, UN
NL
*All
valu
es a
re p
erce
ntag
es b
y w
eigh
t of f
resh
sam
ple e
xcep
t fig
ures
in p
aren
thes
es w
hich
are
give
n on
an
anhy
drou
s bas
is.
t(D
) =dr
ied;
(F)
=fr
ozen
; (H
) = ha
ndfu
ls; (
N) =
no p
rese
rvat
ive;
(T) =
tille
rs.
$(a)
=Pro
tein
val
ue a
ssum
ed;
(b)=
Wat
er c
onte
nt a
ssum
ed f
rom
M33
1/75
; (c)
= A
ssum
ed w
ater
con
tent
fro
m 7
8117
5; (
d)=
Wat
er c
onte
nt a
ssum
ed f
rom
189
7174
; (e
) = F
ibre
val
ue a
ssum
ed.
§Lab
orat
orie
s: N
AL
= N
atio
nal
Agr
icul
tura
l La
bora
torie
s, N
airo
bi.
UN
NL
= U
nive
rsity
of
Nai
robi
Nut
ritio
n La
bora
tory
, D
epar
tmen
t of
Ani
mal
Pro
duct
ion.
W
AR
F =
WA
RF
Inst
itute
, Inc
., M
adis
on, W
isco
nsin
. **
Met
hods
UN
NL
: W
ater
by
air
oven
, ash
by
muf
fle fu
rnac
e 60
0"C
, fat
by
Soxh
elt-d
ieth
yl e
ther
ext
ract
, pro
tein
by
mac
roK
jeld
ahl,
fibre
by
mul
tiple
ext
ract
ion
of
nutri
ents
, car
bohy
drat
es b
y di
ffer
ence
. W
AR
F-I:
Wat
er b
y va
cuum
ove
n A
OA
C 1
22,1
970,
l Ith
editi
on; a
sh by
AO
AC
123
,197
0,l I
thed
ition
; fat
by
AO
AC
128
,197
0,l I
thed
ition
; pro
tein
by
AO
AC
16,
1970
, I 1
th ed
ition
(%N
x 6.
25);
fibre
by A
OA
C 1
29,1
970,
11 th
editi
on; c
arbo
hydr
ates
by
diff
eren
ce; f
olic
acid
by
AO
AC
830
,195
5,8t
hedi
tion;
ribo
flavi
n by
AO
AC
789
,197
0, I
1 th
editi
on; a
scor
bic
acid
by
J. B
iol.
Che
m. 1
47,3
99, 1
943.
W
AR
F-2:
Sam
e as W
AR
F-I
exce
pt ri
bofla
vin
by A
OA
C 8
46, 1
975,
43.1
39.
WA
RF-
3: W
ater
by
AO
AC
394
, 197
5, 1
2th e
ditio
n, 2
2.01
3; a
sh b
y A
OA
C 2
22, 1
975,
12t
h edi
tion,
14.
006;
fat b
y A
OA
C 1
35, 1
975,
12t
h edi
tion,
7.0
47; p
rote
in b
y A
OA
C
15. 1
975,
12t
h ed
ition
, 2.0
49 (%N
x 6.
25);
fibre
by
AO
AC
137
, 197
5, 1
2th
editi
on, 7
.054
; car
bohy
drat
e by
diff
eren
ce; f
olic
aci
d by
AO
AC
770
, 196
5, 3
9.05
9; ri
bofla
vin
by
AO
AC
846
, 197
5, 1
2th e
ditio
n, 4
3.13
9.
WA
RF-
4: s
ame
as W
AR
F-3
exce
pt w
ater
by
AO
AC
282
, 197
5,12
th ed
ition
, 16.
217;
folic
acid
by
AO
AC
830
, 195
5,38
.48;
anti-
tryps
in b
y C
erea
l Che
mis
try 5
1: 3
7638
2,
1974
. WA
RF-
5: sa
me
as W
AR
F-4
exce
pt fo
lic ac
id b
y A
OA
C 7
70, 1
965,
39.0
59; p
-car
oten
e by
Ing
. Eng
. Che
m.,
Ana
l. Ed
., 13
,600
, 194
1.
WA
RF-
6: s
ame
as W
AR
F-5
exce
pt fo
lic ac
id b
y A
m. J
. Clin
. Nut
r. 2
1(10
): 12
02, 1
968;
anti-
trips
in b
y A
OC
S Ba
12-
75,3
rd e
ditio
n, R
ev. 1
976.
N
AL:
AO
AC
197
5, 1
2the
d.
ttA
lmos
t sur
ely
S. s
pica
tus.
N
00
00
Tab
le 2.
Vita
min
s, tr
ypsi
n in
hibi
tor
Fo
b
Spec
ies
colle
ctio
n in
fr
esh
acid
ac
id
Rib
ofla
vin
inhi
bito
r p-
caro
tene
L
abor
ator
y &
R
ef. n
o.
(mo/
day/
yr)
Cod
e Pl
ant p
art*
sa
mpl
e sa
mpl
e (g
/uni
t pg
100g
-l)
(mgl
w-'
)(m
g 10
0g-')
TIU
(1O
Og-
I) (mg lO
O-')
andm
etho
ds$
Dat
e of
U
nits
go
f Fo
lic
Asc
orbi
c T
ryps
in
a 3 3
Aca
cia
tort
ilis
13A
03
/07/
76
TOB
S Fl
ower
s 2A
08
/02/
74 T
OD
U
Gre
en s
eeds
+coa
ts
38A
07
/20/
76 T
OD
U
Gre
en se
eds -
coat
s 39
A
07/2
0/76
TO
YU
G
reen
see
d co
ats
Aca
cia
xant
hoph
loea
5A
09
/28/
75
FTB
S Fl
ower
s 2
08
1
04/2
9/76
FT
DR
D
ry s
eeds
14
A
06/2
5/76
FT
GR
Bro
wn/
Bro
wn-
blac
k gu
m
15A
03
/07/
76
FTG
U
Cle
ar/li
ght a
mbe
r gum
15
AS
06/2
5/76
FT
GU
C
lear
/ligh
t am
ber g
um
Azi
ma
tetr
acan
tha
9A
02/0
9/75
A
TFR
R
ipe
frui
ts
9AS
06/2
9/76
A
TFR
R
ipe
frui
ts
34A
06
/12/
76
AT
FU
Unr
ipe
frui
ts
Chl
orop
hyrw
n sp
. nr b
aker
i 19
A
04/2
9/76
C
BL
U
Gre
en le
aves
C
omm
icar
pus p
lum
bagi
neus
30
A
05/2
9/76
SF
FR
Rip
e fr
uits
C
ynod
on da
ctyl
on
8A
11/2
9/75
CD
LU
G
reen
tille
rs
Ludw
igia
sto
loni
fera
26
A
05/1
5/76
LSW
U
Stem
s Ly
cium
Pur
opae
um '
1 OA
12/0
9/75
T
FBS
Flow
ers
29A
05
/29/
76
TFF
R
Rip
e fr
uits
1l
A
12/2
2/75
T
FLU
Y
oung
gre
en le
aves
200
236
1120
93
1
150
200
375
315 30
5054
325Y
1683
cm
1013
13
64
535
20.0
0 15
.60
39.8
5 38
.50
20.9
0 34
.65
39.3
0 56
.55
49.3
5
47.0
0 12
3.90
81
.00
23.8
0
30.0
0
19.8
0
24.6
5
12.3
0 72
.30
20.3
0
0.10
0.
066
0.03
6 0.04 I
0-14
0.24
0.
33
0.26
0.79
0,00
6
0.06
1
1.90
0.0 1
2 0.053
0.03
8
35.4
<
5-5
2.1
16.8
48.3
51
.7
<5-1
14
.5
t4.2
< 14
.7
9.2
5.6
23.3
33-2
<32.
1
19.5
85.0
16
.1
36.8
178.
3 49
.3
c 3.
9 <
4.8
I 10.
8 12
.1
13.4
<4
.7
4.2
49.0
42
.9
41.5
51.9
12.0
87.8
37.2
35.7
62
.1
< 7.
5
0.5
1.1
0.20
1.
7
1.3
0.90
0.
08
0.02
0.
02
0.14
0.
15
0.19
0.53
0.72
0.49
0.17
1.1
0.4 1
0.
50
WA
RF-
3 W
AR
F-1
3284
.4
WA
RF-
7 52
86.6
W
AR
F-7
WA
RF-
2 7.
35
WA
RF-
4 W
AR
F-5
WA
RF-
3 W
AR
F-5
WA
RF-
2 W
AR
F-5
WA
RF-
5
WA
RF-
4
WA
RF-
5
WA
RF-
2
WA
RF-
4
WA
RF-
2 1.
04t
WA
RF-
5 W
AR
F4
Salv
ador
a pe
rsie
a 4A
09
/28/
75
IA
08/0
2/74
Se
tari
a se
rtir
illat
a 27
A
05/1
6/76
So
lanu
m d
ubiu
m
33A
06
/05/
76
Spor
obol
us r
onsi
ndis
12
A
03/0
6/76
Sp
orob
olus
cor
dofa
nus
3A
08/0
7/74
Sp
oroh
olus
ken
trop
hyllu
s 16
A
04/0
8/76
Tr
iant
liem
a re
rato
sepa
la
7A
10/2
9/75
6A
10
/29/
75
Trib
ulus
terr
estr
is
24A
05
/07/
76
With
ania
som
nife
ra
35A
06
/25/
76
SPFR
SP
LU
SVLU
SIFU
SCT
U
SMC
R
SKC
U
TCB
U
TC
FU
TT
FU
Rip
e fr
uits
Y
oung
leav
es
Gre
en ti
llers
Unr
ipe
frui
ts
Late
ral m
eris
tem
s
Cor
ms
Cor
m co
res
Flor
al b
uds
Frui
ts
Soft
frui
ts
759
478
152
153
132
121
600
26 I
59 1
1445
48.9
5 0.
064
19.0
0 0.
040
31.8
5 0.
21
59.7
0 0.
39
53.4
0 0.
40
8.05
0.
067
8.65
0.
014
36.8
0 0.
14
20.2
0 0.
034
67.0
0 0-
046
8.5
< 4.
2
49.8
7.9
8.6
< 10
.6
< 25
.1
14.9
14
.1
44.2
5.1
127.
5
56.6
11.5
15.5
< 35
.3
< 25
.1
9.3
14.1
58.1
0.1
0.58
0.20
0.23
0.06
0.24
< 0.
25
0.24
0.
87
0.22
WA
R€-
2 W
AR
€-1
WA
RF
4
WA
RF-
5
WA
RF-
3
WA
R€-
I
WA
RF
4
WA
R€-
2 W
AR
F-2
WA
RF
4
9
WSF
R
Rip
efru
its
512
49.0
5 0.
10
8.9
61.8
0.
43
WA
R€-
5
*All
sam
ples
pres
erve
d in
oxa
lic ac
id e
xcep
t 20B
-1 (
no pr
eser
vativ
e).
tEqu
ival
ent t
o 17
45IU
100
g-I.
:S
ee n
ote
(5) in
Tab
le 1
.
s %.
2. 3
N
\o
0
Tab
le 3. E
lem
enta
l ana
lysi
s, by
em
issi
on sp
ectr
osco
py (W
AR
F L
abor
ator
ies)
YO Pa
rts
per m
illio
n
Spec
ies
Ref
. no.
C
ode
Plan
tpar
t P
K
Ca
Mg
Na
A1
Ba
Fe
Sr
B
Cu
Zn
Mn
Cr
Salv
ador
a per
sica
IB
SP
LU
Y
oung
gree
n le
aves
0.
12
0.84
1.
45
0.26
0-
03
59.6
12
.4
48.4
20
6 28
.9
2.5
4.8
26.2
4.
1 A
cari
a fo
rrili
s 2B
T
OD
U+
YU
G
reen
seed
s+co
ats
0.10
0.
37
0.15
0.
10
0.01
5.
3 3.
0 11
.8
18.4
6.
8 2.
4 5.
6 6.
2 1.
3 Sp
orob
olus
cord
ofan
us
3B
SMC
R
CO
~S
0.40
1.38
1.
27
0.69
0.
90
4290
55
.4
4050
19
2 37
.0
10.7
18
.1
216
14.2
cs a 3
rA P 3
Composition of Amboseli plants 29 1
seeds, namely 1.7%, is surely wrong, judging from the literature on leguminous seeds, including other Acacia species (Gwynne, 1969) and consequently so too is the reported very high carbohydrate level, which is calculated by difference. No other analysis of A . tortilis seeds has been found in the literature. We therefore used the mean (23.7% of dry weight) of the four values given by Gwynne (1969) for seeds of other acacias with indehiscent pods. This was converted to a fresh-matter 20.3% protein for seeds with a water content of 14.3%, and the value ofcarbohydrate was then adjusted accordingly. Even this value may be an underestimate, since protein in the pods with seeds of this species was 4.0% of fresh matter, equivalent to 14.6% of dry matter (17.8% of dry matter in Dougall, Drysdale & Glovers’ 1964 report on material collected in nearby Tsavo National Park) and in Gwynne’s 1969 report, the mean ratio of protein in seeds to protein in pods plus seeds for four analyses of indehiscent acacias was 2.0, which suggests 25.3% protein (fresh-weight basis) for the seeds of A . tortilis.
Taxonomic notes We have relied on the East African Herbarium (now the Kenya Herbarium) for identification of plants in Amboseli, and a small herbarium collection is main- tained in the Park. All plants used in this report were identified by the authors by comparison with these Herbarium-identified specimens.
The systematics of East African plants is still imperfectly known. The following taxonomic notes are based on our discussions and correspondence with the Herbarium staff, whose assistance and patience we are happy to acknowledge.
Some lilies recorded as Ornithogalium donaldsonii may have been 0. ecklonii, which also occurs in Amboseli. Several grass species have recently been separated from Cynodun dactylon (Clayton, Phillips & Renvoize, 1979). Material gathered around a waterhole in the Amboseli study area during 1980 and typical of material formerly identified as C. dactylon was identified at the Kenya Herbarium as C. tilemfurtisis. Dusysphuera prostrata is now a synonym of Volkensinia prostratu and Sulunum duhium of S . coaguluns. Plants that S.A. referred to as Sericocomopsis hilrlehrrrtidtii are probably Achyrunthes aspera L. var. pubescens. Psilolemma jurgeri (syn. Odyssea jaegeri) may have been confused with Odyssea puucinervis.
Sumplr descriptions For the material included in each sample, we have prepared detailed descriptions that will be useful to anyone attempting to apply, replicate or compare our results. However, these descriptions are too bulky to publish. Copies of the full text are available from the authors or from the Primate Center Library, Wisconsin Regional Primate Research Center, Madison WI 53715 (tel. 608-263-3512). In addition, a copy has been deposited in the library of the Institute for Primate Research, National Museums of Kenya, Nairobi.
Discussion This report gives the results of 864 biochemical analyses from ninety-one primate foods obtained from thirty-seven species of plants in Amboseli. Many of these foods are eaten by other animals, as well.
292 S . A . Altmann. D . G . Post and D . F. Klein
Several related studies have been carried out in Amboseli. Wrangham & Waterman (1981) analysed total phenols and tannins from many of the vervet monkeys’ plant foods. Post (1978) mapped a large portion of this area into hundreds of plant zones, on the basis of a classification of twenty-three zone types. Altmann (1976) and Western (unpubl.) have been studying demographic changes in the acacia trees of Amboseli. The exudates of these trees have been analysed by Hausfater & Bearce (1976) and the effects of herbivores on them have recently been studied by Michael Milgroom. Klein (1978) has studied phenological changes in the Amboseli flora and effects of these changes on vervet monkey breeding seasons. We are currently monitoring key phenological changes on a routine basis. Long-term changes in t h e h b o s e l i habitat have been described by Western (1973) and by Western & van Praet (1973).
As a result of these studies, the Amboseli region is now one of the world’s most intensively studied natural feeding areas for primates. Much work remains to be done, however. For a number of foods, proximate analyses have not yet been carried out and, to date, few analyses have been made of components within the proximate classes. So, for example, while complementarity between the amino acids of the grasses and legumes is suspected on the basis of related cultivars (Altmann, 1987), no analyses of amino acids have been carried out yet. (Such a study has recently been initiated by S. A. Altmann and Robert Heinrickson.) The only plant toxins that have been analysed to date are trypsin inhibitor, total phenols and tannins, and those only from a few Amboseli plants. Toxins of some of the local species, e.g. Solanum incanum (Watt & Breyer-Brandwijk, 1962) are known from research at other sites. Only a few analyses of micro-nutrients have been carried out (Tables 2 and 3) and at present we have only very limited data on phenological changes in nutrients and toxins.
Acknowledgments Our research was supported by the following grants: MH 19617 from the U.S. Public Health Service to S.A.; grants from the National Science Foundation, the Society of Sigma Xi and the Boise Fund to D.P.; grants from the National Science Foundation, the National Institutes of Health and the Society of Sigma Xi to D.K. Several people worked with us in the field to collect the plant samples: Rachel and Michael Altmann, Glenn Hausfater, the Hans Kummer family, Jane Scott, Peter Stacey, Roxana Jansma, Vicki and Rob Foley. Much advice on field preservation of samples was provided by David Aulick of WARF Laboratories. The staff of the East African Herbarium (now the Kenya Herbarium) identified our numerous plant specimens, a service that was indispensible for this project. We are grateful to the Republic of Kenya for permission to do research in Amboseli and to the two successive wardens of Amboseli, Joseph Kioko and Bob Oguya, for their support and co-operation.
References ALTMANN, S.A. (1976) The Acacia Woodland of Amboseli National Park: Current Status and Future
ALTMANN, S.A. (1987) Foraging for Survival: Yearling Baboons in Amboseli. University of Chicago Press. Developments. Report to Director, Kenya National Parks.
Chicago.
Composition of Amboseliplants 293
ALTMANN, S.A. & ALTMANN, J. (1970) Baboon Ecology: African Field Research. University of Chicago
CLAYTON, W.D., PHILLIPS, S.M. & RENVOIZE,S.A. (1974) Floraof Tropical East Africa, Gramineae (Parr2).
DOUGALL, H.W., DRYSDALE, V.M. & GLOVER, P.E. (1964) The chemical composition of Kenya browse and
GWYNNE, M.D. (1969) The nutritive values of Acacia pods in relation to Acacia seed distribution by
HAUSFATER, G. & BEARCE, W.H. (1976) Acacia tree exudates: their composition and use as a food source by
KLEIN, D.F. (1978) The Die! and Reproductive Cycle of a Popularion of Vervef Monkeys (Cercopithecus
POST, D.G. (1978) Feeding and Ranging Behavior of rhe Yellow Baboon (Papio cynocephalus). Ph.D. disser-
POST, D.G., HAUSFATER, G . & MCCUSKEY, S.A. (1980) Feeding behaviour of yellow baboons (Papio
WATT, J.M. & BREYER-BRANDWIJK, M.G. (1962) The Medicinal and Poisonous Plants of Southern and
WESTERN, D. ( 1 973) The changing face of Amboseli. Africana 5(3), 23-29. WESTERN, D. & VAN PRAET, C. (1973) Cyclical changes in the habitat and climate of an east African
ecosystem. Nalure (Lond.) 241, 104-106. WnANGHAM, R.W. & WATERMAN, P.G. (1981) Feeding behaviour of vervet monkeys on Acacia torlilis and
Acacia xanrhuphloea: with special reference to reproductive strategies and tannin production. J. Animal
Press, Chicago.
Crown Agents for Overseas Governments and Administrations. London.
pasture herbage. E. Afr. Wildl. J . 2,82-125.
ungulates. E. Afr. Wildl. J . 7, 176-178.
baboons. E. Afr. Will/. J . 14,241-243.
aethiops). Ph.D. dissertation, New York University.
tation, Yale University.
cynocephalus): relationship to age, gender and dominance rank. Folia Primarol. 34, I7Cl95.
Easrern A,frica. Livingstone, London.
EcoI. SO, 7 15-73 I .
(Manuscript received 9 September 1984)