G.J.B.A.H.S.,Vol.5(2):15-24
(April-June,2016)
ISSN: 2319 – 5584
Abundance and Species Composition of Indigenous and Exotic Fruit Trees in
the Feeding sites of Fruit Bats: A Study Case of Kasanka National Park and
Kafinda Game Management Area, Central Zambia
1
Chansa Chomba & 2Caludia Sianjobo
1
School of Agriculture and Natural Resources, Disaster Management Training Centre,
Mulungushi University, P. O. Box 80415, Kabwe, Zambia. Email: chansachomba@rocketmail.com
2
C/O Mr. Sianjobo, Lusaka, Zambia
Abstract
We conducted a basic inventory of tree species composition, density and frequency, to establish baseline data on
abundance and phenology of fruiting trees in the known fruit bats’ foraging areas. The study was carried out between
November and December 2014 and January to February 2015, in Kasanka National Park and Kafinda Game Management
Area. This area hosts one of the largest global concentrations of fruit bats estimated at ten million in one single locality
every October – December/January and was therefore, found to be suitable for this study.Plot method using 20 m x 20 m
square quadrats were set up inside the National Park and Kafinda Game Management Area. Each tree in the plot was
identified and DBH taken at 1.3 m above ground. Tree height was determined using a tree height measuring rod and
Haga Car Leiss altimeter. Identification of tree species was done by; i) direct observation, ii) use of a field guide, and iii)
use of an experienced herbarium technician. Results obtained showed that there were 64 species of trees. Of these, 20
(approximately 30 %) were fruit trees, of which 16 (approximately 80 %) were available and accessible to fruit bats.
Mangifera indica (an exotic species) and Uapaca spp were the most abundant fruit trees with relative abundance > 10 %.
The density of the trees was high at 365/ha of which 199/ha (55 %) was covered by fruit trees. The study area had
abundant flowering and fruiting trees (55% by area). Further research is required to; i) investigate other environmental
factors influencing the choice of Kasanka area as a feeding and roosting site for migratory fruit bats, and ii) explore
potential alternative sites for migrant fruit bats which share similar phenology and tree species composition as the
Kasanka area.
Keywords: Foraging, alternative, phenology, colony, flowering, accessible
1. Introduction
The miombo ecoregion is located within the Zambezian Regional Centre of Endemism and covers approximately
3.6 million square kilometres spanning ten countries, Angola, Botswana, Democratic Republic of Congo, Malawi,
Mozambique, Namibia, South Africa, Tanzania, Zambia and Zimbabwe (WWF 2001). The ecoregion is largely
Caesalpinoid woodland that is generally called broad-leaved ‘dystrophic’ savannah woodland (Huntley and Walker
1982). In Zambia, it is defined by the dominance or high frequency of trees belonging to the legume subfamily
Caesalpinoideae and genera Brachystegia, Julbernardia and Isoberlinia spp. Other genera include, Baikiaea,
Cryptosepalum, Colophospermum and Burkea.
The miombo is generally divided between wet and dry miombo and other vegetation communities associated with it
are: mopane, itigi thicket, wetland grassland, Acacia/Combretum complexes, Baikiaea, Burkea/Terminalia/Combretum
complexes, and Cryptosepalum which are largely non-fruit bearing. These vegetation communities usually support low
densities of micro and mega chiroptera, but the Kasanka – Kafinda areas located in wet miombo of central Zambia is an
exception to this rule and is the only known locality in Zambia which hosts one of the largest global concentrations of
migratory fruit bats in a single locality (Ritcher, 2004), comprising eight species viz; Dwarf epauleted fruit bat
(Micropteropus pusillus), Anchieta's broad-faced fruit bat (Plerotes anchietae), Peters's epauletted fruit bat
(Epomophorus crypturus), Wahlberg’s epauletted fruit bat (Epomophorus wahlbergi), Little epauletted fruit bat
(Epomophorus labiatus), East African little epauletted fruit bat (Epomophorus minor), Dobson's epauletted fruit bat
(Epomops dobsonii) and Straw-coloured fruit bat (Eidolon helvum) (sensu Ritcher and Cumming, 2006). Such large
congregations of fruit eating bats in a single locality are only expected in tropical rain forest where fruits are more
abundant (Ritcher, 2004).
It is not known at least by the current state of knowledge how forest dwelling species which mainly subsists on fruit
pulp and juices, nectar, and pollen as well as chewing leaves to obtain special nutrients, which are abundant in tropical
rainforests, would migrate to utilise a dystrophic miombo vegetation community which is generally poor in fruits. Other
researchers, including Keely (2009), suggest that migration are probably genetically determined, instinctive and
influenced also by weather conditions and the availability of food, but nothing is known of how bats recognize migration
goals or how succeeding generations learn their locations. If food is one of the important factors determining their
selection of sites by bats, then it becomes inevitable to investigate the abundance of fruit trees in the known feeding
areas.
In Kasanka National Park and Kafinda Game Management Area for instance, Ritcher (2004), recorded food sources
for bats which included, Musa spp, Magnistipula butayei, Parinari curatelifolia, Syzygium spp, Uapaca kirkiana,
Uapaca nitida, Uapaca benguelensis, Uapaca sansibarica and others which are in fruit during the period coinciding with
the arrival of migratory fruit bats. However, she did not determine the overall tree species composition, abundance and
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phenology of fruit trees as this may, acting together with other environmental factors, influence the number of bats that
can be supported by the habitat, determine their time of arrival, length of stay and departure time.
This study was intended to provide baseline information on the ecological status of the habitat in terms of tree
density and availability of fruit trees, which could be used in monitoring the habitat and in safeguarding such areas
against destructive anthropogenic activities. Preventing disturbances to the habitat is important because it may lead to
low reproductive success as was also recorded in birds by Robinson et al. (1995).
The role of the current impacts of climatic change scenarios combined with wild fire were also considered in this
study because they can affect woody plant communities including fruit trees. Thus, understanding the abundance of
fruiting trees can enable management to measure the degree of departure from baseline information and help in securing
such foraging areas by managing and controlling anthropogenic activities which may fragment or destroy habitat quality
and deprive bats of their food source.
This is important because worldwide, bat populations are declining at a rapid rate, due in large part to the
destruction of feeding and roosting habitats and the misuse of toxic pesticides (Wilson, 2010). Human interactions with
bats have also contributed to their decline. In many nations, bats are unjustifiably earmarked as nuisances or threats to
public health and killed. Between 1963 and 1970, the world's largest bat colony, some 30 million Mexican free-tailed
bats in Eagle Creek Cave in the Apache Sitgreaves National Forest in southeastern Arizona was reduced to 30,000
individuals, a decline of 99.9 percent. Major population losses have been recorded on all continents, and several islanddwelling Megachiroptera, such as the little Mariana fruit bat of Guam, have recently become extinct (Wilson, 2010), this
is no exception to Africa and Zambia in particular where habitat transformation is a major threat to the long-term survival
of bats.
Keely (2009) also showed that the decline in the populations of Old World fruit bats was attributed to excessive
hunting, persecution by farmers, and deforestation. Some species have been hunted to extinction for food and medicinal
potions (IUCN, 1990; Dumont 2000). Many other species are endangered or vulnerable (IUCN, 1990; Horvorka et al,
1998). Unfortunately, there is no legal protection available throughout their range in Africa. Therefore, maintenance of
good quality habitat by protecting fruit trees for food and thick woodland or forests for roosting remains critical in
sustaining bat populations. In Kasanka National Park, for instance, bat migration forms a basis for photographic tourism
and loss of habitat quality and decline in their population size would negatively affect tourism, which is an important
source of income for conservation.
In this study, we conducted a basic inventory to determine tree species composition, establish density and fruiting
season, determine the abundance of fruit trees among other woody plants as an indication of food availability in areas
where Ritcher (2004) conducted her studies and recorded the types of fruits eaten by bats. Information generated by this
study would be utilised by the Department of National Parks and Wildlife (NPW) to maintain a minimum density of fruit
trees as source of food for bats. Knowing species composition and density would also enable NPW to set lower
thresholds of vegetation change below which management intervention may be required to actively manipulate the
habitat and restore its physiognomic status. Understanding the nature of the habitat with respect to species composition,
density and the distribution of fruit trees is critical in ensuring that the human - bat interaction of hunting and habitat
clearance through logging as earlier recorded by Ritcher (2004) is monitored and negative human impacts such as
Chitemene system of agriculture can be prevented. In emphasising the importance of determining tree density and species
diversity, Keely (2009) showed that in some areas, large numbers of Old World fruit bats have been killed by farmers
fearing damage to orchard crops, when in actual fact bats may only feed on crops during droughts, when native trees
provide little food, otherwise most commercial fruit is harvested while it is still too green to attract Old World fruit bats.
2. Material and Methods
2.1 Location and Description of Area
2.1.1 Location
The study was carried out in Kasanka National Park and Kafinda Game Management Area. The National Park and
Game Management Area are located in Serenje district of Central Province of Zambia. It is an appropriate site for
studying the feeding ecology of the fruit bats, because it is the only site in Zambia where such large numbers of up to ten
million gather over the period October/November – December/January every year. The National Park is located at
coordinates (12º30’S 30º14’E) and altitude of 1200 m above sea level (Figure 1).
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Figure 1 Location of Kasanka National Park and the adjacent Kafinda Game Management Area, Zambia. (Source:
Zambia Tourist Board)
2.1.2 Climate and Vegetation
The study area is located in Agro-ecological zone III with annual rainfall ranging from 1,100 mm to 1, 700 mm
(Cosson et al. 1999). The rainy season is between October/November - April. The vegetation communities are composed
of mainly miombo woodlands dominated by trees of the genera Brachystegia, Isoberlinia, and Julbernardia spp
interspersed with trees of the genera such as; Uapaca, Protea, and Faurea intermixed with wide, grassy dambos and
small stands of chipya forest (Chidumayo, 1987).
The common vegetation communities are: Chipya forests comprising wooded grassland dominated by fire resistant
tree species such as Terminalia mollis, Erythrophleum africanum, and Combretum spp. These forests were found in small
patches in mosaic with miombo woodland throughout Kasanka. Chipya is notable due to absence of Uapaca and other
common miombo fruiting species. “Mateshi” forest a dry evergreen forest exits in Kasanka only as small relics (Smith &
Fisher, 2001).
The Mushitu forest located inside the National Park is important because it is the only known roost site in the
Southern Africa region for a large migratory fruit bat colony (Ritcher, 2004). The “Mushitu” is a three-canopy closed
evergreen forest of up to 27m tall and is represented by only two small patches covering an area of about 0.4 km² (Smith
& Fisher, 2001). It is dominated by species such as Khaya nyasica, Parkia filicoidea, and Diospyros mespiliformis
(Smith & Fisher, 2001). Below the upper canopy is a dense thicket of plants consisting mainly of ferns and climbers
(Storrs, 1995). Waterberry (Syzygium cordatum) and African mahogany (Khaya nyasica) dominate the taller canopy
layer, while Swamp fig (Ficus trichopoda) dominates the understory.
2.2 Field Data Collection Methods
2.2.1 Random Sampling
Quantitative data were collected by randomly selecting sampling plots in woodlands located in Kasanka National
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Park and Kafinda GMA. Sample plots in Kafinda GMA, were taken starting from the Serenje/Samfya road (T2). Random
numbers were generated for determining the 35 starting points of each transect from Serenje/Samfya road as baseline.
The orientation of transects was 90° from T2 Road. The choice of which side of the road transects should be set was
decided by tossing a coin, where the tail side corresponded to the left side of the road and head to the right side. A Total
of Seventy-five (20 inside the park and 55 outside the park) sample plots measuring 20 m x 20 m were sampled. Plant
species in each plot were counted, identified and measured. Sample plots inside the National Park were taken at random
surrounding the roosting site within a radius of 5 km (Figure 2).
Figure 2 Location of sample plots in recorded bat feeding areas
Notes: Numbers represent sample location of plots, and blue open water, dark green riparian forest and light green open
woodland and grassland respectively.
2.2.2 Determining Abundance of Fruiting Trees
To determine the abundance of fruiting trees, quantitative and qualitative approaches were used to collect primary
data. Qualitative information was collected through nonstructured interviews with park management staff and local
community members. This provided the researchers with local knowledge on various aspects of human-bat interactions.
Quantitative data were collected by physically taking measurements in the field. Coordinates were taken for each
plot using GPS. Plots were then established using a 50 meter measuring tape and setting the corners of each plot at 90o
using the 3, 4, 5, method. Species name and Diameter at Breast Height (DBH) were taken for all species inside the plot.
Trees ≥ 10 cm DBH stem size were counted in each plot. The DBH was taken at 1.3 metres above ground by passing the
tape around the tree trunk and noting readings in cm on the tape. The DBH was measured and converted using the
equation C = Dπ. Thus D = C/π, (Chomba et al. 2013). Fruit and non-fruit trees were noted and recorded as such. The
fruiting season for each fruiting tree was obtained from literature, particularly Storrs (1995) to see whether the fruiting
season coincided with the arrival of fruit bats.
Where tree stems branched below 1.3 m above ground, individual stems were counted and considered as separate
trees because measurements used to calculate basal area were taken at 1.3 metres above ground level (Chomba et al.
2013). The tools and materials used included; i) GPS for distances, direction and geographic locations, ii) Tape for stem
diameter measurements and marking the dimensions of the plot, and iii) Forms for entering data.
3. Results and Discussion
3.1 Results
3.1.1 Tree Species Composition
The study area had 64 species of which 20 (approximately 30%) were fruit trees (Table 1 and 2).This is a high
species composition which is not typical of miombo woodlands and is attributed to habitat mosaic; Chipya forests
“Mateshi” forest a dry evergreen forest, and Mushitu forest which provided a variety of habitat conditions. The riverine
vegetation for instance was dominated by mainly species of the genera Syzygium and Ficus in swampy areas which also
contributed to the high diversity of species.
Table 1 Trees species recorded in the study area, Kasanka National Park and Kafinda Game Management Area,
Zambia
No. Scientific Name
English Name Local Name (Bemba)
Fruit tree
1
Acacia heteracantha
Kafifi
2
Albizia antunesiana
Musase, Mukoso
3
Anisophyllea boemii
Mufungo
●
4
Anona senegalensis
5
Bobgunia madascariensis (Syn.
Ndale
Swartzia madagascariensis)
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6
7
8
9
Brachystegia boehmii
Brachystegia spiciformis
Burkea africana
Cassia abbreviata
10
11
12
13
14
15
16
17
18
19
20
21
22
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
Capassa violecea (Lonchocarpus
capassa)
Combretum molle
Citrus sinensis
Dalbergia nitidula
Diospyros mespiliformis
Diospyros batocana
Diplorynchus condylocarpon
Erythrina abyssinica
Erythrophleum africanum
Faurea intermedia
Ficus spp
Hymenocardia acida
Isoberlinia angolensis
Julbernardia paniculata
Lannea spp.
Lantana camara
Magnistipula butayei
Mangifera indica
Maprounea africana
Maquesia macroura
Montes africanus
Morus nigra
Musa spp
Ochna schweinfurthiana
Oldifieldia dactylophylla
Ozoroa reticulata
Parinari curatellifolia
Peltophorum africanum
Pericopsis angolensis
Persia americana
Phyllocosmus lemaireanus
42
43
44
45
46
47
48
49
50
51
52
53
Piliostigma thonningii
Pinus kesiya
Pericopsis angolensis
Phyllocosmos lemaireanus
Protea spp
Pseudolachnostylis maprouneifolia
Psidium guajava
Pterocarpus angolensis
Rothmania englerana
Schrebera trichoclada
Strychnos cocculoides
Strychnos spinosa
54
55
Syzygium guineense guineense
Syzygium cordatum
(April-June,2016)
Long pod
cassia
Lilac tree
Ngasa, Musamba, Muombo
Muputu
Kapanga, Mukoso
Munsokansoka, Musambamfwa
Bush willow
African ebony
Lucky bean
African beech
Mango
-
Montamfumu, Mulama, Kaunda
ISSN: 2319 – 5584
Chiya
Bush orange
Elephant
orange,
Monkey ball
Water berry,
Water tree
-
Kalongwe
Muchenja
Muntafita
Mwenge
Mulunguti
Kayimbi, Mukoso
Saninga
●
●
●
Akapempe
Umutobo
Umutondo
Kaumbu, Kabumbu
Mubwilili, Mukwebula
Mango, Umwembe
Kafulamume
Museshi
Chimpampa
●
●●
Inkonde
Kabanga
Kampangwila, Lunda, Muonga
Kulukumo
Umupundu
Mwikalankanga
Mubanga
Kotapela
Mwembembe, Musengameno,
Kampombwe
Mufumbe, Nachimfumbe
●●
Mubanga
Mwembembe, Musengameno
Musoso
Musangati
Mupela
Mulombwa
Mupulupumpi, Mwinebala
Kapande
Kasongole
Sansa
Musafwa
Mufinsa
●
●●
●●
●
●
●
●
Mufinsa
●
Mubobo, Chimpakwa
Namwinshi
58
Uapaca nitida
Musokolobe
●
59
Uapaca sansibarica
Musokolobe
●
60
Uapaca banguelensis
Makonko
●
61
Uapaca kirkiana
Wild loquat
Musuku
●
62
Vitex doniana
Mufutu
●
63
Zahna africana
Chibangalume
64
Ziziphus abyssinica
Kangwa, Kalanangwa
Notes: Exotic fruit trees were only found in the Game Management Area and not inside the National Park. Single dot (●)
denotes indigenous fruit tree and double dots (●●) represents exotic fruit trees.
56
57
Syzygium g. huillense
Terminalia mollis
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We identified 16 (80%) of the fruit trees as being available and accessible when bats arrived. Four (4) of the 20
were either out of season or had hard shells (e.g. Strychnos spp.) that bats were unable to break open and access its juicy
parts (Figure 3 a & b; Table 2).
(a)
(b)
Figure 3 Strychnos spp, a) fruits on a tree, and b) hard shell of Strychnos fruit containing juicy parts, but too hard to be
broken by bats. (Source: http:www.crfg.org).
Table 2 Indigenous and exotic fruit trees in Kasanka and Kafinda Game Management Area, as food supporting millions
of bats during the period November - January/February, Zambia
No.
Species
Season of fruiting
Remarks
1
Anisophyllea boemii
September- November/December
2
Citrus sinensis
Any time of year
3
Diospyros mesipilliformis
April - September
◙
4
Ficus spp
Any time of year
5
Magnistipula butayei
October - January
6
Mangifera indica
Rainy season
7
Musa spp
Any time of year
8
Parinari curatellifolia
September- November/December
9
Persia americana
Any time of year
10
Psidium guajava
Any time of year
11
Strychnos cocculoides
September- November/December
♦♦
12
Strychnos spinosa
September- November/December
♦♦
13
Syzygium cordatum
November - March
14
Syzygium guineense guineense October - January
15
Syzygium g. hullense
December/January-March
16
Uapaca nitida
September – November/December
17
Uapaca sansibarica
September – December
18
Uapaca kirkiana
August-November/December
19
Uapaca benguelensis
August-November/December
20
Vitex doniana
April - August
◙
Notes:
♦♦ Fruit is hard and bats cannot break to access its juicy parts; within bat season; and
◙ out of bat season.
3.1.2 Abundance and Density of Fruit Trees
Of the 16 fruiting trees available to bats, Uapaca spp, Mangifera indica, were the most abundant with relative
abundance being > 10% (Figure 4 a, b & c).Total tree density was high at 365/ha of which more than half 199
(approximately 55 %) /ha were fruit trees.
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(a)
(b)
(c)
Figure 4 a) Relative abundance of fruit trees, b) density of fruiting trees, c) frequency, Kasanka National Park and
Kafinda Game Management Area, Zambia. (Notes: Musa spp, Psidium guajava and Mangifera indica are exotic plants
planted by people living in GMA for food)
3.2 Discussion
Our findings show that the current state of fruit tree density in the bats foraging area was high. This is probably
because the two areas are protected as National Park (IUCN Category II) and Game Management Area (IUCN Categeory
VI). In National Parks, human settlements are not allowed save for isolated illegal activities and wild fire. In the Game
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Management Area, human settlements and activities are strictly regulated in compliance with a subsisting General
Management Plan. Such efforts may have contributed to the high abundance of fruit trees recorded in this study.
Studies carried out in Zimbabwe by Fenton et al. (1998) stressed the importance of knowing species composition
and density in bats’ habitat because the degradation of habitats for bats is a pervasive problem in conservation. Fenton et
al. (1998) however noted that the same picture did not apply to birds, that not all organisms are equally affected by
reduction in density, species composition and tree canopy in woodlands. Species of micro-chiroptera which are largely
insect eating bats persisted even in sites devoid of fruit trees and where tree canopy has been virtually eliminated (Brosset
et al., 1996).
In our study which focused on fruit bats, we emphasize that it is eminent that the conservation one species, would
be affected by other species. Increase in elephant populations for instance, which are prime destructive force of
woodlands (Racey & Entwistle, 2000), may reduce tree density which raises the question of whether Kasanka National
Park management team should focus on species management or the habitat.
3.2.1 Abundance of Fruiting Trees and Fruiting Season
The abundance of fruiting trees would be attributed to the ecological location of the area in Ecological Zone III of
the country which receives the highest amount of rainfall of between 1,200 - 1,700 mm /y-1. Such high rainfall
characterises the abundance of wet dambos and perennial rivers and streams which are suitable habitats for fruit bearing
trees of the genera Syzygium (Figure 5 a & b ) which provide juicy fruits as acknowledged by Cumming & Bernard
(1997). Such habitat mosaic contributes to the high diversity and abundance of fruit trees a factor not earlier investigated
by Ritcher (2004).
(a)
(b)
Figure 5 a) Riverine vegetation and b dambos which are characteristic of wet habitats suitable for Syzygium spp a genus
of fruit trees utilized by bats. Kasanka National Park, Zambia.
Regarding the fruiting season, Ritcher (2004) showed that arrival time of migrant fruit bats was October/November
and departure was around December/January. This period coincides with the rainy season and fruiting season (Storrs,
1995) during which time 18 (approximately 80%) of the recorded 20 fruit trees eaten by bats as recorded by Ritcher
(2004) were fruiting.
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Arriving during the fruiting season (October – January) which coincided with the rainy season ecologically reduced
search time for food. Trees ripening in the rainy season had adequate moisture and their fruits were juicier which may as
noted by Ritcher (2004) eliminated the need for bats to search for water to drink which ultimately conserved energy that
bats needed to fly in search of water.
4. Conclusion and Recommendations
4.1 Conclusion
In this study it has been shown that the current density of trees in areas used by bats is 365/ha and the density of
fruit trees is 199/ha. It would appear from the current status that these are baseline densities needed to maintain feeding
areas in a condition that would support current numbers of migrant fruit bats.
In the mean time, efforts should be made to ensure that such tree densities (365/ha for all tree species combined and
199/ha for fruit trees) are not reduced because that would have a negative impact on food availability when the
physiognomy of the habitat has been transformed as earlier shown by Robinson et al. (1995). Intuitively, these results
only provided baseline information on habitat quality and do not in any way suggest reasons why bats migrate to
Kasanka area. They also do not suggest that this locality has the highest concentration of fruit trees among all the wet
miombo vegetation communities in Zambia, but rather convenient to state that an abundance of fruit trees is inevitable in
fruit bat feeding areas. Further research is required to examine other environmental factors that may influence the choice
of Kasanka National Park and Kafinda GMA as destination area for migrant fruit bats.
4.2 Recommendations
In Kafinda GMA like other similar areas in Zambia, human habitation and other anthropogenic activities related to
primary extraction of resources are practiced. In Central Zambia where the study area is located soils are dystrophic,
acidic, highly leached and poor in macro and micro nutrients. Compounded by high poverty levels, perennial food
shortages, exacerbated by low levels of education, communities are left with no option but to depend on the extraction of
forest based resources for their survival. To cope with poor soils in the absence of artificial fertilizers which is beyond
their reach, communities practice a traditional form of shifting cultivation called the chitemene system. In this practice,
trees are chopped down, lopped or pollarded in the field and the cut branches and stems are collected and piled together
at a central location then burned between September – early November when the moisture content is low and
temperatures are high as to support combustion. The ashes lower the acidity and introduce phosphorous, which increases
soil fertility.
Trees are unselectively cut down for fuel wood for cooking, and some of it is taken for sale at Tuta Bridge for
drying fish. Trees are also cut down during the harvesting of caterpillars contributing further to decrease in tree density
and the number of fruiting trees.
In view of the current increase in human population, it is likely that the demand for wood fuel would increase.
Based on this baseline study the Department of National Parks and Wildlife, should consider formulating and
implementing comprehensive General Management Plans for the National Park and GMA in order to safeguard the
foraging areas against destructive human activities.
Acknowledgements
We extend our sincere thanks to the Acting Director General of Zambia Wildlife Authority (ZAWA) for providing a
letter of authority to enter and collect data in a National Park. We also thank management of Kasanka National Park for
the cooperation and support during the field data collection trips.
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