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DOI 10.1007/s10841-007-9089-2
ORIGINAL PAPER
Received: 22 February 2007 / Accepted: 3 May 2007 / Published online: 30 May 2007
Springer Science+Business Media B.V. 2007
Abstract African elephants and other indigenous megaherbivores have a major impact on local vegetation structure, including aquatic communities, as their big feet and
large mass pound the fringes of water bodies. This disturbance is likely to have a profound influence on the structure
and composition of insect assemblages in these habitats.
We investigated which dragonfly (Odonata) species were
tolerant of trampling by elephants and other game.
Assemblage composition differed according to extremely
high, very high or high disturbance levels. Dragonfly
abundance was greatest where impact was high, and
decreasing when disturbance became very high or extremely high. Several odonate species are well-adapted to
fairly high levels of disturbance, although too much is
impoverishing. Medium and low impact sites were geographically separated, and this, combined with much lower
disturbance levels, had a considerable influence on promoting regional dragonfly diversity. Several regional specialist species only occurred in the geographically
separated, low-impact sites. The full complement of dragonflies is present only when there is a combination of
various disturbance levels combined with spatial variation.
Elephant impact is similar to that of humans, with too
much of either or both, leading to a species-poor, habitatgeneralist dragonfly assemblage.
Keywords Elephant trampling Dragonfly biodiversity
Parallels with human impact
Introduction
Anthropogenic habitat loss is the primary reason for current loss of biodiversity. This loss comes about from
fragmentation of natural ecosystems and the reduction of
remnant patch quality (Fahrig 2003). Prior to the massive
human impact of the last 6,000 years, the European landscape was disturbed by vertebrates (Vera 2000). Similarly,
on the African continent, there always has been, and in
places still is, a major impact from megaherbivores
(Samways and Kreuzinger 2001; Gibson 2004). These
animals include elephant (Loxodonta africana), which can
influence plant assemblage composition and pattern
(Dublin et al. 1990; Van de Vijver et al. 1999; Lombard
et al. 2001), and dung beetle assemblages (Botes et al.
2006).
Dragonflies are considered as valuable indicators of
human disturbance of African riverine systems (Samways
and Steytler 1996). Yet some of these systems are naturally
disturbed by elephants and other large megaherbivores,
characteristic of the savanna. This has probably been going
on for many millennia, long before human agricultural
influence. It can be seen today where human disturbance is
still relatively minimal, as in parts of southern Africa.
Dragonflies and other physically small components of these
ecosystems have adapted to these impacts, which include
substantial trampling and harvesting of plant material.
Dragonflies are particularly prone, being aquatic insects in
a dry landscape, living precisely where the large animals
congregate to drink. This presents an interesting ecological
situation, because it is in these highly disturbed, focal
aquatic areas that some dragonfly species are remarkably
common. Yet not all local areas are affected by elephants
in this way, with these animals preferring some specific
watering locations over others. The local megaherbivore
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pressure puts enormous demands on the dragonfly assemblage and would seem to be a major natural factor affecting
dragonfly assemblage composition and pattern.
The aim here was to determine the extent to which
megaherbivores, especially elephants, influence the relative
abundance and species richness of dragonfly species.
Additionally, the objective was to establish which dragonfly species are tolerant of such extreme natural disturbance and which are not. A final aim was to determine
whether the various elephant population levels are positive
or negative drivers of dragonfly diversity.
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Analysis
Ordination was performed on abundance data using
Canonical Correspondence Analysis (CCA), with interspecies distances, biplot scaling and no transformation,
with the program CANOCO (ter Braak and Smilauer
2003). CCA is robust and combines positive aspects of
indirect gradient ordination with aspects of regression
(Leps and Smilauer 2003). It allows for all axes, independent of each other, to select linear combinations of
environmental variables, which maximize the dispersion of
species scores. CCA is therefore a constrained correspondence analysis in the sense that site scores are restricted to
be a linear combination of measured environmental variables. It is also completely unhampered by high correlations among species or environmental variables. Testing
the significance of the relation with environmental variables was carried out by the Monte Carlo permutation test
included in CANOCO.
To add weight to the graphical summary of data provided by CCA ordination techniques, univariate statistics
were utilized to determine the exact significance of correlations between species and environmental variables.
495
Bank condition
Water condition
Aquatic vegetation
Extremely
high
Dumatau 1
Bare soil
Very high
Dumatau 2
Bare soil
High
Dumatau 3
Medium
Chitabe
Low
Vumbura
Results
A total of 31 Odonata species were recorded (Table 2). The
graphical summary of data in the form of CCA (Fig. 2)
shows distinct separation of species composition at extremely high, very high, and high impact levels from low
and medium impact levels. A summary of eigenvalues and
of the Monte Carlo permutation tests for both CCA ordinations are given in Table 3. The Monte Carlo tests resulted in axes, which are significant in both ordinations,
therefore the null hypothesis that environmental variables
had no effect on species representation is rejected. The
effects of environmental variables are highly significant
(P < 0.002 with 499 permutations).
The number of species and individuals decreased from
high impact levels to extremely high impact levels (Figs. 3,
4). While the number of individuals dropped at medium
and to a much lesser extent at low impact levels, the
greatest number of species was at low impact levels. While
no species had significant, positive correlations with extremely high impact levels, 13 species had significant
negative correlations Urothemis edwardsii, Rhyothemis
semihyalina, Trithemis hecate, Diplacodes lefebvrii, Diplacodes sp.1, Acisoma panorpoides, Hemistigma albipunctum, Orthetrum icteromelas, Agriocnemis victoria,
Ceriagrion glabrum (P < 0.01) and Anax imperator,
Crocothemis erythraea, Agriocnemis exilis at (P < 0.05).
Furthermore, A. victoria, O. icteromelas, A. panorpoides,
T. hecate, (P < 0.01) and R. semihyalina (P < 0.05) continued to have significant negative correlations with very
high impact levels. Conversely, C. glabrum, Ischnura
senegalensis, A. exilis, Orthetrum trinacria, H. albipunctum, Diplacodes sp.1 (P < 0.01) and Brachythemis leucosticta (P < 0.05) had significant positive correlations
with very high impact levels. With regards to high impact
levels only two species were significantly negatively correlated (A. victoria, and O. icteromelas (P < 0.01))
whereas sixteen species were significantly and positively
correlated (C. glabrum, I. senegalensis, A. imperator,
Phyllomacromia contumax, O. trinacria, Diplacodes sp.1,
D. lefebvrii, B. leucosticta, Trithemis annulata (P < 0.01)
and Pseudagrion massaicum, Ictinogomphus ferox, Orthetrum chrysostigma, H. albipunctum, A. panorpoides, C.
erythraea, U. edwardsii (P < 0.05)).
With medium impact levels, I. senegalensis, A. exilis, H.
albipunctum, Diplacodes sp.1, C. erythraea, U. edwardsii
(P < 0.01) and C. glabrum, U. edwardsii (P < 0.05) were
significantly negatively correlated, while Pseudagrion hamoni, A. victoria, A. imperator, O. icteromelas, R. semihyalina (P < 0.01) and Chalcostephia flavifrons (P < 0.05)
were significantly positively correlated. At low impact
levels five species I. senegalensis, A. imperator, O. trinacria, B. leucosticta (P < 0.01) and C. glabrum
(P < 0.05) were significantly negatively correlated. Conversely 14 species Ceriagrion katamborae, Pseudagrion
deningi, A. victoria, O. icteromelas, A. panorpoides,
Diplacodes sp.2, C. erythraea, T. hecate, Trithemis monardi, Trithemis stictica, R. semihyalina, U. edwardsii
(P < 0.01) in addition to Rhyothemis fenestrina and
Aethriamanta rezia (P < 0.01) were significantly and positively correlated to low impact levels. Pantala flavescens
was the only species not positively correlated for any of the
five impact levels.
Discussion
While it is widely recognized that megaherbivores, such as
elephant, can drastically alter landscapes (Buechner and
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496
Species
EH
VH
5
6
7
10
*
*
11
12
13
14
15
16
*
*
17
18
Diplacodes sp.1a
19
Diplacodes sp.2a
20
21
22
23
24
*
*
*
*
*
*
*
*
25
26
27
28
29
30
31
*
*
*
*
* Species present; EH Extremely high; VH very high; H high; M medium; and L low elephant impact. a Since this study was undertaken,
Dijkstra (2006) has done a taxonomic revision of this difficult genus; both these morphospecies are likely to be the actual species Diplacodes
pumila Dijkstra, 2006
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497
1.0
1200
Medium
4
15
1000
800
600
400
200
0
Extremely
High
19
20
High
-0.2
Very High
11
23 14
1
12
10 5
16
22
6 7
17
18
21
29
30
Extremely High
High
Medium
Low
Impact level
13
27
Very High
25
26
24 2
28
31
3 Low
-0.4
0.6
Table 3 Summary of eigenvalues and Monte Carlo test for CCA ordination
Axes
Eigenvalues
Speciesenvironment correlations
Cumulative % variance of species data
Cumulative % variance of environmental data
Monte Carlo test of significance
2
0.721
0.997
52.4
74.5
F-ratio
3
0.140
0.944
0.066
0.846
Total inertia
0.041
1.376
0.816
62.6
67.4
70.4
89.0
95.8
100.0
P-value
49.626
0.0140
26.791
0.0020
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498
No. of speci es
25
20
15
10
5
0
Extremely
High
Very High
High
Medium
Low
Impact level
maximum dragonfly diversity, a combination of intermediate (medium to high) and low disturbance levels appear
essential, which supports the general tenet that habitat
heterogeneity is a general principle in insect and other
biodiversity conservation. Finally, the results here suggest
that very high impact from elephants and humans are very
similar by breaking down vegetation to the detriment of the
dragonfly assemblage. Too many elephants and too many
people are likely to lead to the same outcome: a speciespoor dragonfly assemblage made up of only habitat generalists.
Acknowledgements We thank Russel Friedman of Wilderness
Safaris for making this study possible, and Marlon Samways for
practical support.
References
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Buechner HK, Dawkins HC (1961) Vegetation change induced by
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