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J Insect Conserv (2008) 12:493498

DOI 10.1007/s10841-007-9089-2


Elephant impact on dragonflies

Michael J. Samways Paul B. C. Grant

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

M. J. Samways (&)  P. B. C. Grant

Department of Conservation Ecology and Entomology, Centre
for Agricultural Biodiversity, University of Stellenbosch, Post
Bag X1, Matieland 7602, South Africa
e-mail: Samways@sun.ac.za

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.
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



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.

Sites, materials and methods

The primary research area was the Linyanti River, Botswana, between Kings Pool (1826 S, 2342 E; 954 m
a.s.l.) and Duma Tau (1831 S, 2334 E; 940 m a.s.l.). It
is a slow-moving, meandering, perennial river with fringing reeds, papyrus and grass, and with many marginal
pools and some lagoons. The fringing muddy bottom is
heavily affected by elephants, which trample pools to such
an extent that the river margins become heavily puddled
and in places devoid of vegetation (Fig. 1). Other megaherbivores in the area have a much more localized and
lesser affect, and these include hippopotamus (Hippopotamus amphibius), Cape buffalo (Syncerus caffer), zebra
(Equus burchelli), and red lechwe (Kobus leche). In addition, a large chacma baboon (Papio ursinus) population
removed water lilies on a regular basis. The extent of impact on the grass banks is also great, to the extent that the
grass is grazed down to lawn length, to mostly only a few
centimetres tall. Fringing the bank, mostly about 3040 m
from the rivers edge is a gallery forest of bushes and trees
of various sizes (Fig. 1). The various levels of megaherbivore

Fig. 1 A site at Duma Tau, Botswana, highly impacted by elephants

and other large vertebrates


J Insect Conserv (2008) 12:493498

impact are listed in Table 1, which are also the dependent

variables used in the analyses.
A further site, Chitabe (1931 S, 2322 E; 940 m
a.s.l.), received medium impact from elephants, and had
more extensive water bodies and localized pools (pans),
leaving the flowing system lightly grazed and grass
reaching an average height of about 50 cm (Table 1).
As there was no low-impact site on the Linyanti River
system, the nearest possibility was to include a site on the
Okavango River system, but it is recognized here that this
also implies wide geographical separation, and thus not
providing totally comparable data with the other sites. This
site was at Vumbura (1859 S, 2250 ; 940 m a.s.l.),
where the slow-moving Okavango system is mostly composed of an abundance of reeds, papyrus and tall (1 m)
grass. The large mammals are less abundant and make use
of regular paths, leaving most of the vegetation intact, and
having little stirring effect on the muddy bottom (Table 1).
During April 2002, at each of the five sites (representing
extremely high, very high, high, medium and low impact)
ten transects 100 m long were selected. These transects (50
in total) were each along one side of the waters edge, 3 m
wide, and included moving (<10 cm s1) and still water.
The end of one transect was about 50 m from the beginning
of the next. On warm, sunny days, between 09h00 and
15h00, the transects were slowly walked, and all adult male
Odonata individuals were recorded with close-focus binoculars.

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.

J Insect Conserv (2008) 12:493498


Table 1 Vegetation and water conditions at the five sites

Impact level Site

Bank condition

Water condition

Aquatic vegetation


Dumatau 1

Bare soil

Mud and highly turbid water

Pounded reeds, grass and lilies

Very high

Dumatau 2

Bare soil

Mud and variably turbid water

Pounded reeds grass and lilies with patches

of standing reeds and grass


Dumatau 3

Bare soil and some grass

Variably turbid and clear water



Long grass with bare soil


Mostly clear with turbid patches

Mostly standing reeds and grass with

abundant lilies
Long grass, reeds, sedges and lilies broken
by many channels



Long grass, sedges and reeds

Clear water channels

Based on tests for normality using probability plots in the

program STATISTICA (Statsoft Inc. 2003), data were
shown to be nonparametric, therefore Spearmans rank
correlation coefficient was utilized in STATISTICA for
nonparametric modelling.

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

Extensive stands of reeds and sedges, with

extensive lilies and macrophytes

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.

While it is widely recognized that megaherbivores, such as
elephant, can drastically alter landscapes (Buechner and



J Insect Conserv (2008) 12:493498

Table 2 Species recorded throughout the whole study





Ceriagrion glabrum (Burmeister, 1839)

C. katamborae Pinhey, 1961

Pseudagrion deningi Pinhey, 1961

P. hamoni Fraser, 1955

P. massaicum Sjostedt, 1909


Ischnura senegalensis (Rambur, 1842)

Agriocnemis exilis Selys, 1872

A. victoria Fraser, 1928

Anax imperator Leach, 1815




Ictinogomphus ferox (Rambur, 1842)

Phyllomacromia contumax Selys, 1879


Orthetrum chrysostigma (Burmeister, 1839)


O. icteromelas Ris, 1910

O. trinacria (Selys, 1841)


Chalcostephia flavifrons Kirby, 1889

Hemistigma albipunctum (Rambur, 1842)



Acisoma panorpoides Rambur, 1842


Diplacodes sp.1a


Diplacodes sp.2a


D. lefebvrii (Rambur, 1842)


Crocothemis erythraea (Brulle, 1832)


Brachythemis leucosticta (Burmeister, 1839)


Trithemis annulata (Beauvois, 1807)


T. hecate Ris, 1912





T. monardi Ris, 1931


T. stictica (Burmeister, 1839)


Rhyothemis semihyalina (Desjardins, 1832)


R. fenestrina (Rambur, 1842)


Pantala flavescens (Fabricius, 1798)

Urothemis edwardsii (Selys, 1849)


Aethriamanta rezia Kirby, 1889



* 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

Dawkins 1961; Laws 1970; Dublin et al. 1990; Ben-Shahar

1993; Cumming et al. 1997; Van de Vijver et al. 1999;
Western and Maitumo 2004), their effect on insect biodiversity is little known (but see Rivers-Moore and Samways
1996; Samways and Kreuzinger 2001, Botes et al. 2006).
Some dragonfly species here were very tolerant of the
influences of megaherbivores. Intense elephant activity
opens up habitat to more tolerant or adapted species yet
destroying habitat for more sensitive species. Generally,
these areas of intense impact also have low dragonfly
diversity. Even species which occur in extremely high
impact areas occur in greater numbers where level of impact is less. One exception is Brachythemis leucosticta,
which as an adult is also intimately related to game, as it


follows the wandering animals and feeds on the flushed up

The medium impact site had lower species richness
than the greater impact areas, but the results are not
directly comparable because of there being two variables: site difference and impact level. Together, these
two variables resulted in a change in species turnover,
but with more species being lost than gained. This is
illustrated in Fig. 2, where the medium disturbance site
had a different assemblage composition compared to the
other sites. These results point to the value of high
spatial habitat heterogeneity for maintaining regional
diversity, and which comes about at least partly through
level of disturbance.

J Insect Conserv (2008) 12:493498


No. of indivi dual s









Very High

23 14
10 5
6 7





Extremely High




Impact level

Fig. 3 Total number of individuals recorded at each site



Very High

24 2
3 Low



Fig. 2 Canonical Correspondence Analysis (CCA). Species are

represented by (D). Nominal variables are represented by (().
Environmental variables are given in Table 1. For species names
see Table 2

Certain species were significantly positively correlated

with very high impact, and indeed can be very abundant.
Furthermore, they are indicative of major natural megaherbivore disturbance. In contrast, other species that were
significantly positively correlated for low impact areas are
good indicators of undisturbed or minimally disturbed
natural conditions.
Only seven of the 31 species (Table 2) tolerated all
levels of impact, making them the ultimate habitat generalists. All seven are also widespread and common across
southern Africa and even elsewhere. One of them, Crocothemis erythraea, even benefits from human disturbance

such as removal of indigenous trees (Samways and Steytler

Very few species actually preferred the intensely disturbed habitats, although Ischnura senegalensis, like B.
leucosticta, appeared to thrive in such conditions. Ischnura
senegalensis also inhabits artificial pools and reservoirs
that are unsuitable for most other species, again illustrating
the similarity between elephant and human impact.
Some species preferred intermediate (medium to high)
disturbance (Pseudagrion massaicum, Ictinogomphus ferox, Phyllomacromia contumax, Orthetrum chrysostigma
and Trithemis annulata), and all can be common at artificial reservoirs, as well as natural pools, with a margin of
suitable vegetation. This is another parallel between
intermediate elephant and human impact.
A combination of site separation and low impact benefited certain habitat specialists (Ceriagrion katamborae,
Pseudagrion deningi, Agriocnemis victoria, Orthetrum
icteromelas, Trithemis monardi, T. stictica, Rhyothemis
fenestrina and Aethriamanta rezia). Tentatively, these
species are sensitive to any major effects of elephant. Such
species have great potential as indicators of healthy
freshwater systems in southern Africa (Suhling et al.
All of the 16 species (over 50%) that occurred in very
highly or extremely highly disturbed habitats were also in
lesser disturbed habitats, indicating that intense disturbance
is not a prerequisite for their appropriate habitat. For

Table 3 Summary of eigenvalues and Monte Carlo test for CCA ordination
Speciesenvironment correlations
Cumulative % variance of species data
Cumulative % variance of environmental data
Monte Carlo test of significance





Total inertia










First canonical axis (0.721)



All canonical axes (0.969)





J Insect Conserv (2008) 12:493498

No. of speci es


Very High




Impact level

Fig. 4 Total number of species recorded at each site

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.

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