BioRisk 5: 73-84 (20 | 0) Apeer-reviewed open-access journal

doi: 10.3897 /biorisk.5.843 RESEARCH ARTICLE & B lO R IS k

http://biorisk-journal.com/

Impacts of extreme weather and climate change
on South African dragonflies

Michael J. Samways

Department of Conservation Ecology and Entomology, and Centre for Invasion Biology, University of Stellen-
bosch, P/Bag X1, Matieland 7602, South Africa

Corresponding author: Michael J. Samways (samways@sun.ac.za)

Academic editor: jiirgen Ott | Received 29 July 2010 | Accepted 9 August 2010 | Published 30 December 2010

Citation: Samways MJ (2010) Impacts of extreme weather and climate change on South African dragonflies. In: Ott J
(Ed) (2010) Monitoring Climatic Change With Dragonflies. BioRisk 5: 78-84. doi: 10.3897/biorisk.5.843

Abstract

The absence of ice sheets for many millions of years, yet variable topography and changing climate, has
generated considerable biodiversity in South Africa. There is no evidence to date that anthropogenic
climate change has affected odonate populations in the region. One reason is that the highly varying
weather and climate constitutes considerable background noise against which any effects of modern cli-
mate change must be measured. Evidence is accumulating that the Holocene interglacial and gradual
warming has left some species with isolated populations in montane areas among a matrix of arid land.
Many South African odonate species are remarkably vagile and elevationally tolerant, readily immigrating
into and emigrating from pools during wet and dry phases respectively. Some species take this movement
to greater extremes by moving the southern margins of their geographical range back and forth with
varying climate. After floods, populations of riverine odonates can recover within a year, although where
the riparian corridor has been stripped of its trees, the recovery is very slow. Various synergistic impacts,
particularly from invasive alien woody plants, are having a severe impact on many riverine species, and
reducing their ability to respond positively to changing environmental conditions. Large-scale removal of
these woody aliens is greatly benefiting the odonates’ ability to survive in the short-term and to restore

natural corridors for movement in the face of possible future climatic changes.

Keywords

Climate change, extreme weather, dragonflies, South Africa

Copyright M.J. Samways. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

74 Michael J. Samways/ BioRisk 5: 73-84 (2010)

Introduction

The last 130 000 years has seen three major events in the earth’s climatic history. Dur-
ing the Eemian (130 000 — 110 000 years BP) the climate was a major driver of plant
populations and their evolution. During the glacial period (110 000 — 10 000 years
BP) there were alternating cold and temperate conditions with major shifts in insect
assemblages at any one location (Elias 1994, Coope 1995, Ponel et al. 2003). In the
current interglacial (10 000 years BP to Present) there has been a general warming of
the earth’s climate (with minor cool periods) when human pressure on the landscape
increased considerably, especially in localized areas (Fagan 2004).

This natural background warming has accelerated in recent years through the im-
pacts of the industrial age. The important point here is that in very recent years, it has
been a combination of global climate change and anthropogenic landscape fragmen-
tation that has caused considerable stress on insect populations (Warren et al. 2001,
Travis 2003).

While the southern hemisphere has been subject to the various climatic changes
as in the north, it has not had a major glaciation event for over 200 million years. This
lack of cold sterilization, combined with milder climatic variations, various orographic
conditions and much soil variation, has been the background for considerable specia-
tion and the evolution of many localized endemics (Fjeldsa and Lovett 1997a,b; Gold-
blatt 1997; Jiirgens 1997). Additionally, the historical human impact on the landscape
has only been intensive over wide areas in the last 200 years, but still leaving today
many areas that are only minimally disturbed. These various factors have all played a
major role in determining the local distribution of the biota. Yet there is another fac-
tor that is part of this backdrop when assessing any contemporary climate change on
dragonflies in the region. ‘This is the ‘El Nifio Southern Oscillation’ (ENSO), which
produces alternately droughts and floods, often devastatingly so. ENSO, in addition to
the overall warming during the current interglacial, constitutes the background noise
against which we must measure the localized effects of anthropogenic climate change.

What the botanists have to say

Rutherford et al. (1999) undertook an interesting study on the effect that current an-
thropogenic climate change may have on floristic composition in reserve areas in South
Africa. A climate change scenario of increased temperature but no change in precipita-
tion would result in an increased growing season in some areas and a decrease in others.
In this latter group is a reserve in the Northern Cape (Augrabies Falls National Park)
which could lose a third of its plant species. However, such extinctions depend on the
tolerances of the focal species, and their departure may be partly balanced by immi-
grating species more tolerant of altered conditions. Within the Cape Floristic Region,
most species of Proteaceae are likely to experience geographic range contractions and
even a shift to higher elevations (Midgley et al. 2003). The evidence clearly points to

Impacts of extreme weather and climate change on South African dragonflies 75

the necessity of maintaining landscape linkages to ensure propagule migration. The
challenge however, is that 30% of the currently remaining natural vegetation could
be transformed within the next 20 years, with increased agriculture, urbanization and
invasion by alien woody plants (Rouget et al. 2003). Midgley et al. (2002) go on to
point out that the biome-level approach appears to underestimate the risk of species
diversity loss from the impacts of climate change because many narrow range endem-
ics will experience range dislocation throughout the biome and not necessarily just in
areas identified as biome contractions. Midgley et al. (2002) recommend that targeted
vulnerable plant species be monitored both for early warning signs of enhanced climate
change and as empirical tests of predictions. Williams et al. (2005) then add that the
sensitive and vulnerable species may not necessarily be able to make use of linkages
unless particular and suitable habitat conditions are available for stepping stone events.
Indeed, many Proteaceae species will not even disperse across gaps of unsuitable habi-
tat of only 100 m.

Marginality and the South African odonate fauna

South Africa has a proportionately high number of endemic and globally threatened
odonate taxa compared with neighbouring areas (Samways 1992). These globally
threatened taxa have small and restricted geographical ranges mostly in ecosystems
sensitive to human impact. In contrast, most of the nationally threatened taxa are
marginal, and the threats are largely natural, driven by ENSO events (Samways
2006a). This has an important bearing on practical national Red Listing, because
the issue is intrinsically bound to marginal rarity and associated risks of natural local
extinction. Such local extinctions of even common insects are not unusual (Dempster
1989). Typically, marginal populations are small, sparse and isolated from each other
(Lawton 1993) with considerable variation between the populations in genotype and
phenotype (Shreeve et al. 1996). Little evidence is available on how these odonate
species respond to climatic variations in the core of their geographical range. How-
ever, evidence from East Africa (Clausnitzer 2005) and Seychelles (Samways 2003b)
suggests that in general the forest species are restricted to that particular habitat, and
only following forest corridors, while many of the open habitat species are more vagile
and opportunistic.

Evidence for loss of marginal South African odonate populations is strong. Besides
the well-known migrants such as Anax ephippiger, Pantala flavescens, Sympetrum fon-
scolombii and Tramea spp. (the ‘sweepstake’ species), which readily colonize temporary
pools, there are many species (the ‘stepping stone’ species) which rapidly locate and
colonize newly-created artificial water bodies (Osborn and Samways 1996). Among
this group of species is Anax tristis, with its remarkable flight ability. Most of the species
in this last category are geographically widespread generalists, although not necessarily
strong fliers (e.g. Africallagma spp. and Agriocnemis spp.), although Agriocnemis falcif-

era (Fig. 1) is a localized species and national endemic which readily colonizes artificial

76 Michael J. Samways/ BioRisk 5: 73-84 (2010)

Figure |. Agriocnemis falcifera, a South African endemic, but nevertheless an opportunistic species which

readily colonizes new pools.

water bodies so long as they have a constant water level and a wide and shallow margin
with an abundance of grasses and sedges.

These ‘stepping stone’ species are highly elevationally tolerant and may move ac-
cording to prevailing weather conditions (Samways 1989a; Niba and Samways 2006).
Species such as Diplacodes luminans (Fig. 2), Urothemis edwardsii and Rhyothemis
semihyalina occasionally colonize high-elevation pools when weather conditions are
suitably warm. These ‘stepping stone’ species are highly vagile and habitat-tolerant,
responding to ENSO events with remarkable speed. Even the localized Orthetrum ro-
bustum (Fig. 3) behaves in this way. In wet years, it is common in temporary pools
in iSimangaliso Wetland Park, retreating to permanent lakes such as Lake Bhangazi
North, during dry years.

Most of these ‘stepping stone’ species are inhabitants of still water, which includes
pools (‘kuile’) left behind in dropping rivers as the dry season advances. In the West-
ern Cape, species in this category include the very rare and threatened Metacnemis
angusta (Fig. 4) and Proischnura polychromatica, which despite their extremely limited
geographical distribution, are highly responsive and rapidly locate their preferred habi-
tat (Samways et al. 2005). Furthermore, the riverine ‘stepping stone’ species recover
within one year after an extreme flood event (Samways 1989b).

There is a further category, the ‘range shifting’ species, which change the south-
ern limits of their geographical range margins in accordance with ENSO events.
During the extreme floods in February 2000, two taxa Pseudagrion coeleste coeleste
and P sjoestedti became nationally extinct, as their habitat was changed from tree-
lined river channels to early succession vegetation in the south-eastern corner of the

Impacts of extreme weather and climate change on South African dragonflies fas

Figure 2. Diplacodes luminans, a ‘stepping stone’ species which moves to new pools up elevational gra-
pping p p p g

dients during warm years.

Figure 3. Orthetrum robustum populations expand to new pools in wet years, retreating to permanent

ones in dry years.

Kruger National Park. While P coeleste has recovered by colonizing another location,
P sjoestedti has not yet reappeared. These were not the only species that were affected.
Others, such as Lestinogomphus angustus (Fig. 5) and Pseudagrion sudanicum, suffered
severe population crashes but nevertheless survived on a reduced source population.

78 Michael J. Samways / BioRisk 5: 73-84 (2010)

Figure 4. Metacnemis angusta is an extremely rare species which readily colonizes suitable habitat when,

for example, it is restored by removing invasive alien woody plants.

Figure 5. Lestinogomphus angustus populations dropped substantially during the huge floods of February
2000.

In contrast, some others benefited from these weather-induced habitat changes. The
locally rare Trithemis werneri, which prefers savanna rivers with wide riparian zones,
responded by becoming abundant the following year. Similarly, Crocothemis erythraea
readily colonizes quiet river zones where the natural tree canopy has been thinned or
largely removed.

Impacts of extreme weather and climate change on South African dragonflies iy)

Figure 6. Aciagrion dondoense is a recent invader into South Africa, possibly driven south by extreme floods

to the north, and seemingly partly responsible for the local demise of Agriocnemis ruberrima ruberrima.

The most extreme geographical range shift was by Aciagrion dondoense (Fig. 6),
which prior to the floods of 2000, was not known in South Africa. By 2001, it was
present in large numbers breeding in the iSimangaliso Wetland Park. It appears that
the extensive flooding in southern Mozambique pushed it southwards where it then
established, possibly with the benefit of warm conditions in recent years. This was not
an isolated case, with other species, such as Lestes dissimulans and L. uncifer apparently
also moving south. Both these species appear to oscillate their southern range margins,
as indicated by records over the last few decades. In 1956, L. uncifer was even recorded
as far south as Durban (Pinhey 1984).

Evidence (from the ongoing South African dragonfly database) is accumulating
that these huge shifts in southern range margins may be a relatively common phenom-
enon in species with a wide African distribution. This cautions mapping exercises, and
emphasizes that a distribution map for many of these species can be spurious depend-
ing on the timing of the original data records in the ENSO cycle.

The case for narrow-range endemics

All except two (P coeleste umsingaziense and Agriocnemis ruberrima ruberrima) of the
14 globally Red Listed South African odonate taxa inhabit running water or pools in
river systems. The two species which inhabit still water are subject to both anthro-
pogenic pressure, such as overgrazing and loss of habitat to urbanization, and ex-
treme natural events such as drought. This combination of anthropogenic impact and

80 Michael J. Samways / BioRisk 5: 73-84 (2010)

adverse natural conditions appears to be particularly synergistic, causing permanent
local extinction at many sites, without recolonization, as seen in O. robustum. There
seems to be another twist in the case of A. ruberrima, which went rapidly extinct
when Aciagrion dondoense invaded its stronghold, Mfabeni Swamp in the iSiman-
galiso Wetland Park. It appears that A. dondoense became an intense competitor and
possibly predator, as it became very abundant in the same swamp microhabitats as A.
ruberrima when it disappeared.

Impacts on all the remaining globally Red Listed South African odonates are distinct-
ly anthropogenic, although of course, adverse weather cannot be ruled out as a future syn-
ergistic threat. [he major impact for most of these taxa is from invasive alien trees ([ATs)
which shade out the microhabitats and stress the local populations (Samways and Taylor
2004; Samways 2006b). The reverse also occurs. When the IATs are removed there is re-
markable recovery of populations, and for three species possibly even extinction reprieve
(Samways et al. 2005; Samways and Sharratt 2010). The recovery effect is largely through
re-establishment of suitable sunny habitats, although in some cases there is also recovery
of populations through improvement in the water table when the [ATs are removed.

Evidence points to many of the endemic species having climatic relictual distribu-
tions. Certain outlier populations show this very distinctly. There is an isolated popula-
tion of Chlorolestes tessellatus in Sevenweekspoort, apparently at least a hundred kilom-
eters from the nearest population, with arid and unsuitable habitat in between. It is a
similar situation for Chlorolestes fasciatus (Fig. 7) which is represented by an isolated
population in the mountains of the Mountain Zebra National Park, surrounded by a
particularly arid and unsuitable matrix.

Synthesis of findings to date

Although there has been a claim that recent human-induced climate change has in-
duced range shifts in certain herbivorous insect species in South Africa (Giliomee
1997), the range expansions can be more parsimoniously explained by a switch in host
plant (Geertsema 2000). Nevertheless, there is the potential for major geographical
range shifts, and even local extinctions of insects with climate elevation of only 2°C
(Erasmus et al. 2002). However, empirical evidence from insects suggests that simple
climate envelope models overlook the nuances of biology of a species, making future
predictions of geographical range change extremely tenuous (Samways et al. 1999).
Evidence from dragonflies in the region clearly suggests that they have already been
through substantial climatic bottlenecks, with vicariance being common among many
of the narrow endemics, and great elevational tolerance and vagility being a feature
of the more widespread species. This situation parallels that in plants, where certain
species are at risk of extinction from lack of responsive mobility to climatic change
(Williams et al. 2005), which could also be the case with dragonflies (Samways 2008).
However, there are two mitigating factors. Firstly, evidence from highly threatened
species’ responsiveness to invasive alien plant removal shows that they are remarkably

Impacts of extreme weather and climate change on South African dragonflies 81

Figure 7. Chlorolestes fasciatus is a South African endemic, with climatic relictual populations in isolated
montane areas now in an arid matrix.

resilient, rapidly colonizing and populating re-instated habitats where the invasives
have been removed. Secondly, many of the narrow-range endemic species still have the
opportunity to shift elevation and move up the streams. This assumes of course that the
streams still would have sufficient flow at the higher elevations. ‘This is indeed a prob-
lem in Mayotte (Samways 2003a), while in the Seychelles, the high elevation endemic
species are to some extent tolerant of droughting conditions and minimal flow in the
upland streams (Samways 2003b). Behavioural flexibility to elevation is thus already
built into the genotype of many dragonfly species in this ENSO-prone region.

Probably the greatest threat to South African dragonflies in the current changing
climate scenario is from the synergistic effects of anthropogenic disturbance, particu-
larly from alien plants. This sort of synergism between climate change and habitat dis-
turbance has been described by Travis (2003) as “a deadly anthropogenic cocktail”. For
South African dragonflies, like British butterflies (Warren et al. 2001), this may well be
the case, with the mountains possibly being their saviour, as has been the case during
the Quaternary of Europe (Hewitt 2003). Clearing of the invasive alien woody plants
not only improves the maintenance of biodiversity on site but also opens up riparian
pathways along the elevational gradient. This has long-term evolutionary value as well
as short-term ecological benefit. Bearing this in mind, there does appear to be some-
thing concrete that we can do to lessen the blow of further climate change. ‘The first
results on South African Odonata species suggest that rehabilitating riparian corridors
might well benefit a whole range of aquatic species (Smith et al. 2007; Samways and
Sharratt 2010; Magoba and Samways in press).

82 Michael J. Samways / BioRisk 5: 73-84 (2010)

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