Cimbebasia 18: 207-224, 2003
207
Odonata of African arid regions - are there desert species?
Frank Suhling1, Reinhard Jödicke2 & Wolfgang Schneider3
1
Technische Universität Braunschweig, Department of Geoecology, Langer Kamp 19c, D-38106 Braunschweig,
Germany; e-mail: [email protected]
2
Am Liebfrauenbusch 3, D-26655 Westerstede, Germany; e-mail: [email protected]
3
Department of Zoology, Hessisches Landesmuseum, Friedensplatz 1, D-64283 Darmstadt, Germany;
e-mail: [email protected]
This paper reviews current knowledge of desert-inhabiting Odonata in Africa. By comparing
literature data of countrywide and some regional African Odonata surveys we identify typical
desert dragonflies and analyse traits enabling them to survive in desert conditions. Two criteria are
used in order to differentiate such species, viz. endemism in deserts and widespread occurrence in
desert areas. Very few endemics occur in the Sahara, Arabian and Somalian Deserts of Africa,
almost all occurring in permanent waters. In southern African arid regions no desert endemics are
known, and most of the widespread desert-dwelling species do not appear to be restricted to
deserts, their main habitats being in savanna wetlands. All are shown to be common over most
of the Afrotropical and/or southern Palaearctic Regions. In contrast to most endemics, widespread species colonise a broad spectrum of aquatic habitats, including ephemeral waterbodies.
Some traits allow Odonata to exist in deserts. Typically desert species are highly mobile and
multivoltine, i.e. they have up to four or five, sometimes overlapping, annual generations, allowing them to develop in temporary wetlands. Many species are opportunistic with respect to the
type of aquatic habitat they colonise. A cluster analysis on presence/absence data for 529 dragonfly
species reveals that, among dragonfly assemblages in deserts, most are similar to each other, but
differ from those in non-desert regions in Africa. Thus, it is concluded that there are typical
dragonfly assemblages of desert wetlands that are similar in the African and the Arabian Deserts.
Based on research, Odonata assemblages of some typical wetlands in the Sahara and Namib
Deserts are described.
INTRODUCTION
Odonata are common freshwater-associated organisms, being particularly widespread in the tropics. Having aquatic larval development, Odonata
would be expected to be absent or poorly represented in desert environments. This is indeed the
case in most desert habitats. Consequently, textbooks outlining desert ecology rarely refer to
Odonata (vide Ayyad & Ghabbour 1986; Evenari
et al. 1985; Louw & Seely 1982, 1986). Deserts do,
however, contain wetlands, which are colonised
by a number of aquatic animal groups, including
Odonata (Dumont 1982; Krupp & Schneider
1988). Pinhey (1978) classified a number of Afri-
can Odonata species, including Crocothemis erythraea, Trithemis kirbyi ardens and Paragomphus genei,
as being tolerant of arid conditions.
In deserts, episodic precipitation may establish
ephemeral or temporary rivers or ponds (Jacobson et al. 1995). In mountainous regions, springs
and permanent streams may provide water bodies suitable as habitats for dragonfly larvae (e.g.
Martens & Dumont 1983; Schneider 1988). Similar water bodies may occur along the courses of
ephemeral rivers resulting from resurgence of underground water dependent on geology or topography (Jacobsen et al. 1995). Additionally, there
are man-made habitats created by impounding
Proceedings of the 1st PHAON Meeting presented at the 2nd WDA International Symposium of Odonatology, Gällivare,
Sweden, 26th July 2001.
208
Cimbebasia 18, 2003
Figure 1. Distribution of arid regions of Africa and the Arabian Peninsula (after Shmida 1985). Names refer
to the Odonata checklists mentioned in the text (vide also Table 1).
ephemeral rivers and, particularly in northern Africa and Arabia, by pumping water for irrigation
purposes. Odonata are excellent flyers that are able
to cover long distances enabling them to colonise
even the most isolated habitats (Corbet 1999).
Indeed, almost all desert aquatic habitats are
utilised by Odonata for oviposition and larval development (e.g. Dumont 1978b: Saharan Ahaggar
mountain springs; Kimmins 1950: Saharan oasis;
Prinsloo 1990: Namib ephemeral river; Schneider
1988: Arabian oasis). Moreover, Odonata appear
to often be the dominant predators in desert
wetlands, in some cases preventing even anurans
from colonising ponds (Channing 1976).
In this paper we pose the question: are there species and specific dragonfly assemblages typical of
African deserts? We analyse literature data to (1)
separate desert endemics (for definition vide infra)
from common desert species and (2) to compare
the Odonata fauna of desert and non-desert regions, in order to identify similarities between
Suhling et al. – Odonata of African arid regions
faunas of several desert regions. The traits and
characteristics that allow Odonata to live under
desert conditions are discussed, and some typical
desert wetlands and their Odonata assemblages
are described.
Authors and dates of publication for species occurring in deserts are listed in Tables 2, 4 and 5; for
all others mentioned, authors and years are in the
text.
ODONATA OF AFRICAN DESERTS
Deserts are commonly defined by annual precipitation (Schulz 1995; Shmida 1985), although many
additional factors may further contribute to the
definition of deserts, such as vegetation (Irish 1994;
Rutherford & Westfall 1994). True deserts feature
precipitation below 120 mm per annum, and in
extreme deserts the figure may be below 70 mm
(Shmida 1985). Semi-deserts feature precipitation
between 100 and 500 mm (Schulz 1995; Shmida
1985). Such arid regions, which make up 38% of
the African Continent (Shmida 1985), are situated
in northern and southwestern Africa (Figure 1).
The Sahara is by far the most extensive arid region
covering most of northern Africa. The adjacent
Arabian Desert - although situated in Asia - is
included here, since, with the Sahara and the
Somalian Desert, it forms part of the Saharo-Arabian phytogeographical region. In addition, the
Odonata fauna of Arabia is broadly related to the
African (Schneider 1988; Waterston 1985). Desert
and semi-desert also cover the Somalian horn
southwards to Kenya and adjacent to southwestern Arabia. The arid island Soqotra (12°20’N,
53°00’E) is situated halfway between Somalia and
Yemen. These deserts are part of the southern
palaearctic desert belt which extents into central
Asia, where aridification began at the end of the
Miocene Period, some 8-10 million years ago
(Evenari 1985).
The deserts and semi-deserts of southern Africa
lie between 14° and 34°S. Traditionally, this arid
part of southern Africa has been separated into
three sections: Namib, Karoo and Kalahari (Werger
1986). There is evidence that the Namib in its
present stage was formed during the Pleistocene
209
Period due to the formation of the Benguela
Upwelling System (Walter & Breckle 1984). However, a geological review made it probable that the
Namib tract has not experienced climates more
humid than semi-arid for any length of time during the last 80 million years (Ward et al. 1983).
The Odonata of at least some parts of the northern Sahara and the Arabian Desert are comparatively well known (Jacquemin & Boudot 1999:
Morocco; Jödicke et al. 2000a: Tunisia; Samraoui
& Menaï 1999: Algeria; Schneider & Krupp 1993:
Arabia). For other desert regions, however, although country checklists are available, these do
not easily allow separation of the desert regions
from other ecosystems (e.g. Carfi 1974; Pinhey 1984,
1985; Samways 1999). Table 1 presents a compilation of literature sources referring to the Odonata
of African arid regions. This table also represents
the database here used to analyse differences between African regions (vide infra); so checklists of
non-desert regions are also cited.
DESERT ENDEMICS
Endemic species may provide one possible means
of characterising desert species. Desert endemics
are here regarded as species that exclusively occur,
or almost so, within desert and semi-desert regions (Table 2). Nevertheless, some of these species, such as Ischnura saharensis which is common
in the western half of the Sahara (Figure 2), may
have a wide distribution. Other species are known
from only a few sites. The known sites for
Paragomphus sinaiticus, however, indicate a wide distribution from the Sinai in the north to the Hoggarmountains, Niger, in the southwest and Oman in
the southeast. Agriocnemis sania is recorded from
the Libyan Sahara (Oasis of Gat), the Sinai, Israel,
and Somalia. Many species are restricted to the
southern edge of the Arabian Desert (Table 2).
Enallagma granti is so far known only from the
island of Soqotra. A Somalian-Arabian distribution is shown also by Enallagma somalicum and
Urothemis thomasi. In southern African deserts no
endemics could be identified (but vide infra). At
least for the Namib, this may be an effect of its
minor age or its smaller size as compared, for
example, to the Sahara.
210
Cimbebasia 18, 2003
Table 1. Compilation of literature sources related to Odonata in Africa and Arabia (vide Figure 3). Abbreviations: S Maghreb = Saharan fringe in Morocco, Algeria & Tunisia; S Tunisia = Tunisian Sahara; SW
Sahara = western Sahara, Mauritania, Saharan Mali, Aïr Mountains in Niger & Ahaggar Mountains in
Algeria.
Desert/Country
Morocco
S Maghreb
S Tunisia
Tunisia
SW Sahara
Libya
Egypt
Saudi Arabia
Yemen
Oman
Soqotra
Somalia
Namib
Namibia
South Africa
Gambia
Tanzania
Kenya
Uganda
Ghana
SW Cameroon
Literature source
Reviewed in Jacquemin & Boudot 1999.
Reviewed in Jacquemin & Boudot 1999; Jödicke et al. 2000; Samraoui & Menaï 1999.
Reviewed in Jödicke et al. 2000a.
Reviewed in Jödicke et al. 2000a.
Dumont 1976, 1977, 1978a, 1978b, 1978c.
Dumont 1974; Kimmins 1950; Nielsen 1935a, 1935b; Ris 1911.
Reviewed in Dumont 1980, 1991.
Reviewed in Schneider & Krupp 1993.
Reviewed in Schneider & Krupp 1993; Schneider & Parr 1998.
Reviewed in Schneider & Dumont 1997.
Reviewed in Schneider & Dumont 1998.
Carfí 1974; Dumont & Martens 1984.
Curtis 1991; Martens et al. 2003.
Curtis 1991; Jödicke & Martens in prep.; Martens et al. 2003.
Ris 1921; Samways 1999.
Gambles et al. 1998; Prendergast 1998.
Clausnitzer 2001 and in litt.
Clausnitzer 2001 and in litt.
Clausnitzer 2001 and in litt.
O’Neill & Paulson 2001.
Vick 1999.
Only sparse information exists regarding habitat
requirements of desert endemics (Table 3), and
for the majority of species the larvae and larval
microhabitat preferences remain unknown. The
larval habitat of some species have been described
previously, e.g. Paragomphus sinaiticus (in the Hoggar, Niger) (Martens & Dumont 1983), Agriocnemis
sania (in its northern range, vide Figure 2) (Dumont
1974) and I. saharensis (in Tunisia) (Jödicke et al.
2000a). Most desert endemics belong to the Zygoptera, which require aquatic vegetation such as
reeds, on which to oviposit, but such vegetation
rarely occurs in ephemeral habitats. As far as it is
known, all Zygoptera species are restricted to
permanent wetlands, such as small lakes or permanent springbrooks. Only Ischnura saharensis has
sometimes been encountered at semi-permanent
gueltas (Dumont 1982; Jödicke 1995), which were,
however, no more than 10 km distant from a permanent waterbody. All desert endemics may be
relics from more humid conditions in their respective regions (Dumont 1982).
ISOLATED POPULATIONS IN THE NAMIB
Although no endemic Odonata are known to occur in southern African deserts, something may
be learnt regarding the processes of endemism by
considering some Odonata species with remote,
isolated populations in Namibia. Aeshna minuscula
McLachlan, 1896, is regarded as being endemic to
South Africa (e.g. Chelmick 2001), but is also
known to occur in Namibia, over 1000 km north
of the principle range (Ris 1921), and has recently
been further recorded from a permanent springbrook of the Naukluft River (24°15’45”S,
16°13’50”E) (Martens et al. 2003). Typical habitats
of A. minuscula elsewhere include mountain pools
and deposition zones of streams (Samways 1999).
Pseudagrion kersteni lives at well-vegetated margins
of moderately flowing streams and rivers (Samways 1999), and is recorded from five sites in Namibia, including the Naukluft River. The remaining species are recorded from isolated sites in the
Namib Desert, such as Trithemis stictica, Orthetrum
Suhling et al. – Odonata of African arid regions
julia falsum Longfield, 1955, and Crocothemis
sanguinolenta, which are also normally associated
with permanent running water.
These populations may have become isolated at
suitable habitats (permanent springbrooks)
following the latest progressive aridification, which
occurred in the region about five million years ago,
during the late Miocene Period. Permanent
springbrooks are also the typical habitats of endemic species in northern African deserts. In the
Namib Desert, however, the isolation may be –
for historical or geographical reasons – not strict
enough for new species to evolve from these isolated populations. If other aquatic insects are considered, there are also very few Namib endemics
(vide Curtis 1991). This is chiefly due to the fact,
that the Namib is a mere 200 km in width, and
211
genetic exchange even between isolated populations and the main range are possible. Further
studies of the genetics of such populations may
lead to a better understanding of the processes of
isolation and speciation.
WIDESPREAD DESERT SPECIES
With the notable exception of Ischnura saharensis,
the majority of desert endemics have a very restricted distribution and do not normally constitute typical Odonata assemblages in deserts. Another definition of desert species must therefore
be formulated. If the African desert regions as a
whole are compared, 15 species are shown to occur in almost all regions (vide Table 4). All are widespread, ranging from africotropical to circum-tropical (vide Pinhey 1978), and a further 19 species are
A
C
B
D
Figure 2. Distribution maps of some Odonata desert endemics. A, Ischnura saharensis Aguesse (filled circles)
& Arabicnemis khalidi (Schneider) (open circle); B, Paragomphus sinaiticus Morton (filled circles) & Agriocnemis
sania Nielsen (open circles); C, Urothemis thomasi Longfield (filled circles) & Aeshna yemenensis Waterston
(open circles); D, Arabicnemis caerulea Waterston (filled circles) & Pseudagrion arabicum Waterston (open circles).
212
Cimbebasia 18, 2003
Table 2. Endemic species of African and Arabian Desert regions. No endemics are known from the Namib
Desert and Kalahari Sandveld. For the definition of desert endemics, vide text.
Species
Sahara Sinai Arabia Yemen Oman Soqotra Somalia Ethiopia
NW Sahara
Ischnura saharensis Aguesse
+
-
-
-
-
-
-
-
E Sahara/Arabian Deserts
Paragomphus sinaiticus Morton
+
+
+
-
+
-
-
-
Agriocnemis sania Nielsen
+
+
-
-
-
-
+
-
Somalian Desert/S Arabia
Enallagma granti (McLachlan)
-
-
-
-
-
+
-
-
2
Enallagma somalicum Longfield
-
-
-
-
+
-
+
+
Urothemis thomasi Longfield
-
-
-
-
+
-
+
-
S Arabian Desert
Aeshna yemenensis Waterston
-
-
-
+
-
-
-
-
Arabicnemis caerulea Waterston
-
-
-
+
+
-
-
-
Arabineura khalidi (Schneider)
-
-
-
-
+
-
-
-
Pseudagrion arabicum Waterston
-
-
+
+
-
-
-
-
1
3
1
Also occurring in Israel; 2Arabian populations are treated as Enallagma somalicum ssp. amitium Waterston, 1990; 3Waterston
& Pittaway (1991) consider Urothemis thomasi a subspecies of the oriental Urothemis signata (Schneider, 1988), and
Schneider & Dumont (1997) regard thomasi as a ‘bona fide species’’ with an East African ssp. aethiopica Nielsen, 1957.
regularly encountered in deserts, but are not as
widespread within Africa (Table 4). Clearly, there
are specific traits that allow such species to colonise desert wetlands (vide infra). All of these species also occur in the savanna and/or temperate regions, but not in rainforests, except in cases where
forests have been substantially altered through
human intervention, e.g. logging and farming.
ECOLOGICAL CHARACTERISTICS OF
DESERT SPECIES
Widespread species are mainly referred to here, as
listed in Table 4, as these species are the common
species of deserts. For the ecology of African
Odonata, vide Corbet 2003. As desert endemics are
rare, with the exception of Ischnura saharensis, they
depend on atypical habitats (Table 3) and may not
necessarily show traits required for survival in
desert environments.
LIFE CYCLES
Corbet (1999: 220) placed the Odonata into categories according to life-cycle features. Most species
listed in Table 4 belong to the A.2.2 type of Corbet
(1999). Such species are multivoltine, i.e. they have
up to four or five generations per year and may
breed in ephemeral ponds during the rainy season. Nomadic adults may be carried by rain-bearing weather-fronts (vide Corbet 2003). During the
dry season they may be restricted to a few permanent waters. Sympetrum fonscolombii, Pantala flavescens
and Crocothemis erythraea are typical species of the
Suhling et al. – Odonata of African arid regions
A.2.2. type. A further common type, A.1., is
facultatively multivoltine and unregulated as aquatic
habitats are continuously available, e.g. Anax
imperator. Many species may inhabit several habitat
types, and may prefer either type A.2.2. or A.1.,
depending on the type of habitat. In both types,
embryonic and larval development is short, lasting in total less than 120 days. In artificial ponds in
the Namib Pantala flavescens requires 35 days from
oviposition to emergence (F. Suhling pers. obs.).
In other species, this figure may be as short as 20
days (vide review in Corbet 1999: 630). A few species, such as Lestes pallidus, are univoltine, having
an egg diapause of several months or siccatating
as adults (type 2.1.2). There is no evidence to suggest that partivoltine species are to be found in
deserts, but there may be some that occur in permanent springbrooks. Most species, however, tend
to breed opportunistically in all kinds of wetlands.
In a significant number of species, such as Trithemis
kirbyi, there appears to be little or no reproductive
seasonality. In others, such as Sympetrum fonscolombii, the life cycle may be regulated by the wet and
dry seasons (vide infra). Reproductive seasonality
of species appears, however, to depend on the
213
habitat. All species breeding at ephemeral ponds
are seasonal, whereas at permanent wet-lands the
generations may widely overlap.
DROUGHT RESISTANCE
In Odonata, several cases of siccatation are reported, concerning all life stages. Some larvae may
survive long periods of complete desiccation (vide
review in Corbet 1999: 190). Interestingly, most
such observations are from temperate zones, not
deserts, as one would expect. Reports of living
larvae of Trithemis arteriosa and Orthetrum chrysostigma in dry mud of seasonal pools in the Sahara
(Dumont 1982) do not constitute evidence that
these species are able to survive as larvae until the
end of the dry season (Corbet 1999: 191). Larvae
of at least some species of gomphids and libellulids, including O. coerulescens anceps, are able to
survive in moist sand for at least one month and
emerge successfully (F. Suhling pers. obs.; Suhling
& Müller 1996). However, survival in desert species (Table 4) in the larval stage seems to be unusual. But, this is still an open question (vide infra).
Table 3. Habitat requirements of desert endemics from Africa and Arabia.
Species
Aeshna yemenensis
Agriocnemis sania
Arabicnemis caerulea
Arabineura khalidi
Enallagma granti
Enallagma somalicum
Ischnura saharensis
Paragomphus sinaiticus
Pseudagrion arabicum
Urothemis thomasi
Habitat
Waters in mountains above 2000 m a.s.l. (Waterston 1985).
Richly vegetated standing waters and marshes, occasionally in slow-running water
(Dumont 1974).
Shallow sections of a wadi with rich submerged and immersed aquatic vegetation
(Schneider 1988).
Springwater running over rocks with bushes, algae and mosses (Schneider &
Dumont 1995). Also in a shallow section of a wadi with rich vegetation (Schneider
1988). Prefers fast running waters (Schneider & Dumont 1997).
Wadis with running water, exuviae on vertical rocks (Schneider 1999; Schneider &
Dumont 1998).
Data unavailable.
Permanent streams, ponds and swamps, with dense vegetation, but low salinity;
rarely at temporal waters (Dumont 1984; Jacquemin & Boudot 1999; Jödicke
1995; Jödicke et al. 2000a).
Stagnant or near-stagnant but permanent waters, larvae in dense macrophyte
vegetation, exuviae on reeds (Martens & Dumont 1983; Schneider 1988).
Waters in mountains above 2000 m a.s.l.; perennial wadi pools (Schneider & Krupp
1993).
Stagnant pools in intermittent wadis (Schneider 1988; Schneider & Dumont 1997).
Species
Li Eg Ar Om Ye So
SI Na
World distribution
+
-
-
-
+
-
+
+
+
+
-
+
+
Africa, W Asia
+
+
+
+
+
+
+
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+
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+
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-
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+
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-
+
SW Old World
+
W Old World
+ Africa, Middle East, S Europe
+
W Old World
+
W Old World
+
Africa, Middle East
+
Africa, S Europe, Arabia
+
Circum-tropic
+
Africa, S Europe, Arabia
+
Eurasia, Africa
+
Africa, S Europe, Arabia
+ Africa, Middle East, S Europe
+
Africa, Middle East, India
+
Africa, S Europe, W Asia
-
-
-
-
+
+
+
+
+
+
+
+
-
+
+
+
-
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
-
+
+
+
+
+
Africa, W Asia
+
Africa, Arabia
+ S & central Africa, S & E Asia
S & central Africa, Arabia
+
S & central Africa, Arabia
+
S & central Africa, Arabia
Cimbebasia 18, 2003
Throughout Africa
ZYGOPTERA
Pseudagrion sublacteum (Karsch, 1893)
ANISOPTERA
Anax ephippiger (Burmeister, 1839)
Anax imperator Leach, 1815
Brachythemis leucosticta (Burmeister, 1839)
Crocothemis erythraea (Brulle, 1832)
Diplacodes lefebvrii (Rambur, 1842)
Orthetrum chrysostigma (Burmeister, 1839)
Orthetrum trinacria (Selys, 1841)
Pantala flavescens (Fabricius, 1798)
Paragomphus genei (Selys, 1841)
Sympetrum fonscolombii (Selys, 1840)
Trithemis annulata (Beauvois, 1807)
Trithemis arteriosa (Burmeister, 1839)
Trithemis kirbyi ardens (Gerstaecker, 1891)
Zygonyx torridus (Kirby, 1889)
Africa except NW
ZYGOPTERA
Ceriagrion glabrum (Burmeister, 1839)
Enallagma nigridorsum Selys, 1876
Ischnura senegalensis (Rambus, 1842)
Pseudagrion hamoni Fraser, 1955
Pseudagrion kersteni (Gerstaecker, 1869)
Lestes pallidus Rambur, 1842
Mo Ma ST Tu Sa
214
Table 4. Widespread desert Odonata and their distribution patterns. + species present, - species not recorded. Abbreviations: Mo = Morocco; Ma =
southern Maghreb; ST = southern Tunisia; Tu = Tunisia; Sa = southwestern Sahara (Niger, Mauritania, Algeria, West Sahara); Li = Libya; Eg =
Egypt; Ar = Saudi Arabia; Om = Oman; Ye = Yemen, So = Somalia; SI = Soqotra island; Na = Namib.
Table 4. cont. Widespread desert Odonata and their distribution patterns. + species present, - species not recorded. Abbreviations: Mo = Morocco;
Ma = southern Maghreb; ST = southern Tunisia; Tu = Tunisia; Sa = southwestern Sahara (Niger, Mauritania, Algeria, West Sahara); Li = Libya; Eg
= Egypt; Ar = Saudi Arabia; Om = Oman; Ye = Yemen, So = Somalia; SI = Soqotra island; Na = Namib.
Species
Li Eg Ar Om Ye So
SI Na
World distribution
-
-
-
+
+
+
-
+
+
+
+
+
-
+
+
+
+
+
-
-
+
+
+
+
-
Africa, Arabia
S & central Africa, Arabia
Africa, Arabia
+
+
+
+
+
+
+
+
+
+
+
+
-
+
-
-
-
-
N Africa, Arabia, Middle East
N Africa, Arabia, Middle East
+
+
+
+
+
+
+
+
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+
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+
+
-
+
+
-
+
-
+
+
-
-
N Africa, Europe, S Asia
Europe, W & Central Asia
Europe, N Africa, Arabia
Asia, N Africa
W Asia, Mediterranean, India
Arabia, Mediterranean, India
N Africa, S Europe, W Asia
+
+
+
+
+
+
-
-
-
-
-
-
-
NW Africa
Suhling et al. – Odonata of African arid regions
ANISOPTERA
Crocothemis sanguinolenta (Burmeister, 1839)
Nesciothemis farinosa (Foerster, 1898)
Urothemis edwardsii (Selys, 1849)
N Africa and Asia
ZYGOPTERA
Ischnura evansi Morton, 1919
Ischnura fountaineae Morton, 1905
ANISOPTERA
Anax parthenope Selys, 1839
Orthetrum brunneum (Fonscolombe, 1837)
Orthetrum coerulescens anceps (Schneider, 1845)
Orthetrum sabina (Drury, 1770)
Lindenia tetraphylla (Vander Linden, 1825)
Selysiothemis nigra (Vander Linden, 1825)
Sympetrum sinaiticum Dumont, 1977
Only in NW Africa
ZYGOPTERA
Ischnura saharensis Aguesse, 1958
Mo Ma ST Tu Sa
215
216
Cimbebasia 18, 2003
Siccatation in the egg stage is very common in
Odonata. Most Lestes Leach, 1815 (Lestidae), have
an obligate diapause in the egg stage, which allows them to survive several months (up to some
years?) of drought (Jödicke 1997). Some Libellulidae, such as the Indian species Potamarcha congener (Rambur, 1842), survive the dry season in the
egg stage (Miller 1992). In others, such as some
species of Sympetrum Newman, 1833, there is at
least a facultative egg diapase (Ando 1962; Corbet
1999: 56), thus allowing drought survival of certain widespread species (Table 4). Only Lestes pallidus, since lestid eggs are able to survive drought
(vide supra), may regularly survive dry periods in
the egg stage, although it also siccatates in the adult
stage (vide infra), whereas the Anax (Leach, 1815)
species and perhaps Ischnura saharensis may facultatively do so. The implication is that dry-season
survival of the aquatic stages appears to be uncommon in desert species as well as in endemics, most
of these developing directly and requiring free
water.
Tropical centered Odonata (vide Corbet 1999: Chapter 7) frequently feature a relatively long adult stage,
as compared to temperate species (e.g. Fincke 1992).
Also, some Algerian Odonata spend the duration
of the dry season in mountains as adults (Samraoui
et al. 1998), a habit which is interpreted as aestivation. Such dragonflies remain reproductively immature until they return to their oviposition sites in
autumn. It is to be expected that postponed reproductive maturation is common in desert Odonata,
although relevant information is available only for
Sympetrum sinaiticum, S. fonscolombii and Lestes pallidus, (Corbet 1999: 635; Jödicke in press; Jödicke
et al. 2000a, 2000b). In some regions, however,
Lestes pallidus, may also siccatate in the egg stage, as
reported for Lestes plagiatus from Nigeria (Gambles 1960). Sympetrum fonscolombii, in contrast, normally does not siccatate at all, since it acts opportunistically (vide supra).
THERMOREGULATION
Adult dragonflies are reproductively active only
during the day, some being able to withstand very
high daytime temperatures. Trithemis kirbyi ardens
often perches on rocks or on sandy ground, which
may reach temperatures well above 60°C. At high
temperatures Libellulidae and Gomphidae characteristically rest in the ‘obelisk position’, i.e. orientating the tip of the abdomen directly towards
the sun, so reducing bodily exposure to heat
(Corbet 1999: 285). Some species avoid the hottest hours of the day by restricting flight activity to
the morning and evening (Utzeri & Gianandrea
1989), while others still rest for extended periods
in shady spots, e.g. Crocothemis divisa Karsch, 1898,
which rests on termite mounts in Côte d’Ivoire
(Grabow et al. 1997). Thoracic temperatures in
North American desert dragonflies often exceed
the thermal death point, which may be as high as
53°C, of congeners and conspecifics inhabiting
cooler habitats (Polcyn 1994).
OVIPOSITION
All Zygoptera and Aeshnidae (Anisoptera) generally oviposit endophytically, whereas all other
Odonata are exophytic, although some may oviposit on vegetation. The desert species (Table 4)
include only 10 Zygoptera species, although they
usually equal the Anisoptera in numbers elsewhere.
One probable reason for this is that endophytic
species require reeds or other submerged vegetation for oviposition, and such conditions do not
normally occur in many desert wetlands. Species
of Anax Leach (Aeshnidae), though endophytic,
are able to oviposit elsewhere, e.g. in wet mud or
rotten wood. Most desert species - particularly those
breeding in ephemeral pools such as Pantala
flavescens, Orthetrum chrysostigma and Paragomphus
genei - oviposit in open water.
MOBILITY
Odonata are generally excellent fliers and highly
mobile. Anisoptera are, however, more powerful
flyers than Zygoptera (Rüppell 1989) and thus have
greater potential for colonising desert wetlands.
Some species of Anisoptera have been encountered during migration (Corbet 1999: 409; Schneider 1981). These include some desert species (Table 4), namely Anax ephippiger, A. imperator, A.
parthenope, Diplacodes lefebvrii, Lindenia tetraphylla,
Pantala flavescens, Selysiothemis nigra, Sympetrum
fonscolombii and Zygonyx torridus. Some Zygoptera
Suhling et al. – Odonata of African arid regions
also migrate, including all Ischnura Charpentier,
1840, in Table 4 (Corbet 1999: 411). Some Anisoptera are true long-distance migrants, which may
enter deserts or even cross them in large aggregations (Corbet 1999: 429). Pantala flavescens is an
obligate migrant, which is even known to cross
oceans (Corbet 1999: 405). Sympetrum fonscolombii
and Anax ephippiger sometimes enter northern Europe, probably from northern Africa and/or Arabia (Burbach & Winterholler 1997; Lempert 1997),
and S. fonscolombii is a seasonal migrant in the Cape
Region of South Africa (Barnard 1937). The spatial displacement of dragonflies may be assisted
by weather and storm fronts (vide Corbet 2003).
An important phenomenon is the annual shift
of the Inter-Tropical Convergence Zone (ITCZ),
which determines the season of rainfall in the tropics. Air masses transported with the ITCZ may
contain Odonata (Corbet 1999: 400, 2003) and
transport them into deserts seasonally. Many of
these species form large, conspicuous aggregations
comprising millions of individuals while migrating.
HABITAT REQUIREMENTS
Most desert wetlands are temporary, i.e. they contain water only for a certain period in the year, or
ephemeral, i.e. they are temporary but contain water
at unpredictable times (Williams 1987), which may
be either running or standing waters (Jacobson et
al. 1995; Uys & O’Keefe 1997; Williams 1987).
Permanent waters are usually associated with
springs and may consist of small streams or chains
of pools of a few kilometres in length. Other
permanent waters are large rivers, such as the Nile,
Niger, Orange or Kunene, crossing the desert, and
large man-made reservoirs or impoundments.
Most desert species are opportunistic users of such
wetlands. As many species are multivoltine and
fly over long distances, they are able to produce at
least one generation in ephemeral wetlands and
may produce a further generation if in permanent
wetland. Pantala flavescens, Orthetrum chrysostigma,
Sympetrum fonscolombii and Trithemis kirbyi ardens,
for example, require the presence of water and
probably the absence of predators and competitors in order to complete their development. Predators are an important factor in the structuring of
Odonata assemblages (e.g. McPeek 1990; vide also
217
review in Johnson 1991), and these Odonata species may fail to reproduce in habitats with a high
density of fishes.
Other Odonata species are more restricted in their
habitat preferences. The endophytic Zygoptera and,
to a lesser extent, Anax spp., require vegetated wetlands for oviposition (vide supra). Crocothemis erythraea and Diplacodes lefebvrii also prefer vegetated
wetlands, although they oviposit exophytically (vide
Busse & Jödicke (1996) for discussion of the significance of substrates suitable for development in
ephemeral waters). Paragomphus genei mainly inhabits ephemeral and permanent running waters or
pools in ephemeral rivers, as well as large impoundments. This species shows no particular
preference for vegetation, being also able to develop over a very short period, and yet appears to be
confined to river valleys. Some species occur only
in permanent (often running) waters, e.g. Crocothemis sanguinolenta, Orthetrum coerulescens anceps and
Pseudagrion kersteni. Zygonyx torridus is even more
limited as larvae inhabit rapids and waterfalls,
which are very scarce in desert environments. Brachythemis leucosticta mainly inhabits large open waterbodies, such as rivers and impoundments.
Some Odonata are able to complete their development in ephemeral waters of highly variable ionic
concentrations, which may exceed those of seawater (vide review in Corbet 1999: 193). Australian
desert dragonflies of the genera Anax, Orthetrum
Newman, 1833, and Diplacodes Kirby, 1889, regularly reproduce in salt lakes (Halse 1998; Timms
1993). Of the African desert species, Anax
ephippiger, Lindenia tetraphylla, Orthetrum trinacria
and Pantala flavescens have been recorded as reproducing in brackish waters (Cheng & Hill 1980;
Gallardo-Mayenco 1994; Krupp & Schneider 1988).
In the Namib Desert, Ischnura senegalensis,
Crocothemis erythraea and Orthetrum trinacria have
been observed in water with a conductivity of 42
mScm-1, corresponding to about 150% more Salinity than seawater (F. Suhling pers. obs.).
ODONATA ASSEMBLAGES IN DESERT
REGIONS
We identified a certain number of Odonata species widespread in African and Arabian deserts. If
these make up the major part of desert Odonata
218
Cimbebasia 18, 2003
Figure 3. Cluster of 21 African regions (deserts and countries) based on presence/absence data of 529
species of Odonata (Ward method, squared Euclidian distance). The number of species is in parentheses.
assemblages, the faunas of desert regions should
be highly similar and different to non-desert regions, although the widespread species may also
occur there.
Therefore, as a part of this review, available data
sets from desert, savanna and rainforest regions
are analysed (Table 1). The data set comprises Odonata checklists of 21 African regions (Table 1) and
a total of 529 species. To test for similarities and
differences between the regional faunas, a Clusteranalysis (Ward method, Euclidian distance) on
presence/absence of the species was used.
Results illustrate two main clusters (Figure 3). The
first cluster includes all desert or semi-desert regions, including the Namib Desert, and the second cluster consists of savanna (eastern and southern Africa, including Namibia) and forest regions
(western Africa, parts of Uganda). Similarities in
the ‘desert cluster’ are much closer than in the ‘savanna and forest cluster’. This we ascribe to lower
species numbers at all desert sites: 15 species cooccuring in almost all desert regions, and a further
18 that, although their range is limited, occur in
many deserts (vide Table 4). These species largely
constitute the regional desert assemblages, indicating that there are indeed typical desert Odonata
assemblages, which consist of widespread African
species (vide supra). The forest and savanna regions,
in contrast, have a higher number of species and
degree of endemism. For example, in the genera
Umma, Chlorocypha and Phyllomacromia there are
mostly forest endemics, and the genus Chlorolestes
is endemic to the Cape (vide Clausnitzer 2001;
Samways 1999; Vick 1999).
Examples of Odonata assemblages of most wetland habitats in African and Arabian deserts, based
on observations from the Sahara and Namib
Deserts, are presented in Table 5, including those
from geographically isolated permanent waters.
OPEN QUESTIONS
There are many open questions concerning African desert dragonflies. Some important arid regions have never been systematically surveyed, al-
Suhling et al. – Odonata of African arid regions
though some relevant data may exist, e.g. for all
southern parts of the Sahara and the Sahel, some
parts of the arid Kalahari and the Arabian Desert
(vide Table 1). One important ecological question
concerns drought resistance of the aquatic stages.
Are there desert species, which regularly or at least
occasionally siccatate in the larval or egg stage? Since
‘snapshot’ observations cannot answer that question, further experimental field studies are required,
e.g. by use of artificial ponds. Another topic concerns migration and isolation in desert species. To
what extent are single desert populations really
isolated? This topic shall require, beside detailed
surveys, genetic approaches. Migration cannot
merely be studied using traditional techniques, e.g.
mark-release-recapture methods, as Odonata may
migrate over very long distances (vide supra). New
approaches need to be taken, including molecular
genetics.
CONCLUSIONS
Exact knowledge of dragonfly faunas in African
deserts is not currently available, but this analysis
has shown that desert endemics, i.e. species occurring exclusively in deserts, are not particularly abundant or widespread. Three categories of species
can be identified from the study: (1) species that
appear to be relics from more humid conditions,
occurring in uncommon habitats, mainly permanent springs, streams or lakes; (2) widespread species occurring in almost all desert regions of Africa
and Arabia; (3) more regional species, but also
widespread and abundant in Africa and Arabia,
including only one desert endemic (Ischnura
saharensis). Most of the widespread species also
occur in other ecosystems, mostly savannas, which
represent their preferred habitat. Typical desert
species are regarded, however, as those that are to
be found in most aquatic habitats in deserts these
comprising the typical Odonata assemblages. Most
of the species in these assemblages exhibit characteristics such as high mobility and rapid development, which allow them to survive even in ephemeral aquatic habitats. These traits are pre-adaptations that enable aquatic animals to survive desert
conditions. Consequently, many desert species do
not have very strict habitat requirements. As aquatic
habitats are rare and unpredictable in time and
space, desert species behave opportunistically with
219
respect to habitat selection. Bearing all these facts
in mind, it is not surprising that desert Odonata
are very similar in Africa and Arabia, even if separated from the non-desert regions, and, moreover,
that they contain a low proportion of endemics.
ACKNOWLEDGEMENTS
We thank Eugène Marais (National Museum of Namibia)
and Andreas Martens (Zoologisches Institut, Universität
Braunschweig), who commented on earlier drafts of
this paper. Bernd Kunz supplied some field data on
characteristic desert habitats in Tunisia. The first author
acknowledges research support from the German Ministry of Science, BMBF 01LC0024. We also thank Philip
Corbet (United Kingdom), Viola Clausnitzer (Germany)
and one anonymous referee for improving the manuscript.
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Reservoir/dam
Sahara Namibia
Ephemeral running Vegetated permanent Permanent stream
waters
swamps
Sahara
Namibia
Sahara Namibia
+
+
-
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Ischnura fountaineae
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Anax imperator
Anax parthenope
Anax speratus Hagen, 1867
Ictinogomphus ferox (Rambur, 1842)
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Brachythemis leucosticta
Crocothemis erythraea
Crocothemis sanguinolenta
Diplacodes lefebvrii
Ephemeral pool
220
Table 5. Examples of Odonata assemblages of different types of habitats from the northern Sahara and the Namib Deserts. Note that there are wide
differences in the Zygoptera assemblages, while the Anisoptera are broadly similar. For the description of habitats vide Appendix.
Table 5. cont. Examples of Odonata assemblages of different types of habitats from the northern Sahara and the Namib Deserts. Note that there
are wide differences in the Zygoptera assemblages, while the Anisoptera are broadly similar. For the description of habitats vide Appendix.
Species
Sahara
Namibia
+
+
+
+
+
+
+
-
+
+
+
+
+
+
-
Reservoir/dam
Sahara Namibia
+
+
+
+
+
-
+
+
+
+
+
-
Ephemeral running Vegetated permanent Permanent stream
waters
swamps
Sahara Namibia
+
+
-
+
+
+
-
Sahara
Namibia
+
+
+
+
+
+
+
+
-
+
+
+
+
+
+
+
Sahara Namibia
+
+
+
+
+
+
-
+
+
+
+
+
+
+
+
+
Suhling et al. – Odonata of African arid regions
Orthetrum caffrum
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Orthetrum coerulescens
Orthetrum julia falsum
Orthetrum nitidinerve (Selys, 1841)
Orthetrum sabina
Orthetrum trinacria
Palpopleura lucia
Pantala flavescens
Selysiothemis nigra
Sympetrum fonscolombii
Sympetrum sinaiticum
Trithemis annulata
Trithemis arteriosa
Trithemis furva
Trithemis kirbyi ardens
Trithemis stictica
Urothemis edwardsii
Zygonyx torridus
Ephemeral pool
221
222
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Manuscript received January 2002; accepted May 2002.
Appendix. Description of localities and sites used in the interpretation of Odonata assemblages at different habitats (vide Table 5).
(1). Ephemeral and temporary ponds
Saharan example: TUNISIA, two temporary ponds at the
edge of the Chott el Jerid near Tozeur, 33°55’N, 08°11’E,
and near El Faouar, 33°22’N, 8°39’E.
Namib example: NAMIBIA, artificial ponds at Tsaobis Leopard Nature Park, 22°22’S, 15°44’E.
(2). Reservoirs/impoundments
Saharan example: MOROCCO, Barrage Youssef-Ben-Tachfine,
29°46’N, 09°27’W.
Namib example: NAMIBIA, Von Bach Dam, 22°00’S, 16°57’E.
(3). Ephemeral and temporary running waters
Saharan example:
TUNISIA,
Oued el Melah at road between
Metlaoui and Gafsa, 34°08’N, 8°19’E.
Namib example: NAMIBIA, Gaub River at Gaub pass/road
C14, 23°28’S, 15°45’E.
(4). Well-vegetated permanent swamps
Saharan example: TUNISIA, swamps with Salicornia and reed
fed by ditches from Jemna Oasis, 33°33’N, 09°00’E and
Ghidma Oasis, 33°26’N, 08°49’E.
Namib example: NAMIBIA, Ugab West, 20°57’S, 14°06’E.
(5). Permanent streams
Saharan example: TUNISIA , mountain springbrook at
Tamerza, 34°24’N, 07°58’E.
Namib example: NAMIBIA, Naukluft River, 24°15’S, 16°14’E.