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Global diversity of dragonflies (Odonata) in freshwater

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Hydrobiologia (2008) 595:351–363
DOI 10.1007/s10750-007-9029-x

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of dragonflies (Odonata) in freshwater

Vincent J. Kalkman Æ Viola Clausnitzer Æ
Klaas-Douwe B. Dijkstra Æ Albert G. Orr Æ
Dennis R. Paulson Æ Jan van Tol

Ó Springer Science+Business Media B.V. 2007

Abstract Larvae of almost all of the 5,680 species inhabitants of alpine mountain bogs, seepage areas in
of the insect order Odonata (dragonflies and damsel- tropical rain forests, and waterfalls. They are often
flies) are dependent on freshwater habitats. Both successfully used as indicators for environmental
larvae and adults are predators. The order is relatively health and conservation management. The highest
well studied, and the actual number of species may be diversity is found in flowing waters in rain forests of
close to 7,000. Many species have small distribu- the tropics, the Oriental and Neotropical regions
tional ranges, and are habitat specialists, including being the most speciose. This paper discusses diver-
sity, summarises the biogeography of dragonflies in
the different biogeographical regions and gives the
Guest editors: E. V. Balian, C. Lévêque, H. Segers and total number of species and genera per family per
K. Martens biogeographical region. Examples are given of areas
Freshwater Animal Diversity Assessment of particular diversity, in terms of areas of endemism,
presence of ancient lineages or remarkable recent
V. J. Kalkman (&)
K.-D. B. Dijkstra
J. van Tol
National Museum of Natural History Naturalis, PO Box radiations but no well-based review of areas with
9517, 2300 RA Leiden, The Netherlands high endemism of dragonflies is available so far. The
e-mail: [email protected] conservation status of dragonflies is briefly discussed.
K.-D. B. Dijkstra Species confined to small remnants of forest in the
e-mail: [email protected] tropics are most under threat of extinction by human
J. van Tol activities.
e-mail: [email protected]
Keywords Odonata
Dragonflies
Diversity

V. Clausnitzer
Gräfestraße 17, 06110 Halle/Saale, Germany Endemicity
Biogeography
Conservation
e-mail: [email protected]

A. G. Orr
Introduction
CRC-TREM, AES, ENS, Griffith University, Nathan,
QLD Q4111, Australia
e-mail: [email protected] With 5,680 extant species, dragonflies are a relatively
small order of insects. Their size and colour and their
D. R. Paulson
diurnal and often conspicuous behaviour make them a
Slater Museum of Natural History, University of Puget
Sound, Tacoma, WA 98416, USA popular group for both professional and amateur
e-mail: [email protected] entomologists.

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352 Hydrobiologia (2008) 595:351–363

Dragonflies are among the most ancient of winged with two extant species from Japan and the eastern
insects, dating back well into the Permian (Grimaldi Himalayas. The Anisozygoptera, which have some
& Engel, 2005). They include the largest insect that features recalling Zygoptera, are now often included
ever lived, the griffenfly Meganeuropsis permiana in Anisoptera (as done here), or combined with them
Carpenter, with a wingspan of c. 70 cm. Dragonflies under the new name Epiprocta. Zygopterans have a
are recognised by their long, slender abdomen; large broad head with widely separated eyes and similar
globular eyes, often making up a large portion of the fore and hind wings. Most species rest with wings
head; short antennae; and long wings, which have a closed. The larvae are slender and rely mainly on two
conspicuous nodus and usually a pterostigma. They or three caudal gills for respiration. Anisoptera are on
possess a unique mechanism of indirect sperm average larger and more robust than Zygoptera. Their
transfer: sperm are produced in the testes situated at hind wings are distinctly broader at their base than
the abdomen tip, but the secondary copulatory organs the fore wings and in most families the eyes touch on
that transfer them to the female lie on the ventral side top of the head. At rest most species spread their
of the abdomen base. Sperm must be transferred wings. The larvae are typically much sturdier than
externally to this organ before copulation. This those of Zygoptera and lack caudal gills: oxygen is
copulatory organ is used not only to inseminate, but absorbed through gills in the rectum. A general
also to remove the sperm of the female’s previous outline of odonate diversity is given by Silsby (2001).
mates. Sperm competition in Odonata was first A checklist of all dragonflies including synonyms and
reported by Waage (1979) and stimulated numerous references is found on http://www.odonata.info (van
studies, making dragonflies one of the most studied Tol, 2005)
animal groups in terms of reproductive behaviour. Dragonfly larvae live in freshwater environments
Another unique feature of odonates is the strongly and only a few species tolerate brackish conditions,
modified labium of the larva, which can be extended two of which even live in salt marshes and mangroves.
at great speed to seize prey. Both running and standing waters are used, while a
The extant dragonflies are divided into two few species are semi-terrestrial or inhabit water held
suborders, the Zygoptera or damselflies and the in tree holes, leaf axils and other phytotelmata. Many
Anisoptera or true dragonflies (Fig. 1). Until recently species have small distributional ranges, and are
a third suborder, Anisozygoptera, was recognised, habitat specialists, including inhabitants of alpine
mountain bogs, seepage areas in tropical rain forests,
and waterfalls. Larvae prey on all kinds of small
animals up to the size of tadpoles and small fish.
Larvae take from a few weeks to 7 years to develop.
Emergence takes place above water on plants or on
the shore, after which most species leave the water
edge to mature. Males return to the water to search for
females or to establish territories. Females often only
return to mate and to oviposit. Information on the life
history and behaviour of odonates is thoroughly
summarised in Corbet’s (1999) review of odonate
behaviour and ecology.

Species diversity

Information on the number of species of odonates is

derived from the Global Species Database Odonata
prepared for the Catalogue of Life (van Tol, 2005).
Fig. 1 The damselfly Pseudagrion kersteni and the dragonfly
Orthetrum julia sitting on the same perch. (photo: Viola Taxa were assigned to one or more of the biogeo-
Clausnitzer) graphical regions based on the authors information

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Hydrobiologia (2008) 595:351–363 353

and several key references (e.g., Lieftinck, 1949; conspicuous and many favour open habitats, although
Watson et al., 1991; Westfall & May, 1996; Okudaira in absolute numbers they still represent a large
et al., 1999; Needham et al., 2000; Tsuda, 2000; proportion of species to be described. This is
Wang, 2000). Subspecies were not considered. especially so for the Coenagrionidae in South Amer-
Table 1 enumerates the number of genera and species ica. Since 1970 an average of 38 species have been
per family for each biogeographical region. Family- described annually (Fig. 4). With an undiminished
level classification of odonates is poorly resolved, rate of description an estimated 95% of all species
although most families are broadly accepted. The will be described in 2030.
most recent contribution to the higher classification
of dragonflies was published by Rehn (2003). With
the advent of molecular techniques, revision of Processes influencing diversity of dragonflies
family-level classification may be expected.
In total 5,680 species of Odonata are known, 2,739 Factors influencing the distribution of dragonfly
belonging to the suborder Zygoptera (19 families) diversity can be divided into historical (geological)
and 2,941 to the suborder Anisoptera (12 families). and ecological factors. Both determine current spe-
Table 1 and Fig. 2 show that the tropics support by cies diversity, while composition at family and genus
far the most species of dragonflies. Besides higher level is predominantly determined by the first.
diversity at the species level, the number of families Dragonflies are an ancient group, and present-day
is also much greater in the tropics (Fig. 3). Twelve of distribution reflects the distribution of the families
the 31 families are restricted mostly to running waters before the break-up of Pangaea and subsequent super-
within tropical forest habitats. The two largest continental schisms. However, more well-founded
families (Coenagrionidae and Libellulidae) are be- phylogenetic reconstructions are needed before a
lieved to be relatively recent (Rehn, 2003). Almost all satisfactory synthesis of this subject can be written.
ubiquitous species belong to these two families and Today’s patterns of dragonfly diversity correspond
they dominate in unshaded habitats with stagnant largely with the present climatological zones. Tem-
water (both artificial and natural, e.g., savannas). perature accounts for a sharp increase of diversity
Both families include species with the greatest from the poles to the equator, while precipitation
migratory capacity, including those with distributions obscures this pattern by reducing diversity in areas of
spanning more than one continent and almost all low precipitation, resulting in ‘gaps’ in diversity.
species found on isolated islands. Diversity of tropical odonates is at least partly
It is estimated that between 1,000 and 1,500 explained by the high diversity of aquatic habitats
species of dragonflies await description (Table 1). If in tropical forests (Orr, 2006), especially in montane
this is true, the actual number of extant species may areas (Oppel, 2005). Mountains not only provide a
be close to 7,000. The Oriental, Australasian and greater contemporary diversity of habitats, but also a
especially the Neotropical regions hold the highest greater potential for survival in regional refugia. The
number of undescribed species. In the latter, new relative long-term stability of forest habitats (also in
species are still discovered more rapidly than the short term, the limited seasonality), which
descriptions are published (Paulson, 2004). The fauna provides opportunities for animals with a specialist
of Africa is relatively well known and relatively lifestyle, might also explain the high diversity of
depauperate. Overall the families Platystictidae, Pro- tropical odonates.
toneuridae, Gomphidae and Corduliidae are believed Speciation events in dragonflies can be directly
to hold relatively many undescribed species. They are linked to isolation events in the geological past such
typically inconspicuous odonates with small ranges, as Andean orogeny (De Marmels, 2001) and isolation
often confined to seepages or small runnels in tropical in refugia in southern Europe during the ice ages
forest. Gomphidae, Corduliidae and also Chlorogom- (Sternberg, 1998). Polhemus (1997) showed how a
phidae in Southeast Asia are difficult to collect as single coenagrionid founder evolved into 22 species
they spend little time at the waterside. The two of Megalagrion on the Hawaiian Islands. Speciation
largest families, Coenagrionidae and Libellulidae, are was not only promoted by isolation after the coloni-
relatively well known, because most species are sation of newly formed volcanic islands, but also by

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354 Hydrobiologia (2008) 595:351–363

Table 1 (a) Number of species per family per biogeographical region. (b) Number of genera per family per biogeographical region
PA NA NT AT OL AU PAC ANT World

(a)
Aeshnidae 57 42 127 44 138 76 13 – 441
Amphipterygidae – – 3 2 5 – – – 10
Austropetaliidae – – 7 – – 4 – – 11
Calopterygidae 37 8 61 20 60 4 – – 171
Chlorocyphidae 3 – – 41 80 15 – – 135
Coenagrionidae 95 101 370 197 185 170 88 – 1084
Cordulegastridae 18 9 1 – 27 – – – 49
Chlorogomphidae 5 – – – 40 – – – 41
Corduliidae 20 50 37 17 57 54 12 – 239
Dicteriadidae – – 2 – – – – – 2
Epiophlebiidae 1 – – – 1 – – – 2
Euphaeidae 11 – – – 65 1 – – 69
Gomphidae 127 100 273 149 358 42 – – 966
Hemiphlebiidae – – – – – 1 – – 1
Isostictidae – – – – – 41 5 – 46
Lestidae 17 19 38 17 39 29 3 – 148
Lestoideidae 2 – – – 4 9 – – 13
Libellulidae 121 107 352 245 190 184 32 – 1012
Macromiidae 6 9 2 37 50 16 – – 122
Megapodagrionidae 2 – 130 38 28 57 5 – 260
Neopetaliidae – – 1 – – – 0 – 1
Perilestidae – – 18 1 – – – – 20
Petaluridae 1 2 1 – – 6 – – 10
Platycnemididae 23 – – 33 130 37 – – 210
Platystictidae – 1 42 – 119 29 1 – 191
Polythoridae – – 58 – – – – – 58
Protoneuridae 1 3 94 37 57 52 – – 245
Pseudolestidae 7 – – – 15 – – – 22
Pseudostigmatidae – – 18 1 – – – – 19
Synlestidae 6 – 1 10 17 8 – – 37
Synthemistidae – – – – – 35 9 – 43
Total 560 451 1636 889 1665 870 168 0 5680
Undescribed 75–100 5–10 400–500 100–125 300–400 175–250 30–40 0 1085–1425
(b)
Aeshnidae 14 13 15 6 18 19 7 – 48
Amphipterygidae – – 2 1 1 – – – 4
Austropetaliidae – – 2 – – 2 – – 8
Calopterygidae 8 3 3 3 10 1 – – 16
Chlorocyphidae 3 – – 3 14 4 – – 18
Coenagrionidae 15 15 38 15 23 24 12 – 90
Cordulegastridae 3 1 1 – 5 – – – 5
Chlorogomphidae 1 – – – 1 – – – 1
Corduliidae 6 8 2 6 7 16 3 – 41
Dicteriadidae – – 2 – – – – – 2

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Hydrobiologia (2008) 595:351–363 355

Table 1 continued

PA NA NT AT OL AU PAC ANT World

Epiophlebiidae 1 – – – 1 – – – 1
Euphaeidae 5 – – – 12 1 – – 12
Gomphidae 33 14 26 20 43 9 – – 92
Hemiphlebiidae – – – – – 1 – – 1
Isostictidae – – – – – 11 1 – 12
Lestidae 3 2 2 1 5 3 3 – 8
Lestoideidae 1 – – – 1 2 – – 3
Libellulidae 31 27 44 53 56 45 16 0 143
Macromiidae 2 1 2 1 2 2 – – 4
Megapodagrionidae 2 – 14 6 10 6 3 – 39
Neopetaliidae – – 1 – – – – – 1
Perilestidae – – 2 1 – – – – 3
Petaluridae 1 2 1 – – 2 – – 5
Platycnemididae 4 – – 9 8 11 – – 25
Platystictidae – 1 1 – 5 2 1 – 6
Polythoridae – – 8 – – – – – 8
Protoneuridae 1 2 14 4 8 1 – – 25
Pseudolestidae 1 – –– – 3 – – – 3
Pseudostigmatidae – – 5 1 – – – – 6
Synlestidae 2 – 1 2 2 3 – – 8
Synthemistidae – – – – – 4 1 – 4
Total 137 89 186 132 235 169 47 0 642

habitat specialisation (stagnant water, seepage, included in the Palaearctic. The faunal diversity in
phytotelmata and swift streams) within an island. these areas is at least as high as in North America and
Speciation has also been promoted by the isolation of is far richer than in Europe. In China the Palaearctic
patches of tropical forest due to climatological factors fauna merges gradually into the Oriental fauna. This
(Dijkstra & Clausnitzer, 2006). Large river systems transition zone is very rich compared to the other parts
such as the Amazon and Congo basins, forming an of the Palaearctic and harbours many species not
ever-changing mosaic of land and water, probably occurring elsewhere in the Palaearctic. The large
also facilitated speciation, but distribution patterns in differences in diversity between different parts of the
these regions are known insufficiently to verify this Palaearctic are largely due to the advance of glaciers
hypothesis. during the Pleistocene ice ages, which impoverished
the fauna in the western two-thirds of the Palaearctic.
Here the main mountain ranges and seas run east–west
A brief outline of odonate biodiversity within the (e.g., the Mediterranean Sea, the Pyrenees, Alps and
biogeographical regions Himalayas) thus forming a barrier for northern species
retreating southwards. Similar factors also apply
Palaearctic today as Oriental species can easily penetrate into
the Palaearctic, but northward expansion of African
Large parts of the Palaearctic are relatively species and Oriental species into the western Palaearctic is
poor when compared with areas at the same latitude in hampered by the same barriers as those limiting
North America. Europe for instance has only slightly southward retreat in the past. The ice ages also
more than half the number of species of Texas. promoted speciation by isolating species in various
Exceptions are Japan, Korea and the part of China refugia, especially evident in Europe. Most Palaearc-

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356 Hydrobiologia (2008) 595:351–363

Fig. 2 Diversity of dragonflies per biogeographical region (species number/genus number). PA—Palaearctic, NA—Nearctic, NT—
Neotropical, AT—Afrotropical, OL—Oriental, AU—Australasian, PAC—Pacific Oceanic Islands, ANT—Antarctic

tic species with a more northern distribution are by thorn forest and then desert, but species of the
widespread, several ranging from Europe to eastern moister uplands of the Mexican Plateau have also
Russia or even into the Nearctic. Whether after the ice moved north into the southwestern states. Thus the
ages these wide-ranging species colonised the eastern latter region is a centre of diversity and endemism in
Palaearctic from the western Palaearctic or vice versa North America, as are the north-eastern and
is still a point of debate (Kosterin, 2005). south-eastern coastal plains, Allegheny-Appalachian

Nearctic 100%

The dragonfly fauna of the Nearctic is richer than that 75%

of most of the Palaearctic. As in the Palaearctic, the

eastern part of the Nearctic is richer than the western 50%

part, and most eastern states in the USA have larger

species lists than all of Europe. This is presumably 25%

because the humid East has had a continuous

0%
connection with the wet tropics to the south, and
an
tic

al
tic

fic
ca
ca

nt
rc

rc

si

ci

numerous tropical species have moved into south-

pi
pi

rie

la

Pa
ea

ea

tro
tro

ra
O
la

eo
ro

st
Pa

eastern USA, while the West has gone through arid

Af

Au
N

periods when odonate dispersal was interrupted and

Fig. 3 Percentages of species belonging to a family for the
aquatic faunas were presumably extirpated by glaci- seven different biogeographical regions. The four largest
ation. The species of the wet forests on the west coast families are at the bottom with from bottom to top:
of Mexico are restricted from advancing northward Coenagrionidae, Libellulidae, Gomphidae and Aeshnidae

123
Hydrobiologia (2008) 595:351–363 357

800 ing recent or temporary habitats. The extent of tropical

700 forest in Africa is believed to have contracted
600
substantially during periods with a cooler and drier
500
400 climate. As a consequence the ‘old’ African fauna
300 seems to be largely gone, although relicts remain in
200 isolated areas that were apparently more stable.
100
Examples are the genera Pentaphlebia (Amphipterygi-
0
dae) and Nubiolestes (Perilestidae) in the Cameroon
50

70

90

10

30

50

70

90

10

30

50

70

90
17

17

17

18

18

18

18

18

19

19

19

19

19
highlands and Coryphagrion (Pseudostigmatidae) in
Fig. 4 Rate of description of new taxa in Odonata the East Coast forests, which all have their only
relatives in tropical America. The families Synlestidae
and Megapodagronidae, which have a global but rather
uplands, Ozarks, Great Plains, and Pacific coast. fragmented distribution, are largely restricted to South
Stream-dwelling gomphids are especially likely to Africa and Madagascar, respectively. On the other
show restricted distributions and diversification, and hand, the present-day extent of forest and other tropical
they comprise the largest odonate family in the habitats, such as the continent’s famous savannahs, has
eastern Nearctic. However, many odonate species, allowed remarkable speciation in a few genera (e.g.,
both northern and southern, are wide-ranging over the Chlorocypha, Pseudagrion, Paragomphus, Phyllo-
entire moist eastern half or all across the continent. macromia, Orthetrum and Trithemis). These groups
Others are restricted to the West, often both arid and often have strong Asian affinities, suggesting palaeo-
humid parts of it, as ultimately it is the presence of tropic faunal exchange followed by rapid radiation in
water bodies that determines their distributions. Some periods with a more favourable climate. A small but
of even the largest odonate families appear to show interesting element in the fauna of the eastern coast and
different origins in the Nearctic, for example coe- Indian Ocean islands are genera of probably Papuan-
nagrionids and libellulids mostly from the tropics and Australian origin (Hemicordulia, Teinobasis), that
gomphids and corduliids mostly from northern lati- probably reached Africa by wind-aided trans-oceanic
tudes. There is a substantial boreal fauna; Canada dispersal. The highest odonate diversity, as well as the
holds 208 species, but many of them are restricted to greatest number of range-restricted species, is found in
the southeastern border region, including tropical the Guineo-Congolian forest, which stretches from
genera such as Hetaerina, Argia, and Pantala (Ca- Senegal to western Kenya. The richest area is the
tling et al., 2005). Special features of the Nearctic Cameroon highlands and the surrounding Lower
include the presence of two petalurids, a Pacific Guinea lowland forest. The Upper Guinean forest,
Northwest montane species (Tanypteryx hageni) Congo Basin and Albertine Rift are other core areas
with nearest relative in Japan and a southeastern within this forest belt. Outside it, coastal East Africa
lowland species (Tachopteryx thoreyi); certain genera (including the Eastern Arc Mountains), the Ethiopian
(Tanypteryx, Lanthus, Stylogomphus and Hagenius) highlands and South Africa are notable for their
that show a distinct relationship between eastern Asia endemism. Although the approximately 175 odonate
and eastern Nearctic; and a very recent radiation of species of Madagascar include distinctly Afrotropical
Enallagma (Brown et al., 2000). elements, 60% of Anisoptera and almost 95% of
Zygoptera species are endemic. Endemism and diver-
Afrotropical sity is greatest on the island’s wet eastern coast.

It is notable that, among tropical faunas, the Afrotrop- Oriental

ical fauna is relatively poor and its composition is
nearest that of the Holarctic, with few families and a The Oriental region is, together with the Neotropical
large proportion of Coenagrionidae and Libellulidae region, by far the most species-rich of the eight
(Dijkstra & Clausnitzer, 2006). This may be explained regions recognised here. In China the Oriental and
by the relatively unstable climatological history of the Palaearctic faunas merge gradually along a climatic
continent, which favoured species capable of colonis- gradient. The Chlorogomphidae and Euphaeidae are

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358 Hydrobiologia (2008) 595:351–363

largely confined to the Oriental region although both confined to the region or showing a relict distribution.
have outlying species occurring in the Palaearctic, For several families a large percentage of the world
and several families such as the Chlorocyphidae, fauna is restricted to the Australasian region: Aus-
Platycnemididae, Platystictidae and Pseudolestidae tropetaliidae (36%), Isostictidae (89%), Lestoideidae
are exceptionally well represented. Within the region, (69%), Petaluridae (60%) and Synthemistidae (81%).
several loosely defined subregions, each with a Hemiphlebiidae and Cordulephyinae (Corduliidae) are
characteristic dragonfly fauna, may be recognised: both endemic for continental Australia. The Austrope-
i.e., the Indian subcontinent, Sundaland, the Philip- taliidae are only shared with the southern Andes and
pines, and the main landmass of southeast and east are therefore believed to be of Gondwanian origin. The
Asia (including tropical and subtropical China, but Petaluridae and the Synlestidae are good examples of
excluding the Malaya peninsula). The latter subre- families showing a relict distribution. The majority of
gion exhibits the highest diversity in both species and dragonflies of the Australian continent occur in the
genera of the entire Oriental region, presumably eastern Great Dividing Range and in the adjacent
owing to its large area, numerous mountain ranges narrow coastal strip to the east of this, and in the wetter
intersected by major rivers, and mosaic of forest parts of the southwest. Greatest diversity is to be found
types. Particularly speciose is the area including the in the north-eastern wet tropics of Queensland. The dry
north of Thailand, Laos and Vietnam together with interior of the continent has a depauperate fauna of
tropical China, recognised by some as distinct widespread eurytopic species. The New Zealand fauna
faunistic sub-region (van Tol & Rozendaal, 1995; is poor with only 17 species (Rowe, 1992) including
Wilson & Reels, 2003). Within the Indian sub-region two species of Petaluridae. New Guinea is very
the greatest number of species and endemics occurs species-rich with a high percentage of endemics,
in tropical forest refugia. Richest are the tropical and owing to the perhumid tropical conditions and a highly
subtropical forests to the south of the Himalayas, dissected, mountainous topography that creates
including Sikkim, North Bengal and the Khasi Hills, numerous isolated stream systems, each including a
with other centres of diversity in the Western Ghats wide altitudinal range. New Guinea was formed during
and Nilgiris and the wet south-western and central the mid-late Caenozoic when the northward moving
part of Sri Lanka (Lahiri, 1989; Bedjanič, 2004). Australian plate collided with island arcs to the north,
Extensive semi-arid parts of the subcontinent host a resulting in massive uplifting and orogenesis. The
depauperate and unexceptional fauna. Present-day island arcs were part of a complex archipelago that
Sundaland is divided into several large land masses probably played a part in faunal exchange between the
which were contiguous as recently as 8,000 years ago Oriental region and the Australasian region, resulting
when sea levels were lower. Highest levels of in unexpected affinities between the Philippines and
endemicity and species richness occur in north New Guinea (van Tol & Gassmann, 2005). New
Borneo among forest stream dwellers in montane Guinea and Australia were connected as recently as
and mixed dipterocarp forest, but Java, Sumatra and 8,000 years ago and generally have strong biogeo-
the Malay peninsula all host distinctive faunas. The graphical affinities. They share a high diversity of
fauna of the Philippines has a high number of Megapodagrionidae, Isostictidae and Synthemistidae.
endemics (more than 60% of the named species) However considering their shared history the differ-
sharing elements with both the Oriental and the ences in the dragonfly fauna is remarkable. Especially
Australasian fauna. Its numerous islands have facil- striking is the absence of Platystictidae and Platycne-
itated speciation, resulting in a high number of mididae in Australia and conversely the virtual lack of
endemic species in genera such as Drepanosticta, Gomphidae and Brachytroninae (Aeshnidae) in New
Amphicnemis, Teinobasis, Risiocnemis and Oli- Guinea (Lieftinck, 1949). The Moluccas and Lesser
goaeschna (Hämäläinen & Müller, 1997). Sundas (Nusa Tenggara) have probably never been
connected to either the Oriental region or the Austral-
Australasian region asian region. The islands of Lesser Sundas have a drier
climate than the rest of the Indonesian archipelago but
The Australasian dragonfly fauna is very distinct with a their faunas are generally commensurate with island
strong representation of small families either largely area. Most of the species on these islands, including the

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Hydrobiologia (2008) 595:351–363 359

many endemics, are of Oriental origin (Lieftinck, rather than biogeographical. Nevertheless, the two
1953). The Moluccan fauna is largely derived from faunas are quite distinct, with a strong faunal break
New Guinea, is rather depauperate, and is perhaps most at middle elevations around the Mexican Plateau,
notable for its lack of the genus Neurobasis (Calo- many Nearctic species in temperate habitats on that
pterygidae), present in New Guinea, Sulawesi and the plateau, and tropical species surrounding it in the
Philippines. The island of Sulawesi was formed by the lowlands (Paulson, 1982). Dispersal was apparently
collision of several elements of Laurasian origin and much greater from south to north when Panama
Gondwanian origin. The dragonfly fauna is therefore a emerged in the Pliocene to provide a pathway
blend of species of Australasian and Oriental origin, between the continents, and that dispersal continues
although the latter dominate (van Tol & Gassmann, today. The Polythoridae, Dicteriadidae and Neope-
2005). No current review of Sulawesi dragonflies is taliidae are endemic to the region, the latter confined
available, but it is known that the fauna is less species- to the southern Andes while the former two are
rich than might be expected (van Tol, 1987) The family distributed in the tropics. Largely confined to this
Chlorocyphidae shows exceptional higher-level diver- region are the Austropetaliidae, Perilestidae and
sity, as does Borneo, which perhaps dates back to the Pseudostigmatidae. The latter includes 18 species of
most recent connection of the two land masses 42 mya. very elongate spider-eating, phytotelmata-breeding
damselflies which are among the most remarkable
Pacific odonates. Significant regions of odonate diversifica-
tion include the Mexican Plateau, Chiapas to
As might be expected, the Pacific is species poor. Honduras highlands, Costa Rica-Panama highlands,
Species present can be divided into two groups: those northern Andes, eastern Andean foothills, tepuis of
with a very small area of distribution, being often the Guyana Shield, Guyana lowlands, Atlantic
confined to a single island or island group, and highly forests of Brazil, Rio Paraná basin, and southern
vagile eurytopic species which occur on most Pacific Andes. In the last, Gondwanian groups, including
islands, and which generally also occur throughout the Austropetaliidae, Neopetaliidae, Petaluridae, and
much of the Oriental or Australasian regions (or Gomphomacromia, are prominent. This leaves the
both). Even in Hawaii this phenomenon occurs, huge Amazon basin, poorly known but presumably
although the widespread species originate from the with its own regions of endemism. The Neotropical
Americas. Both the widespread species and the fauna equals that of the Oriental region in both
endemics belong mainly to the Coenagrionidae and modern (species) and ancient (family) diversity. The
the Libellulidae. In the Coenagrionidae the colonisa- complexity of the mountain ranges extending from
tion of an island or group of islands was often Mexico to Chile and the varied climates along their
followed by speciation events leading to a group of length have produced a great variety of odonate
closely related species. This has occurred on Hawaii habitats, as well as providing repeated opportunities
(Megalagrion), Pohnpei (Teinobasis), Fiji (Nesobasis for speciation, and Argia, with 108 named species,
and Melanesobasis) and Samoa (Pacificagrion and is the star of this show. Other characteristic
Amorphostigma). An exception to this pattern is New neotropical genera that have diversified widely in
Caledonia, which drifted away from continental the region include Heteragrion, Palaemnema, Acan-
Australia at the end of the Cretaceous, and is thagrion, Telebasis, Phyllogomphoides, Progom-
moderately species rich. It has an interesting fauna phus, Erythrodiplax and Micrathyria. High
showing distinct affinities with Australia and New biodiversity is the rule for all of the countries in
Guinea and has numerous endemic species and this region, but nevertheless, the Neotropical fauna
several endemic genera (Davies, 2002). is the least known in the world. The highest known
local diversity of odonates is in South America, with
Neotropical 186 species at a single site in southern Peru. Much
of the fauna of the West Indies comes from adjacent
Although North and South America have numerous Mexico and South America, but the large Greater
genera and species in common, this is primarily Antillean islands have numerous endemics, includ-
because the boundary between them is political ing Hypolestes of poorly known affinities.

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Table 2 Examples of areas

Biogeo- Name of area Number of Number of Estimate of
with a high number of
graphical species endemic species endemism (%)
endemic dragonflies
region

Afrotropical Ethiopia 96 12 13
South Africa 160 30 19
Madagascar 175 135 77
Oriental Taiwan 142 21 15
Borneo 272 124 46
Hainan 127 20 16
Sri Lanka 116 53 46
Palaearctic Northwest Africa (Morocco, 70 4 6
Algeria and Tunisia)
Japan 215 74 34
Neotropical Cuba 80 5 6
Costa Rica 265 32 12
Venezuela 489 90 18
Australasian Sulawesi 124 55 44
New Zealand 17 10 59
Pacific Hawaii 36 26 72
New Caledonia 55 22 40

Antarctica faunas with both high absolute and relative endemism

are mainly found in moist tropical forests. Although
No species are known from this region and it is at present there is no sound basis for identifying the
unlikely that any species of dragonfly will reproduce most important areas of endemism, it goes without
there although it is not impossible that some species question that the faunas of the islands of New Guinea,
might be found as vagrants. Sulawesi, Sri Lanka and Madagascar are exception-
ally rich in endemics (see Table 2). It is noteworthy
that the percentage of endemic Zygoptera is almost
Areas of endemicity always much higher than the percentage of endemic
Anisoptera. Examples of this are Madagascar (60% in
No well-based review of areas with high endemism of Anisoptera, 95% in Zygoptera), the Philippines (31%,
dragonflies is available. However, this is intended in 86%) and Sri Lanka (30%, 68%).
the near future as part of a Global Dragonfly
Assessment. Regional projects to identify areas of
endemism carried out so far include an analysis of Human-related issues
endemism in freshwater biotas partly based on
Zygoptera for New Guinea and on Zygoptera and Dragonflies have little economic value, although they
Anisoptera in eastern Africa (Polhemus et al., 2004; are used as food and as magical or medicinal
Darwall et al., 2005) and are presently being prepared resources at a local scale, and to an unknown extent
for southern and western Africa by the IUCN may influence populations of disease vectors. The
Freshwater Biodiversity Assessment Programme. group features prominently in nature management in
Some examples of levels of endemism in different the temperate regions of the world (Westfall & May,
regions are summarised in Table 2, showing large 1996; Kosterin et al., 2004; Sahlén et al., 2004) and
inter-area differences between areas in absolute and they are often used as indicators for environmental
relative numbers of endemic species. Species in the health and conservation management (Corbet, 1999).
temperate region have wide distributions and the Their sensitivity to structural habitat quality (e.g.,
percentage of localised species is low, whereas forest cover, water chemistry) and amphibious habits

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make dragonflies well suited for use in evaluating landscapes are generally better able to colonise new
environmental change in the long term (biogeogra- habitats than species confined to forest, and therefore
phy, climatology) and in the short term (conservation have wider ranges on average and seem to be less
biology), both above and below the water surface influenced by habitat alteration. A clear exception is a
(e.g., Clark & Samways, 1996; Sahlén & Ekestubbe, number of South African endemics which declined
2001; Clausnitzer, 2003; Foote & Hornung, 2005; due to shading of their habitat by invasive alien trees
Osborn, 2005). Dragonflies are often used in both (Kinvig & Samways, 2000), and recovered after
fundamental and applied research because of the removal of the trees (Samways, 2004). Destruction of
relative ease with which they can be observed and tropical forest is probably the most important threat
their broad array of behaviours. In many regions to global odonate diversity, potentially resulting in
reliable identification literature is available, so spe- the extinction of numerous species. Unfortunately
cies can be determined fairly easily by the layman. these species are often poorly known, making it
This enables mapping schemes conducted by volun- difficult to say whether a species is genuinely rare or
teers, facilitating the use of distributional data on merely overlooked. Evaluating the conservation sta-
dragonflies in management. From a global perspec- tus of most naturally rare species is hardly possible.
tive, odonates are among the best known insect Examples of data deficiency are known from Africa
groups with respect to taxonomy and distribution, (e.g., Dijkstra & Clausnitzer, 2006), South America
and, apart from butterflies, probably no other group of (Paulson, 2006), the Oriental region (Orr, 2004) and
insects receives so much attention from the general New Guinea. More fieldwork is thus essential to
public and has so many organisations devoted to its establish the true ranges of these species and to
study. An overview of the conservation and research determine areas of endemism within larger tropical
status of the world’s Odonata can be found in forest areas. There is, however, no doubt that species
Clausnitzer & Jödicke (2004). confined to small remnants of forest in areas under
Many species in the temperate region have shown high human pressure are endangered. Examples of
a dramatic decline in distribution and abundance such sites include many of the Philippine islands,
since the second half of the 20th century (Westfall & Hawaii, the small pockets of forest in the Eastern
May, 1996; Sahlén et al., 2004; Inoue, 2004). This Arc Mountains of East Africa and the Caribbean
has been caused by habitat destruction, eutrophica- islands of Cuba, Hispaniola and Jamaica, but a
tion, acidification and pollution of aquatic habitats in well-founded overview of threatened areas of
general, and the canalisation of streams and rivers. high importance for dragonflies is wanting. Espe-
Most of these species are not under immediate threat cially susceptible are species depending on forest
of extinction as they have wide ranges. A marked on small islands such as those of the Seychelles
exception comes from the Ogasawara Islands, Japan, (Samways, 2003). Here the disappearance of
where five endemic species are on the verge of forest-cover not only results in alteration of the
extinction due to the introduction in the 1980s of an habitat but also may change precipitation pat-
Anolis lizard (Karube, 2005). At least in parts of terns.
Europe, some of the species considered threatened Dragonflies have shown to be useful for nature
recovered since the 1990s as result of improved water management and conservation, and recently an
management. Recently it has become evident that increased effort is being made to make information
many dragonflies of temperate regions are respond- on dragonflies available for both scientists and
ing, both in distribution and phenology, to global policymakers. Important initiatives taken are the
climate change (Ott, 2001). The ranges of common update of the IUCN red list, the ‘Pan-Africa
and widespread southern species are expanding in Freshwater Biodiversity Assessment’ started by the
Europe but there is as yet no strong evidence that IUCN (Darwall et al., 2005), which includes drag-
northern species are decreasing as a result of the onflies among other taxa, and the ‘Global Dragonfly
rising temperatures, as might be expected. Assessment’ initiated in 2005. The latter should
Most of the world’s dragonflies are restricted to hopefully result in a more detailed overview of the
the tropics, especially to forest, where the diversity of areas of endemism and conservation priority in the
the group is greatest. Tropical species of open coming years.

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