Balian E.V., L?v?que C., Segers H., Martens K. (Eds.) Freshwater Animal Diversity Assessment

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Antarctica

No species are known from this region and it is

unlikely that any species of dragonﬂy will reproduce

there although it is not impossible that some species

might be found as vagrants.

Areas of endemicity

No well-based review of areas with high endemism of

dragonﬂies is available. However, this is intended in

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

endemism in freshwater biotas partly based on

Zygoptera for New Guinea and on Zygoptera and

Anisoptera in eastern Africa (Polhemus et al., 2004;

Darwall et al., 2005) and are presently being prepared

for southern and western Africa by the IUCN

Freshwater Biodiversity Assessment Programme.

Some examples of levels of endemism in different

regions are summarised in Table 2, showing large

inter-area differences between areas in absolute and

relative numbers of endemic species. Species in the

temperate region have wide distributions and the

percentage of localised species is low, whereas

faunas with both high absolute and relative endemism

are mainly found in moist tropical forests. Although

at present there is no sound basis for identifying the

most important areas of endemism, it goes without

question that the faunas of the islands of New Guinea,

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

always much higher than the percentage of endemic

Anisoptera. Examples of this are Madagascar (60% in

Anisoptera, 95% in Zygoptera), the Philippines (31%,

86%) and Sri Lanka (30%, 68%).

Human-related issues

Dragonﬂies have little economic value, although they

are used as food and as magical or medicinal

resources at a local scale, and to an unknown extent

may inﬂuence populations of disease vectors. The

group features prominently in nature management in

the temperate regions of the world (Westfall & May,

1996; Kosterin et al., 2004; Sahle

n et al., 2004) and

they are often used as indicators for environmental

health and conservation management (Corbet, 1999).

Their sensitivity to structural habitat quality (e.g.,

forest cover, water chemistry) and amphibious habits

Table 2 Examples of areas

with a high number of

endemic dragonﬂies

Biogeo-

graphical

region

Name of area Number of

species

Number of

endemic species

Estimate of

endemism (%)

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,

Algeria and Tunisia)

70 4 6

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

360 Hydrobiologia (2008) 595:351–363

123

make dragonﬂies well suited for use in evaluating

environmental change in the long term (biogeogra-

phy, climatology) and in the short term (conserv ation

biology), both above and below the water surface

(e.g., Clark & Samways, 1996; Sahle

n & Ekestubbe,

2001; Clausnitzer, 2003; Foote & Hornung, 2005;

Osborn, 2005). Dragonﬂies are often used in both

fundamental and applied research because of the

relative ease with which they can be observed and

their broad array of behaviours. In many regions

reliable identiﬁcation lite rature is available, so spe-

cies can be determined fairly easily by the layman.

This enables mapping schemes conducted by volun-

teers, facilitating the use of distributional data on

dragonﬂies in management. From a global perspec-

tive, odonates are among the best known insect

groups with respect to taxonomy and distribution,

and, apart from butterﬂ ies, probably no other group of

insects receives so much attention from the general

public and has so many organisations devoted to its

study. An overview of the conservation and research

status of the world’s Odonata can be found in

Clausnitzer & Jo

dicke (2004).

Many species in the temperate region have shown

a dramatic decline in distribution and abundance

since the second half of the 20th century (Westfall &

May, 1996; Sahle

n et al., 2004; Inoue, 2004). This

has been caused by habitat destruction, eutrophica-

tion, acidiﬁcation and pollution of aquatic habitats in

general, and the canalisation of streams and rivers.

Most of these species are not under immediate threat

of extinction as they have wide ranges. A marked

exception comes from the Oga sawara Islands, Japan,

where ﬁve endemic species are on the verge of

extinction due to the introduction in the 1980s of an

Anolis lizard (Karube, 2005). At least in parts of

Europe, some of the species considered threatened

recovered since the 1990s as result of improved water

management. Recently it has become evident that

many dragonﬂies of temperate regions are respond-

ing, both in distribution and phenology, to global

climate change (Ott, 2001). The ranges of common

and widespread southern species are expanding in

Europe but there is as yet no strong evidence that

northern species are decreasing as a result of the

rising temperatures, as might be expected.

Most of the world’s dragonﬂies are restricted to

the tropics, especially to forest, where the diversity of

the group is greatest. Tropical species of open

landscapes are generally better able to colonise new

habitats than species conﬁned to forest, and therefore

have wider ranges on average and seem to be less

inﬂuenced by habitat alteration. A clear exception is a

number of South African endemics which decl ined

due to shading of their habitat by invasive alien trees

(Kinvig & Samways, 2000), and recovered after

removal of the trees (Samways, 2004). Destruction of

tropical forest is probably the most important threat

to global odonate diversity, potentially resulting in

the extinction of numerous species. Unfortunately

these species are often poorly known, making it

difﬁcult to say whether a species is genuinely rare or

merely overlooked. Evaluating the conservation sta-

tus of most naturally rare species is hardly possible.

Examples of data deﬁciency are known from Africa

(e.g., Dijkstra & Clausnitzer, 2006), South America

(Paulson, 2006), the Oriental region (Orr, 2004) and

New Guinea. More ﬁeldwork is thus essential to

establish the true ranges of these species and to

determine areas of endemism within larger tropical

forest areas. There is, however, no doubt that species

conﬁned to small remnants of forest in areas under

high human pressure are endangered. Examples of

such sites include many of the Philippine islands,

Hawaii, the small pockets of forest in the Eastern

Arc Mountains of East Africa and the Caribbean

islands of Cuba, Hispaniola and Jamaica, but a

well-founded overview of threatened areas of

high importance for dragonﬂies is wanting. Espe-

cially susceptible are species depending on forest

on small islands such as those of the Seyc helles

(Samways, 2003). Here the d isappearance of

forest-cover not only results in alteration of the

habitat but also may change precipitation pat-

terns.

Dragonﬂies have shown to be useful for nature

management and conservation, and recently an

increased effort is being made to make information

on dragonﬂies available for both scientists and

policymakers. Important initiatives taken are the

update of the IUCN red list, the ‘Pan-Africa

Freshwater Biodiversity Assessment’ started by the

IUCN (Darwall et al., 2005), which includes drag-

onﬂies among other taxa, and the ‘Global Dragonﬂy

Assessment’ initiated in 2005. The latter should

hopefully result in a more detailed overview of the

areas of endemism and conservation priority in the

coming years.

Hydrobiologia (2008) 595:351–363 361

123

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123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of stoneﬂies (Plecoptera; Insecta)

in freshwater

Romolo Fochetti Æ

Jose

Manuel Tierno de Figueroa

 Springer Science+Business Media B.V. 2007

Abstract Plecoptera, or stoneﬂies, is a small order

of hemimetabolous insects: according to our data,

more than 3,497 species have been described so far in

the world. The total number of species has enor-

mously increased in the last 30 years (2,000 species

estimated in 1976) and, if the trend continues, then it

will nearly double in the near future. The order is

divided into the suborders Arctoperlaria and Antarct-

operlaria, and includes 16 families: 12 arctoperlarian

and 4 antarctoperlarian. The Arctoperlaria account

for a total number of 3,179 species, and Antarctop-

erlaria, only 318 species. The total number of genera

is 286. We give in this article the estimated number

of species for each family. The fauna and diversity of

stoneﬂy in North America (650 species reported) and

Europe (426 species) are best known. Nevertheless,

in the last 25 years, a mean of 2.6 Plecoptera species

per year were described in Europe. Stoneﬂy-faunas of

Australia (191 species, Tasmania included) and New

Zealand (104 species) are relatively well-known,

while our knowledge of the Plecoptera of Central and

South America (95 and 378 species respectively) is

poor and still not representative of the real diversity.

Africa has a reduced stoneﬂy fauna (126 species).

Asian stoneﬂy diversity (approximately 1,527 spe-

cies) is much greater than that of Europe or North

America despite the fact that, except for Japan and

Asiatic Russia that have been well studied, our

knowledge of the remaining Asiatic areas is extre-

mely poor. Even though our data indicate the

Holarctic Region as the diversity hot-spot for the

order, the analysis of the speciﬁc diversity divided by

family suggests also an important role of tropical

stoneﬂies.

Keywords Plecoptera  Distribution 

Diversity  Zoogeographical regions

Introduction

Plecoptera is a small order of hemimetabolous

insects, commonly called stoneﬂies, with more than

3,497 described species (this article). Stoneﬂies are

distributed over all continents excep t Antarctica, and

constitute a signiﬁcant ecological component of

running water ecosystems. Their ecological require-

ments greatly limit the dispersal capacity of the

nymphs and, because adults have reduced ﬂight

ability, stoneﬂies show a high percentage of

Guest editors: E.V. Balian, C. Le

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

R. Fochetti (&)

Dipartimento di Scienze Ambientali, Universita

della

Tuscia, Largo dell’Universita

snc, 01100 Viterbo, Italy

e-mail: fochetti@unitus.it

J. M. Tierno de Figueroa

Departamento de Biologia Animal, Facultad de Ciencias,

Universidad de Granada, 18071 Granada, Spain

123

Hydrobiologia (2008) 595:365–377

DOI 10.1007/s10750-007-9031-3

endemism. Due to the growing pollution and alter-

ation of water courses and to the high stenoecity of

Plecoptera, numerous stoneﬂy species are reduced to

small isolated populations and many others have

already gone extinct. Stoneﬂies are probably one of

the most endangered groups of insects (Fochetti &

Tierno de Figueroa, 2004).

Plecoptera can be easily recognized by several

morphological characters: soft body, three segmented

tarsi, elongate ﬁliform antennae, mandibulated

mouthparts, two compound eyes, two or three ocelli,

two usually long cerci, 10-segmented abdomen with

vestiges of the eleventh segment. Adults have two

pair of membranous large wings (sometimes reduced

or absent), and subequal fore and hind wings (hind

wings slightly wider) that fold horizontally over and

around abdomen whe n at rest (hence the name:

plecos = folded; pteros = wings) (Fig. 1A). Nymphs

are similar to adults (Fig. 1B), with a closed tracheal

system with or without ﬁlamentous gills. When

present, gills are located on different parts of the

body.

Plecoptera nymphs are aquatic and live mainly in

cold, well-oxygenated running waters, although

some species can also be found in lakes. Few

species are adapted to terrestrial life in Sub-Antarc-

tic areas: Hynes (1976) reported the tendency of

southern hemisphere nymphs to leave the water.

However, the increasing number of stoneﬂies

described from the tropics and their high rate of

endemicity can modify the common belief that

Plecoptera are cold-water specialists, and suggest

instead that the true hot-spot for Plecoptera diversity

are tropical areas (Zwick, pers. com.). The life cycle

of stoneﬂies lasts for one or more years, but there

are also bi- or tri-voltine species . Nymphal or egg

diapause is not uncommon. The nymphs can moult

up to 33 times before emerging. They feed on

animal or vegetable matter as collectors, scrapers,

shredders or predators.

Fig. 1 A-Adult of Nemoura. and 1 B-Nymph of Nemoura

366 Hydrobiologia (2008) 595:365–377

123

Adults are terrestrial or sub-aerial, with the

exception of Capnia lacustra Jewett, whose adults

have been collected only at a depth of 60–80 m in the

Lake Tahoe (USA) (Jewett, 1963), and the adult

females of Zapada cinctipes (Bank) that can remain

under water for a long time (20–60 min) for ovipos-

iting (Tozer, 1979). Adults lifespan is short, from a

few days to weeks; they display complex behaviours

related to mate searching (such as the use of

intersexual vibrational communication in Arctoper-

larian stoneﬂies) and oviposition (Stewart, 1994). The

adult feeds on pollen, lichens, cyanobacteria, etc.,

although the adults of the largest species do not seem

to feed (Tierno de Figueroa & Fochetti, 2001).

Females produce 25–3,000 eggs, and lay their egg-

clutches, or rarely each egg separately, on the water.

Parthenogenesis and ovoviviparity have been also

described for some species.

Species diversity

According to our data, approximately 3,500 species

of Plecoptera have been reported so far in the world

(Tables 1, 2). They are divided as: North America

650 species (Stark & Baumann, 2005), Central

America 95 species, South America 378 species

(Heckman, 2003), Europe 426 species (Fochetti &

Tierno de Figueroa, 2004), Africa 126 species (31

from Mediterranean Africa, with only 12 endemic

species, 50 from tropical Africa, 33 from South

Africa and 12 from Madagascar), Asia approximately

1,527 species [(about 784 from South-East Asia = 31

Taiwan, 32 Korea, 25 Philippines, 39 Borneo, 36

Indonesia, 17 Malaysia, 29 Thailand, 20 Vietnam,

two Laos, three Myanmar, 28 Bhutan, two Bangla-

desh, 10 Sri Lanka, 11 2 India, 63 Nepal, 20 Pakistan,

nine Afghanistan, 306 Japan, no species from Cam-

bodia; Sivec & Yang 2001), China 350 (?) species

(deduced from Yuzhou & Junhua, 2001, who report

231 species of Perlidae), West Asia about 114 species

(two Syria, two Israel, 25 Iran, 15 Lebanon, 70

Asiatic Turkey), Asiatic Russia 279 species (179

Nemouroidea and about 100 Perloidea). Although a

small number of species are shared by several

countries, we consider the total sum as a good

approximation)], Australia 191 species (Michaelis &

Yule, 1988), New Zealand 104 species (McLellan,

2006).

Very few species are shared among zoogeograph-

ical regions (mainly between Nearctic and Palaearctic

and between Palaearctic and Oriental) or among

continents, i.e. at borders between Africa and Europe

or at the borders between Europe and Asia. Australia

and New Zealand are obviously isolated; deserts or

dry lands prevent stoneﬂy migra tion from North to

Central and South America, and from the Palaearctic

to the Ethiopic Regions. Thus, the total number of

described species shoul d not be too far from the

reported one.

In comparison to the papers by Hynes (1976) and

Zwick (1980) who reported about 2,000 described

stoneﬂy species, the total number has enourmously

increased in the last 25 years and, if the trend

continues then it will nearly double in the near future.

The fauna and diversity of stoneﬂy in North

America and Europe are best known. Nevertheless,

new species have been described from these areas at a

high rate: in the last 25 years, a mean of 2.6

Plecoptera species per year were described in Europe

(Fochetti & Tierno de Figueroa, 2005). Stoneﬂy-

faunas of Australia (Tasmania included) and New

Zealand are relatively well-known, while our knowl-

edge of the Plecoptera of Central and South America

is poor and still not representative of the real

diversity.

Asian stoneﬂy diversity is much greater than that

of Europe or North America. This is true despite the

fact that, except for Japan and Asiatic Russia that

have been well studied, our knowledge of the

remaining Asiatic areas is extremely poor. Our

greatest lack of knowledge and the highest uncer-

tainty regard Asiatic faunas occurs in the Oriental

Region. For instance, according to Yuzhou & Junhua

(2001) over 230 species of Perlidae, the most

speciose family in the country, have so far been

described from China. The identity of many of these

species is questionable and cannot be checked

because most of the types were lost or destroyed;

nonetheless, intensive future research will certainly

lead to the discovery of hundre ds of new species in

this country. In fact, a few intensive collecting trips to

Taiwan almost doubled its stoneﬂy-fauna (Sivec &

Yang, 2001).

In Africa, a reduced number of species were

collected, but we do not expect a great increase in

Plecoptera diversity in the future. The results of

recent studies on the systematics of the tropical tricky

Hydrobiologia (2008) 595:365–377 367

123

genus Neoperla might increase the species list and

some new species could be described from Mediter-

ranean countries (Morocco, Algeria, Tunisia), but the

total stoneﬂy diversity will not increase to the level of

the other continents.

As we expected, the analysis of Plecoptera diver-

sity and distribution shows their well-known

ecological requirements as: they are absent from arid

or semi-arid areas everywhere in the world. There are

almost no records from West Australia, no records at

all for Saudi Arabia, Yemen, Oman, Emirates,

Kuwait and Iraq in Asia, and stoneﬂy diversity in

Africa is greatly limited by deserts and dry lands.

The order is divided into the suborders Arctoper-

laria and Antarctoperlaria (see the phylogeny chapter

below), and includes 16 families: 12 arctoperlarian

and four antarctoperlarian. The Arctoperlaria account

for a total number of 3,179 species, and Antarctop-

erlaria, only 318 species. They are listed in Table 2

with the estimated number of species for each family.

The total number of genera is 286 (Table 3). Even

though Table 1 is incomplete because it lacks details

on the distribution of Chinese families and Far East

Russia species, Perlidae is the most diverse family,

with more than 1,000 described species. Since the last

estimate (Zwick, 1980), the number of perlid species

Table 1 Number of Plecoptera species in each family and each continent

NAm CAm SAm Eu AF AS

NZ AUS

Eustheniidae 2 4 15

Diamphipnoidae 5

Austroperlidae 4 1 10

Gripopterygidae 73 70 134

Pteronarcyidae 10 1 2

Peltoperlidae 22 24

Styloperlidae 8

Perlodidae 125 3 62 3 118

Perlidae 82 71 277 18 55 546

Chloroperlidae 95 2 19 1 89

Scopuridae 8

Taeniopterygidae 34 41 2 65

Notonemouridae 17 43 29 32

Nemouridae 71 9 132 8 413

Capniidae 156 8 21 4 126

Leuctridae 55 1 133 10 128

Total 650 95 378 426 126 1,527 104 191

Asiatic Russia and China data have been arbitrarily subdivided (NAm = North America; CAm = Central American; SAm = South

America; Eu = Europe; AF = Africa; AS = Asia; NZ = New Zealand; AUS = Australia)

Table 2 Number of Plecoptera species in each family and

each zoogeographical region

NA NT AT OL

AU World

Eustheniidae 2 19 21

Diamphipnoidae 5 5

Austroperlidae 4 11 15

Gripopterygidae 73 204 277

Pteronarcyidae 10 1 13

Peltoperlidae 22 46

Styloperlidae 8

Perlodidae 63 125 4 311

Perlidae 18 82 348 52 1,049

Chloroperlidae 20 95 2 206

Scopuridae 8

Taeniopterygidae 42 34 142

Notonemouridae 17 43 61 121

Nemouridae 136 71 9 633

Capniidae 21 156 8 315

Leuctridae 138 55 1 327

Total 438

650 474 95 350? 295 3,497

Palaearctic Asia not included. PA: Palaearctic Region, NA:

Nearctic Region, NT: Neotropical Region, AT: Afrotropical

Region, OL: Oriental Region, AU: Australasian Region.

The

distribution of Oriental Plecoptera cannot be unequivocally

established

368 Hydrobiologia (2008) 595:365–377

123

almost doubled, due to the increase of faunistic

research in Asia and, above all, to the improved

knowledge of the systematics of the tropical genera

Neoperla and Anacroneuria. The family Perlidae

spreads over Holarctic, Oriental, Afrotropical and

Neotropical Regions. The most diverse perlid areas

are Asia, and Central and South America. Presently the

Chinese perlid species represent about 25% of the

world Perlidae, making China one of the most impor-

tant areas of the world for perlid diversity (Yuzhou &

Junhua, 2001). The Neotropical perlid fauna currently

includes more than 350 species: Anacroneuria is the

dominant and most diverse stoneﬂy genus, (Stark,

2001) with almost 300 described species.

Perlodidae presently include s about 300 species,

mainly in the Holarctic region. The most speciose

perlodid genus, Isoperla, has a Holarctic distribution

and has more than 100 species, a third of the total. A

similar distribution is shown by the ancient genus

Arcynopteryx [Aubert (1959) stated it is an Ercinic

relict] and by the genus Diura. A. compacta

(McLachlan) is distributed in North America and

Europe, as well as D . bicaudata (Linnaeus) and D.

nanseni (Kempny).

Chloroperlidae include more than 200 species

mainly in the Holarctic region. The genera Sweltsa

and Alloperla have a Nearctic and East Palaearctic

distribution and include s almost 80 species. In

Europe, only four genera and 19 species are present.

Taeniopterygidae is a family present in the

Nearctic and Palaearctic Regions, with almost 150

species. The genus Taeniopteryx (20 species) is

present in Europe and North America (even though

the morphology of the nymphs slightly differs in the

two continents), as well as the genus Oemopteryx.

The genus Doddsia is shared by North America and

Japan. The West Palaearctic genus Brachy ptera

includes more than 30 species.

Nemouridae was consider ed the most diverse

family, with almost 400 described species (Zw ick,

1980). The number current ly is 633 species, but it is

now the second most diverse family due to the

enormous increase of Perlidae. The family has an

Holarctic distribution with some genera reaching the

Oriental Region; the genera Amphinemura (Holarc-

tic; more than 100 species), Nemoura (Holarctic; 175

species) and Protonemura (Palaearctic; 135 species)

account for 2/3 of the whole family. Nemoura arctica

Esben-Petersen has a circum-arctic distribution.

Capniidae and Leuctridae include a similar num-

bers of species, 311 and 321 respectively, both with

mainly Holarctic distribution and a strict systematic

afﬁnity. The most diverse genera are the Holarctic

Capnia (110 species), Allocapnia (Nearctic; 43

species), Mesocapnia (Nearctic and East Palaearctic,

19 species) among Capniidae, and Leuctra (Holarctic,

reaching Northern India; 190 species) and Rhopalop-

sole (East Palaearctic and Oriental; 20 species)

among Leuctridae. The genus Megaleuctra presently

includes six North American and one North Korea

species; it is also known the fossil record in North

Amer

ica

and in Baltic amber.

Notonemouridae (121 species), is considered the

sister taxo n of Nemouridae (but see below), and thus

is an Arctoperlarian family, but shows a disjunct

South African (included Madagascar), South Amer-

ican and Australian (included New Zealand)

distribution. The family may not be monophyletic

(Zwick, 1990) and its biogeography is not fully

understood yet. Austrocercella (15 species) is the

most diverse genus.

Among the remaining Arctoperlaria, the small

family (eight species) of wingless stoneﬂies Scopu-

ridae is known only from Japan and Korea, and the

family Pteronarcyidae (13 species) is present in North

America and East Asia. Peltoperlidae has a disjunct

North American and East Asian distribution (44

species). Styloperlidae a recently instituted family

(eight species), erected from aberrant Peltoperlidae,

is presently known only from China and Taiwan.

Among Antarctoperlaria, Gripop terygidae is the

most diverse family, with 277 described species. It

has a disjunct South American and Australian

(included New Zealand) distribution, but no genera

are shared between the two continents. Dinotoperla

(33 species), Lepto perla (28 species), Riekoperla (26

species) are the most diverse Australian genera,

Zelandobius (27 species) is the most diverse genus of

New Zealand. The remaining Antarctoperlaria fam-

ilies are scarcely diversiﬁed: Diamphipnoidae (ﬁve

species) is a small South American family, Eusthe-

niidae includes two genera in South America (two

species) one genus in New Zealand (four species) and

three genera in Australia (15 species), while

Austroperlidae includ es 10 species in Australia (ﬁve

genera), a single species, Austroperla cyrene

Tylliard, in New Zealand, and four species (th ree

genera) in South America.

Hydrobiologia (2008) 595:365–377 369

123

Among Antarctoperlaria Eustheniidae are preda-

tors, as are Perloidea in the Nor thern hemisphere

Arctoperlaria; Austroperlidae are shredders as are the

Arctoperlaria Pter onarcyidae. Many Gripopterygidae

are scrapers, as are Brachypterainae in the Arctoper-

laria fauna; other Gripopterygidae gener a are

ecologically similar to genera and families of

Nemouroidea.

Phylogeny and historical processes

Stoneﬂies are Pterygota, Neo ptera, Exopterygota

insects. Although they are usually included among

Polyneoptera (possibly they are the sister-group of

the remaining Polyneoptera), the relationships of

Plecoptera with other orders of this subdivision are

poorly understood (Gullan & Cranston, 2005).

Among Polyneoptera, Plecoptera can be identiﬁed

by a combination of mostly primitive characters.

Moreover, only a few derived characters support the

Plecoptera monophyly (Zwick, 2000): in the adult,

gonads form loops with the anterior ends medially

fused; in the male, there is a complex arrangement of

two superimposed seminal vesicles each of which

forms an anterior loop; in the nymph, strong oblique

intersegmental ventrolongitudinal muscles are pres-

ent that support laterally undulating swimming (this

last character also occurs in Zygoptera). Two other

probably derived charact ers have been reported for

Plecoptera include the absence of an ovipositor

(although it can secondarily appea r in some taxa)

and the existence of an accessory circulatory organ

(cercus heart) in Perlidae and Perlodidae, that can be

present in the other families as well.

The order includes 16 families whose relations hips

have been studied by several authors: we will follow

the more recent and widely accepted classiﬁcation by

Zwick (2000) (Fig. 2), which recognizes two large

groups: Antarcto perlaria, present only in the Southern

Hemisphere, and Arctoperlaria, distributed mostly in

the Northern Hemisphere. The ﬁrst taxon includes

four families, the latter includes 12 families belonging

to two subgroups (Systellognatha and Euholognatha)

with six families each one.

The monophyly of Antarctoperlaria is supported

by morphological characters, while monophyly of

Arctoperlaria is supported by ethological and mor-

phological characters related to mate ﬁnding: the

intersexual vibrational communication (drumming,

tremulation, rubbing or combinations of them) and/or

the existence of structures associated to it in the male

eadigyretpoineaT

eadidolreP

eadiinehtsuE

eadionpihpmaiD

eadilreportsuA

eadigyretpopirG

eadirupocS

eadiinpaC

eadirtcueL

eadiruomeN

eadiruomenotoN

eadiycranoretP

eadilrepolytS

eadilrepotleP

eadilreP

eadilreporolhC

SUE

Fig. 2 Plecoptera

phylogeny according to

Zwick (2000). AN:

Antarctoperlaria; AR:

Arctoperlaria; EUS:

Eusthenioidea; GR:

Gripopterygoidea; EU:

Euholognatha; NE:

Nemouroidea; SY:

Systellognatha; PT:

Pteronarcyoidea; PE:

Perloidea. (redrawn from

Zwick, 1990)

370 Hydrobiologia (2008) 595:365–377

123